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PROCEEDINGS 


of the 


Biological Society of 


Washington 


VOLUME 106 
1993 


Vol. 106(1) published 8 April 1993 Vol. 106(3) published 20 September 1993 
Vol. 106(2) published 11 June 1993 Vol. 106(4) published ?? December 1993 


WASHINGTON 
PRINTED FOR THE SOCIETY 


EDITOR 


C. BRIAN ROBBINS 


ASSOCIATE EDITORS 
Classical Languages Invertebrates 
GEORGE C. STEYSKAL Jon L. NORENBURG 


FRANK D. FERRARI 
RAFAEL LEMAITRE 
Plants Vertebrates 
DAvip B. LELLINGER THOMAS A. MUNROE 


Insects 
WAYNE N. MATHIS 


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 1992-1993 


OFFICERS 
President 
STORRS L. OLSON 


President-Elect 
JANET W. REID 


Secretary 
G. DAVID JOHNSON 


Treasurer 
T. CHAD WALTER 


COUNCIL 


Elected Members 
STEPHEN D. CAIRNS JON L. NORENBURG 
RICHARD C. FROESCHNER LYNNE R. PARENTI 
ALFRED L. GARDNER F. CHRISTIAN THOMPSON 


TABLE OF CONTENTS 


Volume 106 


Alonso de Pina, Gloria M. Linca pinita, a new phoxocephalid genus and species (Crus- 
tacea: Amphipoda) from the Argentine continental She] f eee eeeeeeee eens 
Alvarez, Belinda and R. W. M. Van Soest. A new sponge species, Ceratopsion crustosum 
(Demospongiae: Raspailiidae), from deep waters of the Gulf of Mexico 
Baba, Keiji. Anomoeomunida, a new genus proposed for Phylladiorhynchus caribensis 
Mayo, 1972 (Crustacea: Decapeda:'Galatheidae) =. 3422 See eee ee 
Bamber, Roger N. A new species of Kalliapseudes (Crustacea: Tanaidacea: Kalliapseu- 
didae) from Trimica 22 ee ee cS 
Banford, Heidi M. and Bruce B. Collette. Hyporhamphus meeki, a new species of 
halfbeak (Teleostei: Hemiramphidae) from the Atlantic and Gulf coasts of the United 
Stat: eh ec 
Blake, James A. New genera and species of deep-sea polychaetes of the family Nau- 
tiliniellidae from the Gulf of Mexico and the eastern Pacific cee 
Brooks, Daniel R. and Barbara Holcman. Revised classification and phylogenetic hy- 
pothesis for the Acanthostominae Looss, 1899 (Digenea: Opisthorchiformes: Cryp- 
TO@porininiae) ee 8 ee ee ee ee 
Bruce, A. J. Potamalpheops darwiniensis (Crustacea: Decapoda: Alpheidae), the third 
Indo-west Pacific SpGCieS. .c0. =e is a eee 
Campos, Emesto. Systematics and taxonomic remarks on Pinnotheres muliniarum 
Rathbun, 1918 (Crustacea: Brachyura: Pinmothericae) 2. eee cneeeeeeeneneeee 
Campos, Martha R. and Gilberto Rodriguez. Three new species of Strengeriana from 
Colombia (Crustacea: Decapoda: Pseudothelphusidae) eee 
Casanova, Jean-Paul. Spadella japonica, a new coastal benthic chaetognath from Japan 
Castro, Ricardo M. C. Prochilodus britskii, a new species of prochilodontid fish (Os- 
tariophysi: Characiformes), from the rio Apiaca, rio Tapajos system, Mato Grosso, 
| 5) ¢:V A | cee eet omen nome ON Ee MIM. 0 LS ee eeceem yr eee wR TE ee 
Couri, Marcia Souto and Carlos José Einicker Lamas. Revision of Ylasoia Speiser, 
1920 (Insecta? Diptera: Bombyhidae: Lomatinac) = Eee 
Cumberlidge, Neil. Further remarks on the identity of Sudanonautes orthostylis Bott, 
1955 (Crustacea: Decapoda: Potamoidea: Potamonautidae) with comparisons with 
other species from Nigeria and Cameroon 22.26 ee 
Curino, Alejandro C. and Néstor J. Cazzaniga. A new species of freshwater planarian 
from Chile (Platyhelminthes: Tricladida), with a nomenclatural note on Girardia festae 
(Bocore DDG, USS) asc se a a 
Desbruyéres, Daniel and Lucien Laubier. New species of Alvinellidae (Polychaeta) from 
the north Fiji back-arc basin hydrothermal vents (southwestern Pacific) .................... 
Emmons, Louise H. On the identity of Echimys didelphoides Desmarest, 1817 (Mam- 
malia: Rodentia: Echimyidae) 2... Pe eee ee 2 ee re 
Emmons, Louise H. A new genus and species of rat from Borneo (Rodentia: Muridae) 
Erséus, Christer. A new marine species of Smithsonidrilus (Oligochaeta: Tubificidae) 
from the *PlOrida: Keys co. ee Ses a eo ce alee ae ath sd 
Erséus, Christer and Michael R. Milligan. A new species of Uniporodrilus (Oligochaeta: 
Tubificidae) from the Gulf of Mexico coast of Florida, and a phylogenetic analysis of 
11 (ce 1 er ees apron Pe bk oeeiie, SOHN WIN Sei 
Feduccia, Alan and A. Bradley McPherson. A petrel-like bird from the late Eocene of 
Louisiana: earliest record for the order ProcellariifOrrmes 2......::.:.-c-eeeeeeeeecccccceceeeeeeeeeeeeeeeeeeeeeens 
Ferrari, Frank D. and Deborah K. Steinberg. Scopalatum vorax (Easterly, 1911) and 
Scolecithricella lobophora Park, 1970, calanoid copepods (Scolecithrichidae) associ- 
ated witha pelagic tunicate m Monterey Bay 225 
Fugate, Michael. Branchinecta sandiegonensis, a new species of fairy shrimp (Crustacea: 
Anostraca) from western North America 


497-507 


629-632 


102-105 


122-130 


369-384 


147-157 


207-220 


698-704 


92-101 


508-513 


359-365 


57-62 


450-454 


514-522 


633-644 


225-236 


1-4 
752-761 


587-590 


243-250 


749-751 


467-489 


296-304 


Gardner, Alfred L. and Monica Romo R. A new Thomasomys (Mammalia: Rodentia) 
SEE Efe STINE TW 0 Ui (Gam Pe eae ee ees ee 
Ghosh, H. C. and Raymond B. Manning. A new deep-sea crab of the genus Chaceon 
iamnindia (Crustacea: Decapoda: Geryonidac 225 fn) beet 
Goodman, Steven M. and Florent Ravoavy. Identification of bird subfossils from cave 
surface deposits at Anhjohibe, Madagascar, with a description of a new giant Coua 
“SLT OTe) a as 2 ee ee a 
Graves, Gary R. A new intergeneric wood warbler hybrid (Parula americana x Den- 
RIRSIEAICOT OAL AGES aE Tin PENGaG) (ec 88 Venkatesh et 
Graves, Gary R. A new hybrid manakin (Dixiphia pipra x Pipra filicauda) (Aves: 
Pipridae) from the Andean foothills of eastern Ecuador s. 
Hardy, Laurence M. Activity and reproductive patterns of amphibians and reptiles 
from the Engare Ondare River region of central Kenya, during the dry season _. 
Harvey, Michael B. and Eric N. Smith. A new species of aquatic Bufo (Anura: Bufonidae) 
from cloud forests in the Serrania de Siberia, Bolivia 
Hershler, Robert and France Velkovrh. A new genus of hydrobiid snails (Mollusca: 
Gastropoda: Prosobranchia: Rissooidea) from northern South America Ze) 
Hobbs, H. H., II. Cambarus (Jugicambarus) subterraneus, a new cave crayfish (De- 
capoda: Cambaridae) from northeastern Oklahoma, with a key to the troglobitic 
SHLIeICES OF tHe SUOPCHUS JUCICOMDGIUS 20 ee ba 
Hobbs, Horton H., Jr. and Daniel J. Peters. New records of entocytherid ostracods 
infesting burrowing and cave-dwelling crayfishes, with descriptions of two new species 
Holt, Perry C. and Brent D. Opell. A checklist of and illustrated key to the genera and 
species of the Central and North American Cambarincolidae (Clitellata: Branchiob- 
a NR Br Dirt 2s Me Cla ee eee eS th ted emnn A 3 din ded 
Hotchkiss, Frederick H.C. A new Devonian ophiuroid (Echinodermata: @ésaphiuid® 
from New York state and its bearing on the origin of ophiuroid upper arm plates ... 
Humes, Arthur G. and Geoffrey A. Boxshall. Pseudonicothoe branchialis (Crustacea: 
Copepoda: Siphonostomatoida: Nicothoidae), living on the pandalid shrimp Hetero- 
eanpis sibogac olf northwestern Australia (2.2 2 
Jezerinac, Raymond F. A new subgenus and species of crayfish (Decapoda: Cambaridae) 
of the genus Cambarus, with an amended description of the subgenus Lacunicambarus 
Jezerinac, Raymond F. and G. Whitney Stocker. A new species of crayfish (Decapoda: 
Cambaridae) belonging to the genus Cambarus, subgenus Hiaticambarus, from the 
BppePeIk haver dramare Ol West Virginias 2 8 es te ee 
Komai, Tomoyuki. Two new records of the genus Heptacarpus (Crustacea: Decapoda: 
Lupperytigac) trom Japanese waters, tss2lt sy. sree ee el ee 
Kudenov, Jerry D. A new species of Sphaerodoridae (Annelida: Polychaeta) from ae 
EO De Se ni ee a ee £0 ret eee oe 
Leo L., Mariella and Alfred L. Gardner. A new species of a giant Thomasomys (Mam- 
malia: Sigmodontinae) from the Andes of northcentral Peru . e 
Liao, Yulin and David L. Pawson. Caudina intermedia, a new species = sea cucumber 
from the South China Sea (Echinodermata: Holothuroidea: Molpadiida) 
Lima, Idalina Maria Brasil and Cristiana Silveira Serejo. A new species of Benthana 
Budde-Lund from Brazilian caves (Crustacea: Isopoda: Oniscoidea) 
Loffler, Deborah L. and Michael Vecchione. An unusual squid paralarva (Cephalopoda) 
coe tCHLAGM Aap NeOnOpHOre yet Me rok lo Ses 
Lopez-Gonzalez, Pablo J., Mercedes Conradi, and J. Carlos Garcia-Gomez. Enterocola 
africanus, a new species (Copepoda: Ascidicolidae) associated with a compound as- 
cidian Synoicum species from North Africa (Strait of Gibralter) 000 
Manning, Raymond B. Two new species of Neocallichirus from the Caribbean Sea 
eentisiaced..ecapoda: < aliaiassidae)? 20) amt beeireiye: res tes nue pretest ie he 
Manning, Raymond B. Three genera removed from the synonym of Pinnotheres Bosc, 
1802 (Brachyura: Pinnotheridae) ... seen 
Manning, Raymond B. and David K. ea. Er geint dyailinidch. 5 a new ieerakceiniys 
and Tetrasquillidae, a new family of stomatopod crustaceans 
McCranie, James R., Larry David Wilson, and Kenneth L. Williams. A new species 
of Oedipina (Amphibia: Caudata: Plethodontidae) from northern Honduras _. 


762-774 


714-718 


24-33 


402-409 


436-441 


740-748 


442-449 


182-189 


719-727 


455-466 


251-295 


63-84 


315-324 


532-544 


346-352 


545-553 


582-586 


417-428 


366-368 


490-496 


602-605 


131-136 


106-114 


523-531 


85-91 


385-389 


vi 


Morino, Hiroshi and Reuven Ortal. The identity of Talitroides alluaudi (Chevreux) 
(Crustacea: Amphipoda: Talitridae) with notes on a new locality 20 eeeeeeeeeeeee 
Ng, Peter K. L. On a new genus and species of xantid crab (Crustacea: Decapoda: 
Brachyura) from Chesterfield Island, Coral Sea 2 escsseesseeseesesseeeesseeeseseeesseneeeeeenenenenennenennnee 
Norman, Mark D. Octopus ornatus Gould, 1852 (Cephalopoda: Octopodidae) in Aus- 
tralian waters: morphology, distribution, and life History oe ceeeecccceccecceececeneeeeeeeee 
Opresko, Dennis M. A new species of Sibopathes (Cnidaria: Anthozoa: Antipatharia: 
Antipathidae) from the’ Gulf of Mexico) 222 ee eee ne Secon ee 
Pereira S., Guido A. A description of a new species of Macrobrachium from Peri, and 
distributional records for Macrobrachium brasiliense (Heller) (Crustacea: Decapoda: 
Parle rm Omiaae) ie ecicicsscnceissssasachceetcsenctean ties cceteain nee Ai ae ene ae ee UR ced EKER 
Pérez, Linnette Garcia and W. Ronald Heyer. Description of the advertisement call 
and resolution of the systematic status of Leptodactylus gracilis delattini Miller, 1968 
(Amphibia: Leptodactylidae) (2) 2:25 ie 2 or EE ee ee ee Ee 
Petit, Richard E. and M. G. Harasewych. A new Mericella (Mollusca: Gastropoda: 
Cancellaritdae)'from northeastern”Atrica: 2222 a ee eee ee 
Pettibone, Marian H. Polynoid polychaetes associated with a whale skeleton in the 
bathyal Santa*Catalina*basin) “25% 22 oes Oe 2 ee ee eee 
Pleijel, Fredrik. Taxonomy of European species of Amphiduros and Gyptis (Polychaeta: 
FHOSIOMIGAC) sic cancdesceeseseccatn et onc etc ca rac a ee 
Pleijel, Fredrik. Gyptis crypta, a new hesionid species from the U.S.A. east coast, with 
a redescription of G. vittata Webster & Benedict, 1887 (Annelida: Polychaeta) ........... 
Pleijel, Fredrik and Kristian Fauchald. Scalispinigera oculata Hartman, 1967 (Scali- 
bregmatidae: Polychaeta): senior synonym of Lacydonia antarctica (Lacydoniidae) 
Hartmann-Schroéder & Rosenfeldt, 1988 
Price, Roger D. and Robert M. Timm. Two new species of Gliricola (Phthiraptera: 
Gyropidae) from the spiny tree rat, Mesomys hispidus, wy Per iiii.eccccecccecccenneeeeeeeeeeeeeeeeeenom 
Pyburn, William F. A new species of dimorphic tree frog, genus Hyla (Amphibia: Anura: 
Hylidae), from the Vaupés River of Colombia eee eeeeeeeeeeeeeeeeeeennnnennnnnnnnnnnnneen 
Rausch, V.R.and R. L. Rausch. Karyotypic characteristics of Sorex tundrensis Merriam 
(Mammalia: Soricidae), a Nearctic species of the S. QraNe@US-QTOUP nnn eee 
Reid, Janet W. and Teruo Ishida. New species and new records of the genus Elaphoidella 
(Crustacea: Copepoda: Harpacticoida) from the United States 2 eee 
Remsen, J. V., Jr. Zoogeography and geographic variation of Atlapetes rufinucha (Aves: 
Emberizinae), including a distinctive new subspecies, in southern Peru and Bolivia 
Rice, Mary E. Two new species of Phascolion (Sipuncula: Phascolionidae) from tropical 
and subtropical waters of the cemtral westerm Atlamtac oi. ceeeccceccesneeeecesnesesesneeeeeneeesssneeeee 
Robinson, Harold. A review of the genus Critoniopsis in Central and South America 
@Veronieae:, AStenraceae) ee re i ee ee 
Robinson, Harold. Three new genera of Vernonieae from South America, Dasyandan- 
tha, Dasyanthina, and Quechualia (Asteraceae) oii eeeeeeeeecccennnneeeeeeeeeeeeeeeeeeeeennnnnseeseceeeeseeeeeeeeennnnnes 
Rodrigues, Sergio de Almeida and Rosana Moreira da Rocha. Littoral compound 
ascidians (Tunicata) from Sao Sebastiao, estado de Sao Paulo, Brazil 
Roman-Contreras, Ramiro. Probopyrus pacificensis, a new parasite species (Isopoda: 
Bopyhridae) of Macrobrachium tenellum (Smith, 1871) (Decapoda: Palaemonidae) of 
the ‘Pacific ‘coast of Mexico! 220 a ee Diet eo ere ee, Le Peeve 2 ee 
Rozbaczylo, N. and J. J. Canete. A new species of scale-worm, Harmothoe commensalis 
(Poychaeta: Polynoidae), from mantle cavities of two Chilean clams _ 
Rudjakov, J. A. The first finding of the male of Thaumatocypris echinata Miiller, 1906 
(Crustacea: Ostracoda)... rare ee a ee ee ee ae ee 
Saltzman, Jennifer and Thomas E. Bowman. Boreomysis oparva, a new possum shrimp 
(Crustacea: Mysidacea) from an eastern tropical Pacific SeammOUNt oie eee 
Schotte, Marilyn and Raymond B. Manning. Stomatopod Crustacea from Tobago, West 
ANS, 
Smolen, Michael J., Richard M. Pitts, and John W. Bickham. A new subspecies of 
pocket gopher (Geomys) from Texas (Mammalia: Rodentia: Geomyidae) —.... 
Solis-Weiss, Vivianne. Grassleia hydrothermalis, a new genus and species of Amphar- 


332-338 


705-713 


645-660 


195-203 


339-345 


51-56 


221-224 


678-688 


158-181 


237-242 


673-677 


353-358 


46-50 


410-416 


137-146 


429-435 


591-601 


606-627 


775-785 


728-739 


689-697 


666-672 


305-314 


325-331 


566-581 


5-23 


etidae (Annelida: Polychaeta) from the hydrothermal vents off the Oregon coast (U.S.A.), 
So) NDE EEC EO a sR, Paper el ca ee 
Takeuchi, Ichiro. Caprella arimotoi, a new species (Crustacea: Amphipoda: Caprellidea) 
eer EUS EE Se Es |: 
Thuesen, Erik V. Vampyrocrossota childressi, a new genus and species of black medusa 
from the Bathypelagic zone off California (Cnidaria: Trachymedusae: Rhopalone- 
matidae) a RS ee ee 
Weems, Robert E. and Peter G. Kimmel. Upper Triassic reptile footprints and a coel- 
acanth fish scale from the Culpepper Basin, Virginia aes SAO AT 
Wynn, Addison H. and Alan E. Leviton. Two new species of blind snake, genus Typhlops 
(Reptilia: Squamata: Typhlopidae), from the Philippine Archipelago ____ 


661-665 


115-121 


190-194 


390-401 


34-45 


INDEX TO NEW TAXA 


VOLUME 106 


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


PORIFERA 
Demospongea 
CéeratopsiGH::CFUSTOSUTNE Loc OE. bia ES 8 RS eee ey eek ce ee 629 
CNIDARIA 
Hydrozoa 
VIVA VT OCT OSS OU csc at awstats ec at late cee A ae 190 
CORA OS SE cis ac ssc te ag a nln Sa a ne 191 
Anthozoa 
Sibopathes: GCrOSDUniG ic.) oo 8 oo he ee 195 
PLATYHELMINTHES 
BIGirium: New SUPREMUS Ho.est Foes ee en ate ee eee Oe 214 
Gibsonium, new SUDGENUS::4)... Se ee Be ed he Tn ee 215 
Giratdia CANGt 2.1. ic Mee ies ht ee ee 633 
Maillardiella, new Subse@mus:e 2. 235 a. Re AR pee i ed Be es Dit. 
Overstreetium, N€W SUDBENUS |: 5.8 ee ee ee eee 213 
‘Fimontella:Ostrowskig@e 3 88 Se Re ne 208 
MOLLUSCA 
Gastropoda 
AIA OS IVT SUG ASR Ma tea ON eB ae aa ea 182 
LY (| ean: SAE Hae a RE LLP xh ae On Nivea Soe tea ga Joe ies OWE MRR ee RUE 183 
Mericellla DOZZEtIE nc ae ee Re aS et ee San een ee 221 
SIPUNCULOIDEA 
Phascolion gerard! 2 2 ee ee ee : Br ca Mente A Jed eee oe eee 2 591 
POSGVAVVOPTAUT ATI os eee Se ae rn ee tg 594 
ANNELIDA 
Polychaeta 
Amphiduros fuscescens ni...) See ee a ee ns ee eh ee 176 
lO | G1) (/ a AM eee has oe Ae Rs ee eI 153 
GIVING en a na ee er 153 
GF GSSTCTG 0 ok ase soe a SS rel SA Ne aU ee 662 
hydrotherrrialis nc ee ae a a dda i ce 662 
GPUS: CIV DIG a ee re ee na ee 238 
TUCK TOD a kT Te an reac ce Se 165 
VIVCCILCF ROTO aac kre ee eo eo Ae ac 168 
Harmothoe: COMMeNsSQLS ce 667 
CVQUOSTAIDN oe a eo 683 
Vacydomia ocullataa mci ccc ca ee ea 674 
TLGUDICT US cc a ee eee eee Re ea Pe 151 
TUUUCTONUUS ccs oc cee es ee ifs) 


Miralvinella, new subgenus ASS Ae Ee ee een. Set Be SSeS ee ee 
BEN OE a a ae SE Sn OE Se _..._ 148 
SEE TSE ee ee ee EL eee eee eee suoer AS 
PONCE SUT Ba EIST STS sc eI 0 A a SS 
PSE LE ET ie ss ee See 
unidentata wll lotto Si OE Ne SAL ze IE re eB nS 226 
SRS USS TEI DET TI 7 es A ee aE ae nee eT _. 680 
EG cee NS See oe irae ls ne ee a _ 149 
ESSERE SET TS a TE et nn ene Be ee RRO TE Eee ate, WSO 
UEP PST TSE ee 2 | ES Ns en ee Sn eS 2iisi582 
Oligochaeta 
ULL LESTOSG IR US VEEN) is (1) a See Re eee SI 2 eae eel ST eee £ ee (SRT 
oe LLORES aS eee, ee Ses ee ee _.. 244 
ARTHROPODA 
Crustacea 
ELAR AR FEL ie means RP OTe Ne KE IO Sa ch spate, OD a S72 
BTS EIN ew 8 i Fig i ein raes: sine STD 
STWSIGI ET TSE SRE NR Geeta dri a er ee a Seka eh SS 
Bane Ge AE cA aera a cr nn EO an ae ea a ea See 
IIE DUREPOIEIID TY OUR 2 Ri i Uk en Re eae A ame oe tent nn ren AS at cee a ie NE 0 et = 162 
WEL LE ESES, COO eA RRS ETE Oe OS ONDe NCH hr ER, OE STL wae, 496 
EOE OT a RE ce ie ee ee er RS RE EN Sk. foe ne _ 574 
SUITE STE SST TO Ce a ee eee .. 574 
CE SPE DPR OSDS CUE PE ie I ak oe Sea ede 
aie MC REIET, SEAS PERIEIC PONIES Commis amen ace Be ee 4 OG 
Sa res sea TEMES OE AI THLIGATIAIIATUS) CIR CASTS oe _. 346 
aCerimEa REIS 0 DPIC AMID ATS) SHDICEFANICUS. i rake 
a eeemee PRES (COP ERESCEICAIMIDDANUIS) AGATNCMUT a TC. = 55 
thomai .. ene SUS ES ES ING ey lal RN Se Na RS i=. 330 
Fa TPP PRET YS E CI pmrine i er ira, Mes Saeed eens gee 115 
Chaceon alcocki at ee a nen RU 714 
TEE Ea RR al AE ii Sk oe CR PS 705 
EGS I Ss seme cm pet ht ee ee ee! | if 
ie AONE NUN POL CLOSES ocr i Dea a ee ee 22 B59 
Et PEC EGIE FPR EA UE eee I | 
SEER EL Se OE eA Re ae ee ee aes ee enema LS aM A 
EEE EU DI OYE ee ee eae ere bs? 
2 SET SD NE OD Bo, TE oe Se ee eT ee iL ST 
ana RRRIEE TEMES SGTP EPO) OEE EN acs coe ec oR a A a eet 568 
PIE Eas at Ng WAN SN OR rd. SN SO aT B.S 
IAEA REC EN CRERENO Get Setcfl ce c  h eid ee eeeee at DG 
Kalliapseudes (Mesokalliapseudes) SOMiQADQWIGE ocean neeeenneeenenneeeeeneenennnennennnnennnn 122 
DE cece he SS EER eR ON UY a 
PEE a es I EN een TN Ek sn . 499 
Sper iraReec the ISPPISTI PCIE CONCEIVES ke 339 
ULES TON Sr SEP CC) a SR SE See ee Pe Rh ee 375 
CRIED NSENGIS VEE G ee SN ee ae ee a ee aa Sw 107 
OE EL TREN IE el ENN ee ee Oe ee OTS ee Ae ALS Ly Ie 110 
og E01 es Bo Sea RE ee ee, es STG 8 463 
BERR SERITe CREASE TSU ITEC TINTS Soe er eS eee ee 698 
OS ERGSTS NTS PRETTIEST 2, ee a ee RTE TEES 690 
2 RESIS P07 2 ES NR Ss Si ee es ee een. OW acs ese. om 554 
_ DRESREP ST 2) 00 P20 SMS Ee la TS il a EI SE NCO WL ae 508 
EEE EFA es IR ae ee IN seek RC A NS Oe Ce Siz 


OOD ESS STITT MI a UN Se ESO ES ML Tet ll OR en PP eR Oe ey 


TET RAS OUTED AEE oi cesses cepa 88 


Th DOr CAD ATAUUS ANGI S VAUD CUMS a cea a 534 
RV UO PDMS LONI ss in a ct ON SCP cae ER nee 559 
Insecta 
Co CO ae ase ee a es 354 
WO ODI occ al ce ttl la hee ee cn 353 
CHAETOGNATHA 
Spade la JQ ONC ess scsc dec cnecnsscni a I te PO ences 359 
ECHINODERMATA 
[MUN BLOT UI nasi ate sence ct i en oe 366 
SthataSter -P71QCIVENOTUIN cscs tcc ee aa ee ee 66 
a7 (201 8 9 © nD er Sear RE cet Pa ca eee 66 
CHORDATA 
Pisces 
Hyporhamphus (eek ccc ote ie se 371 
Prochilodus.bAtsKil..... 22. ee ee ee eee 58 
Amphibia 
Bufo: G@inDOrOeCNn sts coo ee ee ed ee 443 
Blvia. kKarenanneae sn ee ian en ete 46 
Oedipina,cephyra oe ee ee 385 
Reptilia 
TyphlopsCQStQNOGUS oe oe cothsccscse nen de ee ee 35 
COW GHAS aoe oo a ae ee ee aed 41 
Aves 
Atlapetes:fufinucha térborghi cco ee ee ee 429 
CDOT DCIS sae Og a ih Rg ed ele es Bee eee 26 
Mammalia 
Geonmiys texensis bakeri. 2  eeee 19 
PEGI CCH CEI ODS acs canines cen tan cceghectaas eo ghrelin US: 
COLT OTA Pi eit yee ket oe Gea i oa Aft 753 
"TPROMASOMYS!GPCCO..5 ie ee 417 
PVUACTOWAS  eosoo css oo cect thal ee Ge Joel tN sr 762 
PLANTAE 
Critoniopsis .cotopaxensis.... 2.0. ee ee 610 
OVI ie oe el cst ede se Fe 612 
VOWESHL cick 2 te cs ar ally cin RN NE er, Oc oer ae a, 615 
[0,01 (061 6\) | eae meer ees CNC a a, Ne ee 617 
QUILL OMONSTS nooo a 8 ee ee dr 620 
StCINDQCHEL oo Secs ee 622 
POTD en eel eS ea 625 
CFIGOMIOPSIS, 38.10 S ee sah oe ee 609-626 
IDOLS OL 01} |/ ee ea MN 778 
CUAL CASASTAMA 10. C osc oa ossce ccccecdecce cece 778 
DO | | i nn ee ee 778 
PAULUS ENS eo a Sine 1 780 


To) 6 21122 | Ce ee eo ee eee one een We eR CO 780 


Quechualia 
cardenasii N.c. 
fulta n.c. 
smithii 
trixiodes n.c. 


= 


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ON THE ID! 
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BRAN 
Abstract: 


PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 14 


NTITY OF ECHIMYS DIDELPHOIDES 
SMAREST, 1817 
¥ anal ODENTIA: ECHIMYIDAE) 

Wi 


ouise H. Emmons 


€arly names for species of the genus Echimys (Makalata), 


E. didelphoides and Loncheres obscura, were erroneously assigned to the genus 
Mesomys by Tate (1935). One of these, didelphoides, is the oldest name for the 
red-nosed tree rats currently known as Makalata armata. 


While revising the genera of arboreal 
echimyids I have encountered chronic er- 
rors in the literature that I wish to correct 
without awaiting completion of systematic 
studies. 


Taxonomic History 


Desmarest (1817:55) described Echimys 
didelphoides, attributing the name to E. 
Geoffroy St.-Hilaire. The description is brief, 
and includes the comment that the tail is 
furred for a portion of its base and naked 
distally. Regrettably, Desmarest named the 
species for the resemblance of the tail to 
those of didelphid opossums, whose tails 
are also naked but densely furred at the base. 
The description of E. didelphoides imme- 
diately follows that of Echimys hispidus, on 
the same page. The latter also is described 
as possessing a naked, scaly tail. Lichten- 
stein (1830) described and illustrated a 
specimen he identified as Mus hispidus Des- 
marest, 1817, said by a dealer to have come 
from Cayenne. Cuvier (1832: Plate 18, fig. 
2) illustrated the teeth of E. didelphoides. 

Geoffroy St.-Hilaire (1838) noted that 
Lichtenstein’s specimen was misidentified, 
and, without seeing the specimen, renamed 
it Nelomys armatus. In his complete revi- 
sion of the echimyid rodents, Geoffroy St.- 
Hilaire (1840) explained that armatus re- 
sembled didelphoides except in details of 
color, length of the haired part of the tail, 
tail length, and width of the spines (p. 11), 


but he later noted that the type of didel- 
phoides was a young individual that had 
been preserved in alcohol and it would be 
larger as an adult and its original color could 
not be known with certainty (p. 43). He said 
that his father (E. Geoffroy St.-Hilaire) had 
acquired it in Lisbon and, although it lacked 
a locality, it probably came from Brazil. It 
is illustrated with a fine color plate (I. Geof- 
froy St.-Hilaire 1940: plate 24). 

Wagner (1843) commented that Echimys 
didelphoides was closely related to Lonche- 
res armata: Wagner, 1843, and perhaps 
simply a young one. Waterhouse (1848) said 
the differences between the two did not seem 
to warrant their separation. 

Tate (1935) without seeing the specimens, 
placed didelphoides in the genus Mesomys, 
where it has remained since (e.g., Cabrera 
1960, Honacki et al. 1982, Corbet & Hill 
1991). At the same time, Tate (1935) also 
placed Loncheres obscura Wagner, 1840, in 
the genus Mesomys. 

Husson (1978) erected a new genus, Ma- 
kalata, with I. Geoffroy St.-Hillaire’s ar- 
mata (Lichtenstein’s Mus hispidus) as its 


type. 


Identity of the Holotype of 
Echimys didelphoides 


Tate (1935) based his decision to include 
didelphoides in Mesomys on the color plate 
in Geoffroy St.-Hilaire (1840). Because the 
teeth illustrated by Cuvier (1832) are not 


2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


those of a Mesomys, he assumed that spec- 
imens had been mixed up and the wrong 
one illustrated. 

I examined the holotype, Museum Na- 
tional d’Histoire Naturelle, Paris (MNHN) 
No. 404, 1805 (lot de montage). It is a 
mounted specimen on a wooden base, on 
the bottom of which is written, ““Type de I. 
Geoffroy pére et fils Nelomys didelphoides 
Is Geoff. (T) Echimys didelphoide Geoff. St. 
N (T) Amerique du Sud.”’ The specimen was 
on display in the Grande Gallerie for over 
a century and is severely darkened. The skull 
has been lost, but fortunately, before its loss, 
the teeth were illustrated by Cuvier (1832: 
Plate 18, fig. 2). The teeth are those of a 
young animal, with the third molar not yet 
erupted through the gumline. 

The holotype of Echimys didelphoides is 
a young Makalata armata as currently de- 
fined. All evidence from the period litera- 
ture, including the color plate (Geoffroy St.- 
Hilaire 1840), the teeth (Cuvier 1832), and 
all original descriptions based on the type 
conform exactly to MNHN 404, and there 
is no evidence that a specimen mixup oc- 
curred. 

The holotype of E. hispidus, MNHN 407, 
is a mounted specimen on a wooden base 
with “Echimys hispidus Geoffro. St. H. (T) 
Type de l’espéce”’ written below. The skull 
attributed to this specimen has an attached 
label from the British Museum (BMNH), 
with, in Oldfield Thomas’ handwriting, 
‘““Mesomys hispidus type of “E. hispidus.” 
This specimen is a Mesomys and it is also 
shown accurately in a quaint but unambig- 
uous color plate in Geoffroy St.-Hilaire 
(1840). 

Echimys didelphoides Desmarest 1817, 
therefore, antedates Echimys armatus Geof- 
froy St.-Hilaire, 1838 (based on Lichten- 
stein 1830). Tate (1935) simply erred in 
treating didelphoides as a Mesomys. 

Echimys didelphoides clearly belongs to a 
group that is almost certainly polytypic 
(Emmons & Feer 1990 and Emmons, un- 
published results) and contains a number of 
other named and perhaps unnamed forms 


that are not currently recognized (Cabrera 
1960, Honaki et al. 1982). It is therefore 
important to determine whether E. didel- 
phoides can be attributed to any geographic 
subset or form of the red-nosed tree rats. 
The skin of the holotype does not corre- 
spond in diagnostic traits to either E. oc- 
casius Thomas, 1921 or E. rhipidurus Tho- 
mas, 1928 as redescribed by Emmons and 
Feer (1990), and the teeth are also distinct 
from the latter. The specimen does agree in 
characters with a group of forms like ar- 
mata. 

The skin of the holotype of didelphoides 
was described and illustrated as having a 
pale venter sharply differentiated from the 
sides, and does so now. The type of armata 
was also described as having a completely 
cream or buff venter (“Isabellfarbe,’’ Lich- 
tenstein 1830). This character is rare in red- 
nosed rats, it is found in the holotype of 
Echimys guianae Thomas, 1888, from Guy- 
ana, and also in a few (but not all) specimens 
from south of the Amazon in the Brazilian 
states of Maranhao and Para, from the rio 
Xingu to the Atlantic coast east of Belém. 
Most other populations have exclusively 
gray-brown venters. However, the hair pig- 
ments of echimyids bleach easily and Ven- 
ezuelan specimens stored in alcohol in the 
National Museum of Natural History, 
Washington (USNM), have in 20 years lost 
much color and their venters are now dirty 
yellowish, while skins from the same col- 
lections are dark gray-brown. Because the 
holotype of didelphoides was originally in 
alcohol, its pale ventral color should not be 
given too much emphasis. 

All juveniles of spiny arboreal echimyids 
lack spines, which get gradually heavier with 
age. Compared to armatus, the narrower, 
smaller spines in didelphoides, a chief char- 
acter used by Geoffroy St.-Hilaire (1840) to 
separate armatus from it, therefore has little 
value. 

Among other characters, Husson (1978) 
used direction of upper toothfold opening 
(lingual or labial) to distinguish Makalata 
from Echimys. Because he apparently only 


VOLUME 106, NUMBER 1 


examined red-nosed tree rats from Surina- 
me, he failed to realize that this character 
is extremely variable. Individuals within 
populations can have different states of 
toothfold pattern, although the within-pop- 
ulation variation is much less than that 
found between populations. The cheekteeth 
of E. didelphoides illustrated by Cuvier 
(1832) have all folds opening labially on the 
first two cheekteeth, with the posterior folds 
opening lingually in the third and fourth 
teeth, which matches the pattern often seen 
in specimens from Venezuela and also that 
of an animal from near Belém (USNM 
460069). Others from Para, the holotype of 
E. guianae, and specimens from Suriname 
(Husson 1978) usually have the posterior 
fold opening labially in all four cheekteeth. 
Both the toothfold patterns and the shape 
and proportions of the teeth of didelphoides 
seem to exclude it from populations ex- 
emplified by specimens from Pert and from 
the Amazon Basin west of the rios Negro 
and Tapajoz. A pale venter would likewise 
exclude it from these populations. 

On current evidence, I assign E. didel- 
Dhoides to the greater Guiana region, as de- 
fined by the regional concordance of species 
of primates and other mammals (Emmons 
& Feer 1990), including the area north of 
the Amazon and east of the rio Negro and 
including the Guianas, and south of the 
Amazon from the rio Xingu eastward. 

The names based on specimens from this 
region that are junior synonyms of Echimys 
didelphoides are: 


Nelomys armatus I. Geoffroy Saint-Hi- 
laire, 1838 

Loncheres guianae Thomas, 1888 

Echimys longirostris Anthony, 1921 

Echimys castaneus Allen & Chapman, 
1893 


Identity of Loncheres obscura 


Wagner (1840) described and figured the 
skull, limb bones, and teeth of Loncheres 
obscura collected by Spix in Brazil. The de- 
scription and figures are clearly of an Echi- 


mys (s.1.). Tate (1935), in an apparent lap- 
sus, placed obscura first in the genus 
Mesomys (p. 413), and then in the genus 
Echimys (p. 432). Subsequent authors (Ca- 
brera 1960, with reservations; Honaki et al. 
1982; Corbet & Hill 1991) followed Tate’s 
first allocation, and retained obscura in 
Mesomys. 

I have not seen the holotype of L. obscura. 
The illustrations of it are crude, but they 
preclude identity with Mesomys, Nelomys 
(Atlantic tree rats), and E. rhipidurus. The 
description (“dunklebraun’”’) most closely 
matches dark, almost blackish animals from 
west of the rio Madeira in Brazil, a region 
visited by Spix. Two years later, Wagner 
(1842) described another species, macrurus, 
from Borba (on the east side of the Ma- 
deira), as reddish yellow (“‘fulvescens’’). 
Pending a better identification of obscurus, 
the large red-nosed rats of the central Am- 
azon basin should retain the name E. ma- 
crurus Wagner. 


Acknowledgments 


Travel to London and Paris to examine 
types was supported by the American Mu- 
seum of Natural History. I thank Michel 
Trainier for his considerable help in locating 
dispersed specimens in Paris, and Paula 
Jenkins and Guy Musser for their hospital- 
ity in The British Museum of Natural His- 
tory and the American Museum of Natural 
History. A. L. Gardner helped resolve some 
technical questions of nomenclature. He and 
J. L. Patton made helpful corrections on the 
manuscript. 


Literature Cited 


Cabrera, A. 1960. Catalogo de los Mamiferos de 
America del Sur. Museo Argentino de Ciencias 
Naturales ““Bernardino Rivadavia.”— Ciencias 
Zoologicas 4:538—-543. 

Corbet, G. B., & J. E. Hill. 1991. A world list of 
mammalian species. British Museum, London. 

Cuvier, F. 1832. Description des charactéres propres 
aux genres Graphiure et Cercomys de l’ordre des 
rongeurs.— Nouvelles Annales du Museum 
d’Histoire Naturelle 1:449-452, pl. 18 fig. 2. 


+ PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Desmarest, A.G. 1817. Echimys, Echimys. Tome X. 
Pp. 54-59 in Nouveau Dictionaire d’Histoire 
Naturelle, Deterville, Paris, 10:1—591. 

Emmons, L. E., & F. Feer. 1990. Neotropical rain- 
forest mammals. University of Chicago Press, 
Chicago, 281 pp. 

Geoffroy Saint-Hilaire, I. 1838. Notice sur les ron- 

geurs épineux désignés par les auteurs sous les 

noms d’Echimys, Loncheres, Heteromys et Ne- 

lomys.—Revue Zoologique 1:99-101. 

1840. Notice sur les rongeurs epineux dé- 
signés par les auteurs sous les noms d’Echimys, 
Loncheres, Heteromys et Nelomys.—Magazin 
de Zoologie, Série 2, 2:1-57, pls. 20-29. 
Honaki, J. H., K. E. Kinman, & J. W. Koppl. 1982. 

Mammal species of the world. Allen Press and 
Association of Systematics Collections, Law- 
rence, Kansas, 694 pp. 

Husson, A. M. 1978. The mammals of Suriname. E. 
J. Brill, Leiden, 569 pp., 151 pls. 

Lichtenstein, M.H.C. 1830. Darstellungen newe oder 
wenig bekannte Saugethiere. 2 Vols. C. G. Lu- 
deritz, Berlin, plate XX XV with text. 

Tate, G. H. H. 1935. The taxonomy of the genera of 
neotropical hystricoid rodents. — Bulletin of the 
American Museum of Natural History 68:295— 
447. 


Thomas, O. 1888. On a new species of Loncheres 
from British Guiana.— Annals and Magazine of 
Natural History, Ser. 6, 2(10):326. 

Wagner, J. A. 1840. II. Stachelmaiise. Abhandlungen 

(Bayerische) Akademie Wissenschaften, Mu- 

nich. Pp. 191-210, plate II. 

1842. Diagnosen neuer Arten brasilischer 
Saugethiere. — Archiv fiir Naturgeschichte 8:356— 
362. 

. 1843. Die Saugethiere in Abbilddungen nach 

der Natur mit beschreibungen von Dr. Johann 

Christian Daniel von Schreber. Leipzig. Sup- 

plementband Erlangen, Expedition das Schre- 

ber’schen Sdugthier- und des Esper’sshen 

Schmetterlingswerkes, und in Commission der 

Voss’schen Buchhandlung in Leipzig, 3:xiv + 

614 pp., pls. 85-165. 

Waterhouse, G. R. 1848. A natural history of the 
Mammalia. Vol. II. Hippolyte Bailliere, London 
2:1—500, 21 pls. 


Division of Mammals, MRC 108, Smith- 
sonian Institution, Washington, D.C. 20560, 
U.S.A. 


PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 5-23 


A NEW SUBSPECIES OF POCKET GOPHER (GEOMYS) FROM 


TEXAS (MAMMALIA: RODENTIA: GEOMYIDAE) 
Michael J. Smolen, Richard M. Pitts, and John W. Bickham 


Abstract. —Two isolated populations of pocket gophers were discovered far 
outside the previously known distribution of geomyids in southern Texas. These 
represent a new taxon, which is related to Geomys texensis in chromosome 
morphology and biochemical properties. The two share identical diploid num- 
ber (2N = 70), fundamental number (FN = 68), and the presence ofa distinctive 
large acrocentric X-chromosome. There are no fixed differences in any of the 
18 loci analyzed with starch gel electrophoresis, when comparing the new taxon 
with G. texensis. However, both of these taxa share alleles at two loci that are 
present as fixed differences when compared to G. bursarius major. Analysis of 
cranial morphology indicates that the new taxon resembles G. texensis, but 
differs significantly in cranial dimensions of size and shape, related primarily 
to measurements of basal and palatal lengths and mastoid breadth. Because of 
the morphological distinctiveness and the extreme spatial separation (120 km) 
from populations of G. texensis confined to the central basin of the Edwards 
Plateau, we conclude that this new taxon is an isolated, relictual population of 
G. texensis, and is a distinctive subspecies. 


During a recent survey of the pocket go- 
phers in southern Texas, populations were 
discovered far outside the previously known 
distributional ranges of Geomys. Geomys 
attwateri and G. personatus are both broad- 
ly distributed species (Fig. 1) that occur over 
much of the eastern and central portions of 
an area herein referred to as South Texas 
and bounded by the Edwards Plateau, San 
Antonio River, Gulf of Mexico, and Rio 
Grande River (Davis 1940; Kennerly 1954, 
1959; Williams & Genoways 1981). Their 
distributions are restricted to sandy and 
sandy-loam soils (Davis 1940, Honeycutt & 
Schmidly 1979) that are dispersed through- 
out this region. The northern and western 
parts of South Texas, however, consist al- 
most entirely of hard indurate soils, with 
high clay content, and caliche. Geomys is 
occasionally found in isolated pockets of 
sandy or sandy-loam soils in this area. 

Survey of these isolated patches of suit- 
able habitats in the northwestern part of the 


region detected previously unknown pop- 
ulations in Medina, Zavala and Uvalde 
counties. Karyotypic, morphometric and 
electrophoretic analyses identify these as re- 
lictual populations that are related to Geo- 
mys texensis, and they are described herein 
as representing a new subspecies. 


Materials and Methods 


Animals were live-trapped (Baker & Wil- 
liams 1972) and returned to the laboratory. 
Metaphase spreads were prepared from bone 
marrow cells, using 0.075 M potassium 
chloride hypotonic and Carnoy’s fixative 
(Baker et al. 1982). Fresh slides were pre- 
pared by dropping (1.5 m) the cells onto 
slides flooded with distilled water. Standard 
karyotypes were analyzed following staining 
with 2% giemsa stain in 0.01 M phosphate 
buffer. Diploid number (2N) was deter- 
mined by counting at least 10 spreads, 
whereas fundamental number and chro- 


101 100 


a 


aa ZAP OA a 
ie LESS D DDI AZZ 


(| 


6 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
do oy 

i 
| ) nl 
|) 

el 

lth 
wl il 
| 


| ll 
(ll 
J 
| 
( | | 
NT 


: y 
li 


dl28 


28 A G. personatus 
0 100 
ee 2? 
km 
27 
102 101 100 99 98 


Fig. 1. Map of the distributions of pocket gophers in southern Texas. Open circles represent localities from 
which specimens were pooled to form samples. 


VOLUME 106, NUMBER 1 


mosome morphology were described from 
photographic prints of selected spreads. 

Samples representing the unknown taxon 
(n = 21), G. texensis (n = 13), and G. bur- 
sarius (n = 10) were used to assay biochem- 
ical variation. Heart and kidney tissues were 
minced in a grinding solution (Tris/EDTA/ 
NADP) and homogenized using a mechan- 
ical homogenizer (Tissue Tearor, Biospec 
Products). Samples were loaded into 12% 
starch gels (Starch Art). The techniques used 
for visualizing the allozymes were those de- 
scribed by Harris & Hopkinson (1976) and 
Honeycutt & Williams (1982). Proteins were 
examined on the following buffer systems: 
Poulik, continuous Tris-citrate II (pH 7.0 
and 8.0), Tris-malate EDTA (pH 7.4), and 
Ridgway (pH 6.7). Eighteen presumptive 
loci and their respective Enzyme Commis- 
sion numbers (Murphy et al. 1990) are as 
follows: glycerol-3-phosphate dehydroge- 
nase (G3PDH; E.C. 1.1.1.8), malate dehy- 
drogenase (MDH-1,2; E.C. 1.1.1.37), isoci- 
trate dehydrogenase (IDH-1,2; E.C. 
1.1.1.42), phosphogluconate dehydroge- 
nase (6-PGDH; E.C. 1.1.1.44), superoxide 
dismutase (SOD-1,2; E.C. 1.15.1.1), purine- 
nucleoside phosphorylase (PNP; E.C. 
2.4.2.1), aspartate aminotransferase (AAT- 
eee 2-6-1:1), creatine kinase (CK; E.C. 
2.7.3.2), phosphoglucomutase (PGM-1; E.C. 
5.4.2.2), esterase (EST-1,2; E.C. 3.1.1.-), 
peptidase (PEP-1 leucyl glycyl glycine, 
PEP-2 leucyl alanine; E.C. 3.4.-.-), aconitase 
hydratase (ACOH; E.C. 4.2.1.3), glucose- 
phosphate isomerase (GPI; E.C. 5.3.1.9). 
The allozyme data were scored in a side by 
side comparison of mobility where identical 
mobilities were recorded as individual al- 
leles. Data were analyzed using BIOSYS-1 
(Swofford & Selander 1981), which pro- 
duced Rogers’ genetic similarity and genetic 
distance values for the three taxa (Rogers 
1972). 

The new taxon was compared to G. attwa- 
teri, G. personatus, and G. texensis using 
univariate and multivariate statistical anal- 


yses of morphological data. Geomys attwa- 
teri was represented by two populations from 
the western extent of its distribution and 
thus in relatively close proximity to the new 
taxon. Geomys personatus streckeri was se- 
lected as a representative of the personatus 
complex because its distribution also ap- 
proaches that of the new gopher. Two pop- 
ulations of G. texensis represent the previ- 
ously recognized subspecies, G. bursarius 
texensis and G. bursarius llanensis. They 
were recently elevated to specific ranking 
and synonymized as the monotypic G. tex- 
ensis (Block & Zimmerman 1991) based on 
patterns of biochemical variation. Although 
neither G. texensis nor G. bursarius are 
closely distributed to the new taxon, they 
are included because they are karyotypically 
identical to it. 

Geomyids show extensive variation in 
secondary sexual characteristics (Baker & 
Genoways 1975; Honeycutt & Schmidly 
1979; Williams & Genoways 1977, 1978, 
1980, 1981), which necessitated separate 
analysis of males and females. Three exter- 
nal measurements were recorded from spec- 
imen labels, and 11 cranial characters were 
measured using dial calipers. Skull mea- 
surements were taken as described by Wil- 
liams & Genoways (1977). Measurements 
we evaluated are as follows: total length (TL), 
length of tail (T), length of hind foot (HF), 
condylobasal length (CBL), basal length 
(BL), palatal length (PL), prefrontal depth 
(PFD), length of nasals (LN), diastema 
(DIAS), zygomatic breadth (ZB), mastoid 
breadth (MB), squamosal breadth (SB), ros- 
tral breadth (RB), interorbital constriction 
(IOC), breadth across third molars (M3), 
length of maxillary toothrow (M1). Only 
adult animals were used and they were iden- 
tified by the degree of ossification of the 
sutures of the skull, particularly the fusion 
of the basisphenoid and basioccipital bones 
(Williams & Genoways 1981). The adults 
were classified into two categories based on 
the degree of fusion and ossification. The 


8 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


skulls of older adults continue to grow, caus- 
ing shape of the skull to become more an- 
gular and massive (Russell 1968); this is 
particularly noticeable in rostral, zygomatic 
arches as well as other cranial measure- 
ments. The oldest adult age class consisted 
of animals with a total obliteration of the 
suture line separating the basisphenoid and 
basioccipital bones. The number of animals 
in this oldest age class varied among pop- 
ulations, with most being males. The largest 
sample sizes were in the younger of the adult 
age classes, and these were used in the mor- 
phological analysis. 

Univariate analysis of cranial characters 
(mean, range, standard error, coefficient of 
variation) was carried out using the UNI- 
VARIATE procedure of SAS (SAS Institute 
1988a, 1988b) for each sex. The relation- 
ships among the taxa were assessed using 
multiple analysis of variance (GLM pro- 
cedure), and Tukey’s studentized range test 
(TUKEY’s option of GLM) was used to 
identify maximally nonsignificant subsets. 
Principal component analysis (PRIN pro- 
cedure) using a correlation matrix of char- 
acters was used to identify the source of 
variation among the characters. Mean ei- 
genvectors were computed for each taxon 
and the first two components plotted. Ca- 
nonical discriminant analysis (CANDISC 
procedure) was used to compute canonical 
variates for multivariate analysis of varia- 
tion among taxa. The centroid for each tax- 
on was plotted on the first two canonical 
variates and the 95% confidence ellipse was 
computed using the method described by 
Owen & Chmielewski (1985). Each individ- 
ual was plotted by its first two canonical 
variates in order to visualize the dispersion 
of individuals in relationship to the cen- 
troids. The contribution of each cranial 
variable used in the canonical discriminant 
analysis was determined as a percentage, us- 
ing the procedure described by Schmidly & 
Hendricks (1976). 

Phenetic relationships were assessed us- 
ing correlation and distance matrices 


generated from the character data using NT- 
SYS (Rohlf 1988, version 1.50), with clus- 
tering using UPGMA (unweighted pair 
group method using arithmetic averages). A 
minimum spanning tree also was computed 
and taxa plotted by the first three canonical 
vectors were connected. 


Results 


The diploid number of 16 specimens rep- 
resenting the new populations in Medina, 
Zavala, and Uvalde counties is 2N = 70, 
and the fundamental number is 68. The 
X-chromosome is a large acrocentric and 
the Y-chromosome is medium-sized and 
acrocentric. The morphology of the chro- 
mosomes is indistinguishable from those 
previously reported for both G. bursarius 
and G. texensis (Baker et al. 1973, Hart 
1978). 

Four of the presumptive loci were mono- 
morphic within and among all three taxa 
(Table 1). Two fixed differences were ob- 
served (PEP-1 and SOD-1) between G. bur- 
sarius and both G. texensis and the new 
taxon. No fixed differences were observed 
between G. texensis and the new popula- 
tion. The mean heterozygosities for the new 
taxon, G. texensis, and G. bursarius are low, 
0.037, 0.041, and 0.017, respectively. The 
percentage of the loci that are polymorphic 
varied from a low in G. texensis of 27.8%, 
to 38.9% in the new taxon, and 44.4% in G. 
bursarius. Rogers’ genetic similarities were 
high when comparing G. texensis to the new 
taxon (0.915), whereas the similarities of 
these two taxa to G. bursarius were much 
lower, 0.690 and 0.648, respectively. The 
genetic distance was low when comparing 
the newly discovered populations and G. 
texensis (0.085), whereas these two taxa were 
both more distantly related to G. bursarius, 
0.352 and 0.310, respectively. 

Coefficients of variation produced in the 
univariate analysis of the nongeographic 
variation showed exceedingly high varia- 
tion in the external characters (TL, T, HF) 


VOLUME 106, NUMBER 1 


for both sexes, and were excluded from fur- 
ther morphometric analyses. The new taxon 
had the lowest measurements of the taxa 
used in this study in 22 of the 28 skull char- 
acters measured in the two sexes (Table 2). 
In the six incidences (female—BL, DIAS, 
IOC; male—CBL, DIAS, IOC) where this 
population did not have the smallest mean 
value, one of the populations of G. texensis 
was the smallest. Analysis of geographic 
variation in individual characters using a 
single classification MANOVA test showed 
significant differences in CBL, BL, PL, 
DIAS, ZB, MB, SB, RB, IOC, M3 and M1 
among the populations of females; PFD and 
LN showed no significant differences in this 
analysis. Males had fewer characters dis- 
playing significant differences among the 
taxa, and these were limited to LN, MB, SB, 
RB, IOC, M3 and M1. Further analysis of 
these data using Tukey’s standardized range 
test identified much of these differences to 
be attributed to relationships between only 
two or three of the taxa, especially regarding 
G. texensis, relating it to G. attwateri and 
G. personatus. Six characters displayed sig- 
nificant differences that involved all taxa in 
both males and females (Table 3): MB, RB, 
SB, M3, M1 and IOC. The new taxon was 
not significantly different from the other G. 
texensis populations in MB, SB, M1 and 
IOC in either females or males. Rostral 
breadth (RB) deviates from this clustering 
pattern in both sexes in that the new taxon’s 
measurements are much smaller than those 
of G. texensis. 

The first three principal components de- 
scribe 76.7% and 85.3% of the variation 
observed in females and males, respective- 
ly. The eigenvectors of component I were 
all positive and range from 0.110 to 0.355. 
Skull length measurements (CBL, BL, PL) 
and mastoid breadth (MB) account for most 
of the variation observed. Zygomatic 
breadth (ZB) accounts for additional vari- 
ation in males. Components II and III also 
are influenced by the variation in condy- 
lobasal length (CBL) and basal length (BL) 


Table 1.—Alleles present in three taxa. Lowercase 
letters denote alleles appearing in frequencies less than 
5%, while uppercase letters represent occurrences greater 
than 5%. 


G. t. bakeri G. texensis G. bursarius 


Due 
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in both sexes; however, squamosal breadth, 
mastoid breadth, and prefrontal depth are 
important characters in females. When 
plotted by the mean values for the first two 
principal components, the relationships of 
the taxa are identical to those seen for a 
similar analysis described below using the 
canonical variates, and thus are not pre- 
sented. 

There were significant differences (Ho- 
telling-Lawley’s Trace: females P < F < 
0.0001; males P < F < Q.0001) in mor- 
phology among the taxa in an analysis of 
geographic variation as detected by the 
MANOVA test. The first canonical variate 
accounts for 67% and 57% of the phenetic 
variation in females and males, respective- 
ly. The second and third variates account 
for 17% and 9%, respectively, in females; 
and 28% and 8% in males. Three clusters 
are evident in the two dimensional plots of 
the first two canonical variates of both sexes 
(Fig. 2). Females of both samples of G. att- 
wateri cluster with the G. personatus. The 
two populations of G. texensis also cluster 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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VOLUME 106, NUMBER 1 


together, with the centroid of each popu- 
lation well within the 95% confidence ellipse 
of the other. The centroid of the new taxon 
is well outside of the 95% confidence ellipses 
of all other taxa and its ellipse includes only 
the two centroids of G. texensis at the ex- 
tremes of the confidence interval (Fig. 2). It 
should be noted here that the new taxon had 
the smallest sample size (Fig. 2), and that 
this is directly observed in the larger ellipse 
resulting from the high F-value component 
of the Owen & Chmielewski (1985) for- 
mulation of the 95% ellipse. 

Males do not show as clear a demarcation 
among the clusters as do the females al- 
though an identical pattern is evident. The 
populations of Geomys attwateri cluster to- 
gether and overlap G. personatus. The two 
populations of G. texensis form overlapping 
clusters with the centroid of the new taxon. 
The analysis of males is affected by smaller 
sample sizes and the increased variation as- 
sociated with the continued growth of their 
skulls throughout adult life. 

Those skull characters representing length 
(CBL, BL) and width (ZB, MB) account for 
most of the variation in canonical variate I 
of males (Table 4). Basal length (BL), palatal 
length (PL), and mastoid breadth (MB) pro- 
vide most of the variation seen in the second 
variate. The third variate is also highly 
weighted to length and width variables. A 
similar pattern is seen in females, but this 
length and width variation is seen to be lim- 
ited to condylobasal length (CBL) and mas- 
toid breadth (MB) in the first variate. Over- 
all skull length accounts for the variation 
seen in the second variate, whereas palatal 
length (PL) and squamosal breadth (SB) ac- 
count for most of the variation seen in the 
third variate. 

The phenograms constructed using the 
correlation matrices are identical between 
the sexes, and the cophenetic correlation 
values are 87% for females and 76% for 
males (Fig. 3). The phenograms based on 
distance matrices differ between both sexes 
and the phenograms derived from the cor- 


13 


relation matrices. The cophenetic correla- 
tion values are high for females (0.84) and 
males (0.87). The new taxon is quite distinct 
in both sexes, with the greatest distinction 
appearing in males. Although the clustering 
relationships vary among the analyses and 
sexes, the new taxon is always distinct when 
comparing the branch lengths to those of 
the other species. A similar relationship is 
seen in the branching pattern resulting from 
the minimum-spanning analysis (Fig. 3). 
Females of the new taxon join G. texensis, 
but G. attwateri joins G. personatus. Males 
also show a close relationship among the 
two populations of G. texensis and the new 
taxon. 


Discussion 


The newly discovered populations of 
pocket gophers at first were suspected to be 
only range extensions of taxa known to oc- 
cur in the region. However, analysis of chro- 
mosomal morphology quickly identified 
them as being quite different from any taxon 
occurring in southern Texas, and more 
closely related to gophers in the G. bursarius 
complex to the north. They share a large 
and distinctive acrocentric X-chromosome 
and 68 acrocentric autosomes with G. tex- 
ensis and race D of G. bursarius major (Ba- 
ker et al. 1973, Hart 1978). They contrast 
with Geomys personatus and G. attwateri, 
both of which have a large, subtelocentric 
X-chromosome. Furthermore, G. persona- 
tus and G. attwateri are reported to share a 
diploid number of 70 in southern Texas and 
to possess a small metacentric autosome not 
seen in the new taxon. 

In order to ascertain the relationship of 
the new taxon to G. texensis and G. bur- 
sarius, and to determine its taxonomic af- 
filiation, we used starch gel electrophoresis 
to assay biochemical variation in 18 loci 
coding for structural proteins. Two fixed dif- 
ferences were observed between G. bursari- 
us and both G. texensis and the new taxon. 
No fixed differences, however, were ob- 


14 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 3.—Six characters that separate taxa when analyzed in single classification MANOVA. Taxa are grouped 
in nonsignificant subsets (Tukey’s studentized range test) represented by the horizontal lines. Age class 4 males 
and females: characters by locality. 


n mean min max 
Males 
MB 
G. t. bakeri 5 21.96 21.4 22.3 
G. texensis 4 22.65 20.8 23.9 
G. texensis - 23225 21.9 23.9 
G. personatus 13 23.98 DOS 26.7 
G. attwateri 11 24.06 22.6 2557 
G. attwateri 5 24.11 23:5 DAN 
SB 
G. t. bakeri 5) 16.94 16.6 17.4 
G. texensis 7 17.52 16.9 18.0 
G. texensis 4 17.63 16.8 18.5 
G. attwateri 11 18.22 17.4 19.5 
G. attwateri 5 18.63 18.1 19.5 
G. personatus 13 18.69 | 725 19.8 
RB 
G. t. bakeri 5 9.19 8.9 9.5 
G. personatus 13 9.37 8.9 10.7 
G. attwateri il 9.43 9.1 9.9 
G. attwateri 5) 9.74 8.9 10.4 
G. texensis 4 9.95 9.8 10.2 | 
G. texensis di 10.17 9.3 10.9 
M3 
Ee boke 5 6.90 6.8 7] 
G. texensis 4 7.48 TD 7.8 
G. texensis a 7.61 133 8.0 
G. attwateri ial qe! Teal 8.1 
G. personatus 13 7.80 7.3 8.8 
G. attwateri 5 AO? 126 8.5 
M1 
G. t. bakeri 5 8.49 8.2 8.8 
G. texensis 4 8.63 8.1 9.0 
G. texensis i 8.91 8.6 9.6 
G. attwateri itll 9.36 9.1 9.8 
G. attwateri 5 9.36 9.1 9.6 
G. personatus 13 9.43 8.4 10.6 
IOC 
G. texensis 4 5.87 55 6.3 
G. texensis T! 5.88 5.6 6.2 
G. t. bakeri 5 5.96 Dei! 6.4 
G. personatus 13 6.18 5.9 6.6 
G. attwateri let 6.40 6.0 (pa 
G. attwateri 5 6.53 6.1 6.9 


served between G. texensis and the new tax- geomyids from central Texas. Genetic sim- 
on. Block & Zimmerman (1991) identified _ilarities determined by them for G. bursari- 
fixed differences between G. bursarius and us and populations of G. texensis ranged 
G. texensis in a study involving species of from 0.607 to 0.648, whereas genetic sim- 


VOLUME 106, NUMBER 1 


Table 3.— Continued. 


n mean min max 
Females 
MB 
G. t. bakeri 6 Tt AA 20:5 21.9 
G. texensis 15 236 19.9 22.8 
G. texensis 9 Zia 20.8 23:0 
G. personatus 13 22.93 DS Be 24.0 
G. attwateri 18 23.30 21.6 24.8 
G. attwateri 3 29:30 22.8 24.1 
SB 
G. t. bakeri 6 16.65 16.3 17.1 
G. texensis 9 16.85 16.5 WE: 
G. texensis 15 16.92 16.0 17.9 
G. attwateri 18 17.97 16.9 18.7 
G. attwateri 3 18.01 LiF 18.3 
G. personatus 13 18.21 173 18.9 
RB 
G. t. bakeri 6 8.40 8.0 8.8 
G. attwateri 18 8.56 8.2 9.7 
G. personatus 13 8.87 8.1 9.5 
G. texensis 15 9.15 8.5 9.6 
G. texensis 9 9.37 9.0 10.1 
G. attwateri 3 9.60 9.0 10.1 
M3 
ere bakers 6 6.88 6.5 73| 
G. texensis 15 7.36 7:0 7.8 
G. attwateri 3 7.50 7.4 7.6 
G. attwateri 18 WS 7 8.0 
G. texensis 9 To2 7.1 8.1 
G. personatus 13 Fae 2 ee 8.2 
Ml 
G. t. bakeri 6 8.25 7.8 8.5 
G. texensis 15 8.51 7.9 9.3 
G. texensis 9 8.66 8.1 9.0 
G. attwateri 18 8.99 8.5 9.4 
G. personatus 18 9.08 8.5 9.5 
G. attwateri 3 9.21 9.0 9.5 
IOC 
G. texensis 9 5.76 5.5 5.9 
G. t. bakeri 6 5.83 ee, 6.3 
G. texensis 15 5.86 5.4 6.3 
G. personatus 13 6.07 5.8 6.6 
G. attwateri 18 6.28 5.6 6.7 
G. attwateri 3 6.49 6.2 6.9 


ilarities produced from intraspecific com- 
parisons involving G. texensis were high 
(from 0.931 to 0.937). The new taxon has 
a high genetic similarity when compared to 
G. texensis (0.915). Similar comparison to 


G. bursarius yields a much lower value 
(0.648). Similarity values of 0.9 are com- 
parable to previous studies of geomyids 
when making intraspecific comparisons of 
populations, and the lower genetic similar- 


16 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Females 


4.0 


—4.0 


—6.0 


Females 


-6.0 -—4.0 —2.0 0.0 2.0 4.0 6.0 


CAN | 


. t. bakeri ° 
. p. streckeri + 
. attwateri oe 
. texensis on 


-6.0 -4.0 —2.0 0.0 2.0 4.0 6.0 


CAN | 


Fig.2. Plots of the centroids of taxa along the first two canonical variates. Top plots include the 95% confidence 
ellipse for each taxon around its centroid. Bottom plots include the dispersion of the mean canonical scores for 


each individual in the populations. 


ity values, seen when comparing the new 
taxon to G. bursarius, are within the range 
associated with interspecific comparisons 
(Block & Zimmerman 1991, Dowler 1982). 

Relationship between G. texensis and the 
new taxon is supported further by the mor- 
phometric analysis. In an analysis of the 
cranial characters that appear to separate G. 
personatus and G. attwateri from G. tex- 
ensis, the new taxon was associated with G. 
texensis 1n mastoid breadth, squamosal 
breadth, interorbital constriction, and length 


of molar toothrow. A relationship to G. tex- 
ensis also is seen in the two phenetic clus- 
tering results. The phenograms based on the 
correlation matrices describe the new taxon 
as similar to the two G. texensis popula- 
tions. Minimum-spanning analysis yields 
similar relationships with a population of 
G. texensis serving as the branching neigh- 
bor to the new taxon. 

These data support a proposal that the 
new taxon is closely related to G. texensis. 
The question then arises, do these new pop- 


VOLUME 106, NUMBER I 


Table 4.—Coefficients for canonical variates and the percent influence of each variable for the variates. 


Canonical Canonical Canonical 
Variate I Vaniate II Variate III 
Variable Variable Variable 
Character coeff. % coeff. % coeff. % 
Males 
CRL — 1.8433 28.91 —0.3363 3.36 —4.3026 41.24 
BL 0.6980 10.14 —4.5368 42.57 1.8989 17.09 
Pi —0.6005 6.09 2.1302 13.95 DALI i7ett 
PFD —0.9800 5.87 —0.6656 2.58 0.9685 3.60 
DIAS 1.1349 6.15 2.1888 VOY —0.5400 1.81 
ZB == lal (SUS) 11.26 0.6130 3.87 O727 3.47 
MB 2.0305 17.87 2.2454 L277 A093 LO 
SB 0.8427 5.67 — 1.8478 8.03 0.3642 52 
RB = O92 5 2.10 0.0785 0.18 OPIATE 0.27 
IOC —0.6612 152 0.8264 1523 0.6400 0.91 
M3 0.8382 2.38 le 2629 MES Ys) 0.6570 as Ke 
M1 0.7154 2.44 0.6592 1.45 0.3881 0.82 
Females 

CBL ~1.9191 35.85 0.7594 15.53 —0.0083 0.28 
BL 0.4529 7.94 ~1.7802 34.17 —0.2815 8.76 
PE O:5315 6.44 0.9833 13.05 1.3095 28.18 
PFD —0.4949 3.63 0.1148 0.92 =0)3503 4.56 
DIAS 0.6719 4.25 —1.1497 7.96 Oe) 2a 8.13 
ZB 0.1029 1.18 0.3581 4.49 0.5166 10.49 
MB 2.4876 26.45 —O5207, 6.06 0.5198 9.81 
SB —0.3687 3.06 0.9749 8.87 AOC, 19.20 
RB —1.0596 4.56 0.7393 3.48 0.6176 4.71 
IOC 0.7276 2.09 = 02932 0.92 0.7603 3.87 
M3 —0.4172 1.50 0.9869 3.88 —0.0849 0.54 
Mil 0.7203 3.05 0.1459 0.68 =0:1930 1.45 


ulations, isolated and 120 km distant from 
the previously known distribution of G. tex- 
ensis, constitute a new subspecies? An anal- 
ysis of cranial morphology identifies the new 
populations as being quite distinct from G. 
texensis. They are smaller in 22 of 28 com- 
parisons involving both sexes. Canonical 
discriminant analysis identifies measure- 
ments reflecting the length of the skull (BL, 
PL) and mastoid breadth accounting for 
much of the variation separating the taxa. 
The centroids produced from plotting these 
taxa along the first two canonical variates 
are widely separated, although the ellipses 
are overlapping. Separation is more clearly 
seen in analysis of females, where the two 
populations of G. texensis are closely as- 
sociated with each other, and each is well 


within the ellipse of the other. The new tax- 
on has a centroid separated from these two 
primarily along the axis of the second ca- 
nonical variate. The centroid is well outside 
the range of the ellipses of the G. texensis 
populations although the two centroids of 
G. texensis occur within the 95% ellipses of 
the new taxon. This distinction is less clear 
in males where the ellipses of both taxa in- 
clude the centroids of the other. The new 
taxon is also seen to be quite distinct from 
the two populations of G. texensis in the 
phenogram produced from the clustering of 
the correlation matrices. 

The populations of G. texensis represent 
two previously recognized subspecies, G. b. 
texensis and G. b. llanensis. Honeycutt & 
Schmidly (1979) identified primarily size- 


18 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Females 


[@) 
i=) 
CAN Ill 


G. texensis 


G.\texenais 


CAN |; 
(oo) 
oO 


Females 


0.99 
dt 


Correlation Matrix 


CAN Ill 


G. p. streckeri 


Males 


0.99 
—_ 


Correlation Matrix 


G. p. streckeri G. p. streckeri 
G. attwateri G. attwateri 
G. attwateri G. attwateri 
G. texensis G. texensis 
G. texensis G. texensis 
G. t. bakeri G. t. bakeri 
Females Males 
shes 2 0.64 0.16 75 1.00 O25 
Pistgaes G. p. streckeri PEATE G. p. streckeri 
G. texensis G. texensis 
G. texensis G. texensis 
G. t. bakeri G. attwateri 
G. attwateri G. attwateri 
G. attwateri G. t. bakeri 


Fig. 3. Plot of the centroids of taxa along their first three canonical variates. The centroids are connected 
by the branching order of the minmum-spanning analysis produced from the variance-covariance matrices of 
measurements of cranial characters. UPGMA phenograms produced from correlation and distance matrices of 


the cranial measurements. 


related differences between these taxa. When 
compared to these populations of G. tex- 
ensis, the new populations seem to have dif- 
ferences in cranial features that have re- 
sulted in alteration of both skull size and 
shape. Cranial morphology has been pro- 


posed to be greatly influenced by both soil 
composition and texture (Hendricksen 
1972, Smith & Patton 1988, Wilkins & 
Swearingen 1990). Wilkins & Swearingen 
(1990) noted an increase in the mean values 
of all cranial characters in populations of G. 


VOLUME 106, NUMBER 1 


personatus in sandy soils when compared 
to other soil types. This difference also ex- 
tended into a multivariate analysis which 
effectively separated populations inhabiting 
fine sandy loams, loamy fine sand, and fine 
sand, with an increase in size from smaller 
to larger along that axis. Geomys texensis 
occurs in sandy-loam soils in the central 
basin region of the Edwards Plateau. These 
are porous, well drained soils. The new tax- 
on from South Texas inhabits a loam sand, 
Atco soil that is denser and less friable. Go- 
phers in the heavier and denser soil are 
smaller and have cranial changes which al- 
ter the skull shape, and conform to the mod- 
els described by Smith & Patton (1988) and 
Wilkins & Swearingen (1990). 

Geomys texensis was previously known 
only from the central basin of the Edwards 
Plateau, and isolated there by barriers of 
stony to gravelly clay, clay, and shallow 
loamy soils (Godfrey et al. 1973). Block & 
Zimmerman (1991) described a scenario in- 
volving a late Wisconsin to Holocene en- 
vironmental change that isolated G. tex- 
ensis aS the warmer and drier conditions 
approximately 9000 B.P. accelerated ero- 
sion. The presence of G. texensis along the 
southern edge of the Edwards Plateau is 
plausible within the framework of this hy- 
pothesis. The distribution of G. texensis once 
could have been more widespread in south- 
central Texas, and probably extended fur- 
ther into southern Texas. A cooler climate 
and deeper soils would have allowed G. tex- 
ensis in the central basin to contact popu- 
lations south of the Edwards Plateau across 
the upper reaches of the Llano River drain- 
ages. Subsequent xeric conditions merely 
separated these populations, isolating those 
to the south and ultimately restricting them 
to the smaller pockets of suitable soils south 
of the newly created indurate soils of the 
plateau. Fossil remains of geomyids from 
cave deposits on the Edwards Plateau 
(Dalquest & Kilpatrick 1973) provide evi- 
dence for a wider distribution, with geo- 
myids ranging over at least the eastern por- 


19 


tions of the plateau from 10,000 to 4000 
B.P. 

The new subspecies described below 
demonstrates close affinities to G. texensis 
in cranial and chromosome morphologies, 
and in biochemical variation. Analyses of 
cranial morphology indicate that these pop- 
ulations resemble G. texensis, but appear to 
differ significantly in having size- and shape- 
related changes. These cranial differences 
alone could indicate a species distinction 
but the presence of a low level of genetic 
differentiation leads us to be more conser- 
vative. We conclude that, based on the ob- 
served variation and the extreme spatial 
separation, this new taxon is related to G. 
texensis and is an isolated, relictual popu- 
lation of this species, forming a distinctive 
subspecies. 


Geomys texensis bakeri, new subspecies 


Holotype. — Adult male, skin, skull, and 
body skeleton, no. 52310, Texas Coopera- 
tive Wildlife Collections (TCWC); from 1 
mi E D’Hanis, Medina Co., Texas; obtained 
on 3 Jan 1987 by R. M. Pitts, original no. 
1998. 

Distribution.—Two isolated populations 
have been found along separated drainages 
of the Frio River. One population occurs 
along the Sabinal and Frio rivers in Uvalde 
and Zavala counties. A second population 
in Medina County is restricted to soils along 
Seco and Parker creeks, tributaries of the 
Frio River. Both populations are associated 
with nearly level Atco soil (Stevens & Rich- 
mond 1976, Dittmar et al. 1977), which has 
a patchy distribution in this region. The soil 
is well drained and consists of sandy surface 
layers with loam extending to as deep as 2 
m. This soil is associated with stream ter- 
races formed by the drainage systems in each 
locality. These two populations are widely 
separated (40 km); however, there may be 
additional populations along Seco and Par- 
ker creeks as they flow southward to join 
the Frio River. The nearest geomyids are 


20 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


G. attwateri (Medina County), G. persona- 
tus streckeri (Dimmit County), and G. ft. tex- 
ensis (Kimble County). 

Description.—Small size which is es- 
pecially evident in the measurements of 
body length, skull length (CBL, BL), nasal 
length (LN), and skull breadth (ZB, MB, 
SB). Pelage coloration is russet brown on 
the dorsum, and grades to a paler color along 
the sides. The basal portions of the hairs are 
gray. A dark dorsal stripe extends from the 
head to the rump. The ventral surface is 
white with gray coloration on basal parts of 
the hairs. The tail is sparsely haired and 
consists of a mixture of brown and white 
hairs. The feet are white haired. Subadult 
pelage is a tawny brown, whereas adult pel- 
age is a darker, richer brown, and appears 
glossy and more reflective. 

Pelage color appears paler in gophers col- 
lected in the more sandy surface soil along 
the Frio River in Uvalde and Zavala coun- 
ties then in those from Medina County. This 
difference in pelage coloration correlates 
with the much paler color of the substrate 
at this locality than the darker color of the 
loamy soil at the Medina locality. 

Karyotypic features.—The diploid num- 
ber is 70 and the fundamental number is 
68. The X-chromosome is a large acrocen- 
tric and the Y-chromosome is a medium- 
sized acrocentric. 

Measurements. —Measurements (in mil- 
limeters), as described in Williams & Geno- 
ways (1977), from 12 adult individuals from 
the two populations are listed in Table 4. 
Measurements of the holotype (TCWC 
52310) are: total length, 227; length of tail, 
66; length of hind foot, 27; length of ear, 6; 
greatest length of skull, 40.8; condylobasal 
length, 40.4; basal length, 38.3; palatal 
length, 26.3; prefrontal depth, 15.5; length 
of nasals, 13.3; length of diastema, 14.3; 
zygomatic breadth, 25.6; mastoid breadth, 
22.4; squamosal breadth, 16.9; rostral 
breadth, 9.4; interorbital constriction, 6.3; 
breadth across third molars, 7.1; length of 
maxillary toothrow, 8.5. 


Comparisons. —Cranial measurements of 
individuals of G. t. bakeri are smaller in size 
than those of G. attwateri, G. p. streckeri 
and G. bursarius major (Baker & Genoways 
1975). This is especially evident in the mea- 
surements reflecting the length (CBL, BL) 
and breadth (ZB, MB, SB) of the skull. Geo- 
mys b. major is a larger gopher in all com- 
parative external and cranial measurements 
as seen when comparing the measurements 
herein to those given by Baker & Genoways 
(1975). Geomys attwateri 1s the taxon near- 
est geographically to G. t. bakeri. It has ex- 
ternal measurements comparable to those 
of G. t. bakeri, but the skull is longer (CBL, 
BL) and wider. Significant differences are 
detectable in mastoid breadth, squamosal 
breadth, and interorbital constriction (G. t. 
bakeri males MB = 21.9, SB = 16.9, IOC 
= 5.9; females MB = 21.1, SB = 16.6, IOC 
= 5.8; G. attwateri males MB = 24.0, SB = 
18.6, IOC = 6.4; females MB = 23.3, SB = 
18.0, IOC = 6.3). Geomys attwateri has a 
paler pelage that is a buffy tan in color, and 
the pelage has a uniform, nonglossy ap- 
pearance. Other populations of G. attwateri 
in the eastern part of southern Texas have 
pelage colors that are similar to that of G. 
t. bakeri. 

The karyotypes of Geomys attwateri and 
G. personatus differ from that of G. t. bakeri 
in that they have a large, submetacentric 
X-chromosome and a small, metacentric 
autosome (Davis et al. 1971, Honeycutt & 
Schmidly 1979, Tucker & Schmidly 1981). 
Geomys texensis (Honeycutt & Schmidly 
1979) and races of Geomys bursarius (Baker 
et al. 1973) have an identical diploid and 
fundamental number, and appear identical 
when standard karyotypes are compared. 

Etymology.—The subspecific name is a 
patronym and is selected to honor Dr. Rob- 
ert J. Baker for his many contributions to 
mammalogy, particularly to the systematics 
and evolution of Geomys, as well as his 
overall research program, activity in pro- 
fessional societies, and involvement in 
graduate education. 


VOLUME 106, NUMBER 1 


Acknowledgments 


We very much appreciate the help of many 
individuals. E. Abbott, J. Berry, J. Fell, J. 
Raney, K. Raney, J. Scharninghausen, E. 
Uptagrafft assisted in doing field work. R. 
Lopez, J. Lopez, and J. Lopez assisted in 
the preparation of specimens. Mr. DeWitt 
of Zavala County graciously allowed us to 
collect on his ranch. We thank S. Anderson 
(American Museum of Natural History), F. 
Stangel (Midwestern University), D. Wilson 
(National Museum of Natural History), H. 
Garner (Tarleton State University), G. 
Baumgardner (Texas Cooperative Wildlife 
Collections), R. Martin (Texas Natural His- 
tory Collection), R. Owen (The Museum, 
Texas Tech University), and their respec- 
tive staff, who provided specimens through 
loan or making them available to us during 
our visit to their museum collections. Dr. 
D. Schmidly, Dr. J. McEachran and Dr. S. 
Johnston were involved in valuable discus- 
sions that helped direct the development of 
this study. We greatly appreciate the many 
valuable suggestions concerning the final 
manuscript that were made by Dr. J. Knox 
Jones, Dr. R. C. Dowler, and M. Carleton. 
Financial support for the field work and lab- 
oratory materials was received from the 
Texas Agricultural Experiment Station Ex- 
panded Research Area funds and Program 
Development funds. This is contribution 
number 6 of the Center of BioSystematics 
and BioDiversity. 


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species of Geomys bursarius (Mammalia: Geo- 
myidae) from Texas and New Mexico.—Occa- 
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University 29:1-18. 

———,, M. W. Haiduk, L. W. Robbins, A. Cadena, & 

B. F. Koop. 1982. Chromosomal studies of 

South American bats and their systematic im- 

plications.—Special Publication Series, Pyma- 

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,&S.L. Williams. 1972. A live trap for pocket 

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1320-1322. 


2A 


: ,& J.C. Patton. 1973. Chromosomal 
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Block, S. B., & E. G. Zimmerman. 1991. Allozymic 
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Southwestern Naturalist 36:29-36. 

Dalquest, W. W., & W. Kilpatrick. 1973. Dynamics 
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Davis, B. L., S. L. Williams, & G. Lopez. 1971. Chro- 
mosomal studies of Geomys. —Journal of Mam- 
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Davis, W. B. 1940. Distribution and variation of 
pocket gophers (genus Geomys) in the south- 
western United States.— Bulletin, Texas Agri- 
cultural Experiment Station 590:1-38. 

Dittmar, G. W., M. L. Deike, & D. L. Richmond. 
1977. Soil survey of Medina County, TX. Soil 
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Printing Office, Washington, D.C. 

Dowler, R.C. 1982. Genetic interactions among three 
chromosomal races of the Geomys bursarius 
complex (Rodentia: Geomyidae).— Unpub- 
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Godfrey, C. L., G.S. McKee, & H. Oakes. 1973. Gen- 
eral soils map of Texas. Texas Agricultural Ex- 
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Harris, H., & D. A. Hopkinson. 1976. Handbook of 
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Hart, E. B. 1978. Karyology and evolution of the 
plains pocket gopher, Geomys bursarius. —Oc- 
casional Papers of the Museum of Natural His- 
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Hendricksen, R. L. 1972. Variation in the plains 
pocket gopher (Geomys bursarius) along a tran- 
sect across Kansas and eastern Colorado.— 
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75:322-368. 

Honeycutt, R. L., & D. J. Schmidly. 1979. Chro- 
mosomal and morphological variation in the 
plains pocket gopher, Geomys bursarius, in Tex- 
as and adjacent states. — Occasional Papers, The 
Museum, Texas Tech University 58:1—54. 

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Kennerly, T. E., Jr. 1954. Local differentation in the 

pocket gopher (Geomys personatus) in southern 


22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Texas.—The Texas Journal of Science 6:297-— 

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Haufler. 1990. Proteins I: isozyme electro- 
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ritz, eds., Molecular systematics. Sinauer As- 
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Owen, J. G., & M. A. Chmielewski. 1985. On ca- 
nonical variates analysis and the construction 
of confidence ellipses in systematic studies.— 
Systematic Zoology 34:366-374. 

Rogers, J.S. 1972. Measures of genetic similarity and 
genetic distance.—Studies in Genetics, VII, 
University of Texas Publication 7213:145-153. 

Rohlf, F. J. 1988. NT-SYS-pc: numerical taxonomy 
and multivariate analysis system, version 1.5. 
Exeter Software, Setauket, New York. 

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581-776. 

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Carolina, 441 pp. 

. 1988b. SAS/STAT user’s guide, release 6.03 

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lina, 1028 pp. 

Schmidly, D. J., & F.S. Hendricks. 1976. Systematics 
of the southern races of Ord’s kangaroo rat, Di- 
podomys ordii.—Bulletin of the Southern Cali- 
fornia Academy of Science 75:225-—237. 

Smith, M. L., & J. L. Patton. 1988. Subspecies of 
pocket gophers: causal bases for geographic dif- 
ferentiation in Thomomys bottae.—Systematic 
Zoology 37:163-178. 

Stevens, J. W.,& D. L. Richmond. 1976. Soil survey 
of Uvalde County, TX. Soil Conservation Ser- 
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United States Government Printing Office, 
Washington, D.C. 

Swofford, D. L., & R. B. Selander. 1981. BIOSYS-1: 
a Fortran program for the comprehensive anal- 
ysis of electrophoretic data in population ge- 
netics and systematics. — Journal of Heredity 72: 
282-283. 

Tucker, P. K., & D. J. Schmidly. 1981. Studies ofa 
contact zone among three chromosomal races 
of Geomys bursarius in east Texas.—Journal of 
Mammalogy 62:258-272. 

Wilkins, K. T., & C. D. Swearingen. 1990. Factors 
affecting historical distribution and modern 
variation in the South Texas pocket gopher Geo- 
mys personatus.—The American Midland Nat- 
uralist 124:57-72. 


Williams, S. L., & H. H. Genoways. 1977. Morpho- 
metric variation in the tropical pocket gopher 
(Geomys tropicalis).—Annals of the Carnegie 
Museum 46:245-264. 

———, & 1978. Morphometric variation in 

the desert pocket gopher (Geomys arenarius). — 

Annals of the Carnegie Museum 47:541-570. 

,& 1980. Morphological variation in 

the southeastern pocket gopher Geomys pinetis 

(Mammalia: Rodentia).— Annals of the Carne- 

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, & 1981. Systematic review of the 

Texas pocket gopher, Geomys personatus 

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gie Museum 50:435-473. 


(MJS and JWB) Department of Wildlife 
and Fisheries Sciences, Texas A&M Uni- 
versity, College Station, Texas 77843-2258, 
U.S.A.; (RMP) ARPERCEN, Attn. OPC, 
9700 Page Blvd, St. Louis, Missouri 63132, 
USS.A. 


Appendix 


Specimens examined.—Two hundred ninety seven 
specimens were used from the following collections: 
American Museum of Natural History (AMNH), Mid- 
western University (MSU), Tarleton State University 
(TSU), Texas Cooperative Wildlife Collections 
(TCWC), Texas Natural History Collection (TNHOQ), 
Texas Tech University (TTU), National Museum of 
Natural History (USNM). 

Geomys attwateri. —(85). Texas: Atascosa Co.: 6 mi 
W Campbellton, 1(TNHC); 2 mi NW Campbellton, 
3(TCWC); 1 mi-E Lytle, 1647 CWO); 7 mr EB Pyle 
16(TNHC); 2.4 mi SE Lytle, 3(TCWOC); 7 mi SE Lytle, 
8(TCWC). Bexar Co.: 15 mi SE San Antonio, 1 (TSU). 
Frio Co.: 1 mi N Moore, 3(TCWC); Pearsall city limits, 
2(TCWC); 2.25 mi S, 1 mi E Pearsall, 1(TCWC); Mc- 
Coy, 1(TNHC); 0.3 mi E McCoy, 1(TNHC); 2 mi N 
Pleasanton, 7/TNHC). Medina Co.: 5 mi W Devine, 
4(TCWC); 7.2 mi E Yancy, 1(TCWO), 1(TSU). Wilson 
Co.: 11 mi NW Floresville on HWY 181, 6(TCWOQ); 
4 mi W Floresville, 1(TNHC); 1 mi W Floresville, 
2(TCWC); %4 mi S, 242 mi E Floresville, 4(TCWQ); 3.2 
mi NW Poth, 1(TNHC); 3.6 mi SSE Poth, 1(TNHQ); 
5.4 mi W San Antonio River, between Floresville and 
Pleasanton, 1(TNHC). 

Geomys personatus streckeri. —(68). Texas: Dimmit 
Co.: 13 mi N Carrizo Springs on HWY 277, 3(TTU); 
13 mi NE Carrizo Springs, 3(TTU); 13 mi NE Carrizo 
Springs on US HWY 277, 5(TTU); 4 mi N, 9 mi W 
Carrizo Springs, 1(TCWC); 1 mi S Carrizo Springs, 
700 ft, 27TCWC); 15 mi S, 11 mi W Carrizo Springs, 


VOLUME 106, NUMBER 1 


5(TCWC); 1'2 mi E Carrizo Springs, 6(TCWC); E Car- 
rizo Springs, 3(TTU); Carrizo Springs, 30(TNHO), 
11(1CWC); near Carrizo Springs on HWY 277, 5(TTU); 
1.0 mi SW Carrizo Springs, HWY 186, 4(TCWOC); 2 
mi S Carrizo Springs, low water crossing Dentonio Rd, 
1(TCWC). 

Geomys texensis bakeri.—(35). Texas: Medina Co.: 
1 mi E D’Hanis, 5(TCWC); D’Hanis, 5(TCWC); 54 
mi W Hondo, 6(TCWC); 6.2 mi W Hondo, 4(TCWC). 
Uvalde Co.: 16 mi S Sabinal on FM 187, 6(TCWOC); 
17 mi S Sabinal on FM 187, 4(TCWC). Zavala Co.: 
18 miS Sabinal on FM 187, 1(TCWC); 7, miS Uvalde 
County line, 4(TCWC). 

Geomys texensis texensis.—(108). Texas: Gillespie 
Co.: 13 mi N Fredericksburg, 1(TSU); 1 mi N Fred- 
ericksburg, 2(TCWC); 0.5 mi N Fredericksburg, 
1(TCWC); 9 mi W Fredericksburg, 1(TNHC). Kimble 
Co.: Junction, 3(TCWC). Llano Co.: 2.6 mi N, 1.8 mi 
E Castell, 5(TTU); 6.4 mi E Castell, 1(TCWC); Castell, 
1(TTU); 6.4 mi E Castell 1(TCWC); 1 mi E Castell 
1(TCWOC); 1.2 mi W Castell on FM 152, 1(TCWC); 8 
mi S, 0.9 mi W Kingsland, 4(TTU); 9.2 mi S, 1.1 mi 
E Kingsland, 1(TTU); 10 mi S, 1.8 mi E Kingsland, 


23 


2(TTU); 2.9 mi NW Llano on HWY 71, 2(TTU); 0.2 
mi N, 8.7 mi W Llano, 3(TTU); Drier Cr at Lone 
Grove, 7 mi W Llano, 10(TCWC); 4 mi W Llano 
1(TCWO), Llano, 1(TCWC); 0.2 mi E Llano, 1(TCWO); 
1 mi E Llano, 2(TCWOC); 2 mi E Llano, 4(TCWC); 7.2 
mi E Llano, 1(TCWC); Oatman Cr, 3 mi S Llano, 
6(TCWC); 51.6 mi W Austin on HWY 71, 2(TTU); 3 
mi S Jct FM 268 and HWY 29 on 29, 2(TTU); 9.3 mi 
N Jct Texas 29, Texas 16 on Texas 16, 1(TTU). Mason 
Co.: 3.1 mi E Art, 1(TCWO); Art city limits, 3(TCWC); 
12 mi N Mason, 6(MWU)); 3.6 mi N, 1.5 mi W Mason, 
1(TTU); 1 mi N, 1.1 mi W Mason, 4(TTU); 12 mi W 
Mason, 2(MWU); 9.4 mi W Mason on US 377, 3(TTU); 
Mason, 1(TCWC); 1 mi E Mason, 6(TCWC); 6.5 mi 
E Mason on Texas 29, 1(TTU); 2.0 mi S, 2.7 mi W 
Castell, 1(TTU); 5 mi S Mason, 1(MWU). in Mason 
Co.: 34% mi W Castell, 3(TCWOC); 0.3 mi S, 1.5 mi W 
Castell, 3(TTU); 0.3 mi S, 0.9 mi W Castell, 1(TTU); 
0.3 mi S, 0.8 mi W Castell, 1(TTU); 2.6 mi S, 3.0 mi 
W Castell, 1(TTU); 0.7 miS, 2.1 mi W Castell, 2(TTU); 
1.0 miS, 2.3 mi W Castell, 2(TTU); 11 mi NE London, 
HWY 377, 2(TCWC); 13 mi NE London, HWY 377, 
2(TCWC). 


PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 24-33 


IDENTIFICATION OF BIRD SUBFOSSILS FROM CAVE 
SURFACE DEPOSITS AT ANJOHIBE, MADAGASCAR, 
WITH A DESCRIPTION OF A NEW GIANT 
COUA (CUCULIDAE: COUINAE) 


Steven M. Goodman and Florent Ravoavy 


Abstract. —A collection of bird subfossils from cave surface deposits in north- 
western Madagascar is described. The majority of specimens represent taxa 
that still occur in the region. The exception is a partial pelvis referable to the 
genus Coua (Cuculidae: Couinae), but which is considerably larger than any 
known species. This specimen is described as a new species. 

Résume.—Une collection d’ossements subfossiles d’oiseau provenant des 
gisements cavernicoles de surface du Nord-Ouest de Madagascar a été décrite. 
La plupart des spécimens représentant des taxons qui’existent encore dans la 
région. La seule exception est constitueé par un bassin incomplet pouvant se 
rapporter au genre Coua (Famille Cuculidae: sous-famille Couinae), mais qui 
est nettement plus grand que toute espéce connue. Ce spécimen est décrit 
comme nouvelle espéce. 

Famintinana. —Nofantarina ireo tahirina taratsiefan-taolam-borona hita tany 
amin’ireo sompitrakoran-johy amin’iny faritra avaratr’andrefan’1 Madagasikara 
iny. Ny ankamaroan’ireo santiona azo dia tsy hafa noho ireo karazana mbola 
fahita ao am-paritra. Ny hany niavaka tamin’ireo dia ilay sila-taola-maoja iray 
izay azo raisina ho an’ny sokajy Cova (Tarika Cuculidae: zanatarika Couinae), 
saingy lehibe lavitra noho izay karazana rehetra fantatra ho misy. Naraikitra 


ho karazana vaovao ity santiona ity. 


The subfossil fauna of Madagascar is well 
known for its remarkable array of lemurs 
(e.g., Vuillaume-Randriamanantena 1982, 
Simons et al. 1990) and elephant birds (Ae- 
pyornithiformes) (Andrews 1894, Lamber- 
ton 1934, Battistini 1965). The study of 
hundreds of animal bones recovered at var- 
ious sites on Madagascar has provided in- 
sight into the Holocene faunas of the island, 
inference about ecological change, as well 
as the reasons that a portion of these taxa 
have gone extinct in the past few thousand 
years (Dewar 1984, MacPhee 1986). In par- 
allel situations on other islands, subfossil 
bird bones have provided important infor- 
mation about paleoenvironments and the 
effects of anthropogenic perturbations 
(Steadman 1989, James & Olson 1991, Ol- 


son & James 1991). While there are nu- 
merous archaeological sites on Madagascar 
that have yielded bird bones, this material 
with the exception of elephant birds has been 
rarely studied and thus not synthesized into 
the current working knowledge of the Ho- 
locene environment of the island. 

In 1983 and 1986 excavations were car- 
ried out in northwestern Madagascar near 
Mahajanga in the Grottes d’Anjohibe (An- 
dranoboka), by the Laboratoire de Prima- 
tologie et de Paléontologie des Vertébrés, 
Service de Paléontologie, Universite d’An- 
tananarivo (formerly Université de Mada- 
gascar), and Duke University Primate Cen- 
ter. The focus of these studies was primates 
(Vuillaume-Randriamanantena et al. 1985, 
Simons et al. 1990), but a wide array of 


VOLUME 106, NUMBER 1 


other animal remains were recovered. Dur- 
ing the 1986 field season approximately 
1100 whole or fragmented subfossil bones 
of non-primate vertebrates were excavated, 
and, excluding the bird material, these have 
been described by Ravoavy (1991). In this 
paper we present information on 94 speci- 
mens of avian bones recovered during the 
1983 and 1986 field seasons at Anjohibe. 


Description of Site 


Anjohibe is located in Mahajanga Prov- 
ince, approximately 80 km NE of Maha- 
janga, and is part of a series of caves gen- 
ceally a neferred to. as. the. Grottes 
d’Andranoboka (Decary 1938, de Saint- 
Ours & Paulian 1953). One cave in partic- 
ular is called the Grotte d’Anjohibe (de 
Saint-Ours 1953). The surface material de- 
scribed herein was collected in the northern 
end of de Saint-Ours & Paulian’s (1953) 
““Grotte Principale no. 1”’ (Ravoavy 1991). 

The cave is over 1200 m long from north 
to south, and with floor to ceiling heights 
in some places exceeding 12 m. There are 
numerous entrances and side passages to the 
cave. It still has “‘active’’ calcite formations 
(de Saint-Ours & Paulian 1953), and has 
been used in recent times by local people 
for a variety of activities (de Saint-Ours 
1953). 

All of the material collected during the 
1983 and 1986 seasons was from surface 
deposits (MacPhee et al. 1984; E. Simons, 
pers. comm.). Bone is washed into the cave 
by floods during the rainy season. Also, some 
animals fall through aven, which are straight 
vertical holes open from the ground surface 
down to the cave floor as much as 60 m 
below. There is also a portion of the cave 
where the ceiling has collapsed (area M of 
de Saint-Ours & Paulian 1953) and forest 
vegetation is found on the floor of the cave. 
No radiocarbon date has been determined 
for any of the material recovered from the 
cave (Godfrey & Vuillaume-Randriama- 
nantena 1986; E. Simons; pers. comm.). 


25 


Table 1.— Bird species and minimum number of in- 
dividuals identified from bones recovered during the 
1983 and 1986 field seasons at Anjohibe. Excludes 
material of Coua berthae. 


Minimum number 
of individuals 


Taxa 1983 1984 


Buteo brachypterus 1 
Falco newtoni 

Coturnix sp. 

Numida meleagris 2 
Turnix nigricollis 

Coracopsis vasa 

Tyto alba 

Otus rutilus 

Merops superciliosus 

Hypsipetes madagascariensis 1 
Foudia madagascariensis 


—e ee NNBEBNAN 


Thus, we have no idea when the material 
was deposited, or the span of years repre- 
sented. Most of the bird bones show no signs 
of permineralization and are probably com- 
paratively recent in age. The only exception 
is a Coua pelvis which has some surface 
mineralization, although the underlying 
structure is bone, and thus may be older 
than the balance of the material. 


Birds Recovered from the 
Surface Deposits 


A total of 12 and 82 bird bones recovered 
during the 1983 and 1986 field seasons (re- 
spectively) was used in this study. Most of 
the species identified (Table 1) are taxa that 
still occur in the area (Langrand 1990). The 
present natural vegetation of the region is 
dry deciduous forest; considerable portions 
of this habitat have been degraded in the 
past few centuries. The majority of birds 
identified from the surface deposits are spe- 
cies associated with open habitats and/or 
the forest edge. The hawk (Buteo brachyp- 
terus), falcon (Falco newtoni), parrot (Cor- 
acopsis vasa), and owl (Tyto alba) presently 
occur in a variety of habitats from grassland 
savanna to wooded environments. A par- 


26 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


tially unossified mid-shaft of a humerus, re- 
ferable to Tyto, was a young individual 
probably incapable of sustained flight, and 
this species almost certainly bred in the cave. 
The falcon may have roosted or even nested 
in the cave. The presence of these two rap- 
tors in the cave would account for some of 
the small mammal and amphibian bones 
recovered from the ground surface of the 
cave (Ravoavy 1991); these would have been 
originally deposited as regurgitated pellets. 
The guineafowl (Numida meleagris), but- 
tonquail (Turnix nigricollis), quail (Cotur- 
nix sp.), bee-eater (Merops superciliosus), 
and fody (Foudia madagascariensis) are 
generally found in grassland savanna, while 
a second species of owl (Otus rutilus) and a 
bulbul (Hypsipetes madagascariensis) occur 
in woodland areas or along the forest edge. 
Of these species, Turnix, Merops, Hypsi- 
petes, and Foudia have been identified from 
Tyto alba pellets collected on Madagascar 
(Langrand & Raxworthy, pers. comm.). 

The most common bird species recovered 
from these surface deposits was Numida. 
Adults of this species weigh over 1100 g 
(Urban et al. 1986) and are too heavy to 
have been carried into the cave by any rap- 
tor known to occur on the island. Numida 
may have been introduced to Madagascar, 
and it is now extensively hunted on the is- 
land by people (Langrand 1990). There was 
no clear sign of carnivore gnawing, butch- 
ering marks nor charring on the Numida 
bones, nor on any of the other bird material 
recovered from the cave. It is not clear what 
agent(s) was (were) responsible for the de- 
position of these bones in the cave. On a 
few occasions during the excavations Numi- 
da were observed flying in and out of the 
cave in an area with a collapsed ceiling and 
extensive vegetation. Thus, the Numida 
bones recovered from the surface deposits 
may be of individuals that naturally died 
within the cave. 

One bone, a pelvis, from the 1983 col- 
lection cannot be identified to any modern 
species. On the basis of numerous osteo- 


logical characters, the bone belongs to an 
exceptionally large coua (Coua), a subfam- 
ily of cuckoos (Cuculidae: Couinae) endem- 
ic to Madagascar. There are nine extant Coua 
spp. on the island. Coua caerulea, reynaudii, 
and serriana are found in humid forests; 
gigas, cursor, ruficeps, and verreauxii in the 
dry thorn scrub or dry deciduous forests; 
and cristata and coquereli in both wet and 
dry forest types (Langrand 1990). A tenth 
species, C. delalandei, once occurred on Ile 
Sainte Marie, 8 km off the northeastern coast 
of Madagascar, and possibly on the main 
island itself, but has gone extinct in the past 
150 years (Langrand 1990, Goodman 1993). 
Coua delalandei and gigas are the largest 
known recent couas, measuring approxi- 
mately 57 and 62 cm (respectively) in total 
length (Langrand 1990). 

Muilne-Edwards & Grandidier (1895) de- 
scribed an undated subfossil tarsometatar- 
sus as a new species, Coua primavea, which 
was excavated from a deposit on the west 
coast of Madagascar at Belo-sur-mer, south 
of Morondava, and about 660 km south of 
Anjohibe. They distinguished the subfossil 
from other modern couas by its size; the 
tarsometatarsus of primavea measured 84 
mm in total length, delalandei 70 mm, and 
gigas 69 mm (the latter two measurements 
are presumably from skin specimens). We 
have examined the type tarsometatarsus of 
primavea (Museum National d’Histoire 
Naturelle, Service de Paléontologie, Paris, 
registration MAD 7078) and it measures 
83.2 mm in total length. On the basis of a 
regression analysis between tarsometatarsus 
and pelvis length of modern couas and pri- 
mavea (see Discussion), the pelvis recov- 
ered from Anjohibe cannot be referred to 
any known species of Coua and we propose 
to call it: 


Coua berthae, new species 
Figss lo 


Holotype. —Left half of pelvis, collections 
of the Laboratoire de Primatologie et de Pa- 


VOLUME 106, NUMBER 1 


27 


Fig. 1. 


léontologie des Vertébrés, Service de Pa- 
léontologie, Université d’Antananarivo, 
UM 6264 (Figs. 1, 2). Collected during the 
1983 field season. 

Locality. —From surface deposits, Grotte 
d’Anjohibe (Andranoboka), Grotte Prin- 
cipale no. 1 (de Saint-Ours & Paulian 1953), 
Fivondronana (subprefecture) of Mahajan- 
ga, Province of Mahajanga, about 80 km 
NE Mahajanga, Madagascar (coordinates: 
1) Laborde system — X = 1172, y = 448 and 
Zits 32'S, 46°53'E). 

Chronology. —No radiometric date is 
available from the site. Presumed to be 
Quaternary, probably Holocene. 

Measurements of holotype. —Length— 
from cranial border of the ilia to the Spinae 
iliocaudales, 68.2 mm; length along the ver- 
tebrae—from most cranial vertebra fused 
with the Os lumbosacrale to the most caudal 
vertebra fused with the Os lumbosacrale, 
58.1 mm; smallest breadth across the Partes 
glutaeae of the ilia, 18.3 mm; and greatest 
breadth across the Partes glutaeae of the ilia, 
29.2 mm. (See von den Driesch, 1976, fig. 
59a, c for illustrations and descriptions of 
these measurements.) 

Etymology.—Named in honor of Ma- 
dame Berthe Rakotosamimanana, Direc- 
teur du Laboratoire d’Anthropologie, and 


Pelvis of Coua berthae, new species, holotype UM 6264, (left) dorsal and (right) ventral views. 


Professor, Service de Paléontologie, Uni- 
versité d’Antananarivo, who for many years 
has helped students and foreign researchers 
working on Madagascar in the fields of pa- 
leontology and zoology, and also for her 
contribution to these disciplines. 

Diagnosis.—Distinctly larger than any 
extant member of the genus Coua (Table 2). 
On the basis of a regression analysis of pel- 
vic and tarsometatarsus measurements (see 
Discussion), C. berthae is larger than C. de- 
lalandei, a large recently extinct ground- 
dwelling species; C. primavea, an undated 
fossil species only known from a single tar- 
sometatarsus; and all living Coua spp. 

Paratype.—Complete tarsometatarsus, 
Laboratoire de Paléontologie, Muséum Na- 
tional d’Histoire Naturelle, Paris, MAD 
5490, collected at Ampasambazimba in 
1911 by A. Grandidier and presented as a 
gift of the Academie Malgache. 

Measurements of paratype. — Total length, 
92.9 mm; proximal width 12.4 mm; and 
distal width 13.1 mm. 

Discussion. — Within the four extant couas 
(C. caerulea, reynaudii, ruficeps, and cris- 
tata) for which there was more than one 
skeletal specimen of each available for study, 
there is little intraspecific variation in three 
pelvic measurements and in the greatest 


28 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Comparison of Coua gigas (FMNH 345635) pelvis (left) and Coua berthae (UM 6264) pelvis (right). 
The three views from the top are ventral, left lateral, and dorsal. 


length of the tarsometatarsus (Table 2). C. 
cristata shows some geographic variation in 
size (Milon 1950), which accounts for the 
greater variability in the range of these mea- 


surements in this species than the other 
three. Within these four species there is no 
discernable sexual dimorphism in the skel- 
etal measurements. A strong linear rela- 


VOLUME 106, NUMBER 1 


Table 2.—Pelvis and tarsometatarsus measurements (mm) of Coua spp. 


Length along 
Species Length' the vertebrae 
berthae, sp. nov.” 68.2 58.1 
primavea [62.5] — 
gigas (n = 1) 49.3 42.3 
caerulea 44.28 38.70 
n=5 n=6 
(42.0-45.8) (37.3-40.1) 
cristata 33.50 29:10 
n=7 n=7 
(30.2-36.5) (26.8-29.8) 
reynaudii 35.98 31.33 
n=4 n=4 
(35.3-36.8) (30.1-—32.3) 
ruficeps 40.15 S228 
n=6 n=6 
(39.3-40.8) (30.6-—34.3) 
serriana (n = 1) 45.9 37.6 


Tarsometatarsus 
Cranial Smallest Greatest 
breadth breadth of ilia length 
ge 18.3 92.9 
_ — B3.2° 
25.8 12:5 68.7 
24.20 13.79 53-20 
n=5 n=6 =5 
(23.6—24.7) (13.3-14.0) (53.1-58.0) 
17.44 10.29 42.96 
is n=7 wa 
(15.6-19.0) (9.4—-11.6) (39.3-47.8) 
L778 9.80 47.60 
n=4 n=4 n=3 
(17.2-18.6) (9°7=11-2) (46.4-48.4) 
21.05 10.97 57.68 
n=6 = 6 n=5 
(20.4—21.6) (9.8-11.8) (55.2-60.2) 
22.6 13:3 61.0 


! See text p. 27 for definitions of pelvis measurements. Measurement in brackets is inferred on the basis of a 


regression analysis. 


2 C. berthae pelvic measurements from type specimen, Service de Paléontologie, Université d’Antananarivo, 
UM 6264; and tarsometatarsus from Muséum National d’Histoire Naturelle, Service de Paléontologie, Paris, 


MAD 5490. 


3 Muséum National d’Histoire Naturelle, Service de Paléontologie, Paris, registration MAD 7078. Milne- 
Edwards & Grandidier (1895) gave the greatest length measurement of this element as 84 mm. 


tionship exists among six Coua spp., for 
which at least one skeleton of each was 
available, between the lengths of the pelvis 
and of the tarsometatarsus (77 = 0.80), and 
species with multiple specimens form dis- 
tinct clusters (Fig. 3). On the basis of this 
relationship, the point at which the 83.2 
mm primavea tarsometatarsus intersects the 
regression line corresponds to a pelvic length 
of approximately 62.5 mm. Similarly, the 
berthae pelvis length of 68.2 corresponds to 
a tarsometatarsus length slightly larger than 
90 mm (Fig. 3). Thus, by extrapolation, the 
primavea tarsometatarsus and the berthae 
pelvis are not the same taxon. 

Further evidence for the distinction be- 
tween C. berthae and C. primavea comes 
from a tarsometatarsus recovered at the fa- 
mous subfossil lemur site of Ampasamba- 
zimba on the High Plateau. This element, 
the paratype of C. berthae, measures 92.9 


mm, close to the length predicted by the 
regression analysis (Fig. 3). Moreover, the 
Ampasambazimba tarsometatarsus is 9.7 
mm larger than the type of C. primavea. 
This difference is greater than any size vari- 
ation found within extant Coua spp. (Table 
2): 

Coua delalandei, a species that has gone 
extinct in the past 150 years, has the longest 
tarsometatarsus of any known recent Coua. 
It is represented by less than 15 skin spec- 
imens in museums, and no skeletal material 
is available. The tarsometatarsus length of 
this species, as measured from museum 
skins, is 70 mm (Milne-Edwards & Gran- 
didier 1895), considerably smaller than the 
tarsometatarsus measurement of primavea 
or berthae. 

In numerous groups of cursorial birds 
there is an outgrowth of the ischium into a 
prominent tuberculum preacetabulare 


30 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


95 
90 
85 
80 
75 
70 
65 
60 
55 
50 
45 
40 
35 
30 


tarsometatarsus length 


25 


30 35 40 45 £50 


primavea 


T 
@ 
=] 
amp 
= 
Q 
® 


e-gigas 
O-serriana 
m@-caerulea 


- ruficeps 
- reynaudii 


O - cristata 


55 60 65 70 75 


pelvis length 


Pig. 3: 


Plot of tarsometatarsus length versus pelvis length in six species of modern Coua. The linear regression 


equation is y = 2.13 + 1.29x (r? = 0.80). Dotted lines are extrapolations of measurements based on the regression 


analysis. 


(=pectineal process), the place the M. am- 
biens arises (Baumel 1979). In ground- 
dwelling cuckoos there is considerable de- 
velopment of this process and the M. abiens 
is present; this muscle is thought to help 
with the “facility of leg movement in run- 
ning” (Berger 1952, 1953). In gigas, a ter- 
restrial swift-running species and the largest 
extant Coua spp., the tuberculum preace- 
tabulare is prominent. Absolutely and pro- 
portionately, this process as well as the an- 
titrochanter is larger in berthae than gigas 
(Fig. 2), and the former was presumably an 


extremely large and swift-running species of 
Coua. 

Weights are available for seven of the 
modern skeletal specimens measured, rep- 
resenting gigas, caerulea (3), cristata, and 
reynaudii (2). When weight is regressed 
against length of pelvis, a clear relationship 
emerges, which is best explained by a log- 
arithmic curve (Fig. 4, 77 = 0.97). However, 
since this curve abruptly flattens out, the 
point at which the berthae pelvis measure- 
ment would intersect the regression line is 
at an exceptionally heavy weight. A more 


VOLUME 106, NUMBER I 


31 


70 berthae 
| 
65 7 
| 
60 | 
| 
Ss 55 : 
S) | 
c 
a I 
mas > e | 
= | 
= | 
oO 45 = 
a | 
~ | 
4O : 
6 e - gigas | 
35 ™ = - caerulea 
A - reynaudii i 
O - cristata | 
30 
100 200 300 400 500 600 700 800 
weight (g) 


Fig. 4. Plot of pelvis length versus body mass in four species of Coua. The two regression lines are based 
on linear and logarithmic analyses. The dotted line is the extrapolation of Coua berthae’s mass based on the 


linear regression analysis. 


conservative approach is to examine the 
same relationship with linear regression (Fig. 
4, r> = 0.92), and thus by extrapolation, the 
pelvis length of berthae would intersect this 
curve at about 740 g, which is the presumed 
approximate minimum weight of this spe- 
cies. Since no portion of the sternum or wing 
bones of berthae is known, it is impossible 
to determine if this species was volant. 
However, given its considerable body mass 
and that all Coua spp. have proportionately 
small wing bones (Milne-Edwards & Gran- 
didier 1879, Berger 1953), at the very least 
berthae almost certainly was not a strong 
flier. 

In modern Madagascar, terrestrial Coua 


spp. are regular victims of human trapping 
and hunting (Langrand 1990), and this is 
one of the causes that has been proposed 
for the demise of C. delalandei. During an 
unsuccessful search in April 1991 of the re- 
maining forests of Ile St. Marie for a rem- 
nant population of C. delalandei, it was 
found that there is still exploitation of wild 
animals and that C. caerulea is extensively 
hunted (Goodman, 1993). Although tempt- 
ing to infer human involvement, it is pre- 
mature to make any suppositions on when 
and why Coua berthae went extinct. The 
study of bird material already recovered 
from sites and new excavations with de- 
tailed stratigraphic control should elucidate 


32 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


some of the missing information on the 
number and timing of Quaternary bird ex- 
tinctions on Madagascar. 

Comparative material examined. —Os- 
teological material of Coua spp. is rare in 
collections and skeletons of C. delalandei, 
coquereli, cursor, and verreauxii are not 
available for comparison. With the excep- 
tion of delalandei, all of these birds are small 
species, and the absence of comparative 
material did not hamper the analysis. The 
pelvis of C. berthae was compared to mod- 
ern skeletal material of the following Coua 
spp. (see Acknowledgments for definitions 
of acronyms): gigas (FMNH 345635; UM 
uncataloged partial specimen), caerulea 
(AMNH 6429, 10070; FMNH 345642, 
345644, 352802; UM four uncataloged par- 
tial specimens; UMMZ 209201), cristata 
(AMNH 6430, 10071; MNHN 1883-512, 
1883-514, 1883-517; FMNH 345639; 
UMMZ 157526; USNM 432197, 432219, 
432238), reynaudii (FMNH 352797, 
352798; UMMZ 208403; USNM 208403), 
ruficeps (MNHN 1883-518, 1883-519, 
1883-521, 1883-522, 1883-523; USNM 
432195), and serriana (UMMZ 209202). 


Acknowledgments 


We are indebted to Madame B. Rakoto- 
samimanana, Université d’Antananarivo 
(UM), for permission to study the Anjohibe 
and modern osteological material under her 
care. For loaning or allowing us access to 
skeletal material in their collections we are 
grateful to G. Barrowclough, American Mu- 
seum of Natural History (AMNH), New 
York; C. Lefevre, Laboratoire d’Anatomie 
Comparée, J.-F. Voisin, Laboratoire de 
Zoologie, and D. Goujet, Laboratoire de Pa- 
léontologie, Muséum National d’Histoire 
Naturelle (MNHN), Paris; R. W. Storer, 
University of Michigan Museum of Zoology 
(UMMZ), Ann Arbor; and S. L. Olson, Na- 
tional Museum of Natural History (USNM), 
Washington, D.C. H. James initially located 
the Coua berthae tarsometatarsus in the 


MNHN. J. Sedlock kindly drew Fig. 2. E. 
Simons graciously provided information on 
the excavations at Anjohibe. The 1983 and 
1986 field seasons at Anjohibe were fi- 
nanced by grants from the Boise Fund of 
Oxford University and the National Geo- 
graphic Society to E. Simons. Goodman’s 
studies were supported by the E. T. Smith 
Fund of the Field Museum of Natural His- 
tory (FMNH) and Conservation Interna- 
tional. For comments on an earlier version 
of this paper we are grateful to H. James, 
O. Langrand, S. Olson and T. Schulenberg. 


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1991. Descriptions of 


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, E. L. Simons, N. A. Wells, & M. Vuillaume- 

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Olson, S. L., & H. F. James. 1991. Descriptions of 
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53 


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PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 34-45 


TWO NEW SPECIES OF BLIND SNAKE, GENUS 
TYPHLOPS (REPTILIA: SQUAMATA: TYPHLOPIDAB), 
FROM THE PHILIPPINE ARCHIPELAGO 


Addison H. Wynn and Alan E. Leviton 


Abstract.—Two new species of 7yphlops from the Philippine Islands are 
described. Typhlops castanotus has a distinctly bicolored pattern without a 
head to vent reduction in the number of pigmented scale rows on the dorsum. 
Typhlops collaris has a collar of unpigmented scales behind the head and a 
high number (>400) of middorsal scale rows. Sexual dimorphism is indicated 
for the total number of middorsal scales in T. castanotus, and for tail length 
in both species. A preliminary key is provided to the Philippine typhlopids. 


The blind snakes of the family Typhlopi- 
dae are among the least tractable snakes to 
study. Their small size, small samples from 
single localities, few readily accessible or re- 
liable external and skeletal characters, 
skewed sex ratios that may result from fac- 
ultative parthenogenesis, potential ease of 
transport (especially in inter-island trade of 
agricultural products), and poor descrip- 
tions with even less reliable illustrations of 
nominal species, all contribute to the veil 
of uncertainty that surrounds these animals. 

Largely due to paucity of material, little 
work has been done on Philippine typhlop- 
ids. Until the mid-1950s fewer than 50 blind 
snakes had been collected throughout the 
whole of the Philippines. Most were ob- 
tained between 1915 and 1921 by Edward 
Harrison Taylor, who also described eight 
of the 18 nominal species attributed to those 
islands (Taylor 1917, 1918, 1919, 1922). 
Moreover, most specimens collected before 
Taylor were poorly documented as to prov- 
enance, although all seem legitimately at- 
tributable to the Philippines, except 7yph- 
lops dichromatus Jan (listed by Taylor [1922] 
in his synonymy of T. ruficaudus). 

More recently, Savage (1950) described 
Typhlops hypogius and Typhlops hedraeus, 
each based on single specimens collected by 
A. W. Herre in 1940. Although McDowell 


(1974) dealt specifically with typhlopids 
from New Guinea and the Solomon Islands, 
he also commented on a number of closely 
related Philippine species. For instance, 
McDowell defined the “‘ruficaudus’’ group 
of Typhlops as those species without a rectal 
caecum (or, if present, a poorly defined rec- 
tal caecum), and having an anterior shift of 
the suture between the second and third up- 
per labials, resulting in a fusion of the dorsal 
portion of the glandular line at the base of 
both of these scales. He referred six species 
to this group (Typhlops hypogius, T. jagorii, 
T. kraali, T. luzonensis, T. ruber, and T. 
ruficaudus), but not a seventh, Typhlops 
canlaonensis, which is also clearly allied. Six 


4 McDowell (1974) states that the gland row lying 
under the posterior edge of the postnasal fuses with the 
gland row under the posterior edge of the preocular 
and second upper labial in all members of the T. rufi- 
caudus group, even those in which the imbricate por- 
tion of the preocular and second upper labial narrowly 
contact. In contrast, we find that in 7. /uzonensis (a 
species with contact between the preocular and second 
upper labial) the glandular lines do not fuse. Rather, 
they are separated by a distance corresponding to the 
width of contact between the preocular and second 
upper labial, as expected if the glandular line forms the 
scale base and mirrors the shape of the imbricate por- 
tion of each scale. Nonetheless, the second upper labial 
is reduced in size, as in other members of the T. ruficau- 
dus group. 


VOLUME 106, NUMBER 1 


of the seven species are endemic to the Phil- 
ippines; the seventh, 7. kraalii, is known 
only from the Kei Islands and Ceram (Mc- 
Dowell 1974). 

During the past 50 years, Harry Hoog- 
stral, Walter Brown, Angel Alcala, and 
Donald Hahn have each added large num- 
bers of specimens so that now there are over 
500 from throughout the Philippines in mu- 
seum collections. During our examination 
of much of this material, it became clear 
that several series of specimens cannot be 
readily assigned to described taxa. Two of 
these are described here as new species. We 
also provide a preliminary key to those spe- 
cies we currently recognize. 

Materials and methods.—All measure- 
ments and observations are based on spec- 
imens stored in 70% ethanol or 42% iso- 
propyl alcohol. If hemipenes were not 
everted, sex was determined by examina- 
tion of gonads or associated structures. To- 
tal length was measured to the nearest 1 
mm, and tail length and body diameter were 
measured to the nearest 0.5 mm. Relative 
eye size was determined with an optical mi- 
crometer. The number of middorsal trans- 
verse scale rows was determined by count- 
ing all middorsal. scales posterior to the 
rostral, including the terminal spine on the 
tail. All midventral scales between the men- 
tal scale and anterior lip of the vent were 
counted for the number of midventral 
transverse scale rows, and all midventral 
scales posterior to the vent, including the 
terminal spine, were counted for the num- 
ber of mid-subcaudals. The number of mid- 
dorsocaudal scales was determined by 
counting the middorsal scales posterior to 
the level of the preanal scales. This method 
was preferred over counting mid-subcau- 
dals because of the numerous irregularities 
in the subcaudals from loss of longitudinal 
scale rows on the underside of the tail, and 
the difficulty in determining the first sub- 
caudal at the posterior edge of the vent. In- 
tercalary scales on the dorsal midline were 
not counted except when occurring in pairs. 


35 


Terminology for head and rostral shape fol- 
lows Thomas (1976). In discussions of the 
number of pigmented scale rows in the dor- 
sal stripe, the middorsal and pigmented lon- 
gitudinal rows to both sides are included. If 
a particular longitudinal scale row is noted, 
it is counted from the middorsal. All body 
length measurements are from the anterior 
tip of the head. Hemipenal orientation and 
morphology follow Dowling & Savage 
(1960), except that medial refers to orien- 
tation toward the midline, and lateral away 
from the midline, with the everted hemi- 
pene oriented perpendicular to the body. 
Museum acronyms follow Leviton et al. 
(1985). 


Typhlops castanotus, new species 
Figs. 1, 2 


Holotype. —CAS-SU 27940, an adult male 


from 8 km west of Pulupandan, Inampu- 


lugan Island, Negros Occidental Province, 
Philippines, collected by Angel Alcala and 
party, 23 May 1967. 

Paratypes (12).—CAS-SU 27934-39, 
27941-45, 28446, same data as holotype, 
except as follows: CAS-SU 27934—36, 27942 
collected 24 May 1967; CAS-SU 27937 col- 
lected 26 May 1967; CAS-SU 27943-44 
collected 25 May 1967; and CAS-SU 28446, 
collector and collecting date unknown. 

Additional material examined (2).—CAS 
127973, from Balabag Barrio, Borocay Is- 
land, Aklan Province, Philippines, collected 
by L. Alcala, 16 May 1970; CAS 139171, 
from Makato, Castillo Barrio, Aklan Prov- 
ince, Panay Island, Philippines, collected by 
L. Alcala and party, 3 May 1973. 

Diagnosis. —A moderate-sized member 
of the Typhlops ruficaudus group (McDow- 
ell 1974) with 28 scale rows around the an- 
terior body; dark dorsal stripe, nine or 11 
scale rows wide, the lateral-most scale row 
continuously pigmented for the entire body 
length, sharply set off from the cream-col- 
ored lateral and ventral scale rows; tail uni- 
formly dark dorsally and laterally. 


36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Description of holotype.—Total length 
(TL), 224 mm; tail length, 7 mm; midbody 
diameter (MBD), 7 mm; body diameter at 
vent, 6 mm; TL/MBD, 32; 318 middorsal 
scale rows; 307 midventral scale rows; 13 
mid-subcaudal scales; 13 mid-dorsocaudal 
scales; 28 scale rows around body anteriorly 
reducing to 24 rows posteriorly; head 
(viewed from above) tapered; in profile, 
snout rounded, projecting anteriorly above 
mouth, without transverse rostral keel; nos- 
trils lateral, near tip of snout; eyes dorso- 
lateral, visible beneath ocular shield im- 
mediately behind posterior edge of 
preocular, diameter of eye 16% of distance 
from anterior edge to tip of snout on right, 
17% on left; rostral oval, extends from up- 
per lip ventrally to a level just anterior to 
eyes on dorsum, about '2 width of head, 
widest just posterior to level of nostrils, bor- 
dered laterally by the nasals and by the pre- 
frontal posteriorly; nasals incompletely di- 
vided into pre- and postnasals, suture 
dividing each nasal originating ventrally at 
second upper labial, then extending dorsally 
and anteriorly to nostril, beyond nostril 
ending in a minute dimple-like depression 
near edge of rostral; prenasal overlaps first 
upper labial and anterior edge of second up- 
per labial; postnasals separated from con- 
tact behind rostral by prefrontal, posterior 
edge overlaps supraocular, in broad contact 
with preocular, ventral edge overlaps sec- 
ond upper labial and anterior edge of third 
upper labial; preocular inserts dorsally be- 
tween postnasal and supraocular, borders 
ocular posteriorly, inserts between postna- 
sal and third upper labial ventrally (exclud- 
ed from contact with the second upper labial 
by postnasal); ocular slightly larger than 
preocular, inserts dorsally between supra- 
ocular and parietal, posteriorly contacts two 
postoculars, inserts ventrally between third 
and fourth upper labials, the ventroanterior 
edge overlapped by third upper labial, the 
ventral edge overlapping fourth upper la- 
bial; four upper labials, first and second 
smallest (second slightly larger than the first), 


third and fourth approximately equal in size 
and more than twice as large as second; four 
middorsal scales posterior to rostral (in- 
cluding prefrontal, frontal, and interpari- 
etal) slightly larger than the succeeding body 
scales; one supraocular, one parietal, and 
two postoculars on each side, each 1.5 to 2 
times the width of a body scale. 

All head scales except rostral have basal 
glands forming a glandular line which is 
overlapped by the posteriorly projecting free 
edge of the preceding scale; glands lying 
along internasal suture between nostril and 
second upper labial expand into a striated 
organ; glands lying beneath posterior edge 
of postnasal and preocular fuse at base of 
third upper labial (under imbricate poste- 
rior edge of postnasal); basal glands of scales 
on body occupy anterior /, to 4 of the scale 
(excluding posterior free edge of scale). 

The number of scale rows around the body 
decreases in two pairs of reductions from 
28 rows anteriorly to 24 rows posteriorly. 
Immediately following the fourth enlarged 
middorsal head scale there are 28 scale rows 
around the body (at 6 mm body length, 
however, the midventral row splits produc- 
ing 29 scales before fusing again four rows 
posteriorly), reducing to 26 rows by fusion 
of first and second para-midventral scale 
rows on left at 109 mm body length and on 
right at 119 mm body length, followed by 
the second pair of reductions to 24 scale 
rows at 207 mm body length. 

Dorsal scales dark brown, densely cov- 
ered with chromatophores except for un- 
pigmented glandular area. All dorsal and 
dorsolateral scales of head, extending pos- 
teriorly to the postoculars, darkly pigment- 
ed except as follows: rostral adjacent to 
mouth under snout; on right, prenasal, first, 
second, and third upper labials, and ventral 
and posterior three-fourths of fourth upper 
labial; on left, area of prenasal anterior to 
nostril and ventral portion bordering first 
upper labial, all of first and second upper 
labials, ventral three-fourths of third and 
fourth upper labials. Behind the head shields, 


VOLUME 106, NUMBER 1 


37 


Fig. 1. 
tongue. The bar represents 5 mm. 


13 dorsal scale rows are darkly pigmented 
(on the seventh row from the middorsal, the 
second scale behind the fourth upper labial 
also has a slight amount of pigment), re- 
ducing to nine pigmented longitudinal scale 
rows at 8 mm body length; these nine dorsal 
rows pigmented for remainder of body 
length; at level of vent, the fifth row from 
the middorsal has a slight amount of pig- 
mentation, nine pigmented rows then con- 
tinuing onto tail; at about half the tail length 
the number of pigmented rows reduces to 
seven; posterior to this, dorsal scale reduc- 
tions reduce the number of pigmented rows, 
but the lateral-most pigmented row on each 
side is continuous to the spine; spine pig- 
mented except for the extreme tip. Dorsal 
stripe uniformly dark, without a lateral de- 
crease in chromatophore intensity. The un- 
pigmented ventral and ventrolateral scales 
of the head, body, and tail are cream in color 
and lack chromatophores. 

Tongue length, 3.5 mm, forked about 1.5 
mm from tip, without lateral papillae (Fig. 
1). The hemipenes are everted on both sides. 
The right hemipene subtends four scales and 


Holotype (CAS-SU 27940) of Typhlops castanotus: left lateral view of head and ventral view of 


is 3 mm in length. It is a single, soft organ. 
The medial base is essentially smooth and 
expands into a pair of smooth sacs, the larg- 
est on the anterior side, a smaller sac on the 
posterior side. At about half the length of 
the hemipene, the lateral side expands into 
a flat papillose disk covering the apicolateral 
surface. The distal tip of the disk is enfolded 
to produce a deep groove that extends onto 
the medial shaft. The sulcus spermaticus is 
a deep groove with smooth lips arising on 
the posterior base and extending along the 
posterior surface to the proximal lip of the 
disk, where it ends in an area contiguous 
with a series of grooves on the surface of 
the disk. The remainder of the hemipene is 
flounced with smooth parallel ridges (Fig. 
2). The left hemipene is similar, but the 
apicolateral disk appears to be flaccid. 
Variation. —The sex ratio in our sample 
is seven males, six females. Two juveniles 
could not be sexed. Total length ranges from 
109-253 mm, with no apparent difference 
between the sexes (Table 1). In contrast, al- 
though our samples are small, both mid- 
dorsal scale number and tail length appear 


38 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. The right hemipene of CAS-SU 27940. Lateral, posterior, and medial views are shown, respectively, 


on the left, middle, and night. 


to be sexually dimorphic. In females, the 
number of middorsal scale rows ranges from 
324-339 (¥ = 331), in males 300-327 (X = 
314), with only one male (CAS-SU 27942) 
in the range of females. Although the total 
number of middorsal scale rows is less in 
most males, males have more mid-dorso- 
caudal scales (12-14 [X = 13] vs. 11-12 [X 
= 11.5] for females) and a tail that is pro- 
portionately longer (as percent total length: 
0.026-0.036 LY = 0.031] in males, 0.020- 
0.027 [X = 0.025] in females). 

In all specimens the number of scale rows 
around the body reduces in two pairs of 
reductions from 28 rows posterior to the 
head to 24 rows anterior to the vent, al- 
though reduction patterns vary (Table 1). 
When reducing from an even to odd number 
of scale rows, reductions involve the two 
para-midventral scale rows, or the midven- 
tral and one of the adjacent rows to either 
side; reductions from an odd to even num- 
ber occur by fusion of the first and second 
rows to either the right or left of the midline, 
or fusion of the first scale row to the right 
with the first row to the left of the midline. 
Only one specimen (CAS 127973) has a re- 
duction (from 25 to 24 scale rows) that in- 
volves other scale rows, rows seven and eight 
to the right of the middorsal. The first pair 
of reductions is offset in 14 of 15 specimens 
(CAS-SU 27939 being the one exception). 
The first reduction occurs on the left side 
of the body in 13 of these specimens and 
on the right side in one specimen (CAS-SU 


27934). The reductions are nearly coinci- 
dent in this specimen, the first occurring on 
the right slightly before the second on the 
left. Displacement of the first pair of reduc- 
tions can occur by as much as 31% TL. 
Moreover, each reduction can be followed 
for a considerable length (up to 32% TL) by 
subsequent splitting and refusing of the scale 
rows. Consequently, at midbody (50% TL 
+ 10%) five specimens have 26 scale rows, 
and ten have 28-26 scale rows, depending 
on exactly where the count is made. 

The second pair of reductions, from 26 
to 24, occurs together (or nearly so) at 89- 
97% TL in 12 specimens. In three speci- 
mens the second pair of reductions 1s offset: 
the reduction from 26 to 25 occurs at 66% 
or 70% TL (followed by a region of splitting 
and refusing up to 16% TL), and the reduc- 
tion to 24 at 84% or 89% TL. 

All specimens from Inampulugan Island 
have nine pigmented dorsal scale rows with 
no dorsal to ventral reduction in chromato- 
phore intensity. Specimens from Panay 
(CAS 139171) and Borocay Island (CAS 
127973) have 11 dorsal pigmented scale 
rows, also with no dorsoventral reduction 
in chromatophore intensity in the four scale 
rows to either side of the middorsal, but 
with a slight reduction in intensity in the 
lateral-most row (row 5) on each side. 

The prefrontal separates the postnasals in 
all but one specimen (CAS-SU 27936), in 
which the postnasals touch behind the ros- 
tral and overlap (right nasal over left) under 


39 


VOLUME 106, NUMBER 1 


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40 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


it. The third upper labial contacts the post- 
nasal in all specimens. The junction of the 
small intestine and colon was examined in 
five specimens (CAS-SU 27934, 27936, 
27937, 27942, and 27944) and none have 
a rectal caecum. CAS-SU 27941 lacks retro- 
cloacal sacs. Both left and right hemipenes 
are everted in CAS-SU 27938, 27945, and 
28446. The hemipenes of CAS-SU 27938 
are similar to CAS-SU 27940, except that 
the basal area on both hemipenes appears 
to be more fully everted with the posterior 
basal sac larger and more pronounced than 
in CAS-SU 27940. CAS-SU 28446 also has 
similar hemipenes except that the terminal 
disks are not as fully everted. In CAS-SU 
27945, only the proximal shaft of each hem- 
ipene is everted, and the hemipenes lack the 
papillose disk. The tapered hemipenes have 
basal swellings and flounces as in CAS-SU 
27940. The sulcus spermaticus extends from 
the base to the tip. Some papillae can be 
seen at the tip of the left hemipene. 

Comparisons.—As a member of the 7. 
ruficaudus group, T. castanotus differs from 
Indo-Australian and Philippine typhlopids 
(excluding members of the TJ. ruficaudus 
group) by the absence of a rectal caecum 
and by fusion of the glandular lines under- 
lying the postnasal and preocular (see Mc- 
Dowell 1974, for a discussion of the char- 
acters he used to define the 7. ruficaudus 
group). 

The distinctive feature of a continuous, 
sharp-edged dark dorsal stripe contrasts with 
other members of the TJ. ruficaudus group, 
in which there is a dorsal to ventral decrease 
in chromatophore intensity in the dorsal 
stripe, and the outermost row of the dorsal 
stripe has unpigmented scales interspersed 
within it that increase in frequency poste- 
riorly resulting in a reduction from head to 
tail in the number of pigmented dorsal scale 
rows. 

Typhlops castanotus is most easily con- 
fused with T. ruficaudus and T. canlaonen- 
sis, which also are distinctly bicolored, with 
a dark dorsum and light venter separated 


by a sharp break. Besides differences noted 
above, JT. ruficaudus and T. canlaonensis 
have 30—32 scale rows behind the head (in- 
stead of 28) and dorsal pigmentation that 
ends abruptly at the level of the vent, with 
an irregular middorsal band only a few scale 
rows wide continuing posteriorly onto the 
tail (in 7. castanotus, the dorsal stripe con- 
tinues past the vent onto the tail, usually to 
the terminal spine). 

Etymology. —The specific name castano- 
tus is masculine, Latinized from the Greek 
kastanea and notos, meaning “‘brown- 
backed.” 

Distribution. —In the central Philippines, 
known only from Inampulugan Island be- 
tween Guimaras Island and Negros Island, 
Borocay Island off the northwest coast of 
Panay Island, and near the northern coast 
of Panay Island in the vicinity of Makato. 
This disjunct distribution is likely a sam- 
pling artifact, and suggests that 7. castano- 
tus may be more widely distributed on Pa- 
nay Island and surrounding islands. 

Habitat data are available for all but one 
specimen (from Inampulugan Island). Spec- 
imens from Inampulugan Island were col- 
lected in either ““hardwood forest” or “orig- 
inal hardwood forest,’ except CAS-SU 
27937, which is from a ““bamboo grove and 
hardwood forest.’’ The specimen from Bo- 
rocay Island was collected “‘along [the] edge 
of [a] coconut grove and rough clearing,” 
and the specimen from Panay Island is from 
a “forest remnant.” 

Discussion. —McDowell (1974) described 
the hemipenes of T. ruficaudus and T. kraali 
as having “‘claw-shaped (but soft) papillae 
on the distal half of the organ,’ apparently 
similar to the hemipenes of 7. castanotus. 
Although a male, the specimen of 7. rufi- 
caudus (MCZ 25594) that McDowell lists 
does not have everted hemipenes, and we 
have not seen his specimens of 7. kraali. In 
contrast, McDowell found a specimen of T. 
luzonensis (MCZ 79698) to have smooth 
hemipenes, suggesting that the 7. ruficaudus 
group 1s composed of two subgroups. Our 


VOLUME 106, NUMBER 1 


observations of MCZ 79698 indicate that 
its hemipenes are not fully everted; rather, 
they are similar to the hemipenes of CAS- 
SU 27945 suggesting that a papillose ter- 
minal disk may be hidden within the ta- 
pering shaft. 


Typhlops collaris, new species 
Rig. 3 


Holotype. —UF 55123, an adult male from 
Anuling Mt. (150 m elev.), Caramoan Mu- 
nicipality, Camarines Sur, Luzon Island, 
Philippines, collected by Walter Auffenberg 
on 29 Jul 1982 (see Auffenberg 1988, for 
additional information on this and the fol- 
lowing localities). 

Paratypes (10).—UF 52866, 23 Jul 1982, 
UF 54186 and USNM 319549 (formerly 
UF 54187), 11 Oct 1982, base camp (Barrio 
Terogo, about 2 km north of Caramoan; 
Auffenberg 1988); UF 54188, 8 Aug 1982, 
Kasini Mt. (250 m elev.); UF 54189, 22 Jul 
1982, Ilawod; UF 54192, 26 Jul 1982, UF 
55644 and USNM 319550 (formerly UF 
55645), 27 Jul 1982, UF 55646, 7 Jul 1982, 
Anuling Mt. (150 m elev.); UF 55648, 29 
Jul 1982, Anuling Mt. (200 m elev.). All 
collected by Walter Auffenberg in Cara- 
moan Municipality, Camarines Sur, Luzon 
Island, Philippines, except UF 52866, col- 
lected by Walter Auffenberg et al. 

Diagnosis.—A slender, moderate-sized 
member of the Typhlops ruficaudus group 
having more than 400 middorsal and more 
than 390 midventral scale rows, and a light 
collar of unpigmented scales behind the 
head. 

Description of holotype.—Total length 
(TL), 226 mm; tail length, 4 mm; mid-body 
diameter (MBD), 5.5 mm; body diameter 
at vent, 4.5 mm; TL/MBD, 41; 427 mid- 
dorsal scale rows; 412 midventral scale rows; 
12 mid-subcaudal scales; 11 mid-dorsocau- 
dal scales; 28 scale rows around the body 
anteriorly reducing to 26 scale rows poste- 
riorly; head (viewed from above) tapered; 
in profile, snout rounded, projecting ante- 


41 


Fig. 3. Holotype (UF 55123) of Typhlops collaris: 
left lateral view (below) and dorsal view (above) of 
head. The bar represents 5 mm. 


riorly above mouth, without transverse ros- 
tral keel; nostrils lateral, near tip of snout; 
eyes dorsolateral, visible beneath ocular 
shield, posterior edge of preocular covering 
anterior third of eye on left and anterior 
quarter of eye on right, diameter of both 
eyes 15% of distance from anterior edge to 
tip of snout; rostral oval, extending from 
upper lip ventrally to a level just anterior 
to eyes on dorsum, about '2 head width, 
widest just posterior to level of nostrils, bor- 
dered laterally by the nasals and by the pre- 
frontal posteriorly; nasals incompletely di- 
vided into pre- and postnasals, suture 
dividing each nasal originating ventrally at 
second upper labial, then extending dorsally 
and anteriorly to nostril, beyond nostril 


42 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ending in dimple-like depression near edge 
of rostral; prenasal overlaps first upper la- 
bial and anterior edge of second upper la- 
bial; postnasals in contact behind rostral and 
overlap (left postnasal over right) under 
posterior free edge of rostral, posterior edge 
overlaps prefrontal and supraocular dorsal- 
ly, laterally in broad contact with preocular, 
ventrally overlaps second upper labial and 
anterior edge of third upper labial; preocular 
inserts dorsally between postnasal and su- 
praocular, borders ocular posteriorly, and 
inserts between postnasal and third upper 
labial ventrally (excluded from contact with 
second upper labial by postnasal); ocular 
about equal in size to preocular, inserts dor- 
sally between supraocular and parietal and 
ventrally between third and fourth upper 
labials, the ventroanterior edge overlapped 
by third upper labial and the ventroposteri- 
or edge overlapping fourth upper labial; 
three postoculars, similar in size to the ad- 
jacent body scales; four upper labials, first 
and second smallest (second slightly larger 
than the first), third and fourth approxi- 
mately equal in size and more than twice as 
large as second; two middorsal scales pos- 
terior to rostral (prefrontal and frontal) 
slightly larger than the succeeding body 
scales, interparietal equal in size to the suc- 
ceeding body scales; one supraocular on each 
side, each about 1.5 times the width of a 
body scale; one parietal on each side, each 
almost twice the width of a body scale and 
oriented obliquely to the body axis. 

All head scales except the rostral have 
basal glands forming a glandular line which 
is overlapped by the posteriorly projecting 
free edge of the preceding scale; glands lying 
along internasal suture between the nostril 
and second upper labial expand into a stri- 
ated organ; glands lying under posterior edge 
of postnasal and preocular fuse at base of 
third upper labial (under imbricate poste- 
rior edge of postnasal); basal glands of scales 
on body occupy anterior 3 of the scaie (ex- 
cluding posterior free edge of scale). 


Posterior to the head shields, there are 26 
scale rows around the body, increasing to 
27 rows at 7 mm body length, and 28 rows 
at 11 mm body length; the number of scale 
rows then reduces in one pair of reductions 
from 28 to 26 rows as follows: at 13 mm 
body length the first and second para-mid- 
ventral scale rows on the right fuse (followed 
by a variable region in which this scale row 
divides and refuses up to 42 mm body 
length), and at 14 mm body length the first 
and second para-midventral scale rows on 
the left fuse; there are 26 scale rows for the 
remainder of the body length, although 
about 3 mm anterior to the vent the two 
para-midventral rows on the right fuse, then 
after three transverse scale rows divide again. 

Light pigmentation, consisting of a fine 
network of chromatophore reticulations, 
covers non-glandular portions of dorsal head 
scales (the three middorsal scales posterior 
to the rostral; supraoculars; parietals; and 
dorsal portions of the rostral, postnasals, 
preoculars, and oculars); posterior to these 
head scales, the scale inserted between the 
interparietal and parietal, and the scale be- 
tween the parietal and ocular are pigmented 
on both sides of the head; posterior to these 
pigmented scales is an unpigmented band 
four scale rows wide middorsally and three 
to four rows wide laterally; there are 15 
lightly pigmented longitudinal rows of dor- 
sal scales posterior to the collar reducing to 
11 pigmented rows before the vent due to 
gradual loss of pigmented scales in the lat- 
eral-most row (row 7) over the body length, 
and reduction in pigmentation density in 
row 6 just before the vent; posterior to the 
vent nine dorsal longitudinal scale rows are 
pigmented on the anterior two-thirds of tail, 
but unpigmented scales in the lateral-most 
row and a band of four unpigmented mid- 
dorsal scales produce a mottled appearance; 
pigmented scales end about four transverse 
scale rows anterior to the terminal spine; 
terminal spine pigmented. 

Scales in the dorsal stripe have a basal 


VOLUME 106, NUMBER 1 43 
Table 2.—Scale row reduction patterns in Typhlops collaris. See Table 1 for discussion. 
# of scale rows 

Museum # Sex TL MD MDC MV MSC Tail/TL 28-27 27-26 26-25 25-24 
UF 54186 Bp es 4 de 5 447 4k 20.0165, 4847 (0.52).L./0.52.(0.55) R.,. 0.96. R -.0:96.L 
UF 54188 Be aot Lk 444 92 3620 0.91.(0.93) R 0.92. L —_ —_ 
UF 54189 P2593 400 Il § 448" 10 0014 O55 ©.75)L.. 0.88 R — — 
UF 54192 Po sy sar tl 429° 12: 017 O93 K 0.93 L — _ 
UF 55644 E227 ©4588 11 448 10° 0.015: 0.59 (0.65) L- 0.74 (0:78) R- 0.97 0.97 
WSN 319550 F 255 434 10 427 9 0.016 0.58(0.89)L 0.93R 0.97R — 
UF 55648 F 232 434 11 426 11 O.017 0.57(0.63)L 0.66(0.74R — — 
UF 52866 M 203 422 12 408 11 0.020 — 0.04R 0.91 0.91 
USNM 319549 M 207 412 12 396 13 0.024 0.62(0.85)R 0.64(0.67)L — — 
WE 55123 Mi 226.427 TY A2I2 12 ~O0.0F8" 0:06 ©:19)R “O06 L _ _ 
UF 55646 M 210 461 13 449 12 0.019 0.54(0:68)L  0.56(00.72)R 0O0.97R — 


gland lightly covered by chromatophores; 
posterior to the basal gland, chromato- 
phores are usually concentrated into a nar- 
row dark line, with a fine network of chro- 
matophores on the remainder of the scale. 
The middorsal and adjacent scale rows are 
most darkly pigmented, with the concen- 
tration of chromatophores decreasing lat- 
erally. The ventral scales lack chromato- 
phores. To the unaided eye, the back is light 
brown and there is no sharp demarcation 
of the dorsal stripe. 

Variation. —The sex ratio in our sample 
is four males, seven females. Total length 
ranges from 203-226 mm (X = 212 mm) 
in males and 227-255 mm (X = 243 mm) 
in females (Table 2). The number of mid- 
dorsal scale rows varies from 412-461 (X 
= 430) in males and 434-460 (X = 449) in 
females; males have 11-13 (X = 12) mid- 
dorsocaudal scales and females 10-12 (¥ = 
11), and tail length (as percent total length) 
is 0.018-0.024 (X¥ = 0.020) in males and 
0.014-0.020 (X = 0.016) in females, sug- 
gesting tail length is sexually dimorphic. 

Ten of 11 specimens have 28 scale rows 
posterior to the head (although UF 54192 
has an irregular ventral scale pattern for 11 
mm posterior to the head with up to 30 scale 
rows, and UF 55648 has an irregular pattern 
in which both ventral and lateral scale rows 


fuse and split before becoming regular at 35 
mm total length). One specimen (UF 52866) 
has only 27 scale rows posterior to the head. 

Reduction patterns are variable but al- 
ways involve the two para-midventral scale 
rows, or the midventral and one of the two 
adjacent scale rows, when the reduction is 
from an even to odd number of rows. The 
first and second scale rows to either the left 
or right of the midline fuse when the re- 
duction is from an odd to even number of 
rows. The number of scale rows around the 
body reduces in two pairs of reductions from 
28 scale rows behind the head to 24 rows 
in front of the tail in only two specimens 
(Table 2). Six specimens have only one pair 
of reductions (from 28 to 26) and two spec- 
imens have one pair (28 to 26) followed by 
a single reduction (26 to 25). In UF 52866 
the single anterior reduction (27 to 26) is 
followed by a pair of reductions to 24. In 
the holotype (UF 55123) and UF 52866 the 
first reduction occurs anteriorly (6% and 4% 
TL, respectively); in all other specimens the 
first pair of reductions occurs near midbody 
or on the posterior half of the body (47- 
93% TL). The first pair of reductions is off- 
set in ten specimens, the first reduction oc- 
curring to the left of the middorsal in six of 
these and to the right in four. All reductions 
from 26 to 25, or 26 to 24 scale rows occur 


+4 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


at 91-97% TL. At midbody (50% TL + 
10%) three specimens have 28 scale rows, 
and two have 26 rows. The remaining six 
specimens have either 28-26 or 28-27 scale 
rows at midbody, depending on exactly 
where the count is made. 

The tongue of UF 54186 is 4 mm in length, 
forked at 1.5 mm, and lacks lateral papillae. 
All specimens have an unpigmented collar 
immediately behind the head, beginning 
middorsally with the fourth, fifth, or sixth 
middorsal scale row and the first or second 
scale behind the parietal and ocular shields. 
Middorsally, the collar can be from one to 
four scale rows wide; laterally, it varies in 
width within individuals from one to five 
scale rows, generally widest at the lateral- 
most extent of the dorsal stripe, and bridged 
in two specimens by pigmented scales. Pos- 
terior to the collar, the dorsal stripe is either 
17 or 15 scale rows wide. Contact between 
the postnasals posterior to the rostral varies. 
In seven specimens there is no overlap, al- 
though the postnasals touch or come close 
to touching behind the posterior edge of the 
rostral in three of these specimens; in four 
the postnasals overlap, the left postnasal 
overlapping the right in two individuals and 
the right overlapping the left in two. In all 
specimens examined, the third upper labial 
contacts the postnasal. 

A rectal caecum is absent in the two spec- 
imens (UF 54192, 55644) examined; retro- 
cloacal sacs are absent in UF 52866 and 
USNM 319549. 

Comparisons.—As a member of the T. 
ruficaudus group, T. collaris differs from 
Indo-Australian and Philippine typhlopids 
(excluding members of the J. ruficaudus 
group) by the absence of a rectal caecum 
and by fusion of the glandular lines under- 
lying the postnasal and preocular. 

Typhlops collaris is similar to many other 
populations of the JT. ruficaudus group in 
having a lightly pigmented dorsal stripe 
without a well defined break between the 
dorsal stripe and the unpigmented ventral 
scales. No other members of this group have 
the high number of middorsal transverse 


scale rows (>400) present in 7. collaris or 
a light collar of pigmentless scales posterior 
to the head. 

Etymology.—The specific name collaris 
is from the Latin co/are, in reference to the 
light collar behind the head. 

Distribution. —Known only from the 
eastern tip of the Caramoan Peninsula, Lu- 
zon Island, Philippines. 


Preliminary Artificial Key to the 
Species of Blind Snakes of the 
Philippine Islands 


The following key to Philippine scoleco- 
phidians should serve as a useful prelimi- 
nary guide to the currently recognized spe- 
cies of Ramphotyphlops and Typhlops 
known from the archipelago. We hesitate to 
comment at this time on the status of sev- 
eral problematic populations since our in- 
vestigations of the 7. ruficaudus group are 
incomplete. For this key, we follow Mc- 
Dowell (1974) in including T. /uzonensis and 
T. hypogius in T. ruber, and T. jagorii in T. 
ruficaudus. We tentatively synonymize T. 
canlaonensis with T. ruficaudus. Although 
McDowell (1974) stated that Typhlops hed- 
raeus might be conspecific with Typhlops 
ater, we recognize T. hedraeus here as a val- 
id species. 


1. Rostral without a sharp horizontal 
ridge 
— Rostral with a thickened horizontal 
ridge; tail at least twice as long as 
broad 
2. Tail short, about as long as broad. 
Glands on head confined to base of 
scales 
— Tailabout two to three times as long 
as broad. Head profusely covered 
with glands, glands not confined to 
base of scales; scales in 18 rows 
around body ...... Typhlops hedraeus 
3. Scales in 20 rows around body; in- 
ternasal suture arising from preocu- 
lar Ramphotyphlops braminus 
— Scales in 26-30 rows around ante- 


VOLUME 106, NUMBER 1 


rior third of body; internasal suture 
arising from second upper labial... 4 

4. Preocular contacts second upper la- 

bial forming a horizontal suture; 

scales in 26 or 28 rows behind head; 

15-21 dorsal rows of darkly pig- 

mented scales, occasionally only the 

centers of the scales are heavily pig- 

mented giving rise to a lineate pat- 
tern T. ruber 

— Preocular separated from second 

upper labial by postnasal, not form- 

ing horizontal suture; scales in 28 or 

30 rows behind head; 9-17 pig- 
mented dorsal scale rows 

5. Light nuchal collar present; trans- 

verse scale rows > 390; 11-17 light- 

ly pigmented dorsal scale rows ... 
Typhlops collaris 

— Light nuchal collar absent; trans- 

verse scale rows <390; 9-15 darkly 
pigmented dorsal scale rows 

6. Scale rows behind head 30-32. 11- 

15 darkly pigmented dorsal scale 

rows; scales on tail without dark pig- 

mentation except for narrow mid- 
dorsal stripe ..... Typhlops ruficaudus 

— Scale rows behind head 28. Nine or 

11 darkly pigmented (usually black) 

dorsal scale rows; tail darkly pig- 
mented above and on sides 
Typhlops castanotus 

7. Scales in 20-22 rows around body .. 
Pee! OL, Ramphotyphlops olivaceus 

— Scales in 24-28 rows around body .. 
tk Ramphotyphlops cumingii 


78 © © © © © © © © © © © © oe oe ee ee ee 


wate ele se ce! © © @ e « © « © « «8 © 


Sw ele @ «a © ¢ 8 « © © ©¢ «© 


Acknowledgments 


We are especially grateful to Walter Auf- 
fenberg and David L. Auth (Florida Mu- 
seum of Natural History), E. N. Arnold and 
A. F. Stimson (British Museum [Natural 
History]), C. J. McCoy (Carnegie Museum), 
Jose P. Rosado and Van Wallach (Museum 
of Comparative Zoology), Harold Voris and 
Hymen Marx (Field Museum of Natural 
History), and Richard Zweifel (American 


45 


Museum of Natural History), for the loan 
of specimens in their care. We would also 
like to thank George R. Zug and Ronald I. 
Crombie for commenting on early drafts, 
and Van Wallach and Donald E. Hahn for 
their careful reviews. 


Literature Cited 


Auffenberg, W. 1988. Gray’s monitor lizard. Uni- 
versity of Florida Press, Gainesville, 419 pp. 

Dowling, H. G., & J. M. Savage. 1960. A guide to 
the snake hemipenis: a survey of basic structure 
and systematic characteristics.— Zoologica 45: 
17-28. 

Reviton, AC E.R) E. Gibps, Jr., E. Heal, & C. E- 
Dawson. 1985. Standards in herpetology and 
ichthyology: Part I. Standard symbolic codes for 
institutional resource collections in herpetology 
and ichthyology.—Copeia 1985(3):802-832. 

McDowell, S. B. 1974. A catalogue of the snakes of 
New Guinea and the Solomons, with special 
reference to those in the Bernice P. Bishop Mu- 
seum, Part I. Scolecophidia.—Journal of Her- 
petology 8(1):1—57. 

Savage, J. M. 1950. Two new blind snakes (genus 
Typhlops) from the Philippine Islands.—Pro- 
ceedings of the California Zoological Club 1(10): 
49-54. 

Taylor, E.H. 1917. Snakes and lizards known from 

Negros, with descriptions of new species and 

new subspecies. — The Philippine Journal of Sci- 

ence 12D(6):353-381. 

1918. Reptiles of Sulu Archipelago.—The 
Philippine Journal of Science 13D(5):233-267. 
. 1919. New or rare Philippine reptiles.— The 
Philippine Journal of Science 14(1):105-125. 

. 1922. The snakes of the Philippine Islands. — 

Manila, Department of Agriculture and Natural 

Resources, Bureau of Science Publication 16:1— 

SPR 

Thomas, J. P. R. 1976. Systematics of the Antillean 
blind snakes of the genus Typhlops (Serpentes: 
Typhlopidae). Unpublished Ph.D. dissertation, 
Louisiana State University, Baton Rouge, 288 
pp. 


(AHW) Department of Vertebrate Zool- 
ogy, Division of Amphibians and Reptiles, 
National Museum of Natural History, 
Washington, D.C. 20560, U.S.A.; (AEL) 
Department of Herpetology, California 
Academy of Sciences, San Francisco, Cali- 
fornia 94118, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 46-50 


A NEW SPECIES OF DIMORPHIC TREE FROG, 
GENUS HYLA (AMPHIBIA: ANURA: HYLIDAE), 
FROM THE VAUPES RIVER OF COLOMBIA 


William F. Pyburn 


Abstract. —Hyla karenanneae is a new species of small color-dimorphic tree 
frog from Amazonian Colombia. Males have a bilobed vocal sac and produce 


spermatozoa with a single tail filament. 


Representatives of a new species of tree 
frog were collected at night from a small 
breeding chorus in lowland rainforest near 
the village of Timbo, Department of Vaupés, 
Colombia, in June 1973. The calls consisted 
of short, irregular, atonal notes emanating 
from frogs that were hidden among leaves 
over the water of a swamp. The frogs were 
conspicuous because of their sounds, but 
they were difficult to locate owing to the 
ventriloquistic quality of their voices. At- 
tempts to record the calls on magnetic tape 
were unsuccessful. On subsequent visits to 
this locality, as well as visits to other similar 
habitats in the Vaupés, no other choruses 
of this frog were heard and no additional 
specimens were obtained. 

Methodology note. — Measurements were 
made to the nearest 0.1 mm using Vernier 
calipers and a dissecting microscope. Slides 
of the testes were prepared following the 
method of Delahoussaye (1966). 


Hyla karenanneae, new species 
Fig. 1 


Holotype.—The University of Texas at 
Arlington Collection of Vertebrates (UTA) 
A-3770, an adult female collected by J. K. 
Salser, Jr. and the author on 7 June 1973 
near Timbo, Department of Vaupés, Co- 
lombia (01°06’N, 70°01'W, elev. 170 m). 

Paratypes. —UTA A-3768 and UTA 
A-3769, both adult males, otherwise same 
data as holotype. The holotype was in am- 


plexus with A-3769 on a leaf about one m 
above water at the time of capture. 

Diagnostic characters (based on holotype 
and paratypes). — A small (Table 1), sexually 
dimorphic member of the genus Hyla. Fe- 
male with white lateral stripe bordered be- 
low by pattern of black reticulations on white 
sides (Fig. 1); males with black reticulations 
on yellow sides. Female slightly larger than 
males and with blue-gray dorsum and blue 
ventral surfaces on limbs, hands, feet and 
digits. Males with yellow bilobed vocal sac, 
yellow-brown dorsum with irregular dark 
brown markings. Both sexes with promi- 
nently elevated nostrils and rounded snouts 
(Fig. 2); small axillary membranes; subar- 
ticular tubercles of fingers not divided, no 
orange or yellow spots on limbs, no sub- 
ocular bars or spots; no thoracic glands. Male 
with no pollical spines or nuptial excres- 
cences on fingers; spermatozoa with a single 
tail filament. 

Description of type series. —A small mem- 
ber of the genus Hy/a with flat snout, round- 
ed in dorsal and lateral views, projecting 
slightly beyond lower jaw; nostrils promi- 
nently elevated in live frog; eyes large with 
transparent palpebrum; tympanum dis- 
tinct, circular, its diameter about half 
length of eye opening; tympanic annulus 
present; a supratympanic fold impinging on 
dorsal rim of annulus. 

Body slender, about as wide as head; skin 
of dorsum and lores smooth with scattered 
small warts, ventral skin granular; anal flap 


VOLUME 106, NUMBER 1 


Fig. 1. 


small, not reaching midpoint of thigh; ax- 
illary membrane small, extending from side 
to a point about one third the distance along 
proximal part of upper arm; no calcar, no 
ulnar or tarsal fold; males with bilobed vo- 
cal sac, no pollical spines, no excrescences 
on fingers. 

Fingers (Fig. 2) long, slender with basal 
webs and expanded terminal discs about 
equal in width to diameter of tympanum 
(Table 1); fingers with prominent, rounded, 
undivided, subarticular tubercles; thenar 
tubercle an elongate oval, palmar tubercle 
large, partially divided; relative length of 
fingers: 3>4 = 2>1. 

Legs long, slender, with heel overlap of 
about four mm when legs flexed and held 
at right angles to body axis; adpression of 
leg placing heel at about midpoint of eye; 
tarsal fold absent, exposed skin of shank 
with small warts like dorsum; dermal folds 
at knee and heel; metatarsal tubercles prom- 
inent, elliptical; subarticular tubercles ellip- 
tical to conical and undivided; webbing be- 
tween first and second toes reduced (Fig. 2), 
about equaling that between third and fourth 
fingers. Extent of webbing between other toes 
varying from proximal end of antepenulti- 
mate phalanx of third toe to distal end of 
penultimate phalanx of fifth toe. 

Vomerine teeth in two short, irregular 
rows very close to midline of upper jaw, 
between obliquely elliptical choanae; vocal 


47 


— 


Fig. 2. Hyla karenanneae: (a) dorsolateral view of 
head showing rounded snout and prominent nares; (b) 
ventral view of foot; (c) ventral view of hand. Bar 
represents 1.5 mm. 


slits of male beneath postero-lateral edge of 
tongue; female lacking vocal slits and vocal 
sac; tongue of preserved frogs round with 
posterior notch. 

Color in life of males yellowish brown 
dorsally with dark brown spots and irregular 
dark brown markings forming mottled pat- 
tern over head and back; dark canthal stripe 
and dark postorbital line; no subocular bars 
or spots; sides yellow with black reticula- 
tions between axilla and groin; vocal sac 
yellow; outer surfaces of forearm and shank 
with three to four dark bars; venter pale 
yellowish cream and unmarked; lower sur- 
faces of limbs, hands, feet and digits gray; 
bones green. 

Color in life of female medium bluish gray 
dorsally, faintly mottled with dark gray; a 
cream white lateral stripe from above arm 
base to groin, bordered below by pattern of 


48 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Measurements in mm of the type series of Hyla karenanneae. 


Head 


Catalog 
no. length 


Sex SVL 
UTA A-3768 ) 
A-3769 ) 
A-3770 Q 


26.6 8.2 8.8 
28.9 8.6 a3 
30.5 


black reticulations on otherwise white sides; 
female (as in males) with dark canthal and 
postorbital line, dark bars on forearm and 
shank, no subocular bars or spots; gula and 
venter pearl white, unmarked; ventral sur- 
faces of limbs, hands, feet and digits blue; 
no yellow pigmentation in color pattern. 

Iris in both sexes bright golden bronze. 

In preservative color of sexes similar; fe- 
male with very little dorsal pigmentation, 
dorsum uniform pinkish gray; white lateral 
stripe indistinct; males without yellow col- 
or, pinkish gray with small brown spots over 
dorsum. Bars on limbs faintly visible in both 
sexes; lateral reticulations pale but visible. 
Males with brown canthal stripe and brown 
postorbital stripe. 


Discussion 


The single tail filament of the spermato- 
zoa places this species in the genus Hy/la, 
rather than in the genus Scinax (see Fou- 
quette & Delahoussaye 1977, Pombal-Ju- 
nior & Gordo 1991, Duellman & Wiens 
1992). 

Hyla karenanneae does not readily fall 
into any of the presently recognized species 
groups of small Amazonian Hy/a. It differs 
from all of these groups in dimorphic color 
pattern and vocal sac structure (see Diag- 
nostic characters). H. karenanneae further 
differs from species in these groups as fol- 
lows: 

From the Hyla parviceps group of Duell- 
man & Crump (1974), Heyer (1977, 1980), 
Duellman & Trueb (1989), it is distin- 
guished by having a rounded, rather than 
bifid, subarticular tubercle on the fourth fin- 


Width 
Nostril third 
totym- finger 
panum disc 


Tym- 

panum 
Tibia Eye diam- 
length length eter 


Width 
fourth 
toe disc 


Eye to 
nostril 


13.6 Died | 1S 3.0 8.3 1.4 2 
14.3 3e1 5 52 8.6 1.4 13 
15.8 2.6 Ley 3:3 a2 1.6 1.4 


ger; a rounded, rather than blunt, snout; no 
pale subocular spots or bars; no orange or 
yellow spots on the legs. Hyla karenanneae 
differs from the Hyla columbiana species 
group of Duellman & Trueb (1983) in hav- 
ing a rounded snout and less extensive web- 
bing between the fingers. From the Hyla 
microcephala group of Duellman & Fou- 
quette (1968), H. karenanneae differs in 
having a rounded snout and in lacking uni- 
formly yellow thighs. Hyla karenanneae dif- 
fers from members of the H. /eucophyllata 
species group of Cochran & Goin (1970) in 
having a pigmented thigh skin, in lacking 
red or orange coloration of the thigh and in 
lacking thoracic glands. From the Hyla var- 
labilis species group of Cochran & Goin 
(1970), H. karenanneae differs in lacking a 
pink or red axillary membrane. 

Other small Amazonian Hyla, of uncer- 
tain species group, differ from H. karen- 
anneae as indicated in Table 2. 


Acknowledgments 


The species is named for my daughter, 
Karen Anne Pyburn, who has found an an- 
cient Mayan city in Belize. Field work in 
Colombia was supported by the late W. 
Frank Blair and the International Biological 
Program. I am grateful to George Stewart 
for the preparation of slides that demon- 
strated sperm structure; to Wanda C. Py- 
burn and Jay K. Salser, Jr. for help with the 
field work; to Jonathan A. Campbell and 
Edmund D. Brodie, Jr. for work space, help 
in locating publications and general en- 
couragement; to Belinda Zollotuchen for 
typing the manuscript and to Darrel R. Frost 


49 


VOLUME 106, NUMBER 1 


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50 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


for permission to examine his unpublished 
manuscript dealing with hylid systematics. 

Officials of INDERENA gave permission 
to collect specimens in Colombia. 


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del Brasil.—Neotropica 8:181-192. 

1964. Dos nuevas especies de Hyla de Ron- 
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. 1966. Notas sobre Hylidae do Espirito Santo 

(Amphibia, Salientia).—Revista Brasileira de 

Biologia 26:29-37. 

——., & Ivan Sazima. 1973. Anfibios da serra do 
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Cochran, D. M., & C. J. Goin. 1970. Frogs of Co- 
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Delahoussaye, A. J. 1966. The comparative sperm 
morphology of the Louisiana Hylidae (Am- 
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Academy of Sciences 29:140-152. 

Duellman, W.E. 1972. The systematic status and life 

history of Hyla rhodopepla Ginther.—Herpe- 

tologica 28:369-375. 

1982. A new species of small yellow Hyla 
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,& M.L. Crump. 1974. Speciation in frogs of 

the Hyla parviceps group in the upper Amazon 

Basin.— University of Kansas Museum of Nat- 

ural History, Occasional Papers 23:1-40. 

, & M. J. Fouquette, Jr. 1968. Middle Amer- 

ican frogs of the Hyla microcephalia group.— 

University of Kansas Museum of Natural His- 

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—, & L. Trueb. 1983. Frogs of the Hyla co- 
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——, & 1989. Two new treefrogs of the 
Hyla parviceps group from the Amazon basin 
in southern Peru.—Herpetologica 45:1-10. 

—, & J.J. Wiens. 1992. The status of the hylid 


frog genus Ololygon and recognition of Scinax 
Wagler, 1830.—Occasional Papers of the Mu- 
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(in press). 

Fouquette, M. J., Jr., & A. J. Delahoussaye. 1977. 
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(Amphibia, Anura, Hylidae) and its bearing on 
generic status. — Journal of Herpetology 1 1:387— 
396. 

Goin, C. J. 1957. Description of two new frogs from 

Colombia.—Journal of the Washington Acad- 

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1966. A new frog of the genus Hyla from 
British Guiana. — Quarterly Journal of the Flor- 
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Haddid, C. F. B., & J. P. Pombal-Junior. 1987. Hyla 
hiemalis, nova espécie do grupo rizibilis do es- 
tado de Sao Paulo (Amphibia, Anura, Hyli- 
dae).—Revista Brasileira de Biologia 47:127-— 
2 

Heyer, W. R. 1977. Taxonomic notes on frogs from 

the Madeira and Purus Rivers, Brasil.— Papéis 

Avulsos de Zoologia 31:141-162. 

1980. The calls and taxonomic positions of 
Hyla gqiesleri and Ololygon opalina (Amphibia: 
Anura: Hylidae). — Proceedings of the Biological 
Society of Washington 93:655-661. 

Lutz, B. 1973. Brazilian species of Hy/a. — University 
of Texas Press, Austin, 260 pp. 

Pombal-Junior, J. P., Jr., & M. Gordo. 1991. Duas 
novas espécies de Hy/a da florista Atlantica no 
estado de Sao Paulo (Amphibia, Anura).— Me- 
morias do Instituto Butantan 53:139-145. 

Rivero, J. A. 1961. Salientia of Venezuela.— Bulletin 

of the Museum of Comparative Zoology 126:1- 

207. 

. 1968. Anew species of Hyla (Amphibia, Sali- 

entia) from Venezuelan Guayana.—Breviora 

307:1-S. 

1971. Tres nuevos records y una nueva es- 
pecie de anfibios de Venezuela.—Caribbean 
Journal of Science 11:1-9. 

Vigle, G. O., & D. C. I. Goberdham-Vigle. 1990. A 
new species of small colorful Hy/a from the low- 
land rainforest of Amazonian Ecuador.— Her- 
petologica 46:467-473. 


Department of Biology, Box 19498, Ar- 
lington, Texas 76019, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 51-56 


DESCRIPTION OF THE ADVERTISEMENT CALL 
AND RESOLUTION OF THE SYSTEMATIC 
STATUS OF LEPTODACTYLUS GRACILIS DELATTINI 
MULLER, 1968 (AMPHIBIA: LEPTODACTYLIDAE) 


Linnette Garcia Pérez and W. Ronald Heyer 


Abstract. —The advertisement call of Leptodactylus gracilis delattini Miller 
is described from Campeche, Santa Catarina, Brazil and compared with calls 
from five populations of L. gracilis from mainland Brazil and calls of the closely 
related L. furnarius and L. plaumanni. Based on comparison of call data and 
re-examination of the holotype, Leptodactylus gracilis delattini Muller, 1968 
is considered to be a strict junior synonym of Leptodactylus gracilis (Dumeéril 


and Bibron, 1841). 


Leptodactylus gracilis delattini was de- 
scribed by Paul Muller in 1968, based on 
three individuals collected on Ilha Cam- 
peche, Santa Catarina, Brazil (27°42’S, 
48°28'W), a tiny island located just off the 
southeastern part of the island of Santa Ca- 
tarina (Fig. 1). Leptodactylus gracilis delat- 
tini was described as a subspecies on the 
basis of general morphological similarities 
with L. gracilis, but Miller (1968) knew of 
no geographically proximate specimens of 
L. gracilis on the large island of Santa Ca- 
tarina or the mainland. Miller based his 
taxonomic decision solely on the basis of 
morphology, as he did not have any re- 
cordings of the advertisement call of the Ilha 
Campeche form. Later (Heyer 1978), L. g. 
delattini was considered a strict junior syn- 
onym of L. gracilis, again based on mor- 
phology, although the Ilha Campeche form 
was considered somewhat morphologically 
distinctive. 

The advertisement call of L. gracilis de- 
lattini and calls of geographically proximate 
populations of L. gracilis are now available. 
The purposes of this paper are to describe 
the call of L. gracilis delattini, to compare 
L. g. delattini’s call with available calls from 
other populations of L. gracilis, and to reas- 
sess the systematic position of L. g. delat- 


tini. In order to facilitate this latter aspect, 
we use the advertisement calls of L. fur- 
narius and L. plaumanni, hypothesized close 
relatives of L. gracilis (see Heyer 1978), for 
comparative purposes. 


Materials and Methods 


Recordings analyzed for this paper are: 

USNM (archives at National Museum of 
Natural History) Tape 247, cut 1, Lepto- 
dactylus gracilis delattini, recorded from 
USNM 319151, Brazil, Santa Catarina, 
Campeche (Fig. 1, locality 1), 22:10 h, 25°C 
air, by W. Ronald Heyer on 31 Dec 1991, 
using a Marantz portable tape recorder; 

ASN (Archivo Sonoro Neotropical) Tape 
AJC 27, cut 10, Leptodactylus gracilis, an 
unvouchered recording from Brazil, Santa 
Catarina, Sao José, Tubarao (Fig. 1, locality 
2), 22:00 h, 23°C air, 24°C water, by Adao 
J. Cardoso, on 13 Feb 1982, using a Uher 
reel-to-reel portable tape recorder; 

USNM Tape 11, cut 1, Leptodactylus 
gracilis, recorded from USNM 217781, 
Brazil, Santa Catarina, Santo Amaro da Im- 
peratriz (Fig. 1, locality 3), 21:10 h, 22°C 
air, by W. Ronald Heyer on 19 Nov 1979; 

ASN Tape IS 3, cut 11, Leptodactylus 
gracilis, an unvouchered recording from 


52 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


45° 


20° 


SP 


Fig. 1. 


Map of Brazilian States of Sao Paulo (SP), Parana (PR), Santa Catarina (SC), and Rio Grande do Sul 


(RS), showing recording sites analyzed. 1 = Leptodactylus gracilis delattini, 2-6 L. gracilis, triangle = L. furnarius, 
dot = L. plaumanni (see methods and materials section for further locality information). 


Brazil, Santa Catarina, Aracatuba (Fig. 1, 
locality 4), 21:00 h, 27.5°C air, 29°C water, 
by Ivan Sazima on 14 Feb 1971; 

ASN Tape AJC 40, cut 5, Leptodactylus 
gracilis, recorded from ZUEC (Universi- 
dade Estadual de Campinas Departamento 
do Zoologia) 5310, Brazil, Rio Grande do 
Sul, Bagé (Fig. 1, locality 5), 19:30 h, 24°C 
air, 25°C water, by Adao J. Cardoso on 19 


Dec 1982, using a Uher reel-to-reel portable 
tape recorder; 

ASN Tape AJC 40, cut 2, Leptodactylus 
gracilis, an unvouchered recording from 
Brazil, Rio Grande do Sul, Rio Grande (Fig. 
1, locality 6), 21:30 h, 24°C air and water, 
by Adao J. Cardoso on 18 Dec 1982, using 
a Uher reel-to-reel portable tape recorder; 

ASN Tape AJC 73, cut 5, Leptodactylus 


VOLUME 106, NUMBER 1 


furnarius, recorded from ZUEC 6360, Bra- 
zil, Sao Paulo, Sao José do Barreiro, Fazen- 
da do Bonito (Fig. 1, triangle), 18:50 h, 19°C 
air, 23°C water, by Adao J. Cardoso on 13 
Feb 1987, using a Uher reel-to-reel portable 
tape recorder; 

ASN Tape AJC 41, cut 7, Leptodactylus 
plaumanni, an unvouchered recording from 
Brazil, Santa Catarina, Nova Teut6nia (Fig. 
1, dot), 20:00 h, 23°C air, 22°C water, by 
Adao J. Cardoso on 21 Dec 1982, using a 
Uher reel-to-reel portable tape recorder. 

From 6 to 46 calls per recording were 
analyzed on a Kay Elemetrics Digital Sona- 
Graph Model 7800 for the parameters of 
call structure (harmonics, pulses), range of 
broadcast frequencies, frequency modula- 
tion, intensity modulation, and call dura- 
tion. Up to 50 second duration portions of 
calls were analyzed with a UNISCAN II 
Model 4600 for call rate and call group data. 

Specimens examined are from the Museu 
de Zoologia da Universidade de Sao Paulo 
(MZUSP) and National Museum of Natural 
History, Smithsonian Institution (USNM). 


Results 


The individual recording made for L. 
gracilis delattini did not come from Ilha 
Campeche, but was recorded immediately 
adjacent to Ilha Campeche on the island of 
Santa Catarina at the town of Campeche. 
We have compared the call voucher (USNM 
319151) with the holotype of L. gracilis de- 
lattini (MZUSP 56589). Morphologically, 
they are virtually identical, differing slightly 
in size and body coloration. The holotype 
is 38.0 mm SVL, the call voucher 37.0. The 
dorsum of the holotype is brown; that of the 
call voucher gray. The ventral surface is 
brownish-cream on the holotype and yel- 
lowish-white on the call voucher. The pre- 
vious statement by Heyer (1978:36) that the 
dorsal surface of the tibia (=shank) of the 
holotype lacks light stripes (Heyer 1978:36) 
is only partially correct. The holotype is 
generally darker than fresh, well-preserved 
specimens of L. gracilis (due to preservation 


53 


artifact), so the light stripes are not pro- 
nounced and the dorsalmost stripe found in 
L. gracilis is not apparent. The more lateral 
light-colored stripe is visible on both shanks 
of the holotype, however. In the call vouch- 
er, the more lateral light-colored shank 
stripes are very distinct, more so than the 
somewhat interrupted more dorsal shank 
stripes. 


Call of Leptodactylus gracilis delattini 


In one 50 second period analyzed, there 
are three well-defined call groups of 2.5—4.5 
sec duration, with 10-17 calls per group, 
and 4.8—7.8 seconds between call groups. 
The range of call rates within call groups is 
3.4-4.0 per sec, and call duration ranges 
from 0.04—0.05 sec. The call is pulsatile, 
with some variation between the recordings 
(calls from the first part of the recording are 
strongly pulsed and weakly pulsed in the 
second; the differences are likely due to 
placement of the microphone on the ground 
in the first part of the recording while in the 
second part the microphone was hand held). 
The call apparently lacks harmonic struc- 
ture. The call broadcast frequencies range 
from 1300-3200 Hz. Calls are frequency 
and intensity modulated; of lowest frequen- 
cy and quietest at the beginning, rapidly ris- 
ing in frequency and achieving greatest in- 
tensity at the end of the call in the range of 
2500-3200 Hz (Fig. 2). 


Comparison With Other Calls 


The call of L. gracilis delattini compared 
with calls from five other populations of L. 
gracilis and calls of the related L. furnarius 
and plaumanni indicate that none of the 
calls appear to have harmonic structure (Ta- 
ble 1). The calls from the mainland popu- 
lations of L. gracilis demonstrate but minor 
variation among themselves (Table 1), at 
the level expected for individual variation 
within populations. The call of L. gracilis 
delattini differs no more from the calls an- 
alyzed from the five populations of L. grac- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


54 


puo pure 1Op AjyI[eoo'T 

Nes WCE 4 suluulsog BUOLIS Ie CaL UC c0'0 0S 6c—0087 OLIC—-O807C luubunyyd “7 

yeom ojsuely AyIpeooT 

Les) 1 S UIOFTUs) IO juSsSqYV Soke v0'0 O8ee-OL TE 06LC-0V9T SNIADUANS "T 

9 AyeRooT 

é A UIOjTU Ls) APOM 9 €0°0 0S6c—-O08L7 OO€ I-OICI SILIDAS “TT 

¢ AypeooT 

€I-s a A suluulsog PIM toe) £0 0-00 090£—-0867¢ OLVI—O0PrI SILIDAS “"T 

p AyyesoT 

CC ts 4 pug YROM Ge ¢0'0-v0'0 OSCC—-OCIC OLOI—0£6 SILIDAS T 

wsojiun 10 ROM € Ayt[eooT 

HEAG|| Sat A suruuldog JO 3U0NS Ongac 90°0-V0'0 OOPC-OLIC O101-068 SILIDAS “TT 

puo pue 7 AypeooT 

L-p tae A suluurdog sUONIS aay el 90°0-€0'0 0887-0697  06€1-OLT1 SI1ODdS “T 

yeom | AqtpeooT 

LI-01 ar 4 puy Jo 3u01g OVv-Ve ¢0 0-00 Of Tt—-0S6C OLVI-O0EI 1UIIDIapP '8 °T 

dnoi3 sdnoda |je9 Aduonbely ul osu Ayisuoj}Uut sosjnd puoosss od (spuodes) ZO] Ul FANG) ® mit uoTlReOyNUopy] 
Jod sjj@p PoOUyop [19M (S) JOMO]S 10 jsoysiy YIM eo 91e1 [2D uoneing Aguonbady Aduonboady 
(4) 1sey Ajouies1xq [[@9 Jo uonji0g suipuq suluuisog 


"UOTIODS SPOYIIY Puv s[eLIg}e|Y Ul Ie LIep ALI[eOO] *] OINSI 0} JoJo sfoquiAs pue sioquinu 
Ayyesoy ‘Munwunnyd “7 pue sniupuanf “7 payejoi Ajasopo oy) pue ‘s7/19p48 “7 JO suoneindod say ‘1UIjJMjap siplapvss SnjAjovpodIT AOJ sigjoweIed [[VO—"] WQeL 


VOLUME 106, NUMBER 1 


eon ni a fi i 


55 


ve 
aa 


H th i a nT 


a ina 


Sem 


KILOHERTZ 


SECONDS 


Fig. 2. Advertisement call of Leptodactylus gracilis delattini. Audiospectrogram made with narrow band (45 
Hz) filter. Wave form of the first call shown on audiospectrogram, entire signal length of wave form analysis 


0.107 sec. 


ilis than the calls from the five populations 
of L. gracilis differ among themselves. In 
contrast, the differences among the record- 
ings of L. gracilis (including delattini), L. 
furnarius, and L. plaumanni are more pro- 
nounced. The calls of L. furnarius differ from 
those of gracilis (including delattini) in hav- 
ing a rapid rise in frequency rather than an 
extremely fast rise in frequency and in fre- 
quency range (L. furnarius 2080-2950 Hz, 
L. gracilis 890-3230 Hz). The L. plaumanni 
calls differ from those of L. gracilis (includ- 
ing delattini) in range of broadcast frequen- 
cies (L. plaumanni 2640-3380 Hz, L. grac- 
ilis 890-3230 Hz) and call rate per second 
(L. plaumanni 20.7—23.1, L. gracilis 1.4— 
4.0). The differences observed among the 
calls of L. gracilis (including delattini) and 
those of L. furnarius and plaumanni are at 
the level that code species information 
(Straughan 1973), whereas the call of L. 
gracilis delattini is essentially indistinguish- 
able from the other recordings of L. gracilis 
analyzed. 


Conclusion 


Based on analysis of the advertisement 
call of L. gracilis delattini, we conclude that 
L. gracilis delattini Miller, 1968 is a strict 
synonym of L. gracilis (Duméril and Bi- 


bron, 1841). We find no morphological or 
call data to suggest that the Ilha Campeche 
population of L. gracilis is distinctive at the 
subspecies level. 


Acknowledgments 


Dr. Adao J. Cardoso made recordings 
from the Archivo Sonoro Neotropical avail- 
able to us. Miriam H. Heyer participated in 
the field work at Campeche. Dr. George R. 
Zug reviewed the manuscript. LGP worked 
on the research and writing for this paper 
while a National Museum of Natural His- 
tory Research Trainee in the summer of 
1992. WRH received support for research 
on this paper from the Museu de Zoologia, 
Universidade de Sao Paulo, especially from 
Dr. P. E. Vanzolini (Director), and the 
Smithsonian Institution’s International En- 
vironmental Sciences Program and Office 
of the Director, National Museum of Nat- 
ural History funding of the Neotropical 
Lowlands Research Program. 


Literature Cited 


Heyer, W. R. 1978. Systematics of the fuscus group 
of the frog genus Leptodactylus (Amphibia, Lep- 
todactylidae).— Natural History Museum of Los 
Angeles County, Science Bulletin 29:1-85. 

Miller, P. 1968. Beitrag zur Herpetofauna der Insel 


56 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Campeche (27°42'S/48°28'’W).—Salamandra 
4:47-55. 

Straughan, I. R. 1973. Evolution of anuran mating 
calls: bioacoustical aspects. Pp. 321-327 in J. L. 
Vial, ed., Evolutionary biology of the anurans: 
contemporary research on major problems. 
University of Missouri Press, Columbia. 


(LGP) University of Puerto Rico, Ma- 
yaguez Campus, Biology Department, P.O. 


Box 5000, Mayagiiez, Puerto Rico 00681; 
(WRH) Division of Amphibians and Rep- 
tiles, Department of Vertebrate Zoology, 
National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 
20560, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 57-62 


PROCHILODUS BRITSKII, A NEW SPECIES OF 
PROCHILODONTID FISH 
(OSTARIOPHYSI: CHARACIFORMES), FROM THE 
RIO APIACA, RIO TAPAJOS SYSTEM, 
MATO GROSSO, BRAZIL 


Ricardo M. C. Castro 


Abstract. —Prochilodus britskii is described from a single locality in the rio 
Apiaca, a tributary of the rio Arinos, of the rio Tapajos system, Brazil. The 
species 1s distinguished from all other Prochilodus species by its slender caudal 
peduncle (its depth 8.8 to 9.4% of SL versus 9.7% or higher in the remaining 
species). Prochilodus britskii is a member of the assemblage of Prochilodus 
species lacking dark marks on the caudal-fin. The 6 or 7 teeth in the inner 
tooth row of each side of the lower jaw of P. britskii distinguish it from all 
other species with plain caudal fins which have 8 to 18 teeth with the exception 
of P. vimboides which has 6 to 13 teeth in that series. Prochilodus britskii differs 
from P. vimboides in having 41 to 44 pored scales in the lateral line instead of 
34 to 39. 

Resumo.—Prochilodus britskii, uma nova espécie da familia Prochilodon- 
tidae é descrita de uma unica localidade no rio Apiaca, afluente do rio Arinos, 
tributario do rio Juruena, pertencente a bacia do rio Tapajos, no Estado de 
Mato Grosso, Brasil. A espécie difere de todas as outras espécies do género 
Prochilodus pelo fato da menor altura do seu pedunculo caudal variar de 8,8 
a 9,4% do comprimento padrao contra valores iguais ou superiores a 9,7% nas 
outras espécies. Prochilodus britskii € parte do grupo de espécies do género sem 
manchas negras na nadadeira caudal. Dentro de tal grupo pode ser distinguido 
das outras espécies por possuir 6 a 7 dentes na fileira interna da metade da 
maxila inferior contra 8 a 18 nas espécies restantes, com excecao de P. vim- 
boides, que possui 6 a 13 dentes na mesma fileira de dentes. Prochilodus britskii 
difere de P. vimboides por possuir 41 a 44 escamas perfuradas na linha lateral 
contra 34 a 39. 


The prochilodontid genus Prochilodus 
Agassiz is composed of medium to large 
sized (up to ~45 cm SL) fish species, widely 
distributed through South American wa- 
ters. They are, wherever they occur, among 
the most important species in inland com- 
mercial and subsistence fisheries (see Mago- 
Leccia 1972; Roberts 1973; Vari 1983; 
Lowe-McConnell 1975, 1987). Despite their 
economic importance and widespread dis- 
tribution, the state of the Prochilodus sys- 
tematics, as well as that of Ichthyoelephas 


Posada Arango and Semaprochilodus Fow- 
ler, the two other recognized genera of the 
family, is very confused, with the only pub- 
lished revisionary study dealing with the 
Prochilodontidae (Mago-Leccia 1972) re- 
stricted to the Venezuelan species of the 
family. 

This paper is part of ongoing phylogenetic 
and revisionary studies of the Prochilodon- 
tidae aiming to, among other things, solve 
the numerous taxonomic problems in the 
family (Castro 1990). This work is based on 


58 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


material of a new species of the genus Pro- 
chilodus sent to me by Heraldo A. Britski 
(MZUSP). The specimens were collected at 
single locality, in the rio Apiaca, north of 
the city of Juara, State of Mato Grosso, in 
central Brazil. The rio Apiaca is a tributary 
of the rio Arinos, which, in turn, drains into 
the rio Juruena, a tributary of the rio Ta- 
pajOs, a southern tributary of the rio Ama- 
zonas. 

Methods and materials. —The methods of 
counting and measuring specimens in this 
paper are those outlined in Fink & Weitz- 
man (1974:1-2). Standard length (SL) and 
other body measurements were taken in mm 
and are expressed as percentages of the stan- 
dard length or, in the case of subunits of the 
head, as percentages of the bony head length. 
Ranges of counts include all specimens, with 
the values in square brackets being those of 
the holotype. Counts of total vertebrae are 
from radiographs and include the four ver- 
tebrae of the Weberian apparatus, and the 
fused PU1+U1 of the caudal skeleton 
counted as a single element. All perforated 
lateral-line scales were counted. In counts 
of fin rays, lower case Roman numerals in- 
dicate unbranched fin rays, and Arabic 
numbers indicate branched fin rays. In the 
dorsal-fin ray counts the predorsal spine is 
treated as an unbranched ray. Tooth counts 
were taken from the left side of the jaws. 
All the specimens examined for this study 
are deposited in the Museu de Zoologia da 
Universidade de Sao Paulo, Sao Paulo 
(MZUSP). 


Prochilodus britskii, new species 
Fig. 1, Table 1 


Holotype. —MZUSP 41519, 221.5 mm 
SL, Brazil, Mato Grosso, rio Apiaca, N of 
city of Juara, upriver from a fall (~ 10°36’S, 
58°04'W), collected by Convénio CEMAT/ 
ENGEVIX, 15-24 Feb 1988. 

Paratypes.—6, MZUSP 38856-61, 195.5— 
238.5 mm SL, same collection data as ho- 
lotype. 


Diagnosis. — Distinguished from all other 
species of the genus Prochilodus by having 
a more slender caudal peduncle, its depth 
equal to 8.8 to 9.4% of standard length (SL) 
instead of values equal to or higher than 
9.7% in all other species. Prochilodus brit- 
skii is a member of the assemblage of Pro- 
chilodus species lacking dark marks on the 
caudal-fin (Castro 1990). The 6 or 7 teeth 
in the inner tooth row of each side of the 
lower jaw of P. britskii distinguishes it from 
all other species with plain caudal fins which 
have 8 to 18 teeth in that series, with the 
exception of P. vimboides which has 6 to 13 
teeth in that series. Prochilodus britskii dif- 
fers from P. vimboides in having 41 to 44 
pored scales in the lateral line instead of 34 
18 whe py de 

Description. —Table 1 gives morphomet- 
rics and meristics of the holotype and para- 
types. Body relatively elongate, sub-cylin- 
drical, greatest body depth at origin of dorsal 
fin. Caudal peduncle notably narrow ver- 
tically. Dorsal profile of head slightly con- 
cave to straight. Dorsal profile of body 
slightly convex predorsally; posteroventral- 
ly slightly slanted along base of dorsal fin; 
slightly concave from posterior termination 
of dorsal fin to adipose fin and slightly con- 
cave along caudal peduncle. Dorsal surface 
of body very slightly keeled predorsally and 
rounded transversely posterior to dorsal fin. 
Ventral profile of body gently convex from 
tip of lower jaw to termination of anal fin 
base, slightly concave along caudal pedun- 
cle. Prepelvic region moderately flattened 
transversely proximate to region of pelvic- 
fin insertion. Slight mid-ventral keel present 
between pelvic-fin insertion and anus. 

Head pointed in profile. Mouth terminal. 
Snout length exceeding horizontal eye di- 
ameter; nostrils of each side close together, 
anterior circular, posterior crescent-shaped. 
Adipose eyelid present but scarcely devel- 
oped, more pronounced anteriorly, but 
leaving most of eye uncovered. First infra- 
orbital greatly enlarged, its ventral border 
together with anterior border of anteroven- 


VOLUME 106, NUMBER 1 


59 


Table 1.—Morphometrics and meristics of holotype (MZUSP 41519) and paratypes of Prochilodus britskii: 
A, range for paratypes (n = 6), MZUSP 38856-61; B, range for all the type specimens. Standard length expressed 
in mm; measurements | to 15 are percentages of standard length; 16 to 20 are percentages of bony head length. 


Character 


Standard length 

. Greatest body depth 

. Snout to dorsal-fin origin 

. Snout to pelvic-fin origin 

. Snout to anus 

. Snout to anal-fin origin 

. Posterior termination of dorsal-fin 
base to adipose-fin origin 
7. Posterior termination of dorsal-fin 

base to end of caudal peduncle 

8. Dorsal-fin base length 
9. Dorsal-fin length 

10. Anal-fin base length 

11. Pectoral-fin length 

12. Pelvic-fin length 

13. Caudal peduncle length 

14. Caudal peduncle depth 

15. Bony head length 

16. Snout length 

17. Horizontal eye diameter 

18. Postorbital length 

19. Least interorbital width 

20. Gape width 


Nn Bh WN = 


Lateral line scales 

Scale rows between dorsal-fin origin 
and lateral-line 

Scale rows between anal-fin origin 
and lateral-line 

Scale rows between pelvic-fin origin 
and lateral-line 

Scale rows around caudal peduncle 

Median predorsal scales 

Median scales between posterior 
termination dorsal-fin base and 
adipose-fin origin 

Vertebrae 


Teeth in interior “V’’-shaped tooth row 


of upper jaw, left side 


Teeth in interior “V-shaped tooth row 


of lower jaw, left side 


Holotype A B 
Morphometrics 
2245 195.5-238.5 195.5-238.5 
29.1 28.9-30.3 28.9-30.3 
47.2 45.6-47.4 45.6-47.4 
54.4 53.9-55.1 53.9-55.1 
73.5 76.5-79.0 75.5-79.0 
77.9 78.2-81.3 77.9-81.3 
29.4 27.5—30.2 27.5-30.2 
42.5 41.7-43.7 41.7-43.7 
14.6 14.3-16.2 14.3-16.2 
24.2 23.3-25.3 23.3-25.3 
10.5 8.4-10.4 8.4-10.5 
20.5 18.3-21.3 18.3—21.3 
16.7 15.1-16.5 15.1-16.7 
13.9 12.5-13.8 12.5-13.9 
9.3 8.8-9.4 8.8-9.4 
26.1 25.7-26.2 25.7—26.2 
38.4 36.4-42.6 36.4-42.6 
18.9 17.4—20.6 17.4—20.6 
44.1 41.2-45.7 41.2-45.7 
47.9 46.2-49.2 46.2-49.2 
39.8 38.7-39.9 38.7-39.9 
Meristics 

44 41-43 41-44 

7 6—7 6—7 

6 5-6 5-6 

7 6—7 6—7 
14 13-14 13-14 
14 13-14 13-14 
14 13-15 13-15 
41 40+41 40-41 
13 10-13 10-13 

7 6—7 6—7 


trally expanded second infraorbital delim- 
iting a triangular notch bordering posterior 
margin of very fleshy lips (see Roberts 1973: 
219, fig. 17 for very similar situation in 
Ichthyoelephas, and Vari 1983:33, 49, for 


phylogenetic significance of second infra- 
orbital form). Fleshy lips form oral disc when 
protruded. Functional teeth in two rows in 
each jaw; internal tooth row of upper and 
lower jaws “‘v’’-shaped. External tooth rows 


60 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. 


ye aa 
x ae. F292 
o> sci 


a = 


Prochilodus britskii, new species, holotype, MZUSP 41519, 221.5 mm SL, Brazil, Mato Grosso State, 


rio Apiaca, N of city of Juara, upriver from a fall (~10°36’S, 58°04'W), 15-24 Feb 1988. 


on both jaws follow margins of lips, with 
about 73 teeth in each half of upper jaw and 
60 in each half of lower jaw in holotype. 
*“V°’-shaped inner tooth row on upper jaw 
with 10 to 13 [13] on left side; “‘v’’-shaped 
inner tooth row on lower jaw with 6 to 7 
[7] teeth on left side. All teeth of similar size 
and spoon-shaped in frontal view (see Mago- 
Leccia 1972, fig. 4A for photograph of Pro- 
chilodus tooth) and movably implanted in 
fleshy lips. Upper and lower lips bordered 
by small globular fleshy papillae. 

Scales ctenoid. Scales along dorsal mid- 
line between posterior termination of dor- 
sal-fin and adipose-fin origin unmodified, 
without tongue-shaped membranous pro- 
cess on posterior border (see Mago-Leccia 
1972:44, 58, concerning conditions in Se- 
maprochilodus and Ichthyoelephas). Lateral 
line completely pored, with 41 to 44 (3 para- 
types with 41, 1 paratype with 42 and 2 
paratypes with 43)[44] pored scales; 6 or 7 
(2 paratypes with 6 and 4 paratypes with 
7)[7] transverse scale rows from origin of 
rayed dorsal fin to lateral-line; 6 or 7 (3 
paratypes with 6 and 3 paratypes with 7)[7] 
horizontal scale rows from the origin of pel- 
vic fin to lateral-line; 5 or 6 (5 paratypes 
with 5 and | paratype with 6)[6] horizontal 
scale rows from origin of anal fin to lateral 
line; 13 or 14 (3 paratypes with 13 and 3 
paratypes with 14)[14] median predorsal 
scales; 13 to 15 (1 paratype with 13, 4 para- 
types with 14 and 1| paratype with 15)[14] 
middorsal scales between posterior termi- 
nation of dorsal- and adipose-fin origins; 13 


or 14 (2 paratypes with 13 and 4 paratypes 
with 14)[14] horizontal scales rows around 
caudal peduncle. 

Dorsal fin preceded by small anteriorly 
bifurcated spine (see Géry 1977:367) con- 
sidered herein as an unbranched ray in fin- 
rays counts. Dorsal-fin rays 111,9 or 111,10 
(11,9 rare)[iii,10]; anal-fin rays 111,8 [i11,8]; 
pectoral-fin rays 1,13 ori,14 G,14 most com- 
mon)[i, 13]; pelvic-fin rays 1,8 [1,8]; principal 
caudal-fin rays 10/9 [10/9]. 

Rayed dorsal-fin truncate distally; pos- 
terior unbranched and anterior branched 
rays longest, subequal; fin origin nearer to 
snout tip than to caudal-fin base. Longest 
length of adipose fin about equal to or slight- 
ly larger than horizontal eye diameter. 
Origin of adipose fin on vertical crossing 
anal-fin base just anterior of its posterior 
termination. Pectoral fin distally pointed; 
when fin depressed tip reaching approxi- 
mately two-thirds of distance between fin 
origin and pelvic-fin origin. Pelvic fin fal- 
cate, its origin along vertical imaginary line 
passing through midpoint of dorsal-fin base; 
when depressed tip of fin reaching approx- 
imately four-fifths of distance to anus. Ax- 
illary scale present, pointed, its length about 
one-third of pelvic-fin length. Posterior un- 
branched and anterior branched rays of anal 
fin longest, subequal. Caudal fin forked. To- 
tal vertebrae 40 or 41 (all paratypes with 
40)[41]. 

Color in alcohol. —Background body col- 
or silvery-yellow to silvery-brown on dorsal 
half of body and head. About seven very 


VOLUME 106, NUMBER 1 


diffuse vertical bands on sides of body be- 
tween head and caudal fin; bands formed 
by chromatophore fields, without definite 
limits. Field of black or brown chromato- 
phores forming irregularly shaped spot on 
dorsal half of opercle. Dorsal fin with irreg- 
ularly distributed diffuse and barely visible 
small dark spots. Adipose dorsal with cen- 
trolateral area dusky and dorsal margin 
black. Pectoral, pelvic, and anal fins mostly 
hyaline, with distal portions somewhat 
dusky. Iris silvery-yellow with dusky dorsal 
and ventral areas. 

Color in life. —When recently collected the 
specimens showed a strong reddish-yellow 
coloration on the pelvic, anal, and caudal 
fins (Heraldo A. Britski, pers. comm.). 

Distribution. — Rio Apiaca, tributary of the 
rio Arinos, a tributary of the rio Juruena, 
upper rio Tapajos system, in the State of 
Mato Grosso, Brazil. 

Etymology. —The species name, britskii, 
is in honor of Dr. Heraldo A. Britski 
(MZUSP), who made the specimens avail- 
able to me, in recognition of his great con- 
tributions to Brazilian ichthyology. 

Remarks. —During the ongoing revision- 
ary and phylogenetic studies of Prochilodus 
the only other species of the genus found in 
the southern portion of the rio Amazonas 
basin was Prochilodus nigricans Agassiz, 
1829 (Castro 1990). Prochilodus nigricans 
is very distinct from P. britskii, most no- 
tably in belonging to the group of Prochilo- 
dus species with black marks on the caudal- 
fin. Whereas P. britskii has a remarkable 
restricted distribution for a Prochilodus spe- 
cies, being known from a single small trib- 
utary of the Amazon, P. nigricans is widely 
distributed through the huge Amazon Ba- 
sin, being probably one of the most widely 
distributed species of South American 
freshwater fishes. 


Acknowledgments 


I thank Heraldo A. Britski (MZUSP) for 
sending me the specimens which served as 
the basis of this description, along with col- 


61 


or information on the recently collected 
specimens. Osvaldo T. Oyakava provided 
assistance at MZUSP. Thomas M. Orrell 
(USNM) helped with the composition of 
Table 1. Hertz F. dos Santos and Luiz F. 
Degani (Faculdade de Filosofia, Ciéncias e 
Letras de Ribeirao Preto-Universidade de 
Sao Paulo) helped to prepare Figure 1 and 
assisted in gathering data. Richard P. Vari 
transported the specimens and provided re- 
search facilities and assistance at USNM. 
Comparative specimens used in this study 
were collected, in part, with funding from 
the I.E.S.P. Neotropical Lowland Program 
of the Smithsonian Institution, which also 
supported research at the National Museum 
of Natural History during the preparation 
of this paper. The manuscript benefitted 
from the comments and suggestions of 
Richard P. Vari and Heraldo A. Britski. The 
author received financial support from the 
Conselho Nacional de Desenvolvimento 
Cientifico e Tecnologico (CNPq) of the Bra- 
zilian Federal Government. 


Literature Cited 


Castro, R. M.C. 1990. Revisao taxonémica da fami- 
lia Prochilodontidae (Ostariophysi: Characi- 
formes). Unpublished Ph.D. thesis, Universi- 
dade de Sao Paulo, Sao Paulo, 292 pp., 43 figs. 

Fink, W. L., & S. H. Weitzman. 1974. The so-called 
cheirodontin fishes of Central America, with de- 
scriptions of two new species (Pisces: Characi- 
dae).—Smithsonian Contributions to Zoology 
172:1-46. 

Géry,J. 1977. Characoids of the world. Neptune City, 
New Jersey, TFH Publications, 672 pp. 
Lowe-McConnell, R. H. 1975. Fish communities in 
the tropical fresh waters. New York, Longman, 

337 pp. 

. 1987. Ecological studies in tropical fish com- 

munities. Cambridge, Cambridge University 

Press, 382 pp. 

Mago-Leccia, F. 1972. Consideraciones sobre la sis- 
tematica de la familia Prochilodontidae (Oste- 
ichthyes, Cypriniformes), con una sinopsis de 
las especies de la Venezuela.—Acta Biologica 
Venezuelica 8(1):35—-96. 

Roberts, T. 1973. Osteology and relationships of the 
Prochilodontidae, a South American family of 
characoid fishes.—Bulletin of the Museum of 
Comparative Zoology 145(4):213-235. 


62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Vari, R. P. 1983. Phylogenetic relationships of the Departamento de Biologia, FFCLRP- 
families Curimatidae, Prochilodontidae, An- Universidade de Sao Paulo. Avenida dos 


ostomidae and Chilodontidae (Pisces: Chara- Ban deirantes 3900, CEP 14049, Ribeiraio 
ciformes).—Smithsonian Contributions to Zo- i 
Preto, SP, Brazil. 


ology 378:1-60. 


PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 63-84 


A NEW DEVONIAN OPHIUROID 
(ECHINODERMATA: OEGOPHIURIDA) FROM 
NEW YORK STATE AND ITS BEARING ON THE 
ORIGIN OF OPHIUROID UPPER ARM PLATES 


Frederick H. C. Hotchkiss 


Abstract.—The genus Strataster Kesling and Le Vasseur, 1971, [Protasteri- 
dae] is revised, and S. maciverorum, a new species, is reported from the Panther 
Mountain Formation (Middle Devonian) near Cooperstown, New York. All 
of the species of Strataster (amended) have upper arm plates; those of S. 
maciverorum resemble the carinal plates of starfish. The prevailing view that 
upper arm plates are absent in the Oegophiurida is abandoned. The arrangement 
of the alternating ambulacral plates in Strataster is identical to the arrangement 
in echinoids, and the madreporite is located in interradius III/IV of Lovén’s 
system. These findings necessitate a revised diagnosis of the Oegophiurida. The 
new data also clarify the history of the upper arm plates of modern ophiuroids. 
These plates are serially homologous with the primary radial plates of the 
ophiuroid calycinal system. Ophiuroid arm segmentation was perfected after 
upper arm plates were brought into serial correspondence with the ambulacral 


vertebrae. 


The Devonian ophiuroids of New York 
State continue to provide important mate- 
rial for the investigation of questions at the 
higher levels of echinoderm classification. 
Ophiuroid fossils collected by Dr. Monroe 
A. McIver and Elizabeth P. McIver of 
Cooperstown, New York, and donated to 
the New York State Museum are described 
here as Strataster maciverorum, new spe- 
cies. The starfish-like aboral appearance of 
these specimens was, for a long time, an 
impediment to their proper classification. 
Eventually, the recognition of counterpart 
halves led to the discovery that the McIver 
fossils belong to the well known family Pro- 
tasteridae. The unusual aboral appearance 
is due to the presence of a distinctive series 
of carinal spines. Further review of pub- 
lished descriptions of the Protasteridae led 
to discovery of similar series of carinal spines 
in Strataster ohioensis Kesling & Le Vasseur 
and Drepanaster wrighti Kesling. The im- 
portance of this morphology to the analysis 


of the origin of ophiuroid upper arm plates 
and to the general question of homologies 
between the crinoid arm and the somaster- 
oid ray is the topic of the general part of 
this paper. The first part concerns the sys- 
tematic paleontology of the genus Strataster 
Kesling & Le Vasseur, and the description 
of S. maciverorum. 


Systematic Paleontology 


Note on methods. —The McIver fossils are 
preserved as molds and were studied from 
rubber casts, utilizing either latex or silicone 
rubber. The “disc radius’ (r) and “arm 
length” (R) were measured from the center 
of the disc. [Because many Paleozoic ophiu- 
roids do not have a well defined circular disc 
and have the general outline of a slender 
armed starfish, W. K. Spencer (1934:464, 
468) established the practice of reporting 
disc radius, rather than disc diameter, for 
measurements on Protasteridae.] The 


64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


““widih of the arm” (w) is measured at the 
edge of the disc and does not include the 
width of the splayed arm spines. 

The material has been closely examined 
for evidence of Lovén’s law by recording 
the arrangement of the ambulacral plates 
(Hotchkiss 1978). Examined from the oral 
surface, the ambulacral series on the anti- 
clockwise side of each arm is either in ad- 
vance (A) or behind (B) the series on the 
clockwise side of the arm. Data are recorded 
as if each specimen presents an oral aspect. 
The arrangement AABAB is Lovénian. If 
the madreporite is not identified, data are 
recorded only if it is possible to determine 
which column is ahead of the other for at 
least four of the five arms. If the madre- 
porite is identified on the specimen, the 
madreporic interradius is designated inter- 
radius III/IV and the labels I, II, HI, 1V, V 
are applied to the arms. Data are recorded 
for each scorable arm (even if only one arm 
can be scored). 

The material has been closely examined 
for evidence bearing on the question of ori- 
entation to substrate (cf. Schuchert 1915: 
30, 1919:6, 7; Fell 1963:429, 430; Smith & 
Jell 1990:766, 775, fig. 53) and for other 
aspects of behavior in life. 


Class Ophiuroidea Gray, 1840 
Order Oegophiurida Matsumoto, 1915 
Suborder Lysophiurina Gregory, 1897 
Family Protasteridae S. A. Miller, 1889 


Strataster Kesling & Le Vasseur, 1971 


Strataster Kesling & Le Vasseur, 1971:305, 
317.—Kesling, 1972-10" tn “part GS: 
ohioensis but not S. devonicus).—Ham- 
mann & Schmincke, 1986:61. 

Drepanaster. —Kesling, 1970:74 in part (D. 
wrighti but not D. scabrosus, D. grayae, 
and D. schohariae). [Not Drepanaster 
Whidborne, 1898] 

Eugasterella.—WHarper, 1985:361, 363 in 
part (E. ohioensis but not E. logani and 
E. devonicus). [Not Eugasterella Schu- 
chert, 1914] 


“new genus’’.— McIver & McIver, 1955:159. 


Type species. —Strataster ohioensis Kes- 
ling & Le Vasseur by original designation. 


Diagnosis (amended herein). —Protaster- 
id brittlestars with upper arm plates and 
carinal spines; upper arm plates are not in 
register with the ambulacrals, and their se- 
ries end before the arm tip. Uppermost ver- 
tical spines project at a high angle from the 
sides of the arms. Aboral outline of proxi- 
mal ambulacrals trapezoidal, accommodat- 
ing large dorsal longitudinal muscles. Disc 
radius up to 8 mm; arm length five to six 
times disc radius. Where arms become free 
of disc, their width (not including splayed 
vertical spines) equals about half the disc 
radius. 

Included species. —Strataster ohioensis 
Kesling & Le Vasseur, 1971, type species; 
Strataster wrighti (Kesling, 1970) new com- 
bination; Strataster maciverorum, new spe- 
cies; not Strataster devonicus Kesling, 1972, 
referred to Eugasterella by Harper (1985). 

Remarks. —No other known genera share 
the full set of characters in the revised di- 
agnosis. In particular, no others are known 
to have carinal spines. Hamling’s Ophiu- 
roid has upper arm plates, but it lacks ca- 
rinal and disc spines (Hotchkiss 1980). The 
splayed uppermost vertical spines and their 
contribution to the overall appearance of 
the arm is not known in any other genus. 

Harper (1985:363) regarded Strataster as 
a subjective synonym of Eugasterella 
Schuchert, 1914, which has proximal am- 
bulacrals deeply excavated for insertion of 
dorsal longitudinal muscles. I accept Har- 
per’s new combination Eugasterella devon- 
icus for Strataster devonicus, but Eugaster- 
ella does not have carinal spines and upper 
arm plates, and it is necessary to retain Stra- 
taster as a distinct genus. 

The arrangement of the ambulacral plates 
of all three species conforms with Lovén’s 
Law for echinoids (see, e.g., Melville & Dur- 
ham 1966:U221—U222). The hypothesis 
that the ophiuroid madreporite is located in 


VOLUME 106, NUMBER 1 


interradius III/IV (Hotchkiss 1978) is sup- 
ported by the new Strataster data. 

About half the specimens on a slab of rock 
are preserved with the oral side up, the oth- 
ers with the oral side down. The carinal 
spines of Strataster are similar to those of 
modern starfish such as Calliaster, Oreaster 
and Asteropsis, all of which have the usual 
eleutherozoan orientation of mouth and 
ambulacra in contact with the substrate. The 
orientation of Strataster in life is inferred 
by the presence of carinal spines, to be “‘oral 
side down.” 

As Strataster has upper arm plates, a re- 
vised diagnosis of the Oegophiurida is re- 
quired. The origin of ophiuroid upper arm 
plates is discussed below in the general part 
of this paper. 


Strataster ohioensis 
Kesling & Le Vasseur, 1971 


Strataster ohioensis Kesling & Le Vasseur, 
1971:305, 317, pls. 1-13.—Halpern, 
1972:8.—Kesling, 1972:10.— Hotchkiss, 
1978:542.— Hotchkiss, 1980:93. 

Eugasterella ohioensis. — Harper, 1985:369, 
a7 A 


Diagnosis. —Carinal spines begin near the 
periphery of the disc; there are approxi- 
mately 13 spines in the space of four am- 
bulacrals. The carinal spines are not pre- 
served in a rigid erect position; instead they 
are recumbent, usually all leaning in the 
same direction, and certainly not fused to 
the upper arm plates. The splay of the up- 
permost vertical spines nearly doubles the 
overall arm width so that it nearly equals 
the disc radius. The disc tends to have con- 
vex interradii, lacks spines, and is covered 
above and below by minute granules which 
conceal underlying plates. [Partly based on 
Kesling & Le Vasseur, pl. 7, fig. 2.] 

Types. —Type-bearing slabs in the Uni- 
versity of Michigan Museum of Paleontol- 
ogy and in the collection of Le Vasseur are 
listed by Kesling & Le Vasseur (p. 330). The 


65 


caption to their plate 4, figure 2, designates 
UMMP 58329a as the holotype. 

Lovén’s Law.—The remarkable preser- 
vation of the specimens, and the clarity and 
scale of the photographs, allow investigating 
whether Lovén’s Law applies to S. ohioen- 
sis. Three specimens have four or five arms 
that can be scored in oral view, and the 
madreporite is not identified on the pho- 
tographs: 


Paratype L-25a 
Paratype L-25f 
Paratype L-25h 


Three specimens have arms that can be 
scored in oral view and the madreporite (*) 
is identified on the photographs: 


Pe iti Ty 
Paratype UMMP 


58332a Aw be tah B 
Paratype L-25e AS. Spat wn) B 
Paratype L-251 5) ala ae Ss is <a 


These data indicate that Lovén’s Law ap- 
plies to S. ohioensis and that the madre- 
porite is located in interradius III/IV. 

Orientation and behavior. —Kesling & Le 
Vasseur (p. 338) inferred that S. ohioensis 
was a filter feeder and could hold nearly the 
full length of the arms vertical while holding 
onto the substrate with the proximal tube 
feet near the mouth. They hypothesized that 
the crest of closely spaced carinal spines may 
have been used to pull surface silt over the 
brittle star’s body and arms (negative pho- 
totactic response). About half of the speci- 
mens were buried with the oral side up and 
the others with the aboral side up (p. 330), 
and Kesling & Le Vasseur suggested that 
currents overturned the animals before 
burial (pp. 338, 339). 

Remarks. —Many authors refer to the pa- 
per by Kesling & Le Vasseur (1971) on S. 
ohioensis when discussing ophiuroid com- 
munities, population densities, arm regen- 
eration frequency, and the fossil record of 
brittlestar beds (Meyer 1984; Aronson & 
Harms 1985; Aronson 1987; Aronson & 


66 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Sues 1987, 1988; Meyer 1988; Aronson 
1989, 1992). The size range is from about 
1.2 mm disc radius to about 4 mm disc 
radius. Weathered specimens show that the 
proximal ambulacrals are trapezoidal in ab- 
oral outline, leaving large spaces for dorsal 
longitudinal muscles. The arrangement of 
the carinal spines suggests that in life they 
may have been interconnected by a web. 
Kesling & Le Vasseur report (p. 318) that 
each spine articulates by a ball-and-socket 
joint, precisely like the spine of an echinoid; 
also, where spines are missing, the tubercles 
on which they articulated are conspicuous 
because they are larger than the papillae, 
and are surrounded by a circular groove. 

Age and locality.—Early Mississippian, 
Meadville Shale. Vicinity of Cleveland, 
Cuyahoga County, Ohio. 


Strataster wrighti (Kesling, 1970), 
new combination 


Drepanaster wrighti Kesling, 1970:75, pls. 
1—2.—Hotchkiss, 1978:542.— Hotchkiss, 
1980:93. 


Diagnosis. —Carinal spines (inferred from 
sockets) begin their series midway between 
center and edge of disc, extend onto the arms, 
and number about five in the space of four 
ambulacrals (based on Kesling’s plate 2, fig- 
ure 1). The carinal spines are not erect, and 
are not fused to the upper arm plates. The 
splay of the uppermost vertical spines near- 
ly doubles the overall arm width so that it 
nearly equals the disc radius. Aboral surface 
of disc ornamented with granules and with 
few scattered spines (presence inferred from 
sockets). Oral interradii with numerous 
spines. 

Types. — The holotype and only specimen 
known is in the University of Michigan Mu- 
seum of Paleontology, UMMP 57497. 

Lovén’s Law.—The holotype has five arms 
scorable in oral view (Kesling 1970:pl. 2, 
fig. 3), and the madreporite is not discern- 
ible (p. 75). The specimen obeys Lovén’s 
Law: 


Holotype UMMP 
57497 A A B A B 


Remarks. —The disc radius of about 2.7 
mm is in the middle of the size range for S. 
ohioensis. The count of about five carinal 
spines in the space of four ambulacrals is 
distinctly different from the count of 13 in 
four in S. ohioensis. It agrees with S. ohioen- 
sis in the obviously loose attachment of its 
carinal spines. In fact, the spines are not 
found on the holotype, but their former 
presence is shown by a line of vacant sockets 
which Kesling (p. 75) described as “‘pustular 
bases for attachment of spines,” and which 
he noted extend from the aboral surface of 
the disc [Note: typographical error in his 
text says “‘oral surface’’] onto the aboral 
proximal part of the arms. The spine sockets 
confirm the presence of upper arm plates. 
The count of five carinal spines in the space 
of four ambulacrals is probably not reliably 
different from the count in the new species 
to be described next. The disc outline of the 
holotype has generally concave interradii. 
The oral interradial spines are 0.3 to 0.4 
mm long. The shape of the aboral surface 
of the proximal ambulacrals is hidden by 
upper arm plates and granules (presumably 
the shape is trapezoidal, as in S. ohioensis). 

Age and locality. — Middle Devonian, Ar- 
kona Shale. Vicinity of Arkona, Middlesex 
County, Ontario, Canada. 


Strataster maciverorum, new species 
Figs. 1—5 


““new genus and species,’” McIver & McIver, 
1955:159. Strataster, n. sp. Hotchkiss, 
1976:12. 

Protaster logani. —MclIver & McIver, 1955: 
159. [Not Protaster logani (Hall, 1868)] 


Diagnosis. —Carinal spines begin their se- 
ries at or near the center of the disc and 
number approximately 13 in the space of 
eight ambulacrals. The carinal spines are 
preserved in a rigid erect position and are 
probably fused to the upper arm plates. 


VOLUME 106, NUMBER 1 67 


Pigs a Ws a “a 4 


Fig. 1. Strataster maciverorum, new species. NYSM 13222, Holotype. Figured by McIver & McIver (1955). 
Counterpart halves. Latex pulls. Bar represents 5 mm and applies to both stereopair figures. 1.1, Aboral view. 
Carinal spines are preserved in a rigidly erect position, probably indicating fusion with the upper arm plates. 
Madreporic interradius at lower left. 1.2, Oral view. Groove spines completely close over the ambulacral groove 
on the oral surface. Madreporic interradius at lower right. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


68 


Carinal spines 


. 


13225, Paratype. Aboral view 


Fig. 2. Strataster maciverorum, new species. 2.1, NYSM 
number approximately 13 in the space of eight laterals. Uppermost vertical spine short, splayed at a high angle 


to the arm axis. Latex pull. Bar represents 2 mm. For less enlargement see Fig. 3.1. For counterpart see Fig. 


Paratype. Aboral view. Carinal spines diminish in height distally. Latex pull. Bar 


> 


S12 2225 INMISMU13223 


represents 4 mm. 


VOLUME 106, NUMBER 1 


get 


ae Ve 
¥ 


i; . ; 5 % iy? Heeb Ve Fey. at ie ‘ c: ieee uy 
Fig. 3. Strataster maciverorum, new species. NYSM 13225, Paratype. Counterpart halves. Latex pulls. 3.1, 
Aboral view. Alternating ambulacrals visible through integument near arm tip at left. Madreporic interradius 
at lower right. Bar represents 4 mm. For greater enlargement see Fig. 2.1. 3.2, Oral view. Madreporic interradius 
at lower right. Bar represents 2 mm. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


»@ 


.* a ’ 
See 


2 


—" 


Pou $4) Wisk 
Fig. 4. Strataster maciverorum, new species. NYSM 13226, Paratype. Counterpart halves. Latex pulls. Bars 
represent 2 mm. 4.1, Aboral view. Most spines are missing; spine sockets locate their former positions. The few 
spines present are rigidly erect; evidently fusion of spine to plate is not complete. 4.2, Oral view. Note groove 
spines on arm at right. 


VOLUME 106, NUMBER 1 


‘ ine ok a 
pve awe 
7 nal 


ee 


Fig. 5. Strataster maciverorum, new species. NYSM 13224, Paratype. Latex pull. Bars represent 2 mm. 
Aboral views. Disc radius (r) 7.5 mm. The disc surface is impressed onto the mouth frame. Note the plump 
cylindrical arms. Carinal spines evident, but less distinct than in smaller specimens. 


72 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Measurements of Strataster maciverorum, new species. NYSM, New York State Museum catalog 
number; r, disc radius; R, distance from center of disc to the arm tip; w, width of arm at edge of disc (does not 
include the width of the splayed arm spines); DS, rock specimen number. 


NYSM r R w 
13222 5.5 mm 34 mm 2.5 mm 
13223 8 26+ 4.5 
13224 1S 29+ 4 
13225 5 24 2:5 
13226 4.5 15+ Be 


Similar plates and spines occupy the rest of 
the aboral surface of the disc, diminish in 
size toward the interradial margins, and are 
smaller still on the oral interradii. Splayed 
uppermost vertical spines are very short, 
less than half the length of an arm segment 
and do not materially contribute to the 
overall width of the arm. Interradial outline 
of disc tends to be concave. 

Types. — There are six type-bearing pieces 
of rock, numbered DS9, DS11, DS17, 
DS17a, DS24, and DS37, collected by 
Mclver and Mclver and given to the New 
York State Museum (NYSM). DS11 and 
DS24 are counterpart halves ofa single piece 
of rock, as are DS17 and DS17a. The single 
brittlestar (Fig. 1.1, 1.2) contained in rock 
specimen DS17 and counterpart DS17a is 
now catalogued as NYSM 13222; it was il- 
lustrated by the MclIvers and is designated 
the holotype of Strataster maciverorum. 
There are four paratypes: one each in DS9 
(Fig. 2.2) and DS37 (Fig. 5.1, 5.2), now cat- 
alogued as NYSM 13223 and NYSM 13224, 
respectively, and two in DS11 and coun- 
terpart DS24. Of the latter two, one is oral 
side up and one is oral side down: NYSM 
13225 refers to the individual (Figs. 2.1, 3.1, 
3.2) preserved as an aboral impression in 
rock specimen DS11, and NYSM 13226 re- 
fers to the individual (Fig. 4.1, 4.2) pre- 
served as an aboral impression in rock spec- 
imen DS24. 

Additional material.—The asteroids and 
ophiuroids found by the MclIvers were iden- 
tified by Prof. John W. Wells of Cornell 


R:r w:r Remarks 

6:1 | Fa aa holotype; DS17 

_ 1:1.8 paratype; DS9 

— 1:1.9 paratype; DS37 

Fy 1:20 paratype; DS11, DS24 
_ 12:3 paratype; DS11, DS24 


University, who pointed out that the spec- 
imens may belong in a new genus and spe- 
cies. Labels show that the ‘“Protaster lo- 
gani” of the MclIvers’ article are contained 
in rock specimens DS1, DS2 (figured by the 
Mclvers), DS3, DS12, DS18, DS25, DS30, 
DS70, and DS75. These rocks contain im- 
pressions of the oral surface of a protasterid 
brittlestar that resembles Eugasterella lo- 
gani (Hall), the single type specimen of which 
is itself an impression of the oral surface 
without counterpart. These specimens are 
most likely oral impressions of S. maciy- 
erorum, an inference based on the obser- 
vations (1) that the specimens which Prof. 
Wells distinguished as a new generic type 
are in every case impressions of the aboral 
surface, and (2) that counterpart impres- 
sions of these last are indistinguishable from 
the oral impressions that do not have coun- 
terparts. Hence these specimens are iden- 
tified here as topotypes of S. maciverorum. 
Remains of Devonaster are present in rock 
specimens DS3, DS4, DS19 and DS24; re- 
mains of Encrinaster sp. are present in DS23. 

Etymology.—The species is named in 
memory of Dr. Monroe A. McIver and Eliz- 
abeth P. McIver of Cooperstown, New York, 
who donated these prize fossils to science. 
[According to ICZN Code Recommenda- 
tion 2la (1985), the prefixes ““Mac,” “Mc” 
or ““M”’ should be spelled “mac,” hence 
‘““maciverorum”; I thank Dr. David L. 
Pawson for pointing this out to me.] 

Measurements. —See Table 1. 

Lovén’s Law.—Two impressions of the 


VOLUME 106, NUMBER 1 


oral surface have a recognizable madrepor- 
ite together with one or more arms that are 
scorable. 


I 
Topotype DS12 A A 
Topotype DS70 eat? 


The data indicate that Lovén’s Law applies 
to S. maciverorum and that the madreporite 
is located in interradius III/IV. 

Orientation and behavior.—The carinal 
spines and plates of S. maciverorum are very 
asteroid-like, and suggest that the orienta- 
tion in life was with the oral side toward the 
substrate. Paratypes NYSM 13225 and 
13226 that are preserved on the same piece 
of rock (DS-11) but in opposite orientation 
to each other probably indicate some sort 
of turbulence at the time of burial. 

The fossils show that S. maciverorum, to- 
gether with the starfish Devonaster eucharis, 
lived on a muddy bottom with brachiopod 
and pelecypod shell life and shell rubble as 
part of the benthic epifauna (for faunal lists 
see Rickard & Zenger 1964). Crevice seek- 
ing behavior may be recorded by the ophiu- 
roid in rock specimen DS-2 which has its 
aboral surface pressed against the inside 
(concave) surface of a clam shell (the mold 
is of course convex). As noted by Parsley 
(1981:K2) for specimens of the Ordovician 
stylophoran Enopleura that seem to have 
taken refuge under brachiopod shells, the 
position of the specimen seems “‘to be de- 
liberate, rather than being the result of for- 
tuitous preservation.”’ Berry (1939) reached 
similar conclusions concerning finding nu- 
merous well preserved specimens of the 
Miocene Ophiura marylandica inside the 
shells of the large gastropod Fulgar corona- 
tum. 

The very straight arms of the holotype 
(Fig. 1.1, 1.2) may be the result of a stiff- 
ening reaction such as occurs in many living 
ophiuroids in response to being disturbed. 
This reaction is attributed by Byrne & Hen- 


a3 


dler (1988) to catch connective tissue which 
Motokawa (1988) believes was also present 
in Paleozoic echinoderms. Byrne & Hendler 
(1988) also propose that Paleozoic ophiu- 
roids with limited arm mobility but large 
podial basins may have been vagile mem- 
bers of the epifauna, walking around on the 
tips of large tube feet, a suggestion based on 
observing Ophiogeron supinus from the 
Johnson-Sea-Link submersible. 

Remarks. — All five specimens of S. mac- 
iverorum are larger than the largest of the 
100 S. ohioensis that were measured by Kes- 
ling & Le Vasseur (1971). The largest S. 
maciverorum (NYSM 13223, Fig. 2.2) is 
twice the size of the largest S. ohioensis. As 
in S. wrighti, the shape of the aboral surface 
of the proximal ambulacrals of S. maciv- 
erorum is hidden by the upper arm plates 
and granules. 

The considerable differences between S. 
maciverorum and S. ohioensis are partly 
bridged by S. wrighti. S. ohioensis has loose- 
ly articulated crowded carinal spines and no 
disc spines. S. maciverorum has rigid un- 
crowded carinal spines and many such spines 
on the disc. S. wrighti bridges the gap by 
having loosely articulated uncrowded spines 
and a few such spines on the disc (inferred 
from spine sockets on the disc and arms). 

The distinguishing marks of Strataster and 
its species are found on the aboral surface. 
The same is true of many other Protaster- 
idae, which makes it difficult to identify a 
protasterid for which only an oral view is 
available. 

What is the function of the upper arm 
plates and carinal spines? It is known (Hen- 
dler & Byrne 1987) that ophiuroid upper 
arm plates may contain structures that focus 
light onto photoreceptor cells. It is likely 
that the upper arm plates and carinal spines 
of Strataster provided it with a degree of 
protection from predation. This would be 
especially valuable if Strataster had gonads 
in the arms, as surmised for the Oegophiur- 
ida by Fell (1963) and Petr (1988), because 


74 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


the loss of an arm then also represents the 
loss of reproduction potential. 

Age and locality. —Middle Devonian (Er- 
ian Series), Panther Mountain Formation. 
Vicinity of Cooperstown, Otsego County, 
New York. From a road cut (McIver & 
Mclver 1955:photograph) on N.Y Route 28, 
1.7 miles west of Cooperstown in the Coo- 
perstown Quadrangle. Designated New York 
State Museum Paleozoic Fossil Locality 
6428. For a geologic map and treatment of 
the stratigraphy of the quadrangle, see Rick- 
ard & Zenger (1964). 


Discussion of Ophiuroid Upper arm Plates 


The present account, and a previous pa- 
per on the early growth stage represented 
by Hamling’s Ophiuroid (Hotchkiss 1980) 
are the first to demonstrate the presence of 
upper arm plates in members of the brit- 
tlestar order Oegophiurida. [Salter (1857: 
323) thought that Lapworthura miltoni 
(Salter) had a double series of upper arm 
plates, but Sollas & Sollas (1912:217) 
showed that the upper surfaces of the am- 
bulacral vertebrae merely create the im- 
pression of a double row of upper arm plates 
and that upper arm plates are absent.| The 
new data allow further discussion of the his- 
tory of the upper arm plates of modern 
ophiuroids. 

Ontogeny. —Studies on living brittlestars 
(e.g., Clark 1914) have shown that upper 
arm plates are the last plates laid down in 
an arm segment, and that arm segments are 
always added at the arm tip. The upper arm 
plates of the species of Strataster do not 
extend to the distal (younger) parts of the 
arms, and this is proof that here too they 
are the last plates of an arm region to be 
laid down. It can therefore be said that not 
only are there more plate systems in com- 
mon between ancient and modern ophiu- 
roids than has previously been recognized, 
but also that there is agreement in the se- 
quence in which the plates are laid down. 

Fewkes suggested a century ago (1887:145) 
that the absence of upper arm plates in 


Ophiohelus (moved from the Ophiacanthi- 
dae to the Ophiomyxidae by Spencer & 
Wright 1966:U89), Ophiambix (Ophiacan- 
thidae), and Astrophyton (Gorgonocephali- 
dae) could be “due to degradation, or the 
genera have not progressed through embry- 
onic stages in which dorsals appear, if, as is 
probably the case, dorsals have never ap- 
peared.”’ Mortensen (1913) showed that the 
upper arm plates in the alleged primitive 
ophiuroid Ophioteresis elegans Bell [shown 
by Mortensen to be Ophiothela tigris Ly- 
man] are being formed at the arm tip but 
are being resorbed while still of microscopic 
size. These possibilities offer a plausible ex- 
planation for the lack of upper arm plates 
in most of the previously described Oego- 
phiurida and Stenurida. 

Morphology.—Upper arm plates in the 
genus Strataster differ radically from those 
of extant ophiuroids and closely resemble 
carinals of starfish. 

Relation to the axial arm skeleton: Upper 
arm plates of extant ophiuroids are perfectly 
correlated with the arm segmentation of the 
free portion of the arm, there being one up- 
per arm plate to each segment. In Strataster 
the halves of ambulacral vertebrae are not 
in pairs simulating vertebrae, and so there 
is nO arm segmentation comparable to that 
of extant ophiuroids. Even so, the upper 
arm plates of the species of Strataster might 
be expected to show serial correlation with 
the ambulacrals, but such is not the case. In 
S. ohioensis there are approximately 13 up- 
per arm plates in the space of four ambu- 
lacrals, and in S. maciverorum approxi- 
mately 13 upper arm plates occur in the 
space of eight ambulacrals. Thus there is no 
one-to-one serial correlation between upper 
arm plates and ambulacrals in these lyso- 
phiurine oegophiurids. Nor is there one-to- 
one correlation between asteroid carinals 
and the ambulacrals of the asteroid arm. 

Spines: Upper arm plates of extant ophiu- 
roids lack carinal spines. [Note: Some ex- 
tant ophiuroids do have granules or spinules 
attached to or surrounding one or more up- 
per arm plates, but, as commented by a re- 


VOLUME 106, NUMBER 1 


viewer, “‘they are not articulated and there- 
fore not homologous with protasterid carinal 
spines’; for examples see Clark 191 1:figs. 
38, 43-49, 100, 101, 106, 126, 131, 134.] 
In contrast, carinal spines are a highly 
conspicuous feature in S. ohioensis and S. 
maciverorum. The carinal spines of S. mac- 
iverorum, particularly, are very “‘asteroid- 
like.” These spines almost certainly indicate 
that these animals lived with their oral side 
down as in modern sea stars. 

Relation to the aboral disc skeleton: Em- 
bryologists (e.g., Murakami 1941:73) have 
found that upper arm plates of Ophiurida 
begin their series with the first arm segment 
(the third if elements of the mouth frame 
are counted). As the animal grows, the up- 
per arm plates associated with those arm 
segments that become included in the disc 
region are displaced and crowded together 
at the place where the arm becomes free of 
the disc (Berry 1934:73, fig. 9). In S. maciy- 
erorum and S. wrighti, however, the upper 
arm plates begin their series well within the 
disc region, an arrangement exactly com- 
parable to that seen in asteroids. 

Homologies.—Examination of the ho- 
mologies of upper arm plates necessitate a 
brief excursion into the homologies of other 
plates as well. A summary of the revised 
homologies is given in Tables 2 and 3. 

Homologies of brachials and upper arm 
plates: Carpenter (1887:309, footnote) con- 
cluded that the brachials of a crinoid are in 
a general way represented in the ophiuroid 
by the upper arm plates. Sladen (1884:30) 
shared this view and applied the term “‘bra- 
chial’’ to the median dorsal line of plates of 
the starfish ray as seen in juvenile Zoroaster 
fulgens. Fell (1963:414), however, conclud- 
ed that the carinal plates of asteroids, and 
the dorsal arm plates of ophiuroids, are late 
structures, not homologous with any bra- 
chial ossicles in crinoids. According to Fell 
(pp. 419, 420) the brachial ossicles of cri- 
noids are homologous with the ambulacral 
ossicles of asteroids and ophiuroids, not with 
dorsal arm plates or carinals. The discovery 
of upper arm plates in Strataster, Hamling’s 


iS 


ophiuroid, and Rhopalocoma \ed Hotchkiss 
(1974) to accept the views of Carpenter and 
Sladen over that of Fell. It is shown below, 
however, that Fell’s homology of brachials 
with ambulacrals is correct, and that upper 
arm plates are not represented in crinoids. 

Homologies of radial shields and upper 
arm plates: Fewkes (1887:130) believed that 
ophiuroid radial shields are homologous 
with the first paired brachials of crinoids. 
This view was discredited by Carpenter 
(1887:308-—309) based on study of crinoids. 

Fell (1963:420) argued that radial shields 
are a late phylogenetic development and are 
not part of the calyx: ““Their development 
in Euryalina shows that they arise from an 
adradial series of platelets equal in number 
and position to those segments of the arm 
which have been incorporated into the disc 
below. Thus they are homologous with the 
dorsal arm-plates of the rest of the radius; 
as indeed their arrangement in many ophiu- 
roids suggests, for the basal dorsal arm-plates 
are often split into two portions, right and 
left.”’ This interpretation, however, does not 
accord with the observations of Lyman 
(1882) concerning the development of ra- 
dial shields or with the fate of the first few 
upper arm plates in genera such as Ophier- 
nus. 

Lyman (1882:157) examined minute 
young of Hemipholis elongata [as H. cor- 
difera| and concluded that “‘radial shields, 
so nearly universal among ophiurans, are 
not special plates, but entirely homologous 
with other disc scales, and by no means the 
first to appear.’ He showed (p. 271, pl. 44, 
figs. 10, 11, /) that in Astrogomphus vallatus 
the radial shields ‘‘are made up of several 
overlapping pieces soldered together.’’ He 
described (p. 167) the genus Ophiocoma as 
having radial shields that “‘are continued 
inward [toward the center of the disc] by a 
broad stripe of large, strongly overlapping 
scales, a feature nowhere so developed as in 
this genus.”” His descriptions and figures 
show that the platelets referred to by Fell 
greatly exceed the number of arm segments 
in the disc, and that the accretion of platelets 


76 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 2.—Revised homologies of plates of the aboral surface (tissues derived at metamorphosis from the right 


side of the larva). 


Asteroids 


Ophiuroids 


Crinoids 


Primary circlet: 


centrale centrale 


primary interradials [cf. Zoro- 
aster] 
primary radials 
carinal plates [=serial homo- 
logues of primary radials] 
dials] 


Extraxial skeleton: 


the aboral skeleton outside 
the circlet of primary radi- 
als [superomarginals are dif- 
ferentiated members of the 
aboral extraxial skeleton] 


primary interradials 


primary radials 
upper arm plates [=serial 
homologues of primary ra- 


the aboral skeleton outside 
the circlet of primary radi- 
als [the radial shields of 
advanced ophiuroids are 
modified aboral disc 
plates] 


terminal stem plate (at- 
tachment disc) of comat- 
ulid larval stalk 

? topmost stem penta- 
meres [cf. Aethocrinus] 

? infrabasals 

—(absent) 


the aboral skeleton [=bas- 
als] outside the circlet of 
infrabasals 


is toward the center of the disc (whereas 
upper arm plates develop in succession dis- 
tally). [I thank a reviewer for the following 
additional information: “It is incorrect to 
infer from Lyman’s illustrations that radial 
shields grow by adding platelets proximally. 
Each of the plates arises from a single spic- 
ule and growth proceeds by the enlargement 
of an ossicle—not by the incorporation of 
additional ossicles. However, this does not 
invalidate the suggestion that they are disc 
scales.’’] 

In Ophiernus, which has been very well 
described by Madsen (1977), the upper arm 
plates that correspond to those segments of 
the arm which have been incorporated into 
the disc certainly do not go into the making 
of radial shields. Instead, they accumulate 
at the edge of the disc. As a consequence 
they are no longer associated with their 
proper arm segments, and so we find ex- 
amples of the first six proximal arm plates 
partially resorbed and compressed into the 
space of just three arm segments. Obser- 
vations by A. M. Clark (1974:443 [fig. 1], 
p.462 [fig. 10a]) show that the upper arm 


plates of Amphilimna cribriformis and 
Ophionephthys lowelli do not contribute to 
the making of radial shields; in the latter 
instance, upper arm plates that are over- 
grown by the disc are found to be divided 
by “erosion of the median part’ into left 
and right halves that are presumably on their 
way to being completely resorbed. 

The upper arm plates of all three species 
of Strataster, Hamling’s Ophiuroid, and 
Rhopalacoma, are entire, not split in two to 
form a double row. Upper arm plates arise 
ontogenetically from a single center of cal- 
cification (e.g., Murakami 1940:32). The two 
(and sometimes more) pieces of upper arm 
plates found in some extant ophiuroids must 
be the result of fragmentation of an initially 
entire plate (Lyman 1882:20), or are evi- 
dence for the existence of additional plates 
lateral to the true upper arm plates. In fact, 
both median and lateral upper arm plates 
were demonstrated in ‘“‘Ophioteresis ele- 
gans’”’ by Mortensen (1913:8). Similar 
‘“‘fragmented”’ plates were described in 
Ophiomyxa, Ophiobyrsa and Ophiogeron by 
Byrne & Hendler (1988). Thus, the sup- 


VOLUME 106, NUMBER 1 


77 


Table 3.—Revised homologies of plates of the oral surface (tissues derived at metamorphosis from the left 


side of the larva). 


Asteroids 


Axial skeleton: 


ambulacral plates 
[evolved from stro- 
matocystitid biserial 
flooring plates] 

terminal (ocular) 
plates 


Adaxial skeleton: 


adambulacral ossicles 
[evolved from stro- 
matocystitid pri- 
mary cover plates] 

adaxial virgalia 
[evolved from stro- 
matocystitid cover 
plate series] [=aster- 


Ophiuroids 


ambulacral plates + sublateral 
plates [sublateral plates are 
part of the ambulacral series 
in Rhopalocoma] 

terminal (ocular) plates 


lateral arm plates [mouth angle 
plates of jaw] 


adaxial virgalia [pinnate 3rd 
virgal reported in Trichaster, 
Asteronyx and Astrophyton 
by Fell (1963)] 


Crinoids 


brachial plates [radials = Ist 
plates of brachial series] 
[pinnules = heterotomous 
arm branching] 

— (absent) 


— (absent) 


— (absent) 


opseid growth gradi- 
ent metapinnules] 
Admarginal skeleton: 


admarginal virgalia — (unknown) 


[=poraniid growth 

gradient metapin- 

nules] [=develop- 

mental homologues 

of inferomarginals] 
Marginal skeleton: 


inferomarginal ossicles 
[homologous with coma] 
the stromatocystitid 
marginal frame] 


posed homology between radial shields and 
upper arm plates is not supported by either 
embryology or the new fossil evidence. 
Homologies of the calycinal plates of 
ophiuroids and crinoids: Carpenter (1884), 
Sladen (1884), Fell (1963), and Hotchkiss 
(1974, 1980) believed that the primary ra- 
dial plates of ophiuroids and asteroids are 
homologous with the radial plates of cri- 
noids. This turns out to be a major source 
of misdirection in inferring correct echi- 
noderm plate homologies. If in crinoids the 
brachials are serially homologous with the 
radial plate, then where are the serial ho- 
_mologues of the primary radial plate in the 
Asterozoa? For Carpenter (1887:309, foot- 


[cf. marginalia in Rhopalo- 


— (absent) 


—(absent) 


note) the answer lay with the ophiuroid up- 
per arm plates. For Fell (p. 419) it lay with 
*“‘asterozoan”’ ambulacral plates. To relocate 
“‘asterozoan”’ ambulacral plates from the 
aboral surface (where the calycinal system 
develops) to the oral surface (where am- 
bulacrals are part of the jaw apparatus), Fell 
postulated a “‘dislocation of the main radial 
growth gradient at the junction of the radial 
calycinal plate and the first brachial ossicle”’ 
(p. 382). This is Fell’s mechanism for the 
evolution of the Asterozoa from a crinoid- 
like ancestor. Although Fell (p. 414) states 
that evidence of a dislocation is directly ob- 
servable in the ontogeny of Recent Astero- 
zoa, this is contradicted by the fact that pri- 


78 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


mary radials develop in tissues derived from 
the right side of the bilateral larva whereas 
the ambulacrals develop in tissues derived 
from the left side (Sladen 1884:37, Murak- 
ami 1937:137, figs. 3—S). 

As pointed out recently by Broadhead 
(1988:255), Hyman (1955:57, 85, 698) broke 
away from this misdirection. In Antedon, 
rudiments of the radial plates are not pres- 
ent until the end of the cystidean stage. Ac- 
cording to Hyman, the radial plates are “not 
true thecal or calycinal plates but the first 
plates of the brachial series.”’ This interpre- 
tation, which is accepted by Broadhead, 
means that the primary radial plates of 
ophiuroids cannot represent the radials of 
crinoids. 

The idea that upper arm plates may be 
serially homologous with ophiuroid pri- 
mary radials does not conflict with the idea 
that brachials are serially homologous with 
crinoid radial plates. The interpretation by 
Philip and Strimple (1971) of the Lower Or- 
dovician archaic inadunate crinoid Aetho- 
crinus leads me to consider that the primary 
radials of Hamling’s ophiuroid (Hotchkiss 
1980) may be equivalent to the infrabasals 
of Aethocrinus. The primary interradials of 
developing Asteroidea lie inside the circlet 
of primary radials (Ludwig 1905:pl. 32, fig. 
190, Blake 1990:347, 351); the correspond- 
ing circlet of ossicles in Aethocrinus are the 
topmost stem pentameres that in fact con- 
tribute to the theca. The centrale of ophiu- 
roid early growth stages therefore seems to 
represent the terminal stem plate (attach- 
ment disc) found in the larval stalk of co- 
matulid crinoids (a homology previously 
advocated by Carpenter 1878:374 and by 
Sladen 1884:35). 

Homologies of the crinoid arm and the 
somasteroid ray: Fell (1963) saw in the 
structure of the somasteroid ray a pattern 
of skeletal growth gradients found elsewhere 
only among crinoids. Cover plates along the 
somasteroid metapinnules seemed to con- 
firm the comparison with crinoid pinnules. 
He therefore considered somasteroid am- 


bulacrals and virgalia homologous with cri- 
noid brachials and pinnulars. Because there 
is no plate series in the crinoid arm that 
corresponds with ophiuroid upper arm 
plates, Fell interpreted the absence of upper 
arm plates in somasteroids and Paleozoic 
ophiuroids (Stenurida and Oegophiurida) as 
confirming evidence of his phylogenetic 
theory. But as shown here, upper arm plates 
were present in Strataster. Upper arm plates 
are not yet known from somasteroids, but 
it can now be supposed that they may have 
been present in at least the early growth 
stages. Thus the crinoid arm and the so- 
masteroid ray do not appear to be compa- 
rable structures, and the comparisons and 
the homologies proposed by Fell are doubt- 
ful. 

The ancestry of sea stars is now sought 
among the edrioasteroids. Detailed analysis 
of the marginal frame and the intermediate 
skeletons of sea stars (Hotchkiss 1974, 
Hotchkiss & Clark 1976) suggests homol- 
ogizing the marginal frame of Archegonaster 
with the marginal frame of stromatocystitid 
edrioasteroids (Termier & Termier 1969, 
Smith & Jell 1990). According to Paul and 
Smith (1984:468) somasteroid ambulacrals 
are homologous with the primary ambula- 
cral flooring plates of early Cambrian echi- 
noderms; somasteroid virgalia are thought 
to be derived from stromatocystitid (Cam- 
braster) cover plate series (p. 469). [The ob- 
servation that crinoid pinnules arise from 
heterotomous arm branching (Paul & Smith 
1984:466) whereas virgalia derive from ed- 
rioasteroid cover plate series is another rea- 
son that the crinoid arm should not be con- 
sidered homologous with the somasteroid 
ray.] A search should be made for anteced- 
ents of upper arm plates in somasteroids 
and in stromatocystitid edrioasteroids. 

Serial homology of primary radials and 
upper arm plates: That the upper arm plates 
of S. maciverorum and S. wrighti begin their 
series well within the disc region confirms 
the conclusion stated by Lyman (1882:270) 
that there is no distinction between the up- 


VOLUME 106, NUMBER 1 


per surface of the arms and that of the disc. 
Important to the present context, it shows 
that it is proper to compare upper arm plates 
with disc plates. Judging from S. maciver- 
orum it is very likely that the plates that 
bear the first carinal spines are in fact the 
primary radial plates of the rosette (cf. Lud- 
wig 1905:pl. 32, fig. 190). Hamling’s ophiu- 
roid and Rhopalacoma pyrotechnica (Salter) 
seem to answer unequivocally which disc 
plates hold the long sought homology. In 
both there is a direct serial relation and a 
virtually identical morphology between the 
upper arm plates and the primary radials. 
Upper arm plates are evidently fashioned 
after the plan prescribed by the genetic in- 
structions for fashioning of primary radials, 
and therefore in every sense are serial ho- 
mologues of the latter (cf. Hubbs 1944:293). 

Antiquity of upper arm plates. —Wherever 
primary radial plates occur there is the po- 
tential for them to be serially repeated along 
each arm. Upper arm plates can therefore 
be as ancient as primary radials. The evi- 
dence from Hamling’s Ophiuroid leads to 
the inference that primary radials are an 
inheritance from the somasteroid stem group 
ancestral to both asteroids and ophiuroids. 
It follows that upper arm plates were at least 
potentially, and perhaps were in fact, also 
present in stromatocystitid edrioasteroids. 

The strong similarities between the upper 
arm plates of Strataster (also of Rhopala- 
coma) and the carinals of starfish can be 
stated as follows: Although the upper arm 
plates of modern ophiuroids lack the com- 
mon asteroid character of bearing a carinal 
series of spines, and although asteroids lack 
the modern ophiuroid character of serial 
correspondence between upper arm plates 
and ambulacrals, these Paleozoic ophiu- 
roids bridge both of these gaps. An obvious 
suggestion is that upper arm plates predate 
the divergence of the asteroid and ophiuroid 
lineages (the “‘asteroid/cryptosyringid di- 
vergence” of Smith 1988:88). 

There is evidence for the antiquity of 
upper arm plates at every stage in the evo- 


79 


lution of the ophiuroid lineage. That the 
stenurid Rhopalacoma (perhaps also Bdel- 
lacoma) has upper arm plates could mean 
that upper arm plates predate the stenurid- 
oegophiurid divergence. That the protas- 
terids Strataster and Hamling’s Ophiuroid 
have upper arm plates could mean that up- 
per arm plates predate the lysophiurine- 
zeugophiurine divergence. That certain 
phrynophiurids have upper arm plates could 
mean that upper arm plates predate the 
phrynophiurid-ophiurid divergence. That 
the Silurian Argentinaster bodenbenderi 
Ruedemann has typical upper arm plates 
(personal observation) may mean that up- 
per arm plates date from the very beginning 
of at least the order Ophiurida. 

Such statements are in complete disagree- 
ment with the conclusion of Ubaghs (1953: 
789) and Fell (1963:414) that upper arm 
plates developed late in ophiuroid phylog- 
eny. They also depart from the conclusion 
of Sollas & Sollas (1912:218) that the late 
appearance of upper arm plates in ontogeny 
recapitulates a late phylogenetic history. In- 
stead it appears that the developmental 
pathway for upper arm plates was present 
in the stem group ancestral to both asteroids 
and ophiuroids, and that the presence or 
absence of upper arm plates as a character 
state in Paleozoic ophiuroids was deter- 
mined by genes that regulated expression of 
the pathway. 

History of upper arm plates. —The history 
of ophiuroid upper arm plates therefore ap- 
pears to be as follows. In the stem group 
ancestral to both asteroids and ophiuroids 
they were not in serial correspondence with 
the ambulacral series. This is quite under- 
standable considering the plump arms of 
early asteroids and ophiuroids. This char- 
acter state was carried over into stenurids 
and oegophiurids, where upper arm plates 
still lack serial correlation with the ambu- 
lacral skeleton. In the line of descent in which 
the ambulacrals of the two sides of the arms 
are staggered (Lysophiurina) perfect arm 
segmentation was an impossibility, al- 


80 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


though remarkable arm flexibility was 
achieved. In the zeugophiurine oegophi- 
urids, the ambulacrals of the two sides of 
the arm are in register, and arm segmen- 
tation was perfected when upper arm plates 
were brought into serial correspondence with 
the vertebrae. This condition is found among 
those phrynophiurids that express upper arm 
plates, and is well known among the 
Ophiurida. 

This history requires that upper arm plates 
were present and exposed to natural selec- 
tion in the somasteroid stem group ancestral 
to both asteroids and ophiuroids. Accord- 
ingly it is also necessary to explain the ab- 
sence of upper arm plates among most of 
the previously described Paleozoic Oego- 
phiurida and Stenurida. As in the case of 
explaining the lack of a primary rosette in 
post-juvenile Oegophiurida and Ophio- 
myxidae (Fell 1963:419), absence of upper 
arm plates in these fossils is best explained 
by resorption. Resorption of calcareous 
matter in ophiuroid growth series was noted 
by Clark (1911:3), and was documented for 
upper arm plates by Mortensen (1913). 


Oegophiurida Rediagnosed 


Matsumoto, the author of this order 
(1915:45, proposed as a subclass), included 
in his diagnosis the statement that upper 
arm plates are absent. Later authors have 
agreed with this statement (e.g., Spencer 
1925:280, Fell 1963:407, Kesling 1970:74). 
On the basis of data provided by the species 
of Strataster and the specimens of Ham- 
ling’s Ophiuroid, the diagnosis of the order 
must be amended, for these otherwise com- 
pletely typical protasterid lysophiurine 
oegophiurids possess upper arm plates and 
carinal spines. As reported elsewhere 
(Hotchkiss 1980), Hamling’s Ophiuroid 
shows that early growth stages of oegophi- 
urids carry a centrale and primary radials. 
Accordingly, earlier generalizations based 
on presumed absence of these plates from 
oegophiurids must likewise be abandoned 


(cf. Sturtz 1899:181-182; Spencer 1914:34— 
35, 1925:242-—243; Philip 1965). Further- 
more, it has only recently been discovered 
(Hotchkiss 1978) that Lovén’s Law applies 
to Oegophiurida with alternating ambula- 
cral plates (Protasteridae and Encrinasteri- 
dae), and data are still being accumulated 
(Harper & Morris 1978:157, Harper 1985: 
367, herein). It therefore seems appropriate 
to conclude with a brief revised diagnosis 
of the order. 

Oegophiurida are distinguished from 
Phrynophiurida and Ophiurida by a list of 
negative characteristics: ventral arm plates 
absent, bursae absent, genital plates and 
scales absent, radial shields absent, oral 
shields absent, adoral shields absent. Like 
phrynophiurids and ophiurids, some oego- 
phiurids are now known to have upper arm 
plates, and early growth stages have been 
shown to have a centrale and primary ra- 
dials. A more detailed report has explained 
that the extant Ophiocanops is not a living 
example of the Oegophiurida and that it is 
properly classified in the Phrynophiurida 
(Hotchkiss 1977). 

Oegophiurida are distinguished from 
Stenurida by podial basins which are en- 
tirely on the distal portion of an ambulacral, 
by the absence of sublaterals, and by pres- 
ence of no more than two fused ambulacral 
elements in the mouth frame. There are a 
few taxa classified as oegophiurids that have 
certain stenurid features, but their overall 
facies is that of a typical oegophiurid (e.g., 
Protaster piltonensis Spencer, and Bunden- 
bachia benecki Sturtz). 

The suborder Lysophiurina obeys Lo- 
vén’s Law (heretofore an echinoid trait). In 
the Lysophiurina the halves of ambulacral 
vertebrae are in offset series, whereas in the 
Zeugophiurina they are in register. Others 
have remarked before that the two lines of 
descent represented by these suborders ef- 
fectively make the order polyphyletic 
(Ubaghs 1953:818). Provided that the lim- 
itations of the present classification are un- 
derstood, there is no need to introduce any 


VOLUME 106, NUMBER 1 


broad changes into the classification until 
more work has been done on undescribed 
material in various existing collections. 


Acknowledgments 


Research was supported by a N.D.E.A. 
Title [IV Graduate Fellowship, NSF Grad- 
uate Traineeship, Connecticut State Schol- 
arship Grant, and Smithsonian Institution 
Fellowship in Systematic and Evolutionary 
Biology. Dr. and Mrs. Monroe A. Mclver, 
to whom the new species is dedicated in 
memorium, were kind hosts and were gen- 
erous in donating their collection of Astero- 
zoa to the New York State Museum. I thank 
Dr. John W. Wells of Cornell University for 
informing me of the existence of this im- 
portant MclIver collection, and for showing 
me his latex casts of this material. I thank 
Dr. Bruce M. Bell and the officials of the 
New York State Museum for the loan of the 
Mclver collection. I thank Dr. David L. 
Pawson of the National Museum of Natural 
History, Smithsonian Institution, for his 
constant help with literature and specimens 
Over many years. I thank Dr. Adrian Hog- 
ben and Dr. D. L. Pawson for help in lo- 
cating the proper 1840 citation for Gray as 
the author of the Class Ophiuroidea [the 
citation given by Spencer & Wright (1966), 
although widely used, is incorrect]. I thank 
R. B. Aronson, D. B. Blake, F. J. Fell, G. 
Hendler, S. Inmora, Y. Ishida, M. Jangoux, 
J. M. Lawrence, V. Petr, and A. B. Smith 
for instructive correspondence and their 
help. I thank Alan Doherty for preparing 
silicone rubber casts. The conclusions on 
homologies presented here differ from some 
of those that I expressed in 1974, primarily 
as a result of incorporating some of the views 
of Paul and Smith (1984) and Broadhead 
(1988). I thank the reviewers for suggesting 
numerous improvements to the manu- 
script; technical points raised by the re- 
viewers have been incorporated and cited 
in the text. I thank my wife Anita Hotchkiss 
for her companionship on field trips and her 


81 


encouragement and help in finishing this re- 
search after a fifteen year hiatus. 


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26 Sherry Road, Harvard, Massachusetts 
01451, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 85-91 


ERY THROSQUILLOIDEA, A NEW SUPERFAMILY, 
AND TETRASQUILLIDAE, A NEW FAMILY OF 
STOMATOPOD CRUSTACEANS 


Raymond B. Manning and David K. Camp 


Abstract.—The superfamily Erythrosquilloidea is erected for the family 
Erythrosquillidae, which contains one genus and species, Erythrosquilla me- 
galops Manning & Bruce, 1984. This new superfamily differs from the Bath- 
ysquilloidea, Gonodactyloidea, and Squilloidea by its member having broad 
and ventrally beaded propodi of the third and fourth maxillipeds; it differs 
from the Lysiosquilloidea by having a distinct dorsal median carina on the 
telson. The superfamily may represent a relict lineage within the Stomatopoda. 
The family Tetrasquillidae is erected in the superfamily Lysiosquilloidea for 
three genera, the monotypic Tetrasquilla Manning & Chace, 1990; Tectasquilla 
Adkison & Hopkins, 1984; and Heterosquillopsis Moosa, 1991, which contains 
three species. This new family can be distinguished from the three families 
now remaining in the Lysiosquilloidea by its members having ovate rather 
than styliform (as in the Lysiosquillidae) distal segments of the endopods of 
the first two walking legs, by lacking an enlarged basal lobe on the dactylus of 
the claw (Coronididae), and by lacking a strong proximal fold on the outer 
margin of the uropodal endopod (Nannosquillidae). The only known pantrop- 
ical stomatopod, Tetrasquilla mccullochae (Schmitt, 1940), is included in this 


family. 


Manning & Bruce (1984:332) tentatively 
placed their newly erected family Erythro- 
squillidae in the superfamily Lysiosquilloi- 
dea based on the presence of broad, ven- 
trally beaded propodi of the third and fourth 
maxillipeds of the only member of the fam- 
ily, Erythrosquilla megalops Manning & 
Bruce, 1984 (Fig. 1). They pointed out, how- 
ever, that the Erythrosquillidae differ from 
other lysiosquilloids by having a sharp, dor- 
sal median carina on the telson, by lacking 
a ventrolateral projection on the sixth ab- 
dominal somite overhanging the articula- 
tion of the uropods, and by having a smaller 
ventral papilla of the antennal protopod. 
The first of those three characters is cur- 
rently considered important in distinguish- 
ing superfamilies of the stomatopods, and 
we use it here to help differentiate the new 
superfamily defined below. Our removal of 


the Erythrosquillidae from the Lysiosquil- 
loidea leaves three families in that super- 
family: Coronididae Manning, 1980, Ly- 
siosquillidae Giesbrecht, 1910, and 
Nannosquillidae Manning, 1980. 

Camp & Kuck (1990:852) pointed out that 
a new family might have to be erected for 
Heterosquilloides mccullochae (Schmitt, 
1940), a species placed in the recently erect- 
ed, monotypic genus TJetrasquilla by Man- 
ning & Chace (1990) and assigned to the 
Lysiosquillidae. Camp & Kuck (1990) not- 
ed that characters of the species fit none of 
the lysiosquilloid families then known 
(Manning 1980), and that it also could not 
be accommodated in the Erythrosquillidae 
Manning & Bruce. A new family is diag- 
nosed here for 7. mccullochae and for the 
related Tectasquilla lutzae Adkison & Hop- 
kins, 1984 (Fig. 2). Keys to the superfami- 


86 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Erythrosquilla megalops (Erythrosquillidae). a, Anterior appendages; b, Claw; c, Distal segments of 
third maxilliped; d, Sixth abdominal somite, telson, and uropod; e, Uropod, ventral view (from Manning & 
Bruce 1984:fig. 1). 


lies of Recent Stomatopoda and to the fam- 
ilies of Lysiosquilloidea are presented below. 

Abbreviations used include mm (milli- 
meters), tl (total length, measured on mid- 
line in mm), and USNM (National Museum 
of Natural History, Smithsonian Institu- 
tion, Washington, D.C.). The specimens il- 
lustrated herein are as follows: Erythrosquil- 
la megalops, male holotype, tl 105, Indian 
Ocean off Somalia, USNM 195339; Coron- 
ida bradyi (A. Milne Edwards, 1869), fe- 
male, tl 33, Annobon Island, Gulf of Guin- 
ea, USNM 151531; Lysiosquilla scabricauda 


(Lamarck, 1818), male, tl 227, Fort Pierce, 
Florida, USNM 152469 (walking legs) and 
female, tl 44, St. Lucie Inlet, Florida, USNM 
256888 (uropod and claw); Nannosquilla 
grayi (Chace, 1958), female holotype, tl 40, 
Bass River, Massachusetts, USNM 100931 
(claw) and female paratype, tl 41, same lo- 
cality, USNM 100932 (walking legs and 
uropod); Tetrasquilla mccullochae, female, 
tl 32, Alligator Reef, Florida, USNM 
111028; Tectasquilla lutzae, male holotype, 
tl 73, Gulf of Mexico, off northwestern Flor- 
ida, USNM 204717. 


VOLUME 106, NUMBER 1 87 


Fig. 2. a, Tetrasquilla mccullochae (from Manning & Chace 1990:fig. 46); b, Tectasquilla lutzae (from Adkison 
& Hopkins 1984:fig. 1a). 


Erythrosquilloidea, new superfamily movable apices. No more than 2 interme- 
diate denticles present on telson. 
Diagnosis. —Propodi of third and fourth Type genus. —Erythrosquilla Manning & 
maxillipeds broad, ventrally beaded. Telson Bruce, 1984, herein designated. 
with distinct dorsal median carina. At most, Included families. —Erythrosquillidae 
submedian marginal teeth of telson with Manning & Bruce, 1984. 


88 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Remarks. —The superfamily Erythro- 
squilloidea can be distinguished from 
Bathysquilloidea Manning, 1967, Gono- 
dactyloidea Giesbrecht, 1910, and Squil- 
loidea Latreille, 1803 by the propodi of the 
third and fourth maxillipeds being broad 
and ventrally beaded rather than being slen- 
der and not ventrally beaded; the superfam- 
ily can be distinguished from the Lysios- 
quilloidea Giesbrecht, 1910 by having a 
distinct dorsal median carina on the telson. 
See Manning & Bruce (1984:332) for a fur- 
ther discussion of the relationship of the 
possibly relict Erythrosquillidae to the other 
stomatopod superfamilies. 

We take this opportunity to correct errors 
in the definitions of the families Coronidi- 
dae and Lysiosquillidae presented in Man- 
ning (1980:368) and in the description of 
Erythrosquilla megalops in Manning & 
Bruce (1984:331). In each instance, refer- 
ence to the lack of a strong fold on the inner 
margin of the uropodal endopod should re- 
fer instead to the outer margin of that ap- 
pendage. 


Superfamily Lysiosquilloidea 
Giesbrecht, 1910 
Tetrasquillidae, new family 


Diagnosis. —Size medium, total lengths 
of adults <75 mm. Body compact, de- 
pressed. Dactylus of claw not inflated ba- 
sally. Endopods of anterior 2 walking legs 
ovate. Uropodal endopod lacking strong 
proximal fold on outer margin. 

Type genus.—Tetrasquilla Manning & 
Chace, 1990, herein designated. 

Included genera. —Three: Tetrasquilla 
Manning & Chace, 1990, containing only 
T. mccullochae (Schmitt, 1940), the only 
known pantropical stomatopod (see Man- 
ning & Chace 1990); Tectasquilla Adkison 
& Hopkins, 1984, containing only Tecta- 
squilla lutzae Adkison & Hopkins, 1984, 
known only from off northwestern Florida 
and Georgia, U.S.A.; and Heterosquillopsis 
Moosa, 1991, containing three species from 
the Indo-West Pacific, H. insueta (Manning, 


1970), H. philippinensis (Moosa, 1986), and 
H. danielae Moosa, 1991, the type species. 

Remarks. — This new family can be readi- 
ly distinguished from the three other fam- 
ilies now remaining in the Lysiosquilloidea. 
The Tetrasquillidae differ from the Lysio- 
squillidae in that the distal segment of the 
endopods of the first two walking legs are 
ovate (Fig. 3j, k, m) rather than slender and 
styliform (Fig. 3d, e). The Tetrasquillidae 
differ from the Nannosquillidae by lacking 
a strong proximal fold on the outer margin 
of the uropodal endopod (compare Fig. 4c 
and 4d, e). The Tetrasquillidae can be dis- 
tinguished from the Coronididae by the claw 
(Fig. 5d, e) having the propodus pectinate 
for all its length and by lacking the basal 
inflation of the dactylus (Fig. 5a). 


Key to Superfamilies of 
Recent Stomatopoda 
(Modified from Manning 1980) 


1. Propodi of third and fourth maxil- 
lipeds slender, not beaded or ribbed 


ventrally o<* -..5 +3). ee 2 
— Propodi of third and fourth maxil- 
lipeds broad, usually beaded or 
ribbed ventrally «.-..... S35 aaeee 4 
2. All marginal teeth of telson with 
movable apices ...... Bathysquilloidea 


— At most, submedian marginal teeth 
of telson with movable apices .... 3 
3. Four or more intermediate denticles 
present on telson Squilloidea 
— Two or fewer intermediate denticles 
present on telson ... Gonodactyloidea 
4. Telson lacking sharp dorsal median 


©, ee. ew wy fey tm 


Canin! Mas ee eee Lysiosquilloidea 
— Telson with sharp dorsal median ca- 
Tina 2. eee Erythrosquilloidea 


Key to Families of Lysiosquilloidea 


1. Dactylus of claw inflated basally. 
Propodus of claw pectinate proxi- 
mally only Coronididae 

— Dactylus of claw not inflated basal- 
ly. Propodus of claw completely 
lined with pectinations 


6) ao “wie fe: (a) eof) oie) ie ete 


VOLUME 106, NUMBER 1 89 


Fig. 3. Walking legs 1-3: a—c, Coronida bradyi (Coronididae); d—f, Lysiosquilla scabricauda (Lysiosquillidae); 
g-i, Nannosquilla grayi (Nannosquillidae); j-/, Tetrasquilla mccullochae (Tetrasquillidae); m—n, Tectasquilla 
lutzae (Tetrasquillidae) (legs 1 and 3; from Adkison & Hopkins 1984:fig. 2h, i). 


2. Proximal portion of outer margin of — Distal segment of endopod of an- 

uropodal endopod with strong fold terior two walking legs broadly ovate 

..: 5 Nannosquillidae pee hos). Wee We a PeteasguEnlhidae 

— Proximal portion of outer margin of 

oe endopod lacking strong ethowledemerts 

Co ee ee 3 
3. Distal segment of endopod of an- The figures were prepared by Lilly King 

terior two walking legs slender, sty- Manning. Studies on stomatopod system- 


SUS De Lysiosquillidae atics have been supported by the Smithson- 


90 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. _Uropods, dorsal view: a, Coronida bradyi (Coronididae); b, Lysiosquilla scabricauda (Lysiosquillidae); 
c, Nannosquilla grayi (Nannosquillidae); d, Tetrasquilla mccullochae (Tetrasquillidae); e, Tectasquilla lutzae 
(Tetrasquillidae) (from Adkison & Hopkins 1984:fig. 2e). 


Fig. 5. Distal segments of raptorial claw: a, Coronida bradyi (Coronididae); b, Lysiosquilla scabricauda 
(Lysiosquillidae); c, Nannosquilla grayi (Nannosquillidae); d, Tetrasquilla mccullochae (Tetrasquillidae); e, Tec- 
tasquilla lutzae (Tetrasquillidae) (from Adkison & Hopkins 1984:fig. 1f). 


VOLUME 106, NUMBER 1 


ian Institution through its Scholarly Studies 
Program and the Smithsonian Marine Sta- 
tion at Link Port, Florida; this is contri- 
bution number 304 from that station. The 
second author (DKC) thanks K. A. Steidin- 
ger, Chief, Florida Marine Research Insti- 
tute (FMRI), for supporting systematics re- 
search on the marine biota of Florida, and 
colleagues at FMRI, L. French, J. Leiby, W. 
G. Lyons, T. H. Perkins, and J. F. Quinn, 
Jr., for suggesting improvements to the 
manuscript. 


Literature Cited 


Adkison, D. L., & T. S. Hopkins. 1984. Tectasquilla 
lutzae, new genus and species (Crustacea: Sto- 
matopoda: Lysiosquillidae) from the Gulf of 
Mexico.— Proceedings of the Biological Society 
of Washington 97:532-537. 

Camp, D. K., & H. G. Kuck. 1990. Additional rec- 
ords of stomatopod crustaceans from Isla del 
Coco and Golfo de Papagayo, eastern Pacific 
Ocean.— Proceedings of the Biological Society 
of Washington 103:847-853. 

Chace, F. A., Jr. 1958. A new stomatopod crustacean 
of the genus Lysiosquilla from Cape Cod, Mas- 
sachusetts.— Biological Bulletin, Woods Hole 
114(2):141-145. 

Giesbrecht, W. 1910. Stomatopoden, Erster Theil. — 
Fauna und Flora des Golfes von Neapel 33:vii 
+ 239 pp., pls. 1-11. 

Lamarck, J. B. P. A. de. 1818. Histoire naturelle des 
animaux sans vertebrés, présentant les carac- 
téres généraux et particuliers de ces animaux, 
leur distribution, leurs classes, leurs familles, 
leurs genres, et la citation des principales espéces 
qui s’y rapportent; précédée d’une introduction 
offrant la détermination des caractéres essentiels 
de l’anomas, sa distinction du végétal et des 
autres corps naturelles, enfin, l’exposition des 
principes fondamentaux de la zoologie 5:612 
pp., Deterville, Paris. 

Latreille, P. A. 1803. Histoire naturelle, générale et 
particuliére, des Crustacés et des Insectes 3:467 
pp., F. Dufart, Paris. 


91 


Manning, R. B. 1967. Preliminary account of a new 
genus and a new family of Stomatopoda.—Crus- 
taceana 13:238-239. 

. 1970. Two new stomatopod crustaceans from 

Australia.— Records of the Australian Museum 

28(4):77-85. 

. 1980. The superfamilies, families, and genera 

of Recent stomatopod Crustacea, with diagno- 

ses of six new families.— Proceedings of the Bi- 

ological Society of Washington 93:362-372. 

—.,&A.J. Bruce. 1984. Erythrosquilla megalops, 
a remarkable new stomatopod from the western 
Indian Ocean.—Journal of Crustacean Biology 
4:329-332. 

—, & F. A. Chace, Jr. 1990. Decapod and sto- 
matopod Crustacea from Ascension Island, 
South Atlantic Ocean.—Smithsonian Contri- 
butions to Zoology 503:v + 91 pp. 

Milne Edwards, A. 1869. Rade de Saint-Vincent du 
Cap-Vert (supplément). Pp. 136-138, pl. 17 in 
L. de Folin & L. Périer, 1867-1872, Les fonds 
de la mer, étude internationale sur les particu- 
larités nouvelles des régions sous-marins, 1, 
Bordeaux. 

Moosa, M. K. 1986. Stomatopod Crustacea. Résul- 

tats du Campagnes MUSORSTOM I & II Phil- 

ippines, 2.—Mémoires du Muséum National 
d’Histoire Naturelle, Paris (Series A, Zoology) 

133:367-414, pl. 1. 

1991. The Stomatopoda of New Caledonia 
and Chesterfield Islands. Pp. 149-219 in B. 
Richer de Forges, ed., Le benthos des fonds 
meubles des lagons de Nouvelle-Calédonie, 1, 
Editions de 1ORSTOM, Paris. 

Schmitt, W. L. 1940. The stomatopods of the west 
coast of America based on collections made by 
the Allan Hancock Expeditions, 1933-38.—Al- 
lan Hancock Pacific Expeditions 5(4):129-225. 


(RBM) Department of Invertebrate Zo- 
ology, National Museum of Natural His- 
tory, Smithsonian Institution, Washington, 
D.C. 20560, U.S.A.; (DKC) Florida Marine 
Research Institute, 100 Eighth Avenue SE, 
Saint Petersburg, Florida 33701-5095, 
UWES-A, 


PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 92-101 


SYSTEMATICS AND TAXONOMIC REMARKS ON 
PINNOTHERES MULINIARUM RATHBUN, 1918 
(CRUSTACEA: BRACHYURA: PINNOTHERIDAE) 


Ernesto Campos 


Abstract.—The new genus Juxtafabia is proposed to receive the clam crab 
Pinnotheres muliniarum Rathbun, 1918. The primary diagnostic features are: 
third maxilliped with carpus and propodus stout, subequal in length; propodus 
suboblong, with distal end obliquely rounded and a lunate and broad dactylus 
inserted near its proximal end; and male abdomen widest at somite 3, narrowing 
at somite 4 which is fused to somite 5, and telson longer than broad. The type 
species, Pinnotheres muliniarum Rathbun, 1918, described from a pre-hard 
stage male is considered to be a senior synonym of P. reticulatus Rathbun, 
1918, which was described on the basis of an adult female. Description and 
figures of the young pre-hard stage, adult hard stage male, adult female, com- 
plete synonymy, known distribution, and host are presented. 

Resumen. —El nuevo género Juxtafabia se propone para recibir al cangrejo 
almejero Pinnotheres muliniarum Rathbun, 1918. Sus caracteristicas diagnos- 
ticas primarias son: tercer maxilipedio con el carpus y propodus iguales en 
longitud y robustos; el propodus suboblongo tiene su final distal oblicuamente 
redondeado y lleva proximoventralmente al amplio y lunado dactilus; y el 
abdomen, en el macho, se ensancha en el somita 3, se adelgaza en el 4 el cual 
esta fusionado al 5, y con un telson mas largo que ancho. La especie tipo, 
Pinnotheres muliniarum Rathbun, 1918, que fue descrita en base a un macho 
en fase predura es aqui considerado como un sinonimo antiguo de P. reticulatus 
Rathbun, 1918, la cual fue descrita sobre la base de una hembra adulta. Des- 
cripcion y figuras de las fases masculinas pre-dura y dura, la hembra adulta, 
la sinonimia completa, la distribuciOn conocida y los huéspedes para esta 
especie, se registran. 


The Pinnotheridae is a group of symbiotic 
and sexually dimorphic crabs that undergo 
a series of morphological changes during 
their postplanktonic development. This has 
resulted in some mistaken identifications 
representing developmental stages of the 
same species. For example, Williams (1965) 
found that Pinnotheres depressus Rathbun, 
1918 was the pre-hard stage male of P. os- 
treum Say, 1817, and Campos (1989b) con- 
cluded that P. pubescens (Holmes, 1894) is 
a young female of Tumidotheres margarita 
(Smith, 1869). Recently, Green (1985) de- 


termined that P. jamesi Rathbun, 1923, de- 
scribed from a hard stage male (sensu Chris- 
tensen & McDermott 1958) is a junior 
synonym of P. reticulatus Rathbun, 1918, 
a species which was described on the basis 
of a post-hard stage female. Postplanktonic 
stages of P. reticulatus collected in the upper 
Gulf of California in 1986-1987 confirmed 
Green’s conclusions, and a comparison of 
the pre-hard stage male with the holotype 
of P. muliniarum Rathbun, 1918 revealed 
no difference. In addition, a morphological 
analysis of these specimens has led to the 


VOLUME 106, NUMBER 1 


proposition of a new genus for this species. 
This new genus most closely resembles Fa- 
bia Dana, 1851. 

Specimens for this study were obtained 
from: National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 
(USNM); Colleccion Carcinoldgica, Insti- 
tuto de Biologia, Universidad Nacional Au- 
tonoma de México (EM) and Facultad de 
Ciencias, Universidad Autonoma de Baja 
California (UABC). The abbreviation cw 
refers to carapace width. 


Systematics and Taxonomic Account 
Juxtafabia, new genus 


Diagnosis. —Female carapace subglobu- 
lar, dorsal regions weakly defined. Third 
maxilliped with ischium and merus indis- 
tinguishably fused, ventral margin angular, 
dorsal margin gently convex; palp slightly 
shorter than ischium-merus, carpus and 
propodus subequal in length, stout, latter 
article suboblong with distal end obliquely 
rounded, dactylus lunate, broad, ventrally 
inserted near proximal end of propodus. 
Abdomen with 7 free somites, laterally 
reaching to coxae of walking legs, distally 
covering buccal cavity. 

Male carapace subglobular, regions weak- 
ly defined, pterygostomian region elevated, 
eyes visible in dorsal view; third maxilliped 
similar to that of female; abdomen widest 
at somite 3, narrowing at somite 4 which is 
fused to somite 5, telson longer than broad. 

Type species. — By present designation and 
monotypy, Pinnotheres muliniarum Rath- 
bun, 1918. Gender feminine. 

Distribution. —Upper Gulf of California, 
México to Costa Rica, Central America. 

Hosts.—Mollusca, Bivalvia: species of 
Chione, Polymesoda, Protothaca and Ta- 
gelus. 

Etymology.—The generic name is de- 
rived from the latin Juxta, close to, at side 
of, and the generic name Fabia. The name 
has been selected to emphasize the resem- 
blance of Juxtafabia and Fabia. 


93 


Remarks.—Other genera in the Pinno- 
theridae which also have the ischium and 
merus of the third maxilliped indistinguish- 
ably fused differ from Juxtafabia as follows: 
Dissodactylus Smith, 1870, Clypeasterophi- 
lus Campos & Griffith, 1990, Parapinnixa 
Holmes, 1894 and Sakaina Seréne, 1964 
have an exopod without flagellum (Fig. 1 A); 
Calyptraeotheres Campos, 1990, Ostraco- 
theres H. Milne-Edwards, 1853 and Xan- 
thasia White, 1846 have a palp with two 
articles (Fig. 1B); Limotheres Holthuis, 1975 
and Orthotheres Sakai, 1969 have the dac- 
tylus inserted distally on the propodus (Fig. 
1C); Pinnixa White, 1846 and Scleroplax 
Rathbun, 1893 have a lobe on the external 
margin of the exopod (Fig. 1D); Fabia Dana, 
1851, Tumidotheres Campos, 1989b, and 
Durckheimia De Man, 1889, have a small 
palp with the dactylus inserted on the mid- 
dle of the propodus (Fig. 1E); and Pinnothe- 
res Bosc, 1802 s. str. has a linear and slender 
dactylus inserted proximally on the inner 
margin of the spatulate propodus (Fig. 1F), 
and seven free abdominal somites. Addi- 
tional differences between Juxtafabia and 
the above genera can be found in Burger 
(1895), Rathbun (1918), Tesch (1918), Se- 
rene (1964, 1967), Holthuis (1975), Cam- 
pos (1989a, 1989b, 1990) and Campos & 
Griffith (1990). 

The new genus Juxtafabia most closely 
resembles the genus Fabia. Males in both 
genera have two or more abdominal somites 
fused. However, there are no morphological 
similarities between females of the two gen- 
era. Larvae of J. muliniarum resemble zoeae 
of Fabia and Pinnixa. Zoeae in these genera 
have a common pattern of setae on their 
appendages, a carapace with moderately de- 
veloped rostral, lateral and dorsal spines and 
a distinctive abdominal shape. These genera 
share a very distinct lateral and posterior 
expansion on the fifth abdominal somite 
(Fig. 2A—-E). The genus Pinnixa, however, 
has an elongated telson which is quite dis- 
tinct from that in Juxtafabia and Fabia. 
Whether these morphological similarities 


94 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ABN 


Fig. 1. Third maxilliped. A, Dissodactylus lockingtoni Glassell, 1935; B, Calyptraeotheres granti (Glassell, 
1933); C, Orthotheres unguifalcula (Glassell, 1935); D, Scleroplax granulata Rathbun, 1893; E, Fabia sp.; F, 
Pinnotheres pisum (Linnaeus, 1767). [F, redrawn from Ingle (1980:52, fig. 25)]. 


VOLUME 106, NUMBER 1 95 


Fig. 2. Dorsal view of zoeal abdomen. A, Juxtafabia muliniarum (Rathbun, 1918); B, Fabia subquadrata 
Dana, 1851; C, Fabia sp.; D, Pinnixa longipes (Lockington, 1877); E, Pinnixa aff. rathbuni Sakai, 1934 [D and 
E redrawn from Bousquette (1980:596, fig. 4A) and Konichi (1983:282, fig. 14C) respectively]. 


96 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


between males and larvae have phyloge- 
netic implications remains to be deter- 
mined. 


Juxtafabia muliniarum (Rathbun, 1918), 
new combination 


Pinnotheres muliniarum Rathbun, 1918: 81, 
pl. 18, figs. 2, 3.—Glassell 1934:301.— 
Schmitt et al. 1973:5, 10, 58.—Campos 
1989b:672. 

Pinnotheres reticulatus Rathbun, 1918:93- 
94, pl. 21, figs. 1, 2.—Glassell 1934:301; 
1935:105.—Schmitt et al. 1973:5, 83.— 
Green 19852 611=614- ings? (hy 
Campos-Gonzalez 1988:385.—Campos 
1989b:672. 

Pinnotheres jamesi Rathbun, 1923:625-626, 
pl. 29, figs. 1, 2, text-figs. 1, 2. Glassell 
1934:301.—Silas & Alagarswami 1967: 
1200, 1223.—Schmitt et al. 1973:50. 


Type locality.—Lower California (Baja 
California, Mexico). 

Distribution. —Gulf of Santa Clara (Gulf 
of California), Baja California, Mexico to 
Costa Rica. 

Hosts. —Mollusca, Bivalvia: Chione cal- 
iforniensis (Broderip), C. fructifaga (Sow- 
erby), C. tumens (Verril), Polymesoda infla- 
ta (Philippi), Protothaca grata (Say) and 
Tagelus affinis (C. B. Adams). 

Material examined.—Holotype of Pin- 
notheres muliniarum, Lower California, 
male, cw 3.6 mm (fide Rathbun 1918), 
USNM 23443.—Holotype of P. reticulatus, 
off San Jose Island, Baja California Sur, 
25°O2' 15"N, 110°43'30°W, 17 tms (31 ma), 
S.Sh., 17 Mar 1889, Station 3002 Albatross, 
female, cw 9.2 mm (fide Rathbun 1918), 
USNM 18217.—1 female, cw 2.9 mm, Gulf 
of Santa Clara, Baja California, 25 Apr 1990, 
in Chione fructifaga, UABC.—12 ovigerous 
females, cw 5 to 10 mm, 4 young males, cw 
1 to 2.2 mm, Campo El Pescador, on coast 
4.5 km North of San Felipe, Baja California, 
1988, in Protothaca grata, Chione califor- 
niensis and C. fructifaga, UABC.—7 young 
males, cw 1 to 2.3 mm, 11 adult males, cw 


2.2 to 3.4 mm, and 30 females, cw 4 to 10 
mm, Laguna Pecebut, about 23 km south of 
San Felipe, 1987-88, in P. grata and Ta- 
gelus affinis, UABC.—1 female, cw 5.7 mm, 
Puertecitos, km 72 road San Felipe-San Luis 
Gonzaga, Aug 1986, in P. grata, UABC.— 
1 ovigerous female, cw 10 mm, San Marcos 
Island, north of La Yesera (27°17'N, 
112°07'W), 16 May 1987, in Chione tu- 
mens, EM. 

Male pre-hard stage.—Carapace trans- 
versely subglobular, soft, posterior margin 
convex; front slightly projected and strongly 
deflexed (Fig. 3). Third maxilliped with palp 
slightly shorter than ischium-merus (Fig. 
5A); carpus subequal to propodus in length, 
latter suboblong, with distal end obliquely 
rounded; dactylus broad, lunate, ventrally 
inserted near proximal end of propodus, 
nearly reaching to distal end of last article. 

Chelipeds stouter than walking legs, me- 
rus widening slightly distally, dorsal margin 
convex, ventral margin straight; fingers sub- 
equal, slightly deflexed, curved at tip where 
they cross; cutting edge of dactylus with 
small tooth. 

Relative length of the walking legs 2 > 3 
> 1 > 4, fourth leg not overreaching carpus 
of third leg; margin of legs hairy, specially 
on meri; dorsal margin of propodi convex, 
ventral margin in legs 1-3 straight, concave 
in leg 4; dactyli 1-3 hook-like; dactylus 4 
less curved than on legs 1-3. 

Abdomen widest at somite 3, narrowing 
at somite 4, telson longer than broad, dis- 
tally rounded. Somites 4 and 5 fused. 

Gonopods simple, curved at base, distally 
straight (Fig. 5B). 

Male hard stage. —Carapace (Fig. 4) 
transversely subglobular, well calcified, lat- 
eral margins with a heavy fringe of hair-like 
setae; pterygostomian region higher than in 
pre-hard stage so that eyes are more con- 
spicuous in dorsal view; frontal region oc- 
casionally produced; posterior margin al- 
most straight. Third maxilliped similar to 
that of male pre-hard stage (Fig. 5C). 

Chelipeds no more stout than walking legs; 


VOLUME 106, NUMBER 1 


“ARS. | as 
SN NOW rH i y q 


SAD 
. 
(ile thd inter p RG ® 
a i / SN 
5» 


G7 


Fig. 3. Juxtafabia muliniarum (Rathbun, 1918); dorsal view of pre-hard stage male. Carapace length = 2.3 


mm. 


manus widening distally; dorsal margin 
convex; fingers subequal, curved and crossed 
at tip; ventral margin almost straight; pollex 
with 2 acute teeth on proximal half of cut- 
ting edge, between them a notch where trun- 
cate tooth of dactylus fits. 

Walking legs stout, dorsal and ventral 
margin of meri fringed with setae; propodi 


tapering distally, dorsally convex, almost 
straight ventrally; dactyli equal to those of 
pre-hard stage. External face of carpus and 
propodus of walking legs 2 and 3 with long 
swimming setae. 

Abdomen similar to that of pre-hard stage, 
but somite 7 occasionally subrectangular 
(Fig. 5D). 


Fig. 4. Juxtafabia muliniarum (Rathbun, 1918); dorsal view of hard stage male. Carapace length = 3.14 
mm. Single setae shown on dorsal and ventral margin of the last walking legs may actually be plumose setae. 


98 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Cc 


Fig. 5. Juxtafabia muliniarum (Rathbun, 1918). Pre-hard stage male: A, third maxilliped; B, gonopod. Hard 
stage male C, third maxilliped; D, abdomen; E, gonopod. F, third maxilliped of last post-hard stage female. [A 
and C-D redrawn from Rathbun (1918:81, fig. 38, and 1923:626, figs. 1 and 2)]. 


Gonopods similar to those in pre-hard 
stage, but longer and slender (Fig. 5E). 

Female (ovigerous).—Carapace (Fig. 6) 
subglobular, broader than long, soft, slightly 
convex; gastric region elevated on medial 
line and separated by broad depression from 
branchial region; a lunate depression near 
hepatic margin. Front slightly convex in 
dorsal view, scarcely projecting beyond 
curve of anterolateral margin. Eyes globu- 
lar, completely filling orbits, visible dorsal- 
ly, cornea evident. Epistome and labium 


similar in length and width. Antenna with 
blocky basal articles, flagellum minute, not 
overreaching upper margin of orbit. Anten- 
nulae placed in wide fossettes not larger than 
orbits. 

First 2 articles of palp of third maxilliped 
broad, subequal in length; dactylus curved, 
lunate, attached proximally and reaching to 
end of suboblong propodus (Fig. 5F). 

Chelipeds and legs short, pubescent; che- 
lipeds hairy on inner faces; manus subellip- 
tical, increasing slightly distally, lower mar- 


VOLUME 106, NUMBER 1 


99 


Fig. 6. Juxtafabia muliniarum (Rathbun, 1918); dorsal view of adult female. Carapace length = 7 mm. (Eyes 
are dorsally visible when abdomen is folded towards carapace sternum). 


gin straight, upper slightly convex; 
pubescence arranged in reticulate pattern, 
the interstices of which are smooth and of 
darker color (in alcohol); fingers moderately 
stout, tips hooked toward each other, pre- 
hensile edges of each armed with 2 teeth of 
which that near base of dactylus is largest, 
other minute teeth between them. 

Walking legs furnished with long setae on 
lower margin, specially on propodus of first 
and fourth leg; relative length of legs 2 > 3 
> 4 > 1, the second leg may be asymmet- 
rical, with propodus and dactylus of left leg 
longer than on right one; ventral margin of 
propodus of second and third leg concave; 
ventral margin of propodus of first and 
fourth leg straight; dactyli 1, 2 and 3 similar, 
curved and with slender tips, first shortest, 


second longest; dactylus 4 much longer than 
those preceding and may be longer than its 
propodus. 

Abdomen with 7 free somites, longer and 
broader than carapace (modified from 
Rathbun 1918). 

Remarks.—The study of males in hard 
stage (host, Zagelus affinis) and females in 
post-hard stage (host, Protothaca grata and 
T. affinis) collected during the spring and 
summer of 1986-1987 at Laguna Percebu, 
south of San Felipe, led to the confirmation 
of Green’s (1985) conclusion that Pinnothe- 
res jamesi is ajunior synonym of P. reticula- 
tus. Subsequent collecting of pre-hard stage 
males (host, Chione californiensis) of a pin- 
notherid crab from Campo El Pescador 
showed no difference with Rathbun’s (1918) 


100 


holotype of Juxtafabia muliniarum. Ex- 
amination of additional males collected 
from T. affinis showed that some specimens 
represented pre-hard stage males which were 
identical to those previously identified as J. 
muliniarum (from El Pescador). The other 
specimens were hard stage males identical 
to P. jamesi. To determine whether both 
types of males examined were conspecific, 
specimens of pre-hard stage males (=J. mu- 
liniarum) were kept alive until they molted 
to a hard stage identical to those identified 
as P. jamesi. These results and the study of 
the complete postplanktonic female stages 
from San Felipe and vicinity, led to the con- 
clusion that J. muliniarum, P. jamesi and 
P. reticulatus represent the same species. 
Juxtafabia muliniarum was the first named 
species and, therefore, is the senior syn- 
onym of the other two taxa. 


Acknowledgments 


Many thanks to Prof. A. Carvacho for 
allowing me to revise the holotypes of J. 
muliniarum and P. reticulatus which were 
kindly loaned to him by Dr. R. B. Manning 
(Smithsonian Institution). I also thank Prof. 
Carvacho and J. F. Bergerault for loaning 
additional specimens of J. muliniarum. My 
deep gratitude goes to my wife Alma Rosa 
for hard field and laboratory work, for pre- 
paring original figures, and for commenting 
on early manuscript versions and most im- 
portantly for continued encouragement to 
see this paper to completion. This work was 
partially supported by the program “‘Siste- 
matica de Crustaceos” of the Facultad de 
Ciencias, Universidad Autonoma de Baja 
California and by agreements SEP-UABC 
087-01-0426 and 089-01-0352 and 
CONACYyT 0482-N9108. 


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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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Facultad de Ciencias, Universidad Au- 
tonoma de Baja California, Apartado Postal 
2300, Ensenada, Baja California, México. 


PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 102-105 


ANOMOEOMUNIDA, A NEW GENUS PROPOSED FOR 
PHYLLADIORHYNCHUS CARIBENSIS MAYO, 1972 
(CRUSTACEA: DECAPODA: GALATHEIDAE) 


Keiji Baba 


Abstract. — Anomoeomunida, a new genus, is established for Phylladiorhyn- 
chus caribensis Mayo, 1972, and is differentiated from Phylladiorhynchus Baba, 
1969, by the presence of male pleopods on the first abdominal segment, the 
rostrum bearing a dorsal ridge in midline, the single spine at the distolateral 
angle of the antennular basal segment, and the lack of ventral spines on the 
sickle-shaped dactyli of the walking legs. It is also distinguished from Pleuron- 
codes Stimpson, 1860, by the pterygostomial flap not visible from a dorsal 
view and the dactyli of the walking legs sickle-shaped without ventral spines. 


In a previous paper, Baba (1991:484) sug- 
gested that Phylladiorhynchus caribensis 
Mayo, 1972 from the Caribbean Sea be re- 
moved from that genus. It was suggested 
that the species was close to Pleuroncodes 
Stimpson, 1860, from the eastern Pacific 
but that a new genus was needed. A more 
thorough comparison now allows a new ge- 
nus, Anomoeomunida, to be proposed to 
include this species, and be discussed. 


Anomoeomunida, new genus 


Diagnosis. —Rostrum relatively narrow, 
dorsally with rounded low ridge, with well- 
developed supraocular tooth on each side. 
Lateral limit of orbit with distinct angle in 
dorsal view. Basal segment of antennule with 
3 small terminal spines, mesial one not bi- 
fid. Third thoracic sternite with 2 convex- 
ities on anterior margin, posterior margin 
contiguous to following sternite on entire 
width. Walking legs lacking spines other than 
terminals on merus and carpus; dactylus 
sickle-shaped, lacking ventral spines. Male 
pleopods present on first and second ab- 
dominal somites. 

Gender. — Feminine. 

Type species.—Phylladiorhynchus cari- 
bensis Mayo, 1972. 


Etymology. —From the Greek anomoios 
(unlike, dissimilar) plus Munida, suggesting 
that the genus is different from Munida. 

Remarks. —Mayo (1972:526) noted that 
Phylladiorhynchus caribensis from 11-38 m 
in the Caribbean Sea was intermediate be- 
tween Galathea Fabricius, 1793, and Muni- 
da Leach, 1820, and placed it in Phylla- 
diorhynchus Baba, 1969. She stressed the 
rostral shape in species of the genus and 
amended the generic diagnosis to allow for 
the lack of spination on the walking legs, 
and for three rather than four or five ter- 
minal spines (counting the double mesial 
terminal spines as two) on the basal anten- 
nular segment. Lewinsohn (1982:298) sug- 
gested that this western Atlantic form should 
eventually be placed in a new genus. Fol- 
lowing the examination of the type material 
deposited at the National Museum of Nat- 
ural History, Smithsonian Institution, 
Washington, D.C. (USNM 140187-8) of P. 
caribensis, I am inclined to believe that 
Lewinsohn was correct. 

Anomoeomunida caribensis has most of 
the characteristics of Munida in the general 
striation of the carapace and abdomen, and 
shapes of the antennular basal segment, an- 
tenna and sternum (the anterior portion in 
particular). But the lateral limit of the orbit 


VOLUME 106, NUMBER 1 


103 


Fig. 1. 


Anterior part of carapace in Anomoeomunida caribensis [= Phylladiorhynchus caribensis Mayo, 1972]: 


a, male holotype (USNM 140187), dorsal view, right supraocular spine partly broken; b, same, left lateral view. 


Scales = 1 mm. 


angled in dorsal view, and the walking legs 
with smooth, sickle-shaped dactyli and meri 
without dorsal spines, differentiate it from 
other species of Munida. 

The unique morphology of the walking 
legs also separates A. caribensis from species 
of Phylladiorhynchus. The rostrum in spe- 
cies of Phylladiorhynchus is broadly trian- 
gular, dorsally excavated, and lacks the dor- 
sal ridge seen in the Caribbean species. Mayo 
(1972:523) noted that the lateral limit of the 
orbit in P. caribensis bears a small spine. 
Examination of the type material discloses 
that this spine can be discernible in dorsal 
view (Fig. la), but barely so in lateral view 
(Fig. 1b); in Phylladiorhynchus, the lateral 
orbital angle is sharply produced (Baba 1991: 
fig. 4). The distomesial margin of the an- 
tennular basal segment in Phylladiorhyn- 
chus bears double spines (see Miyake & Baba 
1965:fig. 6A for P. ikedai; Miyake & Baba 
1967: fig. 6b for P. pusillus; Tirmizi & Javed 
1980:fig. 2D for P. bengalensis; Lewinsohn 
1982:fig. le for P. integrirostris), while that 
of P. caribensis bears a single thin small 
spine (Mayo 1972:525, fig. le). The disto- 
ventral margin of the first (basal) segment 
of the antennal peduncle is strongly pro- 
duced into an anterior prolongation in Phyl- 
ladiorhynchus (see Miyake & Baba 1965:fig. 
4B for P. ikedai; Miyake & Baba 1967:fig. 


6c for P. pusillus; Tirmizi & Javed 1980: 
fig. 2E for P. bengalensis; Lewinsohn 1982: 
fig. 1f for P. integrirostris), while it is a thin 
scallop with a very small median process in 
P. caribensis (Fig. 1b). The male pleopods 
on the first abdominal somite are absent 
from Phylladiorhynchus, but present on the 
Caribbean species. At my request, F. A. 
Chace, Jr. of the Smithsonian Institution 
examined the types of P. caribensis; he wrote 
to me that “‘there definitely is a pair of ap- 
parently uniramous pleopods inserted lat- 
erally on the first abdominal somite but they 
are soft and flexible, not stiffened to perform 
as gonopods like those in lobsters and crabs,” 
(Chace, in litt.). 

Anomoeomunida is also related to the 
eastern Pacific Pleuroncodes Stimpson, 
1860, which contains two species: P. mon- 
odon(H. Milne Edwards, 1837) and P. plan- 
ipes Stimpson, 1860. Comparative material 
from the Smithsonian Institution of these 
two species was examined: P. planipes—4 6 
(12.3-13.3 mm in carapace length excluding 
rostral spine), 1 ovig. 2 (14.0 mm), USNM 
81336, pelagic at 20-40 miles off shore, 
southern Lower California (Magdalena Bay), 
Mar 1940, coll. & id.,E.. F. Ricketts; P. 
monodon—2 6 (13.7, 16.3 mm), 2 2 (14.9, 
17.3 mm), USNM 65710, “Albatross” Sta. 
3396, off Panama, 07°32'N, 78°36'W, 474 


104 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Anterior part of carapace: a, Pleuroncodes planipes Stimpson, male (USNM 81336), dorsal view; b, 
same, ventral view; c, Pleuroncodes monodon (H. Milne Edwards), female (USNM 65710), dorsal view. Scales 


= 1mm. 


m, 11 Mar 1891, id. J. E. Benedict [these 
specimens were reported by Faxon (1895: 
72) as P. monodon?}. 

Haig (1955:34) suggested that P. mono- 
don and P. planipes may be identical but 
this seems not to be so. The specimens of 
P. planipes examined have fringes of long 
setae on the pereopods, especially on the 
walking legs which suggests a pelagic life as 
mentioned by Faxon (1895:72), whereas 
those of P. monodon do not; also the spi- 
nation of the pereopods is less pronounced 
in P. planipes. In P. planipes, the third tho- 
racic sternite is relatively wide, with its lat- 
eral margins convergent posteriorly, while 
in P. monodon it is narrow, with an acute 
anterolateral process directed straight for- 
ward. 

These specimens share the following 
characteristics: the pterygostomial flap (or 
linea anomurica) is largely visible from a 
dorsal view (see Faxon 1895:pl. 15: fig. 3; 
Haig 1955:fig. 7); the antennular basal seg- 
ment bears three prominent terminal spines 
(mesial, lateral, and dorsolateral) and an ad- 
ditional small spine proximal to the dor- 
solateral one (Fig. 2b), as typical in Munida; 


the basal antennal segment is perfectly fused 
with the orbit and bears a well-developed, 
basally wide ventromesial spine (Fig. 2a—c); 
the rostrum is spiniform and dorsally ridged 
on the midline (Fig. 2a, c); male gonopods 
are present on the first and second abdom- 
inal segments; the dactyli of the walking legs 
bear a row of distinct but fine ventral spines. 

Of these characteristics, Anomoeomunida 
and most of the known species of Munida 
share the presence of two pairs of gonopods 
and the dorsally ridged spiniform rostrum. 
The basal segment of the antennal peduncle 
is fused with the orbit in both Pleuroncodes 
and Anomoeomunida, and even in Munida 
speciosa von Martens, 1878. 

Pleuroncodes differs most obviously from 
Anomoeomunida in the pterygostomial flap 
largely visible in dorsal view, the dactyli of 
the walking legs bearing a row of small ven- 
tral spines, and the lateral limit of the orbit 
not angled. 


Acknowledgments 


I thank R. B. Manning for the loan of the 
type and comparative materials in the col- 


VOLUME 106, NUMBER 1 


lection of the Smithsonian Institution, and 
F. A. Chace, Jr. for examining the types of 
Phylladiorhynchus caribensis at my request. 
The manuscript benefited from suggestions 
by M. de Saint Laurent and reviews by J. 
Haig, R. Lemaitre, G. C. B. Poore, and A. 
B. Williams. 


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Faculty of Education, Kumamoto Uni- 
versity, 2-40-1 Kurokami, Kumamoto 860, 
Japan. 


PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 106-114 


TWO NEW SPECIES OF NEOCALLICHIRUS 
FROM THE CARIBBEAN SEA 
(CRUSTACEA: DECAPODA: CALLIANASSIDAE) 


Raymond B. Manning 


Abstract.—Two new shore species of Neocallichirus similar to N. grandi- 
manus (Gibbes) are described, both differing from it in having a much longer 
carpus on the major cheliped. Neocallichirus nickellae, new species, from To- 
bago, has a distinctive triangular tooth on the cutting edge of the dactylus; the 
basal tooth on the dactylus is rectangular in N. /emaitrei, new species, from 
Colombia. A key to the western Atlantic species of Neocallichirus is presented. 


Continuing studies of American calli- 
anassids (Manning & Heard 1986; Manning 
1987, 1988, 1992; Manning & Felder 1986, 
1991, 1992; Rodrigues & Manning 1992a, 
1992b) have revealed the existence of two 
new shore (sensu Briggs 1961) species of 
Neocallichirus from the Caribbean, de- 
scribed here. Their descriptions are accom- 
panied by a key to the four western Atlantic 
species of Neocallichirus. The diagnosis giv- 
en below for each species will distinguish it 
from the other western Atlantic species of 
the genus. 

The western Atlantic species of Neocal- 
lichirus can be divided into two groups based 
on the shape of the telson and the uropodal 
endopod. In one group the uropodal en- 
dopod is longer than broad and tapers dis- 
tally, and the posterior margin of the telson 
is excavate, forming distinct posterolateral 
angles. That group includes N. guara (Ro- 
drigues, 1971), N. guassutinga (Rodrigues, 
1971), N. mirim (Rodrigues, 1971), and N. 
trilobatus (Biffar, 1970). A new genus will 
be recognized for these species (Manning & 
Lemaitre 1993), and they will not be con- 
sidered in the accounts of the new species 
given below. 

In the other group, which also includes 
the type species, NV. horneri Sakai, 1988, from 
Australia, the uropodal endopod is broader 
than long and is flattened distally, and the 


posterior margin of the telson is rounded; 
that margin may be slightly indented, but 
never so much as to form distinct postero- 
lateral angles on the telson. The second group 
comprises the two new species named here 
and N. grandimanus and N. rathbunae. 

In some of the species placed in Neocal- 
lichirus by Manning & Felder (1991) the 
propodus of Mxp3 is distally emarginate, 
i.e., there is an indentation on the opposable 
margin (see Fig. 1d). This is shown by Ro- 
drigues (1971:fig. 67) for N. guara and ap- 
pears to be characteristic of both species 
described here. The indentation is present 
in N. trilobatus but is less well marked or 
even absent in N. grandimanus (see Biffar 
1971:fig. Sf, and Manning 1987:fig. 2e), N. 
rathbunae (see Manning & Heard 1986:fig. 
lb), or N. mirim (see Rodrigues 1971:fig. 
84). It is shown by Rodrigues (197 1:fig. 48) 
but not by Biffar (1971:fig. 9f) for N. guas- 
sutinga, although both figures suggest that 
the opposable margin is irregular in each of 
those species. 

All of the types have been deposited in 
the collections of the National Museum of 
Natural History, Smithsonian Institution, 
Washington, D.C. (USNM). 

Abbreviations include: Al (antennule or 
first antenna), A2 (antenna or second an- 
tenna), cl (postorbital carapace length, in 
mm), leg. (collector or collected by), m (me- 


VOLUME 106, NUMBER 1 107 


Fig. 1. Neocallichirus lemaitrei, new species, female paratype, USNM 256875, cl 18.5 mm, Isla de Baru. a, 
Carapace and frontal appendages, lateral view; b, Carapace and frontal appendages, dorsal view; c, Eye, lateral 
view; d, Mxp3, inner surface; e, Major Pl, outer surface; f, Minor Pl, outer surface; g, Abdomen; h, sixth 
abdominal somite, telson, and uropods, dorsal view. 


ters), Man (mandible), Max1-—2 (first and Family Callianassidae Dana, 1852 
second maxillae), mm (millimeters), Mxp1— Subfamily Callichirinae 
3 (first to third maxillipeds), P1—5 (first to Manning & Felder, 1991 
fifth pereopods), Plp1—5 (first to fifth pleo- Genus Neocallichirus Sakai, 1988 
pods), tl (total length, measured on midline, Neocallichirus lemaitrei, new species 
in mm). Figs. 1-3 
The measurement following the number 
of specimens is carapace length; in some Material.—Colombia: Islas del Rosario 


cases total length also is given. Segments of (10°10'N, 75°46'W), Isla del Rosario, beach 
appendages are measured dorsally. on south side, yabby pump, 17 Jul 1988, 


108 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Neocallichirus lemaitrei, new species, female paratype, cl 18.5 mm, USNM 256875, Isla de Baru. a, 
Man: b, Max: c, Max2:/d, Mxp1:> e) Mxp2: f, P2: ¢, P3:-hy P4: 1 P5545 Pips: 


leg. R. Lemaitre: 1 6 (11.6) (holotype, USNM 
256876). 

Isla de Bara [10°10’N, 75°36'W], Aug 
1986, yabby pump, leg. R. Lemaitre: 1 ¢ 
(1:7), 1 2 GSs= tl 76 mm) (paratypes, 
USNM 256875).—Isla de Bart, Ciénaga de 
Cholon, 0.5 m, yabby pump, 7 Aug 1988, 
leg. R. Lemaitre: 2 6 (8.9, 10.5), 5 non-ovig- 
erous 2 (12.4, 15.7, 17.6, 18.1, 18.4), 2 ovi- 
gerous 2 (12.3, 15.4) (paratypes, USNM 
256877). 

Diagnosis.—Carapace with 3 unarmed 
anterior projections, all obtuse. Mxp3 with 
length and height of propodus subequal; op- 
posable margin of propodus emarginate. 
Major P1 with merus lacking ventral spines, 
carpus shorter than palm, both with ventral 
serrations visible in outer view; dactylus with 
large rectangular tooth basally and smaller 
triangular tooth distal to it on cutting edge. 
Minor P1 with movable finger longer than 
palm. Uropodal exopod with dorsal plate 
shorter than ventral plate. 

Description. —Carapace lengths of adults 
to more than 18 mm, total lengths to at least 
76 mm. 


Front with 3 anterior projections, all ob- 
tuse, median falling well short of cornea, 
laterals irregular, inconspicuous. 

Eye with subterminal, hemispherical, 
darkly-pigmented cornea, situated laterally 
(pigment diffuse in some specimens); an- 
terior margin of eye with distomesial angled 
projection; eyes overreaching end of first 
segment of Al peduncle. 

Al peduncle shorter than A2 peduncle, 
with penultimate segment subequal to ter- 
minal segment. A2 peduncle with penulti- 
mate segment longer than terminal seg- 
ment. 

Mxp3 with ischium-merus subpediform; 
ischium with crista dentata on inner face; 
propodus with length and height subequal, 
opposable margin emarginate (ventral edge 
also emarginate in 1 specimen). Other 
mouthparts as illustrated (Fig. 2a—e). P1 un- 
equal and dissimilar. Major P1 with dac- 
tylus slightly shorter than palm, curved ven- 
trally, tip hooking over inner surface of fixed 
finger; cutting edge of dactylus with en- 
larged rectangular tooth basally and smaller 
triangular tooth about midlength in both 


VOLUME 106, NUMBER 1 


6 


109 


Fig. 3. Neocallichirus lemaitrei, new species, male paratype, USNM 256875, cl 11.7 mm, Isla de Baru. a, 
Pipl; c, Plp2. Female paratype, USNM 256875, cl 18.5 mm. b, Plp1l; d, Plp2. 


sexes; cutting edge of fixed finger with a few 
low teeth proximally as well as a line of 
denticles proximal to cutting edge, visible 
in outer view; palm length and height sub- 
equal, ventral margin serrated to base of 
fixed finger, serrations visible in outer view; 
carpus shorter than propodus but at least 74 
as long, higher than long, ventral margin 
with distal 74 serrated, serrations visible in 
outer view; merus narrower and longer than 
carpus, length about 1.5 times height, ven- 
tral margin evenly convex, serrated, lacking 
ventral spines. 

Minor P1 with dactylus longer than palm, 
curved ventrally, unarmed, tip crossing in- 
ner side of unarmed fixed finger (denticulate 
in smaller specimens); gape hairy (omitted 
for clarity in Fig. 1f); palm slightly higher 
than long; carpus longer than propodus, lon- 
ger than high; merus narrower than but as 
long as carpus, length less than twice height, 
smooth ventrally. 

Other pereopods as figured (Fig. 2f-1). 
Plp1-3 as figured (Figs. 2j, 3). 

Telson broader than long, subtrapezoidal, 
unarmed posteriorly, widest just posterior 
to base, posterior margin rounded, with at 
most a shallow median concavity. 


Uropodal exopod with upper edge of dor- 
sal plate, along its posterior margin, not as 
long as lower edge; endopod broader than 
long, widening posteriorly, posterior margin 
flattened. 

Remarks.—Neocallichirus lemaitrei dif- 
fers from N. rathbunae in that the frontal 
projections of the carapace are unarmed, 
lacking apical spinules, and in having the 
dorsal plate of the uropodal exopod shorter 
than the ventral plate. Neocallichirus le- 
maitrei is very similar to N. grandimanus 
(Gibbes, 1850), the most common western 
Atlantic species of the genus; Biffar (1971: 
666, under Callianassa branneri Rathbun, 
1900, a subjective junior synonym of N. 
grandimanus) noted that N. grandimanus 
was a wide-ranging species, common on the 
intertidal flats of southeastern Florida. Neo- 
callichirus lemaitrei differs from N. gran- 
dimanus in having a much longer carpus 
and a slenderer dactylus on the major che- 
liped; in N. grandimanus the carpus is less 
than half as long as the palm, whereas it is 
two-thirds as long in N. /emaitrei. Further, 
the ventral margin of both carpus and prop- 
odus are strongly serrated ventrally in WN. 
lemaitrei, with the serrations visible in outer 


110 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig.4. Neocallichirus nickellae, new species, male holotype, USNM 256879, cl 15.2 mm, Tobago. a, Carapace 
and frontal appendages, lateral view; b, Anterior part of carapace and frontal appendages, dorsal view; c, Mxp3, 
inner surface; d, Major Pl, outer surface; e, Minor P1, outer surface; f, Sixth abdominal somite, telson, and 


uropods, dorsal view. 


view, and the carpus of the minor cheliped 
(P1) is longer than the palm. In N. grandi- 
manus the carpus and propodus appear 
smooth ventrally in outer view and the car- 
pus of the minor P1 is much longer than the 
palm. This new species also differs from N. 
grandimanus in having the opposable mar- 
gin of the propodus of Mxp3 distinctly 
emarginate; the ventral margin of the prop- 
odus also is emarginate in some specimens. 

All of the material of this species was col- 
lected in the Islas del Rosario, a coral reef 
archipelago situated southwest of Cartage- 
na, Colombia. Their location is shown by 
Werding (1982:fig. 1). 

Etymology.—The species is named for my 
colleague and friend, Rafael Lemaitre, who 


collected the types and made his collections 
available for study, and who initiated stud- 
ies on the callianassid fauna of the region 
(Lemaitre & Rodrigues 1991). 


Neocallichirus nickellae, new species 
Figs. 4-6 


Material. —Republic of Trinidad and To- 
bago: Coral Garden, Buccoo Reef (11°11'N, 
60°49'W), Tobago, 28 Jul 1989, leg. Lois 
Nickell: 2 6 (14.1, 15.2) (smaller 4, t156 mm 
is paratype, USNM 256878; larger 4, tl 58 
mm, is holotype, USNM 256879). 

Diagnosis. —Carapace with 3 unarmed 
anterior projections, all low, obtuse. Mxp3 
with length and height of propodus sub- 


VOLUME 106, NUMBER 1 


111 


TA \ 


\\ 


FA 
ZI 
4 oe 


Fig.5. Neocallichirus nickellae, new species, male holotype, USNM 256879, cl 15.2 mm, Tobago, appendages. 
a, Man; b, Max1; c, Max2; d, Mxp1; e, Mxp2; f, P2; g, P3; h, P4; i, PS. 


equal; opposable margin of propodus emar- 
ginate. Major P1 with merus lacking ventral 
spines, carpus almost as long as palm, both 
with serrated ventral margins visible in out- 
er view; dactylus with large, triangular tooth 
basally on cutting edge. Minor P1 with fin- 
gers much longer than palm. Uropodal ex- 
opod with dorsal plate shorter than ventral 
plate. 

Description. —Carapace lengths of adults 
14-15 mm; total lengths 56 and 58 mm. 

Front with 3 anterior projections, all low 
and obtuse, median not extending to cor- 
nea. 

Eye with subterminal, hemispherical, 
darkly-pigmented cornea, situated laterally; 
anterior margin of eye with distomesial an- 
gled projection; eyes falling short of end of 
first segment of Al peduncle. 

Al peduncle shorter than A2 peduncle, 
with penultimate segment about * as long 
as terminal segment. A2 peduncle with ter- 
minal segment about % as long as penulti- 
mate segment. 

Mxp3 with ischium-merus subpediform; 
ischium with crista dentata on inner face; 


propodus with length and height subequal, 
opposable margin distinctly emarginate. 
Other mouthparts as illustrated (Fig. 5a-). 

P1 unequal and dissimilar. Major P1 with 
dactylus about as long as palm, curved ven- 
trally, tip hooking over inner surface of fixed 
finger; cutting edge of dactylus with large, 
triangular basal tooth and smaller lobe about 
midlength; cutting edge of fixed finger un- 
armed; palm slightly longer than high, ven- 
tral margin serrated to base of fixed finger, 
serrations visible in outer view; carpus 
shorter than propodus, slightly longer than 
high, ventral margin serrated, serrations 
visible in outer view; merus narrower than 
but as long as carpus, length less than 2 
times height, tapering distally, ventral mar- 
gin serrate, lacking ventral spines. 

Minor P1 with dactylus longer than palm, 
curved ventrally, unarmed but with a few 
denticles basally on cutting edge, tips cross- 
ing inner side of unarmed fixed finger; gape 
hairy (omitted for clarity in Fig. 4e); palm 
length and height subequal; carpus slightly 
longer than palm, longer than high (height 
equal to palm length), ventral margin 


112 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 6. Neocallichirus nickellae, new species, male holotype, USNM 256879, cl 15.2 mm, Tobago. a, Plp1; 


b, Plp2; c, Plp3; d, Appendix interna of Plp3. 


smooth; merus narrower than but as long 
as carpus, length about twice height, ventral 
margin denticulate distally. 

Other pereopods as figured (Fig. 5f-1). 
Plp1-3 as figured (Fig. 6). 

Telson broader than long, subtrapezoidal, 
unarmed posteriorly, widest just posterior 
to base, posterior margin rounded, with at 
most a shallow median concavity. 

Uropodal exopod with upper edge of dor- 
sal plate, along its posterior margin, not as 
long as lower edge. 

Remarks. —Neocallichirus nickellae re- 
sembles N. /emaitrei and differs from N. 
grandimanus in having the opposable mar- 
gin of the propodus of Mxp3 indented or 


notched and in having a much longer carpus 
on the major cheliped; in N. nickellae the 
carpus is almost as long as the palm, where- 
as it is less than half as long as the palm in 
N. grandimanus and two-thirds as long in 
N. lemaitrei. As in N. lemaitrei, the carpus 
and palm of the major cheliped are serrated 
ventrally, and these serrations are visible in 
outer view; the ventral surface of the carpus 
and palm of N. grandimanus appear smooth 
in outer view. The major chela of N. nick- 
ellae differs from that of both N. lemaitrei 
and N. grandimanus in having a large, tri- 
angular tooth on the cutting edge of the dac- 
tylus. Finally, N. nickellae differs from N. 
rathbunae in lacking sharp frontal projec- 


VOLUME 106, NUMBER 1 


tions and in having the dorsal plate of the 
uropodal endopod shorter than the ventral 
plate. 

The western Atlantic species of Neocal- 
lichirus can be distinguished as follows: 


Key to Western Atlantic Species of 
Neocallichirus 


1. Frontal projections each armed with 
spinule. Upper plate of uropodal ex- 
opod as long as lower plate. Merus 
of major Pl with erect spines on 
ventral margin 

N. rathbunae (Schmitt, 1935); southern 
Florida and Caribbean (Biffar 
1971, Manning & Heard 1986) 

— Frontal projections lacking anterior 
spinule. Upper plate of uropodal ex- 
opod shorter than lower plate. Me- 
rus of major P1 lacking erect spines 
on ventral margin 

2. Carpus of major P1 less than half as 
long as propodus. Opposable mar- 
gin of Mxp3 propodus usually lack- 
ing notch or indentation ......... 

N. grandimanus (Gibbes, 1850); 
Bermuda, southern Florida and 
Caribbean to Brazil (Biffar 1971). 

— Carpus of major P1 more than half 
as long as propodus. Opposable 
margin of Mxp3 propodus with 
notch or indentation 

3. Dactylus of major P1 with large, tri- 
angular tooth basally on cutting edge 

.... N. nickellae, new species; Tobago 

— Dactylus of major cheliped with 
large, rectangular tooth basally on 
SUS er 

.. N. lemaitrei, new species; Caribbean 
coast of Colombia 


ee © © © © © © © ew ee ee ell 


Acknowledgments 


Studies of American callianassids have 
been supported by the Smithsonian Field 
Station at Link Port, Florida. This is con- 
tribution number 297 from that station. I 


113 


thank Ms. Lois Nickell, University Marine 
Biological Station, Millport, Scotland, and 
my Smithsonian colleague Rafael Lemaitre 
for allowing me to work with material col- 
lected by them. The figures were prepared 
by my wife Lilly. 


Literature Cited 


Biffar, T. A. 1970. Three new species of callianassid 

shrimp (Decapoda, Thalassinidea) from the 

western Atlantic. — Proceedings of the Biological 

Society of Washington 83(3):35-49. 

. 1971. The genus Callianassa (Crustacea, De- 

capoda, Thalassinidea) in south Florida, with 

keys to the western Atlantic species. — Bulletin 
of Marine Science 21(3):637-715. 

Briggs, J. C. 1961. The East Pacific Barrier and the 
distribution of marine shore fishes. — Evolution 
15(4):545-554. 

Dana, J.D. 1852. Macroura. Conspectus Crustaceo- 
rum &. Conspectus of the Crustacea of the Ex- 
ploring Expedition under Capt. C. Wilkes, 
U.S.N.—Proceedings of the Academy of Nat- 
ural Sciences of Philadelphia 6:10-28. 

Gibbes, L.R. 1850. On the carcinological collections 
of the United States, and an enumeration of the 
species contained in them, with notes on the 
most remarkable, and descriptions of new spe- 
cies. — Proceedings of the American Association 
for the Advancement of Science, 3rd meeting: 
167-201. 

Lemaitre, R., & S. de A. Rodrigues. 1991. Lepidoph- 
thalmus sinuensis: a new species of ghost shrimp 
(Decapoda: Thalassinidea: Callianassidae) of 
importance to the commercial culture of pe- 
naeid shrimps on the Caribbean coast of Co- 
lombia, with observations on its ecology.—Fish- 
ery Bulletin, U.S. 89(4):623-630. 

Manning, R. B. 1987. Notes on western Atlantic Cal- 

lianassidae (Crustacea: Decapoda: Thalassinid- 

ea).— Proceedings of the Biological Society of 

Washington 100:386—-401. 

1988. The status of Callianassa hartmeyeri 
Schmitt, 1935, with the description of Coralli- 
anassa xutha from the west coast of America 
(Crustacea, Decapoda, Thalassinidae).—Pro- 
ceedings of the Biological Society of Washington 
101:883-889. 

1992. A new genus for Corallianassa xutha 
Manning (Crustacea: Decapoda: Callianassi- 
dae). — Proceedings of the Biological Society of 
Washington 105:571-574. 

, & D. L. Felder. 1986. The status of the cal- 

lianassid genus Callichirus Stimpson, 1866 


114 


(Crustacea: Decapoda: Thalassinidea).—Pro- 
ceedings of the Biological Society of Washington 
99:437-443. 

——,, & 1991. Revision of the American 
Callianassidae (Crustacea: Decapoda: Thalas- 
sinidea). — Proceedings of the Biological Society 
of Washington 104:764-792. 

——,& . 1992 [1991]. Gilvossius, a new ge- 
nus of callianassid shrimp from the eastern 
United States (Crustacea: Decapoda: Thalassi- 
nidea). — Bulletin of Marine Science 49(1—2):558- 
561. 

—.,&R.W. Heard. 1986. Additional records for 

Callianassa rathbunae Schmitt, 1935, from 

Florida and the Bahamas (Crustacea: Decapoda: 

Callianassidae).— Proceedings of the Biological 

Society of Washington 99:347-349. 

, & R. Lemaitre. 1993. Sergio, a new genus of 

ghost shrimp from the Americas (Crustacea: De- 

capoda: Callianassidae).— Nauplius (Brazil) Gin 
press). 

Rathbun, M. J. 1900. The decapod and stomatopod 
Crustacea. Results of the Branner-Agassiz Ex- 
pedition to Brazil, 1.— Proceedings of the Wash- 
ington Academy of Sciences 2:135-155, pl. 8. 

Rodrigues, S. de A. 1971. Mud shrimps of the genus 
Callianassa Leach from the Brazilian coast 


(Crustacea, Decapoda). — Arquivos de Zoologia, _ 


Sao Paulo 20(3):191-223. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


, & R. B. Manning. 1992a. Two new calli- 

anassid shrimps from Brazil (Crustacea: Decap- 

oda: Thalassinidea).— Proceedings of the Bio- 

logical Society of Washington 105:324—330. 

——, & . 1992b. Poti gaucho, a new genus 
and species of ghost shrimp from southern Bra- 
zil (Crustacea: Decapoda: Callianassidae).— 
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Sakai, K. 1988. A new genus and five new species of 
Callianassidae (Crustacea: Decapoda: Thalas- 
sinidea) from northern Australia.—The Beagle, 
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Schmitt, W. L. 1935. Mud shrimps of the Atlantic 
coast of North America.—Smithsonian Miscel- 
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Werding, B. 1982. Porcellanid crabs of the Islas del 

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Department of Invertebrate Zoology, Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. 20560, 
U.S.A. 


PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 115-121 


CAPRELLA ARIMOTOI, A NEW SPECIES 
(CRUSTACEA: AMPHIPODA: CAPRELLIDEA) 
FROM THE SETO INLAND SEA, JAPAN 


Ichiro Takeuchi 


Abstract.—Caprella arimotoi is described based on the material collected 
from the red alga Pterocladia capillacea (Gmelin) on the Mukaishima Island 
in the Seto Inland Sea. The new species is close to C. verrucosa Boeck, 1872, 
but differs in having few plumose setae on antenna II, forwardly vented pro- 
jection on head and elongate gills. Caprella (Spinicephala) pseudoverrucosa 
(nomen nudum) mentioned in Arimoto’s essay of 1978 is synonymous with 


the present species. 


Caprella is the largest genus of the sub- 
order Caprellidea (Crustacea: Amphipoda), 
widely distributed from temperate to boreal 
regions occurring primarily on seaweeds, 
seagrasses, and hydroids. So far about 130 
species of this genus have been reported (e.g., 
McCain 1968; Laubitz 1970, 1972; McCain 
& Steinberg 1970; Vassilenko 1974; An- 
moto 1976; Takeuchi 1989). During my 
short visit to the Mukaishima Marine Bi- 
ological Station of Hiroshima University in 
June 1989, the author found numerous ma- 
ture individuals of Caprella on the red alga 
Pterocladia capillacea (Gmelin) in the sub- 
tidal zone. A close examination of those ma- 
terials revealed that some of them are iden- 
tical with what has been called ““young male”’ 
of C. (Spinicephala) verrucosa Boeck, 1872 
in Arimoto (1976) and with C. (S.) pseu- 
doverrucosa in Arimoto (1978). In this pa- 
per, the specimens are described as a new 
species. 

The type specimens have been deposited 
in the National Science Museum in Tokyo 
(NSMT), National Museum of Natural His- 
tory in Washington, D.C. (USNM), and Ca- 
nadian Museum of Nature in Ottawa 
(NMCC). The definition of mature stages in 
females was referred to Takeuchi & Hirano 
(1991). 


Caprella arimotoi, new species 
Figs. 1-3 


Caprella (Spinicephala) verrucosa, Arimo- 
to, 1976, 122-129 (in part), fig. 67-D. (non 
Caprella verrucosa Boeck, 1872) 

Caprella (Spincephala [sic.]) pseudoverru- 
cosa Arimoto, 1978, 14, fig. 7C. (nomen 
nudum) 


Material examined.—Holotype (NSMT 
11191), male from Pterocladia capillacea 
(Gmelin) Bornet & Thuret found at the 
highest level of subtidal zone, Mukaishima 
Island, Seto Island Sea (34°22'N, 133°13’E), 
June 6, 1989, coll. I. Takeuchi. Allotype 
(NSMT 11192), female collected together 
with holotype. Paratypes: NSMT 11193 (14 
males and 4 premature females), USNM 
251762 (10 males and 1 mature and 2 pre- 
mature females), and NMCC 1992-0603 (10 
males and | mature and 3 premature fe- 
males), all collected together with holotype. 
Arimoto’s private collection No. 877-8, 1 
male from Sargassum sp., Tsushima Is- 
lands, December 1946 (?). 

Diagnosis.—Head with triangular for- 
wardly pointing projection above eye. Pere- 
onites II to VI each with 1 or 2 rounded 
dorsal projections. Antenna II of large male 
with 4—5 pairs of plumose setae on each of 


116 


peduncular segments II and III. Basis of 
gnathopod II shorter than half of pereonite 
II; propodus oval, palm with pointed pro- 
jection near proximal end and shallow tri- 
angular projection near distal end. Gills 
elongate, small. Propodus of pereopods V 
to VII each with a pair of proximal grasping 
spines. 

Description. —Holotype (Male; Figs. 1A, 
2A—F’, 3A—H). Body length 6.97 mm; length 
of head 0.45 mm; length of pereonites I to 
VII 0.34, 1.42, 1.37, 1.23, 1.05, 0.66, and 
0.47 mm, respectively. Head anteriorly 
round; dorsal projection curved and point- 
ed forward from posterior end. Pereonite I 
with small posterodorsal projection; pere- 
onites II to IV each bearing a mid-dorsal 
and a posterodorsal blunt rounded projec- 
tion. Pereonite V with a minute anterolat- 
eral and a mid-dorsal blunt rounded pro- 
jection. Pereonite VI with a mid-dorsal 
triangular projection. 

Antenna I about 2 of body length. Pe- 
duncular segments longer in the order of II, 
I, and III; flagellum composed of 11 seg- 
ments and somewhat longer than peduncle. 

Antenna II about 7% length of antenna I. 
Peduncular segments III and IV fringed with 
5 pairs of plumose setae and with 5 pairs 
and a plumose setae, respectively; flagellar 
segment I with 4 pairs of short plumose 
setae; flagellar segment II with a short plu- 
mose setae followed by 3 setae on distal end. 

Gnathopod I with propodus twice as long 
as width; palm serriform, setose with a pair 
of grasping spines near proximal end. 

Gnathopod II with vestigial coxa insert- 
ing '4 from anterior margin of pereonite II. 
Basis about % of pereonite II. Propodus 
oval, % of pereonite II. Palm with 2 pro- 
jections; proximal one 4 from proximal end, 
pointing distally, and carrying 4 setae on 
basal part; distal one low triangular and 
sparsely setose. Dactylus stout, distal 7% of 
inner margin serrated. 

Gills small, elongate, 3 times longer than 
width. 

Pereopods V to VII with vestigial coxae. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Basis of pereopod V with laterally expanded 
projection carrying granulations; merus ex- 
panded on outer side; palm of propodus 
convex, coarsely setose, with a pair of grasp- 
ing spines on a small knob near proximal 
end. Features of pereopods VI and VII close 
to that of pereopod V. Pereopods V to VII 
longer progressively; pereopod VII especial- 
ly large, twice as long as pereopod VI. 

Abdomen. Distal segment of appendage 
oval, shallowly divided into 3 apical teeth; 
basal segment with 5 or 6 setae surrounding 
distal segment. Lobes bearing several long 
setae. Penes medial. 

Mouthparts. Inner plate of maxilliped 
round and distally expanded with 2 spini- 
form setae on distal margin and a facial row 
of several plumose setae; outer plate, sub- 
equal to inner plate, round and bearing 2 
curved spiniform setae and 3 long setae on 
inner margin; segment II of palp with 
scarcely setose inner margin; segment III 
expanded distally, scarcely setose on lateral 
face; segment IV, longer than III, with sharp 
claw. Outer plate of maxilla I rectangular 
and slightly curved, with 7 spiniform teeth; 
distal segment of palp rectangular with 4 
spiniform teeth on distal margin, 3 stout 
setae on distal part of inner margin, and a 
row of 5 long setae followed by a short seta 
on lateral face. Maxilla II with oval inner 
and rectangular outer plate; both with 
densely setose margins. Incisor of left man- 
dible divided into 6 teeth; /acinia mobilis 
separated into 5 teeth followed by 3 setae; 
molar with a long seta near outer edge. Right 
mandible with 5-toothed incisor, /acinia 
mobilis carrying minute teeth on middle 
margin followed by 2 setae; molar large, with 
a long seta and bushy bundle of setae. Upper 
lip finely setose. Inner lobe of lower lip 
round, finely setose on distal part. 

Female (allotype, figs. 1B, 2G-I, 31). Body 
length 5.91 mm. Head 0.43 mm. Pereonites 
I to VII 0.22, 1.12, 0.97, 0:86, 0:87) 02 
0.49, and 0.44 mm, respectively. 

Gnathopod II situated near anterior of 
pereonite II. Basis slightly shorter than 


VOLUME 106, NUMBER 1 


117 


(| 
if < 
LL C7) ) 
| = | OY 
\ B Wiens " 
o é i : 
ax Lo : WS 
A a KX : 7 @ ips ¢ 
f = IS 
i 
\ ~S 
\ ~" 
1.0mm // f 
Fig. 1. Caprella arimotoi, n. sp. from Mukaishima Island in the Seto Inland Sea. A, holotype (male), 6.97 


mm; B, allotype (female), 5.91 mm. 


of pereonite II. Propodus oval, subequal to 
basis; palm smooth, convex and setose, with 
a grasping spine near proximal end. 

Oostegite III setose on margin; oostegite 
IV minutely setose on anterior margin and 
moderately on posterior margin. Gills on 
pereonite III oval and those on pereonite 
IV smaller, elliptical. Abdomen with a pair 
of lobes without setae. 

Etymology.—The specific name, arimo- 
toi, is in honor of the late Dr. Ishitaro Ari- 
moto, who made contributions to the tax- 
onomy of the Japanese caprellidean 
amphipods and was the first person to find 
the present species. 

Localities.—Type locality: Mukaishima 
Island, Seto Inland Sea (34°22'N, 133°13’E). 
Others: Tsushima Islands, and Tateyama 


(Arimoto 1976) and Amatsu-Kominato 
(Takeuchi 1989; as Caprella sp. C), Chiba. 

Remarks.—The present new species is 
close to Caprella verrucosa Boeck, 1872 in 
having blunt dorsal projections on pereon- 
ites I to VI, short basis and oval-shaped 
propodus in gnathopod II, and grasping 
spines on pereopods V to VII. 

Caprella verrucosa was first reported from 
somewhere near San Francisco, California 
(Boeck 1872), and later recorded from both 
sides of the North Pacific; from the Queen 
Charlotte Islands, British Columbia to Santa 
Catalina Island, California (Dougherty & 
Steinberg 1953, Laubitz 1970, McCain & 
Steinberg 1970, Martin 1977, Marelli 1981), 
in the east, and from the Tsugaru Straight 
to the Kyusyu Islands along both Japanese 


118 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Pr 


4 


A: 0.20mm 
B: 0.10mm 
C-G: 0.20mm_ H, |: 0.20mm 


ee 
S33 


Fig. 2. Caprella arimotoi, n. sp. from the Mukaishima Island in the Seto Inland Sea. Holotype (male), 6.97 
mm. A, antenna II; B, gnathopod I; C, gnathopod II; D, pereopod V; D’, coxa of pereopod V; E, pereopod VI; 
E’, coxa of pereopod VI; F, pereopod VII; F’, coxa of pereopod VII. Allotype (female), 5.91 mm. G, gnathopod 
II; H, gill and oostegite on pereonite III; I, gill and oostegite on pereonite IV. 


VOLUME 106, NUMBER 1 


rr 


Fig. 3. Caprella arimotoi, n. sp. from the Mukaishima Island in the Seto Inland Sea. Holotype (male), 6.97 
mm. A, maxilliped; B, maxilla I; C, maxilla II; D, right mandible; E, left mandible; F, upper lip; G, lower lip; 


H, abdomen. Allotype (female), 5.91 mm. I, abdomen. 


coasts (Utinomi 1943, 1947, 1964; Arimoto 
1976; Takeuchi 1989) and the south coast 
of the Korean Peninsula (Kim & Lee 1975, 
Lee 1988) in the west. 

Mature males of C. arimotoi can be sep- 
arated from those of C. verrucosa from the 
Japanese coast (Utinomi 1943, 1947; Tak- 


euchi 1989) and British Columbia (Laubitz 
1970) with the following characters: 1) An- 
tenna I equal to '2 of the body length (4 in 
C. verrucosa), 2) pedunclar segments of an- 
tenna I slender, each about 4 times longer 
than width (2 to 3 times in C. verrucosa), 
3) antenna II carrying 4—5 pairs of plumose 


120 


setae on peduncular segments II and III 
(more than 8 pairs of longer plumose setae 
in C. verrucosa), 4) mid-dorsal projection 
on the head curved forward, (straight for- 
ward in C. verrucosa), 5) lacking ventrolat- 
eral projections on pereonites III to IV (C. 
verrucosa with distinct ventrolateral projec- 
tions on pereonites III and IV), 6) in C. 
verrucosa, additional paired projections 
on pereonites IV to V, and 7) gills 3 times 
longer than width (1.5 times longer in C. 
verrucosa). 

The author examined the specimens of C. 
verrucosa collected from Vancouver Island, 
British Columbia, and deposited at the Ca- 
nadian Museum of Nature (NMC 10867). 
The characters given above were found ap- 
plicable also to these Canadian specimens. 

Arimoto (1976) described two types of C. 
(Spinicephala) verrucosa from the Japanese 
coast. His robust type is identical with C. 
verrucosa as described by Utinomi (1943, 
1947) and Takeuchi (1989), while the slen- 
der type belongs to the new species. Ari- 
moto (1976) stated that the slender type rep- 
resented the young stage of C. verrucosa, 
although its body was 6.5 mm long. Later, 
in his essay (Arimoto 1978), he showed the 
lateral view of slender type labelled “‘Ca- 
prella (Spincephala [sic.]) pseudoverrucosa.” 
During the reexamination of Arimoto’s col- 
lection, a specimen labelled “‘C. pseudover- 
rucosa’”’ from the Tsushima Islands was 
found. The specimen, representing a large 
mature male of 8.79 mm long, is identified 
with C. arimotoi. This indicates that Ari- 
moto had come to the same conclusion as 
the present author that the slender type was 
not a juvenile stage of C. verrucosa, but rep- 
resented another species. His essay (Ari- 
moto 1978), however, was written in Jap- 
anese and without any taxonomic account. 
Thus, it does not constitute a valid publi- 
cation as specified by the International 
Commission on Zoological Nomenclature 
(1985; Article 8-(a)-(1) on pp. 12-13). A 
taxonomic account of C. (S.) pseudoverru- 
cosa was never prepared by Arimoto. Ca- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


prella sp. C in Takeuchi (1989) is also syn- 
onymous with C. arimotoi. 

In conclusion, Caprella arimotoi, n. sp. 
clearly differs from C. verrucosa in several 
characters, and the slender “‘young male” 
of C. (Spinicephala) verrucosa in Arimoto 
(1976) and C. (S.) pseudoverrucosa in Ari- 
moto (1978) are junior synonyms of C. ari- 
motol. 


Acknowledgments 


The author thanks Prof. Ju-shey Ho and 
Dr. M. Takeda for reviewing the manu- 
script, Dr. H. Katayama for use of the lab- 
oratory facility at the Mukaishima Marine 
Biological Station of Hiroshima University, 
Dr. S. Yamato for assistance in the field, 
and the family of the late Dr. I. Arimoto 
for the loan of the specimens. Mrs. D. R. 
Laubitz kindly arranged for my short visit 
to the Canadian Museum of Nature (for- 
merly National Museum of Natural Sci- 
ences) in Ottawa. This study was partially 
supported by the Research Aid of Inoue 
Foundation for Science, Tokyo. 


Literature Cited 


Arimoto, I. 1976. Taxonomic studies of caprellids 
(Crustacea, Amphipoda, Caprellidae) found in 
the Japanese and adjacent waters.—Special 
Publications from the Seto Marine Biological 
Laboratory, Series III:i-v + 1-229. 

. 1978. Nippon no Warekara [The Caprellidea 

of Japan].— Dobutsu to Shizen [Animal and Na- 

ture] 8:10—15 (in Japanese). 

Boeck, A. 1872. Bidrag til Californiens Amphipo- 
defauna.—Forhandlinger 1 Videnskabs-selska- 
bet 1 Christiania: 32-51. 

Dougherty, E. C., & J. E. Steinberg. 1953. Notes on 
the skeleton shrimps (Crustacea: Caprellidae) of 
California.— Proceedings of the Biological So- 
ciety of Washington 66:39-49. 

International Commission on Zoological Nomencla- 
ture. 1985. Pp. i-xx + 1-338 in W. D. L. Ride 
et al., eds., International code of zoological no- 
menclature. Third edition adopted by the XX 
general assembly of the International Union of 
Biological Sciences. University of California 
Press, Berkeley and Los Angeles, U.S.A. 

Kim, H.S., & K.S. Lee. 1975. Faunal studies on the 
genus Caprella (Crustacea: Amphipoda: Ca- 


VOLUME 106, NUMBER 1 


prellidae) in Korea.—Korean Journal of Zool- 
ogy 18:115-126. 

Laubitz, D. R. 1970. Studies on the Caprellidae 

(Crustacea, Amphipoda) of the American North 

Pacific.— National Museums of Canada, Pub- 

lications in Biological Oceanography 1:1-89. 

1972. The Caprellidae (Crustacea, Amphip- 
oda) of Atlantic and Arctic Canada.— National 

Museums of Canada, Publications in Biological 

Oceanography 4:1-82. 

Lee, K. S. 1988. Fauna of Caprellidae (Amphipoda) 
of Cheju Island and its adjacent waters, Ko- 
rea.—The Korean Journal of Systematic Zool- 
ogy, Special Issue 2:97-106. 

Marelli, D. C. 1981. New records for Caprellidae in 
California, and notes on a morphological vari- 
ant of Caprella verrucosa Boeck, 1871.—Pro- 
ceedings of the Biological Society of Washington 
94:654-662. 

Martin, D. M. 1977. A survey of the family Caprel- 
lidae (Crustacea, Amphipoda) from selected sites 
along the northern California Coast.— Bulletin 
of the Southern California Academy of Sciences 
76:146-167. 

McCain, J.C. 1968. The Caprellidae (Crustacea: Am- 
phipoda) of the Western North Atlantic. — Unit- 
ed States National Museum Bulletin 278:i-vi + 
1-147. 

—, & J. E. Steinberg. 1970. Amphipoda I. Ca- 
prellidea I. Fam. Caprellidae. Pp. 1-78 in H.- 
E. Gruner & L. B. Holthuis, eds., Crustaceorum 
Catalogus, Pars 2. Dr. W. Junk Publishers, The 
Hague, The Netherlands. 


121 


Takeuchi, I. 1989. Taxonomic and ecological studies 
of the Caprellidea (Crustacea, Amphipoda) in- 
habiting the Sargassum zone. Doctoral thesis, 
Faculty of Agriculture, The University of To- 
kyo, Tokyo, 244 pp (in Japanese). 

—,&R.Hirano. 1991. Growthand reproduction 
of Caprella danilevskii (Crustacea: Amphipoda) 
reared in the laboratory.— Marine Biology 110: 
391-397. 

Utinomi, H. 1943. Caprellids obtained in Onagawa 

Bay, northern Japan.—Science Reports of To- 

hoku University, Series 4. Biology 17:271-279. 

. 1947. Caprellidae of Japan and adjacent wa- 

ters.—Seibutsu Supplement 1:68—82 (in Japa- 

nese). 

1964. Caprellidea. Pp. 11-15, pls 1-3 in T. 
Kikuchi, ed., Fauna and flora of the sea around 
the Amakusa Marine Biological Laboratory. Part 
V. Amphipod Crustacea. Amakusa Marine Bi- 
ological Laboratory, Kyusyu University, Ku- 
mamoto, Japan (in Japanese). 

Vassilenko, S. V. 1974. [Caprellids of the seas of the 
USSR and adjacent waters.]—Opredeleliteli po 
Faune SSSR 107:1-—288 (in Russian). 


Otsuchi Marine Research Center, Ocean 
Research Institute, The University of To- 
kyo, Akahama, Otsuchi, Iwate 028-11, Ja- 
pan. 


PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 122-130 


A NEW SPECIES OF KALLIAPSEUDES 
(CRUSTACEA: TANAIDACEA: KALLIAPSEUDIDAE) 
FROM TRINIDAD 


Roger N. Bamber 


Abstract. —A new species of kalliapseudid tanaidacean, Kalliapseudes sonia- 
dawnae, is described from three specimens collected from the silty-clay benthos 
in 8-10 m of water in the Gulf of Paria, off the northwest coast of Trinidad. 
The new species is similar to congeners from the Bahamas and the Pacific coast 
of Mexico, but can be distinguished by its stouter limb and antennal articles, 
and elongate dactyli of pereopods 2 and 3. It is unique in bearing a stout 
pectinate spine on the basis of pereopod 1. 


Kalliapseudid tanaidaceans have been 
described previously from the Atlantic 
coasts of Central and South America. Lang 
(1956) established the family and described 
several species from Brazil, while most re- 
cently Sieg (1982) described Kalliapseudes 
(Mesokalliapseudes) bahamensis from the 
Caribbean. In Trinidad waters, Bacescu & 
Gutu (1975) described Discapseudes suri- 
namensis from the Caroni Swamp on the 
northern west coast. The present material 
was collected in 1982 during a study of the 
benthos off the Port-of-Spain coastal area, 
Trinidad, in the Gulf of Paria (Agard 1984). 
Three specimens of a hitherto undescribed 
kalliapseudid were taken from two separate 
sites. 

With this limitation on the quantity of 
material, only one specimen was dissected 
and mounted for microscopical examina- 
tion; the holotype (which was missing its 
antennae) and one male paratype were ex- 
amined whole; consequently features of the 
detailed anatomy (e.g., the mouthparts) re- 
late only to the success of the single dissec- 
tion and details of, for example, the epig- 
nath must await further material. The 
conspicuous distinction of the new species 
does not, however, depend on any such sub- 
tleties. 

The specimens had little sclerotization and 


long setae were commonly not straight. All 
figures were drawn with the aid of the cam- 
era lucida and represent the anatomy as it 
exists, rather than any stylized symmetrical 
interpretation. 

The type material is lodged at the Na- 
tional Museum of Wales, Cardiff, UK 
(NMW). 


Description 


Order Tanaidacea Hansen, 1895 
Suborder Apseudomorpha Sieg, 1980 
Family Kalliapseudidae Lang, 1956 
Kalliapseudes (Mesokalliapseudes) 
soniadawnae, new species 


Material. —One 2, 6 mm long, Holotype 
(NMW.Z.1991.099.1); one 6, 3.4 mm long, 
Paratype (NMW.Z.1991.099.2), both from 
Station F4, 10 m depth, 28.5°C, salinity 
20%, pH 7.35. One 6, 6 mm long, mounted 
in polyvinyl lactophenol, Paratype (retained 
in the collection of the author), Station D4, 
8.5 m depth, 26°C, 29%o, pH 6.79. These 
sampling stations were situated 2 to 3 km 
off the Diego Martin shore, Trinidad, ap- 
proximately 10°40’N 61°35’W, 9 Aug 1982, 
in silty-clay, collected by John Agard. 

Body.—(Fig. 1A) elongate (6-times as long 
as wide), unpigmented, with little scleroti- 
zation. 


Fig. 1. Kalliapseudes soniadawnae. A. Holotype female, body, dorsal. B. Paratype male, antenna 1. C. 
Paratype male antenna 2. D. Uropod, entire and E. Basal articles, holotype female. Scale line 1 mm for A and 
D, 0.2 mm for B, C and E. 


124 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Cephalon. — Wider than long, longer than 
first two pereonites together, with a distinct 


semicircular rostrum, no eyes, a single an-- 


terior dorsolateral seta and a pair of pos- 
terolateral setae on each side. Pereonites 1 
to 5 increasing in length, pereonite 6 the 
shortest; each pereonite with | anterolateral 
and | or 2 posterolateral setae, the former 
on a triangular extension on pereonites 2 to 
6. Five free pleonites, of similar length, with 
8 (pleonite 1) to 11 (pleonite 5) lateral setae; 
each pleonite bearing pleopods. Pleotelson 
(Figs. 1A, 4A) wider than long, with a pair 
of terminal setae, 3 posterior setae on each 
side of these, and 6 lateral setae on each side 
anterior to uropod insertion; 1 pair of an- 
terior dorsolateral setae. 

Male antennule.—(Antenna 1) (Fig. 1B) 
peduncle 4-articled; first article less than 3 
times as long as wide; second as long as 
wide; third and fourth wider than long, with 
setation as figured. Accessory flagellum aris- 
ing on fourth peduncle article and consisting 
of 3 articles; third article bearing 2 distal 
setae. Main flagellum of 10 articles; first as 
long as wide, succeeding articles progres- 
sively longer in relation to their width; tenth 
article 5 times as long as wide; eighth and 
tenth articles with 2 and 3 distal setae, re- 
spectively; flagellum articles 1 to 4 with 
dense rows of 6 to 10 aesthetascs distally; 
articles 5 and 6 bearing paired aesthetascs 
and article 9 with a single aesthetasc. Fe- 
male antennule basal peduncle article of 
similar proportions to that of the male; oth- 
er articles not available. 

Antenna. —(Antenna 2) (Fig. 1C) second 
article with a lateral extension bearing 4 se- 
tae, third article with an articulated “‘squa- 
ma’”’ with 5 setae; fourth article naked; fifth 
setose as figured. Flagellum of 8 articles; 
second to sixth articles with single tergal and 
1 or 2 sternal distal setae; eighth article with 
4 long and 1 short distal setae. No conspic- 
uous sexual dimorphism. 

Mouthparts.—Labrum (Fig. 2C) simple, 
setose; labium with very setose distal lobe 
(Fig. 2E) wider than long. Mandibular palps 


uniarticulate; left mandible (Fig. 2A) incisor 
process with a large distal tooth and a row 
of 5 smaller teeth appearing to connect to 
the lacinia mobilis, itself with paired distal 
teeth, lamina with 5 distal setae; right man- 
dible (Fig. 2B) with a simple incisor process 
with 2 or 3 teeth, lacinia mobilis reduced 
or absent (not seen), lamina with 5 distal 
setae. Maxilla 1 (maxillule) (Fig. 2D) inner 
endite with 4 distal setae, outer endite setose 
with a crown of 9 distal spines. Maxilla 2 
(Fig. 2F) inner lobe with 4 plumose setae 
on its outer lobe and 7 fine and 2 stout setae 
on its inner lobe; endite distally with 4 wide 
pectinate setae and 8 finer setae in 2 rows; 
outer lobe with a wide plumose seta within 
a row of 20 finer distal setae, and 3 short 
lateral spines. Maxilliped (Fig. 2G) endite 
typical for the subgenus, with a pair of cou- 
pling hooks, all outer setae plumose; inner 
edge with 3 simple setae and 6 comb-rows; 
remaining articles furnished on their inner 
margins with 2 parallel rows of plumose fil- 
tering setae; 2 distal setae on the distal ar- 
ticle. Epignath not seen in preparation. 
Cheliped. —(Fig. 4C) long, slender filter- 
ing structure typical of subgenus; basis 2.5 
times as long as wide with a single distal 
sternal seta; merus with 3 distal sternal se- 
tae, carpus with 2 sternal rows of 42 and 32 
filtering setae. Propodus slender (6 times as 
long as wide) with a row of 16 filtering setae, 
proximal ones as long as carpal filtering se- 
tae; distally 3 rostral, 1 tergal and 8 caudal 
simple setae a little longer than half length 
of dactylus; distal finger extending only 
slightly in sternal direction, half as long as 
dactylus and with a serrated terminal spine 
and 4 inner teeth. Dactylus with serrated 
distal spine, a group of 3 mid-tergal setae 
and a row of 7 sternal setae. 
Pereopods.—Pereopod 1 (P1) (Fig. 3A) 
proximal articles with few setae, but with a 
conspicuous distal tergal pectinate spine on 
the basis; distal articles with complex se- 
tation and spination (Fig. 3G); merus, car- 
pus and propodus armed distally with 1, 2 
and 3 stout spines tergally and 0, 1 and 2 


VOLUME 106, NUMBER 1 125 


Fig. 2. Kalliapseudes soniadawnae, mouthparts of paratype male. A. Left mandible. B. Right mandible. C. 
Labrum. D. Ist maxilla. E. Distal lobe of labium. F. 2nd maxilla. G. Maxilliped. Scale line is 0.2 mm. 


126 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Kalliapseudes soniadawnae, pereopods of paratype male. A to F. Pereopods | to 6, respectively. G 
to K. Distal articles of pereopods 1, 2, 5 and 6, respectively. Scale line 0.4 mm for A to F, 0.2 mm for G to K. 


VOLUME 106, NUMBER 1 | Pf 


Fig.4. Kalliapseudes soniadawnae. A. Pleotelson of holotype female, dorsal. B. Pleopod (twisted) of paratype 
male. C. Cheliped of paratype male. Scale line 0.26 mm for A, 0.2 mm for B and C. 


128 


spines sternally, respectively; dactylus blunt 
and distally heavily setose (ca. 20 setae). P2 
(Fig. 3B) with merus and carpus similarly 
proportioned, with parallel sides and nearly 
twice as long as wide; dactylus very long, 
1.4 times length of carpus and propodus 
together, with sensory “‘brush”’ at its base 
and an adjacent toothed spine; setation and 
spination of distal articles as in Fig. 3H. P3 
(Fig. 3C) similar to P2; merus subtriangular; 
carpus only 1.5 times as long as wide; dac- 
tylus 1.4 times length of carpus and prop- 
odus together. P4 (Fig. 3D) and P5 (Figs. 
3E, 3J) similar to each other; carpus distally 
with a row of 3 small spines on both caudal 
and rostral edges; propodus with 2 parallel 
combs of 7 spines of progressively increas- 
ing length from proximal (tergal) to distal 
(sternal) edges; dactylus a blunt, setose sen- 
sory organ. P6 (Figs. 3F, 3K) carpus with a 
tergal row of 5 long setae and a long distal 
seta 1.8 times length of propodus; propodus 
with 2 tergal “‘sole-”’ spines and an adjacent 
comblike row of 14 smaller spines; dactylus 
1.75 times length of propodus. 

Pleopod. —(Fig. 4B) basis with 3 setae; en- 
dopod first article with a single seta; second 
article and exopod less than twice as long 
as broad, with 11 and 12 setae, respectively. 
Uropod (Figs. 1D, 1E) biramous, almost as 
long as pleon; endopod with 3 articles, exo- 
pod with numerous articles (15 to 18), some 
irregular. 

Male specimens with conspicuous genital 
cone on sternum of pereonite six, showing 
no obvious (without dismemberment) sig- 
nificant differences in cheliped or antennal 
morphology (one would not expect the dense 
tufts of aesthetascs on the proximal articles 
of the female antennule main flagellum). All 
three type specimens had a cone-like ventral 
process on the sternum of pleonite 1. 

Etymology.—This most attractive spe- 
cies is named after the marine biologist So- 
nia Dawn Batten. 


Remarks 


Kalliapseudes soniadawnae 1s clearly close 
to both K. viridis Menzies, 1953, from the 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Pacific coast of Mexico, and K. bahamensis 
Sieg, 1982 from the Bahamas. With the for- 
mer it shares the rectangular pereonites with 
anterolateral “‘triangular’’ spine-bearing 
processes (a distinction between these de- 
scribed species emphasized by Sieg 1982) 
and a single seta on the first pleopodal en- 
dopod article; with the latter it shares few 
caudolateral setae on the telson and long 
dactyli on pereopods two and three (al- 
though these are even longer in proportion 
to the combined length of carpus and prop- 
odus in the present species— 1.4 times—than 
in K. bahamensis—1.2 times). These long 
dactyli and the distinct rounded rostrum, 
better developed than in either of the other 
two species, are distinguishing features of 
K. soniadawnae evident from whole-animal 
observation. Equally, the new species has 
stouter articles on its limbs and antennae, 
is generally less setose (although the filtering 
setae of the propodus of the cheliped are as 
long as those of the carpus in K. sonia- 
dawnae, proximally shorter K. viridis and 
in K. bahamensis), and, uniquely, a heavy 
pectinate spine on the distal tergal corner 
of the basis of pereopod 1 (only a long sim- 
ple seta in the other two species). 

Examination of 76 paratypes of K. ba- 
hamensis, kindly loaned by the National 
Museum of Natural History, Washington 
(USNM 181901) revealed that, despite the 
description in Sieg (1982), nearly half of 
these specimens were male with genital 
cones. This allowed the examination of sex- 
ual dimorphism and variability in this spe- 
cles. 

The antennae of a male paratype of K. 
bahamensis are shown in Fig. 5. Antenna 1 
has stouter articles than those of the female, 
and bears dense tufts of aesthetascs on the 
proximal 5 articles of the main flagellum. 
In addition, the distal setae on the propodus 
of the male cheliped are elaborately pecti- 
nate, being largely simple in the female (and 
simple in the male of K. soniadawnae). There 
is no Other conspicuous sexual dimorphism. 
The proportions of the articles of antenna 
2 are similar in both sexes (as in K. sonia- 


VOLUME 106, NUMBER 1 


yi 


129 


Fig. 5. Kalliapseudes bahamensis. A. Antenna 1, male paratype. B. Antenna 2, male paratype. Scale line 0.3 


mm. 


dawnae),; it is therefore reasonable to as- 
sume that the males of K. viridis will have 
slender antenna 2 articles as found in the 
female, and distinct from the more robust 
morphology of K. soniadawnae. 

Variability in the lengths of the dactyli of 
pereopods 2 and 3 was analyzed by mea- 
suring them in comparison to the length of 
the adjacent, shorter toothed spine (see Fig 
3H). This proportion showed no significant 
difference between the two limbs. In K. ba- 


hamensis the dactyl was 3 times the length 
of the adjacent spine (mean from 10 spec- 
imens 2.95, range 2.6 to 3.4) while in K. 
soniadawnae the dactyl was 4 times the spine 
length (mean from all three specimens 4.12, 
range 3.75 to 4.45). In K. viridis this pro- 
portion is about 2 (e.g., Sieg 1982:fig. 7). 
Thus, of these three closely related spe- 
cies, K. bahamensis is immediately distin- 
guishable by its characteristic trapezoidal 
pereonite morphology, in having more than 


130 


10 sternal setae on the cheliped dactylus and 
its uropod basis exceeding the posterior tip 
of the telson by about one-third ofits length. 
The remaining two zoogeographically iso- 
lated species have rectangular pereonites, 
less than 10 sternal setae on the cheliped 
dactylus and uropod basal articles not or 
only just exceeding the telson. They are dis- 
tinguishable by the heavy spine on the basis 
of pereonite 1, the elongate dactyli of pere- 
onites 2 and 3 and the stouter antennal ar- 
ticles shown only by K. soniadawnae. 


Acknowledgments 


I am most grateful to J. Gobin for bring- 
ing the specimens to my attention and for 
supplying the data on the sample sites, to 
Janice Clark of the National Museum of 
Natural History, Washington, for the loan 
of paratypes of Kalliapseudes bahamensis, 
and to Dr. D. Holdich for access to relevant 
literature. 


Literature Cited 


Agard, J.B. R. 1984. A baseline study of the effects 
of pollution on the benthos of the nearshore 
Diego Martin to Port of Spain coastal area.— 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Technical Report, Institute of Marine Affairs, 
Trinidad, 79 pp. 

Bacescu, M., & Gutu, M. 1975. A new genus (Dis- 
capseudes n.g.) and three new species of Apseu- 
didae (Crustacea, Tanaidacea) from the north- 
eastern coast of South America.—Zoologische 
Mededelingen 49:95-113. 

Hansen, H. J. 1895. Isopoden, Cumacean und Sto- 
matopoden der Plankton-Expedition. Ergeb- 
nisse der Atlantischer Ozean Plankton Expedi- 
tion Humboldt-Stiftung 2 (G), Lipsius & Tischer, 
Kiel, 105 pp. + pls 1-8. 

Lang, K. 1956. Tanaidacea aus Brasilien, gesammelt 
von Professor Dr. A. Remane und Dr. S. Ger- 
lach.—Kieler Meeresforschungen 12:249-260. 

Menzies, R. J. 1953. The apseudid Chelifera of the 
eastern tropical and north temperate Pacific 
Ocean.— Bulletin of the Museum of Compara- 
tive Zoology 107(9):443-496. 

Sieg, J. 1980. Sind die Dikonophora eine polyphy- 

letische Gruppe?— Zoologischer Anzeiger 205: 

401-416. 

. 1982. Anmerkungen zum Genus Kalliapseu- 

des Stebbing, 1910 mit Beschreibungen einer 

neuen Art Kalliapseudes bahamensis n.sp. 

(Crustacea, Tanaidacea).— Mitteilungen aus dem 

Zoologischen Museum der Universitat Kiel 1(9): 

3-17. 


FAWLEY Aquatic Research Laborato- 
ries, Marine & Freshwater Biology Unit, 
Fawley, Southampton SO4 1TW, Hants., 
United Kingdom. 


PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 131-136 


ENTEROCOLA AFRICANUS, A NEW SPECIES 
(COPEPODA: ASCIDICOLIDAE) ASSOCIATED WITH A 
COMPOUND ASCIDIAN SYNOICUM SPECIES FROM 
NORTH AFRICA (STRAIT OF GIBRALTAR) 


Pablo J. Lopez-Gonzalez, Mercedes Conradi, and 
J. Carlos Garcia-Gomez 


Abstract. —The genus Enterocola van Beneden, 1860 is principally recorded 
from European waters. In this paper Enterocola africanus, new species, from 
the Strait of Gibraltar (North African side) is described from the compound 
ascidian Synoicum sp. Enterocola fulgens, Enterocola clavelinae, Enterocola 
hessei, Enterocola precaria and Enterocola ianthina are morphologically similar 
to the new species. All are discussed and compared with Enterocola africanus. 

Resumen. —La mayor parte de las especies descritas dentro del género En- 
terocola van Beneden, 1860 proceden de las costas Europeas. Se describe una 
nueva especie, Enterocola africanus del Estrecho de Gibraltar (vertiente Norte 
Africana). Enterocola fulgens, Enterocola clavelinae, Enterocola hessei, Enter- 
ocola precaria y Enterocola ianthina son morfologicamente proximas a la nueva 
especie. Todas ellas son comparadas y discutidas con Enterocola africanus. 


Recently the Laboratorio de Biologia Ma- 
rina of the University of Sevilla and Cadiz 
(Spain) initiated a program to study the co- 
pepods associated with marine inverte- 
brates from the coasts of the Strait of Gib- 
raltar and nearby areas. So far, three marine 
biological expeditions “Bahia 90” and “Ba- 
hia 91” in Algeciras Bay (Southern Iberian 
Peninsula) and “‘Ceuta 91” in Ceuta (North 
Africa) were carried out within a more com- 
prehensive program of marine benthos. The 
studies were centered mainly on the cope- 
pod fauna associated with molluscs, ascid- 
ians, anthozoans, and echinoderms. The first 
results have already been reported (Lopez- 
Gonzalez et al. 1992). 

Four female parasitic copepods belonging 
to the genus Enterocola were found in the 
colonies of the compound ascidian, Synoi- 
cum sp. collected during the “Ceuta 91” 
Expedition. They were later determined to 
be new to science. 

Studies of the genus Enterocola have been 
concentrated in European waters with lim- 


ited references to the North American coast 
and the Philippine Islands (Illg & Dudley 
1980). Recently, Ooishi (1987) recorded an 
undetermined species of this genus from 
Okinawa. Shellenberg (1922) reported Bo- 
tryllophilus sp. associated to Polycitor ren- 
jeri from Plattenberg Bucht (South Africa), 
and not Enterocola sp. as Illg & Dudley 
(1980) referred to in their monograph on 
the Ascidicolidae. 

In this work, Enterocola africanus from 
the North African coast (Ceuta) is de- 
scribed. It represents the only member of 
its genus described from this continent. Al- 
though it is not the only species reported, 
Barnard (1955) quotes Enterocola fulgens 
from South Africa. 


Material and Methods 


The compound ascidians were collected 
on stones from the infralittoral zone (6-12 
m deep). They were maintained in separate 
glass bottles. The copepods were removed 


132 


through dissection of hosts and preserved 
in formalin (4% in sea water). The speci- 
mens were stained with cotton blue, dis- 
sected under a stereomicroscope, and semi- 
permanent mounts were made using 
lactophenol. 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. 


Family Ascidicolidae Thorell, 1859 
Subfamily Enterocolinae Della Valle, 1883 
Genus Enterocola van Beneden, 1860 
Enterocola africanus, new species 
Figs. 1-2 


Type material. —4 °° from 3 colonies of 
Synoicum sp. at Ceuta (Spain, North Africa) 
(35°53'430’N; 15°17’W), 18 Aug 1991. The 
holotype has been deposited in the Museo 
Nacional de Ciencias Naturales de Madrid 
(Spain) (MNCNM 20.04/3504). The three 
paratypes (two dissected) in the collection 
of the authors. 

Female.—Body (Fig. la—c) of relaxed 
specimens 0.97 mm total length (based on 
4 specimens). Proportions of cephalosome: 
metasome: urosome, 1:4.2:1.48. Uncon- 
tracted specimen with dorsal sutures and 
urosome folded without segmentation. Body 
covered ventrally with discontinuous rows 
of spinules. 

Antennule (Fig. le) unimerous, apically 
narrowing suggesting 2 possible segments. 
Basal article covered with discontinuous 
rows of spinules. Distal component with 
minute conical apical protuberance. Junc- 
tion of first and second components with 
about 5 setules. Antenna (Fig. 1f) obscurely 
bimerous. Basal segment unarmed, but with 
several rows of minute spinules. Distal seg- 
ment with 1 subterminal and 5 terminal 
setae and several rows of minute spinules. 
Labrum (Fig. 1g) with 2 lateral spinose lobes; 
ventral surface with rows of spinules. Max- 
illule (Fig. 2a, b) bilobed. Basal portion la- 
melliform, somewhat trilobed distally; dis- 
tal third of anterior margin with 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


unornamented lobe (Fig. 2a, b). A promi- 
nence near articulation of palp bearing 
barbed seta and short setule. Palp with 5 
spinulose setae and 1 unornamented seta on 
its distal truncate margin. Maxilla (Fig. 2c) 
with a massive basal segment bearing an 
articulated digitiform, spinulose endite at 
its distal medial corner. Apical segment bi- 
fid distally, with one process somewhat 
shorter than the other, narrower than basal 
segment, but also heavily sclerotized. Junc- 
tion of basal and apical segments with 1 
short seta. 

Intercoxal area of leg 2—4 with pro- 
nounced mammilliform processes (Fig. la, 
b). Legs 1-4 (Fig. 2d—g) biramous. Anterior 
surface of legs bearing rows of spinules. Pro- 
topodites with small seta at distal lateral 
corner. All legs with bimerous protopodites 
and unimerous rami. Exopodites terminat- 
ing in a pointed process and slightly cufved 
laterally. Exopodite of third leg (Fig. 2f) with 
characteristic styliform outline in most spe- 
cies of the genus. Endopodites approxi- 
mately equal to exopodites; lateral margin 
more convex than medial margin; 2 apical 
setae of each endopodite set close together 
and longer than ramus. Outer apical seta 
about 1.7 times the length inner apical seta 
on all legs. Pediform projection (probably 
leg 5) (Fig. 1b) a plate with subcircular mar- 
gin, bearing 2 separate minute setules. Cau- 
dal rami (Fig. la—c) conical and apparently 
forming a definitive articulation with uro- 
some. 

Male unknown. 

Etymology.—The specific name africa- 
nus was chosen because this is the first spe- 
cies of this genus described from Africa. 


Discussion 


There are five species of Enterocola van 
Beneden, 1860 with mammiliform process- 
es present between at least one pair but not 
all pairs of legs: Enterocola fulgens van Be- 
neden 1860, Enterocola clavelinae Chatton 
& Harant 1924, Enterocola hessei Chatton 


VOLUME 106, NUMBER 1 133 


Fig. 1. Enterocola africanus, new species, female: a, habitus, ventral (A); b, habitus, lateral-oblique (A); c, 
habitus, dorsal (A); d, oral region (B); e, antennule (C); f, antenna (C); g, labrum (C). Scale bars, A: 600 um; B: 
50 um; C: 50 um. 


134 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


~~ 
— 
Tne 
pares 
ay 
= 


~ 
~= 


a geht tee, 
\ - 
Vue 

Metis Ste 
- 
i 
—— 
Ta 
yin 
So oe 
Foe 
“— 
pS Se A Aw Sn 


Z 


-~ 
~ 
- 
~ 


‘ 


eo E 


Fig. 2. Enterocola africanus, new species, female: a and b, maxillule (A), ul: unornamented lobe of the distal 


third anterior margin; c, maxilla (A); d, first leg (B); e, second leg (B); f, third leg (B); g, fourth leg (B). Scale 
bars, A: 25 wm; B: 50 um. 


VOLUME 106, NUMBER 1 


135 


Table 1.—Comparison of selected features between Enterocola clavelinae, Enterocola precaria, Enterocola 
ianthina and Enterocola africanus, new species. 


E. clavelinae 


E. precaria 


E. ianthina 


E. africanus 


small apical setules 


Antennule Unsegmented lobe Unimerous with 
with 2 anterior setae 
and 3 terminal setae 
Antenna Strongly bimerous Unimerous 
Ratio Endopodite longer §Endopodite longer 
Endopodite: than exopodite than exopodite 
exopodite 


Obscurely bimerous 
with about 7 setules 


Bimerous 


In the first, second 
and fourth legs exo- 
podite are shorter 


Unimerous with 5 
setules 


Obscurely bimerous 


Endopodite approxi- 
mately as long as 
exopodite 


Length of 2 apical 
setae 


Equal Equal 


pediform projection With single small 


setule on the margin 


Are not delimited 
from urosome 


Caudal rami 


& Harant 1924, Enterocola precaria Illg & 
Dudley 1980, and Enterocola ianthina Ulg 
& Dudley 1980. None of these species have 
a body covered ventrally with discontinu- 
ous rows of spinules like the new species. 

Enterocola fulgens has setae of antenna 
short, hooked, while E. africanus has these 
setae very long and flexible. 

The most important difference between 
E. hessei and the other species is that the 
two apical setae of each endopodite are set 
on the lateral rather than on the apical sur- 
face, and so are diverging from the axis of 
the ramus. 

Differences between the new species and 
the other three species are summarized in 
Table 1. 

The diagnostic feature of Enterocola af- 
ricanus are: presence of mammiliform pro- 
cesses at the bases of legs 2-4, antennule 
being unimerous with five setules, antenna 
obscurely bimerous with six long flexible 
setae, length ratio of endopodite : exopo- 
dite, length of two apical setae of the endo- 


With single setule 
in the middle 


Without apparent 
articulations 


than endopodite but 
in the third leg exo- 
podite is longer 
than endopodite 


Equal Outer apical seta 1.7 
times as long as 


inner 


Not element of 
armature although 
there are 4 shallow 
emarginations 


With 2 separated 
minute setules 


With apparent 
articulations 


With apparent 
articulations 


podite of the leg, armature of pediform pro- 
jection, and caudal rami articulated. 


Acknowledgments 


We thank Dr. P. L. Illg, Dr. P. L. Dudley 
and Dr. V. Gotto for their generous help 
with information and literature in the course 
of this work. Rocio Juan provided valuable 
assistance during the field work. We are 
grateful to Dr. L. Cervera for help during 
this project. 

Appreciation is extended to CEPSA, Se- 
villana de Electricidad, Excmo. Ayunta- 
miento de los Barrios, Mancomunidad de 
Municipios del Campo de Gibraltar and 
Agencia de Medio Ambiente (Junta de An- 
dalucia) for financial support of this work. 


Literature Cited 


Barnard, K. H. 1955. South African parasitic Co- 
pepoda.— Annals of the South African Museum 
41:223-312. 


136 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Beneden, van P. J. 1860. Sur un nouveau genre de 
crustacé Lernéen.—Bulletin de l’Academie de 
Belgique, série 2, 9:151—160. 

Chatton, E., & H. Harant. 1924. Notes sur les Co- 
pépodes ascidicoles. XV. Sur trois formes nou- 
velles du genre Enterocola P. J. van Beneden. 
Etat actuel de la systématique des Enterocolinae 
n. subf.—Bulletin de la Société Zoologique de 
France 49:354—-364. 

Della Valle, A. 1883. Sui copepodi che vivono nelle 
Ascidie composite del Golfo di Napoli.— Atti 
della R. Accademia dei Licei, serie 3, Memoire 
della Clase de Scienze Fisiche, Matematiche e 
Naturali 15:242-253. 

Illg, P. L., & P. L. Dudley. 1980. The family Asci- 
dicolidae and its subfamilies (Copepoda, Cyclo- 
poida) with descriptions of new species.— Mé- 
moires du Muséum National d’Histoire 
Naturelle, ser. A, T. 117:1-192. 

Lopez-Gonzalez, P. J., M. Conradi, S. Naranjo, & J. 
C. Garcia-Gomez. 1992. Anew species of An- 
thessius (Copepoda: Poecilostomatoida) asso- 
ciated with Berthella stellata Risso, 1826 (Gas- 
tropoda: Opisthobranchia).— Proceedings of the 
Biological Society of Washington 105:240-248. 


Ooishi, S. 1987. A preliminary list of copepods as- 
sociated with ascidians collected around Sesoko 
Island, Okinawa. —Galaxea 6:95-98. 

Schellenberg, A. 1922. Neue Notodelphyiden des 
Berliner und Hamburger Museums mit ein 
Ubersicht der ascidienbewohnenden Gattungen 
und Arten.— Mitteilungen aus dem Zoologi- 
schen Museum in Berlin 10:277-362. 

Thorell, T. 1859. Till Kannedomen om vissa par- 
asitiskt lefvande Entomostraceer. — Ofversigt at 
Kongl. Vetenskaps-Academiens Forhandlingar 
16, 8:335-362. 


(PJLG) (JCGG) Laboratorio de Biologia 
Marina, Departamento de Fisiologia y Bio- 
logia Animal, Facultad de Biologia. Univer- 
sidad de Sevilla, Apdo. 1095, 41080 Sevilla, 
Spain; (MC) Laboratorio de Biologia, Fa- 
cultad de Ciencias del Mar. Universidad de 
Cadiz. Apdo. 40, 11510, Puerto Real (Cad- 
iz), Spain. 


PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 137-146 


NEW SPECIES AND NEW RECORDS OF THE GENUS 
ELAPHOIDELLA (CRUSTACEA: COPEPODA: HARPACTICOIDA) 
FROM THE UNITED STATES 


Janet W. Reid and Teruo Ishida 


Abstract. —We list the first record of Elaphoidella wilsonae from New Mexico 
and new records of Elaphoidella bidens from Maryland, Tennessee, Virginia 
and the District of Columbia. Two new species of harpacticoid copepods from 
the eastern United States, Elaphoidella carterae from Virginia and Elaphoidella 
amabilis from Maryland differ from congeners in the shapes of the caudal ramus 
and caudal setae and the spine formulas of the swimming legs. We provide 
keys to the known species of Elaphoidella from North America. 


Collections of harpacticoid copepods 
from springs and streams in the District of 
Columbia, Maryland, New Mexico and Vir- 
ginia included several species, two previ- 
ously undescribed, belonging to the harpac- 
ticoid copepod genus Elaphoidella. We list 
the new records of the previously known 
species and describe the new ones. The de- 
scription of each species was authored by 
its collector. We furnish identification keys 
and a table of distinguishing characters of 
both sexes of the known North American 
species. For taxonomic examination, spec- 
imens were drawn with the aid of drawing 
tubes before dissection in lactic acid and 
after dissection in polyvinyl lactophenol 
with a little chlorazol black E added, or in 
gum-chloral medium. Lengths were mea- 
sured from the anterior tip of the rostrum 
to the end of the caudal ramus. Specimens 
were deposited in the National Museum of 
Natural History, Smithsonian Institution 
(USNM). 

Hamond (1987) returned several generic 
and subgeneric taxa including Elaphoidella 
to the synonymy of the genus Canthocamp- 
tus Westwood, 1836 s. 1. pending eventual 
revision of the family Canthocamptidae. 
However, we have employed the more fa- 
miliar genus name without wishing to imply 


recognition at the generic level of this poorly 
defined group of species. 


Order Harpacticoida G. O. Sars, 1903 
Family Canthocamptidae G. O. Sars, 1906; 
Monard, 1928; Lang, 1948 
Genus Elaphoidella Chappuis, 1929 
Elaphoidella wilsonae Hunt, 1979 


Elaphoidella wilsonae Hunt, 1979:248—253, 
figs. 1-21. 


Material examined. — 1 2, in 70% ethanol, 
Guadalupe River, Jemez National Forest, 
about 40 km NE of San Ysidro, New Mex- 
ico, about 35°45’N 106°50’W, elevation 
about 2100 m, damp moss by streamside, 
26 May 1991, col. E. Warner (USNM 
251152). 

Remarks. —The specimen from New 
Mexico agrees in all respects with the de- 
scription of females from the type popula- 
tion (Hunt 1979). 

Distribution and habitat.—The Guada- 
lupe River is a third-order stream in the 
drainage basin of the Rio Grande. This spe- 
cies was formerly known only from alkaline 
spring-fed ponds in Garfield and Rio Blanco 
Counties, Colorado, in the basins of the Col- 
orado and Green Rivers respectively. The 
find reported herein extends its known dis- 


138 


tribution some 400 km to the south and 
newly includes the Rio Grande drainage ba- 
sin. 


Elaphoidella bidens (Schmeil, 1893) 


Synonymy. —Given by Lang (1948) and 
Apostolov (1985). 

Material examined. —@, ethanol-pre- 
served, Lonaconing Creek, south of Raw- 
lins, Allegany County, Maryland, about 
39°31'N 78°54'38”"W, 2 Feb 1988, col. M. 
C. Swift (USNM 242082). 2 2, ethanol-pre- 
served, Piney Creek, Garrett County, Mary- 
land, about 39°42'21”N 78°57'45’W, 6 Apr 
1988, col. M. C. Swift (USNM 242084). 3 
2, Mountain Lake, Giles County, Virginia, 
3792 1'22”N 80°32'11”W, elevation 1181 m, 
sandy bottom at small swimming beach on 
south shore, 25 May 1990, together with E. 
carterae, col. J. W. Reid (USNM 250448). 
3 2 on 2 slides, and 4 2, 1 copepodid, eth- 
anol-preserved, near-bank sediments of 
Rock Creek, District of Columbia, just east 
of Maryland border, about 38°59’00’N 
77°903'10’W, 5 Oct 1990, col. T. Ishida 
(USNM 251796). 

Remarks. —Elaphoidella bidens is a usu- 
ally parthenogenetic species recorded from 
nearly every continent. The taxonomy of 
the two subspecies, E. bidens s. s. and E. 
bidens coronata(G. O. Sars, 1904), recorded 
from North America was discussed by Wil- 
son (1956, 1975) and Wilson & Yeatman 
(1959). More recently Apostolov (1985), 
citing morphological variations including 
the coronata-form present in topotypic pop- 
ulations of E. bidens, returned several sub- 
species to the nominate species taxon. 

North American records of E. bidens co- 
ronata reviewed by Wilson (1975) include 
Florida, Georgia, Louisiana, Minnesota, 
North Carolina, Ohio, Pennsylvania, and 
Virginia. These records include a report by 
Carter (1944) from the region of Mountain 
Lake. Subsequent records of FE. bidens s. 1. 
from North America include Coahuila, 
Mexico by Reid (1988) and New York by 
Strayer [1988 (1989)]. H. C. Yeatman col- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


lected an ovigerous female E. bidens from 
a small lake at Sewanee, Tennessee on 8 
April 1973; this is anew record for that state 
(H. C. Yeatman, pers. comm.). The new 
records herein, including the first reports 
from the District of Columbia, Maryland 
and Tennessee, are well within the known 
range of this species in North America. 


Elaphoidella carterae Reid, new species 
Figs. 1, 2 


Material examined.—Holotype 2, dis- 
sected and mounted on slide in polyvinyl 
lactophenol (USNM 251767), and paratype 
2, in 70% ethanol (USNM 251768), from 
Mountain Lake, Giles County, Virginia, 
37°21'22”N 80°32'11”W, elevation 1181 m, 
sandy bottom at small swimming beach on 
south shore, 25 May 1990, col. J. W. Reid. 

Female.—Habitus (Fig. la) cylindrical. 
Length of holotype 0.76 mm, of paratype 
0.62 mm. Cephalosome (Fig. la, b) with 
elongate ovoid nuchal organ. Hyaline fring- 
es of posterior margins ofall somites smooth. 
All somites with scattered long hairs and all 
somites except cephalosome with trans- 
verse rows of tiny hairs, surface of all so- 
mites also finely punctate as in area within 
dotted line (indicated by arrow) on lateral 
surface of pediger 2 (Fig. la). Genital seg- 
ment (Fig. la, c) with remnant of division 
visible laterally beneath integument (indi- 
cated by dotted line in Fig. 1a); ornamented 
with short transverse row of small spines 
lateral to genital field; genital field reaching 
midlength of segment. Two urosomites pos- 
terior to genital segment (Fig. la, c) each 
with one row of small spines on ventral and 
lateral margin. Anal somite (Fig. la, c-e) 
with two spines near posteroventral margin 
above each caudal ramus; anal operculum 
smooth, slightly convex. Caudal ramus (Fig. 
la, c-e) about 1.2 times longer than broad, 
ovate, with dorsal, terminally hooked lon- 
gitudinal keel extending about 7% length of 
ramus, and small subdistal medial lobe. Ra- 
mus with basally biarticulate dorsal seta in- 
serted lateral to end of keel, two lateral se- 


VOLUME 106, NUMBER 1 139 


arent 


PON nnn)! HET OM ey 


. ho) ~ 


f,g 
d,e 


a-c 


Gi 
77) Erp iyyin “ON Wiha 
ee SS, 


Fig. 1. Elaphoidella carterae Reid, new species, female, holotype (USNM 251767): a, Habitus, left lateral 
(arrow indicates detail of somitic punctations); b, Cephalosome, dorsal; c, Urosome, ventral; d, Anal somite 
and caudal ramus, left lateral; e, Part of anal somite and right caudal ramus, dorsal (somewhat compressed in 
permanent mount); f, Antennule; g, Antenna; h, Mandible; i, Maxillule (part); j, Maxilla; k, Maxilliped. Scales 
= 50 um. 


140 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Elaphoidella carterae Reid, new species, female, holotype (USNM 251767): a, Left leg 1 and coupler; 
b, Right leg 2 and coupler; c, Left leg 3 and coupler; d, Left leg 4 and coupler; e, Right leg 5 and coupler. Scale 


= 50 um. 


tae, longitudinal row of five spines slightly 
ventral to insertion of distal lateral seta, 
group of fine hairs distal to medial lobe, and 
three terminal setae. Median terminal setae 
broken in both specimens, lacking proximal 
breaking plane and ornamented with few 
spiniform setules; lateralmost terminal seta 
with bulbous base and slender tip; medi- 
almost terminal seta stout, tapering, slightly 
shorter than lateralmost terminal seta; both 
medialmost and lateralmost terminal setae 
ornamented with fine hairs. Holotype bear- 
ing long ovoid spermatophore (Fig. Ic). 
Rostrum (Fig. la, b) short, subtriangular, 


with two sensillae. Antennule (Fig. 1f) of 
eight articles, article 4 with long broad es- 
thetasc reaching past end of antennule, ar- 
ticle 8 with shorter slender esthetasc. An- 
tenna (Fig. lg) biarticulate, exopodite 
uniarticulate with four setae. Exopodite of 
mandible (Fig. lh) biarticulate, proximal 
and distal articles with one and four setae 
respectively. Maxillule (Fig. 11) partly ob- 
scured in mount, basis with three visible 
setae and long terminal claw. Maxilla (Fig. 
1j) also with three setae on basis. Maxilliped 
(Fig. 1h) prehensile, with basis broken but 
lacking seta present on some congeners. 


VOLUME 106, NUMBER 1 


Legs 1—4 (Fig. 2a—d) each with triarticu- 
late exopodite; endopodite of leg 1 triarticu- 
late, longer than exopodite; endopodites of 
legs 2-4 each biarticulate. Formula for ma- 
jor armament as follows: 


Leg 1. basis 1-1 exp 0-1; 1-1; 0,2,2 
enp 1-0; 1-0; 1,2,0 


Leg 2 basis O-1 exp 0-1; 1-1; 1,2,2 


enp 1-0; 1,2,1 

Leg 3 basis O-1 exp 0-1; 0-1; 2,2,2 
enp 0-0; 1,2,1 

Leg 4 basis O-1 exp 0-1; 1-1; 2,2,2 
enp 1-0; 2,1,1 


Major lateral spines of leg 3 exopodite ar- 
ticles 1 and 2 very large, curved posteriorly. 
More distal setae of exopodites of legs 2—4 
and endopodites of legs 3 and 4 unusually 
stout, almost spiniform. Couplers of all legs 
without ornament. 

Leg 5 (Fig. 2e), medial expansion of ba- 
soendopodite reaching less than '2 length of 
exopodite. Basoendopodite and exopodite 
each with four setae of which lateral and 
medialmost setae are very short and two 
medial setae longer, all setae stout, spini- 
form. 

No variation was observed between the 
two specimens. 

Male. —Unknown. 

Etymology. —Dr. Marjorie Estelle Carter 
collected copepods from Mountain Lake and 
its environs for nearly two decades, but pub- 
lished only two articles, one posthumously, 
from those studies (Carter 1944, Carter & 
Bradford 1972). Her collection, which ap- 
parently included numerous undescribed 
species, no longer survives (H. H. Hobbs, 
Jr., pers. comm. to JWR). It is a pleasure 
to pay tribute to Dr. Carter’s contributions 
to knowledge of American harpacticoid co- 
pepods by naming this species for her. 

Comparisons.—The form of the caudal 
ramus, especially the medial protrusion, and 
the terminal caudal setae resemble those of 
no known member of the Elaphoidella- 
group. The major setation of legs 1—4 also 


141 


Table 1.—Number of major setae and spines on 
proximal: distal articles of endopodites of legs 24 (fe- 
males) and legs 2 and 4 (males) and on basoendopod- 
ites: exopodites of leg 5 of species of Elaphoidella re- 
corded from North America. (Males of E. californica, 
E. carterae, and E. kodiakensis are unknown.) 


Species Leg2 Leg3 Leg4 _ LegS5 
Females 
amabilis |e ee te ae 0 a me ee a 
shawangunkensis i305 + O38. 54:3 
carterae 1:4 0:4 1-47 \4:4 
subgracilis [4A 4 4 
californica ieSye AES, O14 S945 
bidens, kodiakensis, 
reedi, wilsonae 135 1:6 Ae yeaeS 
Males 
shawangunkensis i — OB 0:3 
subgracilis k:3 — v, 0:4 
amabilis 13 — 0:3:4.(0:4. 
reedi, wilsonae 1:4 _ 0:37 0:4 
bidens 1:4 — 2a 0:4 


2 The leg 4 endopodite of the male of E. bidens is 


uniarticulate. 


differs from known North American species 
(Table 1). 


Elaphoidella amabilis Ishida, new species 


Material examined.—Holotype °, dis- 
sected and mounted on slide (USNM 
251799). Allotype 6, dissected and mounted 
on slide (USNM 251800). Paratypes: 2 2 
and 1 6, mounted whole together on slide 
(USNM 251798), and 2 2, in 70% ethanol 
(USNM 251797). All from perennial spring, 
southeast corner of Maryland Maintenance 
Facility, Rock Creek Stream Valley Park, 
Montgomery County, Maryland, approxi- 
mately 100 m west of boundary with Wash- 
ington, D.C., 38°59'16”N 77°03'18"W, 5 Oct 
1990, col. T. Ishida. Mounted specimens in 
gum-chloral medium. 

Female. —Habitus (Fig. 3a) cylindrical. 
Length of holotype 0.57 mm, of mounted 
paratypes 0.55 and 0.56 mm. Hyaline fring- 
es of posterior margins of all somites smooth. 
Surface of all somites rather smooth, faintly 


142 


a 


pa ee ee ese 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Elaphoidella amabilis Ishida, new species, female, b—f, holotype (USNM 251799), a, g, paratype 
(USMM 251798): a, Habitus (Somewhat compressed in permanent mount), left lateral; b, Antennule; c, Antenna; 
d, Genital field; e, Penultimate urosomite, anal somite, and caudal rami, ventral; f, Anal somite and caudal 
rami, dorsal; g, Anal somite and caudal ramus, left lateral. Scales = 100 um. 


punctate. Posterior end of seminal recep- 
tacle (Fig. 3d) reaching 7% length of genital 
segment. Penultimate urosomite (Fig. 3a, e) 
with rows of small spines on lateral and 
ventral margin. Anal somite (Fig. 3e—g) with 
transverse row of lateral spines on each side, 
and with two spines near posteroventral 
margin above each caudal ramus; anal oper- 
culum convex, with marginal comb. Caudal 
ramus (Fig. 3e-g) about 1.5 times longer 


than broad, rectangular, with dorsal, ter- 
minally hooked longitudinal keel extending 
about % length of ramus. Caudal ramus with 
basally biarticulate dorsal seta inserted lat- 
eral to end of keel, two lateral setae, each 
with transverse row of four spines at base, 
and three terminal setae. Median terminal 
seta lacking proximal breaking plane, ba- 
sally expanded with ventral knob at inser- 
tion, remaining part of seta slender, about 


VOLUME 106, NUMBER 1 


143 


Fig. 4. Elaphoidella amabilis Ishida, new species, female, holotype (USNM 251799): a, Right leg 1 and 
coupler; b, Left leg 2 and coupler; c, Right leg 3 and coupler; d, Left leg 4 and coupler; e, Left leg 5 and coupler. 
Male, allotype (USNM 251800): f, Anal somite and caudal rami, dorsolateral; g, Left leg 3 and coupler; h, Right 


leg 5. Scale = 100 um. 


1.3 times longer than urosome. Lateralmost 
terminal seta slender, ventrally curved 
proximally, base bulbous with acute dorsal 
process. Medialmost terminal seta stout 
proximally, tapering distally, about '2 length 
of lateralmost terminal seta. Medialmost and 
lateralmost terminal setae ornamented with 
fine hairlike setules, median terminal seta 
ornamented with short stiff setules. 

Rostrum short. Antennule (Fig. 3b) of 
eight articles, article 4 with long esthetasc 
reaching past end of antennule, article 8 with 
shorter esthetasc. Antenna (Fig. 3c) biartic- 
ulate, exopodite uniarticulate with four se- 
tae. 


Legs 1-4 (Fig. 4a—d) each with triarticu- 
late exopodite; endopodite of leg | triarticu- 
late, longer than exopodite; endopodites of 
legs 2-4 each biarticulate. Formula for ma- 
jor armament as follows: 


Leg 1. basis 1-1 exp 0-1; 1-1; 0,2,2 
enp 1-0; 1-0; 1,2,0 

Leg 2 basis O-1 exp O-1; 1-1; 1,2,2 
enp 1-0; 1,1,1 

Leg 3 basis O-1 exp O-1; 0-1; 2,2,2 
enp 0-0; 1,1,1 

Leg 4 basis O-1 exp O-1; 1-1; 2,2,2 
enp 0-0; 1,1,1 


Major lateral spines of leg 3 exopodite ar- 


144 


ticles 1 and 2 large, curved posteriorly. Dis- 
tal medial setae of endopodites of legs 3 and 
4 short. Couplers of all legs without orna- 
ment. 

Leg 5 (Fig. 4e), medial expansion of ba- 
soendopodite reaching less than '2 length of 
exopodite. Basoendopodite with four setae, 
lateralmost seta very short and two medial 
setae longest. Exopodite with three setae, 
medialmost seta very short. 

Male.—Length of allotype 0.52 mm, of 
paratype 0.51 mm. Urosomite 3 with one 
row of small spines on ventral and lateral 
margin. Anal somite (Fig. 4f) similar to fe- 
male, but with one spine near posteroven- 
tral margin above each caudal ramus. Cau- 
dal ramus (Fig. 4f) subrectangular, with 
dorsal keel extending about '4 length of ra- 
mus, dorsal seta biarticulate at base, two 
lateral setae each with transverse row of four 
spines at base. Median and lateralmost ter- 
minal setae without bulbous bases. 

Legs 1, 2, and 4 similar to those of female. 
Leg 3 (Fig. 4g) exopodite, major lateral 
spines of articles 1 and 2 very large, curved 
posteriorly; major setae and spines of article 
3 shorter than those of female. Leg 3 en- 
dopodite triarticulate, modified, spiniform 
process of article 2 reaching only midlength 
of exopodite article 3, article 3 with two 
short apical plumose setae. 

Leg 5 (Fig. 4h) basoendopodite reduced, 
lacking armament; exopodite slightly longer 
than broad and bearing four spines, next 
innermost spine longest. 

No variation was observed between spec- 
imens of either sex. 

Etymology.—Named for the lovely as- 
pect of the body, especially the female cau- 
dal rami. 

Remarks. —The type locality is the same 
as that of Attheyella (Mrazekiella) spinipes 
Reid, 1987. 

Comparisons.—Elaphoidella amabilis, 
like E. carterae is highly distinctive in the 
structure of the caudal rami and caudal se- 
tae. The major setation of the swimming 
legs of the female is the most reduced of 
known North American species (Table 1). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


The male resembles E. subgracilis in seta- 
tion of the swimming legs, as far as the latter 
species has been described, but differs in 
lacking a papilla on the lateral surface of the 
caudal ramus. 


Keys to Continental North American 
Species of Elaphoidella 


The following key to females of conti- 
nental North American species of Elaphoi- 
della represents a considerable departure 
from the previous key of Hunt (1979) in 
that it is based primarily on the setation of 
legs 2-4. These meristic characters are eas- 
ier to interpret than are descriptions of form 
in a key without illustrations. However there 
is always the possibility of variation in num- 
ber of setae and users should consult the 
original species descriptions. As an addi- 
tional aid, a more complete description of 
setation is given in Table 1. 

Hunt (1979) did not furnish a key to males 
of continental North American Elaphoidel- 
la. The males of E. californica, E. carterae 
and E. kodiakensis are undescribed. The 
male of E. subgracilis is incompletely de- 
scribed. 


Key to females: 


1. Leg 2 endopodite article 2 with total 


of three Setae ......... =. eee Z 
— Leg 2 endopodite article 2 with total 
of four or frve setae’... 2.2 322 eee 3 


2. Leg 3 endopodite article 2 with three 
setae .... amabilis Ishida, new species 

— Leg 3 endopodite article 2 with five 
setae 
shawangunkensis Strayer, 1988 (1989) 

3. Leg 3 endopodite article 1 with one 

medial seta, article 2 with five or six 
setae 
— Leg 3 endopodite article 1 naked, 
article 2 with four setae 
4 alone cle carterae Reid, new species 

4. Caudal ramus with small or no dor- 

sal keel, not ending in hook; hyaline 
membranes of somites smooth ... 5 

— Caudal ramus with pronounced 


oe © 2s eae 


VOLUME 106, NUMBER 1 


dorsal keel, ending in large hook; 
hyaline membranes of somites 
coarsely toothed 
eee bidens (Schmeil, 1893) s. 1. 
5. Leg 3 endopodite article 2 with 
six setae, leg 5 exopodite with five 
setae 
— Leg 3 endopodite article 2 with five 
setae, leg 5 exopodite with four setae 
eS uo le subgracilis (Willey, 1934) 
— Leg 3 endopodite article 2 with five 
setae, leg 5 exopodite with five setae 
eee oe californica Wilson, 1975 
6. Caudal ramus 2-3 times longer than 
broad, anal somite not expanded 
dorsally over caudal ramus, most of 
ramus visible in dorsal view 
— Caudal ramus slightly longer than 
broad, anal somite with postero- 
dorsal expansions, most of caudal 
ramus not visible in dorsal view 
2 wilsonae Hunt, 1979 
7. Apex of ramus with three processes, 
two of these digitiform and flexible 
kodiakensis Wilson, 1975 
— Apex of ramus with normal apical 
Sora nly .. reedi Wilson, 1975 


Key to males: 


1. Leg 5 exopodite with four setae; leg 
2 endopodite article 2 with three or 


Jl.0 LEE 2 See ee 2 
— Leg 5 exopodite with three setae; 

leg 2 endopodite article 2 with three 

SE Lt SESS eee shawangunkensis 
2. Leg 2 endopodite 2 with three 

eee eres SU Be 3 
— Leg 2 endopodite 2 with four 

meee ee Se ee 4 


3. Caudal ramus with small distally 
directed papilla at posterior *%4 of 
Peete SUrCe 2. eck: subgracilis 

— Caudal ramus lacking ornament on 
distal % of lateral surface ...amabilis 

4. Leg 4 biarticulate, article 1 with 
none, article 2 with three setae .. 5 

— Leg 4 uniarticulate, with two setae 
Rees re 2S ASSET O. $C A bidens 

5. Caudal ramus tapering distally, 


145 


medial and lateral surfaces slightly 
expanded in dorsal view .... wilsonae 


— Caudal ramus bottle-shaped, me- 


dial and lateral surfaces incurved in 
“aD ci? 1° ¥ 2° hee reedi 


Acknowledgments 


The cordial assistance of many people 
contributed to this report. Members of the 
Department of Biology, Virginia Polytech- 
nic Institute & State University, Blacks- 
burg, conducted an excursion to Mountain 
Lake subsequent to the 1990 meeting of the 
North American Benthological Society, in 
which JWR participated. Mr. William B. 
Yeaman of the U.S. National Park Service 
facilitated collections by both of us in Rock 
Creek Park, District of Columbia and Mary- 
land, in 1990. Mr. and Mrs. William C. 
Warner and Miss Elizabeth Warner guided 
and assisted JWR with collections in New 
Mexico in 1991. Dr. Horton H. Hobbs, Jr. 
provided information on the career of Dr. 
Marjorie Estelle Carter. Dr. Michael C. Swift 
collected the specimens of Elaphoidella bi- 
dens from Maryland and donated them to 
the National Museum of Natural History. 
Dr. Harry C. Yeatman contributed a pre- 
viously unpublished record of E. bidens. 


Literature Cited 


Apostolov, A. 1985. Etude sur quelques copepodes 
harpacticoides du genre E/aphoidella Chappuis, 
1929 de Bulgarie avec une révision du genre. — 
Acta Musei Macedonici Scientiarum Natural- 
ium 17(7/145):133-163. 

Carter, M. E. 1944. Harpacticoid copepods of the 
region of Mountain Lake, Virginia (With de- 
scription of Moraria virginiana n. sp.).—Journal 
of the Elisha Mitchell Scientific Society 60:158- 
166, plates 65-67. 

—, & J. M. Bradford. 1972. Postembryonic de- 
velopment of three species of freshwater har- 
pacticoid Copepoda.—Smithsonian Contribu- 
tions to Zoology 119:1-26. 

Chappuis, P. A. 1929. Révision du genre Cantho- 
camptus Westwood (Note préliminaire).—Bu- 
letinul Societatii de Stiinte din Cluj 4:41-50. 

Hamond, R. 1987. Non-marine harpacticoid cope- 
pods of Australia. I. Canthocamptidae of the 
genus Canthocamptus Westwood s. lat. and Fi- 


146 
bulacamptus, gen. nov., and including the de- 
scription of a related new species of Cantho- 
camptus from New Caledonia.—Invertebrate 
Taxonomy 1:1023-1247. 

Hunt, G. W. 1979. Description of Elaphoidella wil- 


sonae n. sp. (Canthocamptidae: Copepoda) from 
Colorado.—Transactions of the American Mi- 
croscopical Society 98:248-253. 

Lang, K. 1948. Monographie der Harpacticiden. Vols. 
I, II. Nordiska Bokhandeln, Stockholm, 1648 
pp. 

Monard, A. 1928. Synopsis universalis generum har- 
pacticoidarum.—Zoologische Jahrbicher, Ab- 
teilung fiir Systematik, Okologie und Geogra- 
phie der Tiere 54:139-176. 

Reid, J. W. 1987. Attheyella (Mrazekiella) spinipes, 

a new harpacticoid copepod (Crustacea) from 

Rock Creek Regional Park, Maryland.—Pro- 

ceedings of the Biological Society of Washington 

100:694-699. 

1988. Cyclopoid and harpacticoid copepods 
(Crustacea) from Mexico, Guatemala, and Co- 
lombia.— Transactions of the American Micro- 
scopical Society 107:190-202. 

Sars, G.O. 1903-1911. An account of the Crustacea 

of Norway. V. Copepoda Harpacticoida. Bergen 

Museum, Bergen, 449 pp., 284 pls. 

1904. Pacifische Plankton-Crustaceen. (Er- 
gebnisse einer Reise nach dem Pacific. 
Schauinsland 1896/97.)—Zoologische Jahr- 

biicher, Abtheilung fiir Systematik, Geographie 
und Biologie der Thiere 19:629-646 + Tafeln 
33-38. 

Schmeil, O. 1893. Deutschlands freilebende Siis- 

swasser-Copepoden. II. Teil: Harpacticidae. Er- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


win Nagele Verlag, Stuttgart, 100 pp. + Tafeln 
I-VIII. 

Strayer, D. 1988 (1989). Crustaceans and mites (Ac- 
ari) from hyporheic and other underground wa- 
ters in southeastern New York.—Stygologia 
4:192-207. 

Westwood, J. O. 1836. Canthocamptus.—Parting- 
ton: British Cyclopaedia of Natural History, 
London 2:227. 

Willey, A. 1934. Some Laurentian copepods and their 
variations.— Transactions of the Royal Cana- 
dian Institute 20:77—-98 + Plates XIII-XV. 

Wilson, M. S. 1956. North American harpacticoid 

copepods. 1. Comments on the known fresh- 

water species of the Canthocamptidae. 2. Can- 
thocamptus oregonensis n. sp. from Oregon and 

California.— Transactions of the American Mi- 

croscopical Society 75:290-307. 

1975. North American harpacticoid cope- 
pods II. New records and new species of Ela- 
phoidella from the United States and Canada. — 
Crustaceana 28:125-—138. [The article is actually 

part 11 of a series.] 

—, & H.C. Yeatman. 1959. Harpacticoida. Pp. 
815-861 in W. T. Edmondson, ed., Ward & 
Whipple’s Fresh-Water Biology. John Wiley & 
Sons, New York. 


(JWR) Research Associate, Department 
of Invertebrate Zoology, NHB 163, Nation- 
al Museum of Natural History, Smithson- 
ian Institution, Washington, D.C. 20560, 
U:S.A.; (TI) 372 Irifunecho, Yoichimachi, 
Hokkaido, 046 Japan. 


PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 147-157 


NEW GENERA AND SPECIES OF DEEP-SEA 
POLYCHAETES OF THE FAMILY NAUTILINIELLIDAE 
FROM THE GULF OF MEXICO AND THE 
EASTERN PACIFIC 


James A. Blake 


Abstract.—Three new genera and species of deep-sea polychaetes are de- 
scribed. Pilargis mirasetis Fauchald, 1972 is redescribed and referred to a new 
genus Santelma. These species are tentatively referred to the family Nautili- 
niellidae that heretofore has included commensals of bivalve molluscs. One 
species, Laubierus mucronatus, was dissected from a mussel during its collec- 
tion, but the other three species were not associated with bivalves during the 
sample sorting process. Two genera and species, Miura spinosa and Santelma 
miraseta come from the eastern Pacific, while Laubierus mucronatus and Flas- 
carpia alvinae come from cold-seep communities on the Florida Escarpment. 
The genera presently assigned to the Nautiliniellidae are heterogeneous and 
diverse. These genera are compared with previously described nautiliniellids 
and with the closely related family Antonbrunnidae. 


The nautiliniellids are a small group of 
polychaetes that live as commensals or par- 
asites of deep-sea bivalve molluscs. Al- 
though only first reported in 1989, six spe- 
cies have already been described (Miura & 
Laubier 1989, 1990; Blake 1990; Miura & 
Ohta 1991), and there are probably many 
additional species awaiting discovery. In- 
dividual species tend to be small and have 
probably been overlooked by biologists ow- 
ing to their association with molluscs. The 
morphology of the prostomium and the 
types of setae have proven to be quite vari- 
able and the status of this new group is not 
fully understood. 

Four additional genera and species of this 
family have been discovered from North 
America. One species comes from mussels 
at cold-seep sites on the Florida Escarpment 
and is assigned to a new genus Laubierus. 
Another Florida Escarpment species is as- 
signed to a new genus Flascarpia. A third 
species has been identified from a low oxy- 
gen site in the Santa Maria Basin off central 
California, and is assigned to a new genus 


Miura. A fourth species from deep-water off 
Western Mexico and previously described 
as Pilargis mirasetis by Fauchald (1972) is 
redescribed and assigned to a new genus 
Santelma. 

The types of the new species are deposited 
in the collections of the National Museum 
of Natural History (USNM), Smithsonian 
Institution, Washington, D.C. 


Systematic Account 
Family Nautiliniellidae Miura & Laubier 


Diagnosis. — Bodies elongate, cylindrical, 
and smooth. Prostomium variable, with 1l- 
2 pairs of antennae, medial antenna or pa- 
pilla present or absent, or antennae entirely 
absent; eyes absent; palps lacking. Foregut 
expanded into a muscular pharynx, termed 
a proventriculus (Miura & Laubier 1989, 
1990); this pharynx partially eversible (Blake 
1990, this paper). Peristomial segment well 
developed with distinct tentacular cirri 
present in two genera (Sante/ma and Flas- 
carpia), reduced in three genera (Petrecca, 


148 


Laubierus, and Miura), and lacking in three 
other genera (Nautiliniella, Shinkai, and 
Natsushima). Parapodia subbiramous to ses- 
quiramous, with reduced notopodia, usu- 
ally with acicula; neuropodia with internal 
aciculae and one to several simple hooked 
spines. Pygidium simple, lacking append- 
ages. 


Miura, new genus 


Type species.— Miura spinosa, new spe- 
cies. Gender feminine. 

Diagnosis. —Body, widest anteriorly, ta- 
pering posteriorly. Prostomium rounded; 
antennae and eyes lacking. Pharynx weakly 
muscularized, prominent in first 3 seg- 
ments. Peristomial segment achaetous, 
smaller than following setigers; with 2 pair 
of short lobes present. Parapodia sesquira- 
mous, with thick notopodial lobe, acicula 
not observed; neuropodium with fascicle of 
5—6 simple hooks, with single acicula, some- 
times with tip protruding; ventral cirrus 
present. Pygidium unknown. 

Etymology. — This genus is named for Dr. 
Tomoyuki Miura, who was one of the first 
describers of the nautiliniellid polychaetes. 

Remarks. —This genus differs from pre- 
viously described nautiliniellids by lacking 
antennae altogether and in having sesquira- 
mous parapodia where the notopodial lobes 
are large and thick, but apparently lacking 
aciculae. Like Petrecca, Miura has a reduced 
and achaetous tentacular segment; like 
Shinkai it has several neuropodial spines 
instead of one. A single species, Miura spi- 
nosa, 18S known. 


Miura spinosa, new species 
Fig. | 


Material examined.—California: Santa 
Maria Basin, Sta. R-7, 34°52.90'N, 
121°10.30'W, May 1989, 565 m, holotype 
(USNM 148677). 

Description.—A small species, holotype 
an anterior fragment measuring 1.5 mm 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


long, 0.7 mm wide for 10 setigers. Color in 
alcohol: light tan with scattered orange pig- 
ment spots on dorsal and ventral surfaces. 
Body subcylindrical to oval in cross section, 
widest through first 6-7 setigers, then ta- 
pering posteriorly. 

Prostomium twice as wide as long, broad- 
ly rounded anteriorly (Fig. 1A); antennae 
and eyes absent. Mouth with broad ventral 
lip (Fig. 1B); pharynx weakly muscularized, 
prominent in first 3 segments (Fig. 1A). 
Peristomial segment achaetous, smaller than 
following setigers, with 2 pairs of small stub- 
by lateral lobes. Parapodia sesquiramous, 
with thick notopodial lobe lacking acicula 
(Fig. 1C); neuropodium with fascicle of 4— 
6 simple hooks; each hook sharply pointed, 
with subapical notch and boss (Fig. 1D); 
acicula present, distal end slightly protrud- 
ing on some setigers; ventral cirrus thick, 
rounded lobe. Pygidium unknown. 

Biology. — Miura spinosa was collected at 
a depth of 565 m in a sea valley in the Santa 
Maria Basin that has very fine silty sedi- 
ments with high clay content. This depth is 
also characterized by low dissolved oxygen 
concentrations (X = 0.898 ml/l, SD = 
+0.152; n = 4) in the near-bottom water. 
It is not known if hydrogen sulphide or 
methane is also present at this site. This 
specimen was not associated with a bivalve 
during the sorting process, but may have 
been washed from a host animal during pro- 
cessing. 

Etymology. —The specific name refers to 
the neuropodial spines. 

Remarks.—The shape of the setae of 
Miura spinosa is similar to Pilargis mira- 
setis, with which this species was identified 
in the MMS monitoring program where it 
was collected. Pilargis mirasetis has also 
been found to be a species of Nautilinielli- 
dae and has been referred to a new genus, 
Santelma (see below). Santelma miraseta 
differs from Miura spinosa in having short 
notopodial lobes instead of ones that are 
large and thick, in having long protruding 


VOLUME 106, NUMBER 1 


149 


Fig. 1. 
parapodium; D, neurosetae. 


aciculae instead of lacking these spines, and 
in having two distinct tentacular cirri in- 
stead of two pair of short, stubby lobes. 

Distribution. —California, upper conti- 
nental slope, 565 m. 


Genus Santelma, new genus 


Type species.—Santelma miraseta (Fau- 
chald, 1972). Gender feminine. 

Diagnosis. —Body dorsoventrally flat- 
tened. Prostomium rounded anteriorly, 
bearing 2 lateral and 1 medial antenna or 


a 


Miura spinosa (USNM 148677). A, anterior end, dorsal view; B, anterior end ventral view; C, tenth 


papilla (scar only present on S. miraseta); 
eyes and palps absent. Pharynx enlarged, 
muscular. Peristomial segment achaetous, 
bearing 2 pair of tentacular cirri. Parapodia 
sesquiramous, with prominent notopodial 
lobe bearing large internal acicula; neuro- 
podium with single large acicular protrud- 
ing acicula and fascicle of numerous, small, 
simple setae; dorsal and ventral cirri absent; 
branchiae absent. Pygidium a simple 
rounded lobe. 

Etymology. —The name for this genus is 
coined from the town of San Telmo, Mex- 


150 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Santelma miraseta (LACM-AHF 097). A, anterior end, dorsal view; B, eighth parapodium; C, middle 


parapodium; D, neurosetae from an anterior setiger. 


ico, which is the closest point of land near 
the collecting site of Santelma miraseta. 

Remarks. — The presence of two antennae 
and a third median antenna or papilla dis- 
tinguishes Santelma from other genera in 
the nautiliniellid complex. The four small 
tentacular cirri and large, protruding neu- 
ropodial acicula are also unusual. 


Santelma miraseta (Fauchald, 1972), 
new combination 
Fig. 2 


Pilargis mirasetis Fauchald, 1972:59-60, pl. 
8, figs. a-c.—Salazar-Vallejo, 1986:200, 
pl. 2, figs. 9-10. 


Material examined. —Off San Telmo, 


VOLUME 106, NUMBER 1 


Mexico, Velero IV Sta. 13744-70, 15 Jan 
1970, 18°12’00”N, 104°00’00’W, 2340 m, 
holotype (LACM-AHF 097). 

Description.—Holotype complete, only 
known specimen, 25 mm long, 2 mm wide, 
with about 110 segments. Color in alcohol: 
tan. Body generally robust, dorsoventrally 
compressed, elongate. 

Prostomium wider than long, broadly 
rounded on anterior margin; with 2 lateral 
antennae (bases only) and single median an- 
tenna or papilla (scar only), all located near 
border of prostomium and peristomial seg- 
ment (Fig. 2A); palps and eyes absent. Phar- 
ynx muscular, not everted. Peristomial seg- 
ment slightly larger than following segment, 
bearing 2 pair of short, tapering tentacular 
cirri (Fig. 2A). 

Parapodia sesquiramous, with notopo- 
dium formed into thickened lobe, distally 
prolonged in middle and posterior seg- 
ments; notopodia bearing large internal 
acicula, typically bending sharply near tip 
(Fig. 2B); dorsal cirrus absent. Neuropo- 
dium with large protruding acicula and fas- 
cicle of 18-25 simple bidentate setae (Fig. 
2B); ventral cirrus absent. Setae bidentate, 
tapering apically to fine, mucronate tip; sub- 
apical tooth blunt (Fig. 2D); bidentate setae 
mostly lost in middle and posterior para- 
podia, leaving only protruding spine (Fig. 
2C). Pygidium simple, rounded lobe; with- 
out cirri. 

Remarks. — Santelma miraseta is here re- 
moved from the Pilargidae because palps 
are absent. However, the referral of this ge- 
nus and species to the Nautiliniellidae is 
preliminary pending further review of the 
family. The presence of a medial antenna 
or papilla is new to species of the nautili- 
niellid-complex of genera. The very large 
and conspicuous protruding neuropodial 
acicula was not mentioned by Fauchald 
(1972). This acicula accompanies a cluster 
of very minute bidentate setae in anterior 
setigers. In posterior setigers, the bidentate 
setae are absent and the protruding acicula 
is the only visible seta and provides this 
species with a distinct posterior armature. 


151 


Two types of emergent neurosetae have also 
been reported for Natsushima and Laubier- 
us, but these genera are very distinct from 
Santelma (see Discussion for overall sum- 


mary). 


Laubierus, new genus 


Type species.—Laubierus mucronatus, 
new species. Gender masculine. 

Diagnosis. —Body dorsoventrally flat- 
tened, ribbonlike. Prostomium rounded an- 
teriorly, bearing a single pair of antennae; 
eyes lacking. Pharynx muscularized, form- 
ing distinctive proventriculus. Achaetous 
peristomial segment present, bearing pair of 
short cirri homologous to normal ventral 
cirrus. Parapodia sesquiramous, with acic- 
ulae in both noto- and neuropodia; setal 
fascicles limited to neuropodia; including 
large and small types of simple setae. Ven- 
tral cirrus present. Pygidium a simple lobe. 

Etymology. — This genus is named for Dr. 
Lucien Laubier, polychaete systematist, in 
recognition of his first descriptions of nau- 
tiliniellids. 

Remarks. —The appearance of the phar- 
ynx resembles the proventriculus that has 
been reported for the Japanese genera and 
species. The presence of large and small 
simple spines in the neuropodia of Laubier- 
us is similar to that of the genus Natsushima 
Miura & Laubier. In comparing these gen- 
era, Laubierus has an achaetous peristomial 
segment, large notopodial acicula, and small 
simple neurosetae, whereas Natsushima has 
no peristomial segment, thin notopodial 
acicula, and small bifid neurosetae. 


Laubierus mucronatus, new species 
Fig. 3 


Material examined. —Florida Escarp- 
ment, A/vin Dive 1756, 26°01'N, 84°55’W, 
3243 m, 17 Oct 1986, R. Lutz and G. Tien, 
observers, dissected from mussel, 1 frag- 
ment (JAB); Alvin Dive 1758, 26°01.8'N, 
84°54.9'W, 3266 m, Oct 1986, C. Wirsen 
and B. Tilbrook, observers, holotype 
(USNM 148678). 


152 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


100pm 


a A 


7 
}—- 


Fig. 3. Laubierus mucronatus (USNM 148678). A, anterior end, dorsal view; B, middle parapodium; C, 
small neurosetae; D, large neuroseta. 


VOLUME 106, NUMBER 1 


Description.—A small species, holotype 
complete, 5 mm long and 0.4 mm wide for 
34 setigerous segments; fragment larger, but 
anteriorly incomplete. Color in alcohol: 
Opaque white. Body elongate, dorsoven- 
trally flattened, tapering gradually anteri- 
orly and posteriorly. 

Prostomium wider than long, rounded 
along anterior margin, bearing single pair of 
sublateral antennae (Fig. 3A); tentacular 
segment poorly developed, only vaguely 
separated from prostomium, bearing pair of 
short cirri. Anterior part of digestive tract 
modified into proventriculus, with distinct 
musculature apparent through body wall 
(Fig. 3A). 

Parapodia sesquiramous, with notopodia 
bearing only a single, large acicula, with tip 
sometimes protruding; notopodium elon- 
gate, tapering, extended apically into nar- 
row lobe (Fig. 3B). Neuropodia longer, 
broader than notopodium, with fingerlike 
ventral cirrus; bearing large internal acicula 
and 2 types of sirmple neurosetae; 1 type 
small, numbering 15-20 in 2 rows; each with 
fringed tip from which tapering mucronate 
tip emerges (Fig. 3C); second type 1-2 large, 
falcate hooks, with each bearing delicate 
subapical fringe of fine bristles (Fig. 3D). 
Pygidium a simple lobe, lacking cirri. 

Etymology.—The specific name comes 
from the latin, mucros, referring to the fine, 
tapering point of the smaller neurosetae that 
characterize this species. 


Genus Flascarpia, new genus 


Type species.—Flascarpia alvinae, new 
species. Gender feminine. 

Diagnosis. — Body elongate, compressed, 
ventrum flattened, dorsum rounded. Pro- 
stomium with broadly rounded anterior 
margin, lacking eyes, with 2 lateral anten- 
nae. Distinct peristomial segment, bearing 
a pair of tentacular cirri. Pharynx soft, ever- 
sible. Parapodia sesquiramous, with noto- 
podial lobe containing blood loop and ap- 
parently functioning as gill; acicula absent. 


153 


Neuropodium well-developed, with large 
acicula and fascicle of large simple neuro- 
setae of 1 type; ventral cirrus present. Py- 
gidium a simple lobe, lacking cirri. 

Etymology. —The generic name is a com- 
posite of Florida and Escarpment, denoting 
the Florida Escarpment where the type spe- 
cies was collected. 

Remarks. — The notopodium lacks all se- 
tae, including the acicula, and by having a 
blood loop, apparently functions as a gill. 
Flascarpia and Miura are the only nautili- 
niellid-like genera known to lack notoacicu- 
lae and to have the notopodium modified 
into a soft, fleshy lobe. In Flascarpia, a dis- 
tinct blood loop is present, suggesting that 
it functions as a gill. In Miura, no blood 
loop was observed. The relationships of 
Flascarpia with all of the nautiliniellid-like 
genera are compared in the Discussion (see 
below). 


Flascarpia alvinae, new species 
Fig. 4 


Material examined. —Florida Escarp- 
ment, Alvin... Dive, 1754, 26°02.4'N, 
84°55.3'W, 3303 m, 15 Oct 1986, R. Carney 
and B. Hecker, observers, holotype (USNM 
148679). 

Description. —A moderate-sized species, 
holotype complete with 66 segments, mea- 
suring 21 mm long; 1 mm wide anteriorly, 
2 mm wide in middle. Body widest in mid- 
dle of body, tapering anteriorly and poste- 
riorly; body flattened ventrally, rounded 
dorsally. Pygidium a simple lobe lacking 
cirri. Color in alcohol: tan. 

Prostomium wider than long, broadly 
rounded on anterior margin; with 2 short 
antennae on lateral anterior margins of pro- 
stomium (Fig. 4A). Peristomial segment 
present, not distinctly separated from pro- 
stomium, bearing a single pair of long, fin- 
gerlike tentacular cirri (Fig. 4A). Parapodia 
sesquiramous, with notopodium reduced to 
soft lobe bearing internal blood vessel (Fig. 
4B); acicula absent. Neuropodium elongate, 


154 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Flascarpia alvinae (USNM 148679). A, anterior end, dorsal view (broken line indicates outline of 
everted pharynx); B, middle parapodium; C, D, neurosetae. 


thickened, triangular, tapering to point; 
ventral cirrus short, cirriform. A single large 
internal acicula present and fascicle of 7-8 
large protruding simple spines; each with 
curved tip and 2 thick subterminal protu- 
berances (Fig. 4C—D). Pygidum a simple 
lobe. 

Etymology. —This species is named for 
the DSRV Alvin, which was the vehicle used 
to collect specimens. 

Remarks. —The habitat of Flascarpia al- 
vinae is not known. It is likely, however, 
that this species is a commensal of bivalves 
and was washed from its host during the 
sorting process. 


Discussion 


The first published description of a nau- 
tiliniellid was by Miura & Laubier (1989), 
who described Nautilina calyptogenicola 
from a deep-sea vesicomyid clam collected 
from the Japan Trench at a depth of 5960 
m. This species was characterized by having 
two pairs of small antennae, short dorsal 
and ventral cirri, and a single, large pro- 
truding neuropodial hooked spine. The au- 
thors assigned this new genus to a new fam- 
ily, the Nautilinidae. In a subsequent paper, 
the same authors renamed the genus and 
family Nautiliniella and Nautiliniellidae, 


VOLUME 106, NUMBER 1 


because the earlier names were preoccupied 
in the Cephalopoda (Miura & Laubier 1990). 
In this same paper, the authors described 
two additional new genera and species: 
Shinkai sagamiensis Miura & Laubier from 
the bivalve, Calyptogena soyoae; and Na- 
tsushima bifurcata Miura & Laubier from 
Solemya sp. Both of these latter species were 
collected at the Hatsushima cold-seep site 
off Japan in depths of 1130-1170 m. The 
genus Shinkai was superficially similar to 
Nautiliniella, but differed in having up to 
eight hooks per neuropodium instead of one 
and in having only a single pair of antennae 
instead of two. Natsushima differed from 
the other two genera in having two types of 
neuropodial spines instead of one. Blake 
(1990) described another genus and species, 
Petrecca thyasira, from the mantle cavity of 
a thyasirid clam, Thyasira insignis collected 
from seep-like communities off Newfound- 
land at a depth of 3700 m. Petrecca differed 
from the previously described genera in 
having greatly elongated notopodia and an 
achaetous peristomial segment. Miura & 
Ohta (1991) described Shinkai longipeda 
from the mantle cavity of Calyptogena sp. 
collected at active hydrothermal vents in 
the Okinawa Trough in 1400 m. This spe- 
cies appears to be transitional between the 
genera Shinkai and Petrecca in the devel- 
opment of the notopodia. 

The new taxa described in the present pa- 
per do not agree with any of the previously 
described genera. All have an achaetous 
peristomial segment with either small or 
well-developed tentacular cirri. Miura lacks 
antennae, has a thickened achaetous noto- 
podial lobe, and a fascicle of small neuro- 
podial spines. Laubierus has two small an- 
tennae, a long notopodial lobe with a large 
acicula, and both large and small types of 
neuropodial spines. Flascarpia has two an- 
tennae, two well-developed tentacular cirri, 
a reduced notopodium that may function as 
a gill, and a fascicle of heavy neuropodial 
spines. Sante/ma has three antennae, two 


155 


pairs of tentacular cirri, a single large pro- 
truding acicula and a fascicle of very fine 
bidentate setae. 

The Nautiliniellidae now include eight 
genera, the characters of which are com- 
pared and contrasted in Table 1. The group 
is very heterogeneous and it is likely that 
the genera will need to be redefined and 
reorganized after additional species are dis- 
covered. In general, two distinct groups are 
apparent. One group lacks a peristomial seg- 
ment and includes Nautiliniella, Shinkai, 
and Natsushima. The second group has a 
peristomial or achaetous segment and in- 
cludes Petrecca, Miura, Laubierus, Flascar- 
pia, and Santelma. Among the second group, 
the “‘peristomial segment”’ of Petrecca is ac- 
tually a reduced setigerous segment that has 
a ventral cirrus and neuroacicula, but en- 
tirely lacks a notopodium. In Laubierus, the 
““peristomial segment”’ is also a reduced seg- 
ment that bears a ventral cirrus, but no acic- 
ula. Thus, the presence or absence of a peri- 
stomial segment appears to depend upon 
the degree to which the first setiger is re- 
duced and this undoubtedly contributes to 
the variability in this character that is ex- 
hibited by the different genera that have been 
described. All genera are characterized by 
having simple neuropodial spines, and ex- 
cept for Santelma, these setae are usually 
heavier and of a different form than those 
of the closely related family Pilargidae. 

All of these new species are tentatively 
assigned to the Family Nautiliniellidae based 
on absence of notosetae, presence of simple 
spinous neurosetae, absence of anal cirri, 
and lack of palps. None of these species 
agrees with any genus of the Pilargiidae al- 
though they appear to be closely related. 
The nautiliniellids are also related to An- 
tonbruunia viridis Hartman & Boss (1965) 
from a bivalve dredged in the Mozambique 
Channel off Madagascar in 80-90 m. This 
genus and species was assigned to a separate 
family, the Antonbruunidae by Fauchald 
(1977) and to the Pilargidae by Salazar-Va- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


156 


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VOLUME 106, NUMBER 1 


llejo (1986). The characteristics of Anton- 
bruunia are compared with those of the var- 
ious nautiliniellids in Table 1. The setal 
characteristics of A. viridis are more similar 
to those of the Nautiliniellidae than to the 
Pilargidae (See also Miura & Laubier 1990) 
and I prefer to follow Fauchald (1977) in 
placing Antonbrunnia in its own family. 

All members of the Nautiliniellidae are 
believed to be associated with the mantle 
cavities of deep-sea bivalve molluscs. There 
is insufficient data to determine if the worms 
are commensals or parasites with the clams. 
Four nautiliniellid species are associated 
with bivalves at cold-seep communities 
(Miura & Laubier 1989, 1990; Blake 1990, 
this paper), while a fifth comes from a bi- 
valve at an active hydrothermal vent (Miura 
& Ohta 1991). The Santa Maria Basin spec- 
imen was found in an upper slope sea valley 
at a depth of 565 m in an area of low dis- 
solved oxygen. The specimen was not as- 
sociated with a bivalve when it was sorted 
from the mud, but may have been washed 
from a bivalve in the same sample. The 
exact habitats of Santelma miraseta and 
Flascarpia alvinae are not known. Infesta- 
tion rates of nautiliniellids in bivalves may 
be fairly high when the worms are present. 
For example, out of ten specimens of Thy- 
asira insignis collected from off Newfound- 
land, five were found with a specimen of 
Petrecca thyasira (Blake 1990). These clams 
were each relatively small, ranging from 29 
x 30 cm to 37 X 43 cm in width x length. 
The worms themselves were up to 16 mm 
long and thus occupied a considerable space 
among the gill filaments of the clams. In- 
festation data is not available for other spe- 
cies. 


Acknowledgments 


This study was funded in part by the Pa- 
cific Outer Continental Shelf Region of the 
Minerals Management Service (MMS), U.S. 
Department of the Interior, Washington, 
D.C. under Contract No. 14-35-0001- 
30484. The specimen of Miura spinosa was 


ey) 


collected as part of the MMS Phase II Mon- 
itoring Program in the Santa Maria Basin. 
The specimens of Laubierus mucronatus and 
Flascarpia alvinae were provided by Dr. 
Barbara Hecker, Lamont-Doherty Geolog- 
ical Observatory. The holotype of Pilargis 
mirasetis was provided by Dr. Kirk Fitz- 
hugh of the Los Angeles County Museum 
of Natural History (LACM-AHF). The 
manuscript was greatly improved by in- 
sightful comments provided by Dr. Chris 
Glasby and an anonymous reviewer. 


Literature Cited 


Blake, J. A. 1990. A new genus and species of Poly- 
chaeta commensal with a deep-sea thyasirid 
clam.— Proceedings of the Biological Society of 
Washington 103:681-686. 

Fauchald, K. 1972. Benthic polychaetous annelids 
from deep water off western Mexico and adja- 
cent areas in the eastern Pacific Ocean. — Allan 
Hancock Monographs in Marine Biology No. 
7:1-575, 69 pls. 

. 1977. The polychaete worms. Definitions and 

keys to orders, families and genera.— Natural 

History Museum of Los Angeles County, Sci- 

ence Series 28:1-188. 

Hartman, O., & K. J. Boss. 1965. Antonbruunia viri- 
dis, a new inquiline annelid with dwarf males, 
inhabiting a new species of pelecypod, Lucina 
fosteri, in the Mozambique Channel.— Annals 
& Magazine of Natural History, series 13, 8:177- 
186. 

Miura, T., & L. Laubier. 1989. Nautilina calypto- 
genicola, a new genus and species of parasitic 
polychaete on a vesicomyid bivalve from the 
Japan Trench, representing a new family Nau- 
tilinidae.— Zoological Science 6:387-390. 

—_, & 1990. Nautiliniellid polychaetes 
collected from the Hatsushima cold-seep site in 
Sagami Bay, with descriptions of new genera 
and species. — Zoological Science 7:319-325. 

—, & S. Ohta. 1991. Two polychaete species 
from the deep-sea hydrothermal vent in the 
Middle Okinawa Trough.— Zoological Science 
8:383-387. 

Salazar-Vallejo, S. I. 1986. Pilargidae (Annelida: 
Polychaeta) de Mexico: listade especies, nueva 
especie y biografia.— Cahiers de Biologie Marine 
27:193-209. 


Science Applications International Cor- 
poration, 89 Water Street, Woods Hole, 
Massachusetts 02543, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 158-181 


TAXONOMY OF EUROPEAN SPECIES OF 
AMPHIDUROS AND GYPTIS 
(POLYCHAETA: HESIONIDAE) 


Fredrik Pleijel 


Abstract. —The taxonomy of the European species of Amphiduros Hartman, 
1959, and Gyptis Marion & Bobretzky, 1875, is reviewed. Amphiduros is re- 
corded for the first time from European waters. Amphiduros fuscescens (Maren- 
zeller, 1875), new combination (originally described in Oxydromus), G. pro- 
pinqua Marion & Bobretzky, 1875, and G. rosea (Malm, 1874) are redescribed 
based on available types and newly collected specimens from the vicinity of 
the type localities, and A. fuscescens is removed from synonymy with G. pro- 
pinqua. Two new species are described: G. mackiei from Sweden and the Faroes, 
unique within the genus in having setae from segment 4 rather than segment 
5, and G. mediterranea from southern France and Sicily, a species close to G. 
rosea. A. key to the European species is included, and a checklist for species 
and subspecies described in or later referred to the two genera is appended. 


The genus Amphiduros Hartman, 1959, 
is only known for four species, all described 
from the Pacific. Examination of type and 
newly collected Mediterranean specimens 
of Oxydromus fuscescens Marenzeller, 1875, 
indicates that this species also belongs to 
Amphiduros, and constitutes the first record 
of the genus from Europe. Previously O. 
fuscescens has been treated as a junior syn- 
onym of G. propinqua Marion & Bobretzky, 
1875 (Fauvel 1923, Hartman 1959). 

Two European species of Gyptis Marion 
& Bobretzky, 1875 sensu stricto (excluding 
Podarkeopsis Laubier, 1961; see below) are 
recorded in the literature from this century: 
the type species G. propinqua Marion & 
Bobretzky, 1875, and G. rosea (Malm, 
1874). The status of these two species is 
currently uncertain. For example, Eliason 
(1962) treated G. propinqua as a doubtful 
junior synonym of G. rosea, and Hartmann- 
Schroder (1971) referred Fauvel’s (1923) 
description of G. propinqua to G. rosea. 
Haaland & Schram (1982) also treated G. 
propinqua as a possible junior synonym, but 
remarked that the larvae from the Oslofjord 


differed from those described as G. propin- 
qua by Bhaud (1971). 

I believe that these synonymies are in- 
correct, and that the number of species oc- 
curring in the area is underestimated. This 
study aims to describe all the European spe- 
cies of Gyptis (including two new species) 
and the closely related Amphiduros, to re- 
solve their synonymies and to provide a key 
facilitating their correct identification. Apart 
from the interpretation of tentacular cirri 
(see below), the generic delineations follow 
traditional ones (e.g., Fauchald 1977) and 
no phylogenetic considerations are made at 
this point; these will have to await future 
studies treating the relationships within 
family (Pleijel, in prep.). The study is based 
on museum specimens as well as newly col- 
lected material during trips to the northern 
part of the Swedish west coast, northwestern 
Iceland, eastern Sicily, and Banyuls-sur-Mer 
in southern France. A checklist for species 
originally or later referred to the two genera 
is appended. The list also serves as a record 
of other congeneric species examined in the 
course of this study. 


VOLUME 106, NUMBER 1 


Species belonging to Podarkeopsis are ex- 
cluded. These are distinguished from Gyptis 
and Amphiduros by the presence of an an- 
teriorly inserted median antenna (rather than 
medially on the dorsal surface of the pro- 
stomium), by ten terminal proboscideal pa- 
pillae (rather than absence or a larger num- 
ber), and by the presence of furcate notosetae 
(rather than absence). Species recorded from 
European waters which I consider belong to 
the Podarkeopsis-group are: P. arenicolus 
(La Greca, 1946), described from the Gulf 
of Naples, Italy; P. galangaui Laubier, 1961, 
described from Banyuls-sur-Mer, southern 
France; P. capensis (Day, 1963), described 
from South Africa but recorded from south- 
ern England (Gibbs & Probert 1973) and 
the Tyrrhenian Sea, Italy (Gravina and 
Giangrande 1988); and Gyptis helgolandica 
Hilbig & Dittmer, 1979, described from 
Helgoland in the North Sea. 

In hesionids the number of tentacular cir- 
ri has generally been considered diagnostic 
at the generic level (e.g., Fauchald 1977). 
Gyptis and Amphiduros are described as 
having eight pairs of tentacular cirri, but the 
character “‘number of tentacular cirri” 
probably conceals a mixture of several char- 
acters. In the literature the tentacular cirri 
appear to be defined by absence of para- 
podia, i.e. if parapodial lobes are absent they 
are named tentacular cirri, otherwise dorsal 
and ventral cirri. However, the tentacular 
cirri also differ from cirri in “‘normal”’ seg- 
ments in the usually more pronounced cir- 
rophores, and in being stouter and longer. 
Further, many adult hesionids have dorsal 
cirri on the first setigerous segment that are 
of the same shape as the preceding dorsal 
tentacular cirri. In considering these am- 
biguities I prefer to use the absence of no- 
topodia on segment 4 to define Gyptis and 
Amphiduros and presently avoid the char- 
acter “number of tentacular cirri.”’ Gyptis 
mackiei (which has setigerous neuropodia 
on segment 4 and thus may be interpreted 
as having six pairs of tentacular cirri) will 
then also be included in Gyptis. Possible 


159 


reassignement will have to await future phy- 
logenetic analyses. 


Materials and Methods 


The collected specimens were relaxed, ei- 
ther with menthol or magnesium chloride 
(7% in distilled water), studied alive, pre- 
served in formalin (5—10% in seawater) for 
a few days, rinsed in fresh water and trans- 
ferred to 80% alcohol. For SEM specimens 
were similarly relaxed, preserved for a few 
hours in osmium tetraoxide (1% in artificial 
seawater), rinsed in distilled water, trans- 
ferred to 80% alcohol in a graded series, and 
subsequently critical-point dried. All draw- 
ings were made with a camera lucida; those 
of Figs. 1A, B, K, 4A—C, and 9A, B from 
live, relaxed specimens, and remaining ones 
from preserved specimens. 

All measurements were carried out either 
on live, relaxed specimens or specimens re- 
laxed prior to preservation. Width mea- 
surements were taken from median seg- 
ments and include parapodia but exclude 
cirri and setae. 

Institutions and museums are indicated 
by the following abbreviations: BIOFAR 
(Marine benthic fauna of the Faroe Islands, 
Kaldbak), BMNH (The Natural History 
Museum, London), LACM (Los Angeles 
County Museum of Natural History), 
NHMG (Goteborg Natural History Muse- 
um), NHMR (Natural History Museum, 
Reykjavik), NHMW (Naturhistorisches 
Museum Wien), NMCA (National Museum 
of Canada, Ottawa), NMW (National Mu- 
seum of Wales, Cardiff), SMNH (Swedish 
Museum of Natural History, Stockholm), 
USNM (National Museum of Natural His- 
tory, Smithsonian Institution, Washington, 
D.C.), ZMB (Universitat Humboldt, Mu- 
seum fur Naturkunde, Berlin), ZMH (Uni- 
versitat Hamburg, Zoologisches Institut und 
Museum), ZMUU (Uppsala Universitet, 
Zoologiska Muséet). All material is depos- 
ited at SMNH unless otherwise indicated. 


160 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


A B 


© SS NAN NIN SS SIN SN NIN SNES NN SESS SN 


ANNA ANS 


.N 


LH 
tae esscasiee 
COC 


Fig. 1. Gyptis propinqua. A. Anterior end, dorsal view. Setae omitted. B. Anterior end, ventral view. Setae 
omitted. C. Parapodium segment five, posterior view. D. Parapodium segment six, anterior view, ca. half number 
of setae shown. E. Median parapodium, anterior view, ca. half number of setae shown. F. Acicular notoseta. G. 
Spiked capillary notoseta. H. Serrated capillary notoseta. I. Median neuroseta. J. Ventral neuroseta. Specimens 
from Koster area, Sweden. Scales A-E, K, 0.25 mm; F-J, 50 um. 


VOLUME 106, NUMBER 1 


Gyptis Marion & Bobretzky, 1875 
Gyptis Marion & Bobretzky, 1875:50-51. 


Type species. —Gyptis propinqua Marion 
& Bobretzky, 1875, either by monotypy or 
subsequent designation (see Remarks). 

Diagnosis (provisional). —Hesionids with 
two palps and two frontal antennae. Median 
antenna present, inserted dorsally on pro- 
stomium. Proboscis with terminal ring of 
papillae. Jaws absent. Segment 4 without 
notopodia or notosetae, with or without 
neuropodia with neurosetae. Segment 5 with 
or without notopodia or notosetae, with 
neuropodia and neurosetae. Following 
parapodia with well-developed noto- and 
neuropodia. Notosetae include one to sev- 
eral acicular and large number of capillary 
setae; furcate setae absent. Neurosetae nu- 
merous, all usually compound. 

Remarks. —It is not obvious from Mari- 
on & Bobretzky’s original description that 
G. propinqua should constitute the type spe- 
cies of the genus, but the matter is probably 
without practical significance because it has 
been designated subsequently, if not earlier 
then at least by Hartman (1965). 

The name Oxydromus Grube, 1855 has 
variously been treated as confused (Marion 
& Bobretzky 1875), as a senior synonym to 
Gyptis (e.g., Fauvel 1923, Hartman 1959) 
or, more recently (Hartman 1965), as pre- 
occupied (in Aves by Oxydromus Schlegel, 
1854). However, examination of the type 
material for the type species for the genus, 
O. fasciatus Grube, 1855 (ZMB 3825), shows 
it to be a junior synonym to Ophiodromus 
flexuosus (delle Chiaje, 1827). Apart from 
possibly being a junior homonym the name 
is thus also a synonym of Ophiodromus Sars, 
1862, and of no relevance to Gyptis. 

Pending further investigations (see In- 
troduction) the generic diagnosis is provi- 
sionally emended to include G. mackiei, a 
new species with neurosetae present on seg- 
ment 4. 


Gyptis propinqua Marion & Bobretzky, 1875 
Figs. 1-3, 12 


161 


Gyptis propinqua Marion & Bobretzky, 
1875:51-54, pls. 5-6, fig. 15. 

Oxydromus propinquus. —Fauvel, 1923: 
241-242, fig. 90. [Not Oxydromus pro- 
pinquus sensu Ushakov, 1955:196-197, 
fig. 58] 

Gyptis rosea. —Hartmann-Schroder, 1971: 
132-134, fig. 42.—Helgason et al., 1990: 
205. 


Material examined.—Iceland: ca. 200 
specimens (NHMR), Breidafjodur, 65°05'N, 
23°17'W, 43 m, van Veen grab and Agassiz 
trawl, shell gravel, 9 Aug 1979; 3 specimens 
(NHMR), Breidafjordur, 65°05'N, 23°16'W, 
53 m, van Veen grab, shell gravel, 9 Aug 
1979; 15 specimens, Breidafjordur, Selsker, 
65°05.03’N, 23°16.91’W, 43 m, dredge, 
coarse shell gravel, 16 Jul 1991. Sweden: 7 
specimens, Vaderoarna, SE Norra Ragstu- 
ten, 58°32.2’N, 11°5.0’E, 12 m, dredge, shell 
sand and gravel, 7 Oct 1984; 1 specimen, 
Koster area, SW Yttre Vattenholmen, 
58°52.5'N, 11°06.3’E, 50-100 m, dredge, 
mixed sediments, 7 January 1985; 1 spec- 
imen, Koster area, W Yttre Vattenholmen, 
58°52.4'N, 11°06.5’E, 20-30 m, dredge, shell 
sand, 29 Aug 1985; 4 specimens, Koster 
area, SW Yttre Vattenholmen, 58°52.5'N, 
11°06.3’E, 30 m, dredge, shell sand and 
gravel, 30 May 1989; 7 specimens, Koster 
area, E Yttre Vattenholmen, 58°52.6’N, 
11°06.7'E, 20-30 m, dredge, coarse gravel, 
29 Aug 1989; 16 specimens, Koster area, W 
Yttre Vattenholmen, 58°52.4’N, 11°06.5’E, 
20-40 m, dredge, shell sand and gravel, 10 
Sep 1989; 1 specimen, Koster area, Koster- 
grund, 58°52.5’N, 11°05.1’E, 20-40 m, 
dredge, mixed sediments, 11 Apr 1990; 4 
specimens (all mounted for SEM), Koster 
area, W Yttre Vattenholmen, 58°52.4’N, 
11°06.5'E, 30-60 m, dredge, shell sand and 
gravel, 13 Apr 1990; 2 specimens, Koster 
area, SW Yttre Vattenholmen, 58°52.5'N, 
11°06.3'E, 40 m, dredge, mixed sediments, 
28 Jun 1990; 2 specimens, Koster area, Kos- 
terorunmd LasstaZioiN; VLSOSol EB, ikSim, 
dredge, sand and gravel, 18 Oct 1990; 3 
specimens, Koster area, SW Yttre Vatten- 


162 


holmen, 58°52.5’N, 11°06.3’E, 30-50 m, 
dredge, 6 Jun 1991; 4 specimens, Koster 
area, E Krugglo, 58°53.4’N, 11°05.9’E, 10- 
20 m, dredge, coarse shell gravel, 13 Jun 
1991; 7 specimens, Koster area, E Yttre 
Vattenholmen, 58°52.6'N, 11°06.7’E, 30 m, 
dredge, shell gravel, 18 Jun 1991. Faroes: 
1 specimen (BIOFAR), 61°54.78'N, 
06°28.84'W, 77 m, epibenthic sledge, 20 Jul 
1987; 4 specimens, Torshavn, Hggnabodi, 
62°05'N, 06°33’W, 20-40 m, dredge, 21 Nov 
1988. Italy: 1 specimen, Sicily, Acitrezza, 
37°34.4'N, 15°11.7'E, 41 m,; SCUBA, 17 
May 1990; Sicily, Brucoli, 37°16.8'N, 
15°11.7'E, 10 m, SCUBA; 21 May: 1990: 
France: ca. 75 specimens (5 specimens 
mounted for SEM), Banyuls-sur-Mer, Cap 
d’Osne, 42°29.48'N, 03°08.29’E, 24 m, 
dredge, silty coarse sand with shell gravel, 
9 Oct 1991; 4 specimens, Banyuls-sur-Mer, 
42°29.48'N, 03°08.29’E, 18 m, dredge, sand 
and shell gravel, 13 Oct 1991; 3 specimens, 
Banyuls-sur-Mer, Ile Grosse, 42°29.0’N, 
03°08.1’E, 10 m, SCUBA, gravel, 13 Oct 
1991; 1 specimen, Banyuls-sur-Mer, Cap 
Oullestrell, 42°30.13’N, 03°08.18’E, 18 m, 
dredge, 14 Oct 1991; 1 specimen, Banyuls- 
sur-Mer, Ile Grosse, 42°29.0’N, 03°08.1’E, 
10 m, SCUBA, gravel, 18 Oct 1991. 
Description. — Body, excluding parapo- 
dia, cylindrical, posteriorly tapered; venter 
flattened, without distinct median longitu- 
dinal furrow. Median parapodia only slight- 
ly longer than anterior, posterior ones suc- 
cessively shorter, yielding outline of fairly 
equal width with tapering posterior end. 
Prostomium about as wide as long, an- 
teriorly straight, laterally and posteriorly 
with rounded lobes separated by deep pos- 
terior incision (Figs. 1A, 2A, B). Proximal 
parts of palps cylindrical; distal parts widest 
medially, anteriorly rounded (Fig. 1B). 
Proximal and distal parts of equal length. 
Paired antennae situated on small cerato- 
phores, longer and thinner than palps, with 
pointed tips. Median antenna club-shaped, 
widest subdistally, inserted half-way be- 
tween anterior pair of eyes and anterior 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


margin of prostomium on small, rather in- 
distinct ceratophore. Anterior pair of eyes 
rounded to reniform, twice as large as pos- 
terior pair and situated further apart; pos- 
terior pair rounded, both pairs with lenses. 
Nuchal organs well-developed, lateral to 
prostomium but extending and almost co- 
alescing mid-dorsally (Fig. 2A). 

Large, distinct lip glands present (Figs. 
1B, 2C, E). Proboscis smooth, cylindrical, 
sometimes with enlarged opening in pre- 
served, unrelaxed specimens, divided in 
proximal and distal parts (not obvious on 
specimens with strongly extended probos- 
cides), distal part smaller. Number of ter- 
minal papillae size-dependent (Fig. 12), 
usually numbering 35-50. Papillae long (ca. 
125 wm) and thin with ciliated tips, arranged 
in single ring. 

Tentacular cirri distinctly annulated, rings 
basally about two to three times as long as 
wide, distally usually shorter, tips rounded. 
Dorsal tentacular cirri of segment 2 longest, 
reaching to about segment 10-12; ventral 
tentacular cirri of segment 3 shortest, reach- 
ing to about segment 6. Cirri of segments 3 
and 4 more ventrally displaced than ante- 
rior ones (Fig. 2D). Aciculae present in cir- 
rophores of all tentacular cirri, two to three 
in dorsal ones and one in ventral ones. An- 
terior dorsal segmental delineations not dis- 
tinct, several segments fused or reduced. 
Segment 4 with first dorsally fully distin- 
guishable segment, often forming elevated 
ridge anteriorly (Fig. 2A). 

Segment 5 (setiger 1) with from one to 
three notoaciculae within cirrophores of 
dorsal cirri, without setigerous lobes or se- 
tae (Fig. 1C). Dorsal cirri similar to those 
of segment 4. Neuropodia similar to follow- 
ing ones but slightly smaller. Ventral cirri 
similar to following ones. Segment 6 similar 
to median ones but slightly smaller (Fig. 
1D). 

Dorsal cirri of median segments distinctly 
annulated with about 10—15 rings; rings from 
one to three times as long as wide (Fig. 1E). 
Dorsal cirri longer than setae, slightly dif- 


VOLUME 106, NUMBER 1 


Fig. 2. SEM micrographs of Gyptis propinqua. A. Anterior end, dorsal view. B. Prostomium, dorsal view. 
C. Anterior end, ventral view. D. Anterior end, right side. E. Lip glands. F. Median parapodium, right side, 
antero-ventral view. A & F specimens from Koster area, Sweden, B—D specimens from Banyuls, southern France. 
Scale lines A—D, F, 0.1 mm; E, 50 wm. 


0 5 10 15 20 25 30 35 
No. of segments 


Fig. 3: 
number of segments and length. Specimens from Ban- 
yuls, southern France (squares), Koster area, Sweden 
(circles), and Breidafj6rdur, northwestern Iceland (tri- 


angles). 


Gyptis propinqua. Relationship between 


ferentiated in length and orientation, those 
of segment 5, 8, 10, 12, 15, 17, 19, 21 and 
23 slightly longer and oriented more dor- 
sally than other ones (best observed on live 
specimens). Notopodial lobes conical, usu- 
ally with two internal aciculae (one small 
and difficult to detect), and one emerging, 
anteriorly situated, dorsally bent smooth 
acicular seta (Fig. 1F). About 10-15 capil- 
lary setae inserted behind lobe, with smooth 
proximal parts and two alternating rows of 
teeth medially and subdistally (Fig. 1G). A 
few ventrally situated serrated capillaries 
usually present (Fig. 1H). 

Neuropodia of median segments conical, 
slightly asymmetrical, usually with two 
internal aciculae (one larger than other) 
and 20-30 compound setae. Distal part of 
setal shafts internally reticulated (Fig. 1I, J). 
Blades of median and dorsal setae long, ven- 
tral ones very short. Ventral cirri annulated 
with about 5-10 rings, longer than neuro- 
podial lobe, on distinct cirrophores situated 
far back on neuropodia (Fig. 2F). 

Pygidium a rounded cone. Pygidial cirri 
long and annulated, median papilla absent 
(Fig. 1K). 

Color: Live specimens transparent with 
dark brown pigmentation forming segmen- 
tally arranged transverse bands across dor- 
sum that also connect laterally. First dor- 
sally visible segment with distinct dark 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


brown band dorsally. Eyes orange-red. 
Specimens vary from dark to pale, depend- 
ing on amount of pigmentation. Venter al- 
ways unpigmented. Eggs uncolored. Mature 
males whitish. Pigmentation usually fades 
in alcohol. 

Measurements: Up to 7.5 mm long for 32 
segments (see Fig. 3). 

Habitat.—Coarse and fine shell gravel 
from 10-100 m. 

Distribution. —Iceland, Faroes, Swedish 
west-coast, southern France, Sicily. 

Reproduction. —Mature specimens found 
in May and June at Koster, Sweden, in July 
at Iceland. Eggs large, about 175 wm in di- 
ameter. Bhaud (1971) described the larvae 
of G. propinqua. 

Remarks.—Eliason (1962), Haaland & 
Schram (1982) and Helgason et al. (1990), 
treated G. propinqua as a possible junior 
synonym of G. rosea. The two most recent 
studies, however, commented on inconsis- 
tencies between different descriptions. Fur- 
ther, Hartmann-Schroder (1971) and Hel- 
gason et al. stated that Fauvel’s (1923) 
description of G. propinqua referred to G. 
rosea. There is no remaining type material 
of G. propinqua, either at the museum in 
Paris, or at Station Marine d’Endoume in 
Marseille (J.-C. Dauvin and G. Bellan, in 
litt.); itis presumed lost. Nevertheless, new- 
ly collected material from Banyuls-sur-Mer 
was found to correspond well with the ex- 
cellent original description based on speci- 
mens from Marseille. Comparison of the 
Banyuls specimens with both the type and 
newly collected material of G. rosea leaves 
no doubt that G. propinqua is a distinct and 
very different species (see key for the best 
diagnostic characters). Hartmann-Schro- 
der’s (1971) description and examination of 
the specimens of Helgason et al. (1990) 
shows both accounts of G. rosea actually 
refer to G. propinqua. By contrast, the ac- 
counts of G. rosea by Eliason (1962) and 
Haaland & Schram (1982) are considered 
correct and refer to G. rosea as described 
here. Fauvel’s (1923) description of G. pro- 


VOLUME 106, NUMBER 1 


pinqua (as Oxydromus propinquus) 1s in good 
agreement with Marion & Bobretzky’s spe- 
cies. 

Ushakov (1955) reported G. propinqua 
(as Oxydromus propinquus) from the Sea of 
Okhotsk, but judging from his description 
this presumably represents a different spe- 
cies. 


Gyptis mackiei, new species 
Figs. 4-5 


Material examined.—Sweden: 1 speci- 
men, Gullmarsfjord, Skar, 110 m, mud, 25 
May 1963, (NHMG 12805c); 4 paratypes 
(SMNH 4395), Koster area, SW Yttre Vat- 
tenholmen, 58°52.1'N, 11°06.8’E, 100-140 
m, detritus sledge, mud, 8 Aug 1987; 2 para- 
types (SMNH 4396), Singlefjord, 59°04’N, 
11°10’E, 80 m, detritus sledge, mud, 11 Apr 
1990; holotype (SMNH 4397) and 2 addi- 
tional specimens (used for dissection and 
SEM), Koster area, S Yttre Vattenholmen, 
58°52.0’N, 11°06.6’E, 110-130 m, detritus 
sledge, mud, 1 Jan 1991; 1 specimen 
(mounted for SEM), Koster area, S Yttre 
Vattenholmen, 58°52.25'N, 11°06.30’E, 90- 
140 m, detritus sledge, mud, 19 Aug 1991; 
3 paratypes (NMW.Z. 1992.007.1-2) and 
one additional specimen (mounted for 
SEM), Singlefjord, 59°04.5’N, 11°10.6’E, 78- 
84 m, detritus sledge, mud, 26 Aug 1991; 
1 specimen (mounted for SEM), Singlefjord, 
59°04.8’N, 11°10.8’E, 82-83 m, detritus 
sledge, mud, 15 Sep 1991. Skagerrak: 3 
specimens, 58°08'N, 10°07’E, 295 m, grab, 
27 Jun 1933; 1 specimen (ZMUUV), 58°N, 
09°33’E, 271 m, Agassiz trawl, 30 Jun 1933; 
7 specimens (ZMUV), 58°02.5'N, 09°29.5’E, 
478 m, Agassiz trawl, 30 Jun 1933; 1 spec- 
imen (ZMUU), 57°50'N, 08°51'E, 358 m, 
grab, 5 Jul 1933; 1 specimen (ZMUU), 
57°41'N, 08°35’E, 191 m, grab, 6 Jul 1933; 
6 specimens (ZMUUV), 58°22’N, 10°34’E, 
270 m, Agassiz trawl, 14 Jul 1933; 2 spec- 
imens (ZMUU), 58°30’N, 10°26’E, 300 m, 
Agassiz trawl, 15 Jul 1933. Faroes: 4 spec- 
imens (BIOFAR), 62°31.40'N, 05°02.30'W, 


165 


430 m, epibenthic sledge, 17 Jul 1987; 
1 specimen (BIOFAR), 61°41.75'N, 
05°47.71'W, 354 m, epibenthic sledge, 
18 Jul 1987; 1 specimen (BIOFAR), 
61°13.30'N, 04°46.50’W, 780 m, epibenthic 
sledge, 19 Jul 1987; 4 specimens (BIOFAR), 
60°31.34'N, 08°25.07'W, 732 m, epibenthic 
sledge, 22 Jul 1987; 11 specimens (BIO- 
FAR), 62°05.32'N, 10°06.71'W, 859 m, epi- 
benthic sledge, 15 May 1988; 1 specimen 
(BIOFAR), 62°12.30'N, 03°59.54'W, 402 
m, detritus sledge, 27 May 1989; 2 speci- 
mens (BIOFAR), 62°41.31'N, 10°03.90'W, 
500 m, epibenthic sledge, 1 Jun 1989. 

Description. —Body, excluding parapo- 
dia, cylindrical; venter flattened, without 
distinct longitudinal furrow. Median para- 
podia only slightly longer than anterior, 
posterior ones successively shorter, result- 
ing body-outline of fairly constant width, 
tapering slowly posteriorly. 

Prostomium rounded, as wide as long, 
anteriorly straight, posterior incision often 
indistinct (Fig. 4A), often more conspicuous 
on specimens with everted proboscis. Prox- 
imal parts of palps cylindrical, distal parts 
widest medially, anteriorly rounded (Fig. 
4B); distal parts longer than proximal. Paired 
antennae without distinct ceratophores, as 
long as palps but thinner, with pointed tips. 
Median antenna widest medially, without 
distinctly pointed tip, inserted in front of 
anterior pair of eyes. Anterior pair of eyes 
rounded to reniform, slightly larger than 
posterior pair and situated farther apart; 
posterior pair rounded. Nuchal organs lat- 
eral to prostomium, not coalescing dorsally. 

Large, distinct lip glands present (Fig. 4B). 
Proboscis short, smooth, divided into prox- 
imal and distal parts; distal part smaller (Fig. 
4C). Terminal papillae 25-26 (observed in 
two specimens only; one 5.75 mm long for 
26 segments, and one with posterior end 
missing), short (about 60-70 wm) and blunt, 
distally ciliated, arranged in single ring. 

Tentacular cirm distinctly annulated, rings 
from one to four times as long as wide, tips 
rounded. Dorsal ones of segment 2 longest 


166 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Gyptis mackiei, new species. A. Anterior end, dorsal view. Setae omitted. B. Anterior end, ventral 
view. Setae omitted. C. Head end with protruded proboscis, dorsal view. D. Parapodium segment four, anterior 
view, ca. half number of setae shown. E. Parapodium segment five, anterior view, ca. half number of setae 


VOLUME 106, NUMBER 1 


and stoutest, reaching to about segment 12- 
16; ventral ones of segment 3 shortest, 
reaching to about segment 5, similar to fol- 
lowing ventral cirri but larger. Single acicula 
present in all cirrophores of all dorsal ten- 
tacular cirri; not observed in ventral ten- 
tacular cirri (but may be present). Segment 
1 reduced dorsally (possibly fused to seg- 
ment 2), segment 2 fully developed. 

Notopodia of segment 4 (setiger 1) with 
one notoacicula situated in cirrophores of 
dorsal cirri, without setigerous lobes or se- 
tae (Fig. 4D). Dorsal cirri similar to those 
of segment 3. Neuropodia similar to follow- 
ing ones but slightly smaller, with about 15 
compound setae. Ventral cirri similar to fol- 
lowing ones. Segment 5 similar to segment 
4 but slightly larger (Fig. 4E). 

Segment 6 similar to median ones but 
slightly smaller (Fig. 4F). 

Notopodia of median segments with an- 
nulated dorsal cirri with from three to seven 
rings, shorter than notosetae, with pointed 
tips (Fig. 4G). Dorsal cirri without obvious 
differentiation in length and orientation. 
Notopodial lobes conical, with one or two 
(usually one) internal aciculae (smaller one 
difficult to detect), and from one to three 
emerging, anteriorly situated dorsally bent 
acicular setae (Fig. 41). About 20-30 cap- 
illary setae inserted behind lobe, with 
smooth proximal parts and two alternating 
rows of teeth medially and subdistally (Fig. 
4H). Serrated notosetae not observed. 

Neuropodia of median segments conical, 
with single internal acicula and about 30- 
50 compound setae. Distal part of setal shafts 
internally reticulated (Fig. 4J, K). Blades of 
median and dorsal setae long, ventral ones 
short. A few dorsally situated serrated cap- 
illary setae occasionally present. Ventral cir- 
ri smooth or indistinctly annulated, as long 


_ 


167 
7. 
6 (e) Oo O Oo 
E 5 Ooo 
E4 
AS fe) 
m3 eS) 
3 
ae 
1 
) 
0 5 10 15 20 25 30 
No. of segments 
Fig. 5. Gyptis mackiei, new species. Relationship 


between number of segments and length. Specimens 
from Koster area, Sweden. 


as or slightly longer than neuropodial lobe, 
on distinct cirrophores situated far back on 
the neuropodium. 

Pygidium rounded. Pygidial cirri long, 
annulated, median papilla absent. 

Color: Live specimens transparent with 
dark brown pigmentation forming trans- 
verse stripes dorsally on anterior and pos- 
terior sides of each segment, especially con- 
spicuous on first dorsally visible segment. 
Eyes red. Eggs rose-colored, mature males 
whitish. Brown pigmentation fades in al- 
cohol. 

Measurements: Up to 5.75 mm long for 
29 segments (see Fig. 5). 

Habitat.—Found on mud bottoms from 
78-859 m. 

Distribution. —Northern part of Swedish 
west coast, Skagerrak, Faroes. 

Reproduction. — Mature specimens found 
in January in Sweden; not fully mature spec- 
imens found in August. Eggs about 100 um 
in diameter. 

Remarks.—In many hesionids the ante- 
rior parapodia are successively reduced dur- 
ing ontogeny (e.g., Blake 1975; Haaland and 
Shram 1983; Schram and Haaland 1984), 
and the first segment carrying setae and 


shown. F. Parapodium segment six, anterior view, ca. half number of setae shown. G. Parapodium segment 13, 
anterior view, ca. one third of setae shown. H. Acicular notoseta. I. Spiked capillary notoseta. J. Median neuroseta. 
K. Ventral neuroseta. A—-C paratypes (NMW.Z. 1992.007.1-—2), D—K specimen from Koster area, Sweden. Scales 


A-C, 0.25 mm; D-G, 0.1 mm; H-K, 50 um. 


168 


number of tentacular cirri should be com- 
pared between corresponding semapho- 
ronts only. That the description above is 
based on adults is evidenced by the presence 
of sexual products in many of the speci- 
mens. 

Gyptis mackiei is unique among hesio- 
nids in the character combination of “‘me- 
dian antenna with dorsal insertion,’ and 
““neurosetae present but notosetae absent on 
segment 4 and 5.” It is united with other 
species of Gyptis by the place of insertion 
of median antenna, by the presence of a 
large number of proboscideal papillae, by 
the absence of furcate setae, and by the ab- 
sence of noto- but presence of neurosetae 
on segment 5. 

Gyptis mackiei seems closely related to 
G. propinqua and the two species are rather 
similar in general appearance. In addition 
to the fourth setigerous segment G. mackiei 
differs in having a median antenna which 
is widest in its midregion rather than sub- 
distally, in its prostomium being less deeply 
incised posteriorly, in having much shorter 
and more pointed dorsal cirri, and in having 
rose-colored rather than colorless eggs. 

Eliason’s specimens from Gullmarsfjord 
and of the Skagerrak-Expedition 1933 con- 
stitute a mixture of G. rosea and G. mackiei; 
they have now been relabelled. 

Etymology. —This species is named for 
Andrew Mackie, friend and collaborator. 


Gyptis mediterranea, new species 
Figs. 6-8, 12 


Material examined. —France: 3 para- 
types (NMW.Z.1992.007.3), Banyuls-sur- 
Mer, 42°29.92’N, 03°09.22’E, 35 m, dredge, 
sandy mud with detritus, 3 Oct 1991; ho- 
lotype (SMNH 4398), 7 paratypes (SMNH 
4399, 4400), and 4 additional specimens 
mounted for SEM, Banyuls-sur-Mer, Cap 
Oullestrell, 42°30.17'N, 03°09.48’E, 40 m, 
dredge, mud, 7 Oct 1991; 13 paratypes 
(SMNH 4401), Banyuls-sur-Mer, 42°30.00'N, 
03°11.75’E, 80 m, dredge, mud, 13 Oct 1991. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Italy: 1 specimen, eastern Sicily, Brucoli, 
37°17'N, 15°13’E, 60 m, dredge, mud, 21 May 
1990. 

Description. —Body, excluding parapo- 
dia, cylindrical, tapering posteriorly; venter 
flattened without distinct median longitu- 
dinal furrow. Parapodia long, often directed 
anteriorly, median ones longest, yielding el- 
liptical and flattened outline of animal. 

Prostomium trapezoidal with rounded 
corners (Figs. 6A, 7A), with small posterior 
incision (often visible only in SEM). Prox- 
imal parts of palps cylindrical, distal parts 
thinner, narrowing to rounded ends. Prox- 
imal parts slightly longer than distal. Paired 
antennae slightly shorter and narrower than 
palps, with thin tips, situated on small cera- 
tophores. Median antenna cylindrical, end- 
ing without well defined, prolonged tip, in- 
serted on line between anterior pair of eyes. 
Anterior pair of eyes rounded, larger than 
posterior pair and situated farther apart, 
posterior pair rounded, both pairs with lens- 
es. Nuchal organs lateral to prostomium, 
not dorsally coalescing (Fig. 7B). 

Lip glands absent (Fig. 6B). Proboscis di- 
vided in proximal and distal parts. Proxi- 
mal part larger and longer, with small, poor- 
ly defined papillae (Fig. 7A); distal part short 
and smooth. Number of terminal papillae 
size-dependent (Fig. 12), about 20-32. Pa- 
pillae long (ca. 60-70 wm) and thin with 
ciliated tips, arranged in single ring (Fig. 
7C). 

Tentacular cirri thin, annulated (not al- 
ways obvious proximally) with rings (one 
to three times as long as wide), tips distinctly 
pointed. Dorsal tentacular cirri of segment 
2 longest, reaching to about segment 10-12, 
ventral tentacular cirri of segment 3 short- 
est, reaching to about segment 5-6. Cirri of 
segment 3 and 4 more ventrally displaced 
than anterior ones. Aciculae present in all 
cirrophores of tentacular cirri (small acces- 
sory ones not observed but may be present). 
Segment 1 dorsally reduced, segments 2 and 
3 dorsally fused. 

Notopodia of segment 5 (setiger 1) with 


VOLUME 106, NUMBER 1 169 


“7, 


A —— mi B A MN NAN yahs An 


Fig. 6. Gyptis mediterranea, new species. A. Anterior end, dorsal view. Setae omitted. B. Anterior end, 
ventral view. Setae omitted. C. Parapodium segment five, anterior view. D. Parapodium segment six, anterior 
view, ca. half number of setae shown. E. Parapodium segment 14, anterior view, ca. half number of setae shown. 
F. Acicular notoseta. G. Spiked capillary notoseta. H. Serrated capillary notoseta. I. Median neuroseta. H. 
Serrated capillary notoseta. I. Median neuroseta. J. Ventral neuroseta. A-B holotype, C paratype (SMNH 4401), 
D-J paratype (SMNH 4399). Scales A-B, 0.5 mm; C-E, 0.25 mm; F-J, 50 um. 


170 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 7. SEM micrographs of Gyptis mediterranea, new species. A. Anterior end, dorsal view. B. Anterior 
end, right side. C. Terminal ring of proboscis, anterior view. D. Anterior-dorsal view of dorsum, median segments. 
E. Median parapodium, left side, postero-dorsal view. F. Median parapodium, right side, antero-ventral view. 
Specimens from Banyuls, southern France. Scale lines, 0.1 mm. 


VOLUME 106, NUMBER 1 


one notoacicula in cirrophores of dorsal cir- 
ri, without setigerous lobes or setae. Dorsal 
cirri similar to those of segment 4. Neuro- 
podia similar to following ones but slightly 
smaller and with fewer setae (Fig. 6C). Ven- 
tral cirri similar to following ones. Segment 
6 similar to median ones but slightly smaller 
(Fig. 6D). 

Elevated dorsal ridges present across pos- 
terior side of each segment (Fig. 7D, E), less 
distinct on anterior segments and more pro- 
nounced on median and posterior ones. In 
one specimen the ridges are provided with 
distinct cylindrical papillae (length ca. 15 
um). 

Notopodia of median segments with more 
or less distinctly annulated dorsal cirri 
(smooth proximally), with about five rings, 
about three times as long as wide. Dorsal 
cirri shorter than setae, inserted posteriorly 
to those (Fig. 6E). All dorsal cirri thin, of 
approximately similar length, but those of 
Seement), 8, 10,12, 15,17, 19,21; and.23 
oriented slightly more dorsally than other 
ones (best observed on live specimens). No- 
topodial lobes conical, with one or two in- 
ternal aciculae (small one difficult to detect) 
and from zero to four emerging, dorsally 
bent acicular setae, situated anteriorly to 
other setae (Fig. 6F); tapering but termi- 
nated bluntly, often with fine spines distally. 
Large number of long capillary setae in- 
serted behind lobe, median ones twice as 
long as dorsal and ventral ones, with smooth 
proximal parts and two alternating rows of 
medial and subdistal teeth (Fig. 6G). A few 
serrated notosetae usually present, situated 
ventrally (Fig. 6H). 

Neuropodia of median segments conical, 
usually with two internal aciculae, one large 
and one small, about 20-40 compound se- 
tae, and, occasionally, one or two dorsally 
serrated capillaries. Distal part of setal shafts 
with transverse striation internally. Blades 
thin, dorsal side from distinctly serrated to 
almost smooth, median and dorsal ones long 
(Fig. 61), ventral ones shorter (Fig. 6J). A 
few additional serrated capillary setae often 


171 


oOo N © 


Length (mm) 
o- ND WO Lf OW 


0 5 10 15 20 25 30 35 
No. of segments 


Fig. 8. Gyptis mediterranea, new species. Relation- 
ship between number of segments and length. Speci- 
mens from Banyuls, southern France. 


present, situated dorsally. Ventral cirri 
smooth with fine tapering tips (Fig. 7F), 
without cirrophores, situated distally on 
neuropodium. 

Pygidium rounded. Pygidial cirri annu- 
lated, longer than dorsal cirri, with pointed 
tips, median papilla absent. 

Color: Live specimens transparent. Eyes 
red. Small brown pigment spots may be 
present ventrally on posterior side of para- 
podia. Eggs colorless. Preserved specimens 
white, brown spots retained. 

Measurements: Up to 7 mm long for 32 
segments (see Fig. 8). 

Habitat. — Mud and sandy mud from 35-— 
80 m. 

Distribution. — Presently known only from 
southern France and eastern Sicily. 

Reproduction. —Several of the specimens 
collected in Banyuls in October were mature 
females with an egg size of 50-60 um in 
diameter. No males observed. 

Remarks. —Within Gyptis G. rosea and 
G. mediterranea are unique in having dis- 
tally inserted ventral cirri. They share with 
G. hians Fauchald & Hancock, 1981, the 
elliptical and flattened body-shape, but the 
latter differs in having ventral cirri inserted 
subdistally, and in having the distal part of 
setal shafts internally reticulated rather than 
striated. 

Gyptis mediterranea differs from G. rosea 
in being smaller, having red rather than black 
eyes, having a median antenna without ex- 


172 


tended tip, having a smaller number of ter- 
minal proboscis papillae (even when ad- 
justed for size; see Fig. 12), having shorter 
dorsal cirri, having much more pronounced 
dorsal ridges, and in having colorless rather 
than pink eggs. 

Etymology.—Named for the Mediterra- 
nean Sea. 


Gyptis rosea (Malm, 1874) 
Figs. 9-12 


Ophiodromus roseus Malm, 1874. (p. 82) 

Gyptis rosea. —Haaland & Schram, 1982.— 
Eliason, 1962 (p. 238-240, fig. 9), in part. 
[Not Gyptis rosea sensu Hartmann- 
Schroder, 1971:132-134, fig. 42.—Hel- 
gason et al., 1990:205] 


Material examined. —Sweden: Holotype 
(NHMG 901), Gullmarsfjord, 45 fathoms, 
mud; 2 specimens (NHMG 12805a & b), 
Gullmarsfjord, Skar, 110 m, mud, 25 May 
1963; 1 specimen, Singlefjord, 59°04.9'N, 
11°10.8’E, 80 m, detritus sledge, mud, 27 
Dec 1988; 2 specimens, Singlefjord, 
59°04.9'N, 11°10.8’E, 80 m, detritus sledge, 
mud, 14 Jun 1989; 2 specimens, Koster area, 
W Svartskar, 58°54.5’N, 11°05.0’E, 100-150 
m, dredge, mud, 22 Sep 1989; 6 specimens 
(2 mounted for SEM), Singlefjord, 
59°04.9'N, 11°10.8’E, 80 m, detritus sledge, 
mud, 11 Apr 1990; 1 specimen, Koster 
area, S Yttre Vattenholmen, 58°52.1'N, 
11°06.9’E, 50-100 m, dredge, mud, 1 Oct 
1990; 3 specimens (mounted for SEM), Sin- 
glefjord, 59°04.9'N, 11°10.8’E, 83 m, detri- 
tus sledge, mud, 26 Feb 1991; 1 specimen, 
Koster area, S Yttre Vattenholmen, 
58°52.1’N, 11°06.9’E, 80-140 m, dredge, 
mud, 17 Aug 1991; 5 specimens, Sin- 
glefjord, 59°04.5’N, 11°10.6’E, 82-84 m, 
detritus sledge, mud, 2 Sep 1991; 2 speci- 
mens (1 mounted for SEM), Koster area, S 
Yttre Vattenholmen, 58°52.0’N, 11°06.6’E, 
100-110 m, detritus sledge, mud, 7 Sep 
1991; 5 specimens, Singlefjord, 59°04.8'N, 
11°10.8’E, 82-83 m, detritus sledge, mud, 
15 Sep 1991. Skagerrak: 3 specimens 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


(ZMUUV), 58°08’N, 10°07’E, 295 m, grab, 
27 Jun 1933; 2 specimens (ZMUU), 
58°2.5’N, 09°29.5’E, 478 m, grab, 30 Jun 
1933; 1 specimen (ZMUU), 58°02.5’N, 
09°29.5'E, 427 m, grab, 1 Jul 1933; 1 spec- 
imen (ZMUUV), 58°21'N, 08°56’E, 225 m, 
grab, 2 Jul 1933; 1 specimen (ZMUU), 
58°02.7'N, 08°13.5’E, 241 m, grab, 4 Jul 
1933; 4 specimens (ZMUU), 57°50’N, 
08°51’E, 358 m, grab, 5 Jul 1933; 1 speci- 
men (ZMUU), 57°45'N, 08°07’E, 421 m, 
grab, 6 Jul 1933; 3 specimens (ZMUU), 
57°52'N, 08°01’E, 510 m, Agassiz trawl, 6 
Jul 1933; 2 specimens (ZMUU), 57°58’N, 
06°44’E, 384 m, grab, 7 Jul 1933; 1 speci- 
men (ZMUV), 58°59.5’N, 06°27’E, 290 m, 
dredge, 12 Jul 1933; 6 specimens (ZMUU), 
58°22’N, 10°34’E, 270 m, Agassiz trawl, 14 
Jul 1933; 5 specimens (ZMUU), 58°30’N, 
10°26’E, 300 m, Agassiz trawl, 15 Jul 1933; 
1 specimen (ZMUU), 58°30'N, 10°32.5’E, 
175 m, Agassiz trawl, 15 Jul 1933; 1 spec- 
imen, 57°59'N, 08°40’E, 500 m, RP-sledge, 
15 Mar 1990; 1 specimen, 57°49.5'N, 
08°12.5’E, 500 m, RP-sledge, 15 Mar 1990. 

Description. —Body, excluding parapo- 
dia, cylindrical, tapered posteriorly; venter 
flattened, with distinct median longitudinal 
furrow. Parapodia long, median ones lon- 
gest, resulting in elliptical and flattened out- 
line of animal. 

Prostomium rounded rectangular to trap- 
ezoidal, almost twice as wide as long (Fig. 
9A), with small posterior incision (often not 
visible except in SEM; Fig. 10A). Proximal 
parts of palps cylindrical, distal parts thin- 
ner, anteriorly rounded; proximal and distal 
parts of equal length. Paired antennae with- 
out ceratophores, as long as palps but thin- 
ner, with prolonged tips. Median antenna 
similar in shape to frontal ones but smaller, 
inserted just in front of anterior pair of eyes. 
Anterior and posterior pairs of eyes small, 
rounded; anterior pair slightly larger. Nu- 
chal organs lateral to prostomium, not co- 
alescing dorsally. 

Lip glands absent. Proboscis divided into 
proximal and distal parts (not obvious on 


Fig. 9. Gyptis rosea. A. Anterior end, dorsal view. Proboscis partly everted. Setae omitted. B. Anterior end 
with proboscis everted (some papillae lacking), ventral view. Setae omitted. C. Right parapodium, segment 5, 
posterior view. Full number of setae (ca. 30) not shown. D. Right parapodium, segment 6, anterior view. Full 
number of notosetae (ca. 25) or neurosetae (ca. 40) not shown. E. Right parapodium, segment 21, anterior view. 
Full number of notosetae (ca. 40) or neurosetae (ca. 40) not shown. F. Acicular notoseta. G. Spiked capillary 
notoseta. H. Median neuroseta. I. Ventral neuroseta. Specimens from Singlefjord and Koster area. Scales A-B, 
0.5 mm; C-E, 0.25 mm; F-I, 50 um. 


174 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 10. SEM micrographs of Gyptis rosea. A. Anterior end, dorsal view. B. Distal part of proboscis, ventral 
view. C. Insertion of tentacular cirri, right side. D. Parapodia from segment 25-26, right side, antero-dorsal 
view. E. Parapodium from segment 11, left side, antero-ventral view. F. Spiked capillary notoseta from median 
segment. G. Articulation between shaft and blade of median neuroseta from segment 18. Specimens from 
Singlefjord, Sweden. Scales A-E, 0.1 mm; F—G, 10 yum. 


VOLUME 106, NUMBER 1 


specimens with strongly extended probos- 
cides) (Fig. 9B). Proximal part larger and 
longer, covered with poorly defined, dif- 
fusely distributed papillae, about 30-40 um 
in diameter (Fig. 10A) (surface appearing 
slightly rugose in lower magnifications); dis- 
tal part short and smooth. Number of ter- 
minal papillae size-dependent (Fig. 12), 
about 35-80. Papillae long and thin with 
ciliated tips (Fig. 10B), arranged in single 
ring (may appear as several rings in speci- 
mens with proboscis incompletely protrud- 
ed). 

Tentacular cirri thin, annulated (not al- 
ways obvious proximally) with median rings 
about three times as long as wide, tips dis- 
tinctly pointed. Dorsal tentacular cirri of 
segment 2 longest, reaching to about seg- 
ment 12-18; ventral tentacular cirri of seg- 
ment 3 shortest, reaching to about segment 
5-7. Cirri of segment 3 and 4 more ventrally 
displaced than anterior ones (Fig. 10C). Sin- 
gle acicula present in all cirrophores of ten- 
tacular cirri except ventral ones of segment 
3 and 4, which have double ones (smaller 
ones difficult to detect). 

Anterior segmental delineations not very 
distinct. Segment 1 reduced dorsally except 
for small middorsal part (Fig. 10A), seg- 
ments 2 and 3 appearing fused dorsally, fol- 
lowing segments fully developed. 

Notopodia of segment 5 (setiger 1) with 
one notoacicula situated in cirrophore of 
dorsal cirrus, but without setigerous lobes 
or setae. Dorsal cirri similar to those of seg- 
ment 4. Neuropodia similar to following 
ones but shorter (Fig. 9C). Ventral cirri sim- 
ilar to following ones. Segment 6 similar to 
median ones but smaller (Fig. 9D). 

Notopodia of median segments with more 
or less distinctly annulated dorsal cirri 
(smooth proximally), with about eight to ten 
rings; median rings about three times as long 
as wide (Fig. 9E). Dorsal cirri thin, longer 
than notosetae, inserted posteriorly to those 
(Fig. 10D). All dorsal cirri of similar length, 
those-of Scgoment 5; 8) 10) 12; 15, 17, 19, 
21, 23 and 26 oriented slightly more dor- 


F75 


Length (mm) 


0) 5 10 15 20 25 30 35 40 
No. of segments 


Fig. 11. Gyptis rosea. Relationship between num- 
ber of segments and length. Specimens from Koster 
area and Singlefjord, Sweden. 


sally than other ones (best observed on live 
specimens). Notopodial lobes conical, usu- 
ally with two internal aciculae (one small 
and difficult to detect) and from one to six 
(usually from three to five) emerging, dor- 
sally bent acicular setae, situated anteriorly 
to other setae (Fig. 9F), tapering but ter- 
minated bluntly, occasionally with fine 
spines distally. Large number of long cap- 
illary setae inserted behind lobe, median 
ones twice as long as dorsal and ventral ones, 
with smooth proximal parts and two alter- 
nating rows of teeth medially and subdis- 
tally (Figs. 9G, 10F). A few serrated noto- 
setae situated ventrally occasionally present 
(Fig. 9H). 

Neuropodia of median segments conical 
to rounded, usually with two internal acic- 
ulae, one large and one small, and about 
40-50 compound setae (Fig. 10G). Distal 
part of setal shafts with internal transverse 
striation. Blades thin, dorsal side varying 
from distinctly serrated to almost smooth; 
median and dorsal ones long (Fig. 91), ven- 
tral ones short (Fig. 9J). A few additional 
serrated capillary setae may occur dorsally 
(difficult to detect). Ventral cirri smooth with 
fine, evenly tapering tips (Fig. 10E), situated 
distally on neuropodium, without cirro- 
phores. 

Pygidium rounded. Pygidial cirri very 
long, annulated, median papilla absent. 

Color: Live specimens transparent, stom- 
ach yellowish to orange. Mature females 


No. of papillae 
mo oO ff 
(oj (oi t=) 


0 5 10 15 20 25 30 35 40 
No. of segments 


Fig. 12. Relationship between number of segments 
and number of terminal proboscideal papillae in Gyptis 
propinqua (triangles), G. mediterranea, new species 
(squares), and G. rosea (circles). 


pink-orange. Eyes dark brownish-black. 
Dark brown pigment spots may be present 
dorsally and ventrally on posterior side of 
parapodia. Preserved specimens yellowish- 
white. Brown pigment usually retained. 

Measurements: Up to 11 mm long for 36 
segments (see Fig. 11). 

Habitat. —Mud bottoms from 50-510 m. 

Distribution. — Presently known only from 
Skagerrak, Oslofjord and northern part of 
Swedish west coast. 

Reproduction. —Females with eggs found 
in Sweden in April and August, the former 
immature. Mature eggs bright pink-orange, 
small, about 50-60 wm in diameter. The 
holotype is full of eggs which may explain 
Malm’s choice of specific name; “‘rosea”’ in- 
dicating the color of the live mature female. 

Remarks. —Eliason’s specimens from 
Gullmarsfjord and the Skagerrak-Expedi- 
tion of 1933 constitute a mixture of G. rosea 
and G. mackiei, and have not been re-la- 
belled. His description (Eliason 1962), how- 
ever, clearly is based on specimens of G. 
rosea. Haaland & Schram (1982) provided 
descriptions of both adults and juvenile 
stages from Oslofjord. For previous syn- 
onymy with G. propinqua, see Remarks for 
that species. 


Amphiduros Hartman, 1959 


Amphiduros Hartman, 1959:182 (replace- 
ment name for Amphidromus Hessle, 
1925). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Type species. —Amphidromus setosus 
Hessle, 1925, by subsequent designation 
(Hartman 1959:182). 

Diagnosis (provisional). —Hesionids with 
two palps and two frontal antennae. Median 
antenna present, inserted dorsally on pro- 
stomium. Proboscis without terminal pa- 
pillae. Jaws absent. Segment 5 without no- 
topodia or notosetae, with neuropodia and 
neurosetae. Following parapodia with well- 
developed noto- and neuropodia and large 
number of noto- and neurosetae. Furcate 
notosetae absent. 

Remarks. —Amphiduros is presently sep- 
arated from Gyptis solely on the absence of 
terminal ring of proboscideal papillae. One 
of the states (presence or absence of papil- 
lae) is presumably ancestral, and the genus 
defined on that state may be paraphyletic 
unless supported by other characters. The 
matter warrants further investigation. 


Amphiduros fuscescens (Marenzeller, 1875), 
new combination 
Fig. 13 


Oxydromus fuscescens Marenzeller, 1875: 
143-146, pl.2, fig. 1. 


Material examined.—Italy: 1 syntype, 
(NHMW 2446), Trieste, St. Servola, 3—4 m, 
stones with bore-holes; 1 specimen, eastern 
Sicily, Acitrezza, 37°33.5'N, 15°11.1'E, 42- 
45 m, dredge, muddy sand, 5 Apr 1990; 1 
specimen, eastern Sicily, Brucoli, 37°17.1'N, 
15°12.6’E, 24 m, SCUBA, mixed bottom 
with gravel, boulders and algae, 23 Apr 1990. 
France: 1 specimen (NMW), Banyuls-sur- 
Mer, Ile Grosse, 42°29.0'’N, 03°08.1’E, 10 
m, SCUBA, shell gravel, 13 Oct 1991; 1 
specimen, Banyuls-sur-Mer, Ile Grosse, 
42°29.0'N, 03°08.1'E, 10 m, SCUBA, shell 
gravel, 13 Oct 1991; 2 specimens, Banyuls- 
sur-Mer, Cap Oullestrel, 42°30.22’N, 
03°08.30’E, 18 m, dredge, shell gravel, 15 
Oct 1991. Israel: 4 specimens, Elat, 28°30’N, 
34°34'E, 4 m, SCUBA, 8 Mar 1986. 

Description. —Body, excluding parapo- 
dia, cylindrical, venter flattened, with dis- 
tinct median longitudinal furrow (in larger 


VOLUME 106, NUMBER 1 


D pe Ae byte hit zc 


177 


Fig. 13. Amphiduros fuscescens. A. Prostomium, dorsal view. B. Parapodium segment 5, anterior view, ca. 
half number of setae shown. C. Parapodium segment 6, anterior view, ca. half number of setae shown. D. 
Parapodium segment 14, anterior view, ca. one-third of number of setae shown. E. Spiked capillary notoseta. 
F. Serrated capillary notoseta. G. Median neuroseta. H. Ventral neuroseta. A & E-H specimens from eastern 
Sicily, B—D specimens from Banyuls, southern France. Scales A, 0.5 mm; B—D, 0.25 mm; E-H, 50 um. 


specimens only). Median parapodia only 
slightly longer than anterior ones, posterior 
ones successively shorter, resulting body- 
outline of fairly equal width with slowly ta- 
pered posterior end. 

Prostomium wider than long, anteriorly 
Straight, laterally and posteriorly with 
rounded lobes separated by deep posterior 
incision (Fig. 13A). Palps long and thin, 


proximal parts cylindrical, distal parts ta- 
pering to a point. Proximal and distal parts 
of equal length. Paired antennae situated on 
small ceratophores, as long as palps, slightly 
more slender, with drawn-out tips tapering 
to a point. Median antenna pointed, much 
shorter than paired ones, inserted on line 
between anterior side of anterior pair of eyes. 
Eyes very large with lenses; anterior eyes 


178 


twice as large as posterior. Eye pigment 
poorly delineated, spreading across anterior 
part of prostomium. Nuchal organs lateral 
to prostomium, coalescing dorsally, large 
and distinct. 

Lip glands absent. Proboscis smooth, di- 
vided in proximal and distal parts; proximal 
part slightly larger. Terminal ring of papillae 
lacking, dense ciliation present. One small 
specimen (anterior end, 3.0 mm for 11 seg- 
ments) with (probably) 10 papillae in ter- 
minal ring. 

No specimen observed with complete 
tentacular cirri, but dorsal ones of segments 
3 and 4 much stouter than others. Ventral 
tentacular cirri shorter and thinner than 
dorsal. Cirri of segments 3 and 4 more ven- 
trally displaced than anterior ones. Tentac- 
ular cirri without obvious annulation, all 
situated on long and large cirrophores with 
several (2-6) internal aciculae. 

Anterior segmental delineations uncer- 
tain, segment 1 probably reduced dorsally. 
Notopodia of segment 5 (setiger 1) with two 
notoaciculae situated in cirrophores of dor- 
sal cirri, but without setigerous lobes or se- 
tae (Fig. 13B). Dorsal cirri as stout as those 
of segment 4. Neuropodia and ventral cirri 
similar to following ones. Segment 6 similar 
to median ones (Fig. 13C). 

Notopodia of median segments with short 
dorsal cirri, lacking apparent annulation; 
subdistally widened and tapering to a point 
(Fig. 13D). Notopodial lobes conical, with 
one or two internal aciculae, without emerg- 
ing acicular setae. Large number of capillary 
setae (ca. 20-30) inserted behind lobe, with 
smooth proximal parts and double rows of 
small teeth medially and subdistally (Fig. 
13E). A few, ventrally situated serrated no- 
tosetae present (Fig. 13F). 

Neuropodia of median segments conical, 
with one or two internal aciculae and a large 
number of stout compound neurosetae. 
Distal part of setal shafts internally reticu- 
lated. Blades of median (Fig. 13G) and dor- 
sal setae much longer than ventral (Fig. 
13H). Ventral cirri smooth, tapering to a 
point; inserted distally without cirrophores. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Pygidium rounded. Pygidial cirri not ob- 
served, median papilla absent. 

Color: Live specimens with white pig- 
ment spots distally on dorsal cirri. Eyes or- 
ange. Eggs dark red. Preserved specimens 
yellowish, white pigment spots and red col- 
or of eggs disappear. 

Measurements: No entire, non-regener- 
ating Mediterranean specimens observed. 
Measurements for most complete speci- 
mens: length 15 mm, width 3.3 mm for 28 
segments (posterior end lacking); length 6.2 
mm, width 1.5 mm for 27 segments (regen- 
erating). Two of the Gulf of Aqaba speci- 
mens (see Remarks) are complete: length 14 
mm, width 3.2 mm for 40 segments; length 
10 mm, width 2.4 mm for 37 segments. 

Habitat. —Stones and coarse shell gravel 
from 2-45 m. One specimen from Sicily 
found in muddy sand. 

Distribution. —Southern France, eastern 
Sicily, northern Adriatic, Gulf of Aqaba (see 
Remarks). 

Reproduction. —Females with eggs found 
in May at Sicily and March in Gulf of Aga- 
ba, about 100-110 um in diameter. 

Remarks.—The newly collected speci- 
mens are in good agreement with Maren- 
zeller’s type. According to his original de- 
scription this species should have black 
rather than orange eyes. What is remaining 
of eye pigment on his type, however, sug- 
gests reddish (or orange) rather than black 
eyes. 

The Gulf of Aqaba specimens are in good 
condition and could not be distinguished 
from Mediterranean ones. Common Med- 
iterranean and Red Sea distribution may, 
however, not be very common, and until 
further material is available these speci- 
mens are labelled Amphiduros cf. fusces- 
cens. 

Fauvel (1923) and Hartman (1959) syn- 
onymized A. fuscescens (as Oxydromus) with 
Gyptis propinqua. This is obviously incor- 
rect, and the species differ in many respects, 
most notably size, proboscideal papillation, 
and annulation, shape and length of dorsal 
and ventral cirri. 


VOLUME 106, NUMBER 1 


Key to European Species of 
Amphiduros and Gyptis 


1. Prostomium wider than long, lip 
glands absent, ventral cirri inserted 
distally 

— Prostomium as wide as long, lip 
glands present, ventral cirri inserted 
subdistally 

2. Eyes small with well delineated pig- 
ment, adults (> ca. 20 segments) 
with terminal papillae on proboscis, 
dorsal cirri distinctly annulated and 
tapering evenly to a point, acicular 
notosetae present 

— Eyes large with poorly delineated 
pigment, adults without terminal 
papillae on proboscis, dorsal cirri 
subdistally widened, not annulated, 
acicular notosetae absent A. fuscescens 

3. Median antenna with distinct, well 
delineated tip, eyes brownish-black, 
proboscis of adults (> ca. 25 seg- 
ments) with more than 35 papillae 
in terminal ring, dorsal cirri reach- 
ing farther than setae, distinct ele- 
vated dorsal ridges absent ... G. rosea 

— Median antenna without delineated 
tip, eyes red, proboscis of adults with 
less than 35 papillae in terminal ring, 
dorsal cirri not reaching farther than 
setae, distinct elevated dorsal ridges 
PCESEh Oe ee G. mediterranea 

4. Median antenna widest subdistally, 
adults (> ca. 20 segments) with neu- 
rosetae from segment five, dorsal 
cirri much longer than setae ..... 

oo ee a eee G. propinqua 

— Median antenna widest medially, 
adults with neurosetae from seg- 
ment four, dorsal cirri much shorter 
MAPIMESELAGCL. $e2%.. do ile ctr ac G. mackiei 


Acknowledgments 


I wish to thank K. Fauchald and L. Ward 
(USNM), K. Fitzhugh and L. Harris 
(LACM), G. Hartwich (ZMB), G. V. Hel- 
gason, E. Kritscher (NHMW), A. Mackie 
(NMW), A. Muir (BMNH), A. Norrevang 


179 


(BIOFAR), L. Orrhage (NHMG), and L. 
Wallin (ZMUU) for access to material and/ 
or working facilities, the crews of R/V Mi- 
mir and R/V Nereus for field assistance, and 
the staffs of Laboratoire Arago and Tjarno 
Marine Biological Laboratory where part of 
the work was conducted. Special thanks to 
P. Bouchet for admitting a polychaete work- 
er to join the ““Fifth European Malacological 
Workshop” at Sicily, to G. V. Helgason for 
hostship on Iceland, and to A. Mackie for 
field collaboration, discussions as well as 
comments on the manuscript. Financial 
support was provided by Helge Ax:son 
Johnsons Stiftelse and the Swedish Natural 
Science Research Council (contracts 9555- 
302 & -307). 


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Bhaud, M. 1971. La phase larvaire d’Oxydromus pro- 
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Blake, J. A. 1975. The larval development of Poly- 
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Day, J. H. 1963. The polychaete fauna of South Af- 
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1913. Die Polychaeten-Sammlungen der 
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, & D. R. Hancock. 1981. Deep-water poly- 

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Allan Hancock Foundation Monograph 11:1- 

73: 

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Gibbs, P. E. 1971. The polychaete fauna of the Solo- 
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—., & K. Probert. 1973. Notes on Gyptis capensis 
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Gravina, M. F., & A. Giangrande. 1988. A new rec- 
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Grube, A. E. 1855. Beschreibung neuer oder wenig 

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talem et Americam centralem annis 1845-1848 

suscepto legit cl. A. S. Orsted, adjectis speciebus 

nonnullis a cl. H. Kréyero in itineread Ameri- 
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Haaland, B., & T. A. Schram. 1982. Larval devel- 
opment and metamorphosis of Gyptis rosea 
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——, & 1983. Larval development and 
metamorphosis of Ophiodromus flexuosus (Delle 
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Hartman, O. 1959. Catalogue of the polychaetous 

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1961. Polychaetous annelids from Califor- 
nia.— Allan Hancock Pacific Expeditions 25:1- 
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1965. Catalogue of the polychaetous Anne- 
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index.—Allan Hancock Foundation Publica- 
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Hartmann-Schroder, G. 1959. Zur Okologie der 
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na 1:69-183. 

1965. Zur Kenntnis des Sublitorals der chi- 
lenischen Kiiste unter besonderer Beriicksichti- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


gung der Polychaeten und Ostracoden. Teil II. 

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aus dem Hamburgischen Zoologischen Muse- 

um und Institut 62:59-305. 

1971. Annelida, Borstenwirmer, Polychae- 
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marks on some previously recorded species. —Sar- 
sia 75:203-212. 

Hessle, C. 1925. Einiges tiber die Hesioniden und die 
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Zoologi 17(10):1-36. 

Hilbig, B., & J.-D. Dittmer. 1979. Gyptis helgolan- 
dica sp. n. (Hesionidae, Annelida), eine neue 
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schen Bucht.— Ver6ffentlichungen des Instituts 
fiir Meeresforschung in Bremerhaven 18:101- 
110. 

La Greca, M. 1946. Studii sui policheti del Golfo di 
Napoli.— Pubblicazioni della Stazione zoologi- 
ca di Napoli 20:270—280. 

Laubier, L. 1961. Podarkeopsis galangaui n. g., n. 
sp., hésionide des vases cotiéres de Banyuls-sur- 
Mer.— Vie et milieu 12:211-217. 

Malm, A. W. 1874. Annulater i hafvet utmed Sveri- 
ges vestkust och omkring Gdteborg.—G6te- 
borgs K. vetenskaps- och vitterhetssamhalles 
handlingar 14:67-105. 

Marenzeller, E. 1875. Zur Kenntniss der adriatischen 
Anneliden. Zweiter Beitrag. (Polynoinen, He- 
sioneen Syllideen.).—Sitzberichte der Akade- 
mie der Wissenschaften in Wien 72:129-171. 

Marion, A. F., & N. Bobretzky. 1875. Etude des An- 
nélides du Golfe de Marseille. — Annales des sci- 
ences naturelles 2:1-106. 

Sars, M. 1862. Uddrag af en af detaillerade Afbild- 
ningar ledsaget udforlig Beskrivelse over fol- 
gende Norske Annelider.— Forhandlinger 1 Vi- 
denskabsselskabet i Kristiania 1862:87-95. 

Schlegel, H. 1854. Ook en woordje over den dodo 
(Didus ineptus) en zijne verwanten.— Verslagen 
en Mededeelingen der Koninklijke Akademie 
van Wetenschappen 2:233-256. 

Schram, T. A., & B. Haaland. 1984. Larval devel- 
opment and metamorphosis of Nereimyra punc- 
tata (O. F. Miiller) (Hesionidae, Polychaeta).— 
Sarsia 69:169-181. 

Storch, V., & R. Niggemann. 1967. Auf Echinoder- 
men lebende Polychaeten.—Kieler Meeresfor- 
schungen 23:156-164. 

Ushakov, P. V. 1955. Polychaeta of the Far Eastern 
Seas of the U.S.S.R.—Academiya Nauk SSSR, 
Keys to the Fauna of the SSSR 56:1-433 (trans- 
lated from Russian by the Israel Program for 
Scientific Translations, Jerusalem 1965). 


VOLUME 106, NUMBER 1 


Webster, H. E., & J. E. Benedict. 1887. The Annelida 
Chaetopoda, from Eastport, Maine.— Report of 
the United States Commissioner of Fisheries 
1885:707-755. 

Willey, A. 1902. Polychaeta. Pp. 262-283 in Report on 
the collections of natural history made in the Ant- 
arctic regions during the voyage of the “Southern 
Cross.” William Clowes and Sons, Limited, Lon- 
don, 344 pp. 


Swedish Museum of Natural History, 
Stockholm, and (postal address): Tjarno 
Marine Biological Laboratory, Pl. 2781, 
S-452 96 Stromstad, Sweden. 


Appendix 


Checklist of species and subspecies of Amphiduros- 
and Gyptis, with notes on material examined. 


Amphiduros axialensis Blake & Hilbig, 1990. Hydro- 
thermal vents of northeast Pacific, Juan de Fuca 
Ridge. Type material NMCA. No material ex- 
amined. 

A. fuscescens (Marenzeller, 1875), new combination. 
As Oxydromus fuscescens. Muggia, Trieste, 
Adriatic. Syntype (NHMW) and non-types 
(SMNH, NMW) examined. 

A. izukai (Hessle, 1925). As Amphidromus izukai. Ja- 
pan. Syntype (ZMUU) examined. Synonymous 
to A. setosus (Hessle, 1925). 

A. pacificus Hartman, 1961. California. Holotype 
(LACM) and non-types (LACM) examined. 

A. setosus (Hessle, 1925). As Amphidromus setous. Ja- 
pan. Syntype (ZMUU) examined. Synonymous 
to A. izukai (Hessle, 1925). 

Gyptis brunnea (Hartman, 1961). As Oxydromus brun- 
nea. California. Holotype (LACM) examined. 

G. helgolandica Hilbig & Dittmer, 1979. North Sea. 
Non-types (SMNH) examined. Belongs to Po- 
darkeopsis. 

G. hians Fauchald & Hancock, 1981. Off Oregon. Ho- 
lotype (LACM), paratypes (LACM) and non- 
types (LACM) examined. 

G. incompta Ehlers, 1912 (complementary description 
based on other specimens in Ehlers 1913). Kai- 
ser Wilhelm II Land or Victoria Land, Antarc- 
tic. Type material lost? Non-types (ZMH, 
SMNH) examined. 

G. lobata (Hessle, 1925). As Oxydromus lobatus. Ja- 
pan. Syntype (ZMUU) examined. Synonymous 
to G. pacifica. 

G. maraunibinae Gibbs, 1971. Solomon Islands. Ho- 
lotype (BMNH) examined. Belongs to Podar- 
keopsis. 


181 


G. ophiocomae Storch & Niggemann, 1967. Red Sea. 
Type material in author’s collection (Storch). 
Non-type (SMNH) examined. 

G. pacifica (Hessle, 1925). As Oxydromus pacificus. 
Japan. Syntype (ZMUU) examined. Synony- 
mous to G. lobata. 

G. propinqua Marion & Bobretzky, 1875. Marseille, 
France. No type material. Non-types (SMNH) 
examined. 

G. raluanensis (Augener, 1927). As Oxydromus ralu- 
anensis. New Guinea. Syntype (ZMH) exam- 
ined. 

G. rosea (Malm, 1874). As Ophiodromus roseus. Gull- 
marsfjorden, Sweden. Holotype (NHMG) and 
non-types examined (NHMG, NHMR). 

G. vittata Webster & Benedict, 1887. Maine, U.S. Syn- 
types (USNM) examined. 

Ophiodromus roseus Malm, 1874. See Gyptis rosea. 

Oxydromus arenicolus La Greca, 1946. No type ma- 
terial? No material examined. Belongs to Po- 
darkeopsis. 

O. arenicolus glabrus Hartman, 1961. Holotype 
(LACM) examined. Belongs to Podarkeopsis. 

O. aucklandicus Willey, 1902. Auckland Islands. Type 
material lost. No material examined. Genus un- 
certain (not Gyptis or Amphiduros). Nomen du- 
bium. 

O. brevipalpa Hartmann-Schréder, 1959. Type mate- 
rial ZMH. No material examined. Belongs to 
Podarkeopsis. 

O. brunnea Hartman, 1961. See Gyptis brunnea. 

O. capensis Day, 1963. Holotype (BMNH) examined. 
Belongs to Podarkeopsis. 

O. fasciatus Grube, 1855. Trieste, Italy, or Ville- 
franche, southern France. Syntype (ZMB) ex- 
amined. Junior synonym to Ophiodromus flex- 
uosus (Delle Chiaje, 1827). 

O. flaccidus Grube, 1857. St. Croix, West Indies. Type 
material probably lost. No material examined. 
Nomen dubium. 

O. fuscescens Marenzeller, 1875. See Amphiduros fus- 
cescens, new combination. 

O. heteroculatus Hartmann-Schroder, 1965. Valdivia, 
Chile. Type material ZMH. No material ex- 
amined. Genus uncertain (not Gyptis or Am- 
phiduros). 

O. lobatus Hessle, 1925. See Gyptis lobatus. 

O. longisetis Grube, 1857. St. Croix, West Indies. Type 
material probably lost. No material examined. 
Nomen dubium. 

O. pacificus Hessle, 1925. See Gyptis pacifica. 

O. pallidus Claparéde, 1864. No type material. Non-types 
(SMNH) examined. Belongs to Podarke. 

O. raluanensis Augener, 1927. See Gyptis raluanensis. 


PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 182-189 


A NEW GENUS OF HYDROBIID SNAILS 
(MOLLUSCA: GASTROPODA: PROSOBRANCHIA: RISSOOIDEA) 
FROM NORTHERN SOUTH AMERICA 


Robert Hershler and France Velkovrh 


Abstract.—An aquatic hydrobiid cavesnail from Andes Mountains of Co- 
lombia and Ecuador is described (Andesipyrgus sketi, new genus, new species). 
The genus is placed in the Cochliopinae based on females having a sperm tube 
separated from the glandular gonoduct. Diagnostic features of Andesipyrgus 
include a minute, near-pupiform shell with thickened aperture; unpigmented 
animal (except for eyespots); reduced ctenidium; female genitalia featuring 
complex histology of glandular gonoduct, oviduct coiled onto right side of 
albumen gland, and absence of seminal receptacle; and male with simple, 
slender penis. Andesipyrgus does not closely resemble other cochliopinids hav- 
ing a simple penis, and appears remotely related to other South American 


hydrobiid fauna. 


The aquatic prosobranch snails of the 
family Hydrobiidae comprise a large cos- 
mopolitan group of several hundred genera 
and a few thousands of Recent species. While 
the group is highly diverse at lower taxo- 
nomic levels in most of the major physio- 
graphic regions where it occurs, one of the 
few exceptions is South America, where only 
seven genera (and about 120 Recent species) 
are found. (North America has about 40 
genera and over 200 species by compari- 
son.) Of these, each of five are represented 
by less than 10 species on the continent, 
while the remaining two (Heleobia Stimp- 
son, 1865; Potamolithus Pilsbry, 1896) have 
radiated extensively. The apparent paucity 
of hydrobiid fauna in South America surely 
relates in part to hydrographic/geologic and 
historical features of the continent, but also 
probably reflects inadequate sampling of the 
brackish coastal and inland waters of this 
huge land mass. 

One habitat that has been especially ne- 
glected in the search for these animals in 
South America is cave streams, whose hy- 
drobiid fauna often is diverse and strongly 
differentiated from local epigean (surface- 
dwelling) taxa. While caves are not extreme- 


ly numerous in the continent, there are large 
areas of karstic terrain to the north (es- 
pecially in the Andes Mountains and in the 
Amazon basin of Brazil) which are poten- 
tially fertile areas for discovery of cavesnail 
fauna. A recent compendium of subterra- 
nean aquatic mollusks (Bole & Velkovrh 
1986) listed only a single species of hydro- 
biid cavesnail from South America, which 
was collected from the Andes Mountains of 
Ecuador by a Yugoslavian expedition dur- 
ing 1978 (Sket 1985). Restudy of this ma- 
terial and other collections made by a Yu- 
goslavian expedition to the Colombian 
Andes in 1984 (Sket 1988) confirmed the 
uniqueness of this animal, which we de- 
scribe as a new genus and species below. 

Material studied is from the personal col- 
lection of the second author (FV). Types 
have been deposited in the National Mu- 
seum of Natural History, Smithsonian In- 
stitution (USNM). 


Family Hydrobiidae Troschel, 1857 
Subfamily Cochliopinae Tryon, 1866 
Andesipyrgus, new genus 


Type species.—Andesipyrgus sketi, new 
species. 


VOLUME 106, NUMBER 1 


Diagnosis. —Shell minute-small, smooth, 
narrow, with thickened aperture. Opercu- 
lum corneous, thin, with eccentric nucleus 
and slightly thickened ventral attachment 
scar. Radula taenioglossate; central teeth 
with two pairs of basal cusps, marginal and 
lateral teeth with relatively numerous, fine 
cusps. Rectum with bend or loop in pos- 
terior pallial cavity. Stomach without pos- 
terior caecum. Animal unpigmented except 
for eyespots. Ctenidium reduced in length, 
and with relatively few filaments. Females 
Oviparous; glandular gonoduct of complex 
histology. Oviduct coiled behind (onto right 
side) of albumen gland, bursa copulatrix of 
posterior position, seminal receptacle ab- 
sent. Bursal duct opening to sperm tube, 
which has an anterior connection to the cap- 
sule gland. Male prostate gland with prom- 
inent pallial section. Penis slender, simple, 
without lobes or enlarged glands. 

Remarks. —The pupiform shell with 
thickened aperture of Andesipyrgus (similar 
to that of unrelated Bythinella Moquin- 
Tandon, 1855, from Europe) does not re- 
semble that of any other South American 
hydrobiid. A remote affinity with other fau- 
na of the region is further suggested by the 
configuration of female genitalia (notably 
the coiled oviduct), which is unique in the 
Cochliopinae (and substantially different 
from the usual pattern of a single coil of 
oviduct on the left side of the albumen 
gland). The simple penis (without terminal 
constriction or large-sized glands) of An- 
desipyrgus is shared by 10 other cochliopi- 
nid genera, including several other subter- 
ranean forms. This is considered a 
phylogenetically diverse group (almost cer- 
tainly representing a grade of organization), 
whose genera are of uncertain relationship 
both to one another (in most cases) and to 
other cochliopinid groups (Hershler & 
Thompson 1992). Andesipyrgus does not 
closely resemble any of these genera and 
further evaluation of its affinities cannot be 
made at this time. 

Several morphological features of Ande- 


183 


sipyrgus, including pigmented eyespots and 
brown periostracum, suggest that this ani- 
mal only recently invaded cave waters. 

Etymology.—From Andes, referring to 
distribution along the Andes Mountains, and 
Classical Greek pyrgos, meaning tower and 
referring to the moderately elongate shell. 
Gender masculine. 


Andesipyrgus sketi, new species 
Figs. 1-5, Table 1 


Littoridina(?) jumandi Bole & Velkovrh 
1986:196. [Nomen nudem.] 

‘““Hydrobioidea-Gastropoda.”—Sket 1985: 
84. 

““Hydrobioidea.’’—Sket 1988:55, 58. 


Material examined. —Colombia (Santan- 
der Department): La Paz area (6°11’N, 
73°35'W)—Cueva de los Indios, ca. 6 km 
SW of La Paz, 1995 m elevation, VI-1984, 
FV 43575, FV 43689, USNM 860574 (ho- 
lotype), USNM 860575 (paratypes); Hoyo 
(de) Colombia, ca. 6 km SSE of La Paz, 1775 
m (1 broken shell), VI-1984, FV 43691; 
Hoyo del Aire, ca. 6 km SSW of La Paz, 
1800 m, VI-1984, FV 43690. San Gil area 
(6°33’'N, 73°08’W)— Cueva del Indio, ca. 7.5 
km S of San Gil, 1270 m, VI-1984, FV 
43685. Ecuador: Cavernas de Jumandi, near 
Archidona (0°55'S, 77°48'W), 140 km SW 
of Quito, Napo Province, about 500 m, XII- 
1978, FV 38129, FV 38130. 

Description. —Shell (Figs. 1, 2a) narrowly 
conic to pupiform, 1.6-—2.5 mm tall, with 
4.0-4.5 whorls. Protoconch (Fig. 2b) blunt, 
smooth except for very faint pattern of low 
wrinkles. Teleoconch whorls near flat to 
slightly convex, rarely with slight adapical 
shoulders and/or weak adapical angulation; 
sutures very shallow. Teleoconch sculpture 
of moderately strong growth lines. Aperture 
ovate, less than 50% of shell height, broadly 
adnate or very slightly separated from body 
whorl, usually chalk-white (possibly due to 
lengthy preservation in alcohol). Inner lip 
complete, moderately reflected, thickened, 
sometimes markedly so in parietal region. 


184 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Andesipyrgus sketi, holotype, USNM 860574, standard and side views. Shell height, 2.0 mm. 


Outer lip thick, slightly expanded, weakly 
sinuate, near orthocline. Umbilicus absent 
or very narrowly rimate. Periostracum 
brown. 

Measurements (mm) of the shells of the 
holotype and nine paratypes are in Table 1. 


Operculum (Fig. 2c-e) light brown, trans- 
parent, ovate, paucispiral. Attachment scar 
with weakly developed central callus and 
slight thickening along inner margin. 

Radular ribbon about 470 um long and 
70 um wide, of about 66 rows of teeth, 


Table 1.—Shell measurements (mm) for Andesipyrgus sketi. SH = standard height, HBW = height of body 
whorl, SW = standard width, AH = aperture height, AW = aperture width, WH = number of whorls. 


SH HBW SW 
Holotype 2.0 5S p22 
Paratypes Pps | 1.4 | was 
Dee 5 ge, 
2.0 b3 1.1 
DD iS bed, 
oe | 1.4 1:2 
p94 | Ws LS 
pi | 1.4 FZ 
pp! | a) 1.2 
Det B35 12 


20 1.4 12 


AH AW WH SW/SH 
0.87 0.79 4.0 0.58 
0.83 0.81 4.25 0.58 
0.91 0.87 4.5 0.55 
0.75 0.73 4.5 0.55 
0.81 0.73 4.5 0.54 
0.87 0.81 4.25 0.58 
0.89 0.85 4.5 0.58 
0.83 O19 4.5 0.57 
0.85 0.89 4.25 0.54 
0.81 0.83 4.25 0.56 
0.85 0.75 4.0 0.61 


VOLUME 106, NUMBER 1 


185 


Fig. 2. Scanning electron micrographs of shell and opercula of A. sketi, USNM 860575. a. Shell (height, 2.1 
mm). b. shell apex (bar = 100 wm). c. Operculum, dorsal surface (bar = 176 um). d, e. Operculum, ventral 
surface (bar = 200 um). 


scarcely extending beyond edge of well-de- 
veloped buccal mass and without posterior 
coil. Central teeth (Fig. 3a, b) trapezoidal, 
with well indented dorsal edge; lateral an- 
gles narrow, slightly thickened, well ex- 
panded and sometimes broadened distally. 
Lateral cusps of central teeth narrow, 4—5; 
central cusp slightly longer than laterals; 
basal cusps moderate to long, arising from 


face of tooth near origin of lateral angles, 
inner cusp much broader and slightly longer 
than outer. Basal process of central teeth 
narrow, well excavated. Lateral tooth (Fig. 
3d) formula, 2(inner)/1/3—4(outer); cusps 
narrow, curved, with central cusp enlarged. 
Marginal teeth (Fig. 3c, d) with numerous 
(about 17-20) cusps. Rectum broadly over- 
lapping glandular gonoducts, with bend (fe- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


(Ge ae 
Fig. 3. Radula of A. sketi, USNM 860575. a. Central teeth (bar = 6 um). b. Central teeth (bar = 10 um). c. 
Outer marginal teeth (bar = 10 um). d. Lateral and inner marginal teeth (bar = 13.6 um). 


males) or pronounced U-shaped loop cles with narrow central band of elongate 

(males) in posterior portion of pallial cavity; cilia on dorsal surface. 

anus near mantle edge. Ctenidium filling about 66% of pallial 
Animal white (except for black eyespots), cavity length, extending from well anterior 

without melanin pigment. Cephalic tenta- to pericardium almost to mantle edge. Cte- 


VOLUME 106, NUMBER 1 


nidial filaments about 12, short and narrow. 
Osphradium narrowly ovate, about 33% of 
ctenidium length, positioned along poste- 
rior half of ctenidial axis. Kidney with slight 
bulge into pallial cavity; kidney opening 
thickened, white. Hypobranchial gland 
weakly developed. 

Ovary a small unlobed mass filling about 
0.25 whorls and slightly overlapping pos- 
terior stomach chamber. Albumen gland 
(Fig. 4a, Ag) with large pallial section (> 50% 
of length). Capsule gland (Cg) about equal 
to albumen gland in length, of two tissue 
sections (posterior, orange; anterior, white). 
Capsule gland composed of folded glandu- 
lar cells, thick-walled, with central lumen. 
Coiled oviduct (Co) bends posteriorly be- 
hind pallial wall, loops behind albumen 
gland to right side, then twists back to left 
side of gland, where it coils once before 
looping to right side of bursal duct (Dbu) 
before joining the duct at the pallial wall. 
Coiled oviduct with thin muscular coat, 
strongly ciliated. Bursa copulatrix (Bu) small 
(about 33% of albumen gland length), ovate, 
scarcely extending posterior to albumen 
gland. Bursa of tall glandular cells having 
basal nucleii, lined with thick muscular coat. 
Bursal duct emerging from anterior tip of 
bursal copulatrix; distal section of duct 
broadened, with an internal division into 
two sections suggesting an enclosed seminal 
receptacle. (Study of thin sections could not 
confirm this possibility.) Duct to albumen 
gland (Dag) opening posteriorly from point 
where bursal duct and coiled oviduct join. 
Sperm tube (St) broad, thick, muscular, 
strongly ciliated; positioned ventro-laterally 
to capsule gland, joining capsule gland dis- 
tally. Genital aperture (Ga) a short terminal 
slit. 

Testis an unlobed mass, orange-colored, 
filling about 0.5 whorl posterior to stomach. 
Prostate gland narrow, bean-shaped; walls 
thick, lumen slit-like; pallial section prom- 
inent (about 60% of gland length). Posterior 
vas deferens opening to prostate gland just 
behind pallial wall; anterior vas deferens 


187 


Fig. 4. Genitalia of A. sketi, USNM 860575. a. Left 
side of female glandular oviduct (bar = 0.5 mm). The 
thick curving line represents the posterior wall of pallial 
cavity. In the drawing to the right (slightly enlarged), 
the coiled oviduct has been cut and rotated to expose 
the bursa copulatrix and associated ducts. Abbrevia- 
tions: Ag, albumen gland; Bu, bursa copulatrix; Cg, 
capsule gland; Co, coiled oviduct; Dag, duct to albu- 
men gland; Dbu, bursal duct; Ga, genital aperture; St, 
sperm tube. b. Male penis, dorsal surface (bar = 0.25 
mm). 


opening subterminally. Pallial section of vas 
deferens with proximal coil. Penis (Fig. 4b) 
small, tightly coiled, positioned well behind 
cephalic tentacles slightly to nght of center 
of “neck.” Penis vermiform, unciliated, 
slightly broadened distally, with small folds 
on inner curvature near base. Distal tip of 
penis strongly tapered, vas deferens opening 


188 


TROPICAL AMERICA 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Flora Neotropica base map no 1 


© 200 400.600 800 1000km 
SSSS_—__SS= 


ee 
QO 100 200-300 400 500 600 miles 


© 1979 by the University of Utrecht 


Fig. 5. 


= aE 
2 AS 


oa : 
C20 


ae 


Published by the State University of Utrecht, the Netherlands Department of Systematic Botany 


Map showing distribution of A. sketi. 1, Colombian localities near La Paz and San Gil, Santander 


Department; 2, Ecuador locality near Archidona, Napo Province. 


through short terminal papilla. Proximal 
course of vas deferens not discernable. 

Type locality. —Stream flowing into Cueva 
de los Indios, vicinity of La Paz, Santander 
Department, Colombia. 

Distribution.—A trans-Andean species, 
occurring in cave streams of Rio Magdalena 
basin (draining to Caribbean), Colombia; 
and Rio Napo basin (Amazon basin), Ec- 
uador. 

Habitat.—Andesipyrgus sketi occurs in 
small Andean cave streams at elevations be- 
tween 500-1990 m. Water temperatures 
among sites varied from 15—23°C. Some of 
the cave streams represent sinking rivers 


whereas others probably originate from per- 
colating waters. All of the streams contained 
some surface fauna, particularly insect lar- 
vae (Chironomidae, Elminthidae, etc.) and 
cyclopoid Copepoda, and most also were 
inhabited by more or less troglomorphic an- 
imals: variable catfish populations (Tricho- 
myoterus sp. in Colombia, Astroblepus pho- 
leter Collette, in Ecuador) and probable 
troglobiont, amphibious crabs (Neostren- 
geria sketi Rodriguez, in Colombia) or am- 
phipods (Bogidiella gammariformis Sket, in 
Ecuador). Snails were most common in the 
only cave lacking troglomorphic fauna 
(Cueva de los Indios) and in which Tricho- 


VOLUME 106, NUMBER 1 


myoterus was normally pigmented and 
shaped. For additional details, the reader is 
referred to Sket (1979, 1988). 

Remarks.— Available material was un- 
relaxed, and details of shape of head and 
cephalic tentacles could not be discerned. 

Populations varied in several shell fea- 
tures (width, angulation of aperture, whorl 
outline) and while available material does 
not permit meaningful analysis, further 
study may reveal that a species complex is 
involved. 

Etymology.—Named in honor of Dr. 
Boris Sket, for his discovery of this species 
and for his encouragement of this collabo- 
rative study. 


Acknowledgments 


The authors thank Dr. Boris Sket for 
making collections of the Yugoslavian cav- 
ing expeditions available for study and for 
providing useful comments on the manu- 
script. Scanning electron micrographs were 
prepared by Susann Braden of the Scanning 
Electron Microscopy Laboratory, National 
Museum of Natural History. Shells were 
drawn by Molly Ryan, Dept. of Invertebrate 
Zoology, National Museum of Natural His- 


189 


tory. Anatomical illustrations were inked by 
Susan Escher. 


Literature Cited 


Bole, J., & F. Velkovrh. 1986. Mollusca from con- 
tinental subterranean aquatic habitats. Pp. 177- 
208 in Stygofauna mundi, a faunistic, distri- 
butional, and ecological synthesis of the world 
faunas inhabiting subterranean waters (includ- 
ing the marine interstitial). E. J. Brill/Dr. W. 
Backhuys, Leiden, 740 pp. 

Hershler, R., & F.G. Thompson. 1992. A review of 
the genera of the aquatic gastropod subfamily 
cochliopinae (Prosobranchia: Hydrobiidae).— 
Malacological Review Supplement 5:1-—140. 

Sket, B. 1979. Fauna in the Caverna de Jumandi.— 

Nase Jame 20:85-91. 

1985. Bogidiella (s.1.) gammariformis sp. n. 
(Amphipoda) from Ecuador. —Bioloski Vestnik 
33:81-88. 

1988. Speleobiological investigations in the 
Colombian Andes 1984.—Bioloski Vestnik 36: 
53-62. 


(RH) Department of Invertebrate Zool- 
ogy, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 
20560, U.S.A.; (FV) Department of Biolo- 
gy, Biotechnical Faculty, University of Lju- 
bljana, ASker¢éeva 12, 61001 Ljubljana, Slo- 
venia. 


PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 190-194 


VAMPYROCROSSOTA CHILDRESST, A NEW GENUS 
AND SPECIES OF BLACK MEDUSA FROM THE 
BATHYPELAGIC ZONE OFF CALIFORNIA 
(CNIDARIA: TRACHYMEDUSAE: RHOPALONEMATIDAE) 


Erik V. Thuesen 


Abstract.—A new genus and species of deep-sea medusa, Vampyrocrossota 
childressi, is described from the eastern North Pacific. It has been found in San 
Clemente Basin off Baja California, Mexico, and from the waters off Point 
Conception, California, U.S.A., at depths between 600 and 1475 meters. This 
genus is allied to the cosmopolitan rhopalonematid genus Crossota, but differs 
notably in the shape and position of the gonads. It is the only described species 
of hydromedusa with black pigmentation. 


During the course of an ongoing project 
on the physiology and biochemistry of mid- 
water gelatinous organisms off California, a 
distinctive black medusa was commonly re- 
covered in trawls taken deeper than 600 m. 
The animals were collected by a 10 m? 
Mother Tucker trawl using a specially de- 
signed 30 1 insulated cod end to protect the 
animals from heat and light as they are 
brought to the surface (Childress et al. 1978). 
Medusae were captured in very good con- 
dition, and fragile hydromedusae of the 
families Halicreatidae and Rhopalonema- 
tidae were often brought aboard ship with 
tentacles several body heights in length. The 
black medusa is not included in reports of 
Pacific Ocean hydromedusae (Alvarino 
1967, Kramp 1968, Segura-Puertas 1984), 
and it is not one of the several new species 
of mesopelagic rhopalonematid medusae 
currently being described by Mills & Larson 
(C. E. Mills, pers. comm.). 


Vampyrocrossota, new genus 
Figs) 12 


Diagnosis. —Rhopalonematidae without 
gastric peduncle; stomach with four oral lips, 
extending to or just past the velum when 
empty; with eight tubular gonads attached 


longitudinally to the eight radial canals in 
all the specimens observed; exumbrellar 
furrows present; with tentacles all of one 
kind. 

Type species. —Vampyrocrossota chil- 
dressi, new species. 

Etymology.—From Serbian vampira, a 
nocturnal demon supposed to eat the heart, 
blood and soul of its victim, with reference 
to Vampyroteuthis infernalis, the black me- 
sopelagic squid often captured in the same 
trawls as this animal, and Crossota the 
closely allied rhopalonematid genus. 

Relationships. — Both Bigelow (1913) and 
Kramp (1947) discuss the genus Crossota 
Vanhoffen, 1902 in some detail and are ex- 
plicit that the pendant nature of the gonads 
is an important characteristic distinguishing 
this genus from other related genera. Given 
the importance that this characteristic has 
had in conserving the genus Cyvossota, I have 
erected the genus Vampyrocrossota in this 
paper. This genus is closely related to Cros- 
sota in general appearance, lack of a pedun- 
cle, the large number of tentacles and rib- 
bon-like nature of the radial canals. 

Recently, Larson & Harbison (1990) es- 
tablished the new rhopalonematid genus 
Benthocodon which also differs from Cros- 
sota by having gonads attached to the radial 


VOLUME 106, NUMBER 1 


canals. They reported that the gonads in B. 
hyalinus are ribbon-like and run along most 
of the length of the eight gastric canals with 
the most distal portions hanging free. Vam- 
pyrocrossota is also different from Bentho- 
codon in that it lacks a gastric peduncle and 
has numerous exumbrellar furrows. 


Vampyrocrossota childressi, new species 
Figs. 1, 2 


Types. —Holotype: a 12 mm tall speci- 
men (USNM 91883) taken from 777 m 
depth on 31 July 1991 off Point Conception 
from the RV Point Sur. Paratypes: two spec- 
imens, 11 mm (Paratype A: USNM 91884) 
and 6 mm (Paratype B: USNM 91885) from 
984 m depth, captured on 30 July 1991 off 
Point Conception, California. All types are 
deposited in the National Museum of Nat- 
ural History, Smithsonian Institution. 

Description.—This description is based 
upon observations of ~20 living animals 
ranging in size from 6 to 14 mm in height 
and up to 475 mg wet weight. Up to 14 mm 
tall; up to 10 mm in diameter; velum up to 
3 mm; jelly fairly thin, especially at apex; 
mesoglea colorless; numerous exumbrellar 
furrows; inner surface of bell is black with 
pigment fading out posteriorly towards the 
velum (particularly in immature animals); 
velum is black in some specimens; up to 
400 tentacles, all the same kind; tentacles 
and radial canals reddish-orange; eight 
cream-colored tubular gonads attached one- 
eighth from the top to five-eighths the length 
of the radial canal; stomach without pedun- 
cle, reaching past the velum when extended; 
stomach cream colored with a wide hori- 
zontal black pigmented band located half 
way to the four oral lips. The immature 
paratype specimen had a completely orange 
manubrium when it was collected before 
preservation in formalin. Upon first in- 
spection with the naked eye, this species 
looks remarkably like Crossota rufobrunnea 
with black rather than burgundy pigmen- 
tation. Vampyrocrossota childressi is much 


191 


less active after capture and has a lower met- 
abolic rate than many other rhopalonema- 
tids which have been captured in the same 
trawls. These other species include Crossota 
alba, C. rufobrunnea, Pantachogon sp., 
Sminthea eurygaster, and Colobonema seri- 
ceum (Thuesen & Childress, unpublished). 

Etymology. —Named in honor of James 
J. Childress of the Marine Science Institute, 
University of California at Santa Barbara 
who has devoted a considerable part of his 
life to the study of midwater organisms off 
the California coast and is in part respon- 
sible for the discovery of this medusa. 

Distribution. —This animal has been re- 
covered in trawls from San Clemente Basin, 
off Baja California, Mexico where the bot- 
tom depth can be greater than 2000 m to 
northwest of Point Conception, California, 
U.S.A. where the bottom depth is over 4000 
m. The shallowest discrete depth tow in 
which it has been taken was 600 m and it 
has been taken in discrete depth trawls 
reaching to 1475 m. We have not routinely 
fished at depths greater than this and it is 
not known how deep V. childressi occurs. 
Although not abundant (never more than 
three specimens in a trawl), it is routinely 
taken in the above region all four seasons 
of the year. 

Coloration. —The pigmentation of V. 
childressi 1s neither a dark blue nor deep 
burgundy but is truly black. The color does 
not fade in specimens preserved in 10% for- 
malin in filtered seawater after storage in 
the dark for two years. Other bathypelagic 
organisms including fish, crustaceans and 
molluscs are known to have black pigmen- 
tation (Wimpenny 1966, color frontispiece), 
however no other species of hydromedusa 
with black pigmentation is recorded in the 
literature. Kramp (1961) describes the gut 
of the coronate scyphozoan Nausithoé glo- 
bifera as being black, however the guts of 
other coronates, suchas N. rubra, Periphylla 
periphylla, Atolla wyvillei and A. vanhoef- 
feni, are not black but rather a densely-pig- 
mented deep burgundy in living specimens. 


192 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Pigel: 


Side view of Vampyrocrossota childressi, new genus, new species, holotype, collected from 777 m 


depth off Point Conception, California. Inner-bell pigmentation and velum pigmentation is black. Bell is 12 


mm in height. 


Anecdotal accounts reporting large num- 
bers of a black medusa (Semaeostomeae; 
Chrysaora sp.?) in the Los Angeles, Cali- 
fornia, area were prevalent during July—Au- 
gust of 1989, however the organism has yet 
to be described in the scientific literature. 
Other Cnidaria with black pigmentation are 
known. The anemone Metridium senile has 
a black endodermal melanin (Fox & Pantin 
1941), and the siphonophore Erenna ri- 


chardi has black endodermal pigmentation 
which is thought to be acquired by feeding 
on black midwater fishes (Totton 1965). The 
small size of V. childressi suggests that me- 
sopelagic fishes are not the source ofits black 
pigmentation. Some of the burgundy-col- 
ored deep-sea medusae, including Crossota 
rufobrunnea, contain porphyrin pigments 
(Herring 1972, Bonnett et al. 1979). The 
exact nature of the pigment in V. childressi 


VOLUME 106, NUMBER 1 


ae Qe ee 
OR mec ony) Moy 
ET I Ly 


193 


(i 


" 


Fig. 2. Side view of Vampyrocrossota childressi, new genus, new species, drawn with inner-bell pigmentation 
“removed” to reveal the gonads and manubrium. The manubrium can reach past the velum when extended in 


living individuals. 


is not known, although spectrophotometric 
analysis of pigment extracted in ethanol re- 
veals an absorption peak at 479 nm indi- 
cating that it has blue-light absorbing com- 
ponent (Thuesen, unpublished data). 


Acknowledgments 


Iam grateful to J. J. Childress, K. L. Smith 
and the captains, crews and scientists aboard 


the Research Vessels Point Sur and New 
Horizon for their assistance at sea. Research 
cruises were supported through National 
Science Foundation grants OCE 85-00237 
to J. J. Childress and OCE 89-22620 to K. 
L. Smith. I thank C. E. Mills for informative 
discussion of mesopelagic medusae and for 
comments on the manuscript. The figures 
were drawn by P. Schalk of Stichting ter 
Bevordering van de Nederlandse Oceano- 


194 


grafie. I was supported in part by a Califor- 
nia Sea Grant Fellowship through NOAA, 
National Sea Grant College Program, De- 
partment of Commerce, under grant num- 
ber NA89AA-D-SG138, project number 
USDC EG-10-8B, through the California 
Sea Grant College, and in part by the Cal- 
ifornia State Resources Agency. The U.S. 
Government is authorized to reproduce and 
distribute for governmental purposes. 


Literature Cited 


Alvarino, A. 1967. Bathymetric distribution of Chae- 
tognatha, Siphonophorae, Medusae, and Cte- 
nophorae off San Diego, California.— Pacific 
Science 21:474—485. 

Bigelow, H. B. 1913. Medusae and Siphonophorae 
collected by the U.S. Fisheries steamer “‘Alba- 
tross”’ in the northwestern Pacific, 1906.—Pro- 
ceedings of the United States National Museum 
44:1-119, 6 pls. 

Bonnett, R., E. J. Head, & P. J. Herring. 1979. Por- 
phyrin pigments of some deep-sea medusae. — 
Journal of the Marine Biological Association of 
the United Kingdom 59:565-573. 

Childress, J. J., A. T. Barnes, L. B. Quetin, & B. H. 
Robison. 1978. Thermally protecting cod ends 
for recovery of living deep-sea animals. — Deep- 
Sea Research 25:419-422. 

Fox, D. L., & C. F. A. Pantin. 1941. The colours of 
the plumose anemone Metridium senile (L.).— 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Philosophical Transactions of the Royal Soci- 
ety, Series B 230:415-—450. 

Herring, P. J. 1972. Porphyrin pigmentation in deep- 
sea medusae.— Nature 238:276-277. 

Kramp, P. L. 1947. Medusz. Part III. Trachylina 

and scyphozoa.— Danish Ingolf-Expedition Re- 

ports 5:1-66, 6 pls. 

. 1961. Synopsis of the medusae of the world. — 

Journal of the Marine Biological Association of 

the United Kingdom 40:1—-469. 

1968. The hydromedusae of the Pacific and 
Indian Oceans. —““Dana’’—Reports 72:1—200. 
Larson, R.J.,& G.R. Harbison. 1990. Medusae from 

McMurdo Sound, Ross Sea including the de- 
scriptions of two new species, Leuckartiara 
brownei and Benthocodon hyalinus. —Polar Bi- 
ology 11:19-25. 

Segura-Puertas, L. 1984. Morphology and zoogeog- 
raphy of medusae (Cnidaria: Hydrozoa and Scy- 
phozoa) from the eastern tropical Pacific. —In- 
stituto de Ciencias del Mar y Limnologia 
Universidad Nacional Autonoma de Mexico 
Publicaciones Especiales 8:1—320. 

Totton, A.K. 1965. A synopsis of the Siphonophora. 
British Museum (Natural History), London 230 
pp., 40 pls. 

Wimpenny, R. S. 1966. The plankton of the sea. 
American Elsevier, New York, 426 pp. 


Marine Science Institute, University of 
California, Santa Barbara, California 93106, 
WtStAe 


PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 195-203 


A NEW SPECIES OF SIBOPATHES 
(CNIDARIA: ANTHOZOA: ANTIPATHARIA: ANTIPATHIDAE) 
FROM THE GULF OF MEXICO 


Dennis M. Opresko 


Abstract. —A new species of antipatharian (Cnidaria: Anthozoa: Antipathar- 
ia), Sibopathes macrospina, is described from the Gulf of Mexico. The species 
differs from the only other species in the genus, S. gephura van Pesch, by having 
longer spines and anterior pinnules positioned slightly above, rather than below, 
the adjacent lateral pinnules. The taxonomic relationships of Sibopathes with 
Cladopathes, Hexapathes, Taxipathes, Bathypathes, Schizopathes, and Par- 


antipathes are discussed. 


An unusual antipatharian coral was re- 
cently collected in the northern Gulf of 
Mexico during submersible investigations 
conducted by Dauphin Island Sea Lab un- 
der the direction of T. Hopkins. This coral 
proved to be related to a species in the genus 
Sibopathes previously known only from the 
Indo-Pacific. Comparisons with the type 
material of the Pacific species have revealed 
that the Gulf specimen represents a new 
species. The holotype has been deposited in 
the National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 
(USNM). 


Family Antipathidae 
Subfamily Cladopathinae 
Genus Sibopathes van Pesch, 1914 


Type species.—Sibopathes gephura van 
Pesch, 1914:203-205, pl. VI, figs. 3, 5-6, 
15; pl. VII, fig..3. (Type locality: Indo-Pa- 
cific, east of Timor, Indonesia, 8°17.4’S, 
127°30.7’E, 1224 m, Siboga Stn. 280). 

Diagnosis. — Polyps transversely elongat- 
ed with six reduced primary mesenteries, 
no secondary mesenteries, and no actino- 
pharynx. 


Sibopathes macrospina, new species 
Figs. 1, 2, 3, 4A-—C, E 


Holotype.—USNM 91417. Gulf of Mex- 
ico, off Alabama, 29°09'30’N, 88°01'10’W, 


UNCW 9119, Johnson Sea Link Stn. JSL 
3097, 26 Aug 1991, 489-559 m, Coll. W. 
W. Schroeder. 

Diagnosis. —Corallum branched and pin- 
nulate (Fig. 1); pinnules simple, 1—2 cm long, 
arranged in four longitudinal rows and 
grouped in alternating pairs with each pair 
consisting of one lateral and one anterolat- 
eral pinnule (Fig. 2A, B). Spines simple, tri- 
angular, smooth, 0.07-0.12 mm in height 
(Fig. 3); arranged in longitudinal rows; from 
four to six spines per millimeter in each row. 
Polyps elongated, about 2 mm in transverse 
diameter (Fig. 4A); in a single row with four 
polyps per centimeter. 

Description. — Holotype about 36 cm high 
and 16 cm wide (Fig. 1). Basal holdfast ab- 
sent; lowermost part of the stem about 2 
mm in diameter. Corallum branched irreg- 
ularly to the fourth order with branches 0.5— 
2.0 cm apart. Branches and branchlets 
straight or curved, and directed upward 
(branch angle 60—75°). Overall branching of 
corallum planar with branchlets arising pri- 
marily from sides of lower order branches, 
occasionally from front (corresponding to 
polyp-bearing side of stem) and rarely from 
back. 

Stem and branches pinnulated; branch- 
lets developing from elongated pinnules 
which become pinnulated in turn. Pinnules 
simple (without subpinnules), relatively 
straight and stiff; not strictly uniform in size, 


196 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Th ym/ ‘ 


” 


a  / ji! 


I 
8) 


Fig. 1. Sibopathes macrospina. Holotype, USNM 91417; height about 36 cm. 


number, or arrangement on branchlets but 
usually in four longitudinal rows with two 
nearly opposite lateral or posterolateral rows 
and two anterolateral rows (Fig. 2A). An- 
terolaterals occasionally located more lat- 


erally and laterals more posterolaterally. 
Anterolaterals absent near base of some 
branchlets. Pinnules 1.8-—3.5 mm apart 
(mean 2.75 mm, n = 20) with four or five 
per centimeter in each longitudinal row. 


VOLUME 106, NUMBER 1 


197 


Pap. 2. 


A-B. Sibopathes macrospina, holotype, USNM 91417. A, Cross sectional view of branchlet showing 


arrangement of pinnules around axis; scale equals 0.2 mm. B, Frontal (polypar) view of branchlet showing 


arrangement of pinnules; scale equals 0.4 mm. 


Pinnules in lateral rows arranged alternate- 
ly; anterolateral pinnules placed 0.40.7 mm 
above (distal to) adjacent lateral pinnule. 
Overall, pinnules form alternating pairs, 
each consisting of one lateral and one slight- 
ly higher anterolateral member (Fig. 2B). 
Lateral pinnules 0.9-1.9 cm long (mean 1.36 
cm, 7 = 14) and 0.20—0.32 mm in diameter 
(near base). Axial canal of pinnules 0.06— 
0.08 mm in diameter. Pinnules project up- 


ward slightly forming angle of 75° or more 
with branchlet. Anterolateral pinnules usu- 
ally shorter, but occasionally as long as or 
longer than adjacent lateral pinnules. Ad- 
jacent pinnules from different branches 
anastomose near base of corallum. 

Spines usually simple, but occasionally 
bifid; triangular, acute, and smooth-sur- 
faced (Fig. 3); small and relatively unde- 
veloped at tip of pinnules (Fig. 4B) but up 


198 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. 
0.1 mm. 


to 0.12 mm high (from midpoint of base to 
apex) on lower portion of pinnules (Fig. 4C). 
Spines unequally developed around circum- 
ference of pinnules; largest spines usually 
on front or polyp side of axis but, in places 
nearly equal in size on three sides, leaving 
abpolypar side with smallest spines. Poly- 
par spines usually extending out perpendic- 
ular to pinnular axis, but occasionally di- 
rected distally or proximally. Abpolypar 
spines 0.02-0.05 mm high. Spines on stem 
and larger branches rarely more than 0.04 
mm high. Spines on pinnules arranged in 
longitudinal rows; three to five rows visible 
in lateral view (excluding rows in which 
spines only partially visible). Distance be- 
tween adjacent spines in each row 0.20-0.36 
mm; generally from four to six spines per 
millimeter in each row. 

Polyps elongated along transverse axis; 


Sibopathes macrospina, holotype, USNM 91417. Stereo SEM of pinnule with axial spines; scale equals 


distance from distal edge of distal lateral 
tentacles to proximal edge of proximal lat- 
eral tentacles t.8—2.3 mm (Fig. 4A). Polyps 
arranged in a single row with about four 
polyps per centimeter; interpolypar space 
0.5—0.8 mm. Peristomal folds absent. Oral 
cone usually elongated transversely 0.28— 
0.44 mm; sagittal diameter 0.14—0.28 mm. 
Tentacles 0.12—0.25 mm in length. Ova 
present in lateral sections of coelenteron (Fig. 
4E). Polyps with only six rudimentary and 
incomplete mesenteries, no actinopharynx, 
and no evidence of mesogloeal partitions 
separating coelenteron into central and lat- 
eral chambers. 

Etymology.—The specific name is de- 
rived from the Latin “‘macros” and “‘spina”’ 
in reference to the relatively large spines on 
the pinnules. 

Comparisons. —This species is very sim- 


VOLUME 106, NUMBER 1 


“I 


Fig. 4. A-C. Sibopathes macrospina, holotype, USNM 91417. A, Stereo SEM of polyp. B, Distal end of 
pinnule. C, Center of pinnule. D. S. gephura van Pesch, holotype, Siboga Stn. 280, center of pinnule. E. S. 
macrospina, holotype, USNM 91417, polyp with ova. Scale in A and E equals 0.5 mm; scale in B (also for C 
and D) equals 0.1 mm. 


200 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Morphometrics for Sibopathes gephura and S. macrospina. 


Character 


Corallum 
Height (cm) 
Basal stem diameter (mm) 
Highest order of branching 
Rows of pinnules 


Pinnules 


Length of lateral pinnules (cm) 
Basal diameter (mm) 
Diameter of central canal (mm) 
Distance apart in one row (mm) 
Angle between lateral and 
anterior pinnules 

Distal angle with stem 

Spines 
Height, polypar (mm) 
Height, abpolypar (mm) 
Distance apart in one row (mm) 
Average number per millimeter 
Number of rows on pinnules (one view) 


Polyps 
Length (mm) 
Interpolypar space (mm) 
Number per centimeter 
Length of tentacles (mm) 
Oral cone, transverse diameter (mm) 
Oral cone, sagittal diameter (mm) 


S. gephura iS. macrospina 
10 36 
~0.85 —) 
Ie 4 
Ar_5 24° 
0.7-2.2 0.9-1.9 
0.22-0.28 0.20-0.32 
0.10-0.16 0.06-0.08 
2.2-3.0 1.8-3.5 
45-90° 30-45° 
~60° = 
0.03-0.04 0.07-0.12 
<0.03-0.04 0.02-0.05 
0.12-0.22 0.20-0.36 
5-7 4-6 
3-4 3-5 
2.0-2.5 1.8-2.3 
0.5-1.1°¢ 0.50.8 
3-3.5 3.5-4 
<0.5 0.12-0.25 
0.6-0.87° 0.28-0.44 
0.2 0.14-0.28 


a Specimen may only be a branch from a larger colony. 


b Most common condition. 
© Maximum value reported by van Pesch (1914). 


ilar to Sibopathes gephura van Pesch (1914). 
Both species have a branched pinnulated 
corallum with simple pinnules arranged for 
the most part in four longitudinal rows. The 
major differences between the two species 
involve the orientation and arrangement of 
the pinnules and the size of the spines (Table 
1). Pinnules on S. gephura appear somewhat 
less regular in length than those on S. mac- 
rospina, although in both cases they reach 
about the same maximum size of about 2 
cm before becoming pinnulated branchlets. 
In S. gephura the pinnules tend to be more 
curved and more distally directed (relative 
to the branch), whereas in S. macrospina 
they are straight, stiff, and extend out more 
horizontally. The central axial canal in the 


pinnules of S. macrospina is smaller in di- 
ameter than that in S. gephura (Table 1). 
Van Pesch (1914) described the arrange- 
ment of the pinnules in S. gephura as being 
in four longitudinal rows, two lateral, one 
anterior, and one posterior; however, in re- 
examining the type specimen it was found 
that in places the rows are also arranged 
biserially, that is, with two rows on the right 
and two on the left. In such cases the pin- 
nules are placed in alternating pairs with the 
front or anterolateral pinnule of each pair 
located slightly below the adjacent lateral 
(as viewed from the front or polyp-side of 
the corallum). This arrangement differs from 
that found in S. macrospina where the an- 
terolaterals are placed slightly above the ad- 


VOLUME 106, NUMBER 1 


jacent laterals. Furthermore, on the holo- 
type of S. gephura a few additional pinnules 
were found representing a fifth longitudinal 
row. These occurred in front of and slightly 
below the adjacent anterolateral pinnules 
such that each group of three pinnules 
formed a descending series going from the 
side to the front of the branch (as viewed 
from the polyp-side of the corallum). In 
contrast, in S. macrospina the pattern 1s re- 
versed, with the pinnules in each group 
forming an ascending series (Fig. 2B). 

In both species the spines are simple, tri- 
angular and acute; however, in S. gephura 
they measure not more than 0.04 mm in 
height (Fig. 4D), whereas in S. macrospina 
they attain a maximum size of 0.12 mm. In 
addition, the spines in S. macrospina are 
spaced farther apart than those in S. ge- 
phura (Table. 1). In both species the spines 
are largest on the side of the axis bearing 
the polyps; however, this is not easily seen 
in S. gephura because the abpolypar spines 
are only slightly smaller than the polypar 
spines. 

There are no major differences in the ex- 
ternal morphology of the polyps of the two 
species except that the polyps in S. macro- 
spina are slightly smaller and closer together 
and have shorter tentacles than those in S. 
gephura (Table 1). These differences are not 
very great and may be due, in part, to state 
of preservation. In both species the polyps 
are arranged in a single row; on the upper 
side of the pinnules in S. gephura, and on 
the upper and occasionally front and lower 
sides of the pinnules in S. macrospina. In 
neither species is there any sign of peristo- 
mal folds dividing the polyps into central 
and lateral sections. 

Although the specimen of S. macrospina 
was not originally fixed for histological ex- 
amination, several polyps were removed 
from the corallum, sectioned, stained, and 
examined microscopically. Internal features 
of the polyp were difficult to distinguish be- 
cause of the poor condition of the tissue; 
however, as in the case of S. gephura, there 


201 


was no sign of an actinopharynx, and the 
mesenteries were rudimentary and ap- 
peared to be no more than six in number. 
In addition, there was no indication of me- 
sogoeal septa separating the coelenteron into 
central and lateral chambers. Van Pesch re- 
ported the same condition in S. gephura. 
Although nematocysts could not be seen in 
the polyps of S. macrospina, they were re- 
ported as occurring in batteries on the ten- 
tacles of S. gephura polyps (van Pesch 1914). 


Discussion 


The two species of Sibopathes differ from 
all other antipatharians in that the polyps 
lack an actinopharynx and, consequently, 
the six mesenteries are incomplete. Van 
Pesch (1914) created the subfamily Homoe- 
otaeniales to contain Sibopathes and a sec- 
ond genus Cladopathes Brook, whose pol- 
yps also have only six mesenteries; however, 
in Cladopathes the mesenteries are attached 
to a well-developed actinopharynx. Clado- 
pathes was originally included with Schi- 
zopathes, Bathypathes, and Taxipathes, in 
the subfamily Schizopathinae by Brook 
(1889). Polyps in the latter three genera have 
ten mesenteries, six primary and four sec- 
ondary. In establishing the Schizopathinae, 
Brook considered the number of mesenter- 
ies to be of secondary importance when 
compared to the transverse elongation of 
the polyps into what he interpreted as “‘di- 
morphic” structures consisting of “‘gastro- 
zooids” and “‘gonozooids.” In describing the 
polyps of Schizopathes, Brook reported that 
the “‘zooids”’ were isolated from one anoth- 
er externally by peristomal folds and inter- 
nally by mesogloeal partitions extending 
down from the upper, interior surface of the 
coelenteron. These features were not spe- 
cifically described for other genera in the 
subfamily, and Brook’s own illustrations in- 
dicate that peristomal folds are not typical 
of Bathypathes or Taxipathes polyps. Con- 
sequently, the only remaining diagnostic 
character of the Schizopathinae is the ex- 


202 


treme transverse elongation of the polyp. In 
1896, Schultze proposed a new classifica- 
tion of the Antipathidae based on the num- 
ber of mesenteries, rather than on the pre- 
sumed dimorphism of the polyps. He 
therefore established a separate subfamily, 
the Hexamerota for Cladopathes. The sub- 
family was renamed the Cladopathinae by 
Kinoshita in 1910, who added to it a new 
monotypic genus Hexapathes. The polyps 
of H. heterosticha reportedly contain an ac- 
tinopharynx, six complete primary mesen- 
teries and no secondaries; however, the 
skeletal morphology is quite different from 
that of Cladopathes (and Sibopathes); in- 
stead, it is similar to that of species of Bath- 
ypathes. Thus, van Pesch’s Homoeotaeni- 
ales is equivalent to the Cladopathinae and 
contains those genera having polyps with 
six mesenteries, but in which an actino- 
pharynx may be present (Cladopathes and 
Hexapathes) or absent (Sibopathes). 

In describing Sibopathes gephura, van 
Pesch (1914) noted that many of the ana- 
tomical features of the polyps were sugges- 
tive of a very primitive condition. These 
features included: absence of an actino- 
pharynx and secondary mesenteries; small 
size of the sagittal mesenteries and their oc- 
currence only near the base of sagittal ten- 
tacles; thinness of the mesogloea; presence 
of epidermally derived cells within the me- 
sogloea; very wide connection between the 
axis epithelium and the body wall; occur- 
rence of deeply staining gland cells through- 
out the gastrodermis; and absence of mes- 
enteric filaments (defined by van Pesch as 
extensions of the mesenteries at the aboral 
edge of the actinopharynx). Furthermore, 
van Pesch noted that the transverse mes- 
enteries disappeared at the junction of the 
central and lateral parts of the polyp, but 
reappeared, with a club-shaped edge, in the 
lateral chambers. Van Pesch did not con- 
sider the club-shaped edge of the transverse 
mesenteries to represent a mesenteric fila- 
ment. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


If Schultze’s classification is followed, it 
could be argued that the absence of an ac- 
tinopharynx and the presence of other prim- 
itive characters would warrant the taxo- 
nomic recognition of Sibopathes at least at 
the same level as the Cladopathinae, and 
perhaps even at the family level (with the 
recent removal of the Dendrobrachiidae, the 
order currently contains only the single fam- 
ily Antipathidae, see Opresko & Bayer 
1991). However, if one considers the pos- 
sibility that the characters of Sibopathes are 
secondarily derived as a result of the ex- 
treme transverse elongation of the polyp (a 
character which in itself would be difficult 
to view aS a primitive state), and if one eval- 
uates overall similarities in external mor- 
phology, then it might be argued that S7- 
bopathes is indeed related to Cladopathes 
but not to Hexapathes. Like Sibopathes, 
Cladopathes is quasi-sympodial (i.e., with- 
out a single continuous stem), multi- 
branched, and pinnulated. The pinnules are 
simple or forked and arranged in three or 
four longitudinal rows “‘showing a subspiral 
arrangement.” In contrast, Hexapathes het- 
erosticha Kinoshita is clearly monopodial 
with two rows of lateral pinnules and one 
row of anterior pinnules, a pattern identical 
to that occurring in Bathypathes lyra Brook. 

Sibopathes and Cladopathes also show 
external similarities to the genera Taxi- 
pathes and Parantipathes. All four genera 
(Sibopathes, Taxipathes, Parantipathes, and 
Cladopathes) have transversely elongated 
polyps of relatively small size (generally <3 
mm). In contrast, the polyps of Hexapathes 
heterosticha were described as being 5-9 mm 
long, and this is also the case for some spe- 
cies of Bathypathes. Taxipathes and Par- 
antipathes also have pinnules arranged in 
longitudinal rows and alternating semispiral 
groupings, but they differ from Sibopathes 
and Cladopathes in having secondary mes- 
enteries in the polyps. However, this differ- 
ence is certainly not as significant as that 
separating Sibopathes from Cladopathes 


VOLUME 106, NUMBER 1 


(i.e., the absence of an actinopharynx). Fur- 
ther analysis may eventually show that these 
four genera form a natural assemblage. 


Acknowledgments 


The author wishes to thank S. Cairns for 
his helpful suggestions and for taking the 
scanning electron micrographs; T. Hopkins 
and W. W. Schroeder of the University of 
Alabama, Marine Environmental Sciences 
Consortium, Dauphin Island Sea Lab for 
providing the specimen of S. macrospina 
which was collected during field studies con- 
ducted under NSF EPSCOR Grant No. R11- 
8996152 and NOAA/NURC/UNCW Grant 
No. NA88AA-D-URO04; R. W. M. van 
Soest of the Riksmuseum van Natuurlijke 
Historie in Amsterdam for the loan of the 
type specimen of S. gephura; T. Bayer, C. 
Bast, and M. Bogle for reviewing the manu- 
script; S. Braden of the Smithsonian Insti- 
tution for preparing the samples for the 
scanning electron microscope (SEM); and J. 
Wesley of Oak Ridge National Laboratory 


203 


for preparing the histological sections. This 
work was supported in part by the Smith- 
sonian Institution and by Oak Ridge Na- 
tional Laboratory, Oak Ridge, Tennessee. 


Literature Cited 


Brook, G. 1889. Report on the Antipatharia.—Re- 
ports of the Scientific Results of the Voyage of 
the Challenger.— Zoology 32:5—222. 

Kinoshita, K. 1910. On a new antipatharian Hexa- 
pathes heterosticha n. gen. and n. sp.—Anno- 
tationes Zoologicae Japonenses 7:231—234. 

Opresko, D. M., & F. M. Bayer. 1991. Rediscovery 
of the enigmatic coelenterate Dendrobrachia, 
(Octocorallia: Gorgonacea) with descriptions of 
two new species.—Transactions of the Royal 
Society of South Australia 115:1-19. 

Schultze, L.S. 1896. Beitrag zur Systematik der An- 
tipatharien.—Abhandlungen der Senckenber- 
gischen naturforschenden Gesellschaft 23:1—40. 

Van Pesch, A.J. 1914. The Antipatharia of the Siboga 
Expedition. — Siboga-Expeditie Monographe 17: 
1-258. 


Health and Safety Research Division, Oak 
Ridge National Laboratory, P.O. Box 2008, 
Oak Ridge, Tennessee 37831-6050, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 204-205 


INTERNATIONAL COMMISSION ON ZOOLOGICAL 
NOMENCLATURE 


% The Natural History Museum 
Cromwell Road 

London, SW7 5BD, U.K. 

Tel. 071-938 9387 


Applications published in the Bulletin of Zoological Nomenclature 


The following Applications were published on 30 September 1992 in Vol. 49, 
Part 3 of the Bulletin of Zoological Nomenclature. Comment or advice on these 
Applications is invited for publication in the Bulletin, and should be sent to the 
Executive Secretary, I.C.Z.N., % The Natural History Museum, Cromwell Road, 
London SW7 SBD, U.K. 


Case No. 


2806 Zanclea costata Gegenbaur, 1856 (Cnidaria, Hydrozoa): proposed conser- 
vation of both generic and specific names. 

2827 Gebia major capensis Krauss, 1843 (currently Upogebia capensis; Crustacea, 
Decapoda): proposed replacement of neotype, so conserving the 
usage of capensis and also that of G. africana Ortmann, 1894 (cur- 
rently Upogebia africana). 

Podisus Herrich-Schaeffer, 1851 (Insecta, Heteroptera): proposed conserva- 
tion of P. vittipennis Herrich-Schaeffer, 1851 as the type species. 

ANTHRIBIDAE Billberg, 1820 (Insecta, Coleoptera): proposed precedence over 
CHORAGIDAE Kirby, 1819. 

Catocala connubialis Guenée, 1852 (Insecta, Lepidoptera): proposed con- 
servation of the specific name. 

METOPIINAE Foerster, 1868 (Insecta, Hymenoptera), METOPIINI Raffray, 1904 
(Insecta, Coleoptera), and METOPIINI Townsend, 1908 (Insecta, Dip- 
tera): proposed removal of homonymy. 

Acamptopoeum Cockerell, 1905 (Insecta, Hymenoptera): proposed desig- 
nation of Camptopoeum submetallicum Spinola, 1851 as the type 
species. 

Cynolebias opalescens Myers, 1942 and Cynolebias spendens Myers, 1942 
(Osteichthyes, Cyprinodontiformes): proposed conservation of the 
specific names. 

Filimanus Myers, 1936 (Osteichthyes, Perciformes): proposed designation 
of Filimanus perplexa Feltes, 1991 as the type species. 

Rana megapoda Taylor, 1942 (Amphibia, Anura): proposed conservation of 
the specific name. 

Megophrys montana Kuhl & van Hasselt, 1822 (Amphibia, Anura): proposed 
placement of both the generic and specific names on Official Lists, 
and Leptobrachium parvum Boulenger, 1893 (currently Megophrys 
parva): propposed conservation of the specific name. 

Anisolepis grilli Boulenger, 1891 (Reptilia, Squamata): proposed conservation 
of the specific name. 


VOLUME 106, NUMBER 1 


Opinions published in the Bulletin of Zoological Nomenclature 


The following Opinions were published on 30 September 1992 in Vol. 49, Part 


3 of the Bulletin of Zoological Nomenclature. 


Opinion No. 

1689. Epizoanthus Gray, 1867 (Cnidaria, Anthozoa): conserved. 

1690. Helix (Helicigona) barbata Férussac, 1832 (currently Lindholmiola barbata; 
Mollusca, Gastropoda): lectotype designation confirmed. 

1691. Polygyra Say, 1818 (Mollusca, Gastropoda): Polygyra septemvolva Say, 1818 
designated as the type species, and POLYGYRIDAE Pilsbry, 1895 given 
precedence over MESODONTIDAE Tryon, 1866. 

1692. Phyllodoce Lamarck, 1818 and Polyodontes de Blainville, 1828 (Annelida, 
Polychaeta): conserved. 

1693. Coccinella undecimnotata Schneider, [1792] (currently Hippodamia (Sem- 
ladalia) undecimnotata; Insecta, Coleoptera): specific name con- 
served. 

1694. Rhinapion Beguin-Billecocqg, 1905 (Insecta, Coleoptera): conserved. 


£695. 


1696. 


Acanthophthalmus van Hasselt in Temminck, 1824 (Osteichthyes, Cyprin- 
iformes): not conserved. 
HYDROBATIDAE Mathews, 1912 (1865) (Aves, Procellariiformes): conserved. 


PROC. BIOL. SOC. WASH. 
106(1), 1993, p. 206 


REVIEWERS 


The following people reviewed manuscripts for the Proceedings in 1992. P. Alderslade, F. Alvarez, 
A. Alvarino, W. D. Anderson, Jr., M. Aoki, A. Asakura, K. Baba, I. Ball, S. Bandoni, A. M. Bauer, 
F. M. Bayer, J. A. Blake, T. E. Bowman, R. Brinkhurst, J. Burch, S. D. Cairns, E. Campos, M. D. 
Carleton, R. Carney, K. Carpenter, J. Chess, K. Coates, C. O. Coleman, B. B. Collette, M. R. 
Cooper, J. S. Costlow, Jr., N. Cumberlidge, H.-E. Dahms, D. Davis, M. Dillon, M. Dojiri, R.-C. 
Dowler, L. H. Emmons, K. Fauchald, D. L. Felder, R. M. Feldmann, G. Fenton, K. Fitzhugh, J. 
F. Fitzpatrick, O. S. Flint, Jr., J. Friend, R. J. Gagné, A. L. Gardner, S. Gelder, A. C. Gill, C. J. 
Glasby, W. Goldberg, I. Goodbody, J. Goy, M. Grasshoff, G. R. Graves, D. E. Hahn, R. Hanley, 
E. Harada, W. Hartman, P. C. Heemstra, G. Hendler, D. Hendrickson, B. Hilbig, S. Hiruta, J.-s. 
Ho, H. H. Hobbs, Jr., D. Holdich, J. Holsinger, R. W. Holzenthal, T. S. Hopkins, G. D. Johnson, 
N. K. Johnson, J. K. Jones, Z. Kabata, Y. Kikuchi, G. L. Kirkland, Jr.,G. Klassen, L. S. Kornicker, 
R. Kropp, J. Kudenov, S. Lanyon, R. Larson, D. Laubitz, R. Lemaitre, W. G. Lyons, E. Macpherson, 
L. Madin, C. Magalhaes, R. B. Manning, J. W. Martin, W. N. Mathis, J. Mauchline, D. McKinnon, 
G. A. de Melo, H. Michel, M. Milligan, C. Mills, W. L. Minckley, R. F. Modlin, R. D. Mooi, J. 
C. Morse, A. Muir, M. Murano, T. Newberry, P. K. L. Ng, J. L. Norenburg, R. D. Owen, L. R. 
Parenti, K. Parkes, C. Patterson, J. L. Patton, D. L. Pawson, T. Perkins, M. H. Pettibone, F. Pleyel, 
G. Pohle, W. F. Presch, G. Pretzmann, J. Pruski, A. Rea, J. W. Reid, C. B. Robbins, S. de A. 
Rodrigues, G. Rodriguez, C. F. E. Roper, F. Rowe, K. Ruetzler, C. W. Sabrosky, J. Sieg, T. Simpson, 
R. Sluys, A. E. Smalley, J. Smith III, G. S. Steyskal, J. H. Stock, F. C. Thompson, F. G. Thompson, 
M. Thurston, R. Toll, S. Tyler, J. L. Villalobos, R. Vonk, N. Voss, E. Walker, V. Wallach, L. A. 
Ward, M. J. Wetzel, M. K. Wicksten, A. B. Williams, J. T. Williams, K. Wittmann, K. Wouters, 
H. Yeatman, R. Young, W. Zeider, R. Zottoli, G. R. Zug. 


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CONTENTS 


On the identity of Echimys didelphoides Desmarest, 1817 (Mammalia: Rodentia: Echimyidae) 
Louise H. Emmons 

A new subspecies of pocket gopher (Geomys) from Texas (Mammalia: Rodentia: Geomyidae) 
Michael J. Smolen, Richard M. Pitts, and John W. Bickham 


— Identification of bird subfossils from cave surface deposits at Anjohibe, Madagascar, with a 


description of a new giant Coua (Cuculidae: Couinae) 
Steven M. Goodman and Florent Ravoavy 
Two new species of blind snake, genus 7yphlops (Reptilia: Squamata: Typhlopidae), from the 


Philippine Archipelago Addison H. Wynn and Alan E. Leviton 
A new species of dimorphic tree frog, genus Hyla (Amphibia: Anura: Hylidae), from the Vaupés 
River of Colombia William F. Pyburn 


Description of the advertisement call and resolution of the systematic status of Leptodactylus 
gracilis delattini Miiller, 1968 (Amphibia: Leptodactylidae) 

Linnette Garcia Pérez and W. Ronald Heyer 

Prochilodus britskii, a new species of prochilodontid fish (Ostariophysi: Characiformes), from 

the rio Apiaca, rio Tapaj6s system, Mato Grosso, Brazil Ricardo M. C. Castro 

A new Devonian ophiuroid (Echinodermata: Oegophiurida) from New York state and its 

bearing on the origin of ophiuroid upper arm plates Frederick H. C. Hotchkiss 


Erythrosquilloidea, a new superfamily, and Tetrasquillidae, a new family of stomatopod crus- . 


taceans Raymond B. Manning and David K. Camp 
Systematics and taxonomic remarks on Pinnotheres muliniarum Rathbun, 1918 (Crustacea: 
Brachyura: Pinnotheridae) Ernesto Campos 
Anomoeomunida, a new genus proposed for Phylladiorhynchus caribensis Mayo, 1972 (Crus- 
tacea: Decapoda: Galatheidae) Keiji Baba 
, Two new species of Neocallichirus from the Caribbean Sea (Crustacea: Decapoda: Callianas- 
sidae) Raymond B. Manning 
Caprella arimotoi, a new species (Crustacea: puppets Caprellidea) from the Seto Inland 
Sea, Japan Ichiro Takeuchi 
A new species of Kalliapseudes (Crustacea: Tanaidacea: Kalliapseudidae) from Trinidad 
Roger N. Bamber 
Enterocola africanus, a new species (Copepoda: Ascidicolidae) associated with a compound 
ascidian Synoicum species from North Africa (Strait of Gibraltar) 
Pablo J. Lopez-Gonzalez, Mercedes Conradi, and J. Carlos Garcia-Gomez 
New species and new records of the genus Elaphoidella (Crustacea: Copepoda: Harpacticoida) 
from the United States Janet W. Reid and Teruo Ishida 
New genera and species of deep-sea polychaetes of the family Nautiliniellidae from the Gulf 
of Mexico and the eastern Pacific James A. Blake 
Taxonomy of European species of Amphiduros and Gyptis (Polychaeta: Hesionidae) 
Fredrik Pleiyel 
A new genus of hydrobiid snails (Mollusca: Gastropoda: Prosobranchia: Rissooidea) from 
northern South America Robert Hershler and France Velkovrh 
Vampyrocrossota childressi, a new genus and species of black medusa from the Bathypelagic 
zone off California (Cnidaria: Trachymedusae: Rhopalonematidae) Erik V. Thuesen 
A new species of Sibopathes (Cnidaria: Anthozoa: Antipatharia: Antipathidae) from the Gulf 
of Mexico Dennis M. Opresko 
International Commission on Zoological Nomenclature: Applications and Opinions 
Reviewers— 1992 , 


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ys * 


THE BIOLOGICAL SOCIETY OF WASHINGTON 


1992-1993 
Officers 
President: Storrs L. Olson Secretary: G. David Johnson 
President-elect: Janet W. Reid Treasurer: T. Chad Walter 
Elected Council 

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Custodian of Publications: Austin B. Williams 


PROCEEDINGS 


Editor: C. Brian Robbins 


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PROC. BIOL. SOC. WASH. 
106(2), 1993, pp. 207-220 
REVISED CLASSIFICATION AND PHYLOGENETIC 
HYPOTHESIS FOR THE 
ACANTHOSTOMINAE LOOSS, 1899 
(DIGENEA: OPISTHORCHIFORMES: CRY PTOGON 


Daniel R. Brooks and Barbara Holcman 


Abstract. —Specimens of an acanthostome digenean origin identified as.» 
Acanthostomum scyphocephalum and later transferred to TimonieHa-are-de- 
scribed and named as a distinct species of 7imoniella. Acanthostomum Scy-. 
phocephalum sensu strictu is included in an updated phylogenetic analysis of 
the acanthostome digeneans. The new analysis differs from an earlier one by 
Brooks (1980) by allowing reversals (Wagner criterion vs. Camin-Sokal crite- 
rion), producing a more parsimonious representation of character data; no 
transformation series needed re-polarization. Acanthostomum scyphocephalum 
is a member of the clade containing all other species of Acanthostomum oc- 
curring in North, Central and South America. Acanthostomum is paraphyletic 
if Atrophecaecum is excluded from it; accordingly, the two genera are synon- 
ymized. No other changes from the hypothesis of Brooks (1980) resulted. The 
resulting annotated phylogenetic classification, with synapomorphic diagnoses, 
includes Acanthostomum as the sister-group of Caimanicola, Proctocaecum as 
their sister-group, the monotypic Gymnatrema as their sister-group, and 77- 
moniella as the basal sister-group. Four new subgenera are proposed, one in 


Timoniella, one in Proctocaecum, and two in Acanthostomum. 


The acanthostome digeneans (Opisthor- 
chiformes: Cryptogonimidae: Acanthosto- 
minae) inhabit a variety of piscivorous pol- 
kilotherm amniote vertebrates throughout 
the tropical and subtropical regions of the 
world. Brooks (1980) provided the first phy- 
logenetic systematic analysis of the acan- 
thostomes. He recognized six genera, 77- 
moniella Rebecq, 1960, Proctocaecum 
Baugh, 1957, Gymnatrema Morozov, 1955, 
Caimanicola Teixeira de Freitas & Lent, 
1938, Acanthostomum Looss, 1899 and 
Atrophecaecum Bhalerao, 1940, although 
Acanthostomum had no synapomorphy to 
support its recognition as a monophyletic 
group. 

Since that time, publications have de- 
scribed two additional species and dis- 
cussed their phylogenetic relationships 
(Brooks & Caira 1982, Blair et al. 1988), 


and have added data about host and geo- 
graphic distributions, as well as valuable 
taxonomic information (Ostrowski de Nu- 
nez 1984a, 1984b, 1986, 1987). This study 
provides a description of a new species, and 
incorporates the new data provided by Os- 
trowski de Nunez into the phylogenetic data 
base for the acanthostomes, resulting in an 
updated phylogenetic hypothesis and an an- 
notated classification with cladistic diag- 
noses for all taxa. 


Methods 


In addition to the material specified in 
Brooks (1980), Brooks & Caira (1982) and 
Blair et al. (1988), we have examined the 
following material loaned from Dr. Mar- 
garita Ostrowski de Nunez: Acanthosto- 
mum gnerili (19 specimens from Rhamdia 


208 


sapo, Laguna Chis-Chil, Prov. Buenos Ai- 
res, Argentina); Acanthostomum megace- 
tabulum (1 specimen from Drymarchon 
corais melanurus, Villahermosa, Mexico); 
Caimanicola marajoarum (14 specimens 
from Crocodylus intermedius, Caracas, 
Venezuela; 3 specimens from Paleosuchus 
sp., Antioquia, Colombia); Caimanicola 
brauni (13 specimens from Phrynops hilarii, 
Buenos Aires, Argentina); Timoniella loossi 
(3 specimens from Crocodylus intermedius, 
Caracas, Venezuela); Acanthostomum sp. V1 
of Ostrowksi de Nunez (1984b) (2 speci- 
mens from Caiman fuscus, Rio Chagras, 
Panama). All measurements are in wm un- 
less otherwise noted. TBL = total body 
length. Figures were drawn with the aid of 
a drawing tube. 


Results and Discussion 


Timoniella ostrowskiae, new species 
Figs. 1-3 


Synonyms. —Acanthostomum scypho- 
cephalum of Mane-Garzon & Gil, 1961; Ti- 
moniella scyphocephala of Brooks, 1980. 

Description. —(based on holotype and 2 
paratypes) Body 2.98—3.15 mm long by 420— 
540 wide at midbody. Tegument covered 
with spines of uniform size. Oral sucker ter- 
minal, bell-shaped, 410-517 long by 443- 
517 wide, armed with single row of 23 spines 
68-78 long by 20-25 wide. Pharynx 197— 
205 long by 187-230 wide. Prepharynx 377— 
426 long. Ratio of oral sucker width to pha- 
ryngeal width 1:0.42-0.45. Ceca opening 
separately at posterior end of body. Ace- 
tabulum 1.22—1.47 mm from anterior end; 
forebody 41-47% TBL. Acetabulum 140- 
156 long by 131-147 wide; ratio of oral 
sucker width to acetabulum width 1:0.27— 
0.32.Testes tandem, intercaecal, near pos- 
terior end of body; posttesticular space 5.2— 
5.5% TBL. Anterior testis 187-205 long by 
123-139 wide, posterior testis 180-221 long 
by 115-123 wide. Male genitalia consisting 
of coiled external seminal vesicle lying pos- 
terodorsal to acetabulum and musculo- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


glandular pars prostatica and ejaculatory 
duct extending anteriorly dorsal to acetab- 
ulum, opening into genital atrium. Gonotyl 
lacking. Genital pore ventral, medial, im- 
mediately preacetabular. Ovary 90-139 an- 
terior to anterior margin of anterior testis, 
123-139 long by 115-123 wide. Seminal 
receptacle posterodorsal to ovary, between 
Ovary and anterior testis. Mehlis gland pres- 
ent, Laurer’s canal short. Uterine loops ex- 
tending posteriorly to lateral margin of an- 
terior testis, anteriorly to posterior margin 
of seminal vesicle, occupying 38-43% TBL; 
terminal portion of uterus opening into gen- 
ital atrium. Vitellaria follicular, in two lon- 
gitudinal extracecal fields extending from 
level of posterior margin of seminal vesicle 
to slightly posterior to anterior margin of 
anterior testis. Eggs 25-27 long by 10-12 
wide. 

Type host.—Phrynops hilarii (Dumeril 
and Bibron). 

Type locality.—Rio Negro, Paso de los 
Toros, Departamento de Tacuarembo, 
Uruguay. 

Holotype. —URFC Helm. Coll. No. 
11038. Paratypes: URFC Helm. Coll. No. 
11039-11040. 

Etymology.—The species is named in 
honor of Dr. Margarita Ostrowski de Nu- 
Nez, who first recognized its distinct iden- 
tity. 

Brooks (1980) was unable to locate the 
holotype of A. scyphocephalum Braun, 1899 
which had been collected in “‘Testudo ma- 
tamata’”’ from southern Brazil. Based on ex- 
amination of specimens collected in Phry- 
nops hilarii from Uruguay (reported by 
Mane-Garzon & Gil 1961), Brooks (1980) 
transferred the species to Timoniella be- 
cause it possessed preovarian rather than 
postovarian seminal receptacles, a unique 
and unreversed synapomorphy that diag- 
noses Timoniella among the acantho- 
stomes. Ostrowski de Nunez (1986) found 
the holotype of Acanthostomum scypho- 
cephalum Braun, 1899, and showed that it 
was a member of Acanthostomum. The 


VOLUME 106, NUMBER 2 209 


500 


Figs. 1-3. Timoniella (Maillardiella) ostrowskiae. 1. Ventral view of holotype. 2. Ootype region. 3. Terminal 
genitalia. 


210 


Table 1.—Homoplasious changes for 17 characters 
of acanthostomes. Character consistency index = num- 
ber of apomorphic states for each character divided by 
total number of changes postulated on the phylogenetic 
trees. For character identities, see Appendix 1. 


Character 
consistency 
index 


Char- 
acter 
state 


Figure #-Character 
number on figure 


1-0 9-36 50% (1/2) 
2201 49-38 50% (1/2) 
6-0 5-3 50% (1/2) 
TOien Ashes 10, 50% (1/2) 
9-0 8-10 
9-1 4-10, 5-1, 6-3, 7-5, 8-7, 9-32 33% (3/9) 
10-0 6-1 
10-1 5-2, 7-4, 8-2, 8-17, 9-29, 9-35 17% (1/7) 
Giles ages 0 50% (1/2) 
12-1 5-5, 6-11, 8-14 33% (1/3) 
13-1 5-6, 6-6, 8-13, 9-22, 9-33 33% (2/6) 
[422 5-13 6°2 8-1 
14-3 5-14, 9-19 
14-5 4-12, 6-4, 8-6, 9-26 57% (8/14) 
15-1 5-8, 8-18, 9-24 50% (2/4) 
1620" 753 
16-1 4-20, 5-9, 6-8, 9-20, 9-28, 9-34 17% (1/7) 
171 52108 8:3 67% (2/3) 
18-0 8-30 
18-1 5-11, 7-1, 8-9 40% (2/5) 
DD ATeDLGA TA 50% (2/4) 
2311 (62596-1138 9225 33% (1/3) 
29-1 5-15, 6-12, 8-15 50% (2/4) 


specimens reported by Mane-Garzon & Gil 
(1961) therefore appear to represent a pre- 
viously unnamed species of Timoniella, 
which we formally described and named 
above. Timoniella ostrowskiae is most 
closely related to T. incognita Brooks, 1980, 
T. loossi (Perez Vigueras, 1957) Brooks, 
1980, 7. absita Blair et al., 1988, and T. 
unami (Pelaez & Cruz, 1957) Brooks, 1980, 
all of which lack gonotyls. Of those species, 
T. ostrowskiae is the only one exhibiting 
ceca opening separately at the posterior end 
of the body; T. incognita has blindly-ending 
ceca, and the remaining three species have 
ceca Opening into the excretory vesicle near 
the posterior end of the body. 

As indicated in the introduction, new in- 
formation concerning acanthostomes has 
accumulated since Brooks (1980) produced 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


the first phylogenetic systematic hypothesis 
for the group. Brooks (1980) used the Ca- 
min-Sokal (also known as the Weighted In- 
variant Step Strategy [WISS]— Wiley et al. 
1991) criterion, allowing no evolutionary 
reversals, in producing a phylogenetic tree 
for the acanthostomes. He found no syn- 
apomorphy to support the monophyly of 
Acanthostomum, nor any character that 
could link Acanthostomum with either Cai- 
manicola or Atrophecaecum, two genera 
having synapomorphic support that were 
shown forming an unresolved trichotomy 
with Acanthostomum. In this study we re- 
analyzed the data using the less restrictive 
Wagner criterion (Wiley et al. 1991) after 
checking character polarizations using es- 
timates of higher-level digenean phyloge- 
netic relationships (Brooks et al. 1985, 
Brooks et al. 1989, Brooks & McLennan 
1993) not available to Brooks (1980). We 
found support for the original character po- 
larizations of Brooks (1980, see also Brooks 
& Caira 1982, Blair et al. 1988) but, as 
would be expected, we also found more par- 
simonious optimizations for some homo- 
plasious characters when reversals were al- 
lowed (for a summary of transformation 
series, see Appendix 1; for a summary of 
homoplasious changes, including reversals, 
see Table 1). Based on the information pre- 
sented by Ostrowski de Nunez (1986), we 
included Acanthostomum scyphocephalum 
in the phylogenetic framework; it appears 
to be a member of the clade containing all 
the other species of Acanthostomum occur- 
ring in North, Central and South America 
(Fig. 8). Most importantly, the new analysis 
showed that Acanthostomum is paraphylet- 
ic (Fig. 8), and should be combined with 
Atrophecaecum (Figs. 8, 9). 

Systematic theorists have begun investi- 
gating the problems of providing robust 
means for assessing the results of phyloge- 
netic analyses (e.g., Archie 1989, Farris 
1989, Sanderson & Donoghue 1989, Klas- 
sen et al. 1991, Meier et al. 1991). These 
studies have produced some interesting, and 


VOLUME 106, NUMBER 2 


in some cases initially counter-intuitive, 
findings. For example, the minimum sig- 
nificant value for the most commonly used 
indicator, the consistency index (CI— Wiley 
et al. 1991), drops as one adds taxa and 
characters to a study; for example, a study 
using 50 characters for 20 taxa and reporting 
a CI of 65% may actually be more robust 
than using 10 characters for 7 taxa and re- 
porting a CI of 80%. This happens because 
there are often apomorphic character 
changes occurring once within a given taxon 
that also occur once in another taxon. If the 
scope of a study were expanded to include 
both taxa, the estimate of homoplasy would 
increase (and the CI would drop) even if the 
hypothesized phylogenetic relationships of 
the (now) subgroups did not change. Or, to 
use current terminology, we would say that 
a global phylogenetic analysis had discov- 
ered homoplasy that the two /ocal analyses 
failed to recognize. In some cases such glob- 
al homoplasy could affect the hypotheses of 
relationships, so recognizing global homo- 
plasy may play an important role in deter- 
mining robust character polarities during 
outgroup comparisons at the inception of a 
phylogenetic analysis (e.g., Maddison et al. 
1984, Wiley et al. 1991). 

The acanthostomes provide an excellent 
illustration of the relationship between local 
and global parsimony considerations in 
phylogenetic analysis. If we treat all the 
acanthostomes as a single taxon, the con- 
sistency index for the characters reported 
herein is 50.5% (49 apomorphic character 
states and 97 character changes). This value 
is low for digeneans in general, the consis- 
tency index based on global parsimony con- 
siderations for all digeneans being approx- 
imately 72% (Brooks & McLennan 1993). 
By contrast, if we treat each of the clades 
denoted as a genus separately, the estimated 
homoplasy is generally much lower (100% 
for the characters used at the generic level 
by themselves, including the monotypic 
Gymnatrema, for the species of TJimoniella, 
and for the species of Caimanicola, and 


211 


84.6% for the species of Proctocaecum), the 
exception being Acanthostomum, for which 
the consistency index based on local par- 
simony considerations is 51%. This means 
that although there is much homoplasy 
among the acanthostomes as a whole, most 
of it is dispersed among clades rather than 
concentrated within clades. Thus, deter- 
mination of plesiomorphic states by out- 
group comparisons is not problematical, and 
there is presently a single most parsimoni- 
ous tree, for the group. 

Mensural, or continuous variable, char- 
acters are problematical for phylogenetic re- 
construction. The determination of discrete 
character states is often problematical for 
such traits; options seem to range from con- 
sidering almost every species autapomorph- 
ic to recognizing very few states and there 
is NO consensus approach among phyloge- 
neticists. Brooks (1980) adopted a conser- 
vative approach to recognizing character 
states based on his examination of available 
specimens. Taking a conservative approach 
to such traits often results in considerable 
homoplasy. In the present study, the ho- 
moplasy is distributed among half (17 of 34) 
of the transformation series (1, 2, 6, 7, 9- 
18, 22—23, and 29 in Appendix 1), 9 of which 
(10, 12-13, 15-18, 22-23) are mensural in 
nature. Of the 48 homoplasious character 
transformations, 28 (58.3%) stem from the 
mensural characters and 20 (41.7%) from 
the qualitative traits; moreover, the men- 
sural traits that show homoplasy have a 
combined character consistency index of 
28.6% (14/42), while the qualitative traits 
that show homoplasy have a combined 
character consistency index of 48.7% (19/ 
39) (Table 1). While suggestive, these data 
are actually moot with respect to the ques- 
tion of whether or not such characters pro- 
vide adequate phylogenetic information, 
because they exhibit high levels of homo- 
plasy but do not support relationships that 
are contradicted by non-mensural charac- 
ters. A strong test of these characters re- 
quires a search for intrinsically qualitative 


212 


characters whose apomorphic states sup- 
port phylogenetic relationships that conflict 
with the ones supported by the present data 
base. 


Conclusions 


The changes discussed above are reflected 
in the phylogenetic trees, cladistic diagno- 
ses, and classification below (the monotypic 
Gymnatrema has no separate cladogram). 
In the following, italicized numbers refer to 
synapomorphies listed by number on Fig. 
4: other numbers refer to synapomorphies 
listed by number on Figs. 5-8. Four new 
subgenera are proposed. Each is named for 
a digenean systematist who has advanced 
our understanding of acanthostome rela- 
tionships: Dr. Claude Maillard, University 
of Montpellier, France; Dr. David Gibson, 
British Museum (Natural History), London, 
England; Dr. David Blair, James Cook Uni- 
versity, Townsville, Australia; and Dr. 
Robin Overstreet, Gulf Coast Marine Re- 
search Laboratory, Ocean Springs, Missis- 
sippi, USA. 


Subfamily Acanthostominae Poche, 1926 
(Fig. 4) 


Diagnosis. —Cryptogonimidae with ter- 
minal oral sucker (/); armed with single row 
of spines (2); preacetabular pit (3); genital 
pore not in preacetabular pit (4); seminal 
vesicle coiled posteriorly (5); suckerlike 
gonotyl present (6). 


Genus Timoniella Rebecq, 1960 
(Fig. 5) 


Diagnosis. —Acanthostominae preovari- 
an seminal receptacle (7). 


Subgenus 7imoniella Rebecq, 1960 


Diagnosis. —Timoniella having vitelline 
follicles not extending anteriorly to poste- 
rior margin of seminal vesicle* (1); length 
of body occupied by uterine loops more than 
50% TBL* (2); seminal vesicle not coiled 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


posteriorly* (3); prepharynx shorter than 
pharynx* (4); ratio of oral sucker width to 
pharyngeal width 1:0.25—0.40* (5); ratio of 
body length to width averaging 7.5—15:1* 
(6). 


T. praeterita (Looss, 1901) Maillard, 1974 


Diagnosis. —Cyclocoel (7); forebody 10- 
20% TBL* (8); maximum body length 7-16 
mm* (9). 


T. imbutiformis (Molin, 1859) Brooks, 1980 


Diagnosis. — Ratio of oral sucker : acetab- 
ular width 1:0.8—1.3* (10); oral spines av- 
eraging 25-30 in number™* (11). 


Subgenus Maillardiella, new subgenus 


Diagnosis. —Timoniella lacking gono- 
tyls* (12). 


T. incognita Brooks, 1980 


Diagnosis. — With characters of the sub- 
genus. 

Remarks: This species was originally re- 
ported by Nasir (1974) as Acanthostomum 
scyphocephalum inhabiting Caiman croco- 
dilus crocodilus. According to Ostrowski de 
Nunez (1984b), the acanthostomes reported 
as Acanthostomum scyphocephalum from 
Drymarchon corais by Nasir (1974) may be 
a still undescribed species. 


T. ostrowskiae Brooks & Holcman, 1993 


Diagnosis. —Ceca opening separately at 
posterior end of body* (13). 


T. unami (Pelaez & Cruz, 1957) Brooks, 
1980 


Diagnosis. —Ceca opening into excretory 
vesicle* (14); vitelline follicles extending 
posteriorly to middle of posterior testis* (15). 


T. loossi (Perez Vigueras, 1957) Brooks, 
1980 


Diagnosis. —Ceca opening into excretory 
vesicle* (14); vitelline follicles extending 


VOLUME 106, NUMBER 2 


posteriorly to middle of posterior testis* (15); 
vitelline follicles confluent dorsally (16). 


T. absita Blair et al., 1988 


Diagnosis. —Ceca opening into excretory 
vesicle* (14); vitelline follicles extending 
posteriorly to middle of posterior testis* (15); 
constriction in seminal vesicle (17). 


Genus Gymnatrema Morozov, 1955 


Diagnosis. —Acanthostominae having 
some uterine loops lateral to testes but none 
posttesticular (8); vitelline follicles not ex- 
tending anteriorly to posterior margin of 
seminal vesicle* (/0); vitelline follicles con- 
fluent posttesticularly (J/); one cecum at- 
rophied* (12); one cecum opening laterally 
and one cecum ending blindly (/3). 


G. gymnarchi (Dollfus, 1950) Morozov, 
1955 


Diagnosis. — With characters of the genus. 


Genus Proctocaecum Baugh, 1957 
(Fig. 6) 


Diagnosis. —Acanthostominae having 
some uterine loops lateral to testes but none 
posttesticular (8); ceca opening separately 
and laterally at even levels (9); excretory 
vesicle Y-shaped with short stem and con- 
striction of arms in middle (/4); eggs av- 
eraging more than 30 um long (/5); gonotyl 
large, solid-muscular (/6). 


Subgenus Proctocaecum Baugh, 1957 


Diagnosis.—Proctocaecum having rela- 
tive length of uterine loops less than 45% 
TBE? (1). 


P. gonotyl (Dollfus, 1950) Brooks, 1980 


Diagnosis. — With characters of the sub- 
genus. 


P. vicinum (Odhner, 1902) Brooks, 1980 


Diagnosis. —Ceca opening separately and 
laterally at uneven levels (2). 


2d 


P. coronarium (Cobbold, 
1980 


1861) Brooks, 


Diagnosis. —Ceca opening separately and 
laterally at uneven levels (2); vitelline fol- 
licles not extending anteriorly to posterior 
margin of seminal vesicle* (3); one cecum 
atrophied* (4); maximum oral spine length 
more than 100 wm%* (5). 


Subgenus Overstreetium, new subgenus 


Diagnosis.—Proctocaecum having ratio 
of body length to width averaging 7.5-15: 
FG): 


P. productum (Odhner, 1902) Brooks, 1980 


Diagnosis. — With characters of the sub- 
genus. 


P. elongatum (Tubangui & Masilungen, 
1936) Brooks, 1980 


Diagnosis. —Ceca opening separately at 
posterior end of body* (7); maximum body 
length 7-16 mm* (8); ratio of body length 
to width averaging more than 20:1 (9). 


P. crocodili (Yamaguti, 1954) Brooks, 1980 


Diagnosis. —Ceca opening separately at 
posterior end of body* (7); maximum body 
length 7-16 mm* (8); forebody less than 
10% TBL (10). 


P. atae (Tubangui & Masilungen, 1936) 
Brooks, 1980 


Diagnosis. —Ceca opening separately at 
posterior end of body* (7); maximum body 
length 7-16 mm* (8); ratio of oral sucker 
width to pharyngeal width 1:0.25—-0.40* (11). 


P. nicolli Brooks, 1980 


Diagnosis. —Ceca opening separately at 
posterior end of body* (7); maximum body 
length 7-16 mm* (8); ratio of oral sucker 
width to pharyngeal width 1:0.25—0.40* (11); 
vitelline follicles extending anteriorly to 
posterior margin of acetabulum* (12); max- 
imum oral spine length more than 100 uwm* 


(13). 


214 


Genus Caimanicola 
Teixeira de Freitas & Lent, 1938 
(Fig. 7) 


Diagnosis. —Acanthostominae having 
some uterine loops lateral to testes but none 
posttesticular (8); ceca opening separately 
and laterally at even levels (9); excretory 
vesicle Y-shaped with short stem and con- 
striction of arms in middle (/4); eggs av- 
eraging more than 30 um long (/5); gonotyl 
lacking* (/7); esophagus longer than phar- 
ynx (J8); tegumental spines unusually ro- 
bust in mid-forebody (/9); maximum body 
length 7-16 mm* (20). 


C. pavidus (Brooks & Overstreet, 1977) 
Brooks, 1980 


Diagnosis. —Oral spines averaging 25-30 
in number™ (1). 


C. caballeroi (Pelaez & Cruz, 1953) Brooks, 
1980 


Diagnosis. —Eggs averaging less than 30 
um long (2); maximum body length 2-6 mm 


(3). 


C. marajoarus Teixeira de Freitas & Lent, 
1938 


Diagnosis. —Eggs averaging less than 30 
um long (2); length of body occupied by 
uterine loops more than 50% TBL* (4); vi- 
telline follicles not extending anteriorly to 
posterior margin of seminal vesicle* (5). 


C. brauni (Mafie-Garzon & Gil, 
Brooks, 1980 


1961) 


Diagnosis. —Eggs averaging less than 30 
um long (2); length of body occupied by 
uterine loops more than 50% TBL* (4); ratio 
of oral sucker width to acetabular width 
1:0.3-0.7 (6). 

Remarks: According to Ostrowksi de Nu- 
Nez (1984b), Acanthostomum brauni of 
Caballero (1955) is an undetermined species 
of acanthostome, possibly undescribed. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Genus Acanthostomum Looss, 1899 
(Figs. 8, 9) 


Diagnosis. —Acanthostominae having 
some uterine loops lateral to testes but none 
posttesticular (8); ceca opening separately 
and laterally at even levels (9); eggs aver- 
aging more than 30 wm long (/5); gonotyl 
lacking* (/7); excretory vesicle with long 
stem and short arms (2/). 


Subgenus Blairium, new subgenus 


Diagnosis. —Acanthostomum having ceca 
opening separately at posterior end of body* 


(1). 


A. scyphocephalum (Braun, 1899) Hughes 
et al., 1941 


Diagnosis. — With characters of the sub- 
genus. 

Remarks: Ostrowski de Nunez (1986) re- 
described this species from the type mate- 
rial, which had been missing and presumed 
lost (see Brooks 1980). 


A. americanum (Perez Vigueras, 1957) Her- 
ber, 1961 


Diagnosis. —Length of body occupied by 
uterine loops more than 50% TBL* (2). 


A. megacetabulum Thatcher, 1963 


Diagnosis. — Length of body occupied by 
uterine loops more than 50% TBL* (2); ratio 
of oral sucker : acetabular width 1:0.8—1.3* 


(3). 
A. gnerti Szidat, 1954 


Diagnosis.—Testes oblique (4); vitelline 
follicles sparse (5). 


A. minimum Stunkard, 1938 


Diagnosis. —Testes oblique (4); vitelline 
follicles sparse (5); one cecum atrophied* 
(6). 

A. astorquii Watson, 1976 


Diagnosis.— Testes oblique (4); vitelline 


VOLUME 106, NUMBER 2 


Timoniella Gymnatrema Proctocaecum Caimanicola Acanthostomum 


Fig. 4. Phylogenetic relationships among the gen- 
era of the subfamily Acanthostominae. Numbers refer 
to apomorphic traits listed in diagnoses in text. Each 
asterisk (*) indicates the presence of a homoplasious 
character; the particular homoplasious characters are 
denoted by an asterisk in the diagnoses in the text. 


follicles sparse (5); one cecum atrophied* 


(6). 


Subgenus Gibsonium, new subgenus 


Diagnosis.—Acanthostomum having vi- 
telline follicles not extending anteriorly to 
posterior margin of seminal vesicle* (7); oral 
spines averaging less than 20 in number* 


(8). 
A. absconditum (Looss, 1901) Poche, 1926 


Diagnosis. —With characters of the sub- 
genus. 


[ -- Timoniella -- | [ ------------------------ Maillardiella ----------------------- ] 


praeterita imbutiforme incognita ostrowskiae unami loossi absita 


7 (figure 4) 


Fig.5. Phylogenetic relationships among species of 
Timoniella. Numbers refer to apomorphic traits listed 
in diagnoses in text. Each asterisk (*) indicates the pres- 
ence of a homoplasious character; the particular homo- 
plasious characters are denoted by an asterisk in the 
diagnoses in the text. 


[ ------ Proctocaecum ------ ] [ ------------------ Overstreetium ------------------ ] 


coronarium yicinum gonotyl productum elongatum crocodili alae 


16 ( figure 4) 


Fig. 6. Phylogenetic relationships among species of 
Proctocaecum. Numbers refer to apomorphic traits list- 
ed in diagnoses in text. Each asterisk (*) indicates the 
presence of a homoplasious character; the particular 
homoplasious characters are denoted by an asterisk in 
the diagnoses in the text. 


Subgenus Acanthostomum Looss, 1899 


Diagnosis. —Acanthostomum having oral 
spines averaging 25-30 in number* (9); vi- 
telline follicles extending anteriorly to pos- 
terior margin of seminal vesicle* (10); eggs 
averaging less than 30 um long (11); cecal 
bifurcation approximately 10% TBL preac- 
etabular (12). 


A. knobus Issa, 1962 


Diagnosis. — Ratio of body length to width 
averaging 7.5—15:1* (13). 


A. spiniceps (Looss, 1896) Looss, 1899 


Diagnosis. — Ratio of oral sucker width to 
pharyngeal width 1:0.25—0.40* (14). 


C. pavida C. caballeroi C.marajoara_ C. brauni 


18 - 20 ( figure 4) 


Fig. 7. Phylogenetic relationships among species of 
Caimanicola. Numbers refer to apomorphic traits list- 
ed in diagnoses in text. Each asterisk (*) indicates the 
presence of a homoplasious character; the particular 
homoplasious characters are denoted by an asterisk in 
the diagnoses in the text. 


216 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


astorquil 


minimum gnerii megacetabulum americanum sc 


] [ Gibsonium ] [ - Acanthostomum - ] 


absconditum niloticum spiniceps knobus 


Fig. 8. 


21 ( figure 4) 


Phylogenetic relationships among species of Acanthostomum excluding Acanthostomum (Atrophe- 


caecum). Numbers refer to apomorphic traits listed in diagnoses in text. Each asterisk (*) indicates the presence 
of a homoplasious character; the particular homoplasious characters are denoted by an asterisk in the diagnoses 


in the text. 


A. niloticum Issa, 1962 


Diagnosis. — Ratio of oral sucker width to 
pharyngeal width 1:0.25—0.40* (14); vitel- 
line follicles extending anteriorly to poste- 
rior margin of acetabulum* (15). 


Subgenus Atrophecaecum Bhalerao, 1940 


Diagnosis. —Acanthostomum having oral 
spines averaging 25-30 in number™* (9); vi- 
telline follicles extending anteriorly to pos- 


terior margin of seminal vesicle* (10); vi- 
telline follicles terminating preovarially (16); 
length of body occupied by uterine loops 
more than 50% TBL* (17); forebody 10- 
20% TBL* (18). 


A. indicum Sinha, 1942 


Diagnosis. —With characters of the sub- 
genus. 


A. slusarskii Kalyankar, 1977 


[ ---------------------------------------------- Atrophecaecum ---------------------------------------------- ] 


proctophorum 


indicum — slusarskit pakistanensis asymmetricum 

19 - 22 - 
20 ¥* 25. #* 
ae 


28 * 29 


27 


31 ¥* 


simhai burminis lobacetabulare cerberi marinw 
3 35 * 7 d 
ee 3 39 40 
38 * 
36 * 
32 * 
26 * 
21* 


16-18 ** 


Fig. 9. Phylogenetic relationships among species of Acanthostomum (Atrophecaecum). Numbers refer to 
apomorphic traits listed in diagnoses in text. Each asterisk (*) indicates the presence of a homoplasious character; 
the particular homoplasious characters are denoted by an asterisk in the diagnoses in the text. 


VOLUME 106, NUMBER 2 


Diagnosis. —Ceca opening into excretory 
vesicle* (19); maximum body length 7-16 
mm* (20). 


A. pakistanense Coil & Kuntz, 1960 


Diagnosis. —Prepharynx shorter than 
pharynx* (21); forebody 10-20% TBL* (22); 
ratio of body length to width averaging 7.5— 
15:1* (23); oral spines averaging 20—24 in 
number* (24); maximum oral spine length 
more than 100 uwm* (25). 


A. asymmetricum (Simha, 1958) Khalil, 
1963 


Diagnosis. —Prepharynx shorter than 
pharynx* (21); one cecum lost (27); maxi- 
mum body length 7-16 mm* (28). 


A. proctophorum (Dwivedi, 1966) Yama- 
guti, 1971 


Diagnosis. —Prepharynx shorter than 
pharynx* (21); one cecum lost (27); length 
of body occupied by uterine loops more than 
50% TBL* (29); oral spines averaging 20- 
24 in number™* (30); vitelline follicles con- 
fluent preovarially (31). 


A. simhai Khalil, 1963 


Diagnosis. —Prepharynx shorter than 
pharynx* (21); one cecum atrophied* (26); 
vitelline follicles not extending anteriorly to 
posterior margin of seminal vesicle* (32); 
ratio of body length to width averaging 7.5— 
15:1* (33); maximum body length 7-16 
mm* (34). 


A. burminis (Bhalerao, 1926) Bhalerao, 1936 


Diagnosis. —Prepharynx shorter than 
pharynx* (21); one cecum atrophied* (26); 
vitelline follicles not extending anteriorly to 
posterior margin of seminal vesicle* (32); 
length of body occupied by uterine loops 
more than 50% TBL* (35). 


A. lobacetabulare Brooks & Caira, 1982 


Diagnosis. —Prepharynx shorter than 
pharynx* (21); one cecum atrophied* (26); 
vitelline follicles not extending anteriorly to 


217 


posterior margin of seminal vesicle* (32); 
subterminal mouth* (36); lobate acetabu- 
lum (37). 


A. cerberi (Fischthal & Kuntz, 1965) Brooks, 
& Caira 1982 


Diagnosis. —Prepharynx shorter than 
pharynx* (21); one cecum atrophied* (26); 
vitelline follicles not extending anteriorly to 
posterior margin of seminal vesicle* (32); 
subterminal mouth* (36); no oral spines* 
(38); no esophagus (39). 


A. marinum (Coil & Kuntz, 1960) Brooks 
& Caira, 1982 


Diagnosis. —Prepharynx shorter than 
pharynx* (21); one cecum atrophied* (26); 
vitelline follicles not extending anteriorly to 
posterior margin of seminal vesicle* (32); 
subterminal mouth* (36); no oral spines* 
(38); no prepharynx (40); secondary group 
of vitelline follicles surrounding testes (41). 


Acknowledgments 


We gratefully acknowledge the loan of 
specimens from Doctora Margarita Ostrow- 
ski de Nunez, Buenos Aires, Argentina. This 
study was supported by funds from oper- 
ating grant A7696 from the Natural Sci- 
ences and Engineering Research Council of 
Canada (NSERC) to DRB. 


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220 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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Appendix 1 


Transformation series for characters used to for- 
mulate phylogenetic hypotheses for acanthostome di- 
geneans (for outgroup argumentation, see Brooks 1980, 
Brooks & Caira 1982, Brooks et al. 1985, Brooks et al. 
1989). 0 = plesiomorphic; | or higher indicates apo- 
morphic states. U = nonlinear transformation series, 
run unordered in computer-assisted analyses (numbers 
assigned to each apomorphic state are arbitrary). 


1. Oral sucker subterminal (0); terminal (1). 
2. Oral sucker lacking spines (0); armed with single 
row of spines (1). 
3. Preacetabular pit lacking (0); present (1). 
Ventrogenital pit present (0); lacking (1). 
5. Genital pore in preacetabular pit (0); not in pre- 
acetabular pit (1). 
6. Seminal vesicle not coiled posteriorly (0); coiled 
posteriorly (1). 
7. Gonotyl lacking (0); suckerlike (1); large, solid- 
muscular (2). U 
8. Seminal receptacle postovarian (0); preovarian (1). 
9. Vitelline follicles extending anteriorly to posterior 
margin of seminal vesicle (0); not extending an- 
teriorly to posterior margin of seminal vesicle (1); 
confluent dorsally (2); confluent posttesticularly 
GyU 
10. Space occupied by uterine loops less than 45% 
TBL (0); more than 50% TBL (1). 
11. Prepharynx longer than pharynx (0); shorter than 
pharynx (1). 
12. Ratio of oral sucker width to pharyngeal width 
averaging 1:0.5 (0); 1:0.25—0.40 (1). 
13. Ratio of body length to width averaging less than 
7.5210); 75-1521 (): more than 20::0y. U 
14. Ceca ending blindly near posterior end of body (0); 
cyclocoel (1); opening separately at posterior end 
of body (2); opening into excretory vesicle (3); 
opening separately and laterally at even levels (4); 
one cecum atrophied (5); one cecum opening lat- 


sind 


30. 


oF 
Vas 
Bee 
34. 


erally and one cecum ending blindly (6); opening 
separately and laterally at uneven levels (7); one 
cecum lost (8). U 


. Forebody more than 20% TBL (0); 10-20% TBL 


(1); less than 10% TBL (2). 


. Maximum body length less than 7 mm (0); 7-16 


mm (1). 


. Ratio of oral sucker: acetabular width 1:0.6—0.9 


(0); 1:0.8-1.3 (1); 1:0.3-0.7 (2). 


. Oral spines averaging 20-24 in number (0); 25— 


30 (1); less than 20 (2). U 


. Noconstriction in seminal vesicle (0); constriction 


present (1). 


. Posttesticular loops present (0); some uterine loops 


lateral to testes but none posttesticular (1). 


. Excretory vesicle Y-shaped with long stem (0); with 


short stem and constriction of arms in middle (1); 
long stem and short arms (2). 


. Eggs averaging less than 30 wm long (0); more than 


30 um long (1). 


. Maximum oral spine length less than 100 um long 


(0); more than 100 um (1). 


. Esophagus shorter than pharynx (0); longer than 


pharynx (1). 


. Tegumental spines not unusually robust in mid- 


forebody (0); unusually robust in mid-forebody 
(1). 


. Testes tandem (0); oblique (1). 


Vitelline follicles numerous (0); sparse (1). 


. Cecal bifurcation averages 20% TBL preacetabular 


(0); 10% FBL preacetabular (1). 


. Vitelline follicles terminating at least at ovarian 


level and no further posterior than posterior mar- 
gin of the ovary (0); extending posteriorly to mid- 
dle of posterior testis (1); terminating preovarially 
(). 7 

Vitelline follicles not confluent preovarially (0); 
confluent preovarially (1). 

Acetabulum not lobate (0); lobate (1). 

Esophagus present (0); lacking (1). 

Prepharynx present (0); lacking (1). 

Secondary group of vitelline follicles surrounding 
testes lacking (0); present (1). 


PROC. BIOL. SOC. WASH. 
106(2), 1993, pp. 221-224 


A NEW MERICELLA 
(MOLLUSCA: GASTROPODA: CANCELLARIIDAE) 
FROM NORTHEASTERN AFRICA 


Richard E. Petit and M. G. Harasewych 


Abstract.— Mericella bozzettii, new species is described from nine shells trawled 
off Cape Ras Hafun, Somalia in shrimp and lobster nets at a depth of 200-250 
m. This new species may be distinguished from its western Indian Ocean 
congeners by its large size and.coarsely cancellate sculpture. 


Thiele (1929) originally erected Mericel- 


la, as a subgenus of Cancellaria, to contain 


a single Recent species from bathyal depths 
off Tanzania that he had previously attrib- 
uted (Thiele 1925) to Cancellaria (Merica). 
Mericella paschalis (Thiele, 1925), a closely 
related species from the Zanzibar Channel 
that was described in the same pubhcation 
as the type species, has not previously been 
ascribed to this genus. Olsson & Bayer (1972) 
proposed the generic name Gerdiella to ac- 
commodate three newly discovered Recent 
species taken in bathyal depths (516-897 m) 
of the northern Caribbean Sea and the Straits 
of Florida. They recognized that Gerdiella 
was Closely related to, and possibly conge- 


neric with, Mericella, but distinguished these. 


taxa on the basis of size and geographical 
distribution. Subsequent authors have as- 
signed Cancellaria (Merica) corbicula Dall, 
1908, which occurs in bathyal to abyssal 
depths off southern California, to Mericella 
(Abbott 1974:247) or Gerdiella (Kaicher 
1978). Due to the rarity of material, the 
genus Mericella has received little subse- 
quent attention. An additional species of 
Mericella, known from shells of nine spec- 
imens collected by shrimp trawlers off the 
coast of Somalia, is described in this report. 

Abbreviations used in the text: AMNH, 
American Museum of Natural History, New 
York; MNHN, Museum national d’Histoire 
naturelle, Paris; USNM, National Museum 


of Natural History, Smithsonian Institu- 
tion, Washington, D.C. 


Mericella bozzettii, new species 
Figs. 1-3, Table 1 


Diagnosis.—A large species with thick, 
high-spired, coarsely cancellate shell with 
irregularly spaced varices. Aperture exceeds 
half the shell length, with weak denticles 
along flaring, strongly sinuate outer lip. 

Description. —Shell (Fig. 1, Table 1) large 
for genus, reaching 37 mm, heavy, with tall, 
conical spire, rounded anterior. Protoconch 
(Figs. 2-3) conical, of 24 smooth, rounded 
whorls, aligned with coiling axis, increasing 
in diameter from 368 um to 1.87 mm. Tran- 
sition to teleoconch (Figs. 2—3, arrow) 
abrupt, demarcated by weak varix, followed 
immediately by four faint spiral cords, and 
within 4 whorl by axial costae. Teleoconch 
with up to five strongly convex whorls. Su- 
ture impressed. Shoulder indistinct or ab- 
sent. Spiral sculpture of 23-25 strong cords 
on body whorl, 8-9 on penultimate whorl, 
lacking intervening threads. Axial sculpture 
of 25-28 rounded, regularly-spaced, sinu- 
ate, opisthocline, axial costae on body whorl 
(16-18 on first teleoconch whorl) producing 
beaded, cancellate appearance at intersec- 
tions with spiral cords, with four to six fine 
axial lamellae in intervening concave spac- 
es. Varices up to seven in number, initially 


222 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 1-3. Mericella bozzettii, new species. 1, Apertural, lateral and dorsal views of holotype. Scale bar = 1 
cm. 2, Lateral and 3, apical views of the protoconch. Scale bars = 500 um. 


weak and irregularly placed (90—270° apart), 
more prominent and regularly spaced 
(~ 220°) on third and subsequent teleoconch 
whorls. Aperture elongate, elliptical, de- 
flected from coiling axis by 13—-16°. Outer 
lip thickened, forming flaring, strongly sin- 


uate varix, with 12—18 weak denticles lim- 
ited to the base of the varix. Inner lip ad- 
pressed posteriorly, with an angle of 153- 
157° between parietal region and columella. 
Columella with two weak columellar folds 
and siphonal fold, posteriormost fold most 


VOLUME 106, NUMBER 2 


223 


Table 1.—Mericella bozzettii, new species. Measurements of shell characters. Linear measurements in mm (n 


=o. 


Character 


Shell length (SL) 30.6 
Aperture length (AL) 16.9 
AL/SL O53 
No. of whorls, protoconch? 1.92 
No. of whorls, teleoconch 4.72 
No. of varices 7.0 
No. of axial ribs between 

varices 5-6 14.1 
No. of axial ribs between 

varices 6-7 17.4 
No. spiral cords on 

penultimate whorl fies 
No. of spiral cords on 

whorl 5-6 7.0 
No. of teeth on outer lip 13.3 


a Range 
3.9 22.1-37.0 
2.3 12:2—20.3 
0.018 0.512-0.578 
OA 1.75—2.0 
0.27 4.0-5.0 
0.0 7.0 
2.0 11-18 
om! 13-24 
0.6 7-9 
0.8 5-8 
2.0 10-16 


a » = 3 for this character. 


pronounced. Anterior slope of shell round- 
ed, lacking clear distinction between body 
whorl and siphonal canal. Shell color uni- 
formly white. Periostracum, soft parts un- 
known. 

Material examined. —Holotype, USNM 
860315, 28.7 mm; Paratype 1, AMNH 
226453; Paratype 2, MNHN; Paratype 3; 
Petit collection; Paratypes 4-8, Bozzetti col- 
lection; all from the type locality. 

Type locality. —Off Cape Ras Hafun, ap- 
proximately 150 km S of Cape Guardafni, 
Somalia. Trawled in shrimp and lobster nets 
at 200-250 m. 

Etymology.—This species is named in 
honor of Mr. Luigi Bozzetti, who first 
brought it to our attention and kindly pro- 
vided the type material. 

Remarks.— Although this new species 
corresponds more closely in size to species 
of the western Atlantic genus Gerdiella, it 
is placed in Mericella because of its large 
aperture (>'% shell length), smooth proto- 
conch lacking axial sculpture, as well as be- 
cause of its geographic proximity to other 
species of Mericella. It is readily distin- 
guished from Mericella jucunda and M. pa- 
schalis on the basis of its large size, much 
broader axial costae, thick shell and white 


color. Mericella corbicula more closely ap- 
proaches M. bozzettii in size, but differs in 
having a more finely reticulate surface 
sculpture, a lower spire, and a chalky shell 
surface. 

Although Olsson & Bayer (1972:880) 
confirmed the inclusion of the closely re- 
lated genus Gerdiella in Cancellariidae on 
the basis of the morphology of the radula 
of the type species, nothing is known of the 
anatomy or radular morphology of any spe- 
cies of Mericella. Mericella bozzettii occurs 
in somewhat shallower depths (200-250 m) 
than other species of Mericella (404-2012 
m). 


Literature Cited 


Abbott, R. T. 1974. American seashells, second edi- 
tion. Van Nostrand Reinhold, New York, 663 
pp., 24 pls. 

Dall, W. H. 1908. [Reports on the dredging opera- 
tions off the west coast of Central America to 
the Galapagos, to the west coast of Mexico, and 
in the Gulf of California, in charge of Alexander 
Agassiz, carried on by the U.S. Fish Commis- 
sion Steamer “Albatross,” during 1891, Lieut. 
Commander Z. L. Tanner, U.S.N., command- 
ing. XX XVII. Reports on the scientific results 
of the expedition to the eastern tropical Pacific, 
in charge of Alexander Agassiz, by the U.S. Fish 


224 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Commission Steamer “‘Albatross,”’ from Octo- 
ber 1904, to March, 1905, Lieut. Commander 
L. M. Garrett, U.S.N., commanding. XIV] The 
Mollusca and Brachiopoda.—Bulletin of the 
Museum of Comparative Zoology 43(6):205— 
487, pls. 1-22. 

Kaicher, S. D. 1978. Card catalogue of world-wide 
shells. Pack no. 19—Cancellariidae. Privately 
published, St. Petersburg, Florida. Card nos. 
1859-1964. 

Olsson, A. A., & F. M. Bayer. 1972. Gerdiella, a new 
genus of deep-water cancellariids.— Bulletin of 
Marine Science 22:875-880. 


Thiele, J. 1925. Gastropoda der Deutschen Tiefsee- 
Expedition. IJ].—Deutsche Tiefsee-Expedition 
17:35-382 + pls. 13-46. 

. 1929. Handbuch der systematischen Weich- 
tierkunde.— Gustav Fischer, Jenna 1:1-—376. 


Department of Invertebrate Zoology, 
NHB stop 118, National Museum of Nat- 
ural History, Smithsonian Institution, 
Washington, D.C. 20560, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(2), 1993, pp. 225-236 


NEW SPECIES OF ALVINELLIDAE (POLYCHAETA) 
FROM THE NORTH FIJI BACK-ARC BASIN 
HYDROTHERMAL VENTS (SOUTHWESTERN PACIFIC) 


Daniel Desbruyéres and Lucien Laubier 


Abstract. —The polychaete family Alvinellidae Desbruyéres & Laubier, 1986 
comprises two genera, A/vinella and Paralvinella, and ten species or subspecies. 
All species are strictly associated with hydrothermal vents in the Pacific Ocean. 
The genus Paralvinella includes eight species or sub-species, plus one additional 
new species presently being described from North East Pacific hydrothermal 
fields. In 1989, the French research submersible Nautile had 12 successful dives 
in the North Fiji back-arc Basin and explored two active hydrothermal vents. 
Numerous specimens of two additional new species of the genus Paralvinella 
were collected using the manipulator of the submersible. Paralvinella uniden- 
tata, new species, exhibits several features that lead us to erect three different 
subgenera, Paralvinellas. s., Miralvinella, new subgenus and Nautalvinella, new 
subgenus, within the genus Paralvinella. This species lives within the anhydrite 
mass, very close to hot fluid openings. The second new species, P. fijiensis, 
found in the anhydrite mass, but also on basaltic rocks, is closely related to P. 


grasslei, the type species, and to P. palmiformis. 


While the first discovery of hydrothermal 
phenomena occurring at the axes of oceanic 
ridges goes back to 1976 (Lonsdale 1977), 
the exploration of hydrothermal systems in 
back-arc basins is rather recent (Both et al. 
1986, in Manus Basin, Hessler et al. 1988, 
in Marianas back-arc Basin, Fouquet et al. 
1990, 1991, in Lau Basin, Ohta 1990, in 
Okinawa back-arc Basin) (Auzende et al. 
1989, Jollivet et al. 1989). During the early 
summer of 1989, a French-Japanese bio- 
logical cruise, STARMER 2 (30 June 1989 
to 19 July 1989) was devoted to the study 
of biological communities associated with 
deep-sea hydrothermal vents in the South- 
western Pacific, in the North Fiji back-arc 
Basin (Desbruyéeres et al. 1991). The French 
research submersible Nautile, operated from 
R/V Le Nadir, had 12 successful dives in 
these areas. Two different active sites were 
explored. White Lady vent has one large 
diffuser plus several small cylindrical chim- 
neys; the edifice is a few meters high and is 


built up by anhydrite, with hydrothermal 
fluid temperature up to 285°C; the fluid is 
translucent, relatively depleted of metal due 
to subsurface phase separation and shows a 
low content of hydrogen sulfide. The site is 
located at a depth of 2000 m at 16°59’S and 
173°55’E. Another hydrothermal field 
named Mussel Valley consists of Bathy- 
modiolus spp. beds developing on the ba- 
salt; there is no chimney and the hydro- 
thermal fluid diffuses from cracks between 
basaltic rocks; the fluid temperature does 
not exceed 8.5°C. This site is located at a 
depth of 2700 m at 18°49’S and 173°29’E. 
A large collection of polychaetes including 
representatives of the family Alvinellidae 
were collected from the submersible. 

Since our last publication on Alvinellidae 
(Desbruyéres & Laubier 1991), the total 
number of known Paralvinella species in- 
cludes eight species or subspecies (Detinova 
1988). At least one additional new species 
from North East Pacific hydrothermal fields 


226 


is presently being described and additional 
material tentatively identified as Paralvi- 
nella hessleri has recently been collected 
from Okinawa vent fields (Miura & Ohta 
1991). 

During the dives in the North Fiji Basin, 
numerous specimens of two different spe- 
cies of the genus Paralvinella were collected 
using the manipulator of the submersible. 
Morphological study of these animals 
showed that they represent two new species, 
Paralvinella unidentata and P. fijiensis. Par- 
alvinella unidentata exhibits several fea- 
tures that lead us to erect three different sub- 
genera within the genus Paralvinella. This 
species lives in the anhydrite mass, very close 
to hot fluid openings. The second new spe- 
cies, Paralvinella fijiensis, found in the an- 
hydrite mass and on basaltic rocks, is closely 
related to P. grasslei, the type species, and 
P. palmiformis. 


Paralvinella unidentata, new species 
Figs. 1, 3, 4 


Type locality, material examined. —Sev- 
enty-eight specimens collected during Nau- 
tile dives PL 10 (3 specimens, White Lady 
site), PL 11 (5 specimens, White Lady site), 
PL 16 (37 specimens, White Lady site), PL 
20 (32 specimens, White Lady site), PL 21 
(1 specimen, small hydrothermal vent lo- 
cated 150 m from White Lady site in the 
south-west). Extra specimens from dive PL 
10 and PL 20 deep frozen for biochemical 
analyses. Most specimens come from White 
Lady vent site (depth 2000 m, 16°59’50’S 
and 173°55'47’”E). Holotype (dive PL 16, 11 
July 1989, on White Lady site) deposited in 
the collections of the Muséum national 
d@’Histoire naturelle, Laboratoire de Biolo- 
gie des Invertébrés marins et Malacologie 
(n°UC 350). Paratypes from same dive de- 
posited in the collections of the National 
Museum of Natural History, Smithsonian 
Institution, Washington, D.C. (USNM 
157044). 

Etymology. —The specific name refers to 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


the peculiar unidentate uncini. The lack of 
a secondary tooth on the uncini is unique 
within the family Alvinellidae. 

Description. —Holotype 11 mm long and 
1.1 mm wide with 81 setigerous segments. 
Paratypes (14 specimens from PL 16) range 
from 77 to 88 setigerous segments, with the 
majority (11 specimens) having from 79 to 
83. Length of paratypes ranges from 4.8 mm 
to 9.2 mm, with an average of 5.2 mm. 
Color pale grey-pinkish in ethanol, with 
capillary setae and acicular notopodial hooks 
yellow; integuments iridescent. Body grad- 
ually tapering from about setigerous seg- 
ment 50 to the end of the body. A medio- 
ventral row of small shields present. 

Prostomium well developed, with ovi- 
form shield shape, clearly separated from 
buccal segment by deep grooves, with an- 
terior median incision on two thirds of 
length; with small glandular notch visible 
(with SEM) at base of incision. Buccal ap- 
paratus, from dorsum to ventrum, with 
many grooved tentacles in several rows on 
buccal membrane arising from dorsal side 
of buccal cavity; length of tentacles highly 
variable from one individual to the next 
one, possibly due to preservation. Paired 
large ventral tentacles and ventral organ ab- 
sent. 

Buccal segment laterally and ventrally 
visible, well separated from prostomium and 
from segment II. First visible segment (IT) 
achaetous, laterally and ventrally discern- 
ible, clearly separated from peristomium and 
from branchial region. First 25 to 30 setig- 
erous segments with notopodia only. 

Branchial region with four segments, pre- 
ceded by one reduced asetigerous segment, 
visible laterally and ventrally (segment II). 
First branchial segment (segment III) ase- 
tigerous, totally fused with two first setig- 
erous segments. Third setigerous segment 
well separated from others. Following three 
segments setigerous, with notopodia dor- 
sally elevated in laterodorsal row, with no- 
topodia of same size as those of following 
segments. 


VOLUME 106, NUMBER 2 


Branchiae four pairs, all similar, arranged 
as funnel-like structure, with strong basal 
stem bearing small secondary filaments and 
thin terminal tip devoid of secondary fila- 
ments as long as basal stem. Branchial stem 
bearing one, sometimes two, bean-shaped 
vesicles on internal side near base. Second- 
ary filaments inserted along stem on two 
opposite areas; each leaf-shaped, strongly 
flattened, with median ciliated area, and 
pointed tip. These secondary filaments rem- 
iniscent of species of Alvinella. 

Notopodia, from setiger 1 to end of body 
(7th setiger excepted) each cylindrical, bear- 
ing two groups of capillary setae. Notopodia 
without digitiform lobes. Setiger 7 strongly 
modified, with slightly reduced cylindrical 
notopodia, bearing two to three straight short 
acicular notopodial setae on each side. Se- 
tiger 8 not modified. 

Cylindrical notopodia and uncinigerous 
neuropodial tori on each segment from se- 
tigerous segment 26 to 29 (with a majority 
from 28 to 29). Uncini numerous (20 to 50 
per torus), in single rows, with teeth directed 
anteriorly (retrogressive arrangement). Un- 
cini with only a single main tooth, lacking 
a secondary tooth. 

Pygidium rounded, with five conspicuous 
rounded papillae, two ventral paired papil- 
lae and three smaller, dorsal ones. 

Tubes unknown. 

Ecology.—From observations made on 
the White Lady hydrothermal site, the al- 
vinellid worms colonize the whitish mass 
of anhydrite, living close to high tempera- 
ture (285°C) translucent desalinated hydro- 
thermal fluid openings. On videotapes ob- 
tained by the submersible 3 CCD TV 
camera, the branchial fans of Paralvinella 
spp. are visible at the surface of the mass 
of anhydrite. The worms belong to P. uni- 
dentata and to P. fijiensis, another species 
(see below), that are not distinguishable dur- 
ing the sampling procedure. 

Discussion. —Since the discovery of the 
first species of Paralvinella, P. grasslei Des- 
bruyéres & Laubier, 1982, six species or 


9 | 


subspecies have been described within the 
genus: P. palmiformis Desbruyéres & Lau- 
bier, 1986, P. pandorae pandorae Desbru- 
yéres & Laubier, 1986, P. pandorae irlandei 
Desbruyéres & Laubier, 1986 (Desbruyéres 
& Laubier 1986), P. dela Detinova, 1988, 
P. hessleri Desbruyéres & Laubier, 1989 
(Desbruyéres & Laubier 1989) and P. bac- 
tericola Desbruyéres & Laubier 1991 (Des- 
bruyéres & Laubier 1991). 

Among these species, P. pandorae and its 
two subspecies are clearly distinguished by 
the presence of uncini from setigerous seg- 
ments 5 or 6 (depending on the subspecies), 
a unique situation within alvinellids con- 
sidered as a plesiomorphous character. 

The following morphological features can 
be used to assess the relationship between 
species and groups of related species: The 
total number of segments and its range of 
variation (primitive situation: 100 to 150 
segments, with a large range of individual 
variation; apomorphous situation, 60 to 80 
segments, with a reduced range of individ- 
ual variation); the rank of occurrence of the 
anteriormost neuropodial uncinigerous to- 
rus (first uncinigerous torus anterior to the 
modified setigerous segment in plesiomor- 
phous situation, first uncinigerous torus 
from segment 13 to more than segment 60 
in apomorphous situation). The range of 
variation increases with the rank of occur- 
rence; the buccal apparatus, with two dif- 
ferent basic types: the first one bears two 
large paired lateral tentacles in addition to 
the small ciliated dorsal tentacles, and a re- 
versible ventral globular organ; the second 
one lacks the large paired tentacles and has 
an unpaired pointed organ with a longitu- 
dinal slit and a reversible ventral globular 
organ. The development of the large paired 
tentacles adapted for different trophic be- 
haviors is considered as an apomorphic fea- 
ture. P. unidentata exhibits a unique buccal 
structure, reduced to the numerous grooved 
tentacles inserted on a well developed buc- 
cal membrane: this structure basically dif- 
fers from all previously described species of 


228 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


eC 


en 


SAAD 2 
ZZ 
(CG 
i 
\ | (77 


SAAS 


Ss 


Nie= 
©. 


Fig. 1. Paralvinella unidentata, new species. Entire animal in ventro-lateral view. 


Paralvinella; the shape and position of the posite or adjacent longitudinal rows. There 
secondary filaments of the branchiae. These is no obvious reason to consider either of 
can be cylindrical and slender or flattened these structures or situations more primi- 
and leaf-shaped, and inserted on two op-_ tive than the other one. Nevertheless, con- 


VOLUME 106, NUMBER 2 229 


: 2 
nS NLS iis 
9, Ly: j\ fi Hh 
f ; wl SA 
Y Za Yi WZ 
Si AS WZ 
— UIAS ys 2: EE 
EI LL—=_—— 
3 6 Nj y LS 


Fig. 2. Paralvinella fijiensis, new species. Entire animal in ventro-lateral view. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


230 


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VOLUME 106, NUMBER 2 


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232 


sidering the facts that the genus A/vinella 
possesses flattened leaf-shaped secondary 
filaments and is clearly apomorphic to the 
genus Paralvinella (Desbruyéres & Laubier 
1986), the question of the systematic rank 
of P. unidentata, with identical flattened leaf- 
shaped secondary filaments, must be con- 
sidered carefully; and the shape of the un- 
cini, in all alvinellids but P. unidentata, is 
characterized by one main tooth and one 
secondary smaller tooth, while in P. uni- 
dentata there is only one main tooth. Still, 
the general shape of the uncinus is identical 
in both groups and the presence or absence 
of a secondary tooth is not considered an 
important phylogenetic significance. How- 
ever, the absence of a secondary tooth can 
be considered plesiomorphic to the biden- 
tate semiavicular uncini previously known 
for all alvinellids. 

Within the genus Paralvinella, P. uniden- 
tata can be best compared with P. pandorae 
and its two subspecies in the structure of 
the buccal apparatus and the funnel-like ar- 
rangement of the branchiae. However, P. 
unidentata can be easily distinguished by a 
series of characters including the rank of 
occurrence and shape of neuropodial uncini 
and the leaf-shaped secondary filaments of 
the branchiae. 


Paralvinella fijiensis, new species 
Figs. 2, 5 


Type locality, material examined. — 
Ninety-five specimens collected and pre- 
served during Nautile dives PL 11 (3 spec- 
imens, White Lady site), PL 12 (1 specimen, 
White Lady site), PL 14 (47 specimens; 
White Lady site), PL 16 (10 specimens, 
White Lady site), PL 20 (34 specimens, 
White Lady site). All specimens come from 
White Lady vent site (depth 2000 m, 
16°59'50”S and 173°55'47’”E). Holotype 
(dive PL 14) deposited in the collections of 
the Muséum national d’Histoire naturelle, 
Laboratoire de Biologie des Invertébrés 
marins et Malacologie (n°UC 439). Para- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


types from same dive deposited in the col- 
lections of the National Museum of Natural 
History, Smithsonian Institution (USNM 
157043). 

Etymology. —The species is named for its 
geographic origin. 

Description. —Holotype 24 mm long and 
5 mm wide with 63 setigerous segments. 
Paratypes (15 specimens from PL 14) range 
from 50 to 68 setigerous segments. Color 
pale grey-pinkish in ethanol, with capillary 
setae and acicular notopodial hooks yellow; 
integument iridescent. Body maggot-shaped 
in large specimens; small animals with body 
gradually tapering posteriorly; medioven- 
tral area slightly depressed. When viewed 
under scanning microscope, integument of 
intersegmental areas densely covered with 
small circular glandular areas and secretions 
from same. 

Prostomium reduced medially with two 
anterior lobes and two nucal grooves 
obliquely situated near base. Prostomium 
laterally separated from lateral parts of buc- 
cal segment. Buccal segment laterally and 
ventrally visible, well separated from pro- 
stomium and segment II. Buccal apparatus 
comprising many grooved tentacles inserted 
dorsally and two large paired grooved ten- 
tacles inserted ventrally, ending with three 
unequally developed rounded lobes. Edge 
of main lobe provided with several rows of 
small rounded internal papillae. 

Branchial region comprised of four seg- 
ments, preceded by one reduced asetigerous 
segment still visible laterally and ventrally 
(segment IJ). First branchial segment ase- 
tigerous, ventrally visible. Next three bran- 
chial segments (setigerous segments | to 3) 
totally fused. Notopodia of first setigerous 
segment very reduced; notopodia of 2nd se- 
tigerous segment reduced; notopodia of 3rd 
setigerous segment similar to the following 
notopodia. Notopodia of setigers 1 to 3 ad- 
jacent to stem of external pair of branchiae. 
Fourth setigerous segment fused ventrally 
to fourth branchial segment. 

Branchiae four pairs, all similar, with a 


VOLUME 106, NUMBER 2 233 


50um 


500um 


Fig. 5. Paralvinella fijiensis, new species. A, anterior part in lateral view. B, buccal apparatus, showing 
grooved tentacles (right) and the terminal lobes of one of the ventral large tentacles (left). C, left parapod from 
anterior setiger, showing dorsal lobe. D, capillary seta covered with small spines and two rows of larger ones 


on the edge. E, modified notopodium of setiger 7, with large acicular hooks. F, uncinigerous torus, right side of 
body. 


234 


strong basal stem. Secondary filaments very 
abundant, inserted on two opposite areas of 
stem to end of branchia, reminiscent of Par- 
alvinella grasslei and P. palmiformis. 

Notopodia, from setiger 4 to the end of 
body (7th setiger excepted) cylindrical, 
bearing two groups of capillary setae heavily 
coated with filamentous bacteria. Notopo- 
dia from about setiger 9 to setiger 30 bearing 
dorsal and ventral rounded lobes. Setiger 7 
strongly modified, with two reduced dorsal 
and ventral lobes surrounding a very re- 
duced notopodium bearing three to four 
curved acicular notopodial setae on each 
side. Surface of acicular setae covered with 
minute spinelets; these are longer on convex 
side of acicular hooks when viewed under 
SEM. This structure similar to ornamen- 
tation of notopodial setae in other segments. 
Notopodia of setiger 8 with anterior part 
enlarged. 

Uncinigerous neuropodial tori present on 
each segment from setiger 12 to 19 (13 on 
the holotype). Occurrence of first uncini- 
gerous torus appears to be inversely pro- 
portional to total number of setigerous seg- 
ments: on a total of 23 individuals, with 
total number of setigerous segments from 
46 to 68, first uncinigerous segment occurs 
from 19th to 12th setigerous segment. Re- 
gression curve: 


R, = (96 — N)/2.54 


where R, = rank of occurrence of the first 
uncinigerous torus and N = total number 
of setigerous segments of the body, indi- 
cating that the anteriormost uncinigerous 
tori appear latest. 

Uncini numerous (20 to 50 per row) in 
single rows, with teeth facing anteriorly (ret- 
rogressive arrangement). Uncini increasing 
in number posteriorly. Each uncinus with 
one main tooth surmounted by smaller sec- 
ondary tooth, as in other species of Paral- 
vinella (except P. unidentata, see above). 

Pygidium blunt, without conspicuous pa- 
pillae. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Ecology. —Most specimens were collect- 
ed in anhydrite samples together with P. 
unidentata; on one occasion, a tube inhab- 
ited by P. fijiensis was found on a piece of 
basalt. 

Discussion. —Within the genus Paralvi- 
nella, P. fijiensis exhibits clear relationships 
with the stem species P. grasslei and its close 
relative P. palmiformis. These three species 
have in common: the structure of the buccal 
apparatus, with two large trilobate paired 
ventral tentacles; the general shape of the 
branchiae; the rank of occurrence of the un- 
cinigerous tori, starting between setiger 12 
(P. fijiensis, new species) and setiger 31 (P. 
palmiformis),; and the first setigerous seg- 
ment is very reduced, and the second setig- 
erous segment reduced, emphasizing the 
importance of the cephalization processes. 

Within this group of three species, P. fi- 
jiensis 1s characterized by the number of 
setigerous segments (maximum 68 in P. fi- 
Jiensis, compared to 110 in P. grasslei and 
118 in P. palmiformis), the small papillae 
of the main end of the trilobate paired buc- 
cal tentacles and the shape and relative size 
of the prostomium. Additional minor diff- 
ences can be found in the presence of spine- 
lets on the acicular hooks and the enlarged 
base of the 7th notopodium in P. fijiensis. 


Conclusion 


All Paralvinella species except P. hessleri 
and the two new species described herein 
are known from the East Pacific Rise and 
related ridge systems in the Eastern Pacific. 
Within this general framework, two differ- 
ent species groups can be distinguished in 
the Northern and Southern parts of the East 
Pacific ridge system. These parts have been 
separated by the subduction of the Ameri- 
can plate over the oceanic crust off Oregon 
starting 35 MY (Tunnicliffe 1988). P. pal- 
miformis and P. pandorae pandorae live in 
the Northern part, while P. grass/ei and P. 
pandorae irlandei live in the Southern part. 


VOLUME 106, NUMBER 2 


These two pairs of species (P. grasslei and 
P. palmiformis) or subspecies (P. p. irlandei 
and P. p. pandorae) have been regarded as 
examples of sibling species (Desbruyéres & 
Laubier 1986). Two additional species, P. 
bactericola in the Southern area and P. dela 
in the Northern one, have been discovered, 
providing a third example of a closely re- 
lated paired species. More recently, P. hess- 
leri, collected in the Mariana back-arc Basin 
in the Western Pacific, was found to exhibit 
a close relationship with P. bactericola and 
P. dela. Paralvinella fijiensis and P. uniden- 
tata herein described from the North Fiji 
Basin provide additional information about 
the striking similarities which can be found 
between the different groups of species liv- 
ing in these three major areas: 


Evolutionary groups: 


Group 1 Group 2 Group 3 


East Pacific Rise: 


P. grasslei P. p. irlandei P. bactericola 


Juan de Fuca-Explorer: 


P. palmifor-___P. p. pandorae_ P. dela 
mis 
Western Pacific: 
P. fijiensis P. unidentata _ P. hessleri 


It must be emphasized that each evolu- 
tionary group within the genus Paralvinella 
is present in each of the three geographic 
areas. Moreover, morphological differenti- 
ation is not related to the distance between 
the hydrothermal areas: the three species 
from the Western Pacific are much more 
different from one another than they are 
from related species in other biogeograph- 
ical areas. 

We hypothesize that these three different 
groups of Paralvinella species evolved in- 
dependently from three ancestors adapted 
to hydrothermal environmental conditions. 
As aconsequence of this hypothesis, we pro- 
pose the establishment of three subgenera, 
that can be diagnosed as follows: 


235 


Subgenus Paralvinella 


Type species. — Paralvinella (Paralvinella) 
grasslei Desbruyéres & Laubier, 1982 

Other species. —P. (P.) palmiformis Des- 
bruyéres & Laubier, 1986 and P. (P.) fijien- 
SES 

Diagnosis. — Buccal apparatus, compris- 
ing, from dorsum to ventrum, numerous 
buccal grooved retractile tentacles inserted 
on a semi-circular buccal membrane, two 
large paired trilobate appendages and a 
globular eversible ventral organ. Branchiae 
with cylindrical secondary filaments, in- 
serted on two opposite areas on the stem up 
to its end. Digitiform or rounded notopo- 
dial lobes present on some anterior setig- 
erous segments. 


Subgenus Miralvinella, new subgenus 


Type species.—Paralvinella (Miralvinel- 
la) dela Detinova, 1988 

Other species. —P. (M.) hessleri Desbru- 
yéres & Laubier, 1989 and P. (.) bacter- 
icola Desbruyéres & Laubier, 1991. 

Etymology. —Miralvinella, from Alvinel- 
la and MIR, the name of the two deep-sea 
Russian submersibles recently built in a 
Finnish shipyard for the Shirshov Institute 
of the former Academy of Sciences of the 
Soviet Union. 

Diagnosis.—Buccal apparatus complex, 
comprising from dorsum to ventrum nu- 
merous grooved tentacles inserted on a buc- 
cal membrane, two large deeply grooved ta- 
pering paired tentacles and a globular 
eversible ventral organ. Branchiae with cy- 
lindrical secondary filaments, inserted on 
two opposite areas on the stem up to its end. 
Digitiform notopodial lobes present on some 
anterior segments. 


Subgenus Nautalvinella, new subgenus 


Type species. —Paralvinella (Nautalvinel- 
la) pandorae Desbruyéres & Laubier, 1986 
Other species. —P. (N.) unidentata 


236 


Etymology. — Nautalvinella, from Alvi- 
nella and Nautile, the French deep-sea sub- 
mersible of IFREMER which was used dur- 
ing the STARMER cruise and previous 
expeditions on hydrothermal vents areas. 

Diagnosis. — Buccal apparatus comprising 
numerous grooved tentacles inserted dor- 
sally on a buccal membrane. A median hol- 
lowed pointed lobe can be present in some 
species. No large paired ventral tentacles. 
No ventral organ. Branchiae with numerous 
leaf-shaped secondary filaments inserted on 
two more or less adjacent lines on the stem. 
Terminal part of the stem devoid of sec- 
ondary filaments. No digitiform notopodial 
lobes. 


Literature Cited 


Auzende, J. M., etal. 1989. Le cadre géologique d’un 
site hydrothermal actif: la campagne STAR- 
MER 1 du submersible Nautile dans le Bassin 
Nord-Fidjien.—Comptes Rendus de |’Acadé- 
mie des Sciences, Paris, Série II 309:1787-1795. 

Both, R., et al. 1986. Hydrothermal chimneys and 
associated fauna in the Manus Back-Arc Basin, 
Papua New Guinea.—Eos 67:489—490. 

Desbruyéres, D., A. M. Alayse, & S. Ohta. 1991. Deep- 
sea hydrothermal communities in two back-arc 
basins of the south west Pacific (the North Fiji 
and Lau basins): composition, microdistribu- 
tion and food-web.— 6th Deep-Sea Biology 
Symposium, Copenhagen, July 1991, 20 pp. 

——, & L. Laubier. 1982. Paralvinella grasslei, new 
genus, new species of Alvinellinae (Polychaeta: 
Ampharetidae) from the Galapagos rift geo- 
thermal vents.—Proceedings of the Biological 
Society of Washington 95:484—494. 

———,,& . 1986. Les Alvinellidae, une famille 
nouvelle d’annélides polychétes inféodées aux 
sources hydrothermales sous-marines: systé- 
matique, biologie et écologie.—Journal Cana- 
dien de Zoologie 64:2227-—2245. 

, & 1989. Paralvinella hessleri, new 

species of Alvinellidae (Polychaeta) from the 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Mariana back-arc basin hydrothermal vents. — 
Proceedings of the Biological Society of Wash- 
ington 102:761-767. 

—, & 1991. Systematics, phylogeny, 
ecology and distribution of the Alvinellidae 
(Polychaeta) from deep-sea hydrothermal 
vents.— Ophelia Supplement 5:31-45. 

Detinova, N. N. 1988. New species of polychaetous 
annelids from hydrothermal vents of the Juan 
de Fuca ridge (Pacific Ocean). — Zoologichesky 
Zhurnal 57:858-864. 

Fouquet, Y., et al. 1990. Hydrothermal activity in 

the Lau Basin.—Eos, May 1:678-679. 

, etal. 1991. Hydrothermal activity and me- 

tallogenesis in the Lau back-arc basin. — Nature 

349:778-780. 

Hessler, R. R., P. Lonsdale, & J. Hawkins. 1988. Pat- 
terns on the ocean floor.—New Scientist 1605: 
47-48. 

Jollivet, D., et al. 1989. Premiéres observations de 
communautés animales associées 4 l’hydrother- 
malisme arriére-arc du bassin Nord Fidjien.— 
Comptes Rendus de |l’Académie des Sciences, 
Paris, Série III 309:301-308. 

Lonsdale, P. 1977. Clustering of suspension-feeding 
macrobenthos near abyssal hydrothermal vents 
at oceanic spreading centers.—Deep Sea Re- 
search 24: 857-863. 

Miura, T., & S. Ohta. 1991. Two polychaete species 
from the deep-sea hydrothermal vent in the 
middle Okinawa Trough.— Zoological Science 
8:383-387. 

Ohta, S. 1990. Deep-sea submersible survey of the 
hydrothermal vent community on the north- 
eastern slope of the Iheya Ridge, the Okinawa 
trough. —Jamstec Deepsea Research 2:145-156. 

Tunnicliffe, V. 1988. Biogeography and evolution of 
hydrothermal vent-fauna in the eastern Pacific 
Ocean.— Proceedings of the Royal Society of 
London B 223:347-366. 


(DD) Ifremer, Centre de Brest, B.P. 70, 
29263 Plouzané, France; (LL) Institut 
Océanographique, Laboratoire de Physio- 
logie des Etres marins, 195, rue Saint- 
Jacques, France. 


PROC. BIOL. SOC. WASH. 
106(2), 1993, pp. 237-242 


GYPTIS CRYPTA, A NEW HESIONID SPECIES FROM 
THE U.S.A. EAST COAST, WITH A REDESCRIPTION OF 
G. VITTATA WEBSTER & BENEDICT, 1887 
(ANNELIDA: POLYCHAETA) 


Fredrik Pleijel 


Abstract.—Gyptis crypta, new species, is described from North Carolina, 
Florida, and off Mississippi, and several previous records of G. vittata Webster 
& Benedict, 1887 are referred to this species. The new species is unique within 
the genus in having only ten terminal papillae on the proboscis, and in having 
segment 5 equipped with both noto- and neurosetae, rather than neurosetae 
only. The European species G. rosea (Malm, 1874) and G. mediterranea Pleiyel, 
1993 are considered the most closely related. Gyptis vittata is redescribed from 


type material. 


Subsequent to the original account, Gyp- 
tis vittata Webster & Benedict, 1887 has 
been reported on several occasions from the 
U.S.A. east coast (e.g., Pettibone 1963, Day 
1974, Gardiner 1975, Uebelacker 1984). 
Reexamination of the currently available 
specimens on which these records were 
based shows them to represent, in part (Pet- 
tibone 1963), a species of Podarkeopsis, 
possibly P. Jevifuscina Perkins, 1984, and, 
in part (Gardiner 1975, Uebelacker 1984), 
an undescribed species of Gyptis, herein de- 
scribed as G. crypta. Day’s specimens were 
not available for examination, and are con- 
sidered of uncertain status. Except for the 
original description from Maine, no valid 
literature records of G. vittata appear to ex- 
ist. To avoid further confusion, G. vittata is 
redescribed from the syntypes. 

A provisional diagnosis and a brief dis- 
cussion of the genus Gyptis Marion & Bob- 
retzky, 1875 is provided in Pleijel (1993). 

All drawings were made with a camera 
lucida. Width measurements are from me- 
dian segments and include parapodia but 
exclude cirri and setae. Apart from material 
of congeneric species used for comparison 
(detailed in Pleijel 1993), the study is based 
on specimens from the National Museum 


of Natural History, Smithsonian Institu- 
tion, Washington, D.C. (USNM). 


Gyptis vittata Webster & Benedict, 1887 
Fig. 1 


Gyptis vittata Webster & Benedict, 1887: 
Tis it6epl 1 hes. 21-22. al. tie 23. 
[Not descriptions and figures of Gyptis 
vittata sensu Pettibone, 1963:106—107, fig. 
28c—d.— Gardiner, 1975:119, fig. 8l-p.— 
Uebelacker, 1984:28-29 to 28-31, fig. 28- 
28a—g] 


Material examined. — Maine: 3 syntypes 
(USNM 452, including slides 503-506), 
Eastport, low water, rocks; 25—30 fm, shells. 
(Specimens apparently from two different 
but fused samples.) 

Description. —Prostomium rounded, 
about as wide as long, posteriorly with 
rounded lobes separated by posterior inci- 
sion (Fig. 1A). Palpophores cylindrical; pal- 
postyles widest medially, anteriorly round- 
ed (Fig. 1B), longer than palpophores. Paired 
antennae thinner and possibly longer (see 
Remarks) than palps, cylindrical with 
pointed tips. Median antenna short, club- 
shaped, widest subdistally (but see Re- 
marks); inserted half-way between anterior 


238 


pair eyes and anterior margin of prosto- 
mium. Anterior pair eyes rounded to reni- 
form, twice as large as posterior pair and 
situated further apart; posterior pair round- 
ed; both pairs with lenses. Nuchal organs 
not discernible. 

Large, distinct lip glands present laterally 
on the ventral lips (Fig. 1B). Proboscis prob- 
ably divided in proximal and distal parts 
(visible on a mounted specimen only where 
it is partly everted and difficult to observe). 
Number of terminal proboscideal papillae 
difficult to discern, possibly 15-20. Papillae 
in single ring, conical to cylindrical. 

Proximal parts of tentacular cirri indis- 
tinctly annulated, medial and distal parts 
distinctly annulated; rings about as long as 
wide; tips not observed (no complete ten- 
tacular cirri present). At least one acicula 
present in all cirrophores of dorsal tentac- 
ular cirri, not discernible in ventral ones. 
Anterior dorsal segmental delineations not 
distinct; segment 1 probably reduced aid 
following segments fused. 

Notopodia of segment 5 (setiger 1) with- 
out setigerous lobes or setae. Dorsal cirri 
incomplete, slightly stouter than those of 
following segments. Neuropodia similar to 
following ones. Ventral cirri similar to fol- 
lowing ones but slightly smaller. Segment 6 
similar to median ones. 

Notopodia of median segments situated 
on cirrophores, with distinctly annulated 
dorsal cirri, longer than notosetae. Rings ca. 
1.5 times as long as wide. Possible alter- 
nation of length and orientation of dorsal 
cirri not discernible. Notopodial lobes con- 
ical, with one or two internal aciculae. All 
notosetae simple; setal details not discern- 
ible. 

Neuropodia of median segments conical, 
with two internal aciculae, one larger and 
one smaller. All neurosetae compound; dis- 
tal part of shafts internally reticulated. Ven- 
tral cirri apparently smooth, shorter than 
neuropodial lobe, on small cirrophores sit- 
uated far back on the neuropodium (Fig. 
1C). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Pygidium not observed. 

Color: Eyes brown. No other pigmenta- 
tion retained. 

Habitat.—Currently known only from 
rocks, intertidal, and shells, 25-30 fm. 

Distribution. —Known only from the type 
locality. 

Remarks. —Webster & Benedict’s mate- 
rial consists of three syntypes: one in alco- 
hol, two mounted on slides (one dorsally 
and one ventrally), and two additional slides 
with mounted parapodia. All three speci- 
mens lack posterior ends. The unmounted 
specimen consists of an anterior end plus a 
median part, both in very poor condition. 
The mounted syntypes are both females with 
eggs, one approaching maturity with eggs 
about 120-140 um in diameter. A median 
antenna 1s present only on one of the 
mounted specimens, and, being of unusual 
shape for the group, it cannot be excluded 
that it is distorted (e.g., flattened by prep- 
aration). 

As seen from the descriptions as well as 
specimens deposited at USNM, Gardiner’s 
(1975) and Uebelacker’s (1984) descrip- 
tions of G. vittata both refer to G. crypta, 
new species, whereas Pettibone’s (1963) de- 
scription refers to a species of Podarkeopsis, 
possibly P. levifuscina, from Hadley Har- 
bor, Woods Hole, Massachusetts. Day 
(1973) recorded G. vittata from off Beaufort, 
and stated that the specimens were depos- 
ited at USNM and the Duke University Ma- 
rine Laboratory. Since these specimens are 
absent, however, the record could not be 
verified and is considered uncertain. 


Gyptis crypta, new species 
Fig. 2 


Gyptis vittata. —Gardiner, 1975:119, fig. 81- 
p.— Uebelacker, 1984:28—29 to 28-31, fig. 
28—28a—g [not Webster & Benedict, 1887]. 

Gyptis sp.— Taylor, 1971:167—171, fig. 4a-f. 


Material examined.—North Carolina: | 
paratype (USNM 52892) Wrightsville 
Beach, Banks Channel, intertidal in burrow 


VOLUME 106, NUMBER 2 


Fig. I. 


Gyptis vittata, syntypes. Setae omitted. A. Anterior end, dorsal view. B. Same as A but drawn with 


focus moved to ventral side of specimen; arrows indicate lip glands. C. Parapodium. A & B slide 505, C slide 


503. Scale lines 0.25 mm. 


of Notomastus lobatus, 20 Jul 1974; holo- 
type (USNM 52893) and 5 paratypes 
(USNM 157615), Wrightsville Beach, Banks 
Channel, intertidal in burrow of Notomas- 
tus lobatus, 18 Aug 1974; 5 paratypes 
(USNM 52894), Wrightsville Beach, Banks 
Channel, intertidal in burrow of Notomas- 
tus lobatus, 27 Feb 1975. Gulf of Mexico: 
18 paratypes (USNM 45534), Florida, 
Tampa Bay, 1963; 1 paratype (USNM 
75478), off Mississippi, 30°16.27'N, 
88°36.42'W, 3.4 m, 24 Oct 1980; 1 paratype 
(USNM 75319), off Florida, 24°47.5'N, 
82°13.16'W, 24 m, Jul 1981. 

Description. —Body, excluding parapo- 
dia, cylindrical, tapered posteriorly. Venter 


slightly flattened, without distinct median 
longitudinal furrow. 

Prostomium rectangular to trapezoidal 
with rounded corners (Fig. 2A), with small 
posterior incision (often difficult to discern). 
Palpophores cylindrical, palpostyles thin- 
ner, narrowing to rounded ends. Palpo- 
phores as long as or slightly longer than pal- 
postyles. Paired antennae probably situated 
on small ceratophores, slightly longer than 
or as long as palps, with fine tips. Median 
antenna inserted on or just in front of line 
between anterior pair of eyes, cylindrical or 
evenly tapering to a point, without extended 
tip. Anterior pair eyes rounded, larger than 
posterior pair and situated further apart, 


240 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


lS fy oss Sig © 


ae 
SQA 
ST 


Fig. 2. Gyptis crypta. A. Anterior end, dorsal view. Setae omitted. B. Anterior end, ventral view. Setae 
omitted. C. Left parapodium, segment 5, anterior view. D. Right parapodium, median segment, anterior view. 
About half of full number of setae shown. E. Acicular notoseta. F. Spinose capillary notoseta. G. Serrated 


capillary notoseta. H. Median neuroseta. I. Ventral neuroseta. A & C-I paratypes USNM 157615, B paratype 
USNM 45534. Scale lines A-B 0.25 mm, C 0.2 mm, D 0.2 mm, E-G 50 um, H-I 50 um. 


VOLUME 106, NUMBER 2 


posterior pair rounded, both pairs with lens- 
es. Nuchal organs lateral to prostomium, 
not coalescing dorsally. 

Lip glands absent (Fig. 2B). Proboscis 
short, divided in proximal and distal parts, 
without discernible papillae. Distal part 
narrow and smooth. Ten short, conical ter- 
minal papillae in single ring. 

Tentacular cirri thin and smooth, indis- 
tinctly annulated distally, tips distinctly 
pointed. Dorsal tentacular cirri of segments 
3 and 4 reaching to about segment 10 or 11, 
ventral tentacular cirri of segment 3 short- 
est, reaching to about segment 5. Aciculae 
in cirrophores of tentacular cirri not dis- 
cernible. Segment 1 dorsally reduced, seg- 
ments 2 and 3 probably fused dorsally. 

Notopodia of segment 5 (setiger 1) similar 
to following notopodia but slightly smaller 
and with fewer setae (Fig. 2C). Dorsal cirri 
similar to and as long as those of segment 
4. Neuropodia similar to following ones but 
slightly smaller and with fewer setae. Ven- 
tral cirri similar to following ones. 

Elevated dorsal ridges present across pos- 
terior side of each segment (Fig. 2D), most 
distinct on median and posterior segments. 

Notopodia of median segments with more 
or less distinctly annulated dorsal cirri 
(smooth proximally). Dorsal cirri about as 
long as setae, slightly shorter on anterior 
segments, with about five or six rings, each 
about three times as long as wide (Fig. 2D). 
All dorsal cirri slender, possible alternation 
of length and orientation not discernible. 
Notopodial lobes conical, usually with one, 
occasionally two, internal aciculae and three 
kinds of notosetae: 5—10 anteriorly situated, 
slightly curved acicular setae (Fig. 2E), ta- 
pering, distally with fine spines; slender cap- 
illary setae smooth proximally with two al- 
ternating rows of spines medially and 
subdistally (Fig. 2F); and few, rather stout, 
ventrally situated, serrated capillary noto- 
setae (Fig. 2G). 

Neuropodia of median segments conical, 
with one internal acicula, and about 20—40 
compound setae. Distal part of setal shafts 


241 


with transverse striation internally. Blades 
thin, dorsal serrated, median and dorsal ones 
long (Fig. 2H), ventral ones short (Fig. 21). 
Dorsally situated serrated capillary setae 
absent. Ventral cirri smooth with fine ta- 
pering tips (Fig. 2D), without cirrophores, 
situated distally on neuropodium. 

Pygidium rounded, pygidial cirri long and 
thin with pointed tips, longer than dorsal 
cirri, median papilla not observed. 

Color: Live specimens not observed. Pre- 
served specimens yellow with fine brown 
pigment spots, usually denser distally on 
noto- and neuropodia and on all cirri. Eyes 
reddish brown. Small dark spots may be 
present ventrally at parapodial bases. 

Measurements. —Only two complete 
specimens observed; length 7 mm, width 
1.4 mm for 39 segments; length 7.5 mm, 
width 1.2 mm for 39 segments. 

Habitat.—Currently known intertidally 
from burrows of Notomastus lobatus and 
down to 24 m depth. 

Distribution. —North Carolina, west coast 
of Florida and off Mississippi. 

Etymology.—The name crypta is trans- 
literated from the Greek noun “xputrn,” 
meaning haunt and hiding-place and refers 
to the habit of living in burrows of Noto- 
mastus lobatus. 

Remarks.—The holotype is a mature 
male, and the paratypes include several ma- 
ture females with eggs (50-60 um in di- 
ameter). Gametes were observed in speci- 
mens collected from August to February in 
North Carolina, although those collected in 
February did not appear fully mature. 

Gyptis crypta differs from all other known 
members of the genus in having a small and 
fixed number of terminal papillae on the 
proboscis, and in being provided with both 
noto- and neurosetae on segment 5 (rather 
than only neurosetae). Together with G. ro- 
sea Malm, 1874 and G. mediterranea Plei- 
jel, 1993 G. crypta shares the two putative 
derived characters distally inserted ventral 
cirri that taper evenly to a point (rather than 
subdistally inserted ventral cirri with short 


242 


tips as in other Gyptis). Apart from the 
smaller number of proboscis papillae and 
the biramous parapodia of the fifth segment, 
G. crypta is separated from these two species 
in having lighter colored eyes (probably red 
on live specimens) and a more elongated 
body. 

Although Gardiner (1975) examined the 
syntypes of G. vittata, his description of G. 
vittata and that of Uebelacker (1984) clearly 
characterize the new species, and specimens 
from both authors now constitute part of 
the type material of G. crypta. 


Acknowledgments 


I wish to thank H. G. Hansson and Ou- 
rania Papakosta for nomenclatural advice, 
K. Fauchald and L. Ward for loan of spec- 
imens as well as working facilities, B. Hilbig 
and T. Perkins for comments on the manu- 
script, and the Swedish Natural Science Re- 
search Council (contracts 9555-306 and 
-307) for financial support. 


Literature Cited 


Day, J.H. 1973. New Polychaeta from Beaufort, with 
a key to all species recorded from North Car- 
olina.—NOAA Technical Report NMFS 375: 
1-140. 

Gardiner, S. L. 1975. Errant polychaete annelids from 
North Carolina. — Journal of the Elisha Mitchell 
Scientific Society 91:77-—220. 

Malm, A. W. 1874. Annulater i hafvet utmed Sveri- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ges vestkust och omkring G6teborg.—G6te- 
borgs K. vetenskaps- och vitterhetssamhilles 
handlingar 14:67-105. 

Marion, A. F., & N. Bobretzky. 1875. Etude des An- 
nélides du Golfe de Marseille. — Annales des sci- 
ences naturelles 2:1-106. 

Perkins, T. H. 1984. New species of Phyllodocidae 
and Hesionidae (Polychaeta), principally from 
Florida.— Proceedings of the Biological Society 
of Washington 97:555—582. 

Pettibone, M. H. 1963. Marine polychaete worms of 
the New England region. — Bulletin of the Unit- 
ed States National Museum 227:1-356. 

Pleijel, F. 1993. Taxonomy of European species of 
Amphiduros and Gyptis (Polychaeta: Hesioni- 
dae).— Proceedings of the Biological Society of 
Washington 106:158-181. 

Taylor, J.L. 1971. Polychaetous annelids and benthic 
environments in Tampa Bay, Florida. Unpub- 
lished, Ph.D. Dissertation, University of Flor- 
ida, Gainesville, 1332 pp. 

Uebelacker, J. M. 1984. Chapter 28. Family Hesioni- 
dae Sars, 1862. Pp. 28:1-—28:39 in J. M. Uebe- 
lacker and P. G. Johnson, eds., Taxonomic guide 
to the polychaetes of the northern Gulf of Mex- 
ico. Final report to the Minerals Management 
Service, contract 14-12-001-29091. Barry A. 
Vittor & Associates, Inc., Mobile, Alabama. 7 
vols. 

Webster, H. E., & J. E. Benedict. 1887. The Annelida 
Chaetopoda from Eastport, Maine.— Report of 
the United States Commissioner of Fisheries 
1885:707-755. 


Swedish Museum of Natural History, 
Stockholm, and (postal address): Tjarno 
Marine Biological Laboratory, Pl. 2781, 
S-452 96 Stromstad, Sweden. 


PROC. BIOL. SOC. WASH. 
106(2), 1993, pp. 243-250 


A NEW SPECIES OF UNIPORODRILUS 
(OLIGOCHAETA: TUBIFICIDAE) FROM THE GULF OF 
MEXICO COAST OF FLORIDA, AND A 
PHYLOGENETIC ANALYSIS OF THE GENUS 


Christer Erséus and Michael R. Milligan 


Abstract. — Uniporodrilus purus, new species, from subtidal sand near Pen- 
sacola in northwestern Florida, is described. It is distinguished from all con- 
geners by lacking penial setae. A most parsimonious cladistic hypothesis of the 
species of Uniporodrilus (subfamily Phallodrilinae) is presented. It suggests a 
basal dichotomy of the genus, with U. purus and U. granulothecus, both subtidal 
species, constituting one clade characterized by unpaired male and sperma- 
thecal pores, and the four remaining species, all intertidal, representing another 
clade characterized by a high number of somatic setae and enlarged pharyngeal 


cavity. 


The genus Uniporodrilus was established 
for Uniporodrilus granulothecus Erséus, 
1979, a subtidal marine tubificid from the 
east coast of the United States (Erséus 1979). 
Davis (1985) described a second species as- 
signed to this genus, Uniporodrilus vestig- 
ium, from Georges Bank off Massachusetts. 
In a recent revision of the subfamily Phal- 
lodrilinae (Erséus 1992a), however, the lat- 
ter species was transferred to the genus Arf- 
lantidrilus Erséus, 1982, and at the same 
time, Uniporodrilus was enlarged to include 
also three taxa previously placed in Phal- 
lodrilus Pierantoni, 1902: Phallodrilus scir- 
piculus Erséus, 1985 (from Saudi Arabia), 
Phallodrilus nasutus Erséus, 1990(a), and 
Phallodrilus bipartitus Erséus, 1990(a) (both 
from the Caribbean area). A fifth species 
recently described from Hong Kong, Uni- 
porodrilus furcatus Erséus, 1992(b) is also 
regarded as a member of this genus, which 
has been defined by its “‘varying”’ penial se- 
tae (setae different in size and morphology 
within bundles) and its granulated sper- 
mathecal ducts (Erséus 1992a). 

In oligochaete material collected in north- 
western Florida by Mr. Jerry McLelland 
(Gulf Coast Research Laboratory, Ocean 


Springs, Mississippi), an additional species 
of Uniporodrilus was discovered. It is de- 
scribed in the present paper, which also in- 
cludes a cladistic analysis of the species 
within this genus. 

All specimens of Uniporodrilus purus, new 
species, were stained in paracarmine and 
mounted whole in Canada balsam. The type 
series is deposited in the U.S. National Mu- 
seum of Natural History (USNM), Smith- 
sonian Institution, Washington, D.C. For 
comparisons, the whole-mounted speci- 
mens in the type material of U. granulothe- 
cus (also in the USNM) were re-examined. 

Cladograms were constructed under the 
principle of maximum parsimony, using the 
program PAUP (Phylogenetic Analysis Us- 
ing Parsimony, version 3.0) by Swofford 
(1990). 


Uniporodrilus Erséus, 1979 


Uniporodrilus Erséus, 1979:414-415.— 
1992a:22-23. 

Uniporodrilus (partim): Davis 1985:169. 

Phallodrilus Pierantoni (partim): Erséus 
1990b:54. 


Diagnosis (emended after Erséus 


244 


1992a) (assumed autapomorphies under- 
lined).—Small marine tubificids. Prosto- 


mium distinctly longer than its width at base; 
generally well set off from peristomium (see 
Fig. 1). Somatic setae bifid with upper tooth 
thinner and shorter than lower. [Posterior 
dorsal setae sharply single-pointed in U. na- 
sutus.| Penial setae, when present, in fan- 
shaped bundles or in rings, with bifid or 
single-pointed, hooked tips. Penial setae ab- 
sent [in U. purus], or from about 6 to about 
22 per bundle. Spermathecal pores un- 
paired, mid-ventral in posterior part of IX 
[in U. granulothecus and U. purus], or paired 
in line with ventral setae in X [in all other 
species]. Pharyngeal cavity large and much 
ciliated in some species. Vasa deferentia ei- 
ther entering apical end of, or somewhat 
subapical on, atria. Atria small, oval, or 
elongate, curved; when paired, each with 
two large prostate glands [atrium unpaired 
in U. granulothecus and U. purus, with four 
prostates, at least in U. granulothecus]. Atria 
Opening into more or less developed cop- 
ulatory sacs. Spermathecae [unpaired in U. 
granulothecus and U. purus] elongate, with 
thick-walled, granulated ducts, and small 
ampullae. 

Type species. — Uniporodrilus granulothe- 
cus Erséus, 1979. 

Remarks. — The characteristic shape of the 
prostomium was referred to as “‘snout-like”’ 
by Erséus (1992a:23). For differences be- 
tween this diagnosis and the one by Erséus 
(1992a), see Discussion below. 


Uniporodrilus granulothecus Erséus, 1979 
Fig. 1A-B 


Uniporodrilus granulothecus Erséus, 1979: 
415-417, figs. 1-3.-Erséus 1992a:22. 


Material re-examined. —The holotype 
(USNM 56307) and 3 paratypes (USNM 
56308, 56309), all whole-mounted. 

Remarks. —In the character matrix used 
for the cladistic analysis of the Phallodrili- 
nae (Erséus 1992a:table I), the prostomium 
of U. granulothecus was coded as “‘not snout- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


like.”” This was inferred from the original 
description (Erséus 1979), which does not 
include any particular note on the prosto- 
mium, except that it is “rounded, about as 
long as its width at peristomium.”’ The re- 
examination of the type specimens, how- 
ever, showed that the prostomium of this 
species (Fig. 1A—B) is very similar to that 
of any congener. 

All the re-examined specimens have di- 
atoms in their guts, which suggests that U. 
granulothecus selectively feeds on these al- 
gae. 


Uniporodrilus purus, new species 
Figs. 1C—D, 2 


Holotype. —USNM_ 157046, whole- 
mounted specimen. 

Type locality.—Perdido Key, near Pen- 
sacola, Gulf of Mexico coast of Florida; 500 
m from shore, about 6 m depth, sand (Oct 
1989; J. McLelland). 

Paratypes.—USNM_ 157047-157049, 3 
specimens from type locality. 

Other material.—Milligan collection: 2 
specimens from type locality. 

Etymology.—The species epithet purus 
(Latin for ‘simple,’ ‘plain’) refers to the lack 
of penial setae, and to the single unpaired 
atria and spermathecae, in this species. All 
congeners have penial setae, and all but one 
(U. granulothecus) have paired genitalia. 

Description. —Fixed worms stout, 1.5-1.9 
mm long, 0.16-0.23 mm wide at segment 
XI; 25-35 segments. Prostomium (Fig. 1C-— 
D) large, clearly longer than its width at 
base. Clitellum extending over 12X—XII. Se- 
tae (Fig. 2A—B) bifid with upper tooth re- 
duced; reduction more pronounced in pos- 
terior setae (Fig. 2B) than in anterior ones 
(Fig. 2A); lower tooth is prolonged in pos- 
terior setae (Fig. 2B). Setae 30-45 um long, 
about 1—2 um thick, three per bundle 
throughout body; but absent ventrally from 
XI (penial setae absent). Male pore un- 
paired, mid-ventral, posterior to middle of 
XI. Spermathecal pore unpaired, mid-ven- 
tral, in most posterior part of IX. 


VOLUME 106, NUMBER 2 


Fig. 1. 


245 


200 um 


A-B, Uniporodrilus granulothecus, anterior ends of two paratypes (USNM 56308, 56309); C-D, U. 


purus, new species, anterior ends of holotype (C) and one paratype (D). Abbreviation: prm, prostomium. Dense 
shading indicates epidermal lining, light shading (coelomic?) space within prostomium. 


Pharyngeal cavity hollow, but not large. 
Pharyngeal glands in (III)IV—V. Male gen- 
italia (Fig. 2C) partly paired, partly un- 
paired. Vasa deferentia paired, much longer 
than atrium, conspicuously widened (up to 
15-22 um wide) along most parts; cilia few 
(or absent?), but scattered spermatozoa of- 
ten present in vasa. Both vasa deferentia 
entering unpaired atrium somewhat sub- 


l 50 um 


Fig. 2. 


vdo 


apically. Atrium oval, 30—40 um long, 27- 
30 um wide, with thin outer muscular lining 
(1-2 wm thick), and ciliated and somewhat 
granulated inner epithelium. Atrium open- 
ing into unpaired copulatory sac (but details 
not clear in available material). Lobed bod- 
ies of prostate glands located posterio-lat- 
eral to atrium (on both sides). They appear 
broadly attached to atrium, but a division 


Uniporodrilus purus, new species, A, Free-hand drawing of anterior seta. B, Free-hand drawing of 


posterior seta. C, Lateral view of spermatheca and male genitalia in segments X—XI. Abbreviations: a, atrium; 
cs, copulatory sac; pr, prostate gland; pro, prostate gland of other side of worm; s, spermatheca; vd, vas deferens; 


vdo, vas deferens of other side of worm. 


246 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


24 8 


6 


granulothecus 


purus 


furcatus 


nasutus 


4 611 


bipartitus 


scirpiculus 


furcatus 


scirpiculus 


nasutus 


11 bipartitus 


Fig. 3. The two most parsimonious trees (A—B) obtained with the PAUP program for the six ingroup taxa 
of Uniporodrilus (Table 1) and their 13 characters (listed in text, coded in Table 1). Search method: branch- 
and-bound. Length of trees 17 steps, consistency index 0.765. Rooting at outgroup (Nootkadrilus), which is 
excluded from trees. The clade granulothecus/purus is identical in both trees, and therefore only shown in A. 
The numbers refer to characters. Filled rectangle, autapomorphy; open rectangle, apomorphy that is later followed 
by reversal; two parallel lines, convergence; cross, reversal. 


into anterior and posterior glands (as is nor- 
mal for Phallodrilinae) not obvious; either 
anterior or posterior prostates possibly ab- 
sent. Spermatheca (Fig. 2C:s) unpaired, 
elongate pear-shaped, 85-100 um long, 
maximally 33—45 um wide. Outer, duct-like 


part of spermatheca somewhat granulated, 
longer and more thick-walled than inner part 
(ampulla); lumen of duct, however, irregu- 
lar. Sperm scattered throughout duct as well 
as ampulla. 
Remarks. — This new species is closely re- 


er 


VOLUME 106, NUMBER 2 


247 


Table 1.—Data matrix for parsimony analysis of the species of Uniporodrilus (Fig. 3). For characters and 


character states, see text. 


Taxon 


Character states, characters 1-13 


Outgroup: 
Nootkadrilus Baker, 1982 
Ingroup: 
Uniporodrilus granulothecus Erséus, 1979 
U. purus, n. sp. 
U. furcatus Erséus, 1992b 
U. scirpiculus Erséus, 1985 
U. nasutus Erséus, 1990a 
U. bipartitus Erséus, 1990a 


00?00 00000 000 
10001 01011 011 
10??? 210M O11 
11110 01100 011 
11011 00100 001 
11100 10100 101 
11100 10100 101 


lated to U. granulothecus (cf. Erséus 1979). 
Both taxa have somewhat inflated vasa def- 
erentia, as well as unpaired atrium and sper- 
matheca. Moreover, in these two species, 
the spermathecal pore is in segment IX, not 
in X as is normal for Tubificidae. Unipo- 
rodrilus purus is, however, unique within 
the genus by its lack of penial setae. It is 
further distinguished from U. granulothecus 
by its stouter atrium, and its less clearly 
bipartite spermatheca (in U. granulothecus 
the spermathecal duct is much longer than 
ampulla and has densely granulated walls). 

The condition of the prostate glands (with 
one pair possibly absent) needs to be con- 
firmed when new, preferably sectioned, ma- 
terial becomes available. If any of the pros- 
tate glands is absent, this feature is an 
additional autapomorphy for U. purus. 

The guts appear empty in all examined 
specimens. Thus the diet of U. purus is un- 
known (cf. Remarks for U. granulothecus 
above). 

Distribution and habitat.—Known only 
from the type locality in the northeastern 
part of the Gulf of Mexico. Subtidal sand, 
at 6 m depth. 


Phylogenetic Analysis 


Taxa.—The six species of Uniporodrilus 
(Table 1) are the ingroup taxa in the anal- 
ysis. Nootkadrilus Baker, 1982, was selected 
as the outgroup; according to the cladistic 
analysis of the whole subfamily Phallodri- 


linae (Erséus 1992a), this genus possibly is 
the sister group of Uniporodrilus. For the 
coding of most of the outgroup character 
states, the descriptions by Baker (1982) were 
used. For character 9 in the list below, how- 
ever, material of an unidentified Nootkad- 
rilus species and of N. longisetosus (Brink- 
hurst & Baker, 1979) in the senior author’s 
collection was examined. 

Characters. —The following characters 
and character stares were used. In the PAUP 
run, reversals were allowed for all charac- 
ters. 

1. Prostomium not prolonged (0); dis- 
tinctly longer than its width at base (“‘snout- 
like’’), and well set off from peristomium 
(1). 

2. Somatic setae maximally three or four 
(occasionally five) per bundle (0); maxi- 
mally five to seven (occasionally even eight 
or nine) per bundle (1). 

3. Penial setae all single-pointed (0); at 
least some penial setae bifid (1). 

4. Penial setae of about the same size 
within bundle (0); some penial setae dis- 
tinctly larger than others within bundle (1). 

5. Within each bundle, penial setae 
forming a straight row that is parallel to long 
axis of worm (0); penial setae forming ring 
within each bundle (1). 

6. Penial setae one continuous group 
within each bundle (0); penial setae in two 
groups within each bundle (1). 

7. Male pores paired (0); unpaired (1). 


248 


8. Pharyngeal cavity normal, not en- 
larged (0); enlarged (1). 

9. Pharyngeal glands clearly extending 
into segment VI (occasionally into VII) (0); 
not extending further than into V (1). 

10. Atria and spermathecae paired, sper- 
mathecae in segment X (0); atrium and sper- 
matheca unpaired, spermatheca in IX (1). 

11. Atria elongate, not small (0); pear- 
shaped, small (1). 

12. Prostate glands not broadly attached 
to atria (0); at least one prostate gland 
broadly attached to atrium/atria (1). 

13. Spermathecal ducts not granulated 
(O); granulated (1). 

Results.—The branch-and-bound-algo- 
rithm of PAUP 3.0 was used to find the two 
most parsimonious trees (A and B), which 
both are shown in Fig. 3. Both trees are fully 
resolved, with U. granulothecus and U. pu- 
rus forming one clade, and the four other 
species as the sister group of this clade. The 
topologies of the two trees differ with regard 
to the branching pattern within the larger 
sister group: U. furcatus is either grouped 
with U. nasutus and U. bipartitus (tree A), 
or with U. scirpiculus (tree B). Uniporodrilus 
nasutus and U. bipartitus are always togeth- 
er, as they have identical character codings 
(see Table 1). 

However, the character transformations 
indicated in Fig. 3A—B are not the only most 
parsimonious optimizations of the present 
data set. Although not affecting the branch- 
ing pattern, in tree A, the apomorphic state 
of character 4 can be interpreted as con- 
vergent for U. furcatus and U. scirpiculus, 
instead of assuming reversal in the clade 
nasutus/bipartitus (as shown in Fig. 3A). In 
tree A also, character 5 could be regarded 
as a basal apomorphy for the whole ingroup, 
assuming a reversal in the clade furcatus/ 
nasutus/bipartitus, or it could be seen as 
independent autapomorphies for U. gran- 
ulothecus and U. scirpiculus (U. purus was 
coded as “‘?”? for this character as it lacks 
penial setae altogether). In a similar way, in 
tree B, character 3 could be regarded as con- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


vergent apomorphies for U. furcatus and the 
nasutus/bipartitus clade. Still, none of these 
alternative optimizations, or combinations 
of them, will give other tree topologies than 
those in Fig. 3. 


Discussion 


In the previous revision (Erséus 1992a), 
the varying size and morphology of the pe- 
nial setae within the bundles, and the gran- 
ulation of the spermathecal ducts, were 
assumed to be autapomorphies of Unipo- 
rodrilus. The “‘snout-like’’ prostomium 
(character 2 in Erséus 1992a:fig. 4) was then 
interpreted as a synapomorphy of U. scir- 
piculus, U. nasutus, U. bipartitus and U. fur- 
catus (the last-mentioned called “Hong 
Kong sp. 1”’ by Erséus 1992a). The present 
study has shown that U. granulothecus as 
well as the new taxon U. purus have pro- 
longed prostomia (Fig. 1) and thus, this fea- 
ture (character 1 in Fig. 3) can be used to 
define the genus as a whole. 

The character state “‘penial setae of at least 
two different kinds within bundle”’ (char- 
acter 10 in Erséus 1992a), on the other hand, 
is treated in a more restricted sense here 
(character 4 in present paper). In the present 
analysis, only those two species (U. furcatus 
and U. scirpiculus) with distinctly aniso- 
morphic penial setae are coded as apomor- 
phic for this character (character 4 in Fig. 
3). The variation of setal morphology in the 
penial bundles of U. granulothecus, U. na- 
sutus and U. bipartitus is slight, and in U. 
purus, penial setae are absent. 

The spermathecal duct is not as long (in 
relation to the ampulla) in U. purus as in 
the other species, but it is granulated (al- 
though not as heavily as in the congeners), 
and therefore the granulation of the sper- 
mathecal ducts (character 13 in Fig. 3) is an 
apomorphy that still supports the mono- 
phyly of Uniporodrilus. 

The present parsimony analysis suggests 
a basal dichotomy of Uniporodrilus, with U. 
granulothecus and U. purus forming the sis- 


VOLUME 106, NUMBER 2 


ter group of the rest of the genus (Fig. 3). 
Monophyly of the granulothecus/purus clade 
is unequivocally supported by the unpaired 
male pore (character 7), the restricted ex- 
tension of the pharyngeal glands (character 
9), the unpaired atrium and spermatheca, 
with the latter in segment IX (character 10), 
and the broad attachment of the anterior 
prostates (character 12); characters 7 and 
12, however, also (convergently) apomor- 
phic in U. furcatus. Depending on which 
character optimization one prefers, one may 
also use the ring-shaped penial bundles 
(character 5) to define this group, based on 
the assumption that the penial setae were 
forming rings in the most recent ancestor of 
U. granulothecus and U. purus, but not in 
the common ancestor of the whole genus 
(see Remarks for the Phylogenetic analysis 
above). Moreover, both U. granulothecus 
and U. purus have distinctly widened parts 
of the vasa deferentia. This feature appears 
synapomorphic, but it is not conclusive 
whether it is unique to these two species, as 
the vasa deferentia have not been visible in 
their full lengths in the other members of 
the genus. 

Monophyly of the four other species is 
supported by the high number of somatic 
setae (character 2 in Fig. 3) and the enlarged 
pharyngeal cavity (character 8). Further, two 
characters suggest that U. nasutus and U. 
bipartitus are closely related, the disjunct 
distribution of setae within the penial bun- 
dles (character 6), and the small, pear-shaped 
atria (character 11). With regard to the phy- 
logenetic position of U. furcatus and U. scir- 
piculus, the parsimony analysis is less de- 
cisive. Depending on which one of characters 
3 (the bifid penial setae) or 4 (the distinctly 
anisomorphic penial setae) is considered as 
homoplasious, U. furcatus may either be re- 
garded as the sister taxon of U. scirpiculus 
(Fig. 3B), or as the sister taxon of the na- 
sutus/bipartitus group (Fig. 3A). 

Habitat and geographical distribution 
were not used as “characters” in the par- 
simony analysis, but the monophyly of U. 


249 


granulothecus and U. purus seems to be sup- 
ported by their subtidal habitat, as all the 
other four species, as well as the outgroup 
(Nootkadrilus), are intertidal. Furthermore, 
the possibility that U. furcatus (from Hong 
Kong) and U. scirpiculus (from Saudi Ara- 
bia) are endemic to the Indo-West Pacific 
region appears to favor their monophyletic 
Status, i1.e., in accordance with tree B (Fig. 
3); the congeners are all from the Northwest 
Atlantic and Nootkadrilus is so far only 
known from the Northeast Pacific. 


Acknowledgments 


We are indebted to Mr. Jerry McLelland 
for providing the material of U. purus; to 
Ms. Barbro Lofnertz (University of Gote- 
borg), and Ms. Christine Hammar (Swedish 
Museum of Natural History), for technical 
assistance; and to the Swedish Natural Sci- 
ence Research Council, for financial sup- 
port. 


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tralia. Volume 1. Western Australian Museum, 
Perth, 437 pp. 

1992a. A generic revision of the Phallodri- 
linae (Oligochaeta, Tubificidae.— Zoologica 
Scripta 21:5—48. 

1992b. Hong Kong’s marine Oligochaeta: a 
supplement. Pp. 157—180 in B. Morton, ed., The 
marine flora and fauna of Hong Kong and south- 
ern China III. Proceedings of the Fourth Inter- 
national Marine Biological Workshop: the ma- 
rine flora and fauna of Hong Kong and Southern 
China, Hong Kong, 11-29 April 1989. Volume 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


1. Hong Kong University Press, Hong Kong, 
526 pp. 

Pierantoni, U. 1902. Due nuovi generi di Oligocheti 
Marini rinvenuti nel Golfo di Napoli.—Bollet- 
tino della Societa di Naturalisti i Napoli 16:113- 
ey: 

Swofford, D. L. 1990. PAUP. Phylogenetic Analysis 
Using Parsimony. Version 3.0. Computer pro- 
gram distributed by the Illinois Natural History 
Survey, Champaign. 


(CE) Department of Invertebrate Zoolo- 
gy, Swedish Museum of Natural History, 
Box 50007, S-104 05 Stockholm, Sweden; 
(MRM) Mote Marine Laboratory, 1600 
Thompson Parkway, Sarasota, Florida 
34236, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(2), 1993, pp. 251-295 


A CHECKLIST OF AND ILLUSTRATED KEY TO THE 
GENERA AND SPECIES OF THE CENTRAL AND 
NORTH AMERICAN CAMBARINCOLIDAE 
(CLITELLATA: BRANCHIOBDELLIDA) 


Perry C. Holt and Brent D. Opell 


Abstract. —This paper treats the 90 species of Cambarincolidae known from 
Central and North America. Each species is illustrated and its synonyms, tax- 
onomic references, type specimen disposition, and distribution are given. Keys 
are provided to the family’s nine genera and to the 48 species of Cambarincola, 
2 species of Ceratodrilus, 7 species of Ellisodrilus, 4 species of Oedipodrilus, 8 
species of Pterodrilus, and 18 species of Sathodrilus, Magmatodrilus, Tetto- 
drilus, and Triannulata are each represented by a single species. 


The branchiobdellidans are obligate epi- 
zoites on freshwater crustaceans throughout 
the Holarctic region except, apparently, in 
Central Asia between the Ural Mountains 
and the Amur drainage (Holt 1968a). The 
greatest diversity of families and species is 
found in North America. This may be an 
artifact of collecting, as we have less infor- 
mation from other regions except Europe 
where only a few species are found. In Asia 
only Yamaguchi has done any significant 
work on the Japanese and Korean faunas. 
Despite the greater body of work on the 
North American branchiobdellidans, it is 
likely that only a small fraction of this fauna 
has been accorded taxonomic treatment. 
Most genera and species of North American 
branchiobdellidans are assigned to the fam- 
ily Cambarincolidae (Holt 1986). The phy- 
logenetic position and classification of the 
Branchiobdellida is discussed by Gelder & 
Brinkhurst (1990), Holt (1986, 1989a), and 
Sawyer (1986). 


Methods 


Types or topotypes of all taxa of the cam- 
barincolids have been studied in Holt’s lab- 
oratory and detailed methods for preserving 


and studying specimens are found in Holt 
(1986). Additional methods include Gelder 
& Hall’s (1990) use of a mixture of clove 
oil and methyl salicylate (oil of winter- 
green), as branchiobdellidans, particularly 
gill-inhabiting forms, are often twisted and 
distorted when methyl salicylate alone is 
used in the preparation of entire animals. 
The use of Nomarski differential interfer- 
ence contrast optics gives a much clearer 
view of internal structures in whole animals 
than does bright field illumination. When 
time and the availability of specimens per- 
mit, the worms should be relaxed with an 
appropriate agent, such as chloral hydrate 
and, after fixation in alcohol-formalin, 
lightly stained. For field collection, alcohol- 
formalin (Holt 1963) still seems the best and 
easiest preservative to use. 

All of the branchiobdellidan material, in- 
cluding types, studied by Holt and their ac- 
companying field notes have been deposited 
in the collections of the National Museum 
of Natural History, Smithsonian Institu- 
tion, Washington, D.C., U.S.A. These spec- 
imens are identified by the accession num- 
bers (USNM) of the National Museum, or 
by Holt’s accession numbers (PCH ...). A 
few paratypes are in the collection of the 


F 


Fig. 1. A, generalized branchiobdellidan in lateral view; Abbreviations as follows: I-XI, Trunk Segments; 
B, Bursa; BR, Brain; C, Circulatory System; DR, Dorsal Ridge; DRP, Digital Projections of the Dorsal Ridge; 
EBP, Ental Bulb of Prostate; ELS, Ental Lobe of the Seminiducal Gland; EPS, Ental Process of Spermatheca; 
ED, Ejaculatory Duct; EP, Everted Penis; FVE, Funnel of Vas Deferens; HDP, Histologically Differentiated 
Prostate; INT, Intestine; IPR, Incompletely Separated Prostate; J, Jaws; NP, Nephridiopore; OV, Ovary; PR, 
Prostate; PS, Penial Sheath; PT, Peristomal Tentacles; PP, Prostatic Protuberance; PB, Prostate Bulb; RP, 
Retracted Penis; SPG, Seminiducal Gland; SB, Bulb of Spermatheca; SD, Spermathecal Duct; SMR, Super- 
numerary Muscles of Dorsal Ridge; VN, Ventral Nerve Cord; VD, Vas Deferens (from Holt 1969, 1986); B—C, 
eversible penis of Pterodrilus alcicornis, B, withdrawn; C, everted; D-E, eversible penis of Cambarincola pamelae; 
D, withdrawn; C, everted; F—G, protrusible penis of Oedipodrilus oedipus; F, withdrawn; G, protruded, H, 
withdrawn protrusible penis of Oedipodrilus cuetzalanae; I, protruded protrusible penis of Oedipodrilus macbaini. 
Unless otherwise noted, the figures of each species on the following plates are arranged, from left to right, as 
follows: lateral view of whole specimen, lateral view of reproductive system, ventral view of upper jaw (top) 
and dorsal view of lower jaw (bottom); and lateral views of upper jaw (top) and lower jaw (bottom). Abbreviations 
are given above. 


VOLUME 106, NUMBER 2 


Instituto de Biologia, Universidad Auto- 
noma de México, Ciudad México, D.F., 
México, and indicated by the initials IBUM. 

The characters used in the taxonomy of 
the branchiobdellidans (Fig. 1A) have 
changed over the years with the recognition 
that structures once thought to be common 
to all members of the class are not so and 
the discovery of previously unrecognized 
features. Summaries, in each case incom- 
plete, may be found in Yamaguchi (1934), 
Goodnight (1940), Holt (1953, 1960a, 
1960b), Hoffman (1963), and Gelder & 
Brinkhurst (1990). An example of this is the 
use by Gelder & Hall (1990) of the number 
of oral papillae to separate species. Previ- 
ously these putative sensory structures were, 
at best, simply noted as present or absent. 
Further studies should be done to confirm 
the taxonomic usefulness of these papillae. 

In the preparation of this paper all figures 
were either redrawn from the originals or 
from enlarged illustrations in the literature 
in order to achieve uniformity of style. The 
correctly constructed term “seminiducal 
gland”’ replaces “‘spermiducal gland” in the 
taxonomic keys. This paper cites primarily 
taxonomic papers and its literature cited is, 
therefore, not intended to be a complete 
bibliography of the branchiobdellidans. The 
species names of all crayfish host records 
have been updated according to Hobbs 
(1989). As the literature citations for these 
names are provided by Hobbs, they are not 
repeated in this paper. Unless otherwise 
noted, all hosts are crayfish. 

Nomen inquirendum. —Cambarincola 
Okadai Yamaguchi, 1933. This nominal 
species was described from a specimen in- 
troduced with its American host into Lake 
Chuzenji, Nikko, Japan. Unfortunately, the 
location of the type(s) is unknown and the 
description could apply to any one of sev- 
eral American species. 


The Checklist 
Cambarincola Ellis, 1912 
Cambarincola Ellis, 1912:481. 


255 


Type species. —Cambarincola macrodon- 
ta Ellis, 1912, by original designation. 
Gender. — Masculine. 


Cambarincola acudentatus Holt, 1973 
Figs. 2—5 


Cambarincola acudentatus Holt, 1973b:11- 
iS. 


Types. — Holotype and 1 paratype, USNM 
45435, 6 paratypes, PCH 489, 4 paratypes, 
IBUM, on the isopods Speocirolana bolivari 
and S. pelaezi, from Grutas de Quintero. 11 
km SW of Ciudad Monte, Tanaulipas, Mé- 
xico, 14 May 1950, by Alejandro Villalo- 
bos. 

Distribution. —Known only from the type 
locality. 


Cambarincola alienus Holt, 1963 
Figs. 6-10 


Cambarincola aliena Holt, 1963:97—100. 
Cambarincola alienus. —Holt, 1973c:10. 


Types. —Holotype, USNM 30415, 7 
paratypes, USNMI 30416, 7 paratypes, PCH 
1325, on the isopod Asellus alabamensis 
(Stafford), from Wet Cave, Franklin Co., 
Tennessee, by T. C. Barr [date lost]. 

Distribution. —Known only from the type 
locality. 

Notes: Cambarincola steevesi Holt, 1973c, 
may be conspecific with or a local race of 
C. aliena. 


Cambarincola barbarae Holt, 1981 
Figs. 11-14 


Cambarincola barbarae Holt, 1981a:677— 
679. 


Types. —Holotype, USNM 54639, 5 
paratypes, PCH 1101, on Procambarus 
(Scapulicambarus) clarkii (Girard), from ir- 
rigation ditch 3.1 mi E of Salvang, Santa 
Barbara Co., California, by Perry C. and 
Virgie F. Holt, 3 Jul 1960. 

Distribution. —Santa Barbara and Sono- 
ma Counties, California. 

Notes: “Cambarincola barbarae occurs 


254 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


C. acudentatus 


; 
; C. alienus 
: ; 11 
C. barbarae 


a 


C. bobbi 


es 


C. branchiophilus 20 sep 22 


Figs. 2-22. 2-5, Cambarincola acudentatus Holt, 2, ventro-lateral view; 6-10, Cambarincola alienus Holt; 
11-14, Cambarincola barbarae Holt; 15-18, Cambarincola bobbi Holt; 19-22, Cambarincola branchiophilus 
Holt. 


VOLUME 106, NUMBER 2 


upon an introduced species (Hobbs, 1972: 
72) which also serves as a host for C. me- 
sochoreus Hoffman, 1963, and C. fallax 
Hoffman, 1963. The former species is wide- 
spread in the Mississippi Valley and the lat- 
ter in the Appalachians and were possibly 
introduced with their host. This may be so 
for C. barbarae”’ (Holt 1981a:679). 


Cambarincola bobbi Holt, 1988 
Figs. 15-18 


Cambarincola bobbi Holt, 1988b:794-808. 


Types. —Holotype, USNM 101496, 4 
paratypes, USNM 101497-101499, on 
Cambarus bartonii bartonii (Fabricius), from 
a medium-sized stream in Tom’s Brook (a 
town) 5.7 mi S of Strasburg, Shenandoah 
Co., Virginia, by Marvin L. Bobb and Perry 
C. Holt, 22 Jul 1948. 

Distribution. —Known only from the type 
locality. 


Cambarincola branchiophilus Holt, 1954 
Figs. 19-22 


Cambarincola branchiophila Holt, 1954: 
168-172.—Hoffman, 1963:317-319. 
Cambarincola branchiophilus. —Holt, 

1973b:10. 


Types. —Holotype, USNM 25855, 6 
paratypes, PCH 407 USNM, on Cambarus 
bartonii bartonii (Fabricius), and C. sci- 
otensis Rhoades, from Sinking Creek at 
crossing of State Highway 700, Giles Co., 
Virginia, by Frank D. Kiser, Cornelia Tuten 
and Perry C. Holt, 3 Jul 1950. 

Distribution. — Known only from the type 
locality. 


Cambarincola carcinophilus Holt, 1973 
Figs. 23-28 


Cambarincola carcinophilus Holt, 1973b: 
13-14. 


Types. —Holotype, USNM 45439, 1 
paratype, IBUM, 1 paratype, PCH 698, on 


255 


the freshwater crab Pseudothelophusa ve- 
racruzana, from Rio Tapalapa, Santiago, 
Tuxtla, México, by Alejandro Villalobos and 
Horton H. Hobbs, Jr., 1957. 

Distribution. —Two locations in the low- 
lands of Southern Veracruz. 

Notes: The specimens taken in 1957 by 
Villalobos and Hobbs from the Arroyo de 
Zapoapan de Cabana were associated with 
not only the crab P. veracruzana, but also 
with the crayfish Procambarus zapoapensis 
Villalobos. 


Cambarincola chirocephala Ellis, 1919 
Figs. 29-34 


Cambarincola chirocephala Ellis, 1919:263-— 
264.—Goodnight, 1940:37-38.—Holt & 
Hoffman, 1959:103.—Hoffman, 1963: 
348-351. 

Cambarincola chirocephalus. — Holt, 
1973b:9 [unjustified emendation]. 


Types. —Holotype, USNM 17713, on Or- 
conectes virilis Hagen, from Rolla, Phelps 
Co., Missouri, by J. Barley, date unknown. 

Distribution. —From western Tennessee 
north to Indiana, westward to Oklahoma 
and Kansas. 

Notes: Cambarincola chirocephala ap- 
pears to intergrade with C. philadelphicus 
in western Tennessee (Hoffman 1963:345). 


Cambarincola demissus Hoffman, 1963 
Figs. 35-36 


Cambarincola demissa Hoffman, 1963:365— 
367. 
Cambarincola demissus. — Holt, 1973b:10. 


Types. —Holotype and 4 paratypes, 
USNM 29948, on Orconectes erichsonianus 
Faxon, and O. rusticus (Girard), from the 
Powell River at Big Stone. Gap, Wise Co., 
Virginia, by H. H. Hobbs, Jr. and C. W. 
Hart, Jr., 17 Jun 1950. 

Distribution. —Mountain regions of 
southwestern Virginia. 


Cambarincola dubius Holt, 1973 
Figs. 37-42 


Cambarincola dubius Holt, 1973c:234—236. 


Types. —Holotype, USNM 49673, 2 
paratypes, USNM 49674, 4 paratypes, PCH 
2763, on Cambarus laevis Faxon, and Or- 
conectes inermis testii (Hay), from May’s 
Cave, Monroe Co., Indiana, by Horton H. 
Hobbs, III, 20 Sep 1969. 

Distribution. —Caves in Monroe Co., In- 
diana. 


Cambarincola ellisi Holt, 1973 
Figs. 43-48 


Cambarincola ellisi Holt, 1973b:14—-16. 


Types. —Holotype, USNM 45433, on 
Procambarus regiomontanus Villalobos, 
from Rio San Juan, San Juan, Nuevo Leoni, 
Mexico, by Salvador Contreras, 18 Apr 
1964. One paratype, PCH 1844, on P. re- 
giomontanus, from Arroyo de la Cruz, km 
245 de la carretera Ciudad México-Mon- 
terey, by A. Villalobos, 14 Feb 1964. 

Distribution. —Known only from the type 
locality. 

Notes: Cambarincola ellisiis known from 
a tributary on the Rio Grande (Rio Bravo) 
and is geographically nearer the branchiob- 
dellidans of the United States than of Mex- 
ico. 


Cambarincola fallax Hoffman, 1963 
Figs. 49-54 


Cambarincola fallax Hoffman, 1963:256- 
259.—Hobbs et al., 1967:54—55.— Holt, 
1969:207; 1973c:238; 1981a:679-680. 


Types. —Holotype and 4 paratypes, 
USNM 29945, additional paratypes, PCH 
904, on ““Cambarus longulus subsp.”’ [either 
C. longirostris Faxon, or C. longulus Gi- 
rard], from Maiden Spring Creek, Tazewell 
Co., Virginia, by Richard L. Hoffman, 19 
Jun 1959. 

Distribution. —Appalachian Mountains 
from Canada to Georgia; one specimen from 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Sonoma Co., California. In the southern 
parts of its range this species is confined to 
higher elevations. 

Notes: The specimen from California rep- 
resents either an introduction, or less likely 
a new, but closely related species. 


Cambarincola floridanus Goodnight, 1941 
Figs. 55—60 


Cambarincola floridanus Goodnight, 1941: 
73-74.—Hoffman, 1963:368.—Holt, 
1973a:90-93. 


Types. —Holotype, USNM 20570, on 
Procambarus fallax (Hagen), collected 6.8 
mi S of Lamont, Taylor Co., Florida, by 
Horton H. Hobbs, Jr., 18 Mar 1939. 

Distribution. —The type locality and one 
locality in Liberty Co., Florida. 

Notes: In the paragraph beginning “‘The 
spermiducal gland ...”’ (Holt 1973a:92) a 
lapsus resulted in “‘ectally” being written 
instead of the obviously correct “‘entally.” 


Cambarincola goodnighti Holt, 1973 
Figs. 61-66 


Cambarincola goodnighti Holt, 1973a:88- 
90. 


Types. —Holotype, USNM 49706, 1 
paratype, USNM 48707, 1 paratype, PCH 
2607, on Procambarus fallax (Hagen), and 
P. paeninsulanus (Faxon), collected 5 mi N 
of St. Augustine, St. Johns Co., Florida 
(Collector and date unknown). 

Distribution. —Known only from the type 
locality. 


Cambarincola gracilis Robinson, 1954 
Figs. 67—70 


Cambarincola gracilis Robinson, 1954:467— 
468.—Holt & Hoffman, 1959:97-103.— 
Hoffman, 1963:369.—Holt, 1981a:680- 
685.—Gelder & Hall, 1990:2354. 


Types. —Holotype and 10 paratypes, 
USNM 26110, on Pacifastacus leniusculus 
kalamathensis (Stimpson), from creek on 


VOLUME 106, NUMBER 2 257 


On 


C. carcinophilus 


dr. 
SR 


C. chirocephala 


(Sue 


C. demissus 


(> 


C. dubius 


43 


Be 


C. ellisi 44 eA 46 4 


Figs. 23-48. 23-28, Cambarincola carcinophilus Holt; 29-34, Cambarincola chirocephala Ellis; 35-36, Cam- 
barincola demissus Hoffman; 37-42, Cambarincola dubius Holt; 43-48, Cambarincola ellisi Holt. 


258 


campus of Whitman College, Walla Walla, 
Walla Walla Co., Washington, by A. G. 
Rempel, 11 Aug 1952; 2 paratypes, USNM 
26111, 0n P. 1. klamathensis, from Klamath 
River, 1 mi W of Ash Creek, Siskiyou Co., 
California, by Harold Wolf, 27 Oct 1952; 1 
paratype, USNM 26112, on P. /. trowbridgii 
(Stimpson), from Burdette Creek, Burnaby, 
British Columbia, by G. Clifford Carl, 21 
May 1942. 

Distribution. —From southern California 
to southern British Columbia in streams of 
the Pacific versant. 

Notes: Holt’s statement (Holt 1981a:682) 
that the prostate is subequal in length and 
diameter to the spermiducal gland is an in- 
explicable error (see Holt 1981a:681, fig. 2). 
Cambarincola gracilis is the second most 
common species of branchiobdellidan in 
collections from the Pacific drainage in the 
United States and Canada. 


Cambarincola heterognathus Hoffman, 
1963 
Figs. 71-76 


Cambarincola heterognatha Hoffman, 1963: 
362-365. 

Cambarincola heterognathus. — Holt, 1973a: 
95; 1973b:10. 


Types. — Holotype and 1 paratype, USNM 
29947, on Cambarus sp., from a tributary 
to Big Wilson Creek, 4 mi N of Mouth of 
Wilson, on State Highway 16, Grayson Co., 
Virginia, by Horton H. Hobbs, Jr., and C. 
W. Hart, 14 Jun 1950. 

Distribution.—From northwestern Vir- 
ginia and adjacent West Virginia south and 
west to the central eastern parts of Kentucky 
and Tennessee; disjunct populations in Leon 
and Calhoun Counties, Florida. 


Cambarincola hoffmani Holt, 1973 
Figs. 77-82 
Cambarincola hoffmani Holt, 1973b:16—17. 


Types. —Holotype, USNM 45447, on 
Procambarus hoffmanni (Villalobos), from 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Arroyo de Tlatentilojan, at Los Estajos, 6 
km NE of Zihuateutla, Puebla, México, by 
Alejandro Villalobos, 11 Nov 1949; 2 para- 
types, PCH 1622, on Procambarus cabal- 
leroi Villalobos, from Villa Juarez, Puebla, 
México, by Alejandro Villalobos, May 1944. 

Distribution. —Known only from the type 
locality. 

Notes: Cambarincola hoffmani may be an 
inhabitant of the gill chambers of its hosts 
(Holt 1973b:17). The spelling is correct, as 
the species was named from R. L. Hoffman, 
not for the host crayfish. 


Cambarincola holostoma Hoffman, 1963 
Figs. 83-87 


Cambarincola holostoma Hoffman, 1963: 
359-361. 

Cambarincola holostomus. — Holt, 1973b:10 
[unjustified emendation]. 


Types. —Holotype and 4 paratypes, 
USNM 29946, 2 paratypes, PCH 599, on 
““Cambarus bartonii’ [probably C. b. ca- 
vatus Hay] and C. longulus Girard, from 
Crab Run, Highland Co., Virginia, by L. B. 
Holthuis, 25 Oct 1952. 

Distribution.— Western Virginia in the 
headwaters of the Potomac and James Riv- 
ers (Hobbs et al. 1967:57). 


Cambarincola holti Hoffman, 1963 
Figs. 88-89 


Cambarincola holti Hoffman, 1963:314-— 
316. 


Types. — Holotype and paratypes, USNM 
29940, on Cambarus sp., from a stream in 
the southern part of Somerset, Pulaski Co., 
Kentucky, by Perry C. and Virgie F. Holt, 
28 Jul 1958. 

Distribution. —Known only from the type 
locality. 


Cambarincola illinoisensis Holt, 1982 
Figs. 90-93 


Cambarincola illinoisensis Holt, 1982:251- 
259: 


VOLUME 106, NUMBER 2 259 


49 


C. fallax 


a 


C. floridanus 


(am 


C. goodnighti 


Sao 


C. gracilis 


Figs. 49-76. 49-54, Cambarincola fallax Hoffman; 55-60, Cambarincola floridanus Goodnight; 61-66, 
Cambarincola goodnighti Holt; 67-70, Cambarincola gracilis Robinson; 71-76, Cambarincola heterognathus 
Hoffman. 


260 


Types. —Holotype, USNM 65225, 3 
paratypes, USNM 65226, 5 paratypes, PCH 
840, on Orconectes virilis Hagen, from a 
prairie stream (possibly one of the two Sugar 
or two Mud Creeks that drain into the Ir- 
oquois River) N of Stockland, Iroquois Co., 
Illinois, by Perry C. and Virgie F. Holt, 25 
Jul 1958. 

Distribution. —Known only from the type 
locality. 

Notes: The protruded penes of C. i/linois- 
ensis are similar in some respects to those 
of species of Sathodrilus, but there are not 
enough data now to unite these genera (see 
Holt 1982:254—255). 


Cambarincola ingens Hoffman, 1963 
Figs. 94-97 


Cambarincola ingens Hoffman, 1963:333- 
336. 


Types. —Holotype and 2 paratypes, 
USNM 29944, on Cambarus §sciotensis 
Rhoades, from Sinking Creek, Giles Co., 
Virginia, by Ben I. Johns, 27 Jun 1953. 
Topotypes, taken by others, PCH 234, 407, 
499. 

Distribution. —The southern Appala- 
chians in North Carolina, Tennessee, Vir- 
ginia, and West Virginia. 


Cambarincola jamapaensis Holt, 1973 
Figs. 98-103 


Cambarincola jamapaensis Holt, 1973b:17— 
ZO: 


Types. —Holotype and 5 immature para- 
types, USNM 45438, 1 paratype, PCH 1592, 
on Procambarus mexicanus (Erichson), from 
the Rio Jamapa, 7 km NE of Coscomatepec, 
Veracruz, México, by Perry C. and Virgie 
F. Holt, 9 Jul 1962. 

Distribution. —Known with certainty only 
from the type locality; specimens from two 
localities in Puebla, México, have tenta- 
tively been assigned to this species (Holt 
1973b:20). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Cambarincola leoni Holt, 1973 
Figs. 104-106 


Cambarincola leoni Holt, 1973c:226-229. 


Types. —Holotype, USNM 49676, 1 
paratype, USNM 40677, on Procambarus 
orcinus Hobbs & Means, from Gopher Sink, 
Leon Co., Florida, by D. Bruce Means and 
Joseph Halusky, 3 Apr 1971; 5 paratypes, 
PCH 2756, on P. orcinus, from Gopher Sink, 
by D. Bruce Means and J. F. Berry, 26 Feb 
1971. 

Distribution. —Caves in Leon, Alachua 
and Marion Counties, Florida. 


Cambarincola leptadenus Holt, 1973 
Figs. 107-111 


Cambarincola leptadenus Holt, 1973¢:231- 
234. 


Types. —Holotype, USNM 49678, 1 
paratype, USNM 49679, 2 paratypes, PCH 
2739, on Cambarus tenebrosus Hay, from 
Bethel Cave, Perry Co., Tennessee, by Hor- 
ton H. Hobbs, III, 6 Oct 1968. 

Distribution. —Known only from the type 
locality. 


Cambarincola macrocephala Goodnight, 
1943 
Figs. 112-116 


Cambarincola macrocephela (inadvertent 
misspelling) Goodnight, 1943:100-101. 

Cambarincola macrocephala. —Holt & 
Hoffman, 1959:103.—Hoffman, 1963: 
312-314. 

Cambarincola  macrocephelus. —Holt, 
1973b:10 [unjustified emendation]; Holt, 
1981a:685, fig. 3. 

Cambarincola macrocephalus. —Holt, 
1981a:685-689. 


Types. —Holotype, USNM 20598, on Pa- 
cifastacus gambelii (Girard), from Polecat 
Creek, Tenton Co., Wyoming, by Robert C. 
Brown, 16 Aug 1941. 

Distribution. —The upper reaches of the 


VOLUME 106, NUMBER 2 261 


a 


C. hoffmani 


wl 
Sane 


© ® 


C. holostoma 


88 Not 
C. holti 


a 


C. illinoisensis 


| 94 a 
C. ingens 97 < 


Figs. 77-97. 77-82, Cambarincola hoffmani Holt; 83-87, Cambarincola holostoma Hoffman; 88-89, Cam- 
barincola holti Hoffman; 90-93, Cambarincola illinoisensis Holt; 94-97, Cambarincola ingens Hoffman. 


262 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


100 102 


ee ee 1 Bas 


98 


C. jamapaensis 


104 


C. leoni 106 


<4 (|| a 


109 110 
107 
111 


116 


117 ae 119 121 
C. macrodontus 118 120 499 


Figs. 98-122. 98-103, Cambarincola jamapaensis Holt; 104-106, Cambarincola leoni Holt; 107-111, Cam- 
barincola leptadenus Holt; 112-116, Cambarincola macrocephala Goodnight; 117-122, Cambarincola macro- 
dontus Ellis. 


VOLUME 106, NUMBER 2 


Snake River in Idaho and Wyoming (Holt 
1981a:688). 


Cambarincola macrodontus Ellis, 1912 
Figs. 117-112 


Cambarincola macrodonta Ellis, 1912:481- 
484; 1919:257.—Hall, 1914:190.— 
Goodnight, 1940:31.—Holt & Hoffman, 
1959:97.—Hoffman, 1963:352. 

Cambarincola macrodontus. — Holt, 
1973b:9. 


Types. —Holotype and 2 paratypes, 
USNM 53794, on Cambarus diogenes Gi- 
rard, from a stream in Boulder, Boulder Co., 
Colorado, by Max M. Ellis, date unknown. 

Distribution. — The high plains of the cen- 
tral United States, doubtfully from Las Ve- 
gas, Nevada (see Hoffman 1963:353-354). 


Cambarincola manni Holt, 1973 
Figs. 123-128 


Cambarincola manni Holt, 1973a:85-88. 


Types. —Holotype, 9 paratypes, USNM 
48700, on Procambarus fallax (Hagen), from 
Lake Martha, at 612 E. Lake Martha Drive, 
Winter Haven, Polk Co., Florida, by Ches- 
ter A. Mann, 12 Jan 1964; 1 paratype, 
USNM 48701, on P. fallax, from canal be- 
tween Lake Buckeye and Lake Fanny, Win- 
ter Haven, Polk Co., Florida, by Chester A. 
Mann and Perry C. Holt, 20 Apr 1963; 1 
paratype, PCH 1670, on P. fallax, from Lake 
Martha, Winter Haven, Polk Co., Florida, 
by Chester A. Mann and Perry C. Holt, 20 
Apr 1963; 10 paratypes, PCH 1793, from 
the type locality; 5 paratypes, PCH 1673, 
on Procambarus paeninsulanus (Faxon), 
from slough, 13 mi NW of Inglis, Levy Co.., 
Florida, by Perry C. and Virgie F. Holt, 26 
Apr 1963. 

Distribution. — Florida peninsular. 

Notes: Cambarincola manni appears to 
be confined to peninsular Florida and is the 
only branchiobdellidan from the southern 
portion of the state, but overlaps C. osceolai 


263 


in the northern part of the state (see Holt 
1973a:88, fig. 1). 


Cambarincola marthae Holt, 1973 
Figs. 129-134 


Cambarincola marthae Holt, 1973c:221- 
224. 


Types. —Holotype, USNM 49509, 1 
paratype, PCH 2767, on the isopod Asellus 
alabamensis (Stafford), from Carter’s Cave, 
Jackson Co., Tennessee, by John E. and 
Martha R. Cooper, 21 Sep 1968. 

Distribution. —Known only from the type 
locality. 


Cambarincola mesochoreus Hoffman, 1963 
Figs. 135-137 


Cambarincola vitrea (in part). — Ellis, 1919: 
257-258. 

Cambarincola macrodonta (in part). — Ellis, 
1919:257. 

Cambarincola mesochorea Hoffman, 1963: 
307-311. 

Cambarincola mesochoreus. — Holt, 1973b: 
10; 1981a:689. 


Types. —Holotype and 4 paratypes, 
USNM 29934, additional paratypes, PCH 
815, on Orconectes sp., from stream 1.5 mi 
E of Adyville, Perry Co., Indiana, by Perry 
C. and Virgie F. Holt, 28 Jul 1958. 

Distribution. —Central Mississippi drain- 
age with disjunct populations in Massachu- 
setts and California (Hoffman 1963:308, 
Holt 1981a:689). 

Notes: In Holt, 1981a:689, the last entry 
in the synonymy should be “Cambarincola 
mesochoreus. — Holt, 1973b:10” instead of 
“1963:10.” 


Cambarincola meyeri Goodnight, 1942 
Figs. 138-143 


Cambarincola meyeri Goodnight, 1942:272- 
273.— Hoffman, 1963:354.— Holt, 1973d: 
678-682. 


264 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


23 Bi Ie) SUE 


127 


123 
C. manni 


129 


C. marthae 


135 


C. mesochoreus 


138 es 


(\ C. meyer 


144 


C. micradenus 145 JC. 


Figs. 123-149. 123-128, Cambarincola manni Holt; 129-134, Cambarincola marthae Holt; 135-137, Cam- 
barincola mesochoreus Hoffman, 137, dorsal view of reproductive system, showing recurved prostate; 138-143, 
Cambarincola meyeri Goodnight; 144-149, Cambarincola micradenus Holt. 


VOLUME 106, NUMBER 2 


Types. —Holotype, USNM 20597, on 
““Cambarus bartonii’’ [probably C. b. bar- 
tonii (Fabricius)], from Raven’s Creek, Fay- 
ette Co., Kentucky, by Marvin C. Meyer 
(date unknown). 

Distribution. —Raven Run (=“‘Raven’s 
Creek’’), Fayette Co., Kentucky (Holt 1973d: 
681). 


Cambarincola micradenus Holt, 1973 
Figs. 144-149 


Cambarincola micradenus Holt, 1973b:20- 
22. 


Types. —Holotype and 2 paratypes, 
USNM 45448, 1 paratype, PCH 1615, on 
Procambarus paradoxus (Ortmann), from 
La Canada y Tetela de Ocampo, Puebla, 
México, by Alejandro Villalobos, May 1944. 

Distribution. —Known only from the type 
locality. 


Cambarincola montanus (Goodnight), 1940 
Figs. 150-153 


Triannulata montana Goodnight 1940:57- 
58. 

Cambarincola montanus. — Holt, 1974:67- 
70; 1981a:690-691. 


Types. — Holotype, USNM 2056, on Pa- 
cifastacus sp., from the Kalami River, 
Washington (Collector and date unknown). 
Paratypes were left at the University of Il- 
linois in the collections of H. J. Van Cleave 
(Goodnight 1940:58). 

Distribution. —Streams of the Coastal and 
Cascade Ranges of the Pacific drainage in 
western North America from Santa Barbara 
Co., California to northern Washington 
(Holt 1981a:690). 


Cambarincola nanognathus Holt, 1973 
Figs. 154-159 


Cambarincola nanognathus Holt, 1973b: 
22-23. 


Types. —Holotype, USNM 45444, 1 
paratype, PCH 1830, 1 paratype, IBUM, on 


265 


the freshwater crab Potamocarcinus nica- 
raguensis, from Lago de Nicaragua (Isleta 
de Granada), G. Alvilez, 13 Jul 1964. 

Distribution.—From southern Veracruz, 
México, to Nicaragua. 


Cambarincola olmecus Holt, 1973 
Figs. 160-163 


Cambarincola olmecus Holt, 1973b:24—26. 


Types. —Holotype, USNM 45445, 1 
paratype, USNM 45446, 1 paratype, PCH 
201, 1 paratype, IBUM, on Procambarus 
mexicanus (Erichson), from Tomatlan, Ve- 
racruz, México, by Alejandro Villalobos, 3 
Nov 1948. 

Distribution.—San Luis Potosi and Ve- 
racruz, México. 


Cambarincola osceolai Hoffman, 1963 
Figs. 164-165 


Cambarincola osceola Hoffman, 1963:330- 
333. 

Cambarincola osceolai.—Holt, 1973a:93- 
95; 1973b:10. 


Types. —Holotype and 7 paratypes, 
USNM 29943, on Procambarus paeninsu- 
lanus (Faxon) and Fallicambarus uhleri 
(Faxon) from Dry Creek, 3.1 mi N of Iron 
City, Seminole Co., Georgia, by Horton H. 
Hobbs, Jr. and C. W. Hart, Jr., 9 Sep 1955. 

Distribution. —From southeastern Vir- 
ginia to northwest peninsular Florida. 

Notes: Attention may again be called to 
the suspicions of both Hoffman (1963:331) 
and Holt (1973a:94) that C. osceolai and C. 
vitreus Ellis, 1918:51 are conspecific. 


Cambarincola ouachita Hoffman, 1963 
Figs. 166-167 


Cambarincola ouachita Hoffman, 1963: 
303-305. 


Types. —Holotype, USNM 29937, on Or- 
conectes sp., from a small stream 4.3 mi W 
of the Montgomery Co. line in Chautaugua 


266 


Co., Kansas, 8 Jul 1958, by Perry C. and 
Virgie F. Holt. 

Distribution. —Known only from the type 
locality. 

Notes: Holt (1973b:9-10) changed the 
endings of several species to conform to the 
masculine gender of the generic name Cam- 
barincola. He did not emend the name C. 
ouachita which may be considered a noun 
in apposition, referring to the Ouachita 
Mountains. Hoffman gives no derivation for 
the name ouachita, but Holt knows he meant 
it to refer to the mountains. 


Cambarincola pamelae Holt, 1984 
Figs. 168-171 


Cambarincola pamelae Holt, 1984b:544- 
549. 


Types. —Holotype, USNM 080687, 4 
paratypes, USNM 080688-080691, 10 
paratypes, PCH 4065, on Procambarus 
clarkii (Girard), from an irrigation canal that 
drains into the San Joaquin River in the 
western part of Stanislaus Co., California, 
by J. A. Meeuwse, 2 Dec 1982. 

Distribution. —Stanislaus, Santa Barbara, 
Merced and Sonoma Counties, California. 

Notes: Cambarincola pamelae has been 
found only on P. clarkii, a crayfish species 
that has been widely introduced throughout 
the United States (Hobbs 1989). Therefore, 
it is possible that C. pamelae has also been 
introduced into its present range and that it 
may be a geographical variant of C. meso- 
choreus, to which it is similar (Holt 1984b). 


Cambarincola philadelphicus (Leidy, 1851) 
Figs. 172-177 


Astacobdella philadelphica Leidy, 1851:209. 
Branchiobdella philadelphica. —Moore, 
1893:427-428. 

Bdellodrilus philadelphicus. —Moore, 1895: 
497; 1901:542.—Pierantoni, 1912:17. 
Cambarincola philadelphica. —Ellis, 1912: 
484.—Hall, 1914:190.—Ellis, 1918:49; 
1919:260-263.—Goodnight, 1939:11; 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


1940:38.—Holt, 1954:169.—Holt & 
Hoffman, 1959:103.—Hoffman, 1963: 
341-348. | 
Cambarincola 
1973b:9. 


Dhiladelphicus. — Holt, 


Types. — Holotype, lost. Topotypes, PCH 
695, from Wissahicon Creek in Philadel- 
phia, Pennsylvania (Hoffman 1963:342). 

Distribution. —From Minnesota and New 
York south through the Appalachians and 
Piedmont to South Carolina (Hoffman 1963: 
344). 

Notes: This species is reported in the lit- 
erature as unusually variable. Indeed, in 
southwestern Virginia it appears to be so, 
but many references to it undoubtedly apply 
to other related forms. The problem of vari- 
ability in C. philadelphicus should be re- 
studied. 


Cambarincola restans Hoffman, 1963 
Figs. 178-179 


Cambarincola restans Hoffman, 1963:305— 
307% 


Types. —Holotype and 2 paratypes, 
USNM 29938, on Orconectes sp., from Sug- 
ar Creek, 2 mi N of Avoca, Benton Co., 
Arkansas, by Perry C. and Virgie F. Holt, 
6 Jul 1958. 

Distribution. —Known only from the type 
locality. 


Cambarincola serratus Holt, 1981 
Figs. 180-183 


Cambarincola serratus Holt, 1981a:691-— 
693. 


Types. —Holotype, USNM 54638; 3 
paratypes, PCH 795, on Pacifastacus con- 
nectens (Faxon), from Idaho State Fish 
Hatchery, Riley Creek, Gooding Co., Idaho, 
by Perry C. and Virgie F. Holt, 14 Jul 1958; 
2 paratypes, PCH 784, on P. connectens, 
from spring tributary to Snake River, Ha- 
german, Gooding Co., Idaho, by Perry C. 
and Virgie F. Holt, 14 Jul 1958. 


VOLUME 106, NUMBER 2 267 


fan 


C. montanus SS 
156 158 
2) &> 
159 
154 = 


C. nanognathus 


160 
C. olmecus 


C. osceolai 


C 


. ouachita 167 FP Le 


Figs. 150-167. 150-153, Cambarincola montanus Goodnight; 154-159, Cambarincola nanognathus Holt; 
160-163, Cambarincola olmecus Holt; 164-165, Cambarincola osceolai Hoffman; 166-167, Cambarincola 
ouachita Hoffman. 


268 


Distribution. —Tributaries of the Snake 
River, Gooding Co., Idaho. 


Cambarincola sheltensis Holt, 1973 
Figs. 184-189 


Cambarincola sheltensis Holt, 1973c:229- 
230. 


Types. —Holotype, USNM 49683, 2 
paratypes, PCH 1846, on Orconectes aus- 
tralis australis (Rhoades), from Shelta Cave, 
Huntsville, Madison Co., Alabama, by John 
E. and Martha Cooper, 24 Aug 1963; 2 para- 
types, USNM 49684, 1 paratype, PCH 1863, 
on O. a. austraiis, from Shelta Cave, Mad- 
ison Co., Alabama, by James E. Larimer, 
1965. 

Distribution. —Known only from the type 
locality. 


Cambarincola shoshone Hoffman, 1963 
Figs. 190-191 


Cambarincola shoshone Hoffman, 1963: 


319-320.— Holt, 1981a:693. 


Types. —Holotype and 3 paratypes, 
USNM 29941, topotypes, PCH 785, on Pa- 
Cifastacus connectens (Faxon), from Riley 
Creek (Idaho State Fish Hatchery), Hager- 
man, Gooding Co., Idaho, by Perry C. and 
Virgie F. Holt, 14 Jul 1958. 

Distribution. —Known only from the type 
locality. 


Cambarincola smalleyi Holt, 1964 
Figs. 192-195 


Cambarincola smalleyi Holt, 1964:1-4; 
1973b:26—27. 


Types. —Holotype, USNM 20940, 1 
paratype, 30941, 1 paratype in the collec- 
tions of Tulane University, 1 paratype, PCH 
1702, on the freshwater crab Pseudothel- 
phusa tuminenus, from Rio Hondura, 8 mi 
N of San Jeronimo de Moravia, San José 
Province, Costa Rica, by A. E. Smalley, 9 
Jul 1962. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Distribution. —Known only from the type 
locality. 

Notes: Holt (1964:3) postulated that C. 
smalleyi was carried to Costa Rica by cam- 
barines and qualified (Holt 1973b:27) this 
conclusion by suggesting that the transfer to 
crabs could have occurred much further 
north in México. In truth we know too little 
to make any credible conjectures about the 
ancient origin and subsequent wanderings 
of branchiobdellidans. They may as easily 
have originated as symbionts of freshwater 
crabs and shrimps and later taken up a sym- 
biotic relationship with crayfishes. Cam- 
barincola smalleyi is the southernmost 
known species of the family. 


Cambarincola speocirolanae Holt, 1984 
Figs. 196-199 


Cambarincola speocirolanae Holt, 1984a: 
36-38. 


Types. —Holotype, USNM 80221, 1 
paratype, PCH 4054, on the isopod Speo- 
cirolana palaezi, from Sotao del Arroyo, San 
Luis Potosi, México, by Peter Sprouse, 22 
Feb 1980. 

Distribution. —Known only from the type 
locality. 


Cambarincola steevesi Holt, 1973 
Figs. 200-203 


Cambarincola steevesi Holt, 1973¢c:224—226. 


Types. —Holotype, USNM 40680, 2 
paratypes, USNM 40681, 1 paratype, PCH 
1883, on the isopod Asellus alabamensis 
(Stafford), from Glover’s Cave, Todd Co., 
Kentucky, by R. M. Norton, 17 Apr 1964; 
1 paratype, USNM 49682, on Asellus ala- 
bamensis, from Brown Cave, Barren Co., 
Kentucky, by R. M. Norton, 25 Sep 1965. 

Distribution. —The type locality and Tay- 
lor Cave, Trigg Co., Kentucky. 


Cambarincola susanae Holt, 1973 
Figs. 204—207 


Cambarincola susanae Holt, 1973b:27-29. 


VOLUME 106, NUMBER 2 269 


168 aly 
169 : 
C. pamelae | 


172 
} 173 
C. philadelphicus 


| 178 ) 
179 
C. restans 
‘ | 180 


C. serratus 


184 


C. sheltensis 


Figs. 168-189. 168-171, Cambarincola pamelae Holt; 172-177, Cambarincola philadelphicus (Leidy); 178- 
179, Cambarincola restans Hoffman; 180-183, Cambarincola serratus Holt; 184-189, Cambarincola sheltensis 
Holt. 


270 


Types. —Holotype and 2 paratypes, 
USNM 45441, 10 paratypes, PCH 1529, 3 
paratypes, IBUM, on Procambarus acutus 
cuevachicae (Hobbs), from Cueva Chica, El 
Pujal, 3 km NE of Valles, San Luis Potosi, 
México, by Alejandro Villalobos, 9 May 
1950. 

Distribution.—In eastern México from 
Rio San Juan, Nuevo Leon to western Cam- 
peche. 

Notes: Cambarincola susanae may be an 
ectoparasite in the gill chambers of the host 
(Holt 1973b:29). 


Cambarincola toltecus Holt, 1973 
Figs. 208-211 


Cambarincola toltecus Holt, 1973b:29-31. 


Types. —Holotype, USNM 45436, 2 
paratypes, USNM 45437, 2 paratypes, 
IBUM, 2 paratypes, PCH 697, on the fresh- 
water crab Pseudothelphusa veracruzana, 
from Rio Tapalapa, Santiago, Tuxtla, Ve- 
racruz, México, by Alejandro Villalobos and 
Horton H. Hobbs, Jr., 17 Apr 1957. 

Distribution. —Tropical lowlands of Ve- 
racruz, Mexico. 


Cambarincola virginicus Hoffman, 1963 
Figs. 212-215 


Cambarincola virginica Hoffman, 1963: 
322-323. 
Cambarincola virginicus. —Holt, 1973b:10. 


Types. — Holotype and 1 paratype, USNM 
29942, on Cambarus acuminatus Faxon, 
from a small, slow stream 4.7 mi N of Pe- 
tersburg, Chesterfield Co., Virginia, by Mar- 
vin L. Bobb and Perry C. Holt, 31 May 
1949. 

Distribution.—Along the Fall Line in 
eastern Virginia (Hoffman 1963:323) and 
eastern North Carolina. 


Cambarincola vitreus Ellis, 1918 
Figs. 216-217 


Cambarincola vitrea Ellis, 1918:49-51; 
1919:257-258.— Goodnight, 1940:33-34; 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


1943:100.—Holt & Hoffman, 
103.— Hoffman, 1963:324-329. 
Cambarincola vitreus. —Holt, 1973b:9. 


Types. —Holotype, USNM 17667, on Or- 
conectes immunis (Hagen), from Douglas 
Lake, Cheboygan Co., Michigan, by Max 
M. Ellis, Jul 1915. 

Distribution. — Western portions of the 
upper Mississippi valley from north and west 
of Arkansas. 

Notes: See Cambarincola osceola. 


1959: 


Ceratodrilus Hall, 1914 


Ceratodrilus Hall, 1914:190-191. 
Ceratodrilus Stephenson, 1930:901. 
Cirrodrilus Goodnight, 1940:63-—64 [in part]. 


Type species. —Ceratodrilus thysanoso- 
mus Hall, 1914, by original designation. 
Gender. — Masculine. 


Ceratodrilus ophiorhysis Holt, 1960 
Figs. 218-221 


Ceratodrilus orphiorhysis Holt, 1960a:60 
(incorrect spelling). 

Ceratodrilus ophiorhysis. —Holt, 1960a:61; 
1988a:308. 


Types. —Holotype, USNM 29910, 4 
paratypes, USNM 29911, 3 paratypes, PCH 
786, on Pacifastacus gambelii (Girard), from 
the Snake River between Buhl and Wendel, 
Gooding Co., Idaho, by Perry C. and Virgie 
F. Holt and Judson Ford, 14 Jul 1958. 

Distribution.—The Snake River and its 
tributaries in Idaho and Oregon. 


Ceratodrilus thysanosomus Hall, 1914 
Figs. 222-223 


Ceratodrilus thysanosomus Hall, 1914: 
191.—Stephenson, 1930:801.— Yama- 
guchi, 1932:367. 

Cirrodrilus thysanosomus. —Goodnight, 
1940:64—65 [in part]. 

Ceratodrilus thysanosomus. — Holt, 1960a: 
58-60. 


Types. —Holotype, USNM 17708, from 


VOLUME 106, NUMBER 2 27h 


190 


C. shoshone 


192 


(C) 193 


C. smalleyi 


196 


C. speocirolanae 197 


202 
200 xD 
203 
C. steevesi 
Ww ve 206 
208 = ee Se 
C. susanae 
207 


Figs. 190-207. 190-191, Cambarincola shoshone Hoffman; 192-195, Cambarincola smalleyi Holt; 196- 
199, Cambarincola speocirolanae Holt; 200-203, Cambarincola steevesi Holt; 204-207, Cambarincola susanae 
Holt. 


272 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


streams of Great Basin, Salt Lake City, Utah 
(no date). 

Distribution. —Streams of the Great Salt 
Lake Basin. 

Notes: The two species of Ceratodrilus 
differ only in the length of their dorsal ap- 
pendages (Holt 1960a); their distinctiveness 
should be reinvestigated. 


Ellisodrilus Holt, 1960 
Ellisodrilus Holt, 1960b:170. 


Type species. — Ellisodrilus clitellatus Holt, 
1960, by original designation. 
Gender. — Masculine. 


Ellisodrilus carronamus Holt, 1988 
Figs. 224-227 


Ellisodrilus carronamus, 1988b:796—798. 


Types. —Holotype, USNM 119539, 2 
paratypes, USNM 119540-119541, on Or- 
conectes sp., from Carr Creek, Overton Co.., 
Tennessee, ca. 3 miS of Livingston, on State 
Highway 42, by Perry C. and Virgie F. Holt, 
26 Jul 1961. 

Distribution. —Carr Creek and Roaring 
River, Overton Co., Tennessee. 

Notes: There appears to be a N-S gradient 
in Ellisodrilus species with E. carronamus 
the southern-most, EF. durbini the northern- 
most and E. clitellatus found in between. 
This occurs in part in areas scoured by the 
Pleistocene glaciation (Holt 1960b:171, 174; 
1988b:798). 


Ellisodrilus clitellatus Holt, 1960 
Figs. 228-231 


Ellisodrilus clitellatus Holt, 1960b:169-176. 


Types. —Holotype, USNM 29935, 1 
paratype, USNM 29936, 18 paratypes, PCH 
827, on Cambarus distans Rhoades, from 
stream 8.9 mi E of Columbia on Kentucky 
Highway 80, Adair Co., Kentucky, by Perry 
C. and Virgie F. Holt, 28 Jul 1958. 

Distribution. —Kentucky. 

Notes: The first paragraph on page 171 


(Holt 1960b) should be transposed to follow 
the diagnosis of the genus on page 170. 


Ellisodrilus durbini (Ellis, 1918) 
Figs. 232-234 


Pterodrilus durbini Ellis, 1918:49.—Ellis 
1919:254.—Goodnight 1940:61-62. 
Ellisodrilus durbini. —Holt, 1960b:173. 


Types. — Holotype, USNM 17655, on Or- 
conectes barrenensis Rhoades, from White 
River, at Irondale, Anderson Co., Indiana. 

Distribution.—From Anderson Co., In- 
diana, into Michigan (Ellis 1918:50). 


Magmatodrilus Holt, 1967 


Stephanodrilus Goodnight, 1940:55 [in 
part]. 
Magmatodrilus Holt, 1967b:3-4. 


Type species.—Magmatodrilus obscurus 
(Goodnight, 1940), by original designation. 
Gender. — Masculine. 


Magmatodrilus obscurus (Goodnight, 1940) 
Figs. 235-238 


Stephanodrilus obscurus Goodnight, 1940: 
55-56. 
Magmatodrus obscurus. — Holt, 1967b:4—5. 


Types. —Holotype, USNM 20568, on Pa- 
cifastacus nigrescens (Stimpson) (Good- 
night, 1940:55), (collector and date un- 
known), from Fall River, Shasta Co., 
California; 2 topotypes, USNM 45696, 11 
topotypes, PCH 1818, on unknown host, 
from the head of Fall River, Thousand 
Springs Ranch, Shasta Co., California, by 
Perry C. and Virgie F. Holt, Aug 1964. 

Distribution. —Known only from the type 
locality. 

Notes: Some ambiguity may exist as to 
the type locality. Holt was refused permis- 
sion to trap for crayfish in Fall River by the 
local game warden. The river is a deep, ditch- 
like stream that does not lend itself to col- 
lecting with a dipnet. After several futile 
attempts to do so, the Holts obtained per- 


VOLUME 106, NUMBER 2 273 


208 


C. toltecus 


212 


C. virginicus 


216 


C. vitreus 


218 
C. ophiorhysis 


293 STA Ee 


C. thysanosomus 


Figs. 208-223. 208-211, Cambarincola toltecus Holt; 212-215, Cambarincola virginicus Hoffman; 216-217, 
Cambarincola vitreus Ellis; 218-221, Ceratodrilus ophiorhysis Holt; 222-223, Ceratodrilus thysanosomus Hall. 


274 


mission to take a collection from the spring 
from which Fall River arises. The holotype 
and topotypes appear to be identical (Holt 
1967b:4—5). 


Oedipodrilus Holt, 1967 
Oedipodrilus Holt, 1967a:58. 


Type species. — Oedipodrilus oedipus Holt, 
1967a, by original designation. | 
Gender. — Masculine. 


Oedipodrilus anisognathus Holt, 1988 
Figs. 239-242 


Oedipodrilus anisognathus Holt, 1988b: 
798-800. 


Types. —Holotype, USNM 119534, 1 
paratype, USNM 119535, on Orconectes sp., 
from a small stream in Montgomery Bell 
State Park, Dickson Co., Tennessee, by Per- 
ry C. and Virgie F. Holt, 4 Jul 1958; 3 para- 
types, USNM 119536-119537, on Orco- 
nectes sp., from Carr Creek, ca. 3.0 mi S of 
Livingston, Overton Co., Tennessee, by 
Perry C. and Virgie F. Holt, 26 Jul 1961. 

Distribution. —Dickson and Overton 
Counties, Tennessee, the Central (Nash- 
ville) Basin and the eastern Highland Rim, 
respectively (Holt 1988b:800). 


Oedipodrilus cuetzalanae Holt, 1984 
Figs. 243-246 


Oedipodrilus cuetzalanae Holt, 1984a:38- 
41. 


Types. —Holotype, USNM 80223, 8 
paratypes, PCH 4050, on Procambarus 
cuetzalanae Hobbs, from Sima Zoquiapan, 
1.1 km N of Cuetzalan, Puebla, México, by 
L. Wilk, W. Hooper and M. Minton, 2 Jan 
1980. 

Distribution. —Known only from the type 
locality. 


Oedipodrilus macbaini (Holt, 1955) 
Figs. 247-248 


Cambarincola macbaini Holt, 1955:27-31. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Oedipodrilus macbaini.—Holt, 1969:205; 
1984a:39; 1988b:800-804. 


Types. —Holotype, USNM 25952, 6 
paratypes, PCH 134, on Orconectes sp., from 
Charles Creek, 8 mi W of Ashland, on State 
Highway 5, Boyd Co., Kentucky, by Rod- 
ney Macbain, Jul 1948. 

Distribution. —Known only from the type 
locality. 


Oedipodrilus oedipus Holt, 1967 
Figs. 249-252 


Oedipodrilus oedipus Holt, 1967a:58—60. 


Types. —Holotype, USNM 34086, 7 
paratypes, USNM 34087, 4 paratypes, PCH 
756, on Orconectes compressus (Faxon), 
collected 10.2 mi E of Waverly, Humphreys 
Co., by Perry C. and Virgie F. Holt, 5 Jul 
1958. 

Distribution. —Known only from the type 
locality. 


Pterodrilus Moore, 1894 
Pterodrilus Moore, 1894:449—450. 


Type species. —Pterodrilus alcicornus 
Moore, 1894, subsequent designation by 
Goodnight (1940). 

Gender. — Masculine. 


Pterodrilus alcicornus Moore 1894 
Figs. 253-254 


Pterodrilus alcicornus Moore, 1894:450- 
453.—Pierantoni, 1912:25.—Ellis, 1919: 
245.—Goodnight, 1940:58-60.—Holt, 
1968c:6-12. 


Types. —Apparently lost (Holt 1968c:6), 
but was on Cambarus acuminatus Faxon, 
from Johns River at Blowing Rock, Watau- 
ga Co., North Carolina, in the summer of 
1893, presumably by Moore himself (Moore 
1894:453). 

Distribution.—The mountain streams of 
the Southern Appalachians in Virginia, 
North Carolina, Tennessee, and West Vir- 


VOLUME 106, NUMBER 2 DTS 


224  € 


E. carronamus 


ipr 


228 


229 
E. clitellatus 


232 
E. durbini 


235 


M. obscurus 


239 


: f 3) oe 


——— 


O. anisognathus =e I | 242 


Figs. 224-242. 224-227, Ellisodrilus carronamus Holt; 228-231, Ellisodrilus clitellatus Holt; 232-234, El- 
lisodrilus durbini (Ellis), 234, longitudinal section through a dorsal ridge, showing supernumerary muscles; 235- 
238, Magmatodrilus obscurus (Goodnight); 239-242, Oedipodrilus anisognathus Holt. 


276 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


) 

. ve 
O. cuetzalanae 244 \ LL 
: 247 


O. macbaini 


249 
(7 | 2 
O. oedipus 


P. alcicornus 


257 Wy 
255 
P. cedrus 258 of 


Figs. 243-258. 243-246, Oedipodrilus cuetzalanae Holt; 247-248, Oedipodrilus macbaini (Holt); 249-252, 
Oedipodrilus oedipus Holt; 253-254, Pterodrilus alcicornus Moore; 255-258, Pterodrilus cedrus Holt. 


VOLUME 106, NUMBER 2 


ginia (Holt 1968c:12) in which the species 
is abundant. 


Pterodrilus cedrus Holt, 1968 
Figs. 255-258 


Pterodrilus cedrus Holt, 1968c:21-23. 


Types. —Holotype and 5 paratypes, 
USNM 36464, 3 paratypes, PCH 1396, on 
Orconectes placidus (Hagen) and Cambarus 
tenebrosus Hay, from a small stream at the 
intersection of State Highways 51 and 53 in 
Celina, Clay Co., Tennessee, by Perry C. 
and Virgie F. Holt, 25 Jul 1961. 

Distribution. —The eastern Highland Rim 
and Nashville Basin regions of Tennessee. 


Pterodrilus choritonamus Holt, 1968 
Figs. 259-262 


Pterodrilus choritonamus Holt, 1968c:26- 
28. 


Types. —Holotype, USNM 36471, 2 
paratypes, USNM 36472, on Cambarus te- 
nebrosus Hay, from a tributary (Holt Spring 
Branch) ca. 4.5 mi N of Livingston, Overton 
Co., Tennessee, by Perry C. and Virgie F. 
Holt, 24 Jul 1961; 5 paratypes, PCH 1393, 
on C. tenebrosus and Orconectes placidus 
(Hagen), from Little Eagle Creek, Overton 
Co., Tennessee, by Perry C. and Virgie F. 
Holt, 24 Jul 1961. 

Distribution. —Tributaries of the Cum- 
berland River in the eastern Highland Rim 
region in Tennessee (see locality records for 
Pterodrilus manuscript by Holt in the Li- 
brary of Congress and the USNM). Some 
conspecific material is deposited in the Na- 
tional Museum of Natural History, USNM 
36473-36476. 


Pterodrilus distichus Moore, 1894 
Figs. 263-266 


Pterodrilus distichus Moore 1894:453- 
454.—Pierantoni, 1912:25.—Hall, 1914: 
190.—Ellis, 1919:254.—Goodnight, 
1940:60; 1943:100.—Holt, 1968c:12. 


277 


Types.—From western New York, loca- 
tion unknown (Holt 1968c:12). 

Distribution. —Ohio, Mississippi, Great 
Lakes, drainage systems in Kentucky, Ohio, 
Indiana, Illinois, Michigan, and New York. 


Pterodrilus hobbsi Holt, 1968 
Figs. 267-270 


Pterodrilus hobbsi Holt, 1968c:18. 


Types. —Holotype, USNM 36486, 5 
paratypes, USNM 36487, on Cambarus 
rusticiformis Rhoades, Orconectes rusticus 
(Girard), and Orconectes placidus (Hagen), 
from Spring Creek, 1.4 mi N of the Putnam 
Co. line on State Highway 42, Overton Co., 
Tennessee, by Perry C. and Virgie F. Holt, 
26 Jul 1961. 

Distribution. —Cumberland River in 
Tennessee and Kentucky, the upper Ten- 
nessee drainage in Tennessee, the New Riv- 
er drainage in Virginia and North Carolina, 
the Big Sandy drainage in Virginia (Holt 
1968c:20, 38) 


Pterodrilus mexicanus Ellis, 1919 
Figs. 271-272 


Pterodrilus mexicanus Ellis, 1919:254.— 
Goodnight, 1940:63.—Holt, 1968c:15; 
1973b:32. 


Types. —Holotype, USNM 17654, on 
Procambarus mexicanus (Erichson), from 
Mirador, Veracruz, México, by Nelson and 
Goldman. 

Distribution.—Mountains of Arkansas, 
Oklahoma, and Missouri. 

Notes: The apparent disjunct distribution 
of this species is difficult to explain. Hobbs 
(1989) lists the type locality of P. mexicanus 
as “El Mirador de Zacuapan, 8 km NE of 
Huatusco, Veracruz, México.’ However, 
Holt was unable to find P. mexicanus in 
Mexico (Holt 1973b:32). 


Pterodrilus missouriensis Holt, 1968 
Figs. 273-274 


Pterodrilus missouriensis Holt, 1968c:28-32. 


278 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


267 


P. hobbsi 
270 


Z 271 


P. mexicanus 2/2 


273 


P. missouriensis 


Figs. 259-274. 259-262, Pterodrilus choritonamus Holt; 263-266, Pterodrilus distichus Moore; 267-270, 
Pterodrilus hobbsi Holt; 271-272, Pterodrilus mexicanus Ellis; 273-274, Pterodrilus missouriensis Holt. 


VOLUME 106, NUMBER 2 


Types. —Holotype, USNM 36469, 2 
paratypes, USNM 36470, 2 paratypes, PCH 
1476, on Orconectes luteus (Creaser), from 
Whetstone Creek, 5 mi W of Mountain 
Grove, Wright Co., Missouri, by Perry C. 
Holt, 23 Aug 1961. 

Distribution. —Known only from the type 
locality. 


Pterodrilus simondsi Holt, 1968 
Figs. 275-276 


Pterodrilus simondsi Holt, 1968c:23-26. 


Types. —Holotype, USNM 26477, 5 
paratypes, USNM 36478, 4 paratypes, PCH 
989, on Cambarus bartonii bartonii (Fabri- 
cius), from a tributary to the Oconee River 
S of Morganton, Fannin Co., Georgia, by 
Kenneth W. Simonds, 6 Nov 1958. 

Distribution. —Tributaries to the Oconee 
River in Fannin and Union Counties, Geor- 
gia and Cherokee Co., North Carolina. 


Sathodrilus Holt, 1968 
Sathodrilus Holt, 1968b:294. 


Type species.—Sathodrilus carolinensis 
Holt, 1968, by original designation. 
Gender. — Masculine. 


Sathodrilus attenuatus Holt, 1981 
Figs. 277—280 


Sathodrilus attenuatus Holt, 1981b:849- 
853. 


Types. —Holotype, USNM 65227, 26 
paratypes, PCH 1113, on Pacifastacus leni- 
usculus klamathensis (Stimpson), from Elk 
Creek, ca. 12.6 mi S of Cottage Grove, 
Douglas Co., Oregon, by Perry C. and Virgie 
F. Holt, 11 Jul 1960. 

Distribution. —Streams of the Cascade and 
Coastal Ranges in Oregon and Washington 
to the headwater streams of the Snake River 
in Wyoming. 


Sathodrilus carolinensis Holt, 1968 
Figs. 281-284 


279 


Sathodrilus carolinensis Holt, 1968b:296-— 
299. 


Types. —Holotype, USNM 37107, 1 
paratype, USNM 37108, 1 paratype, PCH 
1333, on Cambarus latimanus (Le Conte) 
and Cambarus sp., from a small stream ca. 
11.5 mi SW of Anderson, Anderson Co., 
South Carolina, on U.S. Highway 29, by 
Perry C. and Virgie F. Holt, 21 Mar 1961. 

Distribution. —Known only from the type 
locality. 


Sathodrilus chehalisae Holt, 1981 
Figs. 285-286 


Sathodrilus chehalisae Holt, 1981b:853- 
855. 


Types. —Holotype and 7 paratypes, 
USNM 65228, 3 paratypes, PCH 1813, on 
Pacifastacus leniusculus trowbridgii (Stimp- 
son), from Chehalis River at Adna, Lewis 
Co., Washington, by Perry C. and Virgie F. 
Holt, 15 Aug 1964. 

Distribution. —Known only from the type 
locality. 


Sathodrilus dorfus Holt, 1977 
Figs. 287-290 


Sathodrilus dorfus Holt, 1977:120—122. 


Types. —Holotype, USNM 53643, 3 
paratypes, PCH 1120, on Pacifastacus leni- 
usculus klamathensis (Stimpson), from a 
small tributary to the Yaguina River, 13.4 
km NE of Toledo, Lincoln Co., Oregon, by 
Perry C. and Virgie F. Holt, 12 Jul 1960. 

Distribution. —Known only from the type 
locality. 


Sathodrilus elevatus (Goodnight, 1940) 
Figs. 291-294 


Cambarincola elevata Goodnight, 1940:34— 
35% 

Cambarincola ? elevata.—Hoffman, 1963: 
27S: 

Sathodrilus elevatus. — Holt, 1978:473-48 1. 


280 


Types. —Holotype, on Orconectes virilis 
Hagen, from Leaf River, Illinois; paratypes, 
on O. virilis, from: Macoupin Creek near 
Carlinville, Illinois; Buck Creek, near Pen- 
field, Illinois; Leaf River, near Bryon, Illi- 
nois; Seven Mile Creek, Rock River drain- 
age, Illinois; and Lake Geneva, Wisconsin. 

Distribution. —The upper Mississippi and 
Red Rivers and Great Lakes drainages in 
Illinois, Indiana, Iowa, Michigan, Minne- 
sota, South Dakota, Wisconsin in the Unit- 
ed States and Ontario, Canada (Holt 1978: 
478-481). 

Notes: The redescription of Sathodrilus 
elevatus and the determination that it is con- 
specific with Goodnight’s holotype were 
based upon the comparison of the holotype 
and numerous specimens from the localities 
listed by Holt (1978:479-481). Goodnight’s 
material, including types, was lost (Good- 
night, pers. comm..). 


Sathodrilus hortoni Holt 
Figs. 295-298 


Sathodrilus hortoni Holt, 1973a:97-99. 


Types. —Holotype, USNM 48713, 2 
paratypes, USNM 49714, 2 paratypes, PCH 
2716, on Cambarus diogenes Girard and 
Cambarus sp., from Pond Creek, 2.1 mi NE 
_ of Laurel Hill, Okaloosa Co., Florida, by 
Horton H. Hobbs, III, 10 Aug 1968. 

Distribution. —Known only from the type 
locality. 


Sathodrilus inversus (Ellis, 1919) 
Figs. 299-302 


Cambarincola inversa Ellis, 1919:259-260. 

Cambarincola ? inversa.—Hoffman, 1963: 
294. 

Sathodrilus virgiliae. —Holt, 1977:128-131. 

Sathodrilus inversus. —Holt, 1981b:855- 
856. 


Types. —Holotype, USNM 16780, 5 
paratypes, USNM 17680, Eugene, Oregon, 
on Pacifastacus leniusculus klamathensis 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


(Stimpson), from Eugene, Oregon, by J. E. 
Gutberlet. 

Distribution. —Common in streams of the 
Pacific versant in Oregon and Washington. 


Sathodrilus lobatus Holt, 1977 
Figs. 303-306 


Sathodrilus labatus Holt, 1977:122-125. 


Types. Holotype and 3 paratypes, 
USNM 53644, 5 paratypes, PCH 1117, on 
Pacifastacus leniusculus klamathensis 
(Stimpson), from Mary’s River, 7.7 km E 
of Blodgett, Benton Co., Oregon, by Perry 
C. and Virgie F. Holt, 12 Jul 1960. 

Distribution. —Western Oregon and 
Washington (Holt 1977:125). 


Sathodrilus megadenus Holt, 1968 
Figs. 307-310 


Sathodrilus megadenus Holt, 1968b:302- 
305. 


Types. —Holotype, USNM 37109, 2 
paratypes, USNM 37110, 2 paratypes, PCH 
1346, on Cambarus latimanus (Le Conte), 
from a small stream, 3.1 mi N of Buchanan, 
Haralson Co., Georgia, by Perry C. and Vir- 
gie F. Holt, 25 Mar 1961. 

Distribution. —Known only from the type 
locality. 


Sathodrilus nigrofluvius Holt, 1989 
Figs. 311-314 


Sathodrilus nigrofluvius Holt, 1989b:738- 
741. 


Types. —Holotype, USNM 118199, 3 
paratypes, USNM 118200-118202, on un- 
known host, from a tributary of the Black 
River, 2 mi NE of Lesterville, Reynolds Co., 
Missouri, on State Road 21, by Perry C. and 
Virgie F. Holt, 22 Aug 1961. 

Distribution. —Known only from the type 
locality. 


VOLUME 106, NUMBER 2 281 


t 275 


P. simondsi 


277 


S. attenuatus 


281 282 
/ 


S. carolinensis 


285 


S. chehalisae 


287 LL_E: fas 
286) ES 
S. dorfus 4 a 


Figs. 275-290. 275-276, Pterodrilus simondsi Holt; 277-280, Sathodrilus attenuatus Holt; 281-284, Sath- 
odrilus carolinensis Holt; 285-286, Sathodrilus chehalisae Holt; 287-290, Sathodrilus dorfus Holt. 


282 


Sathodrilus norbyi Holt, 1977 
Figs. 315-318 


Sathodrilus norbyi Holt, 1977:125-128. 


Types. —Holotype and 3 paratypes, 
USNM 53642, 15 paratypes, PCH 920, on 
Pacifastacus leniusculus klamathensis 
(Stimpson), from Union Flat Creek, ca. 13 
km W of Pullman, Whitman Co., Wash- 
ington, by Darwin E. Norby, 11 Jul 1958. 

Distribution. —Idaho and Washington 
(Holt,.197-7:127): 


Sathodrilus okaloosae Holt, 1973 
Figs. 319-325 


Sathodrilus okaloosae Holt, 1973a:99-102. 


Types. —Holotype, USNM 49715, 5 
paratypes, USNM 49716, 2 paratypes, PCH 
2720, on Procambarus evermanni (Faxon) 
and P. versutus (Hagen), collected 1.0 mi E 
of Santa Rosa Co. line on U.S. Highway 90, 
Okaloosa Co., Florida, by Horton H. Hobbs, 
III, 12 Aug 1968. 

Distribution. —Known only from the type 
locality. 


Sathodrilus prostates Holt, 1973 
Figs. 326-331 


Sathodrilus prostates Holt, 1973b:33-36. 


Types. —Holotype, USNM 4532 [sic] 
(should be 45431), on Procambarus acutus 
cuevachicae (Hobbs), from El Ajenjibre, 
Mesa de San Diego, km 262 de la carretera 
México [Tuspan, Puebla, México], by A. 
Villalobos and H. H. Hobbs, Jr., 12 Apr 
1957; 2 paratypes, USNM 45432, on the 
freshwater crab Pseudothelphusa veracru- 
zana, from Rio Tapalapa, Veracruz, Méxi- 
co, by A. Villalobos and H. H. Hobbs, Jr.; 
1 paratype in the IBUM from the latter lo- 
cality; 7 paratypes, PCH 700, from the latter 
locality. 

Distribution. —From the lower slopes of 
the Sierra Oriental in Puebla to the lowlands 
of Veracruz, México. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Sathodrilus rivigeae Holt, 1988 
Figs. 332-335 


Sathodrilus rivigeae Holt, 1988b:804—807. 


Types. —Holotype, USNM 119545, 2 
paratypes, USNM 119546 (PCH 1089), on 
Orconectes palmeri longimanus (Faxon), 
from cool pools in a medium sized stream 
in Ouachita National Forest 3.2 mi E of 
Joplin, Montgomery Co., Arkansas, at 
crossing of U.S. Highway 270, by Perry C. 
and Virgie F. Holt, 23 Jun 1960. 

Distribution. —Known only from the type 
locality. 


Sathodrilus shastae Holt, 1981b 
Figs. 336-338 


Sathodrilus shastae Holt, 1981b:856-859. 


Types. —Holotype, USNM 65230, 4 
paratypes, PCH 1818, on Pacifastacus fortis 
(Faxon), from the headwaters of Fall River, 
Thousand Springs Ranch, Shasta Co., Cal- 
ifornia, by Perry C. and Virgie F. Holt, 19 
Aug 1964. 

Distribution. —Known only from the type 
locality. 


Sathodrilus veracruzicus Holt, 1968 
Figs. 339-344 


Sathodrilus veracruzicus Holt, 1968b:305— 
308. 


Types. —Holotype, USNM 37105, 3 
paratypes, USNM 37106, 3 paratypes, PCH 
1623, on Procambarus hoffmanni (Villalo- 
bos), from waters near Coyutla, Veracruz, 
by Alejandro Villalobos, 16 Apr 1949. 

Distribution. —Known only from the type 
locality. 

Notes: In the description of this species 
Holt (1968b:299, 307) correctly states that 
oral papillae are absent. In the description 
of S. veracruzicus it is incorrectly stated that 
oral papillae are present in both species. 
They are present only in S. villalobosi. 


VOLUME 106, NUMBER 2 283 


291 


S. elevatus 


295 
S. hortoni 


299 


S. inversus 
Spg 


ipr 


303 


S. lobatus 


307 


S. megadenus 308 


Figs. 291-310. 291-294, Sathodrilus elevatus (Goodnight), 292, left lateral view of reproductive system, 293, 
spermatheca, 294, right lateral view of reproductive system; 295-298, Sathodrilus hortoni Holt; 299-302, 
Sathodrilus inversus (Ellis); 303-306, Sathodrilus lobatus Holt; 307-310, Sathodrilus megadenus Holt. 


284 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


311 


S. nigrofluvius 


315 


S. norbyi 


319 


S. okaloosae 


326 


S. prostates 


332 


S. rivigeae 333 ==. AE 335 


Figs. 311-335. 311-314, Sathodrilus nigrofluvius Holt; 315-318, Sathodrilus norbyi Holt; 319-325, Sath- 
odrilus okaloosae Holt, 321, lateral view of everted penis; 326-331, Sathodrilus prostates Holt; 332-335, Sath- 
odrilus rivigeae Holt. 


VOLUME 106, NUMBER 2 


Sathodrilus villalobosi Holt, 1968 
Figs. 345-348 


Sathodrilus villalobosi Holt, 1968b:299-302, 
1973b:36-38. 


Types. —Holotype, USNM 37101, 4 
paratypes, USNM 37102, 4 paratypes, PCH 
208, on Procambarus paradoxus (Ort- 
mann), from Tetela de Ocampo, Puebla, 
México, by Alejandro Villalobos, May 1941. 

Distribution. — The type locality and Aqua 
Fria, Puebla, México. 


Sathodrilus wardinus Holt, 1981 
Figs. 349-352 


Sathodrilus wardinus Holt, 1981b:859-861. 


Types. — Holotype and 1 paratype, USNM 
65229, 5 paratypes, PCH 921, on Pacifas- 
tacus leniusculus klamathensis (Stimpson), 
from Purdy Creek, 6 mi N of Gig Harbor, 
Pierce Co., Washington, by Darwin E. Nor- 
by, 26 Jun 1939. 

Distribution. — Pierce Co., Washington. 


Tettodrilus Holt, 1968 
Tettodrilus Holt, 1968b:312. 


Type species.—Tettodrilus friaufi Holt, 
1968, by original designation. 
Gender. — Masculine. 


Tettodrilus friaufi Holt, 1968 
Figs. 353-357 


Tettodrilus friaufi Holt, 1968b:3 14-317. 


Types. —Holotype, USNM 37099, 1 
paratype, USNM 37100, on Orconectes mi- 
rus (Ortmann), O. rhoadesi Hobbs, Cam- 
barus graysoni Faxon, and C. tenebrosus 
Hay, from a small stream ca. 8.5 mi S of 
Lewisburg, Marshall Co., Tennessee, on U:S. 
Highway 431, by Perry C. and Virgie F. 
Holt, 18 Apr 1960; 1 paratype, PCH 1007, 
on C. graysoni and C. tenebrosus, from a 
stream tributary to the Harpeth River, 2.4 
mi S of Franklin, Williamson Co., Tennes- 


285 


see, on U.S. Highway 432, by Perry C. and 
Virgie F. Holt, 18 Apr 1960; 1 paratype, 
PCH 1008, on C. graysoni, from a small 
stream, 5.3 mi S of Franklin, Williamson 
Co., Tennessee, by Perry C. and Virgie F. 
Holt, 18 Apr 1960. 

Distribution. —Streams of the Nashville 
Basin in middle Tennessee. 


Triannulata Goodnight, 1940 
Triannulata Goodnight, 1940:56. 


Type species.—Triannulata magna 
Goodnight, 1940, by original designation. 
Gender. — Feminine. 


Triannulata magna Goodnight, 1940 
Figs. 358-361 


Triannulata magna Goodnight, 1940:56- 
57.—Holt, 1974:63-66. 


Types. —Holotype, USNM 20567, on Pa- 
cifastacus sp., from Naches, Washington 
[collector and date unknown]. 

Distribution. —Yakima and Cowlitz 
Counties, Washington (Holt 1974:66). 


Key to the Genera of Cambarincolidae 
(modified from Pennak, 1978) 


1. Penis a protrusible muscular 
Cone (hie MB)» 6 boo sc Alan 2 

— Penis eversible (Fig. 1F-I) .. 4 

2(1). Spermatheca absent; bursa 
asymmetrical and rounded 
(rigs: 225. .229)) oo Ellisodrilus 


Spermatheca present (Fig. 3) 3 
Length 2.5 mm or greater; no 
dorsal appendages present (Fig. 
LA Re eee. Cambarincola 
- Length less than 2.0 mm; cy- 
lindrical or fan-shaped dorsal 
appendages usually present on 
dorsal ridges (Figs. 253, 255) 
RE LED Re es Pterodrilus 
Penis a cuticular tube, often 
very long (Fig. 1 F—I) 


4(1). 


286 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


| , 336 


S. shastae 


; 339 


S. veracruZzicus 


345 
S. villalobosi 


349 


S. wardinus 


353 


T. friaufi 


Figs. 336-357. 336-338, Sathodrilus shastae Holt; 339-344, Sathodrilus veracruzicus Holt; 345-348, Sath- 
odrilus villalobosi Holt; 349-352, Sathodrilus wardinus Holt; 353-357, Tettodrilus friaufi Holt, 355, lateral view 
of bursa and penis. 


VOLUME 106, NUMBER 2 


Figs. 358-361. 


5(4). 


6(5). 


7(6). 


8(7). 


spg 


358 


T. magna 


Penis with internal strands 
(Fig. 321) or an epithelial or 
muscular wali (Fig. 355) .... 5 
Body segments II to VIII with 
cylindrical projections on dor- 

sal surface (Fig. 218), upper lip 
with four tentacles . Ceratodrilus 
Without projections on body 
Onientacies on lip ~)....).... 6 
Penis and ejaculatory duct a 
continuous, muscular eversi- 

ble tube (Fig. 359) ..Triannulata 
Penis and ejaculatory duct dis- 
tinct regions, ejaculatory duct 

not eversible (Figs. 350, 354) il 
Bursal atrium much longer 
than penial sheath; penial 
sheath eversible (Fig. 236) 

» ate Magmatodrilus 
Bursal atrium shorter than 
penial sheath; penial sheath not 
emensibie (FIG4278)) 55 eau. 8 
Seminiducal gland small and 
slender, length and diameter 
about equal to that of ejacula- 
tory duct (Fig. 354) ... Tettodrilus 
Seminiducal gland larger than 
ejaculatory duct (Fig. 278) .. 

_ 6 a ta ee Sathodrilus 


Key to the Species of Cambarincola 


Cephalic area conspicuously 
enlarged (Figs. 112, 178) .... 2 
Cephalic area of normal size 
(Fig. 2) 


359 


2(1). 


5(4). 


6(5). 


287 


361 


Triannulata magna Goodnight, 361, ventral view of upper jaw. 


Prostomial tentacles present 
(Fig. 112); spermathecal bulb 
cylindrical (Fig. 113) ....... 
Pe Rem, C. macrocephala 
Prostomial tentacles absent 
(Fig. 178); spermathecal bulb 
Oval (FIGKAY Oy Oe. C. restans 
Prostomial tentacles present 


(Figs 492.83)" 2 Cee 4 
Prostomial tentacles absent 
(Riga) Pier ater ts gen ec. 8 


Spermathecal bulb with cylin- 
drical ental process; semini- 
ducal gland with two lobes (Fig. 
SOWWErtS oe See eh tee oe C. holti 
Spermathecal bulb without en- 
tal process; seminiducal gland 
without lobes (Fig. 50) ...... 3 
Body without prominent dor- 
sal ridges (Fig. 94); prostate 
longer than seminiducal gland 
(Figs. 95, 151) 
Body with prominent dorsal 
ridges (Fig. 49); prostate short- 
er than seminiducal gland 
(Figs. 50, 173) 
Prostomial tentacles small (Fig. 
94); diameter of prostate less 
than that of seminiducal gland 
(HIG TOS eed erp he C. ingens 
Prostomial tentacles large (Fig. 
150); diameter of prostate 
greater than that of semini- 
ducal gland (Fig. 151) ...... 

We Pete eis C. montanus 


8(3). 


9(8). 


10(9). 


11(8). 


12(11). 


1301 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Prostomial tentacles large (Fig. 
49); diameter of prostate three 
times that of seminiducal gland 
(Fig 50)i ee C. fallax 
Prostomial tentacles small (Fig. 
172); diameter of prostate two 
times that of seminiducal gland 
(Fig. 173) C. philadelphicus 
Length of prostate greater than 
that of seminiducal gland (Figs. 
78, 145) 
Length of prostate equal to or 
less than that of seminiducal 
gland (Figs. 7,99) .......... 11 
Length of spermathecal bulb 
three times that of duct (Fig. 
169); lower jaw with four teeth 
(Fig. 171) C. pamelae 
Length of spermathecal bulb 
about 1.5 times that of duct 
(Fig. 78); lower jaw with two 
teeth (Fig. SO) as on a 
Spermathecal bulb oval, di- 
ameter two times that of duct 
(Fig. 78) C. hoffmani 
Spermathecal bulb lobed, di- 
ameter 1.5 times that of duct 
(Fig. 145) C. micradenus 


Length of prostate no more than 
one-third that of seminiducal 


ee we ee 


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10 


eee © © © © © © 


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gland (Figs. 72,191) ........ 12 
Length of prostate at least half 
that of seminiducal gland (Figs. 
LZ LG sa oS SAI EAS oa sae oe 13 


Seminiducal gland not bi- 
lobed; spermathecal duct lon- 
ger than bulb (Fig. 72) ...... 
C. heterognathus 
Seminiducal gland bilobed; 
spermathecal duct shorter than 
bulb: (ig. 191), as. C. shoshone 
Length of prostate about half 
that of seminiducal gland (Figs. 


eo © © © © © © © © © © © 8 


ETO) ee ee ed 14 
Length of prostate at least two- 
thirds that of seminiducal 
gland (Figs: 7, 99) 9... DS 


14(13). 


15(14). 


16(15). 


17(15). 


18(17). 


19(14). 


20(19). 


21(20). 


Seminiducal gland abruptly 
bent and U-shaped (Figs. 12, 


Lay sacks Oe eee 16 
Seminiducal gland not 
U-shaped (Figs. 16, 20) ..... 19 
Diameter of spermathecal bulb 
and duct similar (Figs. 44, 62) 16 
Diameter of spermathecal bulb 
greater than that of duct (Fig. 
12) cig eee eee 17 
Spermatheca oval; length of 
prostate about half body di- 
ameter (Fig. 44) ........ C. ellisi 


Spermatheca spherical; length 
of prostate one-fourth that of 
body diameter (Fig. 62) 
neon (bee C. goodnighti 
Length of spermathecal duct 
three times that of bulb (Fig. 
1 Do, sorter tee C. barbarae 
Length of spermathecal duct 
about 1.3 times that of bulb 
(Fig. 205) 
Spermathecal bulb nearly 
spherical; length of prostate 1.5 
times that of bursa (Fig. 205) 

. BEd RE SEE C. susanae 
Spermathecal bulb elongate; 
length of prostate twice that of 
bursa (Fig. 217) C. vitreus 
Diameter of seminiducal gland 
1.5 times that of prostate (Fig. 
193) C. smalleyi 
Diameter of seminiducal gland 
at least twice that of prostate 
(Figs. 16, 20) 
Seminiducal gland with two 
lobes (Figs. 20, 213) 
Seminiducal gland not bilobed 
(Fig. 16) 
Length of spermatheca about 
one-half body diameter (Fig. 

D1) NOI I C. branchiophilus 
Length of spermatheca nearly 
as great as body diameter (Fig. 
213) C. virginicus 


ee © we © 


18 


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20 


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21 


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22 


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VOLUME 106, NUMBER 2 


22(20). 


23(22). 


24(23). 


25413). 


26(25). 


27(23). 


28(27). 


29(28). 


30(29). 


Length of bursa greater than 
that of seminiducal gland (Fig. 
36) C. demissus 
Length of bursa less than that 
of seminiducal gland (Figs. 16, 
24) 
Spermathecal bulb spherical 
(Fig. 16) C. bobbi 
Spermathecal bulb elongate 
(Figs. 24, 68) 
Diameter of ejaculatory duct 
one-fifth that of bursa: diam- 
eter of prostate one-fourth that 
of seminiducal gland (Fig. 24) 
te. 12) eae ae C. carcinophilus 
Diameter of ejaculatory duct 
one-half that of bursa; diam- 
eter of prostate one-half that 
of seminiducal gland (Fig. 68) 
RMI EP 62. Soe oe C. gracilis 
Seminiducal gland bilobed 


eee) oe @ in Bat te) eo eo Feige a 


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bess, NO?) oth sch... 26 
Seminiducal gland not bilobed 
REISSUE SO) ewe) ewes... « 27 


Prostate and bursa of similar 
length (Fig. 3) ... C. acudentatus 
Length of prostate three times 
that of bursa (Fig. 167) ..... 
C. ouachita 
Diameter of seminiducal gland 
equal to that of prostate (Fig. 


se © © © © © © © ee ee ee eee 


OS he re en ee 28 
Diameter of seminiducal gland 
at least 1.5 times that of pros- 
gate (Bigs. 75:30, S4)): cca: ..: 32 


Spermathecal bulb smaller 
than bursa (Fig. 124) .. C. manni 
Spermathecal bulb equal to or 
larger than bursa (Figs. 105, 
108) 
Spermathecal duct about twice 
as long as spermathecal bulb 
OBig tO) ye C. speocirolanae 
Spermathecal bulb longer than 
spermathecal duct (Fig. 105) 30 
Diameter of spermathecal bulb 


Reg is a «wa a hk elle eee ee ee eS. a 


31(30). 


32(27). 


33632): 


34(33). 


35(34). 


36(33). 


37(36). 


289 


four times that of duct (Fig. 
105) C. leoni 
Diameter of spermathecal bulb 
1.5 times that of duct (Fig. 108) 


Length of spermatheca half 
that of body diameter; length 
of prostate two-thirds that of 
seminiducal gland (Fig. 108) 
C. leptadenus 
Length of spermatheca nearly 
equal to body diameter; length 
of prostate equal to that of 
seminiducal gland (Fig. 136) 
C. mesochoreus 
Diameter of spermathecal bulb 
at least twice that of duct (Figs. 
38, 56) 
Diameter of spermathecal bulb 
no more than 1.5 times that of 
auch (Figs’ 7, 84) ee Se. 
Length of prostate equal to that 
of seminiducal gland (Figs. 56, 
91) 
Length of prostate two-thirds 
that of seminiducal gland (Figs. 
38, 130) 
Lengths of prostate and bursa 
equal (Fig. 91) ... C. illinoisensis 
Prostate twice as long as bursa 
(Figs. 56, 161) 
Prostate straight; diameter of 
spermathecal bulb four times 
that of duct (Fig. 56); lower jaw 
with four teeth (Fig. 58) .... 
C. floridanus 
Prostate J-shaped; diameter of 
spermathecal bulb 1.5 times 
that of duct (Fig. 161); lower 
jaw with two teeth (Fig. 163) 
PET ire has RN ome C. olmecus 
Spermathecal bulb with a small 
ental lobe (Fig. 139) ...C. meyeri 
Spermathecal bulb without an 


a #1 8 he eure, 6) 6) © Bl (6) @ te. je 


Si 


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ne 2 we we 6) le ee ee oe ee 


39 


34 


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« 6 «6, =o «6 & « @ « » « © = « ‘@ = = 


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“2s © © © © © © © we © we we ee ee 


eptallebet Pig. 38) pcccpatee ss: 37 
Seminiducal gland L-shaped 
(Big: GB ies sacar seks C. dubius 


290 


38(37). 


39(32). 


40(39). 


41(39). 


42(39). 


43(42). 


44(43). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Seminiducal gland oval or cy- 
lindrical (Figs. 130, 210) .... 
Spermathecal bulb oval, di- 
ameter 3 times that of duct; 
prostate longer than bursa (Fig. 
130) C. marthae 
Spermathecal bulb spherical, 

diameter four times that of 

duct; lengths of prostate and 

bursa similar (Fig. 201) 


38 


C. steevesi 
Prominent dorsal ridges pres- 


ent, (Figs. 29) 180) reise re 40 
Dorsal ridges absent (Figs. 6, 
Sis is chug Bae eres Ot IP 41 


Bursa longer than prostate; 
spermathecal duct and bulb 
equal in length (Fig. 30) .... 
so. eh eacerpe C. chirocephala 
Bursa shorter than prostate; 
spermathecal duct half as long 
as bulb (Fig. 181) ....C. serratus 
Length of seminiducal gland 
less than that of bursa; sper- 
mathecal bulb with a large en- 
tal lobe (Fig. 7) C. alienus 
Length of seminiducal gland 
equal to or greater than bursa; 
spermathecal bulb without 
large ental lobe (Figs. 84, 99) 
Seminiducal gland L- or 
U-shaped (Figs. 165, 209) 43 
Seminiducal gland straight or 
slightly curved (Figs. 84, 99) 
Bursa cylindrical, length twice 
its diameter (Fig. 118) ...... 
C. macrodontus 
Bursa oval, length no greater 
than 1.5 times its diameter 
(Piss: 1655 209 se een par We ee 
Seminiducal gland U-shaped; 
diameter of spermathecal duct 
less than that of prostate (Fig. 
Toa SR ee C. osceolai 
Seminiducal gland L-shaped; 
diameter of spermathecal duct 
similar to that of prostate (Fig. 
209) C. toltecus 


42 


45 


as 


45(42). 


46(45). 


47(46). 


Upper and lower jaws with 
three teeth (Figs. 85, 86); sper- 
mathecal bulb cylindrical (Fig. 
SA st ea. eh C. holostoma 
Upper jaw with five teeth (Figs. 
100, 156), lower with four or 
five teeth (Fig. 157); sperma- 
thecal bulb oval (Figs. 99, 155) 
Lower jaw with one large and 
four small teeth (Fig. 187); 
length of prostate 1.5 times that 
of bursa (Fig. 185); body length 
2.5 mm C. sheltensis 
Lower jaw with four teeth of 
equal size (Figs. 101, 157); 
length of prostate and bursa 
similar (Fig. 99); body length 
less‘ than: 2:0; mmm .5. aR 
Length of prostate 0.8 times 
that of seminiducal gland; 
length of ejaculatory duct twice 
its diameter (Fig. 99); central 
tooth of upper jaw larger than 
lateral teeth (Fig. 100) 
5¢4i2 Base, canes C. jamapaensis 
Length of prostate 0.6 times 
that of seminiducal gland; 
length and diameter of pros- 
tate equal (Fig. 155); teeth of 
upper jaw similar in size (Fig. 
156) C. nanognathus 


46 


47 


Key to the Species of Ceratodrilus 


Length of tentacles on head 200 
um (Fig. 218); projections on 
segments II-VII 146 um long 
pte LAN C. ophiorhysis 
Length of tentacles on head 90 
um (Fig. 222); projections on 
segments IJ—-VII 50 um long . 
C. thysanosomus 


Key to the Species of E/lisodrilus 


Bursa large and bent, length 
about two-thirds body diam- 
ter (Pig 229). ra E. clitellatus 
Bursa not bent, length about 


VOLUME 106, NUMBER 2 


2(1). 


half body diameter (Figs. 225, | 
233) 
Bursa wider than long; pros- 
tate diameter equal to that of 
seminiducal gland (Fig. 225) 
fe <p a ei ee E. carronamus 
Width and length of bursa 
about equal; diameter of pros- 
tate half that of seminiducal 
pland (Fig. 233) ...... E. durbini 


Key to the Species of Oedipodrilus 


3(2). 


Prominent dorsal ridges pres- 

2 eae eee O. oedipus 

No dorsal ridges present .... 2 

Spermathecal bulb and duct 

slender, diameters equal; sper- 

mathecal bulb bent at right an- 

gle to duct (Fig. 244) 
Me op ah) Say, O. cuetzalanae 

Spermathecal bulb and duct 

robust (Figs. 240, 248) ....... 3 

Length of spermathecal duct 

half that of bulb; bursa straight 

IEE) ae O. anisognathus 

Length of spermathecal duct 

1.5 times that of bulb; bursa 

long and bent (Fig. 248) .... 

O. macbaini 


we, a) wh ie, wale 


Cito ie ch Se 8) we ene |e «i 2 a Ne 


Key to the Species of Pterodrilus 


21): 


Four of five pairs of dorsal, fan- 
like projections on body (Fig. 
253) 
Body with one to seven pairs 
of dorsal, finger-like projec- 
Hous (Piss. 255.265) 22) .2. 3 
Diameter of spermathecal bulb 
1.5 times that of duct; diam- 
eter of seminiducal gland equal 
to that of bursa (Fig. 254) ... 
eS er ae oe P. alcicornus 
Diameter of spermathecal bulb 
three times that of duct; di- 
ameter of seminiducal gland 
half that of bursa (Fig. 276) . 
Ree. ee. P. simondsi 


S(t): 


4(3). 


5(4). 


6(5). 


7(3). 


291 
Length of prostate equal to that 
of seminiducal gland (Figs. 
2x6. 260) Se oe 4 


Length of prostate 0.6 to 0.5 
times that of seminiducal gland 
Wipsh264 5 274) es SIS. 7 
Diameter of bursa twice that 
of seminiducal gland; length of 
bursa half that of body diam- 
eter (Pie, 27 2))"..24."P. mexicans 
Diameter of bursa one to 1.5 
times that of seminiducal 
gland; length of bursa one-third 
or less that of body diameter 
(Figs. 256, 260) 
Length of spermathecal duct 
twice that of bulb (Fig. 256) . 

Dey Se ee P. cedrus 
Length of spermathecal duct 
less than that of bulb (Fig. 260) 


Diameter of spermathecal bulb 
and duct equal; length of sem- 
iniducal gland and bursa equal 
(Figs. 260)'.2P othe: P. choritonamus 
Diameter of spermathecal bulb 
three times that of duct; length 
of seminiducal gland twice that 
of bursa (Fig. 268) .... P. hobbsi 
Finger-like projections on the 
dorsal surface of seven body 
segments (Fig. 263); length of 
bursa about one-fourth that of 
body diameter (Fig. 264) .... 

a Pip pe a P. distichus 
Finger-like projections on the 
dorsal surface of only one body 
segment (Fig. 273); length of 
bursa about half that of body 
grameter Pig: 274) 246 ee... 

Nee Site gare B, See P. missouriensis 


Key to the Species of Sathodrilus 


ACEY: 


Tentacles on dorsal lips (Fig. 


Se Ss are Neh Cigna gel Aieti S. lobatus 
Dorsal lips without tentacles 
(LEVEING TOT | kee yet aelladabeniipenaren 2 


Ejaculatory duct spherical, di- 


2928 


7(6). 


8(7). 


9(8). 


10(8). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ameter about half that of bursa 
(HiGS.2.9)2)\) (ili, Bah ete S. elevatus 
Ejaculatory duct cylindrical, 
diameter less than half that of 


bursa,,(B102. 27,8) vc taeats SE as 3 
Spermathecal bulb spherical or 
oval (Figs. 346, 350) ....... 4 
Spermathecal bulb cylindrical 
(FACS 6 Zhe lS Owe feeced case ove 5 


Length of spermathecal duct 
four times the diameter of the 
bulb; prostate absent (Fig. 346) 
Se en Ty Pee S. villalobosi 
Length of spermathecal duct 
1.3 times the diameter of the 
bulb; prostate present (Fig. 


350) avery ed. We ste: S. wardinus 
Prostate present (Fig. 278) .. 6 
Prostate absent (Fig. 282) ... 12 


Length of spermathecal duct 
three times that of bulb (Fig. 
BA) EE eT Ene ans S. dorfus 
Length of spermathecal bulb 
equal to or greater than that of 
duct (Figs. 278, 308) 
Ejaculatory duct long, diame- 
ter equal to that of bursa (Fig. 
616.) ee a ee S. megadenus 
Ejaculatory duct short, diam- 
eter less than half that of bursa 


(1G 2), 8 Vp eshn es dlig yell oe 8 
Length of prostate about half 
that of seminiducal gland (Fig. 
04) ee ee ee ee et E 9 


Lengths of prostate and sem- 
iniducal gland equal (Fig. 286) 


Bursa spherical, length about 
one-third body diameter (Fig. 
ZS) ae I ee S. attenuatus 
Bursa cylindrical, length about 
half body diameter (Fig. 300) 
JAE. 1 ae eRe: S. inversus 
Length and width of bursa 
equal (Fig. 286) ....S. chehalisae 
Length of bursa two to three 
times its width (Figs. 327, 333) 


11(10). 


12(5). 


13(12). 


14(13). 


15(14). 


16(15). 


17(16). 


Length of spermathecal bulb 
four times that of duct (Fig. 
32-])icwol yet pees S. prostates 
Lengths of spermathecal bulb 
and duct equal (Fig. 333) ... 

i PRON. Eats cog nese ba S. rivigeae 
Seminiducal gland slender and 
looped, length greater than 
body diameter (Fig. 337) 

Thi cx otal Ete ee S. shastae 
Seminiducal gland short, 
length less than half body di- 
ameter (Figs. 282, 316) 
Diameter of seminiducal gland 
equal to that of bursa (Fig. 316) 

S. norbyi 
Diameter of seminiducal gland 
no more than 0.6 that of bursa 
(Figs: 312; 320)... 7. 3) ee 14 
Length of seminiducal gland 
greater than that of bursa; 
length of spermatheca less than 
half body diameter (Fig. 296) 

red sce ba Laseee eek Re hae ot eee S. hortoni 
Length of spermathecal gland 
less than that of bursa; length 
of spermatheca nearly equal to 
or greater than body diameter 
(Figs. 282: °3:12) 3 >...., 2 15 
Spermatheca bent, length 1.5 
times body diameter (Fig. 340) 

PTS ay Ui Sas S. veracruzicus 
Spermatheca not bent, length 
equal to body diameter (Fig. 
282) A 16 
Spermathecal bulb with a slen- 
der ental lobe whose length is 
equal to that of the sperma- 
thecal duct (Fig. 320) ....... 


eo © © © © © © © © © © © 8 ee ew el el ell 


Spermathecal bulb without a 
long ental lobe (Figs. 282, 312) 

oil gaa a ow eae 17 
Diameter of seminiducal gland 
about one-third that of bursa; 
spermathecal duct cylindrical; 
spermathecal bulb without en- 
tal lobe (Fig. 282) S. carolinensis 


VOLUME 106, NUMBER 2 


- Diameter of seminiducal gland 
half that of bursa; spermathe- 
cal duct constricted; sperma- 
thecal bulb with short ental 
lobe (Fig. 312) ....S. nigrofluvius 


Acknowledgments 


We thank Dr. Richard L. Hoffman and 
Dr. Mark Wetzel for detailed comments and 
Dr. Ralph O. Brinkhurst for general sug- 
gestions that helped improve this paper. Mr. 
George C. Steyskal reviewed the paper’s no- 
menclatural citations. Support from Virgin- 
ia Polytechnic Institute and State Univer- 
sity made possible the publication of this 
manuscript. Perry Holt thanks the Virginia 
Academy of Science, the National Science 
Foundation (grants G-4439, G-9828, and 
GB-372), and Virginia Polytechnic Institute 
and State University’s Biology Department 
and Agricultural Research Division for sup- 
porting his collecting trips and research. He 
thanks Dr. Horton H. Hobbs, Jr., for the 
encouragement and advice provided 
throughout his career. Virgie F. Holt and 
Dr. Susan E. H. West provided invaluable 
help on numerous collecting trips. Drs. Jo- 
seph F. Fitzpatrick, John Holsinger, Denton 
W. Crocker, and Harrison R. Steeves, III, 
and others cited in the test also helped with 
collecting or provided specimens. 


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40. 

1973c. Branchiobdellids (Annelida: Clitel- 
lata) from some eastern North American caves, 
with descriptions of new species of the genus 
Cambarincola. —International Journal of Spe- 
leology 5:219-256. 

1973d. An emended description of Camba- 
rincola meyeri Goodnight (Clitellata: Bran- 
chiobdellida).— Transactions of the American 
Microscopical Society 92:677-682. 

1974. An emendation of the genus Trian- 
nulata Goodnight, 1940, with the assignment of 
Triannulata montana to Cambarincola Ellis, 
1912 (Clitellata: Branchiobdellida).— Proceed- 
ings of the Biological Society of Washington 87: 
57-72. 

1977. An emendation of the genus Sathod- 
rilus Holt, 1968 (Annelida: Branchiobdellida), 
with the description of four new species from 
the Pacific drainage of North America.—Pro- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ceedings of the Biological Society of Washington 
90:116-131. 

1978. The reassignment of Cambarincola 
elevatus Goodnight, 1940, (Clitellata: Bran- 
chiobdellida) to the genus Sathodrilus Holt, 
1968.— Proceedings of the Biological Society of 
Washington 91:472-482. 

. 1981a. Aresume of the members of the genus 
Cambarincola (Annelida: Branchiobdellida) 
from the Pacific drainage of the United States. — 
Proceedings of the Biological Society of Wash- 
ington 94:675-695. 

. 1981b. New species of Sathodrilus Holt, 1968, 
(Clitellata: Branchiobdellida) from the Pacific 
drainage of the United States with the synonymy 
of Sathodrilus virgiliae Holt, 1977.—Proceed- 
ings of the Biological Society of Washington 94: 
848-862. 

1982. A new species of the genus Camba- 

rincola (Clitellata: Branchiobdellida) from I[li- 
nois with remarks on the bursa of Cambarincola 
vitreus Ellis, 1919, and the status of Sathodrilus 
Holt, 1968.— Proceedings of the Biological So- 
ciety of Washington 95:251-255. 
. 1984a. On some branchiobdellids (Annelida: 
Clitellata) from Mexico with the description of 
new species of the genera Cambarincola and 
Oedipodrilus.—Proceedings of the Biological 
Society of Washington 97:35-42. 

1984b. A new species of the genus Camba- 
rincola (Clitellata: Branchiobdellida) from Cal- 
ifornia.— Proceedings of the Biological Society 
of Washington 97:544-549. 

. 1986. Newly established families of the order 
Branchiobdellidae (Annelida: Clitellata) with a 
synopsis of the genera. — Proceedings of the Bi- 
ological Society of Washington 99:676-702. 

. 1988a. The correct name of Ceratodrilus or- 
Dhiorhysis Holt, 1960 (Annelida: Branchiob- 
dellida).— Proceedings of the Biological Society 
of Washington 101:308. 

1988b. Four new species of cambarincolids 
(Clitellata: Branchiobdellida) from the south- 
eastern United States with a redescription of 
Oedipodrilus macbaini (Holt, 1955).—Proceed- 
ings of the Biological Society of Washington 101: 
794-808. 

. 1989a. Comments on the classification of the 
Clitellata.—Hydrobiologia 180:1—5. 
1989b. A new species of the cambarincolid 
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posal of a replacement name for Adenodrilus 
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ceedings of the Biological Society of Washington 
102:738-741. 
—, & R. L. Hoffman. 1959. An emended de- 
scription of Cambarincola macrodonta Ellis, with 
remarks on the diagnostic characters of the ge- 


VOLUME 106, NUMBER 2 


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429. 

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1895. The anatomy of Bdellodrilus illumi- 
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5:542. 

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Robinson, D. A. 1954. Cambarincola gracilis sp. nov., 


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a branchiobdellid oligochaete commensal on 
western crayfishes. — Journal of Parasitology 60: 
466-469. 
Sawyer, R. T. 1986. Leech biology and behavior. 
Clarendon Press, Oxford, 500 pp. 
Stephenson, J. 1930. The Oligochaeta. Clarendon 
Press, Oxford, 979 pp. 
Yamaguchi, H. 1932. On the genus Cirrodrilus Pie- 
rantoni, 1905, with a description of a new bran- 
chiobdellid from Japan.— Annotationes Zoolo- 
gicae Japonensis 13:361-367. 
. 1933. Description ofa new branchiobdellida, 
Cambarincola okadai n. sp., parasitic on Amer- 
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9:191-193. 
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perial University, Series VI, 3:177-219. 


Department of Biology, Virginia Poly- 
technic Institute and State University, 
Blacksburg, Virginia 24061, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(2), 1993, pp. 296-304 


BRANCHINECTA SANDIEGONENSIS, A NEW 
SPECIES OF FAIRY SHRIMP 
(CRUSTACEA: ANOSTRACA) FROM 
WESTERN NORTH AMERICA 


Michael Fugate - 


Abstract.—Branchinecta sandiegonensis, a new species of fairy shrimp is 
described from vernal pools on Del Mar Mesa, San Diego County, California. 
The species is found within 50 km of the Pacific Ocean from Santa Barbara, 
California to Valle de las Palmas, Baja California Norte, Mexico. It can be 
distinguished from the other six branchinectids inhabiting southern California 
by the combination of thoracic spine pattern, ovary length, ovisac length and 
shape, and egg morphology of females and the form of the second antenna of 


males. 


Vernal pools, so named for their colorful, 
springtime floral displays as the pools dry 
after winter rains, have long been known for 
their endemic floras (Crampton 1976, Hol- 
land & Jain 1977). Only one fairy shrimp 
however was known to be endemic to the 
extensive vernal pool habitat ranging from 
southern Oregon through the Central Valley 
of California and into northern Baja Cali- 
fornia, Mexico (Dodds 1923, Brtek 1964) 
before a recent monograph on the Anostra- 
ca of California described four endemic, 
vernal pool species (Eng et al. 1990). Here 
I describe another species found primarily 
in vernal pools atop mesas in San Diego 
County, California. 


Methods 


Animals were obtained either from filled 
pools or by hydrating soil samples from dry 
pools. Freshly molted individuals were fixed 
for 3 hours in 3% glutaraldehyde in 0.1 M 
sodium cacodylate at pH 7.4, post-fixed with 
2% osmium tetroxide in sodium cacodylate 
for 2 hours, dehydrated in a graded series 
of 10% glacial acetic acid in absolute etha- 


nol, and transferred to absolute ethanol. The - 


eggs were air-dried from absolute ethanol, 


the mandibles were air-dried after 10 min- 
utes in tetramethylsilane (Dey et al. 1989), 
and all other parts were critically point- 
dried, coated with gold-palladium and ob- 
served on a Philips 515 scanning electron 
microscope. 


Branchinecta sandiegonensis, 
new species 
Figs. 1-13 


Type material. —One 6, holotype, USNM 
256557, 1 2, allotype, USNM 256558, 2 3, 
3 9, paratypes, USNM 256556, 3 4, 5 8, 
paratypes, Los Angeles County Museum of 
Natural History (LACM), LACM 90-356.1, 
2 6, 3 2, paratypes, Hungarian Museum of 
Natural History, Del Mar Mesa, San Diego 
County, California, USA (32°51'N, 
117°15'W), 17 Mar 1990 (coll. M. Simovich 
& M. Fugate); 4 6, 8 °, paratypes, LACM 
89-357.1, 44, 5 2, paratypes Museo Ciencias 
Naturales de La Plata, Kearney Mesa (Mir- 
amar Naval Air Station), San Diego County, 
California (32°50'N, 117°09'W), 8 Feb 1990 


(coll. M. Simovich); 50 4, 50 2, paratypes, 


USNM 294523, Ramona, San Diego Coun- 
ty, California (33°02'’N, 116°52'W), 4 Mar 
1962 (coll. J. E. Lynch); 50 6, 50 9, para- 


VOLUME 106, NUMBER 2 


types, USNM 305974, Poway, San Diego 
County, California (32°55'N, 117°04’W), 3 
Mar 1962 (coll. J. E. Lynch). 

Type locality.—An extensive network of 
vernal pools surrounded by chaparral on 
Del Mar Mesa, San Diego County, Califor- 
nia, USA, 32°51'N, 117°15'W, elev. 100 
meters, south and east of junction of Inter- 
state 5 and Carmel Valley Road (Green- 
wood 1984). 

Etymology. —Named for San Diego 
County, California, USA. 

Male.—Antenna 1 slender, cylindrical, 
approximately 10 times as long as wide, with 
3 long setae and 8 or more shorter aesthe- 
tascs (type 1 and type 2 sensilla, respec- 
tively—Tyson & Sullivan 1979; Fig. 1). An- 
tenna 2 biarticulate, cylindrical, reaching to 
thoracic segment 8 (Fig. 6). Basal and distal 
segments of approximately equal length. 
Basal segment with medial, oval pulvillus, 
near proximal end, of short spines inter- 
spersed with slightly longer, stouter, conical 
spines (Fig. 10a), medial, elevated cluster of 
6-10 short, stout spines, half distance from 
end of basal segment (Fig. 6a), and single 
row containing clusters of papillae, each with 
sensory seta, on distal 0.67 of anterolateral 
surface. Distal segment slightly arcuate, up- 
per 0.25 cylindrical, remainder mediolater- 
ally flattened (Fig. 2a—e). Breadth 0.2 length 
at joint with basal segment expanding to 
0.33 at tip. Mediolateral surfaces 6 times as 
broad as anteroposterior surfaces.. Medial 
surface flat and lateral concave. Anterior 0.5 
of distal portion inflated, turned in medially 
at 45°, ovoid anteriorly, triangular laterally. 
Posterior edge of distal portion with patch 
of raised, ovoid papillae arranged in rows 
(Fig. 10b). 

Body of mandible with spinelike protu- 
berance on posterolateral surface. Molar 
surfaces of mandibles asymmetrical, broad- 
ly oval with dorsal edge concave and ventral 
convex, divided into 2 basic regions, an an- 
teroventral and posterodorsal (Tyson & Sul- 
livan 1981, Fig. 11). Anteroventral region 
of both molar surfaces similar, with ap- 


297 


proximately 45 ridges and furrows, some 
bifurcating, running dorsoventrally. Ridges 
of rectangular cuticular projections bearing 
many conical protuberances, both projec- 
tions and protuberances becoming taller near 
edges of molar surface. Ridges extend to 
dorsal edge on anterior half of right man- 
dible, but only on anterior 0.15 of left. Pos- 
terodorsal region with 1 long, thin spine at 
posterior edge and row of spines, decreasing 
in size anteriorly, along dorsal edge. On right 
mandible, spines sickle-shaped, each bor- 
dering short row of widely spaced cuticular 
projections, and each row in turn bordering 
small area of relatively unadorned cuticle 
dorsal to ridges of anteroventral region. On 
left mandible, spines squat, conical, and 
more widely spaced, becoming longer and 
thinner anteriorly and bordering large area 
of unadorned cuticle extending almost en- 
tire length of molar surface. 

Maxilla 1 with short, stout spine on ven- 
tral edge and 18 setae, approximately 3 times 
as long as ventral spine, on medial edge (Fig. 
3). Proximal 0.33 of seta with 4—7 stout spi- 
nules, arranged in single row, along medial 
surface, distal 0.67 with 2 rows of setules. 
Maxilla 2 small, with 6-10 setae on crown 
(Fig. 4). Setae pliant, with 2 rows of setules 
on distal 0.8. Medial surface of maxilla 2 
with 2-3 short, pliant setae covered with 
setules, not arranged in rows, and 1 small, 
stout spine nearer base. Ventral surface of 
labrum broadly triangular with lateral flap 
on each side, posterior edge of flaps spinose. 
Oral surface with small, distal lobe covered 
with fine setae. 

Thoracic segments appendage-bearing, 
with paired dorsolateral, cuticular papillae, 
each papillus with sensory seta (Figs. 6, 9). 
Papillae not on raised protuberances, lateral 
on segments 1-3, 5—8, and 10—11, dorsal on 
4 and 9 (compare Lynch 1960, fig. 4 and 
Cohen 1983, fig. 22). All 11 pairs of ap- 
pendages similar in form, but those of seg- 
ment 11 reduced (Fig. 8). There is much 
confusion as to which lobes of anostracan 
phyllopodous limbs are homologous with 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


298 


~ ~ 
<x 
PS ST Se ——~ = 
Sy SxS 
= ENS Sieeseas ‘ 
/ 
/ 


SSD 


ae QM ERIN Dy ee 


5a 2 


Branchinecta sandiegonensis, new species male. 1. antenna, 1, distal tip. 2a—e. antenna 2, distal 
segment. a. lateral. b. anterolateral. c. anterior. d. medial. e. dorsomedial. 3. maxilla 1, medial view. 4. maxilla 


Figs. 1-5. 
2, posterior view. Sa, b. right penis. a. ventral view. b. distal tip, lateral view. (scale bars 2, 4, 5, 6: 0.1 mm, 3 


1 mm) 


VOLUME 106, NUMBER 2 


299 


TERS : =S ri} XS D —~ 


eee Bi Ss 3 
i en 
: ea) ee SS 
i “Y § 5 


Est 
f 3 
x 


Fig. 6. Branchinecta sandiegonensis, new species male. a. ventral view. b. lateral view. (scale bar 1.0 mm) 


stenopodous limbs. Here the nomenclature 
of Linder (1941) is employed. Endite | with 
3 centers of origin for its numerous poste- 
rior setae. Middle series with 7—9 posterior 
setae and long, slightly toothed anterior seta. 
Distal series with 11—16 posterior setae and 
2 anterior setae on basal edge, distal seta 
comblike, with single row of stout setules, 
basal seta shorter, spiniform. Appendage 1 1 
with 3-5 posterior setae and short, spini- 
form anterior seta in middle series and 4— 
6 posterior setae in distal series (Fig. 8c). 
Endite 2 similar to distal series of endite 1, 
with 13-18 posterior setae and 2 anterior 
setae. Appendage 11 with posterior setae 
reduced to 2-4. Endites 3, 4, and 5 with 3, 
2, and 2 posterior setae, respectively (Figs. 
8 & 12a). Endites 3 and 4 with 2 anterior 
setae on appendages 2-11 and 4—7 anterior 
setae on appendage 1. Endite 5 with 2-6 
anterior setae on appendages 2—11 and 4-8 
on appendage |. Endopodite large, elongate, 
shaped like broad scimitar (Fig. 8). Setae 


along medial edge with several rows of fine 
setules surrounding distal 0.67 (Fig. 12b), 
becoming comblike toward ventral edge, 
along ventral edge, long and thin, with 2 
rows of fine setules. Exopodite oval to tri- 
angular, surrounded by setae resembling 
those of ventral edge of endopodite. Epi- 
podite smooth, inflated, without setae. 
Preepipodite thin, semicircular, with 
coarsely toothed edge, reduced on append- 
age 11. Early in development with 2 lobes, 
in adults with slight notch along lateral edge. 
Genital segments only slightly expanded, 
paired papillae dorsal, paired penes arising 
ventrolaterally. Non-rectractile portion of 
penis with ventral, fleshy lobe and medially- 
directed, sclerotized spur (Fig. 5a). Distal 
portion of penis eversible with two small, 
sclerotized lobes on lateral surface each with 
six to ten pyramidal teeth (Fig. 5b). 
Post-genital segments with paired papil- 
lae in following positions: one, dorsal, two, 
ventral, three, dorsal, four, lateral, five and 


300 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


» \ 
v \ 
— | 
é J 
SN yy), 
— 4, 
iS Li 
YY 
a a a 
is \ Nats > 
\ \ N a 
) N\ A> 
Via 
ie 
i Ne 
ay \ 
SF 
— 
bh Hy 


NA A . 
X i LA (LAA O) . 


Figs. 7-8. Branchinecta sandiegonensis, new species, 7a—c. thoracic appendages female. a. appendage 1. b. 
appendage 5. c. appendage 11. 8a-c. thoracic appendages male. a. appendage 1. b. appendage 5. c. appendage 
11. (scale bar 0.1 mm) 


VOLUME 106, NUMBER 2 


Pleccaease ie 


S53 SS 


KE 


ae 


a 


Fig. 9. Branchinecta sandiegonensis, new species female, a. dorsal view. b. lateral view. (scale bar 1.0 mm) 


six, dorsal. Cercopods on anal segment 
bearing setae with Z rows of fine setules (Co- 
hen 1983). 

Length of mature individuals, from front 
of head to end of anal segment, excluding 
cercopods, 9.0—16.0 mm. 

Female.—Head similar to that of male, 
except for pair of cuticular papillae on dor- 
sal surface, posterior to mandibular crease, 
and form of antenna 2 (Fig. 9). Antenna 2 
cylindrical, approximately 3 times as long 
as wide basally, gradually tapering to 5 times 
as long as wide at 0.8 its total length, then 
rapidly to sharp point (Fig. 9b). Anterior 
surface with 2 patches of sensory setae, each 
seta borne on a cuticular papillus. 

Thorax with cuticular papillae in same 
locations as male, but segments 3 and 5-8 
with 2 pairs of dorsolateral spines arranged 
above and below papillus, dorsal normally 
smaller than lateral spines, segment 4 with 
1 pair of large bilobed spines (Fig. 9). Tho- 
racic appendages similar to male, but en- 
dopodite lobelike (Fig. 7a—c) with stout, 
comblike setae along entire medial edge (Fig. 
12c). 


Genital segments slightly inflated, with 
paired papillae lateral on segment 1 and 
dorsal on 2 (Fig. 9a). Ovisac fusiform, on 
average ending under post-genital segment 
4, occasionally under segment 5. Paired 
ovaries t-shaped, extending from thoracic 
segment 9 or 10 to post-genital segment 4, 
and into ovisac at junction of two genital 
segments, forming an oviductal pouch. 
Resting eggs spherical, diameter «209 
um, SD = 18 um, range = 227-309 um, n 
= 100, with numerous shallow hemispher- 
ical depressions, approximately 40 um in 
diameter, covering surface (Fig. 13a, b). 

Post-genital segments similar to male, but 
arrangement of papillae as follows: one, lat- 
eral, two, ventral, three, dorsal, four, ven- 
tral, five, dorsal, six, lateral. 

Length of mature individuals, from front 
of head to end of anal segment, excluding 
cercopods, 8.0—15.0 mm. 

Remarks. —The form of the male second 
antenna has served as the cardinal character 
for species identification in the genus and 
although all males are currently distinguish- 
able on that basis, the use of that character 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 10-13. Branchinecta sandiegonensis, new species, 10a, b. antenna 2 male. a. pulvillus on basal segment. 
b. tip of distal segment, dorsolateral view. 1la, b. mandibles male. a. left mandible, molar surface. b. mght 
mandible, transition between posterodorsal and anteroventral regions. 1 2a—c. setae on medial surfaces of thoracic 
appendages. a. appendage 1 male, endites 3-5. b, c. medial edge of endopodite. a. male. b. female. 13a, b. resting 
egg. a. whole egg. b. surface detail. (scale bars, la, b, 3a, 4a: 0.1 mm, 2a, b, 3b, c, 4b: 0.01 mm) 


VOLUME 106, NUMBER 2 


alone can often lead to difficulties. Bran- 
chinecta sandiegonensis has been reported 
as Branchinecta lindahli Packard, 1883, a 
common species throughout western North 
America, by Ebert & Balko (1987) ina study 
of the vernal pools on Kearney Mesa, San 
Diego County, California (D. Belk, pers. 
comm.; vouchers in personal collection of 
D. Belk) and by Dr. J. E. Lynch in two ear- 
lier collections from San Diego County 
(USNM 305974, 3 Mar 1962, Poway, 
32°55'N, 117°04’W; USNM 294523, 4 Mar 
1962, Ramona, 33°02’N, 116°52’W). Five 
additional species in the genus are also 
known from southern California, but of 
those only Branchinecta lynchi Eng et al., 
1990 is likely to be confused with B. san- 
diegonensis. Branchinecta mackini Dexter, 
1956 and Branchinecta gigas Lynch, 1937 
are found in large playa lakes in the Mojave, 
while Branchinecta conservatio Eng et al., 
1990 and Branchinecta longiantenna Eng et 
al., 1990, although found in vernal pools, 
are readily distinguished morphologically. 
In general form, B. sandiegonensis females 
resemble those of B. lindahli due to the fu- 
siform shape of the ovisac, however that of 
B. lindahiiis slightly longer, typically ending 
under post-genital segment 5. Branchinecta 
lindahli differs also in having a longer ovary, 
extending from thoracic segment 4—7 to 
post-genital segment 4, an egg with hemi- 
spherical surface depressions, approximate- 
ly 20 um in diameter, and a single dorso- 
lateral spine, always below the papillus, on 
each side of thoracic segments 3-11. Bran- 
chinecta lynchi females share a similar ova- 
ry length, egg surface (Mura 1991), and dor- 
solateral spine pattern with OB. 
sandiegonensis, but differ in having a short, 
conical ovisac, typically ending under post- 
genital segment 3. 

Males of B. sandiegonensis share the large, 
oval pulvillus on the basal segment of the 
second antenna with B. lindahli, but the me- 
dial series of spines is less diffuse. The distal 
segment is broadest in B. sandiegonensis just 
proximal to the tip, while that of B. lindahli 


303 


is at 0.75 the distance from base. The breadth 
at the tip in B. lindahiiis only slightly larger 
than at the joint with the basal segment and 
the entire tip is bent medially at a right angle 
to the segment of the antenna just preceding 
it (Shantz 1905, Lynch 1964). The second 
antenna of B. /ynchi is quite different; the 
pulvillus is smaller, there is a small apoph- 
ysis, Slightly distal and posterior to the pul- 
villus, the distal segment has a narrower 
breadth, and the entire tip is bent medially 
(Eng et al. 1990). 

Distribution and habitat—Branchinecta 
sandiegonensis is found after winter rains in 
vernal pools from northern Baja California 
Norte, Mexico (Valle de las Palmas, Baja 
California Norte, Mexico 32°28'N, 
116°37'W, 15 Nov 1987, soil sample, coll. 
M. Fugate & G. Pratt) to Santa Barbara, 
California (Isla Vista, Santa Barbara Coun- 
ty, California 34°24'N, 119°51'W, 1, 7 Apr 
1991, coll. J. Kornmeyer). Its current range 
is centered in San Diego County, California 
(Del Mar Mesa, Kearney Mesa, Ramona, 
Otay Mesa, 32°34'N, 116°58’W). All known 
localities are within 50 km of the Pacific 
Ocean and at elevations less than 700 m. 
There are no records from Los Angeles and 
Orange counties and it is unknown if the 
Santa Barbara population is either disjunct 
or the northern end of a formally continu- 
ous distribution. 

The pools in San Diego County are shal- 
low (<30 cm) and often on chaparral cov- 
ered mesas (Greenwood 1984, Ebert & Bal- 
ko 1987). Zedler (1984) found 46 plant 
species common in pools at Kearney Mesa, 
one of which is endangered (Pogogyne 
abramsii) and Ebert & Balko (1987) found 
B. sandiegonensis associated with six cla- 
docerans, one copepod, nine ostracods and 
at least 21 rotifer species at the same site. 
Streptocephalus woottoni Eng et al., 1990 
and B. lindahli have subsequently been 
found at Kearney Mesa, the former in a pool 
containing B. sandiegonensis and the latter 
in a road rut, but not in pools containing B. 
sandiegonensis (M. Simovich, pers. comm.). 


304 


Branchinecta sandiegonensis has been found 
in a disturbed pool with B. lindahli at Del 
Mar Mesa and in two pools with S. woottoni 
and aclam shrimp, CyZicus sp. at Otay Mesa. 


Acknowledgments 


Marie Simovich and her students sup- 
plied me with many field-collected speci- 
mens and soil samples. Clay Sassaman pro- 
vided financial support, lab space, criticism 
and encouragement. Denton Belk provided 
useful comments on the text and figure pre- 
sentations and examined his collections of 
material from San Diego County, Califor- 
nia. Marcia Hartz and Chris Williams as- 
sisted with and the Chancellor’s Patent Fund 
at the University of California, Riverside 
provided funds for the electron microscopy. 
Three anonymous reviewers substantially 
improved the manuscript. 


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Crampton, B. 1976. A historical perspective on the 
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Dey, S., T. S. Basu Baul, B. Roy, & D. Dey. 1989. A 
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of Ecology Publication No. 28, University of 
California, Davis, 280 pp. 

Holland, R. F., & S. K. Jain. 1977. Vernal pools. Pp. 
515-533 in M. Barbour & J. Major, eds., Ter- 
restrial vegetation of California. John Wiley & 
Sons, New York, 1002 pp. 

Linder, F. 1941. Contributions to the morphology 
and the taxonomy of the Branchiopoda Anos- 
traca.— Zoologiska Bidrag fran Uppsala 20:101- 
302 + 1 plate. 

Lynch, J. E. 1960. The fairy shrimp Branchinecta 

campestris from northwestern United States 

(Crustacea: Phyllopoda).—Proceedings of the 

United States National Museum 112:549-561. 

. 1964. Parkard’s and Pearse’s species of Bran- 

chinecta: analysis of nomenclatural involve- 

ment.—American Midland Naturalist 71:466- 

488. 

Mura, G. 1991. SEM morphology of resting eggs in 
the species of the genus Branchinecta from North 
America.—Journal of Crustacean Biology 11: 
432-436. 

Shantz, H.L. 1905. Notes on North American species 
of Branchinecta and their habitats. — Biological 
Bulletin 9:249-263 + plates X—XII. 

Tyson, G. E., & M. L. Sullivan. 1979. Antennular 

sensilla of the brine shrimp, Artemia salina. — 

Biological Bulletin 156:382-392. 

. 1981. A scanning electron microscopic study 

of the molar surface of the mandibles of the 

brine shrimp (Cl. Branchiopoda: Or. Anostra- 

ca).—Journal of Morphology 170:239-251. 

Zedler, P.H. 1984. Micro-distribution of vernal pool 
plants at Kearney Mesa, San Diego County. Pp. 
185-197 in S. Jain & P. Moyle, eds., Vernal 
pools and intermittent streams. Institute of 
Ecology Publication No. 28, University of Cal- 
ifornia, Davis, 280 pp. 


Department of Biology, University of 
California, Riverside, California 92521, 
U.S.A.; (Current Address) Department of 
Biology, University of Oregon, Eugene, Or- 
egon 97403, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(2), 1993, pp. 305-314 


THE FIRST FINDING OF THE MALE OF 
THAUMATOCYPRIS ECHINATA MULLER, 1906 
(CRUSTACEA: OSTRACODA) 


J. A. Rudjakov 


Abstract.—Two adult males and two juveniles of Thaumatocypris echinata 
Miiller, 1906 were found in the near-bottom layer off the west coast of Mad- 
agascar at the depth of 180-1220 m. The first description of male morphology 
is given. The sexual dimorphism manifests itself in the structure of mandibular 
endopodite instead of endopodite of second antenna as it was known for other 
halocypridids previously. The contents of the gut suggest that the species is 


either a carnivore or scavenger. 


The myodocopid ostracod genus Thau- 
matocypris with only species T. echinata was 
established by Miiller (1906) on six females 
derived from a depth of 1100 m near In- 
donesia. Since then, only one female of the 
type species was found nearly in the same 
region at the depth of about 2000 m (Poul- 
sen 1969). 

Two other living genera of the family 
Thaumatocyprididae, all benthic— Thau- 
matoconcha (eight species) and Danielopo- 
lina (six species) were thoroughly investi- 
gated by Kornicker & Iliffe (1989a, 1989b) 
and by Kornicker & Sohn (1976). The latter 
authors also used the opportunity to rein- 
vestigate the specimen of Thaumatocypris 
echinata described by Poulsen (1969). 

Plankton investigation in the near-bot- 
tom layer of the Indian Ocean off the west 
coast of Madagascar (17th trip of R/V Vi- 
tyaz) revealed a few specimens of the genus 
Thaumatocypris. The Madagascar speci- 
mens (two males and two juveniles) were 
preliminarily studied by Dr. Louis S. Kor- 
nicker (National Museum of Natural His- 
tory, Smithsonian Institution, Washington, 
D.C.), who returned them to me with the 
kind suggestion to prepare the first male de- 
scription. Two males have been deposited 
to the U.S. National Museum of Natural 
History and given USNM numbers. 


Materials and Methods 


Ostracods were collected in the 17th cruise 
of R/V Vityaz by the towing underwater 
apparatus “SOUND” equipped with open- 
ing/closing plankton sampler (Biryukov et 
al. 1990). Observations are briefly sum- 
marized below. 

Station (St.) 2649; 3 Dec 1988; 0206-0306 
h; 22°25’S, 43°00”E; 970-950 m; 30 m above 
bottom. 1 male USNM 194110 (1.16 mm), 
1 juvenile specimen (0.65 mm). 

St. 2655:°3°) Dec’ 1988: 1917—2017 h; 
DBS: WARES As Ot 21220 rena k: >) an 
above bottom. | juvenile specimen (0.72 
mm). 

St. 2661; 4 Dec 1988; 1200-1230 h; 
22°13'S, 43°07’E, 260-180 m; 30 m above 
bottom. 1 male USNM 194116 (1.20 mm). 

The specimens were preserved with form- 
aldehyde and after about two-years storage 
transferred into alcohol. The males were 
dissected in water diluted glycerol. Ap- 
pendages were placed on slides moistened 
with Faurés liquid diluted 3-4 times with 
water. After hardening of the fluid with the 
appendages mounted in a necessary order 
and position, a drop of molten glycerol-gel- 
atin was deposited on a membrane formed 
on the fluid, and a cover glass added (Rud- 
jakov 1968). Separated valves were mount- 


306 


ed on the shallow-well slides with glycerol- 
gelatin. 


Results 


The following description of the adult 
male was mostly prepared using the speci- 
men from St. 2661, supplemented with the 
features of the specimen from St. 2649 when 
necessary. 


Thaumatocypris echinata Miller, 1906 
Figs. la—d, 2a—c, 3a—d, 4a-f, 5a, b 


Thaumatocypris echinata Miller, 1906:42, 
pl. 6, figs. 1-18; 1912:54 [listed].—Poul- 
sen, 1969:7, fig. 1.—Kornicker & Sohn, 
1976:34, figs. 14c, d, 15. 


Description. —Shell (Fig. la). Surface 
smooth, without discernible anterior ridges 
and serration along posterior ridge. Each 
valve with short upper and long lower tube- 
like anteroventral horns, appearing broken. 
The shell can firmly stand on its four horns, 
anterior side down, with furcal lamellae just 
above ventral pair of horns and with anten- 
nal and mandibular bristles protruding be- 
tween dorsal and ventral pairs of horns. 
Postero-dorsal ridge of right valve with short 
rounded tubercle (hardly discernible on the 
male from St. 2661). Antero-ventral margin 
of each valve with hairs distally bifurcate 
or split into several branches. Indistinct ad- 
ductor muscle attachment scar (Fig. 1b) does 
not form radial pattern (patterns of left and 
right valves seem to differ significantly). Scar 
consists of about 9-18 muscle segments. 
Length 1.16—1.20 mm, height 0.90-1.05 mm 
(without horns). 

First antenna (Fig. 1d): 8-jointed, third 
and fourth joints being not clearly separat- 
ed. First joint without spines and hairs, with 
1 bare dorsal bristle about as long as ventral 
side of second through fourth joints com- 
bined and | proximally spinous lateral bris- 
tle, as long as 6 distal joints together, di- 
rected dorsally. Second joint with hairs 
dorso-laterally and proximo-ventrally, with 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


1 ventral bristle reaching to eighth joint or 
exceeding it and | dorsal bristle reaching to 
fifth joint, both bristles without discernible 
spines. Third joint with long hairs on dorsal 
and ventral sides, its dorsal margin some- 
what shorter than dorsal margin of fourth 
joint. Fourth joint with ventral group of hairs 
and 2 spinous disto-ventral bristles, short 
bristle reaching or exceeding eighth joint and 
long bristle as long as total length of dorsal 
margin of 7 distal joints. Fifth joint with 
ventral group of hairs and 3 bristles on dis- 
to-ventral corner, | bristle (not the longest) 
medially with short hairs in proximal part, 
long bristle about 2.2—2.8 times total length 
of 7 distal joints, others about 2 times their 
total length. Sixth joint with dorsal group 
of hairs, without bristles. Seventh joint with 
1 short dorsal spinous bristle as long as com- 
bined length of 3—4 distal joints and with 2 
long disto-ventral bristles, the longest of 
them being about 3.1-—3.5 times total length 
of 7 distal joints. Eighth joint with two bris- 
tles, dorsal bristle covered with stout small 
spines along dorsal margin being about 3.9— 
4.3 times total length of 7 distal joints. 
Second antenna: Protopodite with cluster 
of hairs in proximo-ventral area, without 
bristles. Exopodite 9-jointed, first joint di- 
vided into long proximal and short distal 
parts. Joints 2-8 each with 1 long bristle 
with natatory hairs. Ninth joint with 2 bris- 
tles, short and long, both devoid of natatory 
hairs. Endopodite 2-jointed (Fig. lc), but 
very short distal part of second joint may 
be interpreted presumably as third joint. 
First joint with long hairs, 1 ventral and 2 
dorsal bristles, ventral bristle with marginal 
spines. Second joint with 2 transverse rows 
of hairs, | lateral bristle bearing spines along 
ventral margin and 2 pre-terminal bristles. 
Terminal part of second joint (or third joint) 
with 2 bristles, ventral one being the longest. 
Mandible (Fig. 2a—c): Coxale endite seems 
to have the same character set as Thau- 
matoconcha radiata Kornicker et Sohn, 
1976: proximal set of teeth with four broad 
teeth, some of them bifurcate. Distal set with 


VOLUME 106, NUMBER 2 


two large flat teeth bearing several cusps and 
distally flattened spinous bristle. Basale with 
knife-like process and 11 bristles, which can 
be subdivided into 4 groups: 3 posterior 
bristles, 1 anterior bristle, 5 lateral bristles 
and 2 medial bristles, the longest of the lat- 
ter with long hairs. First endopodite joint 
with 1 dorsal bristle, second joint with 4 
bristles on ventral margin and 2 bristles on 
dorsal margin. Third endopodite joint with 
7 bristles, the longest bristle with ca. 10-12 
long spines directed distally in the middle 
part of its posterior side, with ca. 80-90 
short spines along posterior side and sparse 
short spines along anterior side more dis- 
tally. Shorter of the 2 stouter bristles seems 
wavy bent with long spine-like processes (ca. 
10 along each side) in the middle and with 
very short fine spines more distally. Distal 
half of the longest bristle of the terminal 
joint with fine short spines. 

Maxilla (Fig. 3a—d): First endite with 1 
proximal and 7 distal bristles, second endite 
with 7-9 bristles, third endite with 6 bristles 
distal of which being not partitioned off the 
endite body. Basale with 1 long dorsal (cov- 
ered with long hairs along ventral side) and 
1 shorter lateral bristles. There is 1 lateral 
bristle between basale and first endopodite 
joint. First endopodite joint with long hairs 
bearing 5 dorsal bristles and 2-3 disto-lat- 
eral bristles. Second endopodite joint with 
5-6 bristles, of which 1 distal bristle being 
claw-like and not clearly partitioned off the 
joint. 

Fifth limb (Fig. 4a, b): Epipodial append- 
age with 14 bristles arranged in 3 groups of 
5 (dorsal), 5 and 4 (ventral). Protopodite 
and endopodite with total of 19 bristles in 
3 indistinct groups each with 6 (proximal), 
7 and 6 (distal) bristles. First exopodite joint 
hirsute with 1 long disto-dorsal bristle and 
9 ventral bristles. Second joint with hairs 
and 2 midventral bristles. Third joint with 
3 terminal bristles, length of the shortest of 
them (ventral) being about 36% of others. 

Sixth limb (Fig. 4c): Epipodial appendage 
with 15 bristles arranged in groups of 6 (dor- 


307 


sal), 4 and 5 (ventral) bristles. Hirsute pre- 
coxale, coxale and basale each with 2 ven- 
tral bristles. Disto-dorsal process of basale 
(endopodite?) with 1 long bristle and small 
spine. First exopodite joint with 2 disto- 
ventral bristles, second with 1 dorsal and 2 
midventral bristles. Third joint with 3 ter- 
minal bristles, length of the shortest of them 
(ventral) being about 39% of others. 

Seventh limb (Fig. 4d): With 2 finely spi- 
nous bristles of nearly equal length. 

Copulatory organ (Fig. 4e, f) consists of 
elongate anterior part and tapered posterior 
part with 3 hardly discernible transparent 
hair-like appendices. Anterior part with a 
single long tooth-like process not reaching 
distal part of the organ. 

Lip morphology (Fig. 5a) is consistent with 
family diagnosis given by Kornicker & Sohn 
(1976). 

Rod-shaped organ not discernible. 

Furca (Fig. 5b): Each lamella with 2 long 
anterior claws separated from lamella, fol- 
lowed by 6 short spinous claws joined to 
lamella and by 1 bare triangular process ori- 
ented like preceding claws (the latter is ab- 
sent on right lamella of the specimen from 
St. 2649). 

Posterior of body with hook-like process 
proximal to furcal lamellae oriented pos- 
teriorly. 


Discussion 


A comparison of the described specimens 
of T. echinata with descriptions and draw- 
ings by Muller (1906) and Poulsen (1969) 
and with supplemental description and 
summary of characters given by Kornicker 
& Sohn (1976) reveals a set of differences 
(Table 1). Only one of them may be attrib- 
uted to sexual dimorphism for certain: the 
number of ventral bristles of fourth joint of 
first antenna (as in the other Thaumatocy- 
pridid genera). Some differences may be 
sexually dimorphic: presence of the lateral 
bristle of first antenna’s first joint, the bristle 
number of first antenna’s seventh joint, the 


308 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


500 Pree! Sb 
Sep eet ees 


Fig. 1. Thaumatocypris echinata Miller, 1906, adult male: a, Outside view of right valve of the specimen 
from St. 2649; b, Adductor muscle-scar of the right valve of the specimen from St. 2649; c, Endopodite of left 
2nd antenna of the specimen from St. 2661, medial view; d, Left 1st antenna of the specimen from St. 2661, 
lateral view. (Scale in micrometers.) 


309 


VOLUME 106, NUMBER 2 


LT} 
LTT coe 
CO 


LTH 
LT 
— ~ 
~~ => 


ees 


Fig. 2. Thaumatocypris echinata Miller, 1906, adult male: a, 3rd endopodite joint of right mandible of the 
specimen from St. 2649, lateral view; b, Basale of right mandible of the specimen from St. 2661, lateral view; 
c, Endopodite of right mandible of the specimen from St. 2661, 3rd joint missing, medial view. (Scale in 


micrometers.) 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


310 


C 
Cie 
Ss hy 
Sa 
S | 
Y [| 
~ Bisg 4 
L] 
4 ERA HE 
— iy BS 
ay CS ea ane 
= i 0 an | ge H 
ie eel ay 
| ] : E | 
@ | 
/ | \ \ 3 
Thaumatocypris echinata Miller, 1906, adult male, St. 2661: a, Basale and endopodite of left maxilla, 


Pig. 3: 
lateral view (R—bristle found on right limb only); b, 1st endite of right maxilla, medial view; c, 2nd endite of 
right maxilla, medial view (L—bristle found on left limb only); d, 3rd endite of left maxilla, lateral view. (Scale 


in micrometers.) 


VOLUME 106, NUMBER 2 31 


100 


Fig. 4. Thaumatocypris echinata Miiller, 1906, adult male: a, Distal joint of 5th limb of the specimen from 
St. 2649; b, 5th limb of the specimen from St. 2661, distal bristles missing; c, 6th limb of the specimen from 
St. 2661 (dotted lines denote folds which can be erroneously interpreted as sutures between joints); d, 7th limb 
of the specimen from St. 2661; e, Copulatory organ of the specimen from St. 2661; f, Tip of copulatory organ 
of the specimen from St. 2661. (Scale in micrometers.) 


armament of male mandibular endopodite suggestions are not supported by the sexual 
terminal bristles (on two longest bristles the dimorphism in other genera of Thaumato- 
male has spines in place of hairs drawn for cypridids (Kornicker & Sohn 1976, Kor- 
females by Miller and Poulsen) though these _ nicker & Iliffe 1989). Some differences may 


312 


, 20 | 


\ 


Fig. 8: 
crometers.) 


prove to be a result of individual damage 
or variability (total number of bristles on 
mandible basale, of second joint of man- 
dible endopodite or of first joint of fifth limb 
exopodite). Some bristle groups are difficult 
to count: epipodite bristles, bristles of max- 
illae coxale and of fifth limb. In many other 
instances there is much more similarity of 
the male with the female described by Poul- 
sen and reinvestigated by Kornicker & Sohn 
than with Miuller’s non-adult female, though 
judging by Muller’s drawing and by his 
specimen length (1.55 mm) it could not be 
younger than the A-1 developmental stage 
of Kornicker & Sohn (1976). 

As it follows from the comparison above 
there are no morphological differences of 
Thaumatocypris specimens investigated so 
far which could be treated as important on 
the species level. The length of the males 
found is smaller than one could predict on 
the basis of Muller’s and Poulsen’s female 
measurements (1.55 and 1.4 respectively) 
and male: female length ratio (1.06) deter- 
mined by Kornicker & Sohn for Thauma- 
toconcha radiata. But the ratio may be quite 
different for Thaumatocypris and geograph- 
ical variability also may prove to be quite 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


J 


Thaumatocypris echinata Miiller, 1906, adult male, St. 2661: a, Upper lip; b, Furca. (Scale in mi- 


significant. The latter may be the situation 
for the taxonomical importance of the pos- 
terodorsal process of the right valve in the 
males studied. Dr. Louis S. Kornicker 
pointed out in his letter reviewing the 
manuscript that “*. . . there does exist some 
difference in the endopodite of the male and 
female 2nd antenna in that Poulsen and 
Miller show it to be truncate. .. .”’ This ob- 
servation is true but the difference in ques- 
tion may be attributed to sexual dimor- 
phism. These considerations form the basis 
for referring the males described to Thau- 
matocypris echinata, but the final species 
identification will become possible after 
finding and investigation of Thaumatocy- 
pris females from the Madagascar area. 

The diagnosis of the genus Thaumato- 
cypris given by Kornicker & Sohn (1976:34) 
should be changed in several respects to in- 
clude adult male and an emended diagnosis 
follows: 

Each valve with upper and lower long an- 
teroventral protuberances. First antenna: 
first joint with O—1 lateral bristles; seventh 
joint with 2-3 bristles, 1—2 ventral, 1 dorsal; 
eighth joint with 2 bristles. Second antenna: 
first endopodial joint with 3 bristles, 1 ven- 


VOLUME 106, NUMBER 2 


Table 1.—Main morphologicai differences of Thau- 
matocypris specimens known so far (number of bristles 
if other not indicated). 


Female 
Poulsen Miller 
Character 1969 1906 Male 
First antenna 
Ist joint: lateral 0 0 1 
4th joint: ventral 1 1 2 
6th joint ] 0 0 
7th joint: ventral 1 1 2 
Mandible 
Basale 13 12? 11 
Endopodite 
2nd joint: ventral 4 3 4 
2nd joint: dorsal 2 3 2 
3rd joint 7 6 7 
Makxilla 
Basale: dorsal 0 1? 1 
Basale: ventral 1 2? 1 
Endopodite 
lst joint: anterior 4 5 5 
Ist joint: posterior 2 2 2-3 
Fifth limb 
Epipodite 3 nd 14 
Protopodite + 
endopodite 17 nd 19 
Exopodite 
Ist joint: ventral 8 nd 9 
Sixth limb 
Epipodite car 12 nd 15 
Furca 
(no. of short claw- 
like processes) y | 7 6-7 


tral, 2 dorsal; terminal endopodial joint of 
male without hook-like process. Mandible: 
one of male endopodite terminal bristles 
wavy bent and with long spine-like pro- 
cesses in the middle. Maxilla: second en- 
dopodite joint with 6 bristles. Fifth limb: 
second exopodial joint without terminal 
bristle on ventral margin; third exopodial 
joint with 3 bristles. Sixth limb: process on 
dorsal corner of first exopodial joint with 1 
bristle and 1 minute spine; third exopodite 
joint with 3 bristles. Rod-shaped organ is 
minute cone-shaped or not discernible. 
The most striking peculiarity of the spe- 
cies is the absence of the sexual dimorphism 
in the endopodite structure of the second 


313 


antenna: there is no hook-like process typ- 
ical for other halocypridinid males. Instead, 
the sexual dimorphism manifests itself in 
the mandibular endopodite terminal bristle 
morphology, not known in the other genera 
of the Halocyprida order. In other respects 
Thaumatocypris males do not differ from 
other Thaumatocyprididae genera. For ex- 
ample, the copulatory organ of 7. echinata 
differs from those described by Kornicker 
& Sohn (1976:38, fig. 18) for Thaumato- 
concha, but the extent of the difference is 
within the range of variability within the 
latter genus. 

The contents of the gut (crustacean-like 
claws) suggest that the species is either a 
carnivore or scavenger. The species has not 
been found in the vertical plankton catches 
in the localities where the “SOUND” ap- 
paratus tows were made. Therefore the spe- 
cies may be attributed presumably to the 
near-bottom fauna as it was described by 
Heinrich & Rudjakov (1991). 


Acknowledgments 


I thank Dr. Louis S. Kornicker and anon- 
ymous reviewers who offered valuable sug- 
gestions for improvement of the manu- 
script. Thanks to the crew and scientific team 
ofthe R/V Vityaz for help collecting. Special 
thanks to A. K. Heinrich, my wife and son 
who helped with the figures. Manuscript 
preparation was supported by the Soros 
Foundation. 


Literature Cited 


Biryukov, S. G., V. N. Maryatkin, A. S. Matveev, V. 
A. Popov, & J. A. Rudjakov. 1990. Experience 
of usage of towed underwater apparatus 
“SOUND” for study of near-bottom plank- 
ton.—Okeanologiya 30:152-156 (in Russian, 
English summary). 

Heinrich, A. K., & J. A. Rudjakov. 1991. Vertical 
distribution of plankton animals in the near- 
bottom layer and the biological structure of the 
ocean. —Okeanologiya 31:146—150 (in Russian, 
English summary). 

Kornicker, L. S., & T. M. Iliffe. 1989a. New Ostra- 
coda (Thaumatocyprididae, Halocyprididae) 
from anchialine caves in the Bahamas, Palau, 


314 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


and the Yucatan peninsula, Mexico.—Smith- 
sonian Contributions to Zoology 470:1-47. 

——,,& . 1989b. Troglobitic Ostracoda (Cy- 
pridinidae, Thaumatocyprididae) from anchia- 
line pools on the Galapagos Islands.—Smith- 
sonian Contributions to Zoology 483:1-38. 

—, & I. G. Sohn. 1976. Phylogeny, ontogeny, 
and morphology of living and fossil Thauma- 
tocypridacea (Myodocopa, Ostracoda).— 
Smithsonian Contributions to Zoology 219:1- 
24: 

Miller, G. W. 1906. Ostracoda.— Wissenschaftliche 
Ergebnisse der Deutschen Tiefsee-Expedition auf 
dem Dampfer “Valdivia” 1898-1899 8:1-128 
+ pls. 1-31. 


. 1912. Ostracoda.—Das Tierreich 31:i—xxxili 

+ 1-434, figs. 1-92. 

Poulsen, E. M. 1969. Ostracoda—Myodocopa, 3A: 
Halocypriformes—Thaumatocypridae and Hal- 
ocypridae. — Dana-Report 75:1-100. 

Rudjakov, J. A. 1968. A method of making perma- 

nent preparations of small Arthropoda.—Zoo- 

logichesky Zhurnal 47:453—454 (in Russian, En- 
glish summary). 


P. P. Shirshov Institute of Oceanology, 
Russian Academy of Sciences, 23 Krasikov 
St., Moscow 117218, Russia. 


PROC. BIOL. SOC. WASH. 
106(2), 1993, pp. 315-324 


PSEUDONICOTHOE BRANCHIALIS (CRUSTACEA: 
COPEPODA: SIPHONOSTOMATOIDA: NICOTHOIDAE), 
LIVING ON THE PANDALID SHRIMP 
HETEROCARPUS SIBOGAE 
OFF NORTHWESTERN AUSTRALIA 


Arthur G. Humes and Geoffrey A. Boxshall 


Abstract. — Both sexes of Pseudonicothoe branchialis (Siphonostomatoida: Ni- 
cothoidae), living on the marine shrimp Heterocarpus sibogae off the coast of 
northwestern Australia, are described, the male for the first time. Both sexes 
have hyaline flaps (suckers?) on the exopods of legs 1-3. The innermost element 
on the free segment of leg 5 is sexually dimorphic. Pseudonicothoe is recognized 
as a valid genus, to which Paranicothoe procircularis (Carton) is transferred as 


a new combination. 


The siphonostomatoid copepod family 
Nicothoidae Dana, 1852, includes several 
genera whose species live on decapod crus- 
taceans. Except Choniostoma Hansen, 1897, 
all genera living on the decapods belong to 
the Nicothoe group, as recognized by Box- 
shall & Lincoln (1983). These genera are 
Nicothoe Audouin & Milne Edwards, 1826, 
Choniosphaera Connolly, 1929, Choniomy- 
zon Pillai, 1962, Paranicothoe Carton, 1970a 
(see also 1970b), Hadrothoe Humes, 1975, 
and Pseudonicothoe Avdeev & Avdeev, 
1978. In Nicothoe, Choniosphaera, Choni- 
omyzon, and Hadrothoe, the prosome of the 
female is swollen. Only Paranicothoe and 
Pseudonicothoe have a flattened prosome in 
the female. 

All species of Paranicothoe live on the 
gills of penaeid and pandalid shrimps in 
warm regions of the Indo-Pacific. As men- 
tioned by Boxshall & Lincoln (1983), two 
species of Paranicothoe, P. procircularis 
(Carton, 1967) (see also Carton, 1970b) and 
P. cladocera Carton, 1970a, may parasitize 
epicaridean isopods which are in turn par- 
asitic on shrimps. Paranicothoe procircu- 
laris parasitizes Pseudione affinis (Sars) 
(Isopoda) which lives in the branchial cavity 
of the pandalid shrimp Plesionika ensis (A. 


Milne Edwards) in the Java Sea. Parani- 
cothoe cladocera inhabits the brood cavity 
of Orbione natalensis Bourdon (Isopoda) 
found in the gill cavity of the penaeid shrimp 
Hymenopenaeus triarthrus Stebbing, in the 
Mozambique Channel, off Natal, South Af- 
rica. 

Pseudonicothoe branchialis Avdeev & 
Avdeev, 1978, lives on the gills of the pan- 
dalid shrimp Heterocarpus laevigatus Bate 
in the Marshall Islands. 

The purpose of this paper is to describe 
the male of Pseudonicothoe branchialis for 
the first time, and to redescribe certain fea- 
tures of the female. 


Siphonostomatoida Thorell, 1859 
Nicothoidae Dana, 1852 
Pseudonicothoe Avdeev & Avdeev, 1978 


Redescription of the genus Pseudonico- 
thoe, based on both sexes: Body cyclopi- 
form, flattened, relatively unmodified. So- 
mite bearing leg 1 fused with cephalosome. 
Urosome in female 5-segmented, in male 
6-segmented. Caudal ramus with very long 
inner terminal seta. 

Rostrum weakly developed. Antennule 
11-segmented. Antenna 4-segmented with 
small exopod bearing | seta. Siphon short 


316 


with terminal sucking disk. Mandible slen- 
der blade. Maxillule with 2 lobes. Maxilla 
small, strongly sexually dimorphic. Maxil- 
liped 5-segmented with terminal claw. Ven- 
tral surface of cephalosome sexually di- 
morphic, with maxillules and maxillae much 
more widely separated in female than in 
male, and with transverse arched scleroti- 
zation in front of maxillae in female. 

Legs 1-4 with 3-segmented rami. Leg 1 
with inner spine on basis. Second and third 
segments of exopods of legs 1-3 in both 
sexes with small round hyaline flaps. Second 
segment of endopods in legs 1—3 with | in- 
ner seta but this segment in leg 4 with 2 
such setae. 

Leg 5 with elongate free segment bearing 
4 setae in both sexes, but innermost of these 
setae sexually dimorphic, short and spini- 


form in male, but long and setiform in fe- 


male. 


Pseudonicothoe branchialis 
Avdeev & Avdeev, 1978 


Material. —2 22, 5 66, 3 copepodids from 
the pandalid shrimp Heterocarpus sibogae 
de Man, in shrimp trawl in 392-400 m, 
FRV Soela, station NWS-7, 18°33.2'S, 
117°30.9’E, 25 Apr 1983. One adult 92, 3 
adult 66, and 3 copepodids (2 99, 1 4) de- 
posited in the Northern Territory Museum, 
Darwin, Northern Territory, Australia; 1 9, 
1 6, and 1| dissected 6in The Natural History 
Museum, London, England, BM(NH) Reg. 
Nos. 1992.1067-1069. 

Male. — Body (Fig. 1a) with flattened pro- 
some subcircular in dorsal view. Length 1.20 
mm (1.05—1.32 mm) and greatest width 0.61 
mm (0.51-—0.66 mm), based on 4 specimens 
in lactic acid. Somite bearing first pair of 
legs fused with cephalosome. Epimera of 
metasomal somites rounded. Tergum of so- 
mite bearing leg 3 arched forward medially, 
exposing broad tergum of somite bearing leg 
4, this tergum crenulated posteriorly and 
showing median longitudinal sclerotization. 
Ratio of length to width of prosome 1.15: 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


1. Ratio of length of prosome to that of 
urosome 1.45:1. 

Somite bearing leg 5 (Fig. 1b) 65 x 170 
um. Genital somite (Fig. 1c) rectangular in 
dorsal view, 83 < 135 um (length including 
leg 6 but not its setae). Four postgenital so- 
mites from anterior to posterior 70 x 104, 
78 X 92,78 X 78, and 47 x 81 um. Anal 
somite indented medially. 

Caudal ramus (Fig. 1d) elongate, outer 
side 91 wm, inner side 127 wm, and width 
at midregion 29 um. Ratio of outer length 
to width 3.14:1. Ratio of inner length to 
width 4.38:1. Outer lateral seta, placed dor- 
sally, 77 wm, dorsal seta 44 wm, outermost 
terminal seta 67 wm, and innermost ter- 
minal seta, placed subterminally, short, 28 
um, all these setae smooth. Two long me- 
dian terminal setae 122 um (outer) and 890 
um (inner), both with very small serrations 
along their midregions (Fig. le). Rami with 
thick sclerotized inner and outer walls. 

Rostrum (Fig. 1f) weakly developed. An- 
tennule (Fig. 2a) 1 1-segmented, 450 um long, 
first 2 segments stout, remaining 9 segments 
slender. Lengths of its segments (measured 
along their posterior nonsetiferous mar- 
gins): 56 (52 um along anterior margin), 117, 
16, 21, 42, 39, 39, 36, 36, 37, and 18 um, 
respectively. Formula for armature: 3, 14, 
2,2,2, 2, 2, 2, 2, 2, and’ 7 +) aesthetessnx 
setae on second segment, associated with 
conspicuous, incomplete, segmental scler- 
otizations, noticeably stronger than other 
setae. All setae smooth. In 4 males segmen- 
tal sclerotizations in segments 4—11 stronger 
in left antennule (Fig. 2b) than in right an- 
tennule (Fig. 2a). 

Antenna (Fig. 2c) short, 130 wm long in- 
cluding terminal seta (compare length with 
that of antennule, 450 wm). Four segment- 
ed, but terminal seta showing trace of sub- 
division. First segment (coxa) unarmed. 
Second segment (basis) with minute exopod 
4 x 4.5 um with 1 seta 30 wm and orna- 
mented with patch of spinules. Endopod 
with large first segment having prolonged 
inner distal corner and dense field of small 


VOLUME 106, NUMBER 2 ST7 


Fig. 1. Pseudonicothoe branchialis Avdeev & Avdeev, 1978. Male. a, dorsal (scale A); b, urosome, dorsal 
(B); c, genital double somite and first postgenital somite, ventral (C); d, anal somite and caudal ramus, dorsal 
(C); e, detail of longest seta on caudal ramus, dorsal (D): f, cephalosome, ventral (E). 


318 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Pseudonicothoe branchialis Avdeev & Avdeev, 1978. Male. a, antennule, anterodorsal (scale C); b, 
segments 3-11 of antennule, ventral (C); c, antenna, antero-outer (F); d, end view of siphon, ventral (G); e, 
mandible, posterior (G); f, maxillule, ventral (G); g, maxilla, ventral (G); h, maxilliped, anterior (H); 1, endopod 
of maxilliped, anterior and slightly inner (F); j, endopod of maxilliped, posterior (F). 


VOLUME 106, NUMBER 2 


spines on inner ventral surface; second seg- 
ment small with 2 smooth inner setae and 
1 terminal seta with slight indication of di- 
vision proximally; terminal seta 44 um with 
small lateral spinules. 

Siphon very short, in ventral view ap- 
pearing as round sucking disk 60 um in di- 
ameter (Fig. 2d). Mandible (Fig. 2e) simple 
blade 117 um long. Maxillule (Fig. 2f) with 
2 lobes, outer lobe with 2 setae, inner lobe 
with 3 setae, all setae smooth. Maxilla (Fig. 
2g) situated far posterior to maxillule (see 
Fig. 1f). Subrectangular first segment 70 um 
long and stout. Second segment 52 um long, 
bearing 1 proximal seta; first third partially 
and indistinctly separated from second third, 
terminal third slightly clawlike and weakly 
separated from preceding third. Maxilliped 
(Fig. 2h) 5-segmented, 450 um long includ- 
ing claws. First segment (syncoxa) with 1 
smooth inner seta, second segment (basis) 
elongate with 1 smooth inner seta and or- 
namented with large inner field of long spi- 
nules and small spinules along outer surface. 
Three endopodal segments, first with outer 
seta and 2 small setules, second with | spine 
18 wm, and third with 1 spine 29 um, and 
terminal claw 90 um (Fig. 21, j). 

Ventral region between maxillipeds and 
first pair of legs as in Fig. If. 

Legs 1+ (Figs. 3a-c, 4a) with 3-segmented 
rami. Formula for armature as follows: 


P, coxa O-1 basis 1-I exp I-1; I-1; IL,1,3 
enp 0-1-'0-1- 15 


P, coxa 0-1 basis 1-0 exp I-1; I-1; IL1,4 
enp 0-1; 0-1; 1,1,4 
P; coxa 0-1 basis 1-0 exp I-1; I-1; II, 1,3 
enp 0-1; 0-1; 1,1,3 
P, coxa Q-1 basis 1-0 exp I-1; I-1; IL1,4 
enp 0-1; 0-2; 1,1,2 


Inner spine on basis of leg 1 minutely 
barbed and 34 um long. Outer seta on basis 
of legs 1-3 long and feathered, but in leg 4 
short and smooth. Terminal element on ex- 
opod of leg 1 intermediate between spini- 
form and setiform, barbed along outer side, 
with long coarse setules along inner side. 


319 


Leg 2 with 2 outer elements on third en- 
dopodal segment intermediate in form be- 
tween spine and seta, with short spinules 
along outer side and longer setules along 
inner side (Fig. 3b). Exopods of legs 1-3 
with small hyaline flaps (suckers?), 1 on seg- 
ment 2 and 3 on segment 3 (Fig. 3a-c). 

Leg 5 (Fig. 4b, c) with elongate free seg- 
ment, in ventral view 86 X 25 um, ratio 
3.4:1, bearing 4 setae, 2 terminal (outer 117 
um, inner 130 wm) and 2 subterminal; 1 
subterminal seta long and barbed (85 um), 
other subterminal seta (inner) short, 43 um 
stout, smooth, mucronate. Dorsal seta on 
body 86 um. 

Leg 6 (Fig. 4d) with 3 setae from outer to 
inner 65, 92, and 99 um, innermost seta 
with small proximal inner protuberance. 

Color of living specimens unknown. 

Female.—Body (Fig. 4e) with flattened 
prosome (as shown in Avdeev & Avdeev, 
1978: fig. 2) more elongate and tapered an- 
teriorly than in male. Length 1.93 mm (1.93- 
1.94 mm) and greatest width 1.00 mm (1.01- 
0.98 mm), based on 2 specimens in lactic 
acid. Terga of somites bearing legs 2—4 part- 
ly fused. Ratio of length to width of prosome 
1.58:1. Ratio of length of prosome to that 
of urosome 2.59:1. 

Somite bearing leg 5 (Fig. 5a) 47 x 122 
um. Genital double somite 83 um long, 100 
um wide in anterior rounded half, 73 um 
wide in abruptly narrowed posterior half. 
Genital areas located dorsolaterally just an- 
terior to junction of 2 halves of double so- 
mite. Each area (Fig. 5b) with 2 small del- 
icate setae. Three postgenital somites from 
anterior to posterior 47 x 65,50 x 55, and 
31 xX 49 um. 

Caudal ramus similar to that of male but 
slightly larger, 130 um long on outer side, 
39 wm wide at midregion, ratio 3.33:1. 

Rostrum, antennule, antenna, siphon, 
mandible, and maxillule, arranged as in Fig. 
5c, like those of male. Maxilla (Fig. 5d) small, 
55 wm long, apparently 2-segmented, with 
terminal spine. First segment with 2 small 
setae. Maxilliped as in male. Maxillules and 


320 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Pseudonicothoe branchialis Avdeev & Avdeev, 1978. Male. a, leg 1 and intercoxal plate, anterior 
(scale H); b, leg 2 and intercoxal plate, anterior (H); c, leg 3 and intercoxal plate, anterior (H). 


VOLUME 106, NUMBER 2 52) | 


Fig. 4. Pseudonicothoe branchialis Avdeev & Avdeev, 1978. Male. a, leg 4 and intercoxal plate, anterior 
(scale H); b, leg 5, dorsal (C); c, leg 5, ventral (C); d, leg 6, ventro-outer (C). Female. e, dorsal (A). 


322 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


foe 
—<— 


<P ILA AM a 


ee S TS << en 7 fe * my YE: 


© - 


Wn 


4 


—<— 
Pye PD + 

CLL 

ile RR 


< 


, dorsal (scale E); b, genital 
exopod of leg 1, anterior (H); f, leg 2, 


urosome 


Pseudonicothoe branchialis Avdeev & Avdeev, 1978. Female: a, 


Fig: 5. 
area, dorsal (C); c, cephalosome, ventral (A); 


3) 


e€ 


d, maxilla, ventral (G); 


anterior (H); g, exopod of leg 3, anterior (H); h, leg 5, dorsal (H). 


> 


VOLUME 106, NUMBER 2 


makxillae widely separated, with transverse 
sclerotized bar between them, as in Fig. 5c. 

Legs 1-4 segmented and armed as in male. 
Legs 1-3 with exopods having small round 
hyaline flaps, 1 on second segment and 3 on 
third segment (Fig. 5e—-g), as in male. 

Leg 5 (Fig. 5h) with free segment 133 x 
52 um, ratio 2.56:1. Four setae from inner 
to outer 125, 140, 133, and 122 um. Dorsal 
seta 146 um. 

Leg 6 represented by 2 setae on genital 
area (Fig. 5b). 

Only 1 somewhat damaged egg sac seen, 
separated from female, containing approx- 
imately 12 flattened, discoidal, linearly ar- 
ranged eggs, measuring 1045 x 330 um. 

Color unknown. 

Remarks. —We have been unable to make 
a direct comparison of the specimens from 
Heterocarpus sibogae with the type speci- 
mens of Pseudonicothoe branchialis. The 
types consist of two females, holotype and 
paratype, deposited in the Laboratory of 
Parasitology of Marine Animals, Pacific 
Ocean Scientific Research Institute of Fish- 
eries and Oceanography, Vladivostok, Rus- 
sla. 

Although the description and figures of P. 
branchialis published by Avdeev & Avdeev 
(1978) conform in major respects to our 
specimens from Australia, a few minor dif- 
ferences may be observed: (1) few relatively 
long setae on the antennule (Avdeev & 
Avdeev’s fig. 5), (2) the antenna with a 
feathered seta on the exopod and the fourth 
segment with one of the two small setae 
feathered (their fig. 10), (3) the outer branch 
of the maxillule with three setae (their fig. 
6), (4) the first segment of the maxilla in the 
female with one curved claw (their fig. 7), 
and (5) the free segment of leg 5 relatively 
short, ratio 1.64:1 (their fig. 7). We believe 
that these small differences may be attrib- 
utable to the relatively small number of 
specimens studied, the difficulties of obser- 
vation, and to the style of illustration. 

The exopods of swimming legs 1—3 of both 
male and female P. branchialis are orna- 


323 


mented with marginal structures referred to 
as hyaline flaps in the present account, and 
as suckers by Avdeev & Avdeev (1978). 
When viewed from the side, as in Fig. 3a— 
c, these structures appear to be hyaline ex- 
tensions of the lateral margin of the exo- 
podal segments. When they are reflexed 
across the surface of the exopodal segment, 
radial surface striations are visible and their 
appearance is more suckerlike. Similar 
structures were figured on legs 1-3 of male 
P. procircularis (Carton) by Carton (1967). 

The discovery of the male of P. bran- 
chialis prompted us to reconsider the valid- 
ity of the genus Pseudonicothoe which was 
treated as a subjective synonym of Parani- 
cothoe Carton by Boxshall & Lincoln (1983). 
The type species of Paranicothoe, P. cla- 
docera, differs from Pseudonicothoe in the 
segmentation of the antennules, in the seg- 
mentation and armature of the maxilliped, 
and in the spine and setal formula of the 
swimming legs. According to Huys & 
Boxshall (1991) the antennule is 10-seg- 
mented in P. cladocera, the distal part com- 
prises relatively short segments, and the api- 
cal segment is longer than the two preceding 
segments combined, whereas in P. bran- 
chialis the antennule is 11-segmented, the 
distal part comprises relatively long seg- 
ments, and the apical segment is markedly 
shorter than the subapical segment. The en- 
dopod of the maxilliped of P. cladocera is 
2-segmented and is armed with one claw on 
the first segment and two on the second. In 
P. branchialis, by comparison, the endopod 
is 3-segmented, with the first segment car- 
rying three short setae, the second segment 
bearing one claw, and the third two claws. 
Finally, the endopod of leg 4 has a setal 
formula of 0-1; 0-2; 1,1,2 in P. branchialis 
and 0-1; 0-1; 1,1,2 in P. cladocera. 

These differences justify the generic level 
separation of Pseudonicothoe and Parani- 
cothoe, as represented by its type species, 
and we propose to recognize Pseudonico- 
thoe as a valid genus again, thereby revers- 
ing the proposal made by Boxshall & Lin- 


324 


coln (1983). The type species of 
Pseudonicothoe is P. branchialis. Parani- 
cothoe is also a valid genus, containing only 
the type species, P. cladocera. We propose 
to transfer Paranicothoe procircularis (Car- 
ton) to Pseudonicothoe, as a new combi- 
nation, Pseudonicothoe procircularis (Car- 
ton, 1967). This transfer is based on the 11- 
segmented condition of the antennule, and 
on the spine and setal formula of the swim- 
ming legs. 


Acknowledgment 


We thank Dr. A. J. Bruce, Northern Ter- 
ritory Museum, Darwin, Australia, for 
sending specimens of the nicothoid to us for 
study. The study of the copepods was aided 
by a grant (BSR 88 21979) to AGH from 
the National Science Foundation of the 
United States. 


Literature Cited 


Audouin, V., & H. Milne Edwards. 1826. Mémoire 
sur la Nicothoe, animal singulier qui suce le sang 
des homards.— Annales de Sciences Naturelles 
9:345-358. 

Avdeev, G. V., & V. V. Avdeev. 1978. Pseudoni- 
cothoe branchialis gen. et sp. n. (Crustacea, Co- 
pepoda) from gills of Heterocarpus laevigatus 
from the Pacific Ocean.— Zoologicheskii Zhur- 
nal 57:1893-1897. 

Boxshall, G. A., & R. J. Lincoln. 1983. Some new 
parasitic copepods (Siphonostomatoida: Ni- 
cothoidae) from deep-sea asellote isopods.— 
Journal of Natural History 17:891-900. 

Carton, Y. 1967. Description de Nicothoe procircu- 
laris n. sp. (Crustacea, Copepoda) discussion sur 
la forme male. — Videnskabelige Meddelelser fra 
Dansk Naturhistorisk Forening 130:143-152. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


1970a. Description de Paranicothoe n. gen. 
un nouveau représentant de la famille des Ni- 

cothoidae.—Galathea Report 11:239-246. 

1970b. Le genre Paranicothoe, un nouveau 
représentant de la famille des Nicothoidae.— 

Second International Congress of Parasitology, 
Journal of Parasitology 56(II):47-48. 

Connolly, C.J. 1929. Anew copepod parasite Choni- 
osphaera cancrorum, gen. et sp. n., representing 
a new genus, and its larval development. —Pro- 
ceedings of the Zoological Society of London 
1929:415-427. 

Dana, J. D. 1852. Conspectus crustaceorum quae in 
orbis terrarum circumnavigatione Carolo Wilkes 
e classe reipublicae foederatae duce, lexit e de- 
scripsit Jacobus D. Dana. Part 2.— Proceedings 
of the American Academy of Arts and Sciences 
2:9-61. 

Hansen, H.J. 1897. The Choniostomatidae. A family 
of Copepoda, parasites on Crustacea Malacos- 
traca. Copenhagen, Denmark. Pp. 1-205. 

Humes, A. G. 1975. Hadrothoe crosnieri n. gen., n. 
sp. (Crustacea, Copepoda), from a penaeid 
shrimp (Crustacea, Decapoda) in Madagas- 
car.— Zoologischer Anzeiger 195:21-34. 

Huys, R., & G. A. Boxshall. 1991. Copepod evolu- 
tion. The Ray Society 159:1-468. 

Pillai, N. K. 1962. Choniomyzon gen. nov. (Cope- 
poda: Choniostomatidae) associated with Pan- 
ulirus.—Journal of the Marine Biological As- 
sociation of India 4:95—99. 

Thorell, T. 1859. Till kannedomen om vissa parasi- 
tiskt lefvande Entomostracer.—Ofversigt af 
Kongliga Akademiens Forhandlingar 16(8):335- 

362. 


(AGH) Boston University Marine Pro- 
gram, Marine Biological Laboratory, Woods 
Hole, Massachusetts 02543, U.S.A.; (GAB) 
The Natural History Museum, Cromwell 
Road, London, England SW7 5BD. 


PROC. BIOL. SOC. WASH. 
106(2), 1993, pp. 325-331 


BOREOMYSIS OPARVA, A NEW POSSUM SHRIMP 
(CRUSTACEA: MYSIDACEA) FROM AN EASTERN 
TROPICAL PACIFIC SEAMOUNT 


Jennifer Saltzman and Thomas E. Bowman 


Abstract.— Boreomysis oparva is described from near-bottom low oxygen 
waters surrounding an inactive seamount, Volcano 7. It is characterized by an 
upturned rostrum, large eyes without papillae, and a male pleopod 3 exopod 
with 3 spiniform setae on each of the last eight segments. It is abundant slightly 
below the seamount summit in near-bottom low oxygen water, but absent from 
water near the seamount base (with higher oxygen) and from low oxygen pelagic 


waters. 


Volcano 7 is an inactive seamount 20 km 
in diameter in the eastern tropical Pacific at 
13°23'N, 102°27'W. It arises from a depth 
of 3400 m to a summit at 730 m; the latter 
penetrates the pronounced oxygen-mini- 
mum zone of the region (Wishner et al. 1990, 
Levin et al. 1991). In November 1988 the 
plankton and benthos of Volcano 7 were 
sampled from shipboard on the R/V Atlan- 
tis IJ and from the submersible D.S.R.V. 
Alvin. The samples collected from the sub- 
mersible contained more than 400 speci- 
mens of an undescribed species of the mys- 
idacean genus Boreomysis, described and 
illustrated herein. 


Methods 


A multiple opening-closing 8-net system 
(183 wm mesh) mounted on the Alvin 
(Wishner & Gowing 1987) was used to col- 
lect zooplankton 1-3 m above the bottom 
during daytime dives. The 1-liter Plexiglas 
cod-end chambers were equipped with 
spring-loaded front and rear doors that were 
fixed open during the tow and were snapped 
shut at the end of the tow, making the cham- 
ber watertight. Simultaneously with the 
chamber closing a spring-loaded needle 
punctured a rubber balloon within the 
chamber, releasing its contents of glutaral- 
dehyde and fixing the sample in situ. Plank- 


ton samples were also collected from ship- 
board with a 1 m? MOCNESS plankton net 
system (333 um mesh) in vertically strati- 
fied tows to 1200 m depth. 


Order Mysidacea Boas, 1883 
Suborder Mysida Boas, 1883 
Family Mysidae Dana, 1850 
Subfamily Boreomysinae Holt & Tattersall, 
1905 
Genus Boreomysis G. O. Sars, 1869 
Boreomysis oparva, new species 
Figs. 1-20 


Material. —See Table 1. 

Types. — Holotype 6, 22.5 mm, Dive 2145, 
Net 4, USNM 251923. Paratypes: Dive 
2139, Net 8, 14 specimens, USNM 251918: 
Dive 2142, Net 8, 100 specimens, USNM 
251919; Dive 2143, Net 4, 2 specimens, 
USNM 251920; Dive 2144, Net 8, 4 spec- 
imens, USNM 251921; Dive 2145, Net 4, 
6 specimens, USNM 251922. 

Etymology.—From “o,” the chemical 
symbol for oxygen, plus the Latin “‘parvus”’ 
(little, slight), referring to the occurrence of 
this mysid in low-oxygen water. 

Description. —Length up to about 25 mm. 
Anterior margin of carapace slightly con- 
vex. Rostrum acute, reaching 3 to '2 length 
of lst segment of antenna 1, directed dor- 
sally at angle of about 45° to lateral axis of 


326 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Boreomysis oparva collected from Volcano 7 by the submersible D.S.R.V. Alvin in November 1988. 
Collections marked with an asterisk are deposited in the National Museum of Natural History, Smithsonian 


Institution. 
Dive-Net ) Q Juv. 
2139-1 — — 7 
4 1 2 1 
3 1 i 2 
6 2 4 — 
8* 5 8 1 
2140-1 2 3 — 
2 — = — 
3 8 4 3 
4 1 2 = 
5 ] 4 _ 
6 — 3 = 
v + | 7 
8 1 12 13 
2142-1 3 + 3 
2 1 4 — 
3 10 hi 4 
4 26 72 116 
5 6 8 6 
7 6 18 6 
8* 9 56 35 
2143-2 2 1 1 
3 - 3 3 
4* — l 1 
5 _ 3 2 
8 3 6 6 
2144-1 = 1 a 
3 _ 1 1 
= 1 1 — 
i 1 — — 
8* _ 1 3 
2145-3 a ue 2 
4* 4 1 
8 1 1 — 


Maximum 

length Mid-depth 
(mm) (m) Location 
18.0 790 summit 
26.5 852 summit 
22S 790 summit 
OAS 810 summit 
25.4 852 summit 
19.8 804 summit 
20.0 793 summit 
23.0 815 summit 
18.8 874 summit 
20S 804 summit 
18.3 793 summit 
23.9 817 summit 
18.7 876 summit 
23.0 797 summit 
21.4 782 summit 
23.4 788 summit 
250 797 summit 
22.2 797 summit 
DOS 788 summit 
2S 808 - summit 
27M 1300 flank 

— 1297 flank 

— 1308 flank 
20.2 1300 flank 
22.0 1309 flank 

— 1289 flank 
14.3 1179 flank 

— 1250 flank 
20.8 1195 flank 
[gis 1259 flank 

9.6 1323 flank 
2295) 1352 flank 
20.3 1304 flank 


carapace. Anteroventral corner of carapace 
acute; cervical groove well developed. Car- 
apace covering laterally 1st segment of ex- 
opods of thoracopods and posteriorly about 
’) of pleonite 1. Eye with rather short, nar- 
row stalk and broad cornea, reaching well 
beyond midlength of lst segment of antenna 
1, without ocular papilla. 

Antenna 1, lst segment of peduncle nar- 
rower than 2nd and 3rd segments, nearly 
3x as long as wide, with medial papilla at 


distal 4 bearing 4 setae and distolateral pa- 
pilla with 5 setae. 2nd segment short, with 
2 long dorsal setae, shorter seta on disto- 
medial corner, and 3 setae on distolateral 
papilla. 3rd segment slightly more than '2 
length of 1st segment, with cluster of 7 setae 
at distomedial corner and 3 setae on quad- 
rate process at midwith of distal margin. 6 
antennular brush very dense and long; if 
present, 6 lobe obscured by setae of brush. 
Ist segment of lateral flagellum densely 


VOLUME 106, NUMBER 2 SAF 


Figs. 1-9. Boreomysis oparva. 1, Head and thorax, dorsal; 2, Same, lateral; 3, Eye and rostrom, dorsal; 4, 
Antenna, 1, 2, dorsal; 5, Peduncle of ¢ antenna 1, ventral; 6, Antenna 2, dorsal; 7, Right mandible, gnathal 
surface; 8, Left mandible, same, but molar omitted; 9, Maxilla 1, outer ramus. 


328 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 10-16. Boreomysis oparva. 10, Apex of mandibular palp; 11, Maxilla 2; 12, Endites of maxilla 1, Ist 
endite (below) enlarged; 13, Maxilliped; 14, Thoracopod 2; 15, Thoracopod 5; 16, Penis, lateral. 


VOLUME 106, NUMBER 2 


ey, 
—————— 


Figs. 17-20. Boreomysis oparva. 17, Exopod of 6 pleopod 3, distal segments; 18, Telson, dorsal; 19, Apex 
of telson, dorsal; 20, Right uropod, dorsal, with enlarged detail of spines of exopod and endopod. 


armed with setae about as long as Ist seg- 
ment of peduncle; only proximal parts of 
setae shown in Fig. 4. 

Antenna 2 peduncle not quite reaching 
midlength of scale; segment 1 produced dis- 
tolaterally into spiniform process. Scale 
nearly 4x as long as broad; terminal spine 
reaching beyond nearly truncate apex, which 
slants laterally at about 15° to axis of scale. 

Left mandible with bicuspid incisor and 
lacinia; spine-row of 8 spines. Right man- 
dible with tricuspid incisor; lacinia dichot- 
omous, dorsal ramus a curved pointed tooth, 
ventral ramus quadrate, bearing 3 spines; 
spine-row formed of a single serrate spine 
separated by gap from 9 simple spines with 
common base. Palp 2nd segment broad 
proximally, narrowing distally, with scat- 


tered marginal setae; 3rd segment about 0.7 
length of 2nd segment, distal half with close- 
set plumose marginal setae and 2 long apical 
setae. 

Maxilla 1 outer ramus with 7 setae on 
surface and 15 apical spines. 

Maxilla 2 protopod margin with dense 
covering of fine setae interspersed with 9 
shorter and stouter setae. lst endite with 16, 
lobes of 2nd endite with 12 and 15 apical 
setae respectively. Endopod segments sub- 
equal in length; lst segment with 9 setae on 
medial margin; 2nd segment with about 30 
marginal setae. Exopod reaching slightly be- 
yond Ist endopod segment, with about 33 
marginal setae. 

Maxilliped (endopod of thoracopod 1) 
endite of basis reaching distal margin of me- 


330 


rus. Carpus nearly as long as propus and 
dactyl (excluding nail) combined. Medial 
margin of all segments densely setose. 

Endopods of thoracopods 2-8 with pro- 
pus divided by incomplete suture. Dactyl of 
thoracopod 2 without nail. Anterior margin 
of propus of thoracopods 3-8 with 3 clusters 
of 5—7 setae in which 1 seta is much longer 
than others. Exopods of thoracopods with 
17-20 segments. 

Penis about as long as basiopod of thora- 
copod 8, widening distally; posterior margin 
with right-angled bend proximal to rounded 
setose apex. 

3rd 6 pleopod with 18 segmented endo- 
pod and longer 23 segmented exopod. On 
exopod paired plumose setae of proximal 
segments replaced on last 8 segments by 3 
simple spiniform setae on each segment, ex- 
cept 2 such setae on apical segment; 2 of the 
3 setae at distomedial corner, | at distolater- 
al corner. 

Telson slightly more than 3 x as long as 
width at base, slightly longer than pleonite 
6, posterior half with slightly concave mar- 
gins. Apical cleft about % length of telson, 
each side with about 30 teeth, margins grad- 
ually diverging posteriorly; base not dilated 
but with narrow slit. Each lobe of apex armed 
with long spine flanked laterally by 1 spine 
and medially by 2 spines; flanking spines 
half length of long spine. Lateral margins of 
telson each armed with 20—23 spines, some 
shorter than others but not arranged in reg- 
ular pattern. 

Endopod of uropod slightly longer than 
telson, with 2 spines (O—1 in immatures) in- 
serted ventrally near medial margin adja- 
cent to statocyst. Exopod '/, longer than en- 
dopod, lateral margin naked for proximal 
74, 2 spines on outer margin at distal end of 
naked part. 

Comparisons. —Nearly 40 species of Bor- 
eomysis are recognized currently, but it is 
uncertain how many of them will prove 
eventually to be valid. Some are based only 
on females or immature males, hence the 
structure of the mature male exopod of 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


pleopod 3, a character of high taxonomic 
value, 1s unknown. The condition in B. 
oparva, the 8 distal segments each with 3 
simple setae, 1s at present unique; other spe- 
cies in which this pleopod has been de- 
scribed have 2 such setae. This feature, 
combined with the upturned rostrum, the 
large eyes lacking papillae, and the 2 spines 
on both the endopod and exopod of the uro- 
pod, readily distinguish B. oparva from sim- 
ilar species. 

Birstein & Tchindonova (1958) key out 
27 species and 2 varieties of Boreomysis, 
and li (1964) gives a key to the 10 species 
and 2 varieties that he recognized from the 
northwestern Pacific. 

Ecology. —The summit of Volcano 7 (730 
m) lies in the oxygen minimum zone (Wish- 
ner etal. 1990), and B. oparva was not found 
there. Slightly deeper at the lower summit 
(ca. 790 m), associated with the increase in 
oxygen concentration from 0.08 to 0.88 ml/ 
liter (Levin et al. 1991), B. oparva became 
abundant, reaching a maximum of 114/m;>. 
At the flank of the seamount (1185-1310 
m) B. oparva was less abundant and absent 
from most of the samples. It was absent 
from near-bottom waters at the base of the 
seamount (3400 m) and from pelagic waters 
(O—1200 m) surrounding the seamount. The 
abundance peak at the lower summit was 
similar to that found for many infaunal taxa 
(Levin et al. 1991). 

Food. —Gowing & Wishner (1992) found 
dense numbers of gram-positive bacteria- 
like bodies in the guts of some specimens 
of this mysid, suggesting that they oppor- 
tunistically consume bacterial aggregates or 
mats at the lower boundary of the oxygen 
minimum zone. 


Acknowledgments 


We thank Dr. Karen F. Wishner for pro- 
viding us with the specimens of the new 
species of Boreomysis and for reviewing the 
manuscript. Ship time for the collection of 
the specimens was provided by NSF grants 


VOLUME 106, NUMBER 2 


OCE 87-16564 to Dr. Wishner and OCE 
87-0193 to Dr. Marcia M. Gowing, and 
ONR contract NO0014-84-K-0081 to Dr. 
Lisa A. Levin. 


Literature Cited 


Birstein, Ya. A., & Yu. G. Tchindonova. 1958. Deep- 
sea mysids from the northwestern part of the 
Pacific Ocean. — Trudy Instituta Okeanologii 27: 
258-355. [in Russian] 

Boas, J. E. V. 1883. Studien tiber die Verwandt- 
schaftsbeziehungen der Malakostraken.— Mor- 
phologisches Jahrbuch 8:485-579. 

Dana, J. D. 1850. Synopsis generum crustaceorum 
ordinis “Schizopoda”’ J. D. Dana elaboratus, et 
descriptiones specierum hujus ordinis quae in 
orbis terrarum circumnavigatione, Carolo Wilkes 
e Classe Reipublicae Faederatae Duce, auctore 
lectae (pars I).—American Journal of Sciences 
and Arts (2) 9:129-133. 

Gowing, M. M., & K. F. Wishner. 1992. Feeding 
ecology of benthopelagic zooplankton on an 
eastern tropical Pacific seamount.— Marine Bi- 
ology 112:451-467. 

Holt, E. W. L., & W. M. Tattersall. 1905. Schizop- 
odous Crustacea from the north-east Atlantic 
slope.— Report on the Sea and Inland Fisheries 
of Ireland, 1902-1903, part 2, Appendix 4:99- 
152, pls. 15-25. 


331 


li, N. 1964. Fauna Japonica, Mysidae. Biogeograph- 
ical Society of Japan, Tokyo, 610 pp. 

Levin, L. A., C. L. Huggett, & K. F. Wishner. 1991. 
Control of deep-sea benthic community struc- 
ture by oxygen and organic matter gradients in 
the eastern Pacific Ocean.—Journal of Marine 
Research 49:763-800. 

Sars, G. O. 1869. Undersogelser over Christiania- 
fjorden Dybvandsfauna anstillede paa en i Som- 
meren 1868 foretagen Zoologisk Reise.—Nytt 
Magasin for Naturvidenskapene 16:305-362. 

Wishner, K. F., & M. M. Gowing. 1987. In situ fil- 

tering and ingestion rates of deep-sea benthic 

boundary-layer zooplankton in the Santa Cat- 

alina Basin.— Marine Biology 94:357—366. 

, L. Levin, M. Gowing, & L. Mullineaux. 1990. 

Multiple roles of the oxygen minimum in ben- 

thic zonation on a deep seamount. — Nature 346: 

57-59. 


(JS) Graduate School of Oceanography, 
University of Rhode Island, Narragansett, 
Rhode Island 02882-1197, U.S.A.; (TEB) 
Department of Invertebrate Zoology (Crus- 
tacea), National Museum of Natural His- 
tory, Smithsonian Institution, Washington, 
D.C. 20560, U.S.A. 

The sequence of the authors is according 
to the recency of their birthdates. 


PROC. BIOL. SOC. WASH. 
106(2), 1993, pp. 332-338 


THE IDENTITY OF TALITROIDES ALLUAUDI 
(CHEVREUX) (CRUSTACEA: AMPHIPODA: TALITRIDAE) 
WITH NOTES ON A NEW LOCALITY 


Hiroshi Morino and Reuven Ortal 


Abstract. —A lectotype of Talitroides alluaudi (Chevreux) is designated from 
the type-series and described to remove taxonomic confusion on the identity 
of this species. The discovery of this species in Israel is briefly discussed. 


Examination of terrestrial talitrids col- 
lected from southern parts of Israel revealed 
two genera and three species, one of which 
proved to be Talitroides alluaudi. This spe- 
cies is one of the most widespread landhop- 
pers, known from the tropics to warm-tem- 
perate regions and hothouses in Europe and 
North America (Friend & Richardson 1986), 
though so far not known from the Levant. 
Although some authors have remarked on 
various characters, especially concerning the 
pleopods, of this species (Medcof 1940, Pal- 
mén 1949, Andersson 1962), no full de- 
scriptions or figures have been published 
since the original description of Chevreux 
(1896, 1901). Close examination of material 
from Israel, with reference to Chevreux’s 
descriptions, revealed a few minor but dis- 
tinct discrepancies between them as well as 
between previous descriptions of 7. alluau- 
di. 

Chevreux (1896) described this species 
three times from two localities. He gave a 
brief original description of this species on 
the basis of material procured from hot- 
houses at the Paris Museum (“les serres du 
Muséum de Paris’’). In this paper he men- 
tioned that the Paris material was identical 
to that collected from the Seychelles. Al- 
though he did not specify the locality of the 
type, the title of his paper suggests that Paris 
is the probable type locality. Shoemaker 
(1936), however, assumed that the Sey- 
chelles was the type locality. In 1901, Chev- 
reux treated the Seychelles material in de- 


tail. The Paris material was described again 
in 1925 (Chevreux & Fage 1925) with il- 
lustrations. A study of these three descrip- 
tions discloses the following apparent un- 
conformity in some characters: The Paris 
1896 and Seychelles material have well-de- 
veloped brood plate on female gnathopod 
2, though this is lacking in the Paris 1925 
material, and; the Seychelles material has 
l-articulated and 3-articulated inner ramus 
on pleopods 1 and 2, respectively, though 
the Paris 1925 material has 4-articulated 
and 1|-articulated inner ramus, respectively 
(no description of pleopods for the Paris 
1896 material). If these descriptions are 
taken authentically, we could recognize three 
species in the Chevreux’s concept of J. al- 
luaudi: two from Paris and one from the 
Seychelles. And if the Paris 1896 material 
was from the Seychelles, as assumed by 
Shoemaker (1936), two species may be dis- 
cernible, from Paris and the Seychelles, re- 
spectively. Medcof (1940) has also pointed 
out some confusion in the earlier descrip- 
tions of this species, which he attributed to 
geographical variation (“based on exami- 
nations of specimens collected from areas 
sometimes widely separated’’). Since Med- 
cof’s analysis of variation is limited to the 
pleopod structure, he did not notice the 
variation in the brood plate. In addition, 
material from Israel at hand displays a pe- 
culiar feature of the pereopod dactyl, which 
was described in the Seychelles material dif- 
ferently, and was not mentioned for the Par- 


VOLUME 106, NUMBER 2 


is material. Thus it is necessary to deter- 
mine the original concept of 7. alluaudi 
through direct reference to the type-series. 

The type material of Talitrus alluaudi de- 
posited in Muséum National d’Histoire Na- 
turelle in Paris consists of preserved spec- 
imens from four localities (MNHWN Paris 
Am 4500, 4501, 4502, 4503) and slides 
(MNHN Paris Am 4504). The label on the 
slides (“Serre du Muséum’’) suggests that 
they comprise the type specimen. Unfor- 
tunately the condition of the slides is so bad 
that it is not possible to discriminate the 
pertinent characters, and all the preserved 
specimens are from other than the Paris 
Museum. Thus the lectotype is selected from 
the locality nearest to the Paris Museum and 
described to resolve the problem of T. al- 
luaudi. Material from Israel and Hawaii is 
also examined. 


Systematic Account 


Talitroides alluaudi (Chevreux, 1896) 
Figs. 1, 2 


Talitrus Alluaudi Chevreux, 1896:112, figs. 
1-4; 1901:389, figs. 1-6.—Chevreux & 
Fage, 1925:270, figs. 280-281. 

Orchestia senni Menzel, 1911:438, figs. 4-9 

Talitroides alluaudi. —Palmén, 1949:61, figs. 
1-12.—Andersson, 1962:211, figs. 1-3.— 
Bousfield, 1984:210. 


Material examined.—Two females (5.5 
mm-—lectotype, 5.0 mm—paralectotype), 
from Serres de la Ville de Paris, Boulogne 
sur Seine, France (MNHN Paris Am 4500); 
1 female (5.0 mm), from Iles Séchelles, 
Mahe, Auct. det 1901 (MNHN Paris Am 
4502); 2 females (6 mm and 7 mm), from 
Ponta Delgada, Ile S. Michel, Acores, Aug 
1930, M. Mequignon collector, (MNHN 
Paris Am 4503); 5 females (up to 5.5 mm), 
from Ben Gurion Univ. campus, Beersheba, 
Israel, bamboo stands, 10 Dec 1987, Y. 
Margalit collector (Morino Cat. No. T.766), 
4 Mar 1989, R. Ortal collector (Morino Cat. 
No. T1836); 1 female (4.5 mm), from Hal- 


333 


awa, Molokai Is., Hawaii, 9 Mar 1968, Si- 
nonaga collector (Morino Cat. No. T.868); 
1 female (4.8 mm), from Kokee, Kauai Is., 
Hawaii, 11-12 Mar 1968, Sinonaga collec- 
tor (Morino Cat. No. T.869). 

Description of lectotype.—Eye small in 
size, subround. Inferior antennal sinus me- 
dium deep. Head longer than deep. 

Antenna 1 exceeding mid-point of pe- 
duncular article 5 of antenna 2, peduncular 
articles subequal in length; flagellum a little 
shorter than peduncle, 6-articulated. An- 
tenna 2: peduncle weakly spinose, article 5, 
1.79 times as long as article 4; flagellum 
subequal to peduncle in length, 1 1-articu- 
lated. 

Upper lip: as wide as deep. Mandible: in- 
cisor chitinized strongly, brown in color, 
5-dentate; left lacinia chitinized as incisor, 
4-dentate; right lacinia not strongly chitin- 
ized, with 3 cusps; spine rows of 2 broad 
plumose bristles. Lower lip normal. Maxilla 
1: innermost spine of outer plate leaning 
medially, outermost 2 spines without den- 
ticles; palp small, joint of article 2 indistinct. 
Maxilla 2: inner plate distolaterally pro- 
duced into pointed tip, several medio-distal 
setae plumose. Maxilliped: inner plate api- 
cally truncate, with 3 weak spine-teeth, in- 
creasing in size laterally, and with several 
plumose stiff setae; outer plate weakly ar- 
cuate with rounded apex, weakly setose; palp 
articles 2 and 3 slender, with a few stiff setae 
distally, article 2 with remnant of medial 
lobe represented by elongate setae, article 4 
partially fused to article 3. 

Gnathopod 1: coxa truncate distally, low- 
er margin spinose, inner shelf weak, with a 
spine; basis broadened distally; carpus 1.50 
times as long as propod; propod simple, 
gradually narrowed to base of dactyl, pos- 
terior margin with 3 stiff long spines and 
several shorter submarginal spines on inner 
and outer sides, anterior margin with 2 spine 
clusters; dactyl cuspate anteriorly, base with 
a stiff seta posteriorly, nail as long as base. 
Gnathopod 2: coxa cuspate posteriorly, as 
deep as wide, lower margin rounded and 


334 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Bip b 
(female, 5.5 mm body length). A, lateral view; B, antenna 1; C, antenna 2; D, left mandible; E, right mandible; 
F, maxilla 2; G, maxilla 1; H, maxilliped; I, distal part of right palp of maxilliped; J, upper lip; K, lower lip. 


weakly spinulose; basis slender, anterior 
margin spinulose; merus and carpus with 
tumescence posteriorly, carpus as long as 
propod; propod anterior margin weakly se- 
tose, posterior tumescent lobe well devel- 
oped, exceeding dactyl by half of propod 
length. 

Coxae 3 and 4 shallower than wide, with 


Talitroides alluaudi (Chevreux). Boulogne sur Seine, France. The lectotype, MNHN Am 4500-1 


a stiff seta posteroventrally, and a few spi- 
nules ventrally. Coxa 5 anterolobate, ante- 
rior lobe deeper than that of pereopod 4. 
Coxa 6: posterior lobe truncate distally, an- 
terior margin of the lobe vertical. Coxa 7 
shallow. 

Pereopods 3-7 cuspidactylate, propod 
lacking hinge spine, dactyl base with a stiff 


VOLUME 106, NUMBER 2 335 


Fig. 2. Talitroides alluaudi (Chevreux). Boulogne sur Seine, France. A, gnathopod 1; B, dactyl of gnathopod 
1; C, gnathopod 2; D, coxal gill of gnathopod 2: E, pereopod 3; F & G, dactyl of pereopod 3; H, pereopod 4; I, 
dactyl of pereopod 4; J, pereopod 5; K, dactyl of pereopod 5; L, pereopod 6; M, coxal gill of pereopod 6; N, 
dactyl of pereopod 6; O, pereopod 7; P, dactyl of pereopod 7; Q, abdominal side plates 1-3; R, pleopod 1; S, 
pleopod 2; T, pleopod 3; U, uropod 3; V, uropod 2: W, uropod 1; X, telson. G = Azores, MNHN Am 4503-1 
(female 6.0 mm); M = Boulogne sur Seine, France, paralectotype MNHN Am 4500-2 (female 5.0 mm); others 
= Boulogne sur Seine, France, lectotype MNHN AM 4500-1 (female 5.5 mm). 


336 


seta. Pereopod 3: dactyl base weakly 
pinched, nail with slight prominence at 
middle on posterior margin. Pereopod 4 
shorter than pereopod 3, basis weakly ar- 
cuate, dactyl base pinched, nail with sharp 
dentition posteriorly. Pereopod 5 longer than 
pereopod 4, basis narrow with a submar- 
ginal spine at posterodistal corner, lacking 
posterodistal lobe. Pereopod 6 similar to 
but much shorter than pereopod 5, dactyl 
nail almost straight. Pereopod 7 slightly lon- 
ger than pereopod 6, basis expanded pos- 
teriorly, as deep as wide. 

Coxal gill of gnathopod 2 as long as basis, 
distal margin with crenulations. Coxal gills 
of pereopods 3-5 small, constricted at mid- 
dle. Coxal gill of pereopod 6 longer than 
basis, reverse L-shaped. Brood plates con- 
fined to pereopods 3-5, small and slender 
with 2-3 simple setae apically. 

Abdominal side plate 1 rounded postero- 
ventrally. Plates 2 and 3 bluntly pointed 
posteroventrally, posterior margins weakly 
spinulose. Pleopod 1: peduncle weakly ar- 
cuate, with 3 plumose setae on outer mar- 
gin, with 2 retinaculae; outer ramus 
7-articulated; inner ramus shorter than half 
of outer ramus, 2-articulated. Pleopod 2 a 
little shorter than pleopod 1, peduncle outer 
margin with a plumose seta, outer ramus 
6-articulated, inner ramus |-articulated with 
apical setae. Pleopod 3 small and slender, 
0.4 times as long as peduncle of pleopod 2, 
with 2 spinules subapically, lacking rami. 

Uropod 1: peduncle with marginal spines 
on both edges, distolateral spine strong, 
closely set to distomarginal spine; rami sub- 
equal in length and shorter than peduncle, 
with elongate apical spines; outer ramus 
marginally bare; inner ramus with 3 mar- 
ginal spines. Uropod 2: peduncle with mar- 
ginal spines distally; rami subequal to each 
other and to peduncle in length, with elon- 
gate apical spines; outer ramus marginally 
bare; inner ramus with spines proximally. 
Uropod 3 very small, not reaching tip of 
telson, subtriangular in shape; peduncle 
broad at base with a strong spine ventrally; 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ramus small, conical, partially fused to pe- 
duncle, with a spinule at tip. Telson broad, 
apically rounded with apical and 3-4 lateral 
spines. 

Male. —Not known. 

Variations. —Eye in the Israeli material is 
somewhat larger than that of the lectotype. 
Number of flagellar articles ranges from 4 
to 6 on antenna | and from 8 to 9 on antenna 
2 in the material examined, tending to in- 
crease with body size. Marginal setae on 
propod anterior margin of gnathopod 2 also 
show variation in number: Seychelles and 
Hawaiian Islands material lacks them, Is- 
raeli material with 1-2 setae, Azores ma- 
terial with 0—2. Swiss material (Menzel 1911, 
as Orchestia senni) also has one seta. Inner 
rami of pleopods 1 and 2 fluctuate in the 
number of articles: from 1 to 4 (mostly 2) 
articles in pleopod 1; 1, exceptionally 3 (in 
Azores material, 7 mm body length) articles 
in pleopod 2. Only this larger specimen from 
the Azores has reduced ramus in pleopod 
3, otherwise lacking them. For each speci- 
men examined, article number of inner ra- 
mus of pleopod | is higher than, rarely same 
as that of pleopod 2. This range and pattern 
of variation in pleopods is in accord with 
those in the other geographical populations: 
Basel, Switzerland (Menzel 1911); Urbana, 
Illinois, U.S.A. (Medcof 1940); Finland 
(Palmén 1949) and the Canary Islands (An- 
dersson 1962). Dactyl nail of pereopod 3 
has usually very weak prominence at the 
posterior middle. The Azores material (6.0 
mm) has, however, distinct prominence (Fig. 
2G) which is rather similar to the dentition 
of pereopod 4. 


Discussion 


Most of the variations mentioned above 
occur within a given population rather than 
between geographically separated popula- 
tions. Thus it is not possible to recognize 
more than one species amongst the material 
examined. All the material lacks a brood 
plate on female gnathopod 2, and has dac- 


VOLUME 106, NUMBER 2 


tylar dentition on pereopod 4 (rarely also 
on pereopod 3). Chevreux (1901) men- 
tioned the dactylar dentition on pereopod 
5, not on pereopod 4, in his Seychelles ma- 
terial. At the same time, in our specimens, 
the inner ramus of pleopod 1 has more ar- 
ticles than that of pleopod 2, and never less, 
as by Chevreux (1901). Thus it is possible 
that Chevreux treated an exceptional spec- 
imen or more probably some confusion oc- 
curred during the preparation of his mate- 
rial. 

The present species is most closely related 
to Talitroides topitotum (see Bousfield 1984). 
Friend & Lam (1985) listed several discrim- 
inating characters between TJ. alluaudi and 
T. topitotum. In addition, the following ones 
are to be mentioned: T. al/uaudi has gnatho- 
pod 2 without brood plate, strongly chitin- 
ized mandible, maxilla 2 with distally 
pointed inner plate, and pereopod 4 with 
dactylar dentition. These features are rather 
peculiar among landhoppers. Talitroides al- 
luaudi also resembles Arcitalitrus sylvaticus, 
a non-cuspidactylate landhopper, in having 
simple gnathopod 1, weakly setose anterior 
margin in gnathopod 2, reduced and plu- 
mose-setose peduncle of pleopods, similar 
spination in uropods 1 and 2, etc. But the 
latter is distinguished from the former by 
the possession of a strongly arcuate outer 
plate and less setose maxilliped. It is inter- 
esting that 7. al/uaudi has a weakly arcuate 
inner plate. 

The distribution of 7. alluaudi in conti- 
nental areas is believed to be due to syn- 
anthropic dispersal (Friend & Lam 1985), 
and the occurrence is thought to be limited 
by the humidity (Andersson 1962). Thus it 
may be astonishing that 7. a//uaudi has been 
discovered in an arid part of Israel with pre- 
cipitation as little as 200 mm annually. So 
far in Israel the present species is known 
solely from the bamboo stands growing on 
a small “island’’ in the middle of a small 
pool in the University campus at Beersheba. 
The bamboo, now commonly utilized for 
gardening in Israel, is an exotic plant for 


337 


this country, most of which were imported 
from West Europe. Thus it is highly prob- 
able that the population of 7. al/uaudi in 
Beersheba was introduced accidentally from 
European countries with the bamboo and 
has survived on this small “‘island,’’ where 
a humid microclimate may have prevailed. 


Acknowledgments 


We are grateful to D. Defaye in Muséum 
National d’Histoire Naturelle, Paris for the 
loan of the type material treated here. 
Thanks to J. Aoki in Yokohama National 
University who provided us with compar- 
ative material. Thanks also to K. Iwatsuki 
of The University of Tokyo who helped lo- 
cate the type material. 


Literature Cited 


Andersson, A. 1962. Ona collection of Amphipoda 
of the family Talitridae from the Canary Is- 
lands.— Arkiv for Zoologi 15(11):211-218. 

Bousfield, E. L. 1984. Recent advances in the sys- 
tematics and biogeography of landhoppers (Am- 
phipoda: Talitridae) of the Indo-Pacific re- 
gion.—Pp. 171-210 in F. J. Radovsky, P. H. 
Raven & S. H. Sohmer, ed., Biogeography of 
the tropical Pacific, Bishop Museum Special 
Publication 72. 

Chevreux, E. 1896. Recherches zoologiques dans les 

serres du Muséum de Paris IV.—Sur un Am- 

phipode terrestre exotique, Talitrus Alluaudi nov. 
sp., acclimaté dans les serres du Jardin des 

Plantes de Paris.— Feuille des Jeunes Natural- 

istes 26:112-113. 

. 1901. Mission scientifique de M.Ch.Alluaud 

aux iles Séchelles. Crustacés Amphipodes.— 

Mémoires de la Societé Zoologique de France 

14:388-438. 

—, & L. Fage. 1925. Faune de France 9. Am- 
phipodes. Paul Lechevalier, Paris, 486 pp. 

Friend, J. A., & P. K. S. Lam. 1985. Occurrence of 
the terrestrial amphipod Talitroides topitotum 
(Burt) on Hong Kong Island.— Acta Zootaxon- 
omica Sinica 10(1):27-33. 

——, & A.M. M. Richardson. 1986. Biology of 
terrestrial amphipods.— Annual Review of En- 
tomology 31:25-48. 

Medcof, J.C. 1940. Variations in the pleopod struc- 
ture of the terrestrial amphipod Talitrus alluau- 
di Chevreux.—Lloydia 3(1):79-80. 

Menzel, R. 1911. Exotische Crustaceen im botan- 


338 


ischen Garten zu Bazel.— Revue Suisse de Zool- 
ogie 19:433-444. 

Palmén, E. 1949. Talitroides alluaudi (Chevreux) 
(Amphipoda, Talitridae) in Finnland gefun- 
den.—Archivum Societatis Zoologicae Botani- 
cae Fennicae ‘Vanamo’ 2:61-64. 

Shoemaker, C. R. 1936. The occurrence of the ter- 
restrial amphipods, Talitrus alluaudi and Tali- 
trus sylvaticus, in the United States.— Journal of 
the Washington Academy of Sciences 26(2):60- 
64. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


(HM) Department of Biology, Ibaraki 
University, Mito 310, Japan; (RO) Depart- 
ment of Ecology, Systematics and Evolu- 
tion, The Hebrew University of Jerusalem, 
Givat Ram, Jerusalem 91904, Israel. 


PROC. BIOL. SOC. WASH. 
106(2), 1993, pp. 339-345 


A DESCRIPTION OF A NEW SPECIES OF 
MACROBRACHIUM FROM PERU, AND DISTRIBUTIONAL 
RECORDS FOR MACROBRACHIUM BRASILIENSE (HELLER) 
(CRUSTACEA: DECAPODA: PALAEMONIDAE) 


Guido A. Pereira S. 


Abstract. —A systematic study is made on a collection of freshwater shrimps 
(Palaemonidae) from northern Pert collected by Dr. R. W. McDiarmid during 
the years 1977-1980. A new species is described Macrobrachium depressi- 
manum, and Macrobrachium brasiliense (Heller) is reported. The diversity of 
palaemonids in the Amazon region is discussed. 

Restimen. —Se realiza un estudio sistematico sobre una coleccion de cama- 
rones de agua dulce del norte del Peru, colectados por el Dr. R. W. McDiarmid 
durante los anos de 1977 y 1980. Se describe una nueva especie, Macrobrachium 
depressimanum, y se reporta la especie Macrobrachium brasiliense (Heller). Se 
discute acerca de la alta diversidad de camarones palaemonidos en la region 


del Amazonas. 


The present work is based on collections 
made primarily by Dr. R. W. McDiarmid 
in northern Pert during 1977 and 1980. The 
specimens were collected during field re- 
search in ethnobiological anthropology, 
conducted in the region of the Alto Mara- 
non River by scientists from the University 
of California. The collection has many sam- 
ples of two species of the freshwater shrimp 
genus Macrobrachium Bate, 1868 and sev- 
eral species of crabs of the families Tricho- 
dactylidae and Pseudothelphusidae. Only 
the shrimps are reported in this paper. De- 
scriptions are based on adult males. Total 
length (TL) is taken from the tip of the ros- 
trum to the posterior border of telson. Car- 
apace length (CL) is taken from the ocular 
margin to the posterior border of carapace. 
Only partial synonymy is given for M. brasi- 
liense (Heller, 1862). All material is depos- 
ited in the National Museum of Natural 
History, Smithsonian Institution, Washing- 
ton D.C., U.S.A. (USNM). 


Macrobrachium depressimanum, 
new species 
(Figs. 1, 2) 


Type material.— Holotype, 1 6, USNM 
230032, 35.5) mm), 6.2 mm (CL); Pert, 
Amazonas Department, Santiago River, La 
Poza; 4°O1'S. 77°47’ W., 26 Jan 1980, coll. 
R. W. McDiarmid. Paratypes, 2 2 (1 ovig.), 
USNM 230030; 4 2 (2 ovig.), 14 6, USNM 
230031 (locality and date, same as for ho- 
lotype). 

Additional material examined.—108 8, 
133 6, from the following localities in Peru: 
Amazonas Department, Santiago River, La 
Poza, 4°01’S, 77°47'W; Caterpiza River, 
3°55'S, 77°42'W; Cenepa River near Huam- 
pami, 4°28’S, 78°10’W; Santiago River at 
Pinglo, at confluence between Santiago Riv- 
er and Maranon River, 4°26’S, 77°39’'W: 
Maranon River across from Santa Maria de 
Nieva, and at confluence of Nieva River 
with Maranon River, 4°35'S, 77°77'W; 


340 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


F 
B 
1mm 
G 
H 
Cc 
N 
\ 
S 
l 
E 
1mm 


Fig. 1. Macrobrachium depressimanum, new species. A, carapace and cephalic appendages; B, carapace; C, 
posterior region of telson; D, telson; E, appendix masculina and appendix interna; F, first pereiopod; G, second 
pereiopod; H, third pereiopod; I, fifth pereiopod. 


Ucayali, Tahuallo River, Pucalpa, 8°23’S, reiopod smooth; ratio palm length/finger 

74°32'W; Ucayali, Tahuallo River, Pucalpa, length about 0.6; palm flattened anterodor- 

Neshuya, 8°23’S, 74°32'W; Madre de Dios, _ sally (Fig. 1G). 

Tambopata Natural Reservoir, La Torre Description. —Rostrum nearly straight, 

River, 11°58’S, 69°11’W. slightly arched over eyes, reaching (or slight- 
Diagnosis — Adult males with second pe-__ly overreaching) distal border of scaphoce- 


VOLUME 106, NUMBER 2 341 


\- > ? 
| \ ! 
i 


16675 4 KU Sau 


« 


—e 


~ 


Fig. 2. Detail of appendix masculina. A, Macrobrachium depressimanum, new species, anterior view; B, 
Macrobrachium jelskii (Miers, 1877), lateral view. Same scale for A and B. 


342 


rite with apex, tip pointing upwards. Upper 
border bearing 11 (9-11 in paratypes) teeth, 
2 proximal teeth more spaced behind the 
orbit. Lower margin bearing 5 (4—6 in para- 
types) teeth (Fig. 1A, B). Carapace smooth. 
Scaphocerite 2.5 times longer than wide. 
Abdomen smooth, posteroventral angle of 
fifth pleuron acute; sixth abdominal seg- 
ment 1.5 times length of fifth, same length 
as telson. Telson with 2 pairs of dorsal spines 
situated at 14 and /4 its length from base. 
Posterior margin, tapering abruptly to me- 
dian apex bearing 2 pairs of lateral spines. 
Inner pair of spines overreaching median 
apex. Four plumose setae present on pos- 
terior margin between inner spines (Fig. 1C, 
D). First pereiopods slenderer than second 
pair, reaching scaphocerite with tips of fin- 
gers; palm cylindrical, 0.9 times length of 
dactyl. Carpus 3.1 times length of palm, 1.1 
times that of merus (Fig. 1F). Second pair 
of pereiopods equal, smooth, reaching an- 
terior border of scaphocerite with distal car- 
pus. Fingers thin, closing over entire length 
of chela, without conspicuous teeth. Palm 
flattened anterodorsally, 2.1 times longer 
than high, 0.6 times length of dactyl. Carpus 
1.5 times length of palm, 0.9 times that of 
merus (Fig. 1G). Third pair of pereiopods 
failing to reach distal border of scaphocer- 
ite. Propodus with longitudinal row of 4 
spines on inner margin, 1.3 times length of 
dactyl, 1.7 times that of carpus (Fig. 1H). 
Fifth pair of pereiopods reaching distal bor- 
der of scaphocerite with tip of dactyl. Prop- 
odus with longitudinal row of 6 spines on 
inner margin, 2 times length of dactyl, 1.7 
times iength of carpus (Fig. 11). Appendix 
masculina 2.1 times longer than appendix 
interna (Fig. 1E), with acute apex, and 18 
short, smooth spines. 

Fecundity. —One ovigerous female, TL = 
38 mm, CL = 7.1 mm, with 23 eggs, 1.2 
mm largest diameter. 

Distribution. —Dr. Célio Magalhaes from 
Instituto Nacional de Pesquisas da Ama- 
zonia (INPA), Brazil, has informed me that 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


there are specimens of this species in the 
invertebrate collection of his Institute, from 
the following localities in Brazil: Amazonas 
State, Tefé River, Solimoes, 3°35’S, 
64°47'W; Solimoes River, Catalao coast, 
near Manaus, 3°8’S, 60°1’W; Acre State, 
Tarauaca River, Tarauaca, 6°42’S, 69°48’W; 
Rondonia State, Madeira River, fall of Teo- 
tonio, 3°22’S, 58°45'W; Mamoré River, near 
Guajara-Mirim, 10°23’S, 65°23'W; Madei- 
ra River, near Guajara-Mirim, 10°23’S, 
65°23’W. Based on these data the species is 
distributed over the upper Amazon Basin, 
from Pert to northeastern Brazil. 

Etymology.—The specific name is de- 
rived from a composite of the Latin words 
depressus meaning dorsoventrally flattened 
and manus meaning hand. It stresses the 
typical shape of the anterior part of the palm 
in this species. 

Remarks.—The present species is mor- 
phologically related to Macrobrachium jel- 
skii (Miers, 1877). M. depressimanum has 
more teeth on the dorsal border of the ros- 
trum. The formula for M. jelskii is 6-9 (1 
post-ocular) whereas it is 10-12 (2 post-oc- 
ular) in M. depressimanum. The relative 
proportion of segments in the second pe- 
reiopods is very distinctive, the ratio be- 
tween palm length/dactyl length is 0.6 in M. 
depressimanum whereas it is 1.5 in M. jel- 
skii. The anterodorsally flattened shape of 
the palm is a typical feature of M. depres- 
simanum. 

The shape of the appendix masculina is 
not of common usage in the taxonomy of 
Macrobrachium. However, the appendix 
masculina of M. depressimanum is very dis- 
tinctive. It is 2.1 times longer than the ap- 
pendix interna, whereas in M. jel/skii it is 
about 1.5 times longer. The appendix in- 
terna of M. depressimanum is also more 
slender compared to that of M. jelskii (Fig. 
2). The two species are allopatric but there 
is a possibility of some overlap since M. 
Jelskii was reported by Holthuis (1966) as 
occurring in north-east Brazil. 


VOLUME 106, NUMBER 2 


Macrobrachium brasiliense (Heller, 1862) 


Palaemon brasiliensis. —Heller, 1862:419. 

Macrobrachium brasiliense. —Holthuis, 
1952:79, pl. 19.—Holthuis, 1959:93.— 
Rodriguez, 1982:382. 


Material examined. —63 2, 114 6, 112 ju- 
veniles, from the following localities in Peru, 
Amazonas Department: Santiago River, La 
Poza, 4°01’S, 77°47'W; Castillo Creek 
(mouth enters Santiago River at 1.5 km 
south of La Poza), 4°02’S, 77°47'W;; at Ping- 
lo, confluence at Santiago River and Mara- 
non River, 4°26’S, 77°39'W; way to Galilea, 
creek in the forest, 4°15’S, 77°49’ W; Galilea, 
4°01'S, 77°49'W; Yutapis River, Shiringa, 
Awies: 77 51'W; Caterpiza River, 3°55'S, 
77°43'W; Cenepa River, 3°55’S, 77°43'W. 

Diagnosis. — Adult males with short spines 
on carapace, specially on anterolateral bor- 
ders. The larger of the second pair of pereio- 
pods has a finger/palm length ratio of 2.0- 
2.3. Fixed finger bears a row of 4 to 11 
tubercles along inner border. 


Discussion 


Since the revision of the Palaemonidae 
by Holthuis (1952), several new species of 
palaemonid shrimps have been described 
from the Amazon Region (Tiefenbacher 
1978; Ramos-Porto 1979; Kensley & Walk- 
er 1982; Rodriguez 1982; Pereira 1985, 
1986, 1991). The latter authors report spe- 
cies that, judged by the small brood and 
somewhat large size of the eggs, can be con- 
sidered to have abbreviated larval devel- 
opment and a strictly land-locked fresh- 
water habitat (Sollaud 1923a, 1923b; 
Shokita 1973, 1977, 1979; Gamba 1984; 
Magalhaes 1985, 1986, 1988; Chong & 
Khoo 1987a, 1987b). The Amazon Region 
has the largest number of strictly freshwater 
species (24) of palaemonids in the world 
(Pereira 1989). This agrees with the tradi- 
tional view of taxonomists and biogeogra- 


343 


phers that consider the Amazon Region as 
an area with high endemism and diversity, 
probably as a result of both environmental 
heterogeneity and the long and complex pa- 
leogeographic history of the basin (Prance 
1982). The upper Amazon Basin (east and 
northeast portion) was formed most re- 
cently during Pliocene times, whereas the 
middle basin was formed during the Cre- 
taceous (Putzer 1984, Rodriguez & Pereira 
1992). The complexity and long history of 
the basin led to the emergence of strictly 
freshwater palaemonids in at least four gen- 
era, Macrobrachium, Palaemonetes Heller, 
1869, Euryrhynchus Miers, 1877, and Pseu- 
dopalaemon Sollaud, 1911, according to 
Magalhaes & Walker (1988), and Pereira 
(1989). The latter two genera are endemic 
to South America, and all four are strictly 
freshwater with abbreviated larval devel- 
opment. 

An interesting argument arises when con- 
sidering the phylogenetic history of the fam- 
ily (Pereira 1989). There is evidence of an 
early freshwater origin of this group because 
the closest relatives of the primitive genus 
Euryrhynchus occur in West African fresh- 
waters (Powell 1976). This raises questions 
about the tempo and mode of evolution of 
the family and their invasion into the ma- 
rine and freshwater habitat (Pereira 1989). 
Magalhaes & Walker (1988) considered ab- 
breviated development as a derived con- 
dition. They concluded that there has been 
massive intergeneric convergence toward 
this type of life cycle in the Palaemonids 
mainly because of selective pressure to live 
in plankton-poor waters. However, a dif- 
ferent point of view arises if the abbreviated 
development is considered the primitive 
condition. It could be argued that the life 
cycle with complete larval development, and 
acquired independently only by Macrobra- 
chium and Palaemonetes, is the derived 
condition. The complete development 
eventually permitted these freshwater 
shrimps to be successful in colonizing more 


344 


productive environments like estuaries and 
mixohaline waters. 


Acknowledgments 


I wish to thank Dr. M. Reaka, Dr. R. W. 
McDiarmid, and Dr. C. Magalhaes for re- 
viewing the manuscript. Special thanks to 
Dr. R. B. Manning for his advice and pro- 
viding working space at the Smithsonian 
Institution, Division of Crustacea, during 
1984-1988. Thanks to Dr. C. Magalhaes 
from INPA, Brazil, and Lic. H. Ortega from 
Universidad de San Marcos, Pert, for gen- 
erously providing additional data on the 
species. Sheila Pauls kindly helped me in 
the final editing of this manuscript. Support 
for present research was provided by Con- 
sejo de Desarrollo Cientifico y Humanisti- 
co, Universidad Central de Venezuela. 


Literature Cited 


Bate, C. S. 1868. On a new genus, with four new 
species, of freshwater prawns.— Proceedings of 
the Zoological Society of London 1868:363-368. 

Chong, S. S. C., & H. W. Khoo. 1987a. Abbreviated 
larval development of the freshwater prawn, 
Macrobrachium pilimanus (De Man, 1879), 

(Decapoda, Palaemonidae), reared in the labo- 

ratory.—Journal of Natural History 21:763-774. 

1987b. The abbreviated larval development 
of the freshwater prawn, Macrobrachium ma- 
layanum (Roux, 1934), (Decapoda, Palaemon- 

idae), reared in the laboratory.—Crustaceana 53: 

29-42. 

Gamba, A.L. 1984. Different egg-associated and lar- 
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chium jelskii and Macrobrachium amazonicum 
(Arthropoda: Crustacea) in a Venezuelan con- 
tinental lagoon.—International Journal of In- 
vertebrate Reproduction and Biology 7:135—142. 

Heller, C. 1862. Beitrage zur naheren Kentniss der 

Macrouren.—Sitzungs Berichte der Akademie 

Wissenschaften in Wiens 45:389-426. 

1869. Zur naheren kenntniss der in den siis- 
sen gewassern des sudlichen Europa vorkom- 
mendem Meerescrustaceen.—Zeitschrift wis- 
senschaffliche Zoologie 19:156—-162. 

Holthuis, L. B. 1952. A general revision of the Pa- 
laemonidae (Crustacea, Decapoda, Natantia) of 
the Americas. II. The subfamily Palaemoni- 
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dation Publications, 12, 396 pp. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


. 1959. The Crustacea Decapoda of Suriname 
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44:1-296. 

1966. A collection of freshwater prawns 
(Crustacea, Decapoda, Palaemonidae) from 
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lier.— Bulletin Institute Royale Sciences Natu- 
relle, Belgian 42:1-11. 

Kensley, B., & I. Walker. 1982. Palaemonid shrimps 
from the Amazon Basin, Brazil (Crustacea: De- 
capoda: Natantia).—Smithsonian Contribu- 
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Magalhaes, C. 1985. Desenvolvimento larval obtido 

em laboratorio de palaemonideos da Regiao 

Amazonica. I. Macrobrachium amazonicum 

(Heller, 1862) (Crustacea, Decapoda).—Ama- 

zoniana 9:247-274. 

1986. The larval development of the palae- 
monid shrimps from the Amazon region reared 
in the laboratory. IV. Abbreviated development 
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tacea: Decapoda).—Amazoniana 10:63-78. 

. 1988. The larval development of Palaemonid 

shrimps from the Amazon Region reared in the 

laboratory. II. Extremely abbreviated larval de- 
velopment in species of the genus Euryrhynchus 

Miers, 1877 (Crustacea, Decapoda, Euryrhyn- 

chinae).—Crustaceana 55:39-52. 

,& I. Walker. 1988. Larval development and 

ecological distribution of central amazonian Pa- 

laemonid shrimps (Decapoda, Caridea). —Crus- 
taceana 55:279-292. 

Miers, E. J. 1877. Ona collection of Crustacea, De- 
capoda and Isopoda, chiefly from South Amer- 
ica, with descriptions of new genera and spe- 
cies.— Proceedings of the Zoological Society of 
London 1877:653-679. 

Pereira, G. 1985. Freshwater shrimps from Vene- 
zuela III: Macrobrachium quelchi (De Man) and 
Euryrhynchus pemoni n. sp. (Crustacea, Decap- 

oda, Palaemonidae) from La Gran Sabana.— 

Proceedings of the Biological Society of Wash- 

ington 98:615-621. 


. 1986. Freshwater shrimps from Venezuela I: 
seven new species of Palaemoninae. — Proceed- 
ings of the Biological Society of Washington 99: 
191-213. 


. 1989. Cladistic, taxonomy, biogeography and 
the evolutionary history of the shrimp family 
Palaemonidae (Crustacea, Decapoda, Caridea). 
Unpublished Ph.D. Dissertation, University of 
Maryland, College Park, 417 pp. 


. 1991. Camarones de agua dulce de Venezuela 
II: neuvas adiciones en las familias Atydae y 
Palaemonidae (Crustacea, Decapoda, Carid- 
ea).—Acta Biologica Venezuelica 13:75-88. 

Prance, G. T. (ed.). 1982. Biological Diversification 


VOLUME 106, NUMBER 2 


in the Tropics. Columbia University Press, New 
York, xvi + 714 pp. 

Powell, C. B. 1976. Two new freshwater shrimps from 
West Africa: the first euryrhynchinids (Decap- 
oda, Palaemonidae) reported from the old 
world.—Revue de Zoologie Africaine 91:649- 
674. 

Putzer, H. 1984. The geological evolution of the Am- 
azon Basin and its mineral resources.: Pp. 14— 
46 in H. Sioli, ed., The Amazon. Limnology and 
landscape ecology of a mighty tropical river and 
its basin. Dr. W. Junk Publisher, The Nether- 
lands, ix + 763 pp. 

Ramos-Porto, M. 1979. Pseudopalaemon amazo- 
nensis, especie nova de camarao de Bacia Ama- 
zonica (Crustacea, Decapoda, Palaemonidae). — 
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cia, 3la Reuniao Anual, Resumos, Suplemento 
de Ciencia e Cultura 31:693. 

Rodriguez, G. 1982. Fresh-water shrimps (Crustacea, 
Decapoda, Natantia) of the Orinoco Basin and 
the Venezuelan Guayana.—Journal of Crusta- 
cean Biology 2:378-391. 

—, & G. Pereira. 1992. New species, cladistic 
relationships, and biogeography of the genus 
Fredius (Decapoda: Brachyura: Pseudothelphu- 
sidae) from South America.—Journal of Crus- 
tacean Biology 12:298-311. 

Shokita, S. 1973. Abbreviated larval development of 
the freshwater prawn, Macrobrachium shokitai 
Fujino et Baba (Decapoda, Palaemonidae) from 


345 


Iriomote Island of the Ryukyus. — Annotationes 
Zoologicae Japonenses 46:11 1-126. 

1977. Abbreviated metamorphosis of land- 
locked freshwater prawn, Macrobrachium as- 
perulum (Von Martens) from Taiwan.—Anno- 
tationes Zoologicae Japonenses 50:1 10—122. 

1979. Early life history of the genus Mac- 

robrachium (Decapoda, Palaemonidae).— Uni- 
versity of the Ryukyus 1:9-17. 
Sollaud, E. 1911. Pseudopalaemon bouvieri, nouveau 
genre, nouvelle espéce, de la famille des Palae- 
monidae.— Bulletin de Museum de Histoire Na- 
turelle, Paris 17:12—16. 

1923a. Le developpement larvaire des Pa- 
laemoninae. I. Partie descriptive. La conden- 
sation progressive de l’ontogenése.— Bulletin 
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510-603. 

1923b. Recherches sur l’embryogénie des 
Crustacés Décapodes de la sous-famille des Pa- 
laemoninae.— Bulletin Biologique de la France 
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Instituto de Zoologia Tropical, Univer- 
sidad Central de Venezuela, Aptdo. 47058, 
Caracas 1041-A, Venezuela. 


PROC. BIOL. SOC. WASH. 
106(2), 1993, pp. 346-352 


A NEW SPECIES OF CRAYFISH (DECAPODA: 
CAMBARIDAE) BELONGING TO THE GENUS CAMBARUS, 
SUBGENUS HIATICAMBARUS, FROM THE UPPER 
ELK RIVER DRAINAGE OF WEST VIRGINIA 


Raymond F. Jezerinac and G. Whitney Stocker 


Abstract. —A new species of crayfish, Cambarus (Hiaticambarus) elkensis, is 
described from the upper Elk, Holly, and Birch rivers of West Virginia. The 
species is most closely related to C. (H.) chasmodactylus but differs from it in 
having shorter fingers, a much narrower gape between the fingers, and moderate 
impressions at the base of the fixed finger, especially on the ventral surface. 


While conducting a crayfish survey of the 
state of West Virginia in 1988, we captured 
what we thought to be C. (H.) chasmodac- 
tylus James (1966) from the upper Elk River 
drainage. We reported this (Jezerinac & 
Stocker 1989:3) as a new drainage record 
for the species. Additional collections were 
made in 1989 from the upper Elk basin, the 
Elk River and its tributaries below Sutton 
Lake, as well as in surrounding watersheds, 
notably the Greenbriar, Gauley, Cherry, and 
Cranberry rivers. Representatives of the 
species were caught only in the upper Elk 
River drainage. With the additional mate- 
rial, we noted that these crayfish differ in 
several respects from C. (H.) chasmodac- 
tylus and are described herein as members 
of a new species. 


Cambarus (Hiaticambarus) elkensis, 
new species 
Fie 1s Tablet 


Cambarus bartonii veteranus. —Faxon, 
1914:389 [in part, Elk River, Cogar’s 
Mills, West Virginia]. 

Cambarus (Hiaticambarus) chasmodacty- 
lus.—Jezerinac & Stocker, 1989:3 [in 
part]. 


Diagnosis.—Body pigmented, eyes well 
developed. Carapace subovate, dorsoven- 
trally flattened. Rostrum with weakly con- 


vergent, slightly concave, thickened mar- 
gins, lacking spines or tubercles and 
terminating in upturned corneous tubercle. 
Areola 3.5 to 5.5 (X = 4.6) times as long as 
wide and comprising 35.1 to 38.1% (X¥ = 
36.6%) of entire length of carapace, and 
bearing 5 to 7 punctations across narrowest 
part. Cervical spine absent or represented 
by small tubercle. Suborbital angle obtuse 
to obsolete. Postorbital ridges terminating 
cephalically in sharp spine or tubercle. 
Branchiostegal spine very small. Antennal 
scale 2.5 times longer than broad, with me- 
sial and lateral margins subparallel near and 
at midlength; basiopodite with very small 
spine; ischiopodite lacking spine. Epistome 
subtriangular, zygoma with about 120° arch. 
Chela smooth and bearing one row of very 
indistinct tubercles along mesial margin of 
palm; lateral margin of fixed finger smooth; 
both fingers with very poorly defined dor- 
somedian longitudinal ridges; fixed finger 
moderately impressed at base especially on 
ventral surface; dactyl 1.8 to 2.3 (X = 2.0) 
times longer than mesial margin of palm; 
palm length 28.4 to 32.3% (X = 30.8%) of 
chela length; elongated setae at base of fixed 
finger very sparse or absent (best seen on 
young specimens); gape of fingers subequal 
to width of dactyl in first form males, less 
so in second form males, and almost non- 
existent in females. Mesial margin of carpus 


VOLUME 106, NUMBER 2 


of chela with large spiniform tubercle and 
smaller conical one proximally; ventral sur- 
face with conical knob on distal articular 
rim. Ventrolateral ridge of merus with 2 to 
4 (X = 2.5) spines; ventromesial one with 
6 to 12 (X = 9) spines. Ischium of only third 
pereiopod with simple hook not reaching 
tubercle on corresponding basis. Boss on 
ischium of fourth pereiopod prominent. 
First pleopod of Form I male (Fig. 1B, F) 
with short terminal elements; corneous cen- 
tral projection truncate distally, recurved at 
greater than 90° to main shaft of appendage, 
with subapical notch; mesial process in- 
flated, tapering, rounded to acute distally, 
directed caudolaterally at angle slightly 
greater than 90° to main shaft of appendage. 
Female with annulus ventralis shallowly 
embedded in sternum, asymmetrical, and 
subrhomboidal. (Additional morphometric 
and meristic data, such as simple descrip- 
tive statistics, ratios, and regression analy- 
sis, may be obtained from the authors or 
the library at The Ohio State University at 
Newark.) 

Holotypic male, Form I.—Cephalothorax 
(Fig. 1A, J) subovate in cross section, dor- 
soventrally compressed. Abdomen narrow- 
er than thorax (21.0 mm and 17.4 mm); 
greatest width of carapace distinctly greater 
than height at caudodorsal margin of cer- 
vical groove (21.0 mm and 15.2 mm). Are- 
ola 4.5 times as long as broad with 5 to 7 
crowded punctations across narrowest part; 
length of areola 37.5% of total length of car- 
apace. Rostrum with thickened, weakly 
convergent, elevated, concave margins de- 
void of spines or tubercles; dorsal surface 
of rostrum slightly concave with many 
punctations, rather sparse punctations on 
and at base of indistinctly delineated acu- 
men. Subrostral ridges well developed and 
evident in dorsal aspect to base of acumen. 
Suborbital angle obtuse. Postorbital ridge 
moderately prominent, grooved dorsolater- 
ally, and terminating cephalically in acute 
corneous tubercule. Branchiastegal spine 
represented by a spiniform tubercle. Cer- 


347 


vical spine reduced to small rounded tu- 
bercle on left side, absent on right. Carapace 
densely punctate dorsally except in gastric 
region, distinctly sculptured over attach- 
ment of mandibular muscle; lateral surface 
with many small granules in branchiostegal 
region; larger granules in mandibular and 
ventral half of hepatic region. 

Abdomen slightly shorter than carapace 
(38.6 mm and 40.0 mm respectively); pleu- 
ra rounded to subtruncate ventrally with an- 
gular caudoventral extremities on third 
through fifth segments. Cephalic section of 
telson with 2 spines on each caudolateral 
corner, mesial spines movable; borders of 
caudal section evenly rounded. Proximal 
podomere of uropod with both lobes ter- 
minating in corneous-tipped blunt spine: 
mesial ramus with low submedian dorsal 
keel ending in small, blunt, premarginal 
spine not extending beyond distal edge of 
ramus. 

Cephalomedian lobe of epistome (Fig. 11) 
broadly rounded with small cephalomedian 
projection, margin somewhat thickened and 
elevated ventrally; main body with distinct 
median groove and paired slitlike grooves 
immediately cephalic to arched epistomal 
zygoma. Ventral surface of proximal podo- 
mere of antennular peduncle with small 
spine at base of distal third. Antennal pe- 
duncle with strong lateral spine on basis, 
remaining podomeres lacking spines. An- 
tennal scale (Fig. 1G) about 2.5 times as 
long as broad with mesial and lateral mar- 
gins subparallel for some distance proximal 
and distal to midlength; strong distolateral 
spine reaching beyond midlength of ulti- 
mate podomere of antennular peduncle. 
Ventral surface of ischium of third maxil- 
liped with broad, longitudinal band of long 
setae laterally and with submarginal lateral 
row of smaller plumose ones, few additional 
short plumose setae in area between; mesial 
margin with 21 denticles. 

Right chela (detached) (Fig. 1L) 2.4 times 
as long as broad, mesial margin of palm 
30.6% of chela length; dactyl 2.1 times palm 


348 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


length; mesial margin of palm with one row 
of 7 indistinct tubercles; remainder of palm 
with widely spaced large, deep punctations 
on mesial region, more crowded on lateral 
region, lateral surface rounded. Gape be- 
tween fingers subequal to dactyl width, and 
proximal half of opposable surface of fixed 
finger without tufts of setae; both fingers 
with poorly defined median longitudinal 
ridge on dorsal surface, absent on ventral 
surface, both with conspicuous deep punc- 
tations; opposable margin of fixed finger with 
row of 11 small (seventh enlarged) rounded 
corneous tubercles, single row of minute 
denticles present slightly ventral to tuber- 
cular row along distal fourth of finger; op- 
posable margin of dactyl with row of 12 
tubercles, minute denticles forming single 
row below level of tubercles along distal 
third. Lateral surface of fixed finger and me- 
sial surface of dactyl non-costate, punctate. 

Carpus of cheliped (Fig. 1L) longer than 
broad with deep dorsal furrow flanked by 
sparse punctations mesially and laterally; 
mesial surface with | spiniform tubercle and 
more proximally situated smaller conical 
one; ventral surface very sparsely punctate 
and bearing 1 tubercle on distal articular 
rim. Merus with 1 premarginal tubercle dor- 
sally; ventrolateral ridge with 2, ventrome- 
sial ridge with 9 tubercles. Ventromesial 
margin of ischium with 2 very small tuber- 
cles. . 

Hook on ischium of third pereiopod only 
(Fig. 1K), hook overreaching basioischial 
articulation and opposed by small tubercle 
on basis. Coxa of fourth pereiopod with 
prominent caudomesial boss ventrally dis- 
posed, and somewhat flattened caudally. 
Boss on coxa of fifth pereiopod vestigial. 
First pleopods (Fig. 1B, C) reaching coxae 
of third pereiopods, symmetrical, with very 
small gap between bases (Fig. 1 D). (See “Di- 
agnosis” for descriptions of the pleopods 
and Fig. 1B, F.) 

Allotypic female.—Excluding secondary 
sexual characteristics, differing from holo- 
type in following respects: areola 5.0 times 


as long as broad and constituting 36.6% of 
total length of carapace; very small cervical 
tubercle present on both sides; mesial mar- 
gin of palm of right chela 30.9% of chela 
length; gape between fingers about % width 
of dactyl; few plumose setae present at base 
of fixed finger; no enlarged tubercle on op- 
posable surface of fixed finger; 10 (right) and 
11 (left) tubercles on opposable margin of 
dactyl; 2 dorsal premarginal tubercles on 
merus; ventrolateral ridge of merus with 3 
(right) and 2 (left) tubercles; ventromesial 
ridge of merus with 11 (right) and 12 (left) 
tubercles. 

Annulus ventralis subrhomboidal (Fig. 
1H), wider than long, slightly movable, with 
caudal wall weakly developed. Postannular 
sclerite about half as wide and about 4 as 
long as annulus. First pleopods reaching 
midlength of annulus when abdomen flexed. 

Morphotypic male, Form II.—Differing 
from holotype in following respects: areola 
4.2 times as long as broad; areola length 
36.2% of carapace length; cervical tubercle 
absent on both sides; spine on ventral sur- 
face of proximal podomere of antennuar pe- 
duncle near distal margin; spine on right 
antennal scale damaged, antennal scale 2.3 
times longer than broad; right chela regen- 
erated; left chela 2.3 times as long as broad; 
mesial surface of palm occupying 29.7% of 
chela length; dactyl length 2.2 times palm 
length; gape about *4 width of dactyl; setae 
at base of fixed finger; ventrodistal margin 
of carpus with 2 tubercles; merus with 2 
premarginal tubercles dorsally, 2 large and 
1 small (left) and 4 (right) tubercles on ven- 
trolateral ridge of merus; ventromesial ridge 
with 9 (right) and 7 (left) tubercles; ven- 
tromesial margin of ischium with 4 (right) 
and 3 (left) tubercles; hook on ischium of 
third pereiopod much reduced, not over- 
reaching basioischial articulation and op- 
posed by small tubercle on basis; first pleo- 
pod (Fig. 1C, E) with juvenile suture on shaft, 
central projection inflated. 

Color notes.— Basic color of dorsal sur- 
faces of chela, carpus, merus, legs, and car- 


VOLUME 106, NUMBER 2 349 


Fig. 1. Cambarus (Hiaticambarus) elkensis, new species. All from holotype male, Form I, except C, E, from 
morphotype male. Form II, and H. from allotype female): A, lateral view of carapace: B, C. mesial view of first 
pleopod:; D, caudal view of first pleopods; E, F, lateral view of first pleopod: G. antennal scale. H, annulus 
ventralis; I, epistome; J, dorsal view of carapace; K. proximal podomeres of third, fourth, and fifth pereiopods; 
L, dorsal view of distal podomeres of cheliped. (See Table 1 for precise measurements.) 


350 


apace brown; abdomen darker brown; tu- 
bercles on opposable margins of fingers and 
mesial spiniform tubercle of corpus yellow- 
ish; distinct black band lining cervical 
groove; mandibular adductor region with 
black reticulated pattern; rostral margins and 
dorsal tubercles on chela reddish; under- 
surface cream. 

Types. —The holotype, allotype, and 
morphotype (USNM 260038, 260039, and 
260040, respectively) are deposited in the 
National Museum of Natural History, 
Smithsonian Institution, Washington D.C. 
A small series of paratypes (1 61, 8 4 II, 10 
?)is at The Ohio State University at Newark 
Crayfish Museum. 

Type locality.—The Laurel Fork of the 
Left Fork of the Holly River (Holly River 
— Elk River — Kanawha River drainage) 
at Holly River State Park campground, 
Webster County, Hacker Valley District, 
West Virginia. (1.1 (air) km NNE of Hacker 
Valley.) At this site, the stream is about 10 
m wide, 0.5 m deep, and flows over a sub- 
strate of sandstone boulders, cobbles, grav- 
els, and sand. The specimens were captured 
from under cobbles lying on sands and grav- 
els where there was considerable current. 
The adjacent terrestrial vegetation included 
hemlock (Tsuga), birch (Betula), alder (A/- 
nus), and rhododendron (Rhododendron). 

Range and specimens examined.—The 
species is endemic to the upper Elk Basin, 
specifically in the Elk River above Sutton 
Lake, and in the Holly and Birch rivers. The 
collectors were R. F. Jezerinac, G. W. Stock- 
er and T. Jones (Coll 1) and GWS and TJ 
(Coll 2) unless otherwise stated. 

We have examined 22 collections con- 
taining a total of 95 specimens—6 (Form 1) 
males, 46 (Form II) males, and 43 females 
from the following localities: Nicholas 
County: Birch River Intersection (Inters) 
County Road (Co Rd) 10 and Co Rd 1/9 
(0.8 (air) km E of Birch River), 10 Sep 1988, 
Coll 1 (1 6 II); Pocahontas County: Slaty 
Fork (Fk.) United States Route (U S Rte) 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


219, 0.3 km (0.4 mi) S of Co Rd 219/12 
(0.3 (air) km SE of Slaty Fk.), 21 Aug 1988, 
Coll 2 (1 4 I, 6 9); Old Field Fk. U S Rte 
250, 12.8 km (8.0 mi) NW of Co Rd 219/ 
25 (at Marys Chapel), 19 Jul 1989, Coll 1 
(2 6 II, 2 2); Old Field Creek (Ck.) Inters U 
S Rte 219 and Co Rd 219/1 (3.0 (air) km S 
of Slaty Fk.), 27 May 1989, GWS, RFJ (1 
2); Webster County: Laurel Fk. Holly River 
State Park Campground (1.1 (air) km NNE 
of Hacker Valley), 17 Jun 1988, GWS, RFJ 
(4 611, 5 2); same locality 23 Aug 1988, Coll 
2(1 61,3 4Il, 3 2); and 20 Jun 1990, GWS, 
Chelsey Stocker, Vicky Stocker (1 6 I, 2 6 
II, 3 2); Right Fk. Inters St Rte 20 and Co 
Rd 5 (at Diana), 3 Sep 1988, GWS, RFJ (1 
2); 20 Jul 1989, Coll 1 (1 4 II, 1 2); (10) Birch 
R. Co Rd 30, 6.1 km (3.8 mi) N of St Rte 
20 (4.6 (air) km RW of Cowen), 23 Aug 
1988, Coll 2 (1 4 II, 1 2); Elk R. Co Rd 26, 
5.6 km (3.5 mi) NE of Co Rd 15 (3.0 (air) 
km NW of Bergoo), 22 Aug 1988, Coll 2 (4 
6 II, 4 9); Back Fk. Co Rd 24, 5.1 km (3.2 
km) E of St Rte 20 (2.2 (air) km E of Webster 
Springs), 20 Jul 1989, Coll 1 (1 61,64 I, 
6 2); Left Fk. Co Rd 3 just W of St Rte 20 
(1.3 (air) km SW of Hacker Valley), 21 Jul 
1989, Coll 1 (3 6 II, 1 2); Right Fk. Inters 
Co Rd 5/1 and Co Rd 15 (at Guardian), 21 
Jul 1989, Coll 1 (2 6 ID); Left Fk. Inters Co 
Rd 3 and Co Rd 8 (at Polling), 21 Jul 1989, 
Coll 1 (3 6 ID); Elk R. Inters Co Rd 26/1 
and Co Rd 26/7 (1.9 (air) km NE of Bergoo), 
20 Jul 1989, Coll 1 (1 6 I, 4 46 II, 1 9); Elk 
R. Co Rd 26, 5.6 km (3.5 mi) NE of Co Rd 
15 (3.0 (air) km NW of Bergoo), 20 Jul 1989, 
Coll 1 (2 6 I, 3 9, 1 2 ovig.); Elk R. Co Rd 
15/3, 0.8 km (0.5 mi) E of St Rte 15 (0.5 
(air) km SW of Curtin), 20 Jul 1989, Coll 1 
(1 3 II); Birch R. Inters Co Rd 30 and Co 
Rd 40 (at Boggs), 21 Jul 1989, Coll 1 (2 4 
II, 2 2); Leatherwood Cr. Co Rd 26/4, 0.8 
km (0.5 mi) S of Co Rd 26 (1.1 (air) km S 
of Bergoo), 20 Jul 1989, Coll 1 (3 6 II, 3 9); 
Elk R. Co Rd 7 at Webster Springs Water 
Treatment Plant (2.4 (air) km W of Webster 
Springs), 22 Jul 1989, Coll 1 (1 6 I); USNM 


VOLUME 106, NUMBER 2 


43706 Elk R., Cogar’s Mills, 30 Jul 1899, 
Collectors unknown (1 ¢ J) [locality un- 
known, Braxton County on USNM label]. 

Variations. —Most specimens examined 
have concave rostral margins that taper to 
the acumen and the rostral length is greater 
than the rostral width. Some specimens have 
subparallel rostral margins, others have 
margins that constrict rather sharply to form 
the acumen, and some have rather broad 
rostra. The suborbital angle varies from ob- 
solete to obtuse. These variations are not 
restricted to any specific watershed. 

Size. —The largest specimen examined is 
a Form II male from the Birch River in 
Webster County with a carapace length (CL) 
of 45.6 mm. The largest female has a 39.0 
mm CL and those of the largest and smallest 
Form I males are 41.2 mm and 36.7 mm, 
respectively. For measurements of the types 
see Table 1. 

Life history notes. —Form I males were 
captured on 20 June (1 specimen), 20 July 
(2), and 21 and 23 August (1 each). The only 
ovigerous female was caught on 21 July and 
carried 112 eggs having diameters of 2.2— 
2.6 mm. This female has a CL of 37.6 mm. 
Specimens are not available for the other 
months of the year. 

Ecological notes.— As has been observed 
with other members of the subgenus Hia- 
ticambarus (Hobbs, 1981:147), C. (4) elk- 
ensis is found under loose rocks in riffles, 
or pools that have currents. The species was 
sought for unsuccessfully in small, head- 
water streams. In these smaller tributaries 
C. (Cambarus) bartonii carinirostris Hay 
(1914) was abundant. We also sampled large 
rivers, especially below Sutton Lake, and 
failed to capture it. Cambarus (Puncticam- 
barus) robustus Girard (1852) was more 
common in these bodies of water. Since the 
species is found in the Birch River below 
Sutton Lake, we suspect that suitable hab- 
itat for this species was probably destroyed 
when Sutton Lake was constructed. 

Relationships.—Cambarus (H.) elkensis 


351 


Table 1.—Measurements (in mm) of types of Cam- 
barus (Hiaticambarus) elkensis, new species. 


Morpho- 
Character Holotype Allotype type 
Carapace 
Height i562. 12-5 14.2 
Width DAO P\ 16.1 18.7 
Length 40:0" ° 32:8 37.3 
Areola 
Length 15:0 - 12:0 [35 
Width 3.3 2.4 5-2 
Rostrum 
Length 3 ee 8.0 8.8 
Length to anterior 
postorbital ridges 7.4 6.5 7.6 
Width between eyes 4.5 eae) 4.6 


Postorbital ridge 
Width 9.0 fhe 8.3 
Chela (right) 
Length lateral margin 
Length mesial margin 
of palm 
Width of palm 
Length of dactyl 
Thickness of palm 


40.8 38 heal 
125 8.3 9.4 
1 Jed 14.0 
26.0 20) 5 
10.2 6.7 8.4 


Abdomen 
Length 38.6 34.0 38.2 
Width 172g 7139 16.1 
Gonopod 
Length 8.3 _ 7.8 
Antennal scale 
Length 6.6 5.4 62* 
Width 2.6 Did, LW ae 


* Left antennal scale. 


appears to be most closely related to C. (H.) 
chasmodactylus but differs from it by hav- 
ing the dactyl length/palm length ratio <2.3, 
less gaping fingers, and moderately strong 
impressions at the base of the fixed finger 
of the chela, especially on the ventral sur- 
face. 

Crayfish associates.—Collected with C. 
(H.) elkensis at one or more sites were Or- 
conectes (Crockerinus) sanbornii sanbornii 
(Faxon, 1884), O. (Procericambarus) spi- 


352 


nosus (Bundy, 1877), C. (C.) b. carinirostris, 
and C. (P.) robustus. 

Etymology. —We name this crayfish after 
the Elk River of West Virginia to which it 
appears to be endemic. 


Acknowledgments 


We thank all individuals who helped in 
the collection of specimens for this study, 
especially T. Jones. Special thanks are ex- 
tended to Dr. H. H. Hobbs, Jr. of the Smith- 
sonian Institution, Dr. J. F. Fitzpatrick, Jr., 
of the University of South Alabama, Dr. H. 
H. Hobbs, III of Wittenberg University and 
two anonymous reviewers for their com- 
ments concerning the manuscript. Funding 
for this project was provided, in part, by the 
West Virginia Division of Natural Re- 
sources, Non-game Program. Dave Dennis 
and Susan Hendrix of The Ohio State Un1- 
versity prepared the illustrations and funds 
for them were provided by The Ohio State 
University, Department of Zoology. 


Literature Cited 


Bundy, W. F. 1877. On the cambari of northern In- 
diana. — Proceedings of the Academy of Natural 
Sciences of Philadelphia 29:171-174. 

Faxon, W. 1884. Descriptions of new species of Cam- 
barus, to which is added a synonymical list of 
the known species of Cambarus and Astacus. — 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Proceedings of the American Academy of Arts 

and Sciences 20:107-158. 

1914. Notes on the crayfishes in the United 
States National Museum and the Museum of 
Comparative Zoology with descriptions of new 
species and subspecies to which is appended a 
catalogue of the known species and subspe- 
cies. — Memoirs of the Museum of Comparative 
Zoology at Harvard College 40(8):35 1-427. 
Girard, C. 1852. A revision of the North American 

astaci, with notes on their habits and geographic 
distribution. — Proceedings of the Academy of 
Natural Sciences of Philadelphia 6:87-91. 

Hay, W.P. 1914. Cambarus bartonii carinirostris Hay. 
In Walter Faxon, Notes on the crayfishes in the 
United States National Museum and Museum 
of Comparative Zoology.— Memoirs of the Mu- 
seum of Comparative Zoology at Harvard Col- 
lege 40(8):384—385. 

Hobbs, H. H., Jr. 1981. The crayfishes of Georgia. — 
Smithsonian Contributions to Zoology 318:viii 
+ 549, 

James, H. A. 1966. Range and variations of the sub- 
species of Cambarus longulus (Decapoda, As- 
tacidae). — Proceedings of the United States Na- 
tional Museum 119(3544):1024. 

Jezerinac, R. F., & G. Whitney Stocker. 1989. Dis- 
tribution of the stream crayfishes of the genus 
Cambarus (Decapoda: Cambaridae) in West 
Virginia.—Ohio Journal of Science 89(2):2-3. 


(RFJ) Department of Zoology, The Ohio 
State University at Newark, University 
Drive, Newark, Ohio 43055, U.S.A.; (GWS) 
13773 Bodle Road, Newark, Ohio 43055, 
U.S.A. 


PROC. BIOL. SOC. WASH. 
106(2), 1993, pp. 353-358 


TWO NEW SPECIES OF GLIRICOLA (PHTHIRAPTERA: 
GYROPIDAE) FROM THE SPINY TREE RAT, 
MESOMYS' HISPIDUS, IN PERU 


Roger D. Price and Robert M. Timm 


Abstract. — Two new species of Gliricola, G. woodmani and G. halli (Phthir- 
aptera: Gyropidae), are described and illustrated for specimens from the spiny 
tree rat, Mesomys hispidus (Rodentia: Echimyidae), in Peru. 

Resumen. —Se describe e ilustra dos nuevas especies de Gliricola, G. wood- 
mani y G. halli (Phthiraptera: Gyropidae) que fueron encontradas en una rata 
espinosa arborea, Mesomys hispidus (Rodentia: Echimyidae), en Peru. 


Thirty-three species of chewing lice of the 
genus Gliricola Mjoberg currently are rec- 
ognized, these being found on members of 
the rodent families Echimyidae, Caviidae, 
and Capromyidae. The majority, or 23, of 
these louse species occur on echimyids, with 
5 each on caviids and capromyids. In the 
most recent work on Gliricola, Emerson & 
Price (1975) describe five of these taxa as 
new, provide illustrations and brief reviews 
for seven described earlier, and give liter- 
ature citations that include all previously 
known taxa. 

We recently obtained two important se- 
ries of Gliricola from the spiny tree rat, Me- 
somys hispidus (Desmarest) (Rodentia: 
Echimyidae), in Peru. Our study of these 
specimens confirms that two species are 
present and their unique features indicate 
that the specimens we have represent two 
undescribed species. It is our purpose here 
to describe and illustrate these new species. 

The locality of capture for the host, Me- 
somys hispidus, is Reserva Cuzco Amazon- 
ico, 14 km E of Puerto Maldonado, Dept. 
Madre de Dios, in extreme southeastern 
Peru at an elevation of 200 m [12°33’S, 
69°03'’W]. Reserva Cuzco Amazonico is a 
national wildlife reserve located on the north 
bank of the Rio Madre de Dios, approxi- 
mately 300 km east-northeast of the city of 
Cuzco. For details of the habitat, climate, 


and history of the reserve see Duellman & 
Koechlin (1991). Woodman et al. (1991) 
provide an annotated listing of the mam- 
mals at this site. 


Gliricola woodmani, new species 
Figs. 1-5 


Type host.—Mesomys hispidus (Desma- 
rest). 

Female. —As in Fig. 1. Head longer than 
wide, with numerous short dorsal setae. 
Thorax as shown; mesonotum with row of 
22 short setae. Marginal abdominal tergal 
setae: I, 21; II-V, 25-26; VI-VII, 23; VIII, 
14; with markedly longer group of setae lat- 
erally on IV—VI. Anterior abdominal tergal 
setae: I, 2; II, 14; WI-V, 20-23; VI-VII, 17- 
18; VIII, 14. Last tergite with total of 8 an- 
terior setae and marginally each side with 
medium setae flanking pair of very long se- 
tae. Pleura II-VI each with 14-16 marginal 
and anterior setae, including medium to long 
seta at outer corner; pleuron VIII with 6-9 
setae, including single very long corner seta. 
Large spiracles on pleura III—VII. Marginal 
abdominal sternal setae: II, 6; HI—VII, 9- 
11. Anterior abdominal sternal setae: II, 5; 
II-VI, 6—10. Sterna II-III as in Fig. 3. Ster- 
num VIII with total of 10 setae. Ventral 
terminalia (Fig. 4) with posterior margin 
bearing 3+2 short setae on each side and 


354 


anteriorly with three prominent lobes, each 
of outer pair of lobes with two slender spat- 
ulate setae and inner lobe with medium seta 
laterad of shorter broader spatulate seta. 

Male. —As in Fig. 2. Much as for female, 
except as follows. Mesonotum with row of 
22-23 setae. Marginal abdominal tergal se- 
tae: I, 19-20; H-V, 23-29; VI-VII, 20-23; 
VIII, 13-14; all setae short and of essentially 
similar length. Anterior abdominal tergal 
setae: II, 16-18; III-V, 25-27; VI-VII, 21- 
30; VIII, 14-17. Last tergite with total of 
8-9 anterior setae and marginally each side 
with single very long seta flanked by several 
short setae. Pleura II—VII each with 12-17 
marginal and anterior setae, including me- 
dium seta at outer corner; pleuron VIII with 
9-11 setae, including very long corner seta. 
Marginal abdominal sternal setae; II, 6; IJ- 
VIII, 7-10. Anterior abdominal sternal se- 
tae: II-VII, 8-12; VIII, 12-14. Chaetotaxy 
of subgenital plate as shown. Genitalia (Fig. 
5) relatively simple, with straight parameres 
slightly swollen basally, endomeral plate 
posteriorly flattened to slightly rounded, 
transverse bridge near paramere base, slen- 
der tapered basal apodeme, and small light- 
ly spiculate sac. 

Dimensions (in mm).— Temple width, fe- 
male 0.23, male 0.21—0.22; prothorax width, 
female 0.20, male 0.19; metathorax width, 
female 0.28, male 0.26; abdominal width at 
V, female 0.43, male 0.31—0.33; head length, 
female 0.27, male 0.26; total length, female 
1.68, male 1.39-1.41; male genitalia width 
0.08, length 0.40-0.41, paramere length 
0.07-0.08. 

Type material. — Holotype male, ex Me- 
somys hispidus, Peru: Dept. Madre de Dios, 
14 km E Puerto Maldonado, Reserva Cuzco 
Amazonico, el. 200 m, 14 June 1989; in the 
Snow Entomological Collection, University 
of Kansas, Lawrence. Paratypes, 1 female, 
1 male, same data and depository as holo- 
type. 

Remarks.—This species is readily sepa- 
rated from all other species of Gliricola on 
the basis of its large dimensions, relatively 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


simple male genitalia, the unique arrange- 
ment of the spatulate setae on the female 
ventral terminalia, the longer lateral mar- 
ginal tergal setae on female abdominal seg- 
ments IV—VI, the single very long seta on 
pleuron VIII and the medium to long seta 
on each of pleura IJ-VII, the pair of very 
long setae on each side of the last female 
tergum and a single such seta on the male, 
and the large number of setae on sterna IJ— 
III. Although some other Gliricola may share 
features similar to some of the above, none 
has all of them. In fact, G. woodmani differs 
so significantly from all other congeneric 
species that it is difficult to select even a 
closely related species. 

Etymology.—This species is named in 
honor of Neal Woodman, University of 
Kansas, in recognition of his efforts in ob- 
taining these and other valuable specimens 
of ectoparasites and their hosts from the 
Neotropics, and his efforts in unraveling 
systematic problems in Neotropical mam- 
mals. 


Gliricola halli, new species 
Figs. 6-9 


Type host.—Mesomys hispidus (Desma- 
rest). 

Female.—Head and thorax near to G. 
woodmani. Abdomen as in Fig. 6. Meso- 
notum with row of 20-21 short setae. Mar- 
ginal abdominal tergal setae: I, 18-26; II- 
VI, 29-37; VII, 25-30; VIII, 21-22; with 
lateral setae only slightly longer than me- 
dian setae. Anterior abdominal tergal setae: 
II, 9-17; III-V, 16-24; VI-VII, 21-28; VIII, 
18-23. Last tergite with total of 6-14 an- 
terior setae and marginally each side with 
medium setae flanking pair of very long se- 
tae. Pleural marginal and anterior setae, in- 
cluding longer seta at outer corner: II, 11- 
13; II, 9-15; IV—VII, 11-14; VIII, 7-10 
setae, including 2 very long posterior setae. 
Large spiracles on pleura IIJ-VII. Marginal 
abdominal sternal setae: II, 0; HI, 6-8; IV- 
V, 9-11; VI-VII, 11-14. Anterior abdom- 


VOLUME 106, NUMBER 2 355 


Figs. 1-9. 1-5, Gliricola woodmani: (1) female; (2) male; (3) female sterna II-III; (4) female ventral terminalia; 
(5) male genitalia; 6-9, Gliricola halli: (6) female abdomen; (7) female sterna II-III; (8) male genitalia; (9) female 
ventral terminalia. 


356 


inal sternal setae: II, 4—5; III-IV, 1-8; V, 4— 
8; VI, 6-8; VII, 11-14. Sterna II-III as in 
Fig. 7. Sternum VIII with total of 10-17 
setae. Ventral terminalia (Fig. 9) with each 
side of posterior margin bearing two minute 
setae medially and five slender spatulate se- 
tae laterally, each side anteriorly with two 
medium setae laterad and two short setae 
mediad of lobe bearing single seta and broad 
spatulate seta. 

Male. —Much as for female, except in cer- 
tain quantitative chaetotaxy. Marginal ab- 
dominal tergal setae: I, 23; II-VI, 28-32; 
VII, 27; VII, 16. Anterior abdominal tergal 
setae: II, 6; HJ-IV, 16-17; V—VII, 20-23; 
VIII, 17. Last segment with five anterior 
setae. Genitalia (Fig. 8) much as for G. 
woodmani, but with evenly rounded en- 
domeral plate and shorter parameres. 

Dimensions (in mm).— Temple width, fe- 
male 0.20-—0.21, male 0.20; prothorax width, 
female 0.17—0.19, male distorted; metatho- 
rax width, female 0.21—0.27, male distort- 
ed; abdominal width at V, female 0.33-0.40, 
male distorted; head length, female 0.22- 
0.24, male 0.22; total length, female 1.35-— 
1.51, male 1.24; male genitalia width 0.08, 
length 0.38, paramere length 0.05-0.06. 

Type material. —Holotype female, ex 
Mesomys hispidus, Peru: Dept. Madre de 
Dios, 14 km E Puerto Maldonado, Reserva 
Cuzco Amazonico, el. 200 m, 14 June 1989; 
in the Snow Entomological Collection, Uni- 
versity of Kansas, Lawrence. Paratypes, 3 
females, 1 male, same data and depository 
as holotype. 

Remarks. —Even though G. halli occurs 
on the same host as G. woodmani, the for- 
mer is easily separated by its smaller di- 
mensions, the arrangement of the spatulate 
setae on the female ventral terminalia, the 
shorter lateral marginal tergal setae on fe- 
male abdominal segments IV—VI, the pair 
of very long setae on pleuron VIII, the small 
number of setae on sterna II-III, and the 
generally larger number of tergal setae and 
small number of pleural setae. 

A number of other Gliricola species have 
a similar type of setal configuration asso- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ciated with the female ventral terminalia, 
but they differ significantly in other aspects. 
Probably the closest species morphologi- 
cally to G. halliis G. humilis Werneck from 
Proechimys albispinus (1. Geoffroy); how- 
ever, the latter has smaller dimensions, a 
different length and shape of the male gen- 
italic parameres, a markedly truncate fe- 
male abdomen, as well as a different ab- 
dominal chaetotaxy. 

Etymology. —This species is named in 
honor of the late E. Raymond Hall, Uni- 
versity of Kansas, in recognition of his nu- 
merous contributions to mammalogy and 
conservation and his establishment of an 
outstanding research collection and library 
at the University of Kansas for the study of 
mammalogy. 


Discussion 


The Neotropical rodent family Echimyi- 
dae is the most speciose and ecologically 
diverse of the living caviomorph rodents. It 
includes some 15 genera and 70 to 100 spe- 
cies. The family was already diverse by the 
Oligocene (Reig 1986). The genus Meso- 
mys, the spiny tree rats, is one of the poorest 
known genera in the family. Four species of 
Mesomys currently are recognized; how- 
ever, these names are based on few speci- 
mens and the relationships of the known 
populations in the genus are in need of re- 
view (Emmons & Feer 1990, Nowak 1991). 
In a recent preliminary analysis of the re- 
lationships of several groups of echimyids, 
Patton (1989) found Mesomys [hispidus] to 
occupy a basal but somewhat ambiguous 
position with respect to the other lineages. 
However, Mesomys is clearly an old and 
distinct lineage within the Echimyidae. The 
most widespread species within the genus 
is Mesomys hispidus, which is found in the 
northern and western Amazon Basin, oc- 
curring in southern Colombia and Vene- 
zuela, eastern Ecuador and Peru, and west- 
erm Brazil. 

Although the occurrence of two conge- 
neric species of chewing lice on the same 


VOLUME 106, NUMBER 2 


host taxon is not typical, it is also not un- 
usual. Within the G/iricola, a number of host 
taxa share two or more species of lice. This 
may in part be due to confusion in the host 
taxonomy, but there are sufficient examples 
of co-occurrence that we are not suspicious 
of the material we are using as the basis of 
the description of our two new species. Ad- 
mittedly, longer series of the lice would be 
preferable, but the differences between the 
two are clearcut and there is no way they 
can be confused. 

The discovery of two congeneric species 
of chewing lice on a single host individual 
confirms our suspicions that species of G/ir- 
icola can be truly sympatric. How these two 
congeners are distributed on the host and 
precisely what they are feeding upon remain 
to be documented. 

To date, 42 species of chewing lice in 4 
genera (Gliricola, Gyropus Nitzsch, Harri- 
sonia Ferris, and Hoplomyophilus Méndez) 
have been described from 25 host species 
representing 8 genera of echimyids. Within 
these Gliricola, we find that each genus of 
host has its own fauna of lice and there are 
no shared species with any other host genus. 
Species of Gliricola have been described 
from Diplomys (1 species), Echimys (3 spe- 
cies), Kannabateomys (1 species), Euryzy- 
gomatomys (2 species), Hoplomys (1 spe- 
cies), Isothrix (1 species), and Proechimys 
(14 species). The discovery of these two new 
species (G. woodmani and G. halli) on Me- 
somys 1s consistent with this, as no lice have 
been described previously from this host 
genus. 

Given the diverse radiation of the echi- 
myids in South America and the paucity of 
lice available from these rodents, we strong- 
ly suspect that numerous new species of 
Gliricola have yet to be collected and much 
remains to be learned about the systematics 
and host relationships in this diverse genus. 


Acknowledgments 


Field work at Cuzco Amazonico by RMT 
was supported by National Geographic So- 


351. 


ciety Grant 4016-89 and the Museum of 
Natural History. Blgo. José Purisaca P. of 
the Direccion General Forestal y de Fauna, 
Ministerio de Agriculture, Lima, issued per- 
mits for our work. José E. Koechlin, of Cuz- 
co Amazonico Lodge, provided excellent fa- 
cilities and support for our studies there. 
Anthony B. Luscombe and the Asociacion 
de Ecologia y Conservacion (ECCO) were 
most instrumental in assisting with logis- 
tics, equipment, and supplies in Peru and 
contributed immeasurably to making our 
work successful. William E. Duellman and 
Philip S. Humphrey’s efforts in securing 
funds and aiding in logistics are most ap- 
preciated. Rosa Arana, Errol D. Hooper, 
Cecilia Pacheco, Victor Pacheco, Cheryl A. 
Schmidt, and Neal Woodman provided 
outstanding assistance with the fieldwork. 
We thank Gloria Arratia for translating our 
abstract into Spanish for the resumen, and 
Barbara L. Clauson for her constructive 
comments on this manuscript. Our study 
has been partially supported by Project No. 
Min-17-015 of the Minnesota Agricultural 
Experiment Station and has been assigned 
paper no. 20,157 of the Scientific Journal 
Series. 


Literature Cited 


Duellman, W. E., & J. E. Koechlin. 1991. The Re- 
serva Cuzco Amazonico, Peru: Biological in- 
vestigations, conservation, and ecotourism.— 
Occasional Papers of the Museum of Natural 
History, The University of Kansas, 142:1-38. 

Emerson, K. C., & R. D. Price. 1975. Mallophaga of 
Venezuelan mammals.—Bngham Young Uni- 
versity Science Bulletin, Biological Series, 20, 
No. 3:1-77. 

Emmons, L. H., & F. Feer. 1990. Neotropical rain- 
forest mammals: a field guide. University of 
Chicago Press, Chicago, 281 pp. 

Nowak, R.M. 1991. Walker’s mammals of the world. 
Vol. II. 5th edition. Johns Hopkins University 
Press, Baltimore, pp. 643-1629. 

Patton, J. L., & O. A. Reig. 1989. Genetic differen- 
tiation among echimyid rodents, with emphasis 
on spiny rats, genus Proechimys. Pp. 75-96 in 
K. H. Redford and J. F. Eisenberg, eds., Ad- 
vances in neotropical mammalogy. Sandhill 
Crane Press, Inc., Gainesville. 

Reig, O. A. 1986. Distribution patterns and differ- 


358 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


entiation of high Andean rodents. Pp. 404-439 
in F. Vuilleumier and M. Monasterio, eds., High 
altitude tropical biogeography. Oxford Univer- 
sity Press, Oxford. 

Woodman, N. etal. 1991. Annotated checklist of the 
mammals of Cuzco Amazonico, Peru.—Occa- 
sional Papers of the Museum of Natural History, 
The University of Kansas, 145:1-12. 


(RDP) Department of Entomology, Uni- 


versity of Minnesota, St. Paul, Minnesota 
55108, U.S.A. (Current address) 4622 Kin- 
kead Ave., Fort Smith, Arkansas 72903, 
U.S.A.; (RMT) Museum of Natural History 
and Department of Systematics and Ecol- 
ogy, University of Kansas, Lawrence, Kan- 
sas 66045-2454, U.S.A. (direct reprint re- 
quests to RMT). 


PROC. BIOL. SOC. WASH. 
106(2), 1993, pp. 359-365 


SPADELLA JAPONICA, A NEW COASTAL 
BENTHIC CHAETOGNATH FROM JAPAN 


Jean-Paul Casanova 


Abstract. — Spadella japonica, a new benthic chaetognath, is described from 
the coastal waters of Japan. It differs from all known species of the genus 
Spadella, mainly by the opening of the female genital orifices at the bottom of 
a cupel and the small number of hooks. The distribution of the three Spadella 
species living in Japanese waters is presented. 


When Dr. Taichiro Goto (Mie Univer- 
sity, Japan) gave me specimens of a new 
Paraspadella during the first workshop on 
chaetognaths (University of Surrey, Sep- 
tember 1988), P. gotoi Casanova, 1990, there 
were also two individuals of a Spadella that 
he thought to be new. They were collected 
in Misaki on 6 August 1987. In November 
1991, I received 16 additional individuals 
caught in Kominato on 12 September 1991 
which, with the other two, differ by many 
characters from the species of Spadella hith- 
erto described. 


Spadella japonica, new species 
Figs. 1-3, Table 1 


The holotype and two paratypes are de- 
posited with the National Science Museum 
Tokyo (NSMT—Ch. 20 and 21-22 respec- 
tively). Three other paratypes are deposited 
in the Muséum national d’Histoire Natu- 
relle, Paris (UC 366) and three other ones 
in the National Museum of Natural History, 
Washington, D.C. (USNM 157572). All are 
from Kominato. 

Description. —Eighteen specimens stud- 
ied. Body stumpy when adult (Fig. 1) and 
transparent. Length up to 3.75 mm not in- 
cluding tail fin. Tail constitutes 48.6 to 
51.8% of total length. 

Number of hooks increasing with age, 
from six (small specimen: 3.25 mm) to eight 
(larger ones: 3.60-—3.75 mm). Anterior teeth 
three or four, short (Fig. 2a, b). No posterior 


teeth. Pigment cell of eyes more enlarged in 
the second set of specimens received (Fig. 
3a). Corona ciliata on neck, oval and trans- 
versely elongated (Figs. 1, 3b). Collarette 
very wide at level of posterior part of head 
and neck, then narrower on trunk (Figs. 1, 
3b). Sensory tufts symmetrically arranged 
on whole body (Fig. 1). Numerous adhesive 
papillae on ventral part of body (Fig. 3c, d), 
from head to tail, and on both ventral and 
dorsal sides of fins. Gut with small intestinal 
diverticula at level of neck, not always vis- 
ible on preserved specimens. Transverse 
musculature thin, stretching from neck al- 
most to transverse septum. Ventral ganglion 
about middle third of trunk. 

Lateral fins beginning on posterior part 
of trunk (about 15-19% of the trunk length) 
and reaching posteriorly to seminal vesicles 
(Fig. 1). Tail fin spatulate. Rayless zone 
wholly absent on all fins. Left lateral fin of 
a small specimen bearing dorsally a small 
area with tiny papillae (Fig. 3e) such as those 
found on adhesive organs of Paraspadella 
gotoi. Ovaries reach to about midlength of 
ventral ganglion; their aperture lateral, at 
bottom of brown colored elongate cupel (Fig. 
3f, g), of which largest diameter (0.20—0.25 
mm) is the same as that of mature ova. 
These cupels are the sole colored parts of 
the body. Ovaries with from one to five ma- 
ture ova and other smaller ones. Seminal 
vesicles small, hook-shaped when empty, in 
close contact with both lateral and tail fins, 


CC 
VG 
€ 
7 
U 
Y 


cies. CC = corona ciliata, SV = seminal vesicles, VG 
= ventral ganglion. 


opening at posterolateral extremity (Fig. 3h, 
i). 

Remarks.—Two specimens of Spadella 
Japonica are interesting. One has a curious 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


‘““papillated organ” arising from the epi- 
dermis on the ventral right side of the tail, 
not far from the transverse septum. Itis 0.15 
mm long and constructed of a short stalk 
ending in a swelling provided with papillae 
(Fig. 3c, d). A second one shows two of these 
“‘organs’’ budding on the head and neck: the 
papillated swellings are visible but there are 
no stalks (Fig. 3), k). 

These observations are pertinent to the 
taxonomic position of Spadella moretonen- 
sis Johnston & Taylor, 1919. Indeed, since 
the revision of the benthic family Spadel- 
lidae Tokioka, 1965 by Bowman & Bieri 
(1989), the genus Spadella is restricted to 
species lacking adhesive organs, those pro- 
vided with such organs constituting the ge- 
nus Paraspadella. It has been demonstrated 
that these organs are modified parts of fins 
(Casanova 1990). Spadella moretonensis was 
described from East Australia from a single 
specimen having two club-shaped papillat- 
ed bodies on the posterior half of the tail, 
situated ventrally on the right side. But the 
authors added: “‘Though they became 
stained like the tissues of the animal [when 
using haematoxylin] yet their asymmetrical 
arrangement and general appearance sug- 
gest that they are foreign bodies— perhaps 
of an algal nature.’ Later on, Tokioka & 
Pathansali (1964) wrote: “Species of schi- 
zoptera-group [=Paraspadella| are easily 
distinguishable by their characteristic ad- 
herent organ, while S. moretonensis ... 1s 
devoid of any complete palm-shaped ad- 
herent organ ... It is possible this might 
rather be a form of schizoptera-group bear- 
ing no complete adhesive organs vut rudi- 
mentary ones.”’ Alvarino (1981) agreed with 
this view. According to Salvini-Plawen 
(1986), who first proposed splitting the large 
genus Spadella into three smaller ones, some 
uncertainty concerns this species with re- 
spect to the asymmetry of these bodies. 
Nevertheless he placed it in the genus Ge- 
phyrospadella (now included in Paraspa- 
della). Lastly, because of this uncertainty, 
this species was not compared with other 


VOLUME 106, NUMBER 2 


a Tt « 


2d ge Nee 
ae ee 


Fig. 2. SEM photographs of Spadella japonica, new 


teeth (< 500). 


ones in recent papers, neither with Paraspa- 
della nor with Spadella (Casanova 1990, 
1991). 

It appears that the bodies described in S. 
moretonensis are the same as those found 
in S. japonica. Because in the latter they are 
present in only two specimens and on the 
head and neck of one of them, they cannot 
be considered rudimentary adhesive organs. 
Thus S. moretonensis is not a Paraspadella 
but a Spadella. According to Goto’s obser- 
vations (pers. comm.), during culture the 
epidermis of S. japonica sometimes.changes 
in appearance, probably as a result of food 
or age. Especially when fed with Artemia 
nauplii, which are not very good for Spa- 
della although they eat them, the epidermis 
becomes thin and seems to be deformed. 
Similarly, the papillae-like structures oc- 
curring in a few specimens living in the sea 
might be the result of bad environmental 
conditions. 

Comparisons with other species. —The 
main differential characteristics of the eight 
species of Spadella known before the pres- 
ent study have been given recently (Casa- 
nova 1991). Four of them always have pos- 


361 


species: a, Ventral part of head (x 180); b, Details of 


terior teeth: the cave species Spadella 
ledoyeri Casanova, 1986, the two deep spe- 
cies Spadella birostrata Casanova, 1987 and 
S. equidentata Casanova, 1987, and S. ant- 
arctica Casanova, 1991. The other four, as 
well as S. moretonensis, may or may not 
have posterior teeth. In addition, as they 
live in neritic temperate or tropical waters, 
comparisons will be made only with them. 

The unusual structure of the area sur- 
rounding the female genital opening easily 
separates Spadella japonica from all other 
species of the genus. In more particular re- 
gard to those being compared: S. cephal- 
optera Busch, 1851 has a prominent cement 
gland close to each opening; S. angulata To- 
kioka, 1951 and S. gaetanoi Alvarino, 1978 
are devoid of this gland; the opening is not 
described in S. bradshawi Bieri, 1974 and 
thus there is probably a simple orifice as in 
the two latter species; as for S. moretonen- 
sis, the aperture is trilobed and situated on 
a well-marked prominence. Furthermore, 
all these species have more numerous hooks 
than S. japonica (Table 1). 

Other main features show the specificity 
of the new Spadella by comparison with 


Fig. 3. Spadella japonica, new species: a, Right eye; b, Anterior part of body in dorsal view showing the 
corona ciliata (arrow); c, Anterior part of the tail of an original specimen in ventral view, showing a curious 
““papillated organ” (arrow) and adhesive papillae; d, Enlargement of the “‘papillated organ” area; e, Area with 
tiny papillae on the ventral side of the left lateral fin (other original specimen); f and g, Two aspects of the female 
genital opening (arrow) in dorsal view; h, Empty seminal vesicles; 1, Mature seminal vesicles; j, Head and neck 
of another original specimen with two “‘papillated organs” (arrows) in dorsal view; k, Enlargement of the organ 
on neck (arrow), just under a sensory spot. Scale bars: 0.1 mm (b, c, e-j), 0.05 mm (d) and 0.02 mm (a, k). 


VOLUME 106, NUMBER 2 


1. Tomioka 

2. Aitsu 

3. Mitsu 

4. Yashima 

5. Setozaki 

6. Misaki 

7. Kominato 

8. Otsuchi 

9. Nanao 
10. Yoron I. 


% Spadella angulata 


1364 
ale 


pees Se 
@ Spadella cephaloptera 


A Spadella japonica n. sp. 


363 


40°N 


36° 


Freon 
6 


° (“4 
22 % 10 


Okinawa 
140° 129°E 


Fig. 4. Distribution of the three species of Spade/la known from neritic waters of Japan. 


each of the five species mentioned above. 
Indeed, it differs from: 


— Spadella cephaloptera, in which lateral fins 
begin at level of the transverse septum 
(Japanese specimens) or slightly before 
(some Mediterranean specimens) and 
seminal vesicles are more or less rounded; 

— Spadella angulata, which has a special 
orange-brown color pattern on the ventral 
and dorsal sides of the body and on the 
digestive tract (Tokioka & Bieri 1966, 
Bieri et al. 1987); according to Goto and 
Thuesen (pers. comm.) who observed the 
two species in the Japanese waters (spec- 
imens alive as well as preserved ones), S. 
angulata is opaque (creamy white) while 
S. japonica is transparent; 

— Spadella bradshawi, in which the corona 
ciliata is massive and lateral fins do not 
touch the seminal vesicles; 


— Spadella gaetanoi, which bears a pair of 
cup-like structures between tip of tail and 
seminal vesicles, and moreover, the latter 
are reniform; 

— Spadella moretonensis, in which the ven- 
tral ganglion is very short (less than 25% 
of the trunk length). 


The characteristics of the six species of 
Spadella which are always or sometimes de- 
void of posterior teeth are summarized in 
Table 1. 

Distribution. —Three species of Spadella 
live in the neritic waters of Japan (Fig. 4). 
Spadella cephaloptera has been found in 
Misaki, Mitsu and Yashima. According to 
Yosil & Tokioka (1939), the two minute 
tentacles on the head are found in small 
specimens but not on large ones, whereas 
in European specimens they also exist on 
adults. Spadella angulata has been men- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


364 


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VOLUME 106, NUMBER 2 


tioned in many localities: Otsuchi, Nanao, 
Setozaki, Tomioka, Aitsu and near Okina- 
wa (Yoron Island, southwest of Japan). Jap- 
anese specimens, first described by Tokioka 
(1951) as S. cephaloptera forma angulata, 
differ from Malay specimens (Tokioka & 
Pathansali 1964) in having, usually, one 
posterior tooth. Spadella japonica, accord- 
ing to Goto (pers. comm.), has been col- 
lected among the Zostera belt in Misaki and 
in tide pools of Kominato and Tomioka and 
is more common than S. angulata. 


Acknowledgments 


I thank very sincerely Dr. Taichiro Goto 
of Mie University (Japan) who gave me the 
specimens of the second new benthic species 
he sampled in Japan. 


Literature Cited 


Alvarino, A. 1978. Spadella gaetanoi, a new benthic 
chaetognath from Hawaii.— Proceedings of the 
Biological Society of Washington 91:650-657. 
1981. Spadella legazpichessi, a new benthic 
chaetognath from Enewetak, Marshall Is- 
lands. — Proceedings of the Biological Society of 

Washington 94:107-121. 

Bieri, R. 1974. A new species of Spadella (Chaetog- 
natha) from California. — Publications of the Seto 
Marine Biological Laboratory 21:281-286. 

—, M. Terazaki, E. V. Thuesen, & T. Nemoto. 
1987. Colour pattern of Spadella angulata 
(Chaetognatha: Spadellidae) with a note on its 
northern range extension. — Bulletin of Plankton 
Society of Japan 34(1):83-84. 

Bowman, T.E.,&R. Bieri. 1989. Paraspadella anops, 
new species, from Sagittarius cave, Grand Ba- 
hama Island, the second troglobitic chaeto- 
gnath.— Proceedings of the Biological Society of 
Washington 102:586—-589. 

Busch, W. 1851. Beobachtungen tiber Anatomie und 
Entwickelung einiger wirbellosen Seethiere. 
Chaetognatha.— Berlin 4:93-100. 


365 


Casanova, J.-P. 1986. Spadella ledoyeri, chaeto- 
gnathe nouveau de la grotte sous-marine ob- 
scure des Trémies (calanques de Cassis). — Rap- 
ports de la Commission internationale pour 
Exploration Scientifique de la mer Méditer- 

ranée 30(2):196. 

1987. Deux chaetognathes benthiques nou- 
veaux du genre Spadella des parages de Gibral- 
tar. Remarques phylogénétiques.— Bulletin du 
Muséum national d’Histoire naturelle, Paris, 4é 
sér., 9, section A(2):375-390. 

1990. A new species of Paraspadella (Chae- 
tognatha) from the coastal waters of Japan.— 
Proceedings of the Biological Society of Wash- 
ington 103:907-912. 

. 1991. The first record ofa benthic polar chae- 

tognath: a new Spadella from the Antarctic. — 

Journal of Natural History 25:1355-1362. 

Johnston, T. H., & B. B. Taylor. 1919. Notes on 
Australian chaetognatha.— Proceedings of the 
Royal Society of Queensland 31:28—41. 

Salvini-Plawen, L. Von. 1986. Systematic notes on 
Spadella and on the chaetognatha in general. — 
Zeitschrift fir Zoologische Systematik und Evo- 
lutionsforschung 24(2):122-128. 

Tokioka, T. 1951. Pelagic tunicates and chaetognaths 

collected during the cruises to the New Yamato 

Bank in the Sea of Japan.— Publications of the 

Seto Marine Biological Laboratory 2:1-25. 

1965. The taxonomical outline of Chaetog- 
natha.— Publications of the Seto Marine Biolog- 
ical Laboratory 12:335-357. 

— ., & D. Pathansali. 1964. Spadella cephaloptera 
forma angulata raised to the rank of species. — 
Publications of the Seto Marine Biological Lab- 
oratory 12:145-148. 

——, & R. Bieri. 1966. The colour pattern of Spa- 
della angulata Tokioka.—Publications of the 
Seto Marine Biological Laboratory 14:323-326. 

Yosii, N., & T. Tokioka. 1939. Notes on Japanese 
Spadella (Chaetognatha).—Annotationes Zoo- 
logicae Japonenses 18:267-273. 


Laboratoire de Biologie animale (Planc- 
ton), Université de Provence, 13331 Mar- 
seille Cedex 3, France. 


PROC. BIOL. SOC. WASH. 
106(2), 1993, pp. 366-368 


CAUDINA INTERMEDIA, A NEW SPECIES OF SEA 
CUCUMBER FROM THE SOUTH CHINA SEA 
(ECHINODERMATA: HOLOTHUROIDEA: MOLPADIIDA) 


Yulin Liao and David L. Pawson 


Abstract.—Caudina intermedia, new species is described. The bodywall os- 
sicles of this species are exclusively tables; knobbed buttons or plates are absent. 
The tables are typical of the genus Caudina Stimpson, 1853 in the strict sense, 
but the absence of other ossicle types suggests some affinities with the genus 
Hedingia Deichmann, 1938. A revised key to the known species of Caudina 


is presented. 


Subsequent to the publication of our re- 
cent paper on the molpadiid sea cucumbers 
of China (Pawson & Liao 1992), an addi- 
tional new molpadiid species was found in 
the collections of the Institute of Oceanol- 
ogy, Academia Sinica, Qingdao (IOAS). 


Family Caudinidae Heding, 1931 
Caudina Stimpson, 1853 


For a summary of the caudinids of China, 
see Pawson & Liao (1992). 


Caudina intermedia, new species 
Fig. 1A-—G 


Material examined. —Holotype, IOAS 
E1056. off eastern Guangdong, 21°45'N, 
115°30’E, 9 Jan 1960, 107 m, muddy sand 
bottom. Paratype, IOAS E1057, same lo- 
cality as Holotype. 

Diagnosis. —Ossicles of body wall exclu- 
sively tables with solid spires derived from 
four pillars fused together, terminating in a 
few teeth. Knobbed buttons, perforated 
plates, and phosphatic deposits absent. 

Description. — Body more or less barrel- 
shaped, with conspicuous narrow tail. Ho- 
lotype 19 mm long, 10 mm in diameter, tail 
7 mm long; Paratype approximately the 
same size. Body wall thin, translucent. Anus 
surrounded by five minute anal teeth. Color 
in alcohol dirty gray to whitish. Radial piec- 


es of calcareous ring with short bifid pos- 
terior projections; interradial pieces slightly 


_ wider than radials (Fig, 1G). Stone canal 


and polian vesicle single. 

Body wall ossicles exclusively tables, dif- 
fering slightly in various parts of the body. 
In anterior body wall, disc more or less cir- 
cular or irregular in outline, 130-180 um in 
diameter (X 148 wm, SD 9.3), with four large 
central perforations and 5-10 peripheral 
ones. Spire tall, average height 150 um, sol- 
id, derived from four converging and fused 
pillars, terminating in three or four spines 
(Fig. 1A, B). In median and posterior body 
wall ossicles, disc more or less square in 
outline, average diameter 150 wm, com- 
monly with four large perforations; spire 
solid, average height 100 um, ending in three 
blunt teeth (Fig. 1C, D). Tables in tail (Fig. 
1E, F) smaller, with numerous perforations 
and small knobs on periphery and elsewhere 
on disc. Spire low, solid; in a few cases a 
single crossbar present. 

Remarks.—This new species is distinct 
from other Caudina species in lacking ac- 
cessory ossicles in the form of buttons or 
plates (see key below). In possessing only 
tables in the body wall, C. intermedia ap- 
proaches Hedingia Deichmann, 1938, but 
the tables of Hedingia are usually consid- 
erably larger (200-300 um), have more per- 
forations, and typically have two-pillared or 


VOLUME 106, NUMBER 2 367 


Fig. 1. Caudina intermedia, new species. A, tables from anterior body wall; B, same in profile view; C, tables 
from median and posterior body wall; D, same, in profile view; E, tables from tail; F, same, in profile view; G, 
radial and interradial pieces of calcareous ring. The scale measures 200 wm for A—D, 100 um for E-F, and 1.2 
mm for G. 


368 


(more commonly) three-pillared spires. Both 
specimens of C. intermedia are only 19 mm 
long. It is conceivable that buttons or plates 
may develop with further growth, but not 
likely, for in other species of Caudina the 
full complement of ossicle types is reached 
at an early age and small size. 


Key to the Known Species of Caudina 
(modified after Pawson & Liao 1992) 


1. Tables inconspicuous, scarce, re- 

duced to rods or plates, or lacking 
meee ae 8s tec arenicola (Stimpson, 1853) 

— Tables conspicuous, numerous, with 


well-developed spire and disk .... 2 
2. Tables accompanied by knobbed 
buttons or perforated plates ...... 5 


— Only tables present; buttons and 
plates absent .. intermedia, new species 
3. Tables accompanied by knobbed 
buttons 
— Tables accompanied by perforated 
plates“ MOlIDULIONS op ue eae 5 
4. Buttons not abundant, weakly 
knobbed, usually oval in outline, 
with two large elliptical and two 
siitall circulanihOres,. 33. she =. &. 
Eh Tes TR Cah A arenata Gould, 1841 
— Buttons abundant, strongly 
knobbed, usually with irregular out- 
line, the four holes more or less alike 
im Size and fort &.24 238 
nO Ot: SARE similis (Augustin, 1908) 
5. Discs of tables 180-280 um in di- 
ameter; perforated plates very vari- 
able in form and in number of holes, 
lacking knobsstais.s Wh oe WAL 
atacta Pawson & Liao 1992 


oe a a a ee 8) a 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


— Discs of tables 150-180 ym in di- 
ameter; perforated plates vary only 
slightly in form and in number of 
holes, often possessing a few low 
KHODS 0) 5 oes «de eee 

. Zhejiangensis Pawson & Liao 1992 


Literature Cited 


Augustin, E. 1908. Uber Japanische Seewalzen.— 
Abhandlungen der K6niglichen Bayerischen 
Akademie der Wissenschaften. II. Supplement 
i: 1-44. 

Deichmann, E. 1938. New holothurians from the 
western coast of North America and some re- 
marks on the genus Caudina.—Proceedings of 
the New England Zoological Club 16:103-115. 

Gould, A. A. 1841. Report on the Invertebrata of 
Massachusetts. Cambridge, Massachusetts, xiii 
+S 2appe 

Heding, S.G. 1931. On the classification of the mol- 
padids. — Videnskabelige Meddelelser fra Dansk 
naturhistorisk Forening i Kjobenhavn 92:275- 
284. 

Pawson, D. L. 1977. Molpadiid sea cucumbers (Echi- 

nodermata: Holothuroidea) of the Southern At- 

lantic, Pacific and Indian Oceans.— Biology of 
the Antarctic Seas VI. Antarctic Research Series 

26:97-123. 

, & Y. Liao. 1992. Molpadiid sea cucumbers 

of China, with description of five new species 

(Echinodermata: Holothuroidea).—Proceed- 

ings of the Biological Society of Washington 105: 

373-388. 

Stimpson, W. 1853. Synopsis of the marine inver- 
tebrates of the Grand Manan.—Smithsonian 
Contributions to Knowledge 5:6-17. 


(YL) Institute of Oceanology, Academia 
Sinica, 7 Nan-Hai Road, Tsingtao, People’s 
Republic of China; (DLP) Department of 
Invertebrate Zoology, National Museum of 
Natural History, Smithsonian Institution, 
Washington, D.C. 20560, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(2), 1993, pp. 369-384 


HYPORHAMPHUS MEEKT, A NEW SPECIES OF 
HALFBEAK (TELEOSTEI: HEMIRAMPHIDAE) FROM 
THE ATLANTIC AND GULF COASTS OF THE 
UNITED STATES 


Heidi M. Banford and Bruce B. Collette 


Abstract.—Hyporhamphus meeki is described from the Atlantic and Gulf of 
Mexico coasts of the United States. It has been confused previously with H. 
unifasciatus (Ranzani) which occurs from southern Florida, Bermuda, the West 
Indies, and Mexico south to southern Brazil. Hyporhamphus meeki has more 
gill rakers, usually 33-39 on the first gill arch and 26-29 on the second arch 
compared to 28-32 on the first arch and 19-25 on the second arch in H. 
unifasciatus. Pectoral-fin rays are usually 11 or 12 vs. 10 or 11 in H. unifasciatus. 
The ratio of preorbital length to orbit diameter is usually greater than 0.70 in 
H. meeki, less than 0.70 in H. unifasciatus. 


There has been considerable confusion 
regarding the taxonomy and systematics of 
the New World halfbeaks (Meek & Goss 
1884, Miller 1945). Although the status of 
some species has been clarified, that of Hy- 
porhamphus unifasciatus (Ranzani, 1842), 
the common inshore halfbeak, has been 
questioned for more than 100 years (Meek 
& Goss 1884, Collette 1978) but not re- 
solved. The name Hyporhamphus unifas- 
ciatus (Ranzani) has been used for inshore 
halfbeaks in the western Atlantic, eastern 
Atlantic, in several parts of the Indo-West 
Pacific, and the eastern Pacific. The eastern 
Atlantic Hyporhamphus were shown to be 
H. picarti (Valenciennes, 1846) by Collette 
(1965); the Indo-West Pacific halfbeaks H. 
limbatus (Valenciennes, 1846) by Parin et 
al. (1980). This paper presents morpho- 
metric and meristic analysis of western At- 
lantic populations of halfbeaks referred to 
H. unifasciatus, with the description of a 
new species. Our objective is to describe this 
new species of Hyporhamphus, and com- 
pare it with the true H. unifasciatus. The 
new species will be included in further study 
of New World halfbeaks and other publi- 


cations, particularly the halfbeak section for 
‘Fishes of the western North Atlantic.” 

Populations referred to H. unifasciatus 
range in the western Atlantic from Uruguay 
in the south, northward along the coast of 
the Americas, through the Caribbean (Jor- 
dan & Evermann 1896), Gulf of Mexico 
(Hoese & Moore 1977), and around Ber- 
muda (Beebe & Tee-Van 1933) to Cape Cod, 
Massachusetts. Strays have been collected 
as far north as Chamcook, Passamaquoddy 
Bay, New Brunswick (Leim & Day 1959). 
In the eastern Pacific they range from Peru 
(Hildebrand 1946) to Baja California and 
around the Galapagos Islands (Meek & Hil- 
debrand 1923). The range of H. unifasciatus 
s. S. (type locality, Brazil) is from Bermuda 
and peninsular Florida southward through 
the Caribbean to Uruguay. Those popula- 
tions referred to as H. unifasciatus from out- 
side this range constitute superficially sim- 
ilar undescribed species of Hyporhamphus 
(Collette 1978). This paper deals only with 
the western Atlantic populations of the H. 
unifasciatus species group. The eastern Pa- 
cific populations will be addressed in future 
publications. 


370 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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Fig. 1. Distribution of Hyporhamphus meeki, n. sp. (closed circles) and H. unifasciatus (open circles) based 


on material examined. 


Materials and methods.—A total of 568 
specimens of western Atlantic Hyporham- 
phus was examined for 24 morphometric 
and meristic characters; an additional 1088 


specimens were examined only for meristic 
characters. The number of specimens ex- 
amined for each character varies due to the 
condition of material. Due to the poor con- 


VOLUME 106, NUMBER 2 


dition of some specimens only some char- 
acters could be observed reliably. Material 
was chosen to represent the entire geograph- 
ical range of what has been considered H. 
unifasciatus in the western Atlantic (Fig. 1). 
The majority of material examined was ob- 
tained from the following institutions (ab- 
breviations from Leviton et al. 1985): 
AMNH, ANSP, CBL (Chesapeake Biolog- 
ical Laboratory, specimens now at VIMS), 
CAS, MCZ, MZUSP, SIO, UF, VIMS, 
UMMZ, and USNM. Additional material 
was collected by the first author in the York 
River at Gloucester Pt., Virginia, and is 
housed at VIMS. Following the description 
of the new species only the material that 
was examined for both morphometrics and 
meristics is listed. 

Most characters examined follow Collette 
(1965) and Parin et al. (1980). Measure- 
ments were made to the nearest tenth of a 
millimeter (mm). Abbreviations and de- 
scriptions of characters examined are as fol- 
lows: SL (standard length); LJL (lower jaw 
length, tip of upper jaw to tip of lower jaw); 
HDL (head length, from tip of upper jaw to 
posterior margin of opercle membrane); UJL 
(upper jaw length, from tip of upper jaw to 
where upper jaw bends); UJW (upper jaw 
width, where upper jaw bends); P,-P> (dis- 
tance from base of upper pectoral ray to base 
of anterior pelvic ray); P,-C (distance from 
base of anterior pelvic ray to caudal base); 
P,-CX (P,-C distance extended anteriorly 
from base of anterior pelvic ray to a point 
on the body or head); BD-P, (body depth 
at origin of pectoral fin); BD-P, (body depth 
at origin of pelvic fin); ABASE (length of 
anal-fin base); DBASE (length of dorsal-fin 
base); P, L (pectoral-fin length, distance from 
base of uppermost pectoral ray to tip of lon- 
gest ray); ORB (soft orbit length); PREORB 
(preorbital length, from corner of mouth to 
anterior margin of orbit); ANA (number of 
anal-fin rays); DOR (number of dorsal-fin 
rays); P,; (L, R, number of pectoral-fin rays); 
PRED (number of predorsal scales in me- 
dian row in front of dorsal fin); RGR, (num- 


a7 4 


ber of gill rakers on first arch (upper + lower 
= total); RGR, (number of gill rakers on 
second arch (upper + lower = total). VERT 
(number of precaudal plus caudal vertebrae, 
including the hypural plate = total number 
of vertebrae). 

Statistical analyses utilized SAS software 
(SAS Institute, Inc. 1985). Frequency dis- 
tributions of counts were compared be- 
tween geographic populations and are pre- 
sented in summary tables. If two populations 
in close geographic proximity did not have 
significantly different counts, counts were 
combined to form a single population in 
subsequent statistical analysis. Infraspecific 
geographic variation is discussed herein. 
Analysis of Variance (ANOVA) was per- 
formed on five data sets of meristic char- 
acters. If the F value for an ANOVA was 
significant (P < .05), Tukey’s Studentized 
Range Test (Tukey-Kramer method) (SAS 
Institute, Inc. 1985) was performed to de- 
termine which means were significantly dif- 
ferent from the others. 

Values of morphometric characters were 
first plotted against SL, and then plotted 
against one another to inspect visually for 
separation between populations. Only those 
plots that proved to be diagnostic for the 
species are presented. Regression equations 
were generated for the plots of morpho- 
metric characters. Residual plots were in- 
spected for homogeneity of variance. Due 
to heteroscedasticity of variance, all mor- 
phometric data were log transformed for re- 
gression analysis. 


Hyporhamphus meeki, new species 
Fig. 2A 


Synonymy. —At least 10 different com- 
binations of names have been used for this 
species. A complete synonymy will be in- 
cluded in the halfbeak section of ‘“‘Fishes of 
the western North Atlantic.’’ The common- 
est names used have been Hyporhamphus 
(or Hemiramphus) roberti (about 30 refer- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


372 


(Js WU p91) 9996 WNSN ‘euteueg ‘uooD ‘snzp0snfiun “FT: (TS WU 091) 69€h6T WNSN ‘edA10]0y ‘satoads mou ‘ryaau snydupmysoddy WY *7 “Bt4 


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Reta 


VOLUME 106, NUMBER 2 


10 


PREORBITAL LENGTH (mm) 


0 ) 


8738 


10 15 


ORBIT DIAMETER (mm) 


Fig, 3. 
(triangles). 


ences, mostly 1862 to 1892) or unifasciatus 
(about 95 references, 1870 up to the pres- 
ent). Bruce (1986) prematurely used the 
name H. meeki as a nomen nudum in his 
treatment of isopod parasites of the genus 
Mothocya. 

Diagnosis.—A member of the subgenus 
Hyporhamphus distinguished from H. rob- 
erti (Valenciennes) by having the dorsal and 
anal fins covered with scales. This species 
is distinguished by the following combina- 
tion of characters: gillrakers on the first arch 
31-40 (Table 1); gill rakers on the second 
arch 20-30 (Table 2); pectoral-fin rays 10- 
13 (Table 5); ratio of preorbital length to 
orbit diameter is usually greater than 0.70 
(>0.70 in 92% of 265 specimens examined; 
Fig. 3). 

Description. —Gill rakers on upper limb 


Relationship of preorbital length to orbit length in Hyporhamphus meeki (squares) and H. unifasciatus 


of first arch 8 to 12, usually 9 to 11, mean 
9.8; lower limb 20 to 29, usually 24 to 27, 
mean 25.1; total of upper and lower limbs 
31 to 40, usually 33 to 37, mean 34.6 (Table 
1). Gill rakers on upper limb of second arch 
2 to 6, usually 4 or 5, mean 4.3; lower limb 
20 to 26, usually 22 or 23, mean 22.3; total 
20 to 30, usually 25 to 28, mean 26.2 (Table 
2). Dorsal-fin rays 12 to 17, usually 14 or 
15, mean 14.5 (Table 3); anal-fin rays 14 to 
18, usually 15 to 17, mean 15.9 (Table 4); 
and pectoral-fin rays 10 to 13, usually 11 or 
12, mean 11.4 (Table 5). Predorsal scales 
(n = 87) 34 to 39, usually 35 to 37, mean 
36.1. Vertebrae (n = 88) 31-35 precaudal 
+ 16-19 caudal = 49-53 total. 
Morphometric data for H. meeki is sum- 
marized in Table 6, for H. unifasciatus in 
Table 7. Lower jaw length 0.79 to 1.54 of 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


374 


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VOLUME 106, NUMBER 2 


35 


Table 2.— Numbers of total gill rakes on second arch in populations of Hyporhamphus meeki and H. unifas- 


clatus. 
Population 19° sp" 21 2? 23 24 25 26 a7 28 29 30 n ¥ 
H. meeki 
MA to GA a a —_ 2 11 53 80 33 8 189 26.8 
E. coast FL 2 4 8 Ps 2 11 29 40 10 6 114 25.9 
W. penin. FL 12 50 60 40 14 1 179 26.0 
FLA pan. to TX l 4 11 26 44 53 10 5 2 156 26.2 
Yucatan 1 3 6 10 5 2 24 23:5 
Species total 4 4 9 6 30 104 196 218 69 20 2 662 26.2 
H. unifasciatus 
Florida Z 4 616 18 17 5 59 22.9 
West Indies y = = it ps 46 a7 11 5 1 141 24.1 
Cent. America 2 5 10 i 6 - 1 35 2a 5 
Carib. S. Am. 6 27 33 a3 24 1 114 24.3 
South America * LG 31 33 21 2 1 109 235 
Yucatan 1 1 5 4 1 12 23-3 
Bermuda Sr>-34--3F iy 5 2 100 21.9 
Species total 2 4f-—§2---$6.--—-133---—-149 95 37 8 1 570 23-5 


head length with 83% of 240 specimens 
greater than or equal to 1.0. Ratio of LJL 
to SL 0.20 to 0.35 with 95% of 240 speci- 
mens 0.22 to 0.30. PREORB to ORB ratio 
0.61 to 1.0, usually 0.70 to 0.90 (92% of 
265 specimens). Distance from base of an- 
teriormost pelvic ray to caudal base extends 
anteriorly to mid-eye, usually between pos- 


terior portion of eye and posterior margin 
of opercular membrane. Dorsal- and anal- 
fin bases about equal, ratio of ABASE to 
DBASE 0.83 to 1.07, mean of 0.96 for 265 
specimens. Origin of dorsal fin over that of 
anal fin. Bases of dorsal and anal fins cov- 
ered with scales. Distance from anterior- 
most pectoral ray to origin of pelvic fin less 


Table 3.— Numbers of dorsal-fin rays in populations of Hyporhamphus meeki and H. unifasciatus. 


Population 12 13 14 
H. meeki 
MA to GA 2 100 
E. coast FL 8 40 
W. penin. FL BY 
FL pan. to TX l 5 90 
Yucatan 15 
Species total 1 15 297 
H. unifasciatus 
Florida 14 
West Indies 1 42 
Cent. America 6 
Carib. S. Am. 16 
South America l 32 
Yucatan l 
Bermuda 5 48 


Species total 7 


15 16 17 n BS 
56 1 1 160 14.4 
34 3 85 14.4 
95 10 157 14.7 
65 4 165 14.4 
12 27 14.5 
262 18 1 594 14.5 
28 6 48 14.8 
77 7 127 14.7 
35 3 44 14.9 
92 10 118 14.9 
76 11 120 14.8 

9 2 12 15.1 
27 2 82 14.3 
344 41 551 14.8 


376 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 4.—Numbers of anal-fin rays in populations of Hyporhamphus meeki and H. unifasciatus. 


Population 14 15 

H. meeki 

MA to GA 1 18 

E. coast FL 1 31 

W. penin. FL i 

FE pan. to TX. 10 59 

Yucatan 5 
Species total 12 124 
H. unifasciatus 

Florida 

West Indies i) 

Cent. America 4 

Carib. S. Am. 8 

South America 1 9 

Yucatan 

Bermuda 2 
Species total 1 38 


than the distance from the pelvic fin origin 
to the caudal base. Median pore of preor- 
bital canal usually posterior, rarely medial. 

Color. —Coloration in life is a silvery light 
tan-green. The fleshy tip of the lower jaw is 
a bright orange red. 

Size. — Adults attain a maximum size of 
179 mm SL (USNM 90798, Cape Charles, 
Virginia). 

Habitat.—All specimens observed were 


ZOE 


16 17 18 n x 
107 36 1 163 16.1 
43 10 85 5:7 
98 45 1 157 16.2 
80 16 165 15.6 
pA 3 27 16.0 
349 110 2 597 15.9 
23 22 1 48 16.5 
83 29 1 128 16.1 
2 13 44 16.2 
63 47 118 16.3 
63 45 1 119 16.3 
10 2 12 16.2 
Jad 46 12 81 16.8 
204 15 550 16.4 


collected near the surface of inshore or es- 
tuarine waters. Specimens collected by the 
first author in Chesapeake Bay and the Gulf 
of Mexico (Florida) were in areas with a 
sandy substrate and in proximity of sub- 
merged aquatic vegetation (eel grass, Zos- 
tera). As is often the case in estuarine con- 
ditions, the water inhabited by H. meeki is 
generally turbid. 

Early life history.—Larvae of H. meeki 


Table 5.—Numbers of pectoral-fin rays in populations of Hyporhamphus meeki and H. unifasciatus. 


Population 9 10 
H. meeki 
MA to GA 2 
E. coast FL 19 
W. penin. FL 
FL pan. to TX 1 
Yucatan 
Species total 22 
H. unifasciatus 
Florida 19 
West Indies 39 
Cent. America 2 Ds 
Carib. S. Am. 9 
South America 14 
Yucatan 
Bermuda 104 
Species total 5 187 


11 12 13 n xe 
75 59 3 158 114 
70 V7 106 11.0 
49 98 147 PR 
V7 41 1 120 11.4 
18 9 ZF 173 
289 224 4 558 11.4 
45 5 69 10.8 
94 3 136 10.7 
39 2 41 10.9 
108 3 120 11.0 
104 4 122 10.9 
11 1 12 1 
107 10.0 

397 18 607 10.7 


VOLUME 106, NUMBER 2 


377 


Table 6.—Summary of morphometric data in percent standard length in populations of Hyporhamphus meeki, 


except for SL in mm. 


Atlantic Gulf 
ae n Min Max Mean SD SE n Min Max Mean SD SE 
SL 169 48.1 179 9697) 27akbi< 2.09 124 63.6 176 128:9 DSsli 2.14 
Bsr; 169.) 31:8 37.8 34.5 1.01 0.08 124+ Si 38.1 34.6 1209" . @:10 
P,-C 169 39.2 46.2 43.0 1.04 0.08 124 41.3 45.7 43.6 G91, 70:08 
| Ws bE for 20:6 Sa. 2G 2:32 048 107° «205 29:8 259 1.88 0.18 
HDL £69" * 19.1 34.4 24.4 roy ONG 124 219 26.0 2300 0:76 0.07 
UJL 169 D2 4.8 4.2 Hs2) 0.02 124 an 4.8 4.3 O:231 'O.02 
UJW 169 4.3 6.2 D2 O32, 0.02 124 4.7 5.8 52 OPA G.02 
BD-P, 169 8.6 13.1 117 On 9e. (O06 124° VO 1333 19 0.62 0.06 
BD-P, 169 72 15.9" ' eA G62) OZ 124 9.0 1523 1229 20) (QI! 
ABASE 169 = 12.3 17.3 14.4 OS” 006 124. ~ 12-5 16.0 3e9 O:72; 0:06 
DBASE SS “13-5 16.9 14.8 0.67 0.05 124° sb32 16.4 14.6 0.67 0.06 
Bae 161 12.5 16.4 14.6 O72 0:06 109° 13h 16.3 14.5 0.64 0.06 
ORB 169 4.8 vel 5:9 0.41 0.03 124 5.0 6.9 ai) O39) 0.03 
PREORB 169 3.4 6.1 4.5 023. 0:02 124 3.4 3.2 4.2 O30) 0:03 


have been described (as Hyporhamphus sp.) 
from Chesapeake Bay, which may indicate 
utilization of estuarine waters as nursery ar- 
eas (Hardy & Johnson 1974). Larvae were 
collected along the Gulf coast of Florida 
most frequently during spring and summer 
in less than 30 m of water (Houde et al. 
1979). The particulars of spawning are un- 
known. Eggs have been attached to floating 
Zostera blades (by their adhesive filaments) 
Over vegetated habitats during summer 
months in Chesapeake Bay (Olney and 
Boehlert 1988). 

Distribution. — Atlantic coast of the Unit- 
ed States from Miami, Florida to Cape Cod, 
Massachusetts and rarely north to Cham- 
cook, Passamaquoddy Bay, New Brunswick 
(Leim & Day 1959), and in the Gulf of Mex- 
ico from the Everglades to Galveston, Texas 
(Fig. 1). Also occurs in Yucatan. The ob- 
served water temperature range is 13.7 to 
34.9°C, so H. meeki has a subtropical to 
temperate distribution. Sympatric with H. 
unifasciatus on the east coast of Florida from 
St. Lucie Inlet south to Miami and on the 
west coast from the Everglades to Tampa 
Bay. 

Etymology.—Named after Seth E. Meek 
who first separated the two species we rec- 


ognize here (Meek and Goss 1884:223)... 
“‘all the specimens... thus far taken on the 
Atlantic coast of the United States north of 
the Florida Keys ... belong to a species 
differing from the West Indian unifasciatus, 
in the slenderness of body and in the greater 
length of the lower jaw,” but misapplied the 
name H. roberti to the northern species. 

Comparisons. — Meristic characters allow 
for statistical separation of H. meeki from 
H. unifasciatus (alpha = 0.05; Tables 8-12), 
though infraspecific variation exists within 
populations of both species. Hyporhamphus 
meeki is discernable from the southern spe- 
cies H. unifasciatus in usually having more 
gill rakers on both the first and second arch- 
es (Tables 1 and 2). Ninety two percent of 
the 780 H. meeki specimens examined have 
total RGR, counts from 33 to 40, whereas 
in H. unifasciatus 91% of 657 specimens 
examined have total RGR, counts from 28 
to 32. Second arch gill rakers in 76% of 662 
specimens of H. meeki range from 26 to 29, 
whereas 92% of 570 specimens of H. uni- 
fasciatus range from 19 to 25. Pectoral fin 
usually with 11 or 12 rays, 10 or 11 in H. 
unifasciatus. 

Regressions of morphometrics also allow 
us to distinguish the two species. This is best 


378 


Table 7.—Summary of morphometric data in percent standard length in populations of //yporhamphus unifasciatus, except for SL in mm. 


Bermuda 


Central and South America 


West Indies and Florida 


SE 


Mean SD 


Max 


Min 


SE 


SD 


Max Mean 


Min 


SE n 


SD 


Min Max Mean 


n 


Character % SL 


2.36 
0.10 


0.11 


109.3 19.57 


3.63 09° 79:7 *166 


[29:4 633.62 


205 


62.4 


73 


a 


l19.4~ 82.55 


195 


ee) 


76 
76 
76 
59 


76 


SL 


0.86 
0.91 
ae 
0.86 
0.27 
0.37 
Ol 
1.38 
0.78 
0.78 
0.52 
0.39 
0.19 


36.7 


39.0 


38.2 35.4 


47.2 


30.8 


40.1 


44.7 41.7 


39.5 


69 
50 


el OFle 
1.86 
0.96 69 


0.27 
0.53 


45.8 43.2 


86 38.9 


0.15 
0.39 
0.12 
0.04 
0.05 
0.14 


1.33 
3.0 


43.7 


a 


0.39 
0.10 
0.03 
0.04 
0.09 
0.17 
0.09 
0.09 
0.07 
0.05 
0.02 


22.9 


30.9 


19.7 


23.8 0.21 
0.10 
0.03 
0.06 


aS 


3 81 19.8 


oME(6) PJ) 


15.0 


L 


ie 


24.7 22.8 


19.5 


25.6 24.1 


21.0 


$6 
86 


26.0 22.6 1.06 


19.6 


IL 


3.8 


3.1 4.5 


4.0 


69 


4.9 4.1 


3.4 
4.4 


ofall 


5.8 


69 


5.5 


6.8 


86 


0.40 


5.3 


76 
76 


76 


—_—_=_=—_= 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


= —_— —_—=— —_— — 


0.26 
0.11 


2.41 
0.99 


11.9 


69 


12.4 


64 
69 


0.10 
0.04 
0.03 


0.91 
0.37 


Ce 


86 


$512.1 


0.12 


= —_—=— —_— 


= —_— = 


7.9 


BD-P, 


BD-P, 


ABASE 
DBASE 
PL 
ORB 


58 
76 
76 


5.8 
otf 


6.7 


5.0 


5.8 


7.0 


0.04 86 4.9 
0.03 85 


0.39 


5.8 


6.7 


oil 


4.3 


2.9 


oil 


PREORB 


illustrated in the preorbital on orbit rela- 
tionship (Fig. 3). The slopes of the regres- 
sions are significantly different (P > F > 
.001). Ratio of preorbital length to orbit di- 
ameter is usually greater than 0.70 in H. 
meeki (92% of 265 specimens examined), 
but less than 0.70 in H. unifasciatus (75% 
of 224 specimens). Lower jaw length on 
standard length tends to be greater in H. 
meeki (Fig. 4), however, there is consider- 
able infraspecific variation within popula- 
tions of the two species especially in H. un- 
ifasciatus. 

Geographic variation in morphology is 
not reported in detail herein. Analysis to 
date indicates that morphology of Atlantic 
and Gulf populations of H. meeki differs 
slightly. Southern Florida and Yucatan ap- 
pear to be areas of sympatry between H. 
meeki and H. unifaciatus. Collection data 
indicate that sympatry of the two species in 
Florida may be largely avoided temporally, 
because both species migrate northward up 
the Florida coasts when the waters warm 
during summer and autumn; consequently 
H. unifasciatus moves into areas occupied 
by H. meeki in the winter. The dynamics 
of their sympatry in Yucatan is probably 
different than in peninsular Florida. Indi- 
vidual collections from Yucatan (UMMZ 
143085 and MCZ 32881) contain both spe- 
cies. Yucatan H. meeki have a preorbital/ 
orbit ratio typical of H. unifasciatus. Yu- 
catan may be an area of hybridization or 
introgression, and needs further study. 

Comparisons of meristic means by the 
Tukey-Kramer method indicate some geo- 
graphic variability in populations of both 
H. meeki and H. unifasciatus. There ap- 
pears to be little consistency in trends in H. 
meeki populations across the different me- 
ristic characters. The only trend with any 
consistency is the position of the west coast 
of the peninsula of Florida population. It is 
usually at the extreme of the range of means, 
being closest to mean values of H. unifas- 
ciatus populations. The one trend that stands 
out in populations of H. unifasciatus is that 


VOLUME 106, NUMBER 2 3719 


Table 8.—Comparison of means for dorsal-fin ray counts for populations of Hyporhamphus meeki and H. 
unifasciatus, and between H. meeki and H. unifasciatus. Populations joined by the same line are not significantly 
different by the Tukey-Kramer method, alpha = 0.05. 


Population n XG TK 
H. meeki 
West peninsula of FL 157 14.7 | 
Yucatan 26 14.5 | | 
FL panhandle to TX 165 14.4 | 
Atlantic coast of FL 85 14.4 | 
MA to GA 160 14.4 | 
H. unifasciatus 
Yucatan 12 15Al | 
Caribbean South America 118 14.9 | | 
Central America 44 14.9 | | 
Florida 48 14.8 | | 
South America (Brazil) 120 14.8 | | 
West Indies 127 14.7 | 
Bermuda 82 14.3 
H. meeki 594 14.5 | 
H. unifasciatus 551 14.8 | 


Bermuda is always at anextreme oftherange  ciatus (62.4—205) from 54 collections, with 
of means, usually at the lower extreme. almost complete morphometric and meris- 

Material examined.—338 specimens of tic dataare listed. Additional specimens used 
H. meeki (48.1-179 mm SL) from 50 col- mainly for meristic data are in the second 
lections, and 230 specimens of H. unifas-  author’s files. 


Table 9.—Comparison of means for anal-fin ray counts for populations of Hyporhamphus meeki and H. 
unifasciatus, and between H. meeki and H. unifasciatus. Populations joined by the same line are not significantly 
different by the Tukey-Kramer method, alpha = 0.05. 


Population n X TK 
H.. meeki 
West peninsula of FL 157 16.2 | 
MA to GA 163 16.1 | 
Yucatan 26 16.0 | | 
Atlantic coast of FL 85 15.7 | | 
FL panhandle to TX 165 15.6 | 
H. unifasciatus 
Bermuda 81 16.8 | 
Florida 48 16.5 | | 
Caribbean South America 118 16.3 | 
South America (Brazil) 119 16.3 | 
Central America 44 16.2 | 
Yucatan 12 16.2 | 
West Indies 128 16.1 | 
H. meeki 597 15.9 


H. unifasciatus 550 16.4 | 


380 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


50 


LL 
>) 


LOWER JAW LENGTH (mm) 
No 
fo) 


—. 
O 


0 30 100 


150 


200 


250 


STANDARD LENGTH (mm) 


Fig. 4. Relationship of lower jaw length to standard length in Hyporhamphus meeki (squares) and two 
populations of H. unifasciatus, Central and South America (triangles) and West Indies (diamonds). 


Hyporhamphus meeki 


Holotype. —USNM 294369 (1, 160) 
Morehead City, NC; 5 Sept 1964. 

ATLANTIC U.S. (paratypes): USNM 
132257 (4, 55.7-115) Woods Hole, MA; 2 
Sep 1876. USNM 68368 (5, 54—-65.8) Great 
South Bay, NY; 1898. USNM 187214 (2, 
73.6-98.9) Deep Cove, Indian R., DE; 7 
Sept 1956. VIMS-CBL 173 (2, 108-123) 
CBL Pier, MD; 21 Jul 1936. VIMS-CBL 
1523 (11, 73.6-91.1) Broomes I., Patuxent 
R., MD; 13 Aug 1958. USNM 90798 (1, 
179) Cape Charles, VA. VIMS 61 (21, 87.6- 
123) Lynnhaven Inlet, Broad Bay, VA; 29 
Sept 1954. VIMS 607 (7, 78.5—159) York 
R., VA; 18 Jul 1970. USNM 131146 (1, 
147) Cape Charles, VA. USNM 131146 (4, 
102-117) Morehead City, NC; 5 Sept 1964; 


taken with the holotype. VIMS-CBL 3571 
(1, 51.5) Morehead City, NC; 9 Jul 1967. 
USNM 51878 (1, 153) Beaufort, NC. VIMS 
7873 (3, 71.9-115) Carteret Co., Atlantic 
Ocean beach, NC; 12 Sept 1976. USNM 
294396 (1, 150) SC, 32°26'N, 79°50'W; 6 
Aug 1953. USNM 149969 (1, 107) Georgia 
coast. USNM 294377 (2, 99.7-104) St. Si- 
mons I., GA; 15 Mar 1956. USNM 294426 
(1, 119) St. Simons I., GA; 14 Apr 1960. 
USNM 294427 (1, 105) St. Simons I., GA; 
16 Nov 1955. USNM 294441 (1, 160) St. 
Simons I., GA; 11 Nov 1956. USNM 
294435 (1, 55.6) off Savannah, GA, 31°41'N, 
80°35’; 21 Oct 1953. 

Other material examined: ATLANTIC 
U.S.: VIMS uncat. (56, 48.1—146), Glouces- 
ter Pt., York R., VA; 19 Jul 1989 to 17 Sept 
1989. 


VOLUME 106, NUMBER 2 


381 


Table 10.—Comparison of means for pectoral-fin ray counts for populations of Hyporhamphus meeki and H. 
unifasciatus, and between H. meeki and H. unifasciatus. Populations joined by the same line are not significantly 


different by the Tukey-Kramer method, alpha = 0.05. 


Population n 
H. meeki 
West peninsula of FL 147 
MA to GA. 158 
FL panhandle to Texas 120 
Yucatan 26 
Atlantic coast of FL 106 
H. unifasciatus 
Yucatan 12 
Caribbean South America 120 
South America (Brazil) 122 
Central America 41 
Florida 69 
West Indies 136 
Bermuda 107 
H. meeki 558 
H. unifasciatus 607 


ATLANTIC FLORIDA: UF 62140 (1, 
125) 4.5 miles N. of Jupiter Inlet; 21 Aug 
1964. UF 77037 (2, 111-127) Matheson 
Hammock; 11 Jul 1970. UF 83999 (20, 
59.4-97.8) Indian R., Brevard Co.; 16 Aug 
1976. USNM 294365 (4, 55.8-141) off New 


>< 


TK 


ey | 

11.4 | | 
11.4 | 
ie | 
11.0 


Lr 
1t8 
109 
10.9 
10.8 
10.7 
10.0 


11.4 
10.7 | 


Smyrna Bch., 29°00’N, 80°32’'W; 14 Oct 
1953. USNM 294397 (19, 52.3-119) 
29°40'N, 81°06’W; 15 Oct 1953. USNM 
294494 (16, 91.4-145) 27°52'N, 80°26’W; 
20 Jan 1961. 

GULF COAST FLORIDA: UF 1010 (9, 


Table 11.—Comparison of means for total first arch gill raker counts for populations of Hyporhamphus meeki 
and H. unifasciatus, and between H. meeki and H. unifasciatus. Populations joined by the same line are not 
significantly different by the Tukey-Kramer method, alpha = 0.05. 


Population n 
H. meeki 
Yucatan 26 
FL panhandle to TX 207 
Atlantic coast of FL 101 
MA to GA 25) 
West peninsula of FL 194 
H. unifasciatus 
Caribbean South America 119 
West Indies 170 
South America (Brazil) 124 
Central America 53 
Yucatan 2 
Florida 66 
Bermuda 113 
H. meeki 780 


H. unifasciatus 657 


X TK 


353 | 

35:2 | 

35.1 | | 
34.5 | 
33.6 


31.1 
3i2I 
30.6 | 
30.5 | 
30.0 | | 
30.0 | 
29.5 | 


34.6 
30.6 | 


382 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 12.—Comparison of means for total second arch gill raker counts for populations of Hyporhamphus 
meeki and H. unifasciatus, and between H. meeki and H. unifasciatus. Populations joined by the same line are 
not significantly different by the Tukey-Kramer method, alpha = 0.05. 


Population n 
H. meeki 
MA to GA 189 
FL panhandle to TX 156 
West peninsula of FL 17z. 
Atlantic coast of FL 114 
Yucatan 38 


H. unifasciatus 


Caribbean South America 114 
West Indies 141 
South America (Brazil) 109 
Central America 35 
Yucatan 12 
Florida 59 
Bermuda 100 
H. meeki 662 
H. unifasciatus 570 


63.6-88.1) Cedar Key; 24 Jul 1948. UF 
51083 (5, 71.8-135) Alligator Harbor; 7 Sept 
1954. UF 52066 (5, 146-170) Alligator 
Harbor; 16 May 1954. UF 68483 (6, 114— 
148) St. Andrew’s Bay, Panama City; 8 Oct 
1968. UF 76904 (24, 50.1-74.2) 1 mile N 
of Lee-Collier Co. line; 18 Jun 1966. USNM 
125446 (2, 139-153) Tarpon Springs; 5 Nov 
1896. USNM 184268 (4, 133-146) Johns 
Pass, Madeira Bch.; 29 Sept 1958. USNM 
294431 (5, 103-136) Sarasota; 24 Aug 1967. 
USNM 294452 (16, 119-134) Sanibel I.; 11 
Aug 1959. USNM 294489 (6, 149-171) 
Sanibel I.; 8 Aug 1964. 

GULF COAST U.S., ALABAMA-TEX- 
AS: USNM 187122 (2, 151-152) S. Mobile, 
AL; 7 Jul 1960. VIMS 5095 (1, 113) Pelican 
Bay, Dauphin I., AL; 29 Aug 1974. USNM 
147781 (1, 176) Mississippi Gulf coast; 
1948. USNM 187120 (2, 123-128) inside 
Chandeleur Sound, LA; 19 Aug 1959. 
USNM 187123 (1, 73.9) Grande Isle, LA; 
21 Jul 1930. USNM 94546 (1, 141) Corpus 
Christi, TX. USNM 103390 (1, 69.1) near 
Corpus Christi, TX; 1937. USNM 120056 
(2, 128-155) Galveston, TX; 1941. USNM 


) 
Meee 
n 


SS) 
‘ea 
Ue 


21.9 


26.2 
235 | 


187119 (2, 168-172) Corpus Christi, Sham- 
rock Cove, TX; 7 Apr 1927. USNM 294440 
(26, 100.5-152) Aransas Pass, Institute of 
Marine Science pier, TX; Mar 1959. 

YUCATAN: UMMZ 143085 (28, 94.2-— 
145) west of Progreso; 28 Mar to 1 Apr 
1936. UMMZ 143087 (15, 64.2—152) 
Chicxulub, near Progreso; 1 Apr 1936. MCZ 
32881 (3, 142-183) Yucatan; 1906. 


Hyporhamphus unifasciatus 


FLORIDA: UF 56209 (1, 143) Virginia 
Key; 26 Nov 1959. UF 62140 (11, 108-134) 
N. of Jupiter Inlet; 21 Aug 1964. USNM 
34999 (4, 182-192) Key West; Dec 1883. 
USNM 158069 (3, 108-129) Snipe and 
Content Keys; June 1956. USNM 38544 (3, 
138-144) Key West; 15-27 Apr 1884. 
USNM 187121 (4, 75.0-79.9) St. Joseph’s 
Bay22 Febsi9s9: 

WEST INDIES: USNM 5802 (2, 146- 
186) Barbados. USNM 5847 (1, 103) Ja- 
maica. USNM 8803 (1, 111) Jamaica. 
USNM 10730 (2, 79.5-89.9) Bahia Honda, 
Cuba. USNM 34938 (2, 118-123) St. 


VOLUME 106, NUMBER 2 


Thomas. USNM 38537 (2, 120-127) Ja- 
maica; 1-11 Mar 1884. USNM 38601 (6, 
119-158) Curacao; 10-18 Feb 1884. USNM 
50111 (1, 194) San Juan Mkt., Puerto Rico; 
14 Jan 1899. USNM 82366 (3, 77.2-86.3) 
Los Arroyos, Cuba; 19 May 1914. USNM 
94077 (4, 112-139) Jamaica. USNM 
107428 (2, 135-165) Bahia Honda An- 
chorage, Cuba; 5 Apr 1937. USNM 130652 
(1, 195) Cuba. USNM 132524 (3, 109-147) 
Port-au-Prince, Haiti; 22 Oct 1945. USNM 
294364 (5, 90.5—129) Sable Bay, Dominica; 
13 Nov 1964. USNM 294493 (6, 78.6-119) 
Jobes Harbor, Greater Antilles; 20 Feb 1966. 
USNM 294515 (11, 81.9-167) 17°56'30’N, 
66°13'12”W; 18 Feb 1966. 

BERMUDA: AMNH 18711 (2, 87.3- 
90.2) Bermuda. ANSP 96626 (2, 114-117) 
Somerset, Bermuda; 12 Jun 1952. ANSP 
109562 (15, 82.5—135) Somerset, Bermuda; 
4 Jun 1952. ANSP 123715 (4, 100-141) 
Bermuda; Jun 1930. MCZ 34890 (5, 110- 
168) Bermuda. MCZ 40757 (6, 106-118) 
Bermuda; 1872. UMMZ 172321 (3, 99.2- 
154) Reach at Bio. Sta., Bermuda; 31 May 
1951. UMMZ 172369 (3, 81.6—-97.9) St. 
George’s I., Bermuda; 5 Jun 1951. UMMZ 
172418 (13, 87.9-145) St. George’s I., Ber- 
muda; 8 Jun 1951. UMMZ 175957 (2, 85.9- 
108) Reach at Bio. Sta., Bermuda. UMMZ 
175967 (3, 124-140) Jetty at Bio. Sta., Ber- 
muda; 19 Mar 1957. UMMZ 175974 (1, 
105) Ferry Reach at Bio. Sta., Bermuda; 21 
Mar 1957. UMMZ 175981 (1, 79.7) Ferry 
Reach at Bio. Sta., Bermuda; 24 Mar 1957. 
UMMZ 176014 (1, 148) Ferry Reach at Bio. 
Sta., Bermuda; 14 Apr 1957. UMMZ 
176154 (1, 114) Ferry Reach at Bio. Sta., 
Bermuda; 4 Jun 1957. USNM 294439 (7, 
87.7-110) Bermuda Harbor, Bermuda; 9 
Mar 1963. 

CENTRAL AMERICA: MCZ 32881 (1, 
183) Yucatan, Mexico; 1906. UF 7107 (2, 
165-181) Veracruz, Mocambo, Mexico; 20 
Jan 1958. UMMZ 143085 (10, 94.2—140) 
W. of Progreso, Yucatan, Mexico; 28 Mar— 
1 Apr 1936. UMMZ 143087 (1, 71) Chicxu- 
lub, Yucatan, Mexico; 1 Apr 1936. USNM 


383 


79658 (1, 160) Colon Mkt., Panama; 23 Jan 
1912. USNM 187843 (9, 62.4-85.5) off 
Bluefields, Nicaragua; 3-4 Jun 1962. 

SOUTH AMERICA: USNM 203826 (11, 
104-121) Gulf of Uraba, Colombia; 11 Jul 
1966. USNM 206658 (17, 121-205) Baru 
I., Colombia; 26 Sept 1969. USNM 38574 
(1, 135) Sabanilla, Colombia; 16-22 Mar 
1884. USNM 94764 (1, 188) near Puerto 
Colombia, Barranquilla, Colombia. USNM 
128286 (3, 121-129) Gulf of Venezuela, 
Venezuela; 5 Apr 1925. USNM 198404 (3, 
86.1-108) off French Guiana; 30 Jul 1956. 
MZUSP 5206 (7, 131-148) Ubatuba, Bra- 
zil; 1967. MZUSP 41092 (3, 184-191) Praia 
de Itapenia, Brazil; Jul 1965. MZUSP 41094 
(12, 131-179) Pontal, Ilheus, Brazil; 25 Oct 
1971. USNM 107220 (1, 151) Recife, Bra- 
Zi 1932, 


Acknowledgments 


For access to specimens mentioned in this 
paper, we thank E. B. Bohlke and W. F. 
Smith Vaniz (ANSP), G. H. Burgess (UF), 
W. N. Eschmeyer and D. Catania (CAS), 
M. N. Feinberg (AMNH), K. E. Hartel 
(MCZ), N. M. Menezes (MZUSP), D. W. 
Nelson (UMMZ), and R. Rosenblatt (SIO). 
We thank the staff of the many institutions 
that house the material we have examined 
for their help, Mildred Carrington for draw- 
ing the figures of Hyporhamphus (Fig. 2), 
and Janet Nestlerode for assistance with 
summarizing data and for plotting the dis- 
tribution map (Fig. 1). Carole Baldwin, Bar- 
ry Chernoff, Thomas Munroe, and Lynne 
Parenti made valuable comments on drafts 
of the manuscript. The second author thanks 
Frederick Berry, formerly of the National 
Marine Fisheries Service, for an introduc- 
tion to the study of halfbeaks and co-dis- 
covery of this new species in the early 1960's. 
This is Contribution No. 1776 from the Vir- 
ginia Institute of Marine Science. 


Literature Cited 


Beebe, W., & J. Tee-Van. 1933. Field book of the 
shore fishes of Bermuda. G. P. Putman’s Sons, 
New York, 337 pp. 


384 


Bigelow, H. B., & W. C. Schroeder. 1953. Fishes of 
the Gulf of Maine.— U.S. Fish & Wildlife Ser- 
vice, Fishery Bulletin 53:1-577. 

Bruce, N. L. 1986. Revision of the isopod crustacean 
genus Mothocya Costa, in Hope, 1851 (Cymo- 
thoidae: Flabellifera), parasitic on marine fish- 
es.—Journal of Natural History 20:1089-1192. 

Collette, B. B. 1965. Hemiramphidae (Pisces, Synen- 

tognathi) from tropical West Africa. — Atlantide 

Report 8:217-235. 

1978. Hemiramphidae. in W. Fischer, ed., 
FAO species identification sheets for fishery 
purposes, western Central Atlantic (fishing area 
31), FAO, Rome. 

Hardy, J. D., Jr., & R. K. Johnson. 1974. Descrip- 
tions of halfbeak larvae and juveniles from 
Chesapeake Bay (Pisces: Hemiramphidae).— 
Chesapeake Science 15(4):241-246. 

Hildebrand, S. F. 1946. A descriptive catalog of the 
shore fishes of Peru. Bulletin, U.S. National Mu- 
seum 189, 530 pp. 

Hoese, H. D., & R. H. Moore. 1977. Fishes of the 
Gulf of Mexico, Texas, Louisiana, and adjacent 
waters. Texas A & M University Press, College 
Station, 327 pp. 

Houde, E. D., J. C. Leak, C. E. Dowd, S. A. Berkeley, 
& W. J. Richards. 1979. Ichthyoplankton 
abundance and diversity in the eastern Gulf of 
Mexico.— U.S. Department of Commerce PB— 
299-839. 

Jordan, D. S., & B. W. Evermann. 1896. The fishes 
of North and Middle America.— Bulletin, U-S. 
National Museum 47, pt. 1:1-1240. 

Leim, A. H., & L. R. Day. 1959. Records of uncom- 
mon and unusual fishes from eastern Canadian 
waters, 1950-1958.—Journal of the Fisheries 
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Leviton, A. E., R. H. Gibbs, Jr., E. Heal, & C. E. 
Dawson. 1985. Standards in herpetology and 
ichthyology: part I. Standard symbolic codes for 
institutional resource collections in herpetology 
and ichthyology.—Copeia 1985(3):802—832. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Meek, S. E., & D. K. Goss. 1884. A review of the 
American species of the genus Hemirham- 
phus. —Proceedings of the Academy of Natural 
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of Panama. Part I.—Field Museum of Natural 
History, Zoological Series 15:1—330. 

Miller, R.R. 1945. Hyporhamphus patris, a new spe- 
cies of hemiramphid fish from Sinaloa, Mexico, 
with an analysis of the generic characters of Hy- 
porhamphus and Hemiramphus.—Proceedings 
U.S. National Museum 96:185-193. 

Olney, J. E., & G. W. Boehlert. 1988. Nearshore 
ichthyoplankton associated with seagrass beds 
in the lower Chesapeake Bay.— Marine Ecology 
Progress Series 45:33-43. 

Parin, N. V., B. B. Collette, & Yu. N. Scherbachev. 
1980. Preliminary review of the marine half- 
beaks (Hemiramphidae, Beloniformes) of the 
tropical Indo-West Pacific.—Trudy Instituta 
Okeanologii 97:7—173 (in Russian). 

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sertatio iv.—Novi Commentarii Academiae 
Scientiarum Instituti Bononiensis 5:339-365. 

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(HMB) College of William and Mary, 
Virginia Institute of Marine Science, 
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vice Systematics Laboratory, National Mu- 
seum of Natural History, Washington, D.C. 
20560, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(2), 1993, pp. 385-389 


A NEW SPECIES OF OEDIPINA 
(AMPHIBIA: CAUDATA: PLETHODONTIDAE) FROM 
NORTHERN HONDURAS 


James R. McCranie, Larry David Wilson, and Kenneth L. Williams 


Abstract. —A new species of Oedipina from a cloud forest locality in northern 
Honduras is described and illustrated. The new species, O. gephyra, shows 
several morphological characteristics that bridge those of the two species groups 
of Oedipina recognized in the most recent revision of the genus. However, 
based on derived features such as the uniform dorsal coloration, short limbs, 
small feet, and long tail, the new species fits more readily into the uniformis 


group than the parvipes group. 


Two species of Oedipina (cyclocauda 
Taylor 1952 and stuarti Brame 1968) have 
heretofore been recorded from Honduras 
(Brame 1968, Meyer & Wilson 1971). Re- 
cently, we collected salamanders of this ge- 
nus from a single hardwood cloud forest 
locality in the western portion of the Cor- 
dillera Nombre de Dios in northern Hon- 
duras. An examination of these specimens 
demonstrated that the population repre- 
sents an undescribed species. 


Methods 


All measurements are in millimeters, 
made to the nearest tenth with dial calipers 
with the aid of a dissecting microscope. 
Measurements taken, method of expressing 
ratios, and the way of counting costal groove 
numbers follow the methods of Brame 
(1968). Abbreviations used are SVL (snout 
vent length [=standard length of Brame)), 
MVZ (Museum of Vertebrate Zoology, Uni- 
versity of California at Berkeley), and 
USNM (National Museum of Natural His- 
tory). An X-ray of the holotype of the new 
species (USNM 316535) confirmed the 
number of trunk vertebrae. Maxillary and 
vomerine tooth row counts are both sides 
summed. Comparative data for the other 
species of Oedipina were taken from Brame 
(1968) and Brame & Duellman (1970), 


however, data for O. cyclocauda were sup- 
plemented by those from the following re- 
cently collected Honduran specimens: AT- 
LANTIDA: mountains behind La Ceiba, 
260 m elev., USNM 316539. YORO: 6.6 
km S Yoro, ca. 1000 m elev., MVZ 171078; 
32.0 km W Yoro, MVZ 167772. 


Oedipina gephyra, new species 
Fig, 1 


Holotype.—USNM 316535, an adult fe- 
male, from 2.5 airline km NNE La Fortuna 
(15°26'N, 87°18'W), 1690 m elev., Cordille- 
ra Nombre de Dios, Departamento de Yoro, 
Honduras, collected 14 Aug 1991 by James 
R. McCranie, Kenneth L. Williams, and 
Larry David Wilson. Original number LDW 
9597. 

Paratypes. —-USNM 316536-37, adult fe- 
males and USNM 316538, adult male, col- 
lected 14-16 Aug 1991 at the type locality, 
1690-1810 m. 

Diagnosis. — Oedipina gephyra can be dis- 
tinguished from all other species of Oedi- 
pina by the following combination of char- 
acteristics: 17 or 18 costal grooves per side; 
18 or 19 trunk vertebrae; 10-11 costal folds 
not covered by adpressed limbs; adult SVL 
52.3-57.8 (X¥ = 55.6); feet small, adult hind 
foot width 1.4-1.6 (¥ = 1.5), SVL/hind foot 


386 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Piet. 


width 34.9-39.6 (X = 37.1); feet extensively 
webbed (Fig. 2); coloration uniform black- 
ish-brown with tiny iridophores present on 
all surfaces. 

Description of holotype.—Adult female 
with SVL of 56.7. Snout bluntly rounded in 
dorsal aspect and in profile; head width 5.4; 
SVL/head width 10.50; head length 8.1; 
SVL/head length 7.00; nostrils small, situ- 
ated near tip of snout; distinct nasolabial 
groove extending from lower-posterior 
margin of each nostril to the lip; labial pro- 
tuberances absent; canthus rostralis mod- 
erately arched; eyes not protuberant, not 
visible beyond margin of jaw when viewed 
from below; suborbital groove distinct; 
postorbital groove shallow, extending pos- 
teriorly from eye before turning sharply 
ventrally to connect with the gular fold, an- 
other branch proceeding sharply ventrally 
just posterior to mandible, extending irreg- 
ularly across throat anterior to gular fold; 


Oedipina gephyra, new species, holotype, USNM 316535, SVL 56.7 mm. 


no dermal glands on head or body; 43 max- 
illary teeth, extending posteriorly to a point 
two-thirds distance through length of orbit; 
two premaxillary teeth, located posterior to 
lip; 22 vomerine teeth, in long, single, arched 
series, extending beyond outer edge of in- 
ternal nares; axilla-groin length 37.3; 17 
costal grooves per side; 18 trunk vertebrae; 
tail length 123.0; SVL/tail length 0.46; tail 
thick, nearly round at base, somewhat lat- 
erally compressed for last half of its length, 
barely constricted at base; tail width 3.6; 
SVL/tail width 15.75; tail depth 3.7; SVL/ 
tail depth 15.32; postiliac gland round, 
prominent; limbs short, 11 costal folds not 
covered when limbs adpressed to sides of 
trunk; hindlimb length 9.4; SVL/hindlimb 
length 6.03; hind foot width 1.5; SVL/hind 
foot width 37.80; digits one and two and 
three and four on forelimbs fused and digits 
one and two and four and five on hindlimbs 
fused, rest of digits on both fore- and hind- 


VOLUME 106, NUMBER 2 


limbs fully webbed; digits on forelimbs in 
order of decreasing length 3-2-4-1, those on 
hindlimbs 3-4-2-5-1. 

Coloration in life: Head, body, and limbs 
blackish-brown throughout; tail black on all 
surfaces; tiny iridophores (visible under 
magnification) present on all surfaces. 

Variation. — Variation in morphology and 
coloration of the paratypes is minimal. The 
premaxillary teeth pierce the lip, labial pro- 
tuberances are weakly developed, and a 
mental gland is slightly indicated in the adult 
male (USNM 316538). There are 17 costal 
grooves per side in two and 18 in one 
(USNM 316537). Both females have 11 cos- 
tal folds not covered by the adpressed limbs, 
whereas the male has 10. Digits four and 
five on the hindlimbs of USNM 316538 are 
not fused. Other variation in the entire se- 
ries (including the holotype) is as follows 
(characteristics for the male separated by a 
comma from that of the females; means in 
parenthesis): SVL 55.5-57.8 (56.7), 52.3; 
axilla-groin length 35.6—-38.6 (37.2), 33.8; 
head width 5.2-5.5 (5.4), 5.4; SVL/head 
width 10.50—10.67 (10.56), 9.69; head length 
8.1-8.4 (8.2), 8.4; SVL/head length 6.61- 
7.05 (6.89), 6.23; tail length 112.0-129.0 
(121.3), 115.0; SVL/tail length 0.45-0.50 
(0.47), 0.45; tail width 3.4—3.8 (3.6), 3.5; 
SVL/tail width 15.21-16.32 (15.76), 14.94; 
tail depth 3.4—3.7 (3.6), 3.2; SVL/tail depth 
15.21-16.32 (15.62), 16.34; hind foot width 
1.4-1.6 (1.5), 1.5; SVL/hind foot width 
36.13-39.64 (37.86), 34.87; hindlimb length 
7.6-9.4 (8.6), 8.4; SVL/hindlimb length 
6.03-7.30 (6.66), 6.23; maxillary teeth 43- 
54 (49.0), 48; vomerine teeth 16-22 (18.7), 
18; premaxillary teeth two in all. 

Natural history notes.— Oedipina gephyra 
was collected from inside rotten logs and 
stumps and underneath a plank within 
hardwood cloud forest (Lower Montane Wet 
Forest formation of Holdridge 1967) from 
1690 to 1810 m elev. Several specimens of 
another salamander, Nototriton barbouri 
(Schmidt 1936), were also found inside rot- 
ten logs at the type locality while another 


387 


TRA 


Fig. 2. Right hind foot of a paratype of Oedipina 
gephyra (USNM 316538). Line equals 1.0 mm. 


salamander, Bolitoglossa conanti McCranie 
& Wilson 1993, was common in arboreal 
situations. 

Comparisons. — Brame (1968) recognized 
two species groups of Oedipina, the parvipes 
group with four species and the uniformis 
group with 11 species and Brame & Duell- 
man (1970) described an additional species 
belonging to the uniformis group. Oedipina 
gephyra shows several characteristics dis- 
tinctive of each group, and therefore cannot 
be easily placed in either group. Oedipina 
gephyra agrees with the parvipes group and 
can be distinguished from each species in 
the uniformis group by having 17 or 18 cos- 
tal grooves per side, 18 or 19 trunk verte- 
brae, and extensively webbed feet (19-22 
costal grooves per side, 20-23 trunk ver- 
tebrae, feet usually not extensively webbed 
in the uwniformis group). On the other hand, 
O. gephyra resembles the uniformis group 
species and can be distinguished from those 


388 


in the parvipes group by lacking a white face 
mask and large whitish markings over much 
of the dorsal surfaces, in having relatively 
short limbs with 10-11 costal folds not cov- 
ered by the adpressed limbs, and in having 
relatively narrow feet with the hind foot 
width 1.4—1.6 mm (dorsal surfaces with large 
whitish spots or patches and usually a white 
face mask, 6—10 costal folds not covered by 
adpressed limbs, and hind foot width 1.6— 
3.8 mm in the parvipes group). Addition- 
ally, O. gephyra has a very long tail (SVL/ 
tail length: 0.45—0.50) similar to most spe- 
cies in the uniformis group, whereas the par- 
vipes group members have shorter tails 
(SVL/tail length 0.57—0.92 for the four spe- 
cies combined; data extrapolated from 
Brame 1968). Also, most members of the 
uniformis group have numerous maxillary 
teeth like O. gephyra, whereas most species 
in the parvipes group have zero to few max- 
illary teeth (Brame 1968). 

According to Brame (1968), the two char- 
acters O. gephyra shares with members of 
the parvipes group (reduced number of trunk 
vertebrae and extensively webbed feet) are 
ancestral traits whereas, four of the five 
characters most similar to members of the 
uniformis group (uniform dorsal coloration, 
relatively short limbs, relatively small feet, 
and a relatively tong tail) are derived states. 
The fifth character most similar to the wni- 
formis group (number of maxillary teeth: 
although one species in the parvipes group, 
O. complex [Dunn 1924] also has numerous 
maxillary teeth) is an ancestral trait. Thus 
in all derived features, the new species is 
most similar to the members of the unifor- 
mis group. Therefore, we believe that the 
closest relationships of O. gephyra lie within 
the uniformis group. 

In addition to the group characteristics 
discussed above, O. gephyra can be further 
distinguished from the two Honduran 
members of the wniformis group as follows: 
from cyclocauda by larger size (SVL 52.3- 
57.8 versus maximum of 50.2) and having 
longer limbs (SVL/hindlimb length 6.0-7.3, 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


X = 6.5 versus 7.2-9.0, ¥ = 8.2) and from 
stuarti by having a slightly wider head (SVL/ 
head width 9.7-10.7, ¥ = 10.3 versus 11.1- 
12.3, ¥ = 11.7, longer limbs (SVL/hindlimb 
length 6.0-7.3, ¥ = 6.5 versus 8.7-9.5, X = 
9.1), narrower feet (SVL/hind foot width 
34.9-39.6, X = 37.1 versus 30.6-30.8, ¥ = 
30.7), and in lacking dermal glands (dermal 
glands numerous on head and dorsum in 
stuarti). Three species in the O. uniformis 
group (collaris [Stejneger 1907], poelzi 
Brame 1968, and pseudouniformis Brame 
1968) are like O. gephyra in having long 
limbs compared to the remaining species in 
the group. Oedipina gephyra can be distin- 
guished from each of these species, in ad- 
dition to the number of trunk vertebrae and 
amount of webbing on the feet, as follows: 
from collaris by size (52.3-57.8, X = 55.6 
versus 57.7-77.1, X¥ = 69.2), snout shape 
(bluntly rounded versus elongated), head 
width (SVL/head width 9.7-10.7, ¥ = 10.3 
versus 9.1-9.9, ¥ = 9.3), and maxillary tooth 
number (43-54, ¥ = 48.8 versus 80-98, X 
= 87.5); from poelzi by coloration (uniform 
blackish-brown versus broad brownish dor- 
sal band bordered with a cream or yellow- 
ish-white thin lateral stripe), head width 
(SVL/head width 9.7-10.7, ¥ = 10.3 versus 
5.1-6.7, X = 6.1), and hind foot width (SVL/ 
foot width 34.9-39.6, X = 37.1 versus 25.0- 
28.9, X = 26.5); from pseudouniformis by 
head width (SVL/head width 9.7-10.7, ¥ = 
10.3 versus 8.7-9.7, X = 9.3) and hind foot 
width (SVL/foot width 34.9-39.6, X¥ = 37.1 
versus 30.2-32.1, ¥ = 31.4). 

Etymology. —The word gephyra is trans- 
literated from Greek (meaning bridge) and 
refers to the species showing several mor- 
phological characteristics that bridge the two 
species groups recognized by Brame (1968) 
in the most recent revision of the genus. 


Acknowledgments 


Collecting and exportation permits were 
provided by R. Soto Rivera of the Corpora- 
cion Hondurena de Desarrollo Forestal, Te- 


VOLUME 106, NUMBER 2 


gucigalpa. J. Porras Orellana, Tegucigalpa, 
has provided much valuable help to us 
through the years. Northwestern State Uni- 
versity provided grant money that aided 
KLW in his field work. The X-ray of the 
holotype was provided by J. M. Savage for 
which we are very grateful. D. Wake and B. 
Stern, University of California at Berkeley, 
loaned comparative material. 


Literature Cited 


Brame, A. H., Jr. 1963. A new Costa Rican sala- 
mander (genus Oedipina) with a re-examination 
of O. collaris and O. serpens.—Los Angeles 

County Museum, Contributions in Science 65: 

1-12. 

1968. Systematics and evolution of the Me- 
soamerican salamander genus Oedipina. —Jour- 
nal of Herpetology 2:1-64. 

—, & W.E. Duellman. 1970. A new salamander 
(genus Oedipina) of the uniformis group from 
western Panama.—Los Angeles County Muse- 
um, Contributions in Science 201:1-8. 

Dunn, E.R. 1924. New amphibians from Panama. — 
Occasional Papers of the Boston Society of Nat- 
ural History 5:93-95. 

Holdridge, L.R. 1967. Life zone ecology. Second ed. 


389 


Tropical Science Center, San José, Costa Rica, 
206 pp. 

McCranie, J. R., & L. D. Wilson. 1993. A review of 
the Bolitoglossa dunni group (Amphibia: Cau- 
data) from Honduras with the description of 
three new species. — Herpetologica 49:1-15. 

Meyer, J. R., & L. D. Wilson. 1971. A distributional 
checklist of the amphibians of Honduras.— Los 
Angeles County Museum, Contributions in Sci- 
ence 218:1-47. 

Schmidt, K. P. 1936. New amphibians and reptiles 
from Honduras in the Museum of Comparative 
Zoology. — Proceedings of the Biological Society 
of Washington 49:43-50. 

Stejneger, L. 1907. A new salamander from Nica- 
ragua. Proceedings of the United States—Na- 
tional Museum 32:465-466. 

Taylor, E. H. 1952. The salamandar and caecilians 
of Costa Rica.—University of Kansas Science 
Bulletin 34:695-791. 


(JRM) 10770 SW 164th Street, Miami, 
Florida 33157, U.S.A.; (L.DW) Department 
of Biology, Miami-Dade Community Col- 
lege, South Campus, Miami, Florida 33176, 
U.S.A.; (KLW) Department of Biology, 
Northwestern State University of Louisi- 
ana, Natchitoches, Louisiana 71497, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(2), 1993, pp. 390-401 


UPPER TRIASSIC REPTILE FOOTPRINTS AND A 
COELACANTH FISH SCALE FROM THE 
CULPEPER BASIN, VIRGINIA 


Robert E. Weems and Peter G. Kimmel 


Abstract.—Three ichnotaxa (Chirotherium, Brachychirotherium, and Ple- 
siornis) are reported for the first time from the Culpeper basin. They occur 
near the base of the Manassas Sandstone and represent the oldest vertebrate 
faunal assemblage from the basin. A fish scale from the overlying Balls Bluff 
Siltstone, referable to Diplurus, is the first report of a coelacanth from Triassic 


strata of the Culpeper basin. 


Palynological studies have established that 
strata in the Culpeper basin (Fig. 1) range 
from Late Triassic to Early Jurassic in age 
(Cornet et al. 1973, Cornet 1977, Litwin et 
al. 1991). Fish remains are abundant locally 
in the Jurassic column of the basin (Schaef- 
fer & McDonald 1978) but are rare and usu- 
ally disarticulated in the Triassic column 
(Gore 1986). Tetrapod remains are found 
more rarely. Parasuchian bones and teeth 
have been reported in the Triassic column 
from River Road in Montgomery County, 
Maryland, near Dulles Airport in Fairfax 
County, Virginia, and from the Culpeper 
Stone Company quarry, Culpeper County, 
Virginia (Weems 1979, Weems & Wiggs 
1991, Weems 1992). Triassic dinosaur foot- 
prints are documented from the Culpeper 
Stone Company quarry, Virginia (Weems 
1987, 1992), and Jurassic dinosaur foot- 
prints have been reported from the region 
near Aldie, Loudoun County, Virginia (Gil- 
more 1924, Roberts 1928, Pannel 1985). 
This paper documents a fifth tetrapod lo- 
cality in the Culpeper basin and the first 
evidence of a Triassic coelacanth fish. 


Locality Data 


The new footprint occurrences are from 
a locality found by Peter Kimmel in July of 
1983 in the Manassas 7.5’-quadrangle along 
Compton Road east of Virginia Route 28 


in Fairfax County (Fig. 2). Located about 
365 m west of the eastern margin of the 
Culpeper basin, this site was excavated to 
construct a brick pumping station. Among 
numerous stone slabs which were un- 
earthed, five contained lightly impressed © 
footprints. Four represent matching part and 
counterpart prints, the fifth is an unmatched 
counterpart print. 

The locality is in the lower Poolesville 
Member of the Manassas Sandstone at a 
horizon less than 50 m above the base of 
the exposed sequence in the Culpeper basin. 
This is the lowest (oldest) horizon in the 
basin that has yielded vertebrate remains. 
Although some workers have assumed the 
Poolesville to be Carnian in age, a definitive 
palynoflora indicates that at least the upper 
part of this unit is early Norian (Litwin et 
al. 1991). The lower Poolesville Member 
remains undated palynologically, but it is 
considered here to be early Norian until 
positive evidence is found for any Carnian 
strata in the basin. 

An impression of a large coelacanth scale 
was found by Juergen Reinhardt and Wayne 
Siglio (U.S. Geological Survey) near the site 
that yielded bones of Rutiodon cf. R. man- 
hattanensis (Weems 1979). The fish scale 
locality lies immediately east of Dulles In- 
ternational Airport in Loudoun County, 
Virginia, near the eastern county border (Fig. 
3). This single scale, the first record of a 


VOLUME 106, NUMBER 2 


N 


Fig. 1. 
locality (1) and fish scale locality (2) of this report are indicated. Detailed locality data are in Figs. 2 and 3. 


sarcopterygian fish in the Triassic column 
of the Culpeper basin, comes from near the 
base of the Balls Bluff Siltstone, which is 
early Norian in age (Lee & Froelich 1989, 
Litwin et al. 1991). 


Footprint Material 


One pair of part and counterpart slabs 
(USNM 412533) contains a lightly im- 
pressed pes impression with a large digit 
located far to the rear and to the side of the 
track (Fig. 4). The position and large size of 
this digit readily characterize this print as 
that of a Chirotherium. The relative pro- 
portions of this track are almost identical 
to those of Chirotherium lulli Bock, but in 
absolute size it is twice as large as the type 
of that species. Because Baird (1954) ob- 
served that the type (and only other de- 
scribed specimen) of C. /u//i was unusually 
small compared to other species of Chi- 
rotherium, we presume that our specimen 
represents an adult of the described Newark 
species rather than a new and larger species 
of nearly identical proportions. Therefore 
we assign our specimen to the described spe- 
cies Chirotherium lulli. Footprints assigned 


VIRGINIA 


Map of Virginia showing area underlain by early Mesozoic rocks of the Culpeper basin. Footprint 


to Chirotherium probably were made by 
pseudosuchian rauisuchids (Charig et al. 
1976). 

A third track-bearing slab (USNM 
412534) contains a counterpart impression 
of a left manus and pes set (Fig. 5). The pes 
is large and shows the impression of three 
forwardly directed blunt-tipped toes. The 
manus is proportionally much smaller than 
the pes and has three slender digits that are 
very nearly the same length. The general 
proportions of these prints are closer to those 
of Grallator, ““Atreipus,”> and Brachychi- 
rotherium than they are to any other de- 
scribed Triassic ichnogenera. Olsen & Baird 
(1986) named “‘Atreipus”’ for Grallator-like 
tracks which also had manus prints. How- 
ever, a recent review of tridactyl theropod 
tracks from the Newark Supergroup (Weems 
1992) has shown that the pes of each of the 
three described species of ““Atreipus”’ has an 
exactly analogous pes in three species of 
Grallator. As the manus of Grallator is (by 
definition) unknown, the distinction be- 
tween “‘Atreipus” and Grallator’ rests on a 
non-character (the absence of manus prints 
in Grallator). ““Atreipus’’ most likely is a 
Grallator walking on four feet instead of 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


392 


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VOLUME 106, NUMBER 2 393 


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Fig. 3. Map showing location of fish scale locality 2 (Loudoun County). Symbols and patterns are the same 
as in Fig. 2. 


two, so the distinction between these two’ was retained. Thus a choice of placement 
genera becomes meaningless taxonomical- for the specimen here in question falls be- 
ly. For this reason, these two genera were tween Grallator and Brachychirotherium. 

synonymized and the earlier name Grallator The absolute size of our print is close to 


— 
Fig. 2. Map showing location of footprint locality 1 (Fairfax County). Geology and border fault located from 


Lee (1979). Stratigraphic column on left shows relative age and sequence for units shown on map. Jurassic 
diabase is a sill that baked the immediately surrounding Triassic rocks. ““T”’-shaped symbols and numbers 
represent strike and dip of sedimentary rocks. 


394 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


0 2 
ek 


cm 


Fig. 4. 


(middle) Outline drawing of right pes of type specimen of Chirotherium lulli (after Baird 1954), mirror- 


reversed to left pes. Overprinted on this pes outline is the best fit outline of the Chirotherium footprint from 
Culpeper basin (stippled areas). Photographs of the left pes imprint of Chirotherium lulli (USNM 412533) from 
Culpeper basin are shown in strong (right) and subdued (left) light. Other linear features in photographs are 


mudcrack boundaries. 


the size of an adult Gral/ator and much 
smaller than an adult Brachychirotherium 
(Fig. 6). But placement of the outline of our 
print on published drawings of a Grallator 
print (Olsen & Baird 1986) and a Brachy- 
chirotherium print (Baird 1954) shows that 
the toe proportions, toe shapes, and appar- 
ent placement of the animal’s weight on the 
ground are what would be expected from 
the foot of Brachychirotherium, specifically 
Brachychirotherium parvum (C. H. Hitch- 
cock). Possibly because the print is lightly 
impressed, no impression was left of digit I 
or of the small nails. The proportions of the 
toes and the placement of weight repre- 
sented by our footprint are not so similar 
to Grallator. Moreover, the very large and 
sharply pointed toes of Gra/lator should have 
left some evidence of their presence even 
though the print is lightly impressed. There- 
fore, we assign this print to Brachychirothe- 
rium. The small size leaves open the pos- 
sibility that our specimen represents a new 
and smaller species of this genus. But our 
specimen easily could have been made by 
a juvenile animal, and fossil footprints can 
display considerable variability (Weems 
1992). Thus for now we are content to assign 
this specimen to Brachychirotherium par- 


vum. Haubold (1971) considered the track- 
maker of Brachychirotherium to be an ae- 
tosaur, but the proportions of the pes track 
also are notably similar to those of the plan- 
tigrade rear foot of the crocodylotarsan Pos- 
tosuchus (Chatterjee 1985). 

The fourth and fifth (part and counter- 
part) slabs (USNM 412535) contain im- 
pressions of digits II] and IV, and a probable 
faint impression of digit II, of a small bi- 
pedal animal (Fig. 7). The proportions and 
relative straightness of the toes suggest that 
the trackmaker was a small (around 0.5 m 
long) functionally tridactyl archosaur. Al- 
though imperfectly preserved, this track is 
fully comparable to the ichnotaxon Plesior- 
nis pilulatus E. Hitchcock (Fig. 8), originally 
described from the Portland Formation 
(Lower Jurassic) of Massachusetts (Lull 
1953). The Culpeper basin occurrence rep- 
resents a significant downward extension of 
its known range. 

Ellenberger (1972) and Lockley et al. 
(1992) have described very bird-like tracks 
(Trisauropodiscus) from rocks of Early Ju- 
rassic age. Lockley et al. (1992) also noted 
that several Early Jurassic Newark taxa show 
strongly bird-like characteristics (Si//imani- 
us tetradactylus, Argoides macrodactylus, A. 


VOLUME 106, NUMBER 2 


cm 


395 


cm 


Fig. 5. (right) Counterpart left manus and pes set (USNM 412534) referable to Brachychirotherium parvum, 
(left) closer photograph of manus print of same specimen of Brachychirotherium. 


minimus, Triaenopus lulli, and T. emmon- 
sil). Similarly, Plesiornis pilulatus (as its ge- 
neric name suggests) is markedly bird-like. 
The prints are small, evidence of footpads 
is often lacking (Lull 1953), the toes are long 
and narrow, and there appears to be at least 
some indication of a rearwardly rotated hal- 
lux in the type. The only characteristic that 
debars Plesiornis from the avian ichnofam- 
ily Trisauropodiscidae is its II-IV digit di- 


varication, which is about 70°. Trisauro- 
podiscidae should have a II-IV digit 
divarication of 90° or greater. 

But even though the IJ-IV digit divari- 
cation of Plesiornis is less than that of typ- 
ical birds, it is greater than that of typical 
Triassic dinosaurs (30° to 50°). Considering 
that this print comes from rocks of early 
Norian age, and that ancestral birds ulti- 
mately should converge back toward a more 


396 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 6. Outline drawings of right pes of Grallator tuberosus (=“‘Atreipus mildfordensis’’) (after Olsen & Baird 
1986) (left) and Brachychirotherium parvum (after Baird 1957) (right) with mirror-reversed outline of print shown 


in Fig. 4 (shaded) superimposed upon them. 


dinosaur-like foot pattern as the two lin- 
eages approach a common ancestor, it 
should not be surprising if footprints of ear- 
ly Norian bird-like animals were rather in- 
termediate in appearance between those of 
typical birds and typical dinosaurs. As os- 
teological remains of a Late Triassic (early 
Norian) bird (Protoavis) recently have been 
described from Texas by Chatterjee (1991), 
this raises the possibility that our small track, 
of nearly identical age and estimated size, 
was made by a protoavid. Therefore we are 
inclined to view Plesiornis pilulatus as a 
primitive bird or near-bird, possibly a pro- 
toavid. 


Fish Material 


Fish remains are locally abundant in the 
Jurassic column of the Culpeper basin 


(Schaeffer & McDonald 1978), but they are 
rare in the Triassic column (Gore 1986). 
Therefore, it is noteworthy that a scale im- 
pression of a large coelacanth (USNM 
421762) (Fig. 9) was found in the lower part 
of the Balls Bluff Siltstone near the locality 
at Dulles Airport that yielded parasuchian 
bones and armor (Weems 1979). Other re- 
ports of coelacanths in the Newark Super- 
group have been from lacustrine beds, but 
the beds in the Dulles Airport area appear 
to be fluvial, rather than lacustrine, in ori- 
gin. This suggests that Diplurus probably 
inhabited river systems as well as lakes dur- 
ing the Late Triassic. 

Two coelacanth taxa are known from the 
Norian and Jurassic portions of the Newark 
Supergroup. Osteopleurus newarki (Bryant) 
occurs in the Triassic part of the Newark 
basin and the Danville basin (Olsen 1988). 


397 


VOLUME 106, NUMBER 2 


Stereophotographs of part (top) and counterpart (bottom) slabs containing left footprint of Plesiornis 


Pig..7. 
pilulatus (USNM 412535). 


398 


Fig. 8. 
basin print (mirror-reversed) superimposed upon it. 


Diplurus longicaudatus Newberry is known 
from the Jurassic part of the Hartford and 
Culpeper basins and from the Triassic and 
Jurassic parts of the Newark basin (Schaef- 
fer 1952, Lull 1953, Schaeffer & McDonald 
1978). Because specimens of Osteopleurus 
newarki are not known to exceed 20 cm in 
total length (Schaeffer 1952:54), their scales 
usually are much smaller than those of Di- 
plurus. Also, Osteopleurus flank scales bear 
only a few widely spaced longitudinal ridges 
(around 8—10), while scales of Diplurus bear 
numerous (20-30) closely spaced longitu- 
dinal ridges. Our specimen has about 25 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


= 1cm 


Outline drawing of a type footprint of Plesiornis pilulatus (after Lull 1953) with outline of Culpeper 


closely spaced longitudinal ridges and a size 
that is comparable only with scales from 
larger specimens of D. longicaudatus. 
Therefore, it probably pertains to Diplurus. 
A species assignment is inadvisable because 
the few fragmentary specimens of Diplurus 
known from the Triassic portion of the 
Newark Supergroup are too incomplete to 
be sure if they pertain to D. longicaudatus 
or to an unnamed antecedent species of this 
genus. 

The occurrence of a large coelacanth in 
fluvial Triassic sediments raises the inter- 
esting possibility that this scale could per- 


VOLUME 106, NUMBER 2 


tain to Chinlea (Schaeffer 1967). Chinlea is 
known so far only from the Chinle and 
Dockum formations of the American 
Southwest (Murry 1986), but the fact that 
it occurs in beds that are comparable in age 
and depositional setting to those of the low- 
er Balls Bluffis suspicious. Chinlea reached 
a size comparable to that of Diplurus 
(Schaeffer 1967), and it also had numerous, 
closely spaced, longitudinal striations on its 
scales (up to 40). Therefore, no obvious size 
or ornamentation characteristics preclude 
the possibility that this genus could be rep- 
resented by this specimen. But because 
Chinlea has never been reported from the 
eastern United States, and because the stri- 
ation count on our specimen is more typical 
of Diplurus than of Chinlea, Diplurus is the 
more logical assignment based on current 
knowledge. 


Discussion 


On the basis of palynological correlations 
(Cornet 1977, Litwin et al. 1991) the Ma- 
nassas Sandstone and Balls Bluff Siltstone 
are correlated with the Upper Triassic Pas- 
saic Formation of the Newark basin. The 
Passaic is known to contain at least 20 kinds 
of vertebrates. Fish remains include Semio- 
notus sp., Synorichthys sp., Diplurus sp., and 
Osteopleurus sp. (Schaeffer 1952, Olsen 
1988). Osteological remains of reptiles in- 
clude a parasuchian (Rutiodon?), the pro- 
colophonid Hypsognathus fenneri, and the 
aetosaur Stegomus arcuatus (Huene 1913, 
Colbert 1960, Baird 1986). Reptilian 
footprint taxa include Apatopus lineatus, 
Grallator parallelus, G. tuberosus, Brachy- 
chirotherium eyermani, B. parvum, Chi- 
rotherium lulli, Coelurosaurichnus sp., 
Gregaripus bairdi (=“‘Genus Incertum”’ of 
Baird 1957), Gwyneddichnium majore 
(probably made by Gwyneddosaurus), G. 
minor, Procolophonichnium sp. (probably 
made by Hypsognathus), Rhynchosauroides 
brunswickii and R. hyperbates (Baird 1957, 
1986; Olsen & Baird 1986; Olsen 1988). 


Fig. 9. 
probably Diplurus (USNM 421762), showing rounded 
outline and numerous, closely spaced longitudinal stri- 
ations. Length about 2 cm. 


Impression of a large coelacanth fish scale, 


The Manassas Sandstone and Balls Bluff 
Siltstone together have yielded 13 kinds of 
vertebrates. Osteological remains from the 
Balls Bluff Siltstone represent a parasuchian 
(Rutiodon?), Diplurus sp., and Semionotus 
sp. (Weems 1979, Olsen 1988). The Balls 
Bluff also has yielded seven kinds of reptile 
footprints: Grallator tuberosus, Grallator 
sillimani, Gregaripus bairdi, Agrestipus hot- 
toni, Kayentapus minor, and Eubrontes sp. 
from the Culpeper Crushed Stone quarry 
(Weems 1987, 1992), and Gwyneddichnium 
mayjore from Manassas National Battlefield 
Park (considered by Olsen as Rhynchosau- 
roides in Gore 1988). The Manassas Sand- 
stone has yielded footprints of Brachychi- 
rotherium parvum, Chirotherium lulli, and 
Plesiornis pilulatus. 

Collectively the Passaic, Manassas, and 
Balls Bluff contain 25 known kinds of ver- 
tebrates. Of these, 8 occur in both basins 
(Semionotus, Diplurus, Rutiodon?, Gwyned- 
dichnium majore, Grallator tuberosus, 
Gregaripus, Brachychirotherium, and Chi- 


400 


rotherium). Because neither the Passaic nor 
the Manassas/Balls Bluff are richly fossil- 
iferous, the high number of apparently en- 
demic forms is not surprising. It is more 
significant that the Passaic and Manassas/ 
Balls Bluff have many more forms in com- 
mon with each other than either does with 
any other interval within the Newark Su- 
pergroup. Thus the correlation of these in- 
tervals in the Culpeper and Newark basins 
is supported by the known vertebrate re- 
mains. 

Three vertebrate taxa (Semionotus, DI- 
plurus, and Grallator), reported from both 
Triassic and Jurassic strata in the Culpeper 
basin, are known elsewhere in the Newark 
Supergroup from both Triassic and Jurassic 
strata. Other ichnotaxa reported from Low- 
er Jurassic strata of the Culpeper basin, Eu- 
brontes giganteus and Eubrontes minuscu- 
lus from the Aldie locality in the Turkey 
Run Formation (Pannel 1985, taxonomi- 
cally updated per Weems 1992), and a Ba- 
trachopus sp. found by Tucker F. Hentz from 
the same area, are known elsewhere in the 
Newark Supergroup only from beds dated 
as Early Jurassic (Olsen 1988). Similarly, 
other fish from the Early Jurassic portion 
of the Culpeper basin (Redfieldius and 
Ptycholepis) are known elsewhere in the 
_ Newark Supergroup only from beds of Early 
Jurassic age (Schaeffer & McDonald 1978). 
Thus a common pattern of faunal succes- 
sion can be documented by vertebrate fos- 
sils in both the Newark basin and the Cul- 
peper basin. This pattern is in accord with 
the palynofloral correlations made between 
the basins by Cornet (1977), and thus sup- 
ports those correlations. 


Acknowledgments 


The authors wish to thank D. Baird, P. 
Dodson, A. J. Froelich, N. Hotton III, J. P. 
Smoot, and two anonymous reviewers for 
their comments, insights and suggestions. 
We also thank D. Bryan Stone III for taking 
the stereophotos of Plesiornis. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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. 1957. Triassic reptile footprint faunules from 

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(REW) Mail Stop 928, U.S. Geological 
Survey, Reston, Virginia 22092, U.S.A; 
(PGK) 9713 Damascus Drive, Manassas, 
Virginia 22110, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(2), 1993, pp. 402-409 


A NEW INTERGENERIC WOOD WARBLER HYBRID 
(PARULA AMERICANA X DENDROICA CORONATA) 
(AVES: FRINGILLIDAE) 


Gary R. Graves 


Abstract.—A new intergeneric wood warbler hybrid (Parula americana xX 
Dendroica coronata) is described from a male in first basic plumage taken 
during fall migration on the Gulf coast of Florida. The hybrid is nearly inter- 
mediate between the parental species in plumage pattern and color, but is more 
similar to its smaller parent, P. americana, in size and shape. 


When Gray’s compendium of avian hy- 
brids was published in 1958, eight hybrid 
combinations (according to the latest tax- 
onomy, (American Ornithologists’ Union 
1983) had been reported among wood war- 
blers (Fringillidae: Parulinae). Since then the 
number of hybrids has more than doubled: 
no fewer than 20 hybrid combinations, nine 
of them intergeneric, are now known (Bled- 
soe 1988, Graves, unpubl.). The purpose of 
this paper is to describe a previously unre- 
ported intergeneric hybrid wood warbler. 

The late Henry M. Stevenson collected 
an unusual hybrid wood warbler on St. 
George’s Island, Franklin County, Florida, 
on 24 October 1970. Stevenson’s penciled 
notation on the specimen label (Tall Tim- 
bers Research Station No. 2881) identified 
it as a hybrid, Parula americana X Den- 
droica coronata. Here I confirm Stevenson’s 
identification and present a diagnosis of the 
hybrid specimen. 


Materials and Methods 


The specimen, sexed as a male, has nar- 
rowly pointed rectrices and dull, weakly 
patterned plumage, indicative of first basic 
plumage in the Parulinae (Pyle et al. 1987). 
I compared it with series of immature males 
in first basic plumage of all North American 
species of wood warblers that breed in the 
United States and a specimen of the hybrid, 
Parula americana < Setophaga ruticilla, in 


the National Museum of Natural History, 
Smithsonian Institution. Measurements of 
wing chord, wing tip length (longest primary 
minus longest secondary), tail length (from 
point of insertion of central rectrices to tip 
of longest rectrix), tarsus length, and bill 
length (from anterior edge of nostril), were 
made with digital calipers to the nearest 0.1 
mm. Color comparisons were made under 
Examolites (Macbeth Corp.). 

Diagnostic assumptions and methods of 
hybrid diagnosis based on plumage color 
and pattern and external morphology follow 
Graves (1990). Although the specimen was 
collected in coastal Florida, I considered all 
migratory species of wood warblers as po- 
tential parents of the hybrid. The analysis 
followed a two-step procedure. First, the 
presumed parental species of the hybrid were 
determined by the comparative analysis of 
plumage pattern and color. This hypothesis 
was then examined with morphometric data. 
Concordance of results are interpreted as 
strong support for the presumed parentage 
of the hybrid (see Graves 1990, Graves & 
Zusi 1990). 

I used principal components analysis 
(PCA) on untransformed variables to re- 
duce dimensionality of data and to facilitate 
the analysis of morphology in two dimen- 
sions. Unrotated principal components were 
extracted from correlation matrices (SYS- 
TAT): 


VOLUME 106, NUMBER 2 


. 
aa 
m 
a 
i 
E 
Fig. 1. 


Dorsal view of Parula americana (left), a presumed P. americana x D. coronata hybrid (Tall Timbers 
Research Station No. 2881), and Dendroica coronata (right). 


Results (contrasting tips of the greater and middle 
wing coverts); (4) spots on two outermost 

Plumage characters.—Prominent pattern pairs of rectrices (rectrix 5 and 6); and (5) 
elements possessed by the hybrid include: dark streaks on the flanks and sides of the 
(1) a semiconcealed coronal patch; (2) short breast. Distinctive color characters of the 
superciliary and subocular spot; (3) wing bars hybrid include: (1) gray dorsal plumage with 


403 


404 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


\) 


a 


Fig. 2. Lateral view of Parula americana (left), hybrid P. americana x D. coronata, and Dendroica coronata 


(right) (see Fig. 1). 


a triangular olive-brown patch on the man- 
tle; (2) pale ventral plumage with a yellow- 
ish wash on the breast; (3) buffy or pale 
chestnut spots on breast and sides of lower 
breast; (4) buffy flanks with blackish shaft 
streaks; (5) pale lower mandible (in dried 


skin); and (6) brownish-black legs (in dried 
skin) (Figs. 1-3). 

The pool of potential parental species can 
be quickly reduced by concentrating on 
characters that the hybrid shares with just 
a few species. Of the many possible color 


VOLUME 106, NUMBER 2 


Fig. 3. 
(right) (see Fig. 1). 


and plumage characters present in the hy- 
brid, only one appears to be synapomorphic 
(shared derived)—the olive mantle which 
contrasts with the neutral gray dorsal plum- 
age. Among the potential parental species 
this character is shared only with Parula 


405 


Ventral view of Parula americana (left), hybrid P. americana x D. coronata, and Dendroica coronata 


americana and the largely sedentary P. pi- 
tiayumi of southern Texas. Phenotypic ex- 
pression of this pattern element has also 
occurred in other hybrids of P. americana 
(P. americana X Setophaga ruticilla, see 
Burleigh 1944; P. americana x Dendroica 


406 


Table 1.—Ranges and means (+one standard de- 
viation) of measurements of fall juvenile male Parula 
americana, Dendroica c. coronata, and the hybrid (Tall 
Timbers Research Station No. 2881). 


P. americana D. coronata 

Character (n = 10) (n= 1 Hybrid 

Wing chord 55.4-62 .0 66.2-74 .4 64.7 
6051 = 24 ID3s2 25 

Wing tip 12.1-16 .9 15.6-19 .8 14.2 
IAG EAS | ay get: [ea 

Tail 39.9-46 .5 51.0-58 .6 44.7 
43.4 + 2.0 33-4 23 

Tarsus 16.3-17 .9 17.3-19 .4 16.3 
164.2265 1822 a= 

Bill 7.2-8. 2 6.5-7. 8 8.0 
127 3203 Tt 204 


dominica, see Haller 1940). Other charac- 
ters of the hybrid that are shared with P. 
americana include whitish superciliary and 
subocular spot, yellowish wash across the 
breast, small buffy or chestnut spots on the 
breast and sides of the lower breast, pale 
unmarked belly and undertail coverts, white 
tail spots (rectrix 5 and 6), well-developed 
wing bars, and pale lower mandible. 

By a process of elimination, the remain- 
ing diagnostic characters of the hybrid, in- 
cluding its streaked buffy flanks, concealed 
pale coronal spot, and dark legs, must have 
been contributed by the other parental spe- 
cies. Among wood warblers in first basic 
plumage, a well-developed white or yellow 
coronal spot, similar to that possessed by 
the hybrid, is present in Dendroica coronata 
and to a lesser extent in D. fusca and D. 
cerulea. Dendroica fusca can discarded as a 
parental choice because the hybrid lacks 
traces of the dark yellow or yellowish-or- 
ange superciliary and throat found in that 
species. Several other wood warblers have 
yellow crowns or concealed coronal spots 
(Vermivora pinus, V. chrysoptera, Dendroi- 
ca virens, and D. occidentalis) or a central 
crown stripe (Mniotilta varia) that could po- 
tentially produce a hybrid with a semicon- 
cealed whitish coronal spot. None of the 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 2.— Factor loadings for the first two principal 
components from analysis of males in first basic plum- 
age of Parula americana, Dendroica c. coronata, and 
the hybrid (see Fig. 4). 


Principal component axes 


Variable I II 
Wing chord 0.96 —0.02 
Wing tip 0.87 0.08 
Tail 0.94 0.02 
Tarsus 0.78 0.51 
Bill ST 0.66 
Percent variance 

explained TES 13.9 


aforementioned species, however, with the 
exception of D. coronata, has dark buffy 
flanks with blackish shaft streaks. The pre- 
dominately white throat and restricted dis- 
tribution of white tail spots in the hybrid 
suggest the eastern subspecies, D. c. coro- 
nata. 

In sum, plumage characters of the hybrid 
can be accounted for by the two most prob- 
able parental species, Parula americana and 
Dendroica coronata (see Appendix). Other 
pairs of species lack the range of pattern 
elements and plumage colors exhibited by 
the hybrid and, barring atavism or some 
unrecognized genetic phenomenon, could 
not have produced the hybrid. 

External morphology.—The hypothesis 
of parentage derived from plumage char- 
acters was tested with an analysis of mor- 
phological size and shape. Because size and 
shape characters are presumably controlled 
by many genes, the mensural dimensions of 
the hybrid are expected to fall within the 
cumulative ranges of parental characters. 
Four of the five measurements of the hybrid 
fall within the range of those for Parula 
americana, but all five are outside the rang- 
es for Dendroica coronata (Table 1). Thus, 
the hybrid is much more nearly the size and 
shape of P. americana, the smaller of the 
two presumed parental species. The bill of 
the hybrid is intermediate in structure be- 
tween the long, rather slender bill of P. 


VOLUME 106, NUMBER 2 


3 


2 - PR. americana 
& 


PCA Il 


407 


D. coronata 


PCA | 


Fig. 4. Bivariate plot of factor scores from a principal components analysis of measurements of Parula 
americana, Dendroica coronata, and their presumed hybrid (filled circle). 


americana and the shorter, wider bill of D. 
coronata. 

The morphological similarity of the hy- 
brid to P. americana is further demonstrat- 
ed by a principal components analysis (Fig. 
4, Table 2). Factor scores of the hybrid fall 
within the envelope of those for P. ameri- 
cana. Under the assumptions used here 
(Graves 1990), had the hybrid’s factor scores 
occurred outside the region of multivariate 
space circumscribed by the combined scores 
of the presumed parental species, the P. 
americana X D. coronata hypothesis could 
have been rejected — provided that the PCA 
axes described a large percentage of the total 
variance and samples of the parental species 
were large. Although this interpretation 
seems to be confirmed by a few case studies 
of avian hybridization (Graves 1988, 1990, 
1992), this method has not been tested with 
large samples of hybrids of known parentage 
(e.g., Vermivora pinus X V. chrysoptera). 

In conclusion, the parentage of the hybrid 
can be attributed, with a high degree of cer- 
tainty, to Parula americana and Dendroica 


c. coronata. The breeding ranges of these 
two species overlap extensively from west- 
ern Minnesota and Ontario east through the 
Great Lakes to the northern Appalachians 
and the maritime provinces of Canada. 


Acknowledgments 


I thank Storrs Olson for bringing the hy- 
brid to my attention, Todd Engstrom of Tall 
Timbers for loaning it, and Richard Banks, 
Ralph Browning, George Hall, and Town 
Peterson for comments on the manuscript. 


Literature Cited 


American Ornithologists’ Union. 1983. Check-list of 
North American birds, 6th ed. A.O.U., Wash- 
ington, D.C., 877 pp. 

Bledsoe, A. H. 1988. A hybrid Oporornis philadel- 
phia x Geothlypis trichas, with comments on 
the taxonomic interpretation and evolutionary 
significance of intergeneric hybridization. — 
Wilson Bulletin 100:1-8. 

Burleigh, T. D. 1944. Description of a new hybrid 
warbler. — Auk 61:291-293. 

Graves,G.R. 1988. Evaluation of Vermivora x Opo- 


408 


rornis hybrid wood-warblers. — Wilson Bulletin 

100:285-289. 

1990. Systematics of the “green-throated 
sunangels”’ (Aves: Trochilidae): valid taxa or hy- 
brids?— Proceedings of the Biological Society of 
Washington 103:6-25. 

1992. Diagnosis of a hybrid antbird (Phle- 
gopsis nigromaculata x Phlegopsis erythrop- 
tera) and the rarity of hybridization among sub- 
oscines. — Proceedings of the Biological Society 
of Washington 105:834-840. 

——, & R. LL. Zusi. 1990. An intergeneric hybrid 
hummingbird (Heliodoxa leadbeateri x Helian- 
gelus amethysticollis) from northern Colom- 
bia.— Condor 92:754-760. 

Gray, A. P. 1958. Bird hybrids. Commonwealth Ag- 
ricultural Bureaux, Bucks, England, 390 pp. 

Haller, K. W. 1940. A new wood warbler from West 
Virginia. — Cardinal 5:49-53. 

Pyle, P.,S. N. G. Howell, R. P. Yunick, & D. F. DeSante. 
1987. Identification guide to North American 
passerines. Slate Creek Press, Bolinas, Califor- 
nia, 278 pp. 


Department of Vertebrate Zoology, Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. 20560, 
U.S.A. 


Appendix 


Comparative descriptions of the hybrid, Parula 
americana x Dendroica c. coronata, and its parental 
species in first basic plumage (males) (see Figs. 1-3). 

The crown, hindneck, mantle, and scapulars of coro- 
nata are dark brown; central crown feathers are sub- 
terminally yellow, forming a semiconcealed coronal 
patch; feathers of the mantle and scapulars have black- 
ish shaft streaks. In americana, the respective parts are 
gray; feathers of the crown, hindneck, and scapulars 
are lightly tipped with olive yellow; the mantle is dark 
olive yellow forming a triangular patch that contrasts 
with the adjacent grayish plumage. Crown feathers of 
americana lack subterminal spots although the basal 
portion of the feathers near the rachi are pale. The 
respective parts of the hybrid are intermediate in ap- 
pearance; feathers of the crown, hindneck, and scap- 
ulars are gray, faintly tipped with brownish-olive; crown 
feathers possess white subterminal spots with scattered 
yellow barbs; the mantle, which contrasts noticeably 
with the adjacent grayish plumage, exhibits a contrast- 
ing patch of pale olive brown plumage with faint darker 
shaft streaks (contrast between the gray hindneck and 
olive mantle of P. americana and the hybrid appears 
indistinct in Fig. 1). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


In coronata, the lower back is brownish gray with 
darker shaft streaks, the rump is pale yellow, and the 
upper tail coverts are black with broad dark gray mar- 
gins. The lower back, rump and upper tail coverts in 
americana are gray, faintly tipped with olive yellow. 
The hybrid is intermediate in appearance; the lower 
back is gray with dark shaft streaks; feathers of the 
rump exhibit olive yellow tipping; the upper tail co- 
verts are black with broad gray margins tinted with 
olive yellow. 

The superciliary region of coronata is slightly paler 
than the dark brown crown; the broken eyering is buffy 
white; the lores and auriculars are dark brown; the neck 
is Slightly paler. In americana, the short superciliary is 
yellowish white anteriorly, turning white over the eye; 
the subocular spot is white (this and the superciliary 
form a broken eyering); the lores, auriculars, and neck 
are gray. The hybrid is intermediate; the superciliary 
and subocular spot are dull white; the lores, auriculars 
and neck are gray (superciliary somewhat obscured in 
Rip.) 

In coronata, the chin and throat are buffy white; the 
breast, sides, and flanks are buffy white to buff with 
dark brown or black shaft streaks; margins of feathers 
at the side of the breast are pale yellow; the belly, vent 
and undertail coverts are white. The chin and throat 
of americana are yellow; feathers of the lower throat 
and pectoral area are dark brown to chestnut brown 
with yellow margins; the pectoral area is bordered pos- 
teriorly by an unmarked yellow band; the sides and 
flanks are pale gray suffused with pinkish buff feathers 
that are occasionally tipped faintly with olive yellow; 
a few buff or chestnut spots occur below the unmarked 
yellow breast band; the belly, vent, and undertail co- 
verts are white, tinted with olive yellow near the vent. 
The venter of the hybrid is somewhat intermediate but 
marked less than either of the parental species. The 
chin and throat are white tinted with pale yellow; the 
breast is very pale yellow, some feathers have buffy or 
pale chestnut subterminal spots; feathers at the sides 
of the breast (near the bend of the wing in the specimen) 
have black shaft streaks; the sides and flanks are buffy 
with dark shaft streaks; the lower breast, abdomen, 
vent, and undertail coverts are white; a few buffy spots 
occur on the sides of the lower breast. 

The remiges and wing coverts are dark grayish brown 
in coronata; greater and middle wing coverts are broad- 
ly tipped with buffy white or buff; outer webs of remiges 
are margined with buff or grayish-brown. In ameri- 
cana, the remiges and wing coverts are gray; greater 
and middle wing coverts are broadly tipped with white; 
remiges have olive-tinted gray margins. The remiges 
and wing coverts of the hybrid are nearly intermediate 
in color and pattern. 

Rectrices of coronata are dark brown with progres- 
sively larger white spots on the inner webs of rectrix 
four, five, and six. In americana, the white area on 


VOLUME 106, NUMBER 2 


rectrix four is limited to a thin stripe along the margin 
of the inner web. The tail pattern of the hybrid is in- 
termediate; rectrix four is similar to that of americana, 
while the two outer pairs of rectrices (5 & 6) are nearly 
identical to those of coronata. 

The bill of coronata is dark brownish black (in dried 


409 


skins). In americana, the bill is yellowish-brown, darker 
near the nostrils and along the culmen. Bill color of 
the hybrid is intermediate. Leg color is blackish-brown 
in coronata and the hybrid and medium brown in 
americana. 


PROC. BIOL. SOC. WASH. 
106(2), 1993, pp. 410-416 


KARYOTYPIC CHARACTERISTICS OF SOREX TUNDRENSIS 


MERRIAM (MAMMALIA: SORICIDAE), A NEARCTIC 
SPECIES OF THE S. ARANEUS-GROUP 


V.R. Rausch and R. L. Rausch 


Abstract. — The karyotype of Sorex tundrensis Merriam, 1900 is redefined on 
the basis of a female specimen from Alaska, and compared with those of taxa 
in Eurasia that have been regarded as conspecific. The diploid number (2N = 
32, FN autosomes = 58) comprised 28 bi-armed autosomes (14 homologous, 
nonpolymorphic pairs) wherein centromeres are median to subterminal, 2 au- 
tosomes (1 pair) with centromeres nearly terminal, and the X-chromosomes 
(the two largest elements in the complement). The diploid numbers of Eurasian 
shrews referred to S. tundrensis have been found (from the literature) to range 
from 31 to 40 in males (with the male trivalent sex-chromosomes typical of 
the S. araneus-group); their karyotypes usually include a larger component of 
acrocentric elements, and the fundamental numbers (autosomes) have been 
calculated to be 52 or 54. The Eurasian taxa appear to make up a complex of 
morphologically similar sibling species. The significant differences between the 
North American and Eurasian taxa in number of major chromosomal arms 
and other characteristics indicate that S. tundrensis is limited geographically 
to the Nearctic. The taxonomic problems involving S. tundrensis and S. arcticus 
Kerr, 1792, both nearctic members of the araneus-group, are briefly reviewed. 


The mammalian fauna of North America 
north of ca. lat. 45°N includes two species 
of shrews placed in the Sorex araneus-group 
(characterized by trivalent sex-chromo- 
somes in the male), Sorex arcticus Kerr, 
1792 and S. tundrensis Merriam, 1900, of 
which the latter has been considered to have 
an holarctic distribution (see Junge et al. 
1983, for review). That these two taxa rep- 
resent independent species was denied for 
some years because of differing opinions 
concerning the significance of macromor- 
phological and morphometric characters. 
Chromosomal comparisons have made clear 
that S. arcticus does not occur in Eurasia. 
Some uncertainty exists concerning the re- 
lationships of taxa designated S. tundrensis 


in the Holarctic, since the karyotype of the 


nominate taxon in northwestern North 
America has not been fully described. 
Herein, we define the chromosomal char- 


acteristics of S. t. tundrensis from Alaska, 
based on one female specimen. The findings 
supplement the observations and conclu- 
sions of Meylan & Hausser (1991), who pre- 
sented the diploid chromosomal comple- 
ment of a male S. tundrensis collected in 
the Yukon Territory. 


Materials and Methods 


A female Sorex tundrensis was captured 
in August 1990 in the Matanuska Valley of 
south-central Alaska (approx. 61°39'N, 
149°12’W). After colchicine and hypotonic 
treatment, cells from marrow and lym- 
phatic tissue were centrifuged, fixed, and 
placed on slides in the field; the slides were 
later stained in the laboratory at the Uni- 
versity of Washington. Procedures applied 
in the preparation of mammalian chro- 
mosomes have been described in detail else- 


VOLUME 106, NUMBER 2 


where (Rausch & Rausch 1975). To produce 
Giemsa-banding, the method of Seabright 
(1972) was used. Chromosomes were count- 
ed and evaluated in more than 50 intact cells 
in metaphase stage; 25 cells were photo- 
graphed, from which, in non-banded com- 
plements, chromosomal measurements were 
made as suggested by Levan et al. (1964). 
In 15 karyograms constructed for compar- 
isons, arm-ratios and size provided the ba- 
sis for assembling pairs of non-banded com- 
plements, and those with G-bands were 
sorted by size and banding-pattern. The 
fundamental number (FN) of major chro- 
mosomal arms was determined following 
the procedure of Matthey (1945) and by di- 
rect measurement of arm-lengths. The skin 
and skeleton of the shrew (orig. No. 47985) 
were prepared by standard methods and de- 
posited in the collection of the Section of 
Mammals, Burke Memorial Washington 
State Museum, University of Washington, 
No. 38109. 

Coordinates (approximate) for localities 
mentioned in the literature cited were es- 
tablished from Atlas SSSR (Glavnoe uprav- 
lenie geodezi i kartografii pri Sovete Minis- 
terov SSSR, Moskva, 1969); in some cases, 
spellings in the published papers differed 
slightly from those of the Atlas. 


Results 


The diploid complement (32) consisted 
of the following: 24 chromosomes with cen- 
tromere median to submedian in location 
(pairs 1-12; range of arm-ratio 1.1 to 2.6); 
6 chromosomes with centromere subme- 
dian to subterminal (pairs 13-15; arm-ratio 
2.7 to 6.1); and 2 chromosomes with cen- 
tromere in the terminal area (pair 16; arm- 
ratio > 10). The findings of Meylan & Haus- 
ser (1991) and other published data (e.g., 
Ivanitskaia & Kozlovskii 1983) concerning 
closely related taxa belonging to the ara- 
neus-group in Eurasia indicate clearly that 
the sex-chromosomes of the female studied 
are the two submetacentric elements of 


411 


greatest size in the complement (first pair 
in the karyograms, Fig. 1A, B). The FN (au- 
tosomes plus sex-chromosomes) was deter- 
mined as 62. We conclude that the shrew 
studied by Meylan & Hausser (1991) is 
karyotypically identical with our specimen 
from Alaska, except for the obvious sex- 
related difference. 


Discussion 


Jackson (1928) defined morphological 
characters that distinguish Sorex arcticus 
and S. tundrensis in North America, but 
later investigators were of the opinion that 
those taxa could be differentiated only at 
the infraspecific level (Rausch 1953, Bee & 
Hall 1956, Hall & Kelson 1959). With 
placement of S. tundrensis in synonymy with 
S. arcticus, the latter would have a contin- 
uous distribution in North America from 
the Atlantic coast in southeastern Canada 
to the northern and western coasts of Alaska 
(as shown by Hall 1981). Youngman (1975) 
reassessed the cranial characters of the two 
taxa and confirmed Jackson’s (1928) con- 
clusion that S. arcticus and S. tundrensis are 
separate species. He also determined that 
the two are allopatric, their geographic rang- 
es separated by a relatively narrow area in 
the western part of the Yukon Territory. A 
disjunct region in southeastern Canada (New 
Brunswick and Nova Scotia) is occupied by 
a taxon designated S. a. maritimensis Smith, 
1939, whose chromosomal characteristics 
suggest that 1t may represent an indepen- 
dent species (Volobouev & van Zyll de Jong 
1988). 

In Eurasia, Stroganov (1936) compared 
S. araneus ultimus G. M. Allen, 1914, de- 
scribed from Nijni Kolymsk (=Nizhneko- 
lymsk) on the lower Kolyma River (68°30'N, 
161°E), and concluded that it was conspe- 
cific with S. tundrensis. He recognized two 
additional subspecies, S. t. petshorae Og- 
nev, 1922 (type locality: the lower reaches 
of the Pechora River, Arkhangel’sk Oblast’, 
lying in northern European Russia, ca. 68°N, 


412 


19 8K wa 


xx ; ax 


pA 


a& 


Ka as 


G 4 8 «aa 
B 


Fists): 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


aa sa 


o & 


Karyotype of Sorex tundrensis, female. 2N = 32. Scale-line has value of 5 um. A. Chromosomes 


consist of 12 pairs of metacentric-submetacentric, 3 pairs subtelocentric, and 1 pair acrocentric. The largest 
elements in the complement (first pair) are the X-chromosomes. Standard Giemsa stain. B. Karyogram with 
chromosomes (arranged as in A) banded by the Giemsa-method. 


54°E) and S. t. middendorfi Ognev, 1933 
(type locality: on the Angara River, Irkutsk 
Oblast’, south-central Siberia, ca. 56°N, 
103°E), and described a third, S. ¢. euro- 
paeus Stroganov, 1936, from Chun Lake on 
the Kola Peninsula (68°N, 36°E). Stroganov 
stated (p. 131) (our translation) that ‘““Here 
it is interesting to note that the Anadyr’ So- 
rex tundrensis ultimus on the basis of sys- 
tematic characters (structure of the skull and 


measurements) stands much closer to the 
American S. t. tundrensis than to any other 
palaearctic forms such as S. t. middendorfi 
and others.” Ognev (1941, cited in Okho- 
tina 1983) reached the same conclusion af- 
ter comparing specimens from the valley of 
the Anadyr’ (ca. 65°N, 171°E) and from 
Alaska. Allen also stated (1914:52), in his 
remarks concerning S. araneus ultimus with 
reference to pelage-color, that ““The same 


VOLUME 106, NUMBER 2 


condition is found in S. tundrensis of north- 
ern Alaska, which is clearly a New World 
derivative of the present species.” 

In accordance with the taxonomic con- 
cept of Hall & Kelson (1959) and others in 
North America, mammalogists in the So- 
viet Union accepted the name arcticus for 
the Eurasian taxa that previously had been 
designated S. tundrensis. Not until the dip- 
loid number of chromosomes and FN for 
S. arcticus had been defined in Canada 
(Meylan 1968) were more definitive com- 
parisons possible. Kozlovskii (1971) dis- 
cussed findings in some Eurasian shrews 
with reference to the karyotype of S. arcti- 
cus, and determined that the palaearctic taxa 
were distinct and apparently represented 
morphologically similar sibling species. On 
the basis of all data, Vorontsov & Liapu- 
nova (1976) concluded that S. arcticus is 
not an holarctic species, and their judgment 
was confirmed by the detailed comparisons 
made by Ivanitskaia & Kozlovskii (1983), 
who pointed out that Sorex tundrensis was 
the applicable name for all of the Eurasian 
taxa that had been erroneously designated 
S. arcticus. Other investigators reached the 
same conclusion on the basis of macro- 
morphological criteria (Junge & Hoffmann 
1981, Okhotina 1983, Junge et al. 1983). 
The allozyme electrophoretic study by 
George (1988) also indicated specific dis- 
tinction. The karyotype of S. arcticus in 
Canada was described in detail by Meylan 
& Hausser (1973). 

Shrews referred to S. tundrensis occur 
widely in Eurasia: northeastern Europe; 
northern and middle Asia, including the 
northern part of Mongolia; northeastern 
regions of China; and the northern part of 
Korea (Okhotina 1984). Information con- 
cerning the distribution of nominal subspe- 
cies and synonymies has been provided by 
Gureev (1981), Junge et al. (1983), and Pav- 
linov & Rossolimo (1987). Okhotina (1984) 
recognized four subspecies of S. tundrensis 
in the Far East of the former Soviet Union: 
S. t. tundrensis Merriam, 1900 (=S. arcticus 


413 


borealis Kashchenko, 1905 = S. a. buxtoni 
J. A. Allen, 1903); S. t. baikalensis Ognev, 
1913 (=S. arcticus baikalensis Ognev, 1913); 
S. t. stroganovi Okhotina, 1983 (=S. arcti- 
cus ssp. nov. Stroganov, 1957); and S. t. 
parvicaudatus Okhotina, 1976 (=S. arcticus 
parvicaudatus Okhotina, 1976). According 
to Okhotina’s concept, the distribution of 
the nominate subspecies of S. tundrensis in 
Eurasia would include northeastern Siberia 
from Chukotka southward to the western 
shore of the Amur River (Primorsk region). 

Chromosomal characteristics have been 
defined for shrews designated S. tundrensis 
from several of the Eurasian localities: vi- 
cinity of Uskovo, Novokuznetsk region 
(53°40’N, 87°E), Kemerovsk Oblast’, and 
vicinity of Irkutsk (52°15’N, 104°E) (Koz- 
lovskii 1971); Boguchan, on the Chun River 
(57°40'N, 96°E), Krasnoiarsk Krai, and 
Bakchar (57°6'N, 82°E), Tomsk Oblast’ (Fe- 
dyk & Ivanitskaia 1972); Novosibirsk 
(55°4'N, 83°E) (Kral & Radjabli 1976); vi- 
cinity of Main (53°N, 91°30’E), Krasnoiarsk 
Krai (Aniskin & Volobuev 1980); vicinity 
of Razdol’noe (43°34'N, 132°E), Primorsk 
Krai; shore of Chaunsk Gulf (68°45’N, 
170°E) and vicinity of Stokovyi Ten’kinsk, 
Magadansk Oblast’; Katon-Karagai 
(49°10'N, 85°30’E), East Kazakh Oblast’: 
and Moneron Island (at the southern end of 
Sakhalin Island), Sakhalinsk Oblast’ (Iva- 
nitskaia & Kozlovskii 1983); two localities 
in the Seleginsk Aimak in northern Mon- 
golia (ca. 5O°N, south of Lake Baikal) (Iva- 
nitskaia & Malygin 1985); and Zveringo- 
lovskoe (54°30'N, 64°45’E), Kurgansk 
Oblast’, and vicinity of Abakan (53°38’'N, 
91°30’E), Krasnoiarsk Krai (Ivanitskaia et 
al. 1986). Ivanitskaia & Kozlovskii (1983, 
table 1) summarized the data on diploid 
number, fundamental number, and mor- 
phological characteristics of autosomes for 
shrews from the aforementioned localities, 
with exception of those reported by Ivan- 
itskaia & Malygin (1985) and Ivanitskaia et 
al. (1986). 

The diploid number of chromosomes in 


414 


males of the Eurasian shrews referred to So- 
rex tundrensis has been found to range from 
31 to 40, with fundamental numbers (au- 
tosomes) of 52 or 54. Because of the tri- 
valent sex-chromosome in males, diploid 
numbers in females were typically one less. 
In some Eurasian taxa, polymorphism has 
been observed in chromosome-pairs 1, 2, 
4, and 6 (see Ivanitskaia & Kozlovsku 1983, 
table 1). Aniskin & Volobuev (1980) and 
Ivanitskaia et al. (1986) suggested that such 
variation in the designated homologues was 
due to Robertsonian translocations. 

While the diploid number of chromo- 
somes of S. ¢t. tundrensis from the Yukon 
Territory and Alaska is the same as that of 
animals from some populations in the 
Palaearctic, e.g., Novosibirsk (middle Asia) 
and Magadansk Oblast’ (northeasternmost 
Siberia), the morphological differences in the 
chromosomal complements between Eur- 
asia and North America are well defined. 
The autosomal complement of the Alaskan 
female (Fig. 1) consisted of 14 pairs of bi- 
armed chromosomes ranging from meta- 
centric to subtelocentric, and a single pair 
of acrocentrics, whereas complements of the 
Siberian specimens may have a larger com- 
ponent of acrocentric elements and/or a dis- 
parate diploid number (e.g., those from Mo- 
neron Island and Primorsk Krai). The 
fundamental number, established on the ba- 
sis of arm-ratios, of S. tundrensis from Alas- 
ka is thus greater (58, as compared with 52 
or 54 in the Eurasian shrews). For the male 
specimen from the Yukon Territory, Mey- 
lan & Hausser (1991) obtained an FN of 54, 
but their preparations (Meylan & Hausser, 
fig. 1) probably did not permit accurate 
measurement of all elements of the com- 
plement, and they designated three pairs as 
acrocentric. The value of the FN as deter- 
mined by them would explain their conclu- 
sion that taxa referred to S. tundrensis in 
Eurasia must be conspecific with that in 
North America. With relation to our find- 
ings, should future studies demonstrate that 
an additional autosomal pair (chromo- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


somes with arm-ratio of about 6) is better 
classified as acrocentric, the FN of the North 
American taxon would be reduced to 56, 
still greater than any recorded in Eurasian 
animals. The difference in total number of 
chromosomal arms indicates a significant 
degree of evolutionary distance between the 
North American and Eurasian taxa, and in 
combination with morphological dissimi- 
larities existing between the respective 
karyotypes, permits the conclusion that So- 
rex tundrensis is limited in occurrence to 
the Nearctic. The taxa in Eurasia that pre- 
viously have been referred to S. tundrensis 
appear to make up a complex of sibling spe- 
cies, as has been suggested by Kozlovskii 
(1971) and others, among which karyotypic 
differences seem smaller than those between 
them and S. tundrensis. Further compari- 
sons using Giemsa-banding and other 
methods will be required to determine ho- 
mologues of chromosomes in the different 
populations, including S. tundrensis in 
North America. 

Rand (1954) perceived that Sorex arcti- 
cus and S. tundrensis evidently have been 
derived from two separate Pleistocene dis- 
persals of shrews of the araneus-group into 
North America via Beringia. The present 
geographic range of S. arcticus, the earlier 
migrant, is a consequence of its northward 
expansion during post-glacial time, whereas 
that of S. tundrensis is still approximately 
within the North American portion of the 
former Beringian Refugium. 

We undertook field-work at various lo- 
calities in south-central and eastern Alaska 
during 1988-1992. Shrews were numerous 
in south-central Alaska during August 1989— 
1990, but according to our findings, popu- 
lations consisted mostly of S. cinereus Kerr 
and S. monticolus Merriam; we obtained 
only the single specimen of S. tundrensis. 
Shrews decreased in numbers after 1990, 
and in 1992 were at the lowest density ob- 
served. A similar pattern of change in num- 
bers was seen in eastern Alaska where, with 
the exception of a single specimen of S. hoyi 


VOLUME 106, NUMBER 2 


(Baird), the same two species made up our 
collections. Factors that influence changes 
in numerical densities of shrews are not un- 
derstood, but interspecific competition in 
probably marginal habitat at the southern 
limits of the range of S. tundrensis was per- 
haps significant. 


Acknowledgments 


We express our sincere gratitude to Mr. 
L. Alan LeMaster and Ms. Shirley Le- 
Master, Gakona, Alaska, whose generous 
provision of space for laboratory-proce- 
dures contributed significantly to our work. 


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Washington State Museum, DB-10, and 
Department of Comparative Medicine, SB- 
42, School of Medicine, University of 
Washington, Seattle, Washington 98195, 
USS.A. 


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INFORMATION FOR CONTRIBUTORS 


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CONTENTS 


Revised classification and phylogenetic hypothesis for the Acanthostominae Looss, 1899 (Di- 
genea: Opisthorchiformes: Cryptogonimidae) Daniel R. Brooks and Barbara Holceman 
A new Mericella (Mollusca: Gastropoda: Cancellariidae) from northeastern Africa 
Richard E. Petit and M. G. Harasewych 
New species of Alvinellidae (Polychaeta) from the North Fiji back-arc Basin hydrothermal 
vents (southwestern Pacific) Daniel Desbruyéres and Lucien Laubier 
Gyptis crypta, a new hesionid species from the U.S.A. east coast, with a redescription of G. 
vittata Webster & Benedict, 1887 (Annelida: Polychaeta) Fredrik Pleiel 
A new species of Uniporodrilus (Oligochaeta: Tubificidae) from the Gulf of Mexico coast of 
Florida, and a phylogenetic analysis of the genus 
Christer Erséus and Michael R. Milligan 
A checklist of and illustrated key to the genera and species of the Central and North American 


Cambarincolidae (Clitellata: Branchiobdellida) Perry C. Holt and Brent D. Opell 
Branchinecta sandiegonensis, a new species of fairy shrimp (Crustacea: Anostraca) from western 
North America Michael Fugate 


The first finding of the male of Thaumatocypris echinata Miller, 1906 (Crustacea: Ostracoda) 
J. A. Rudjakov 


Pseudonicothoe branchialis (Crustacea: Copepoda: Siphonostomatoida: Nicothoidae), living on’ - 


the pandalid shrimp Heterocarpus sibogae off northwestern Australia 
Arthur G. Humes and Geoffrey A. Boxshall 
Boreomysis oparva, a new possum shrimp (Crustacea: Mysidacea) from an eastern tropical 


Pacific seamount Jennifer Saltzman and Thomas E. Bowman 
The identity of Talitroides alluaudi (Chevreux) (Crustacea: Amphipoda: Talitridae) with notes 
on a new locality Hiroshi Morino and Reuven Ortal 


A description of a new species of Macrobrachium from Peru, and distributional records for 
Macrobrachium brasiliense (Heller) (Crustacea: Decapoda: Palaemonidae) 
Guido A. Pereira S. 
A new species of crayfish (Decapoda: Cambaridae) belonging to the genus Cambarus, subgenus 
Hiaticambarus, from the upper Elk River drainage of West Virginia 
Raymond F. Jezerinac and G. Whitney Stocker 
Two new species of Gliricola (Phthiraptera: Gyropidae) from the spiny tree rat, Mesomys 
hispidus, in Peru Roger D. Price and Robert M. Timm 
Spadella japonica, a new coastal benthic chaetognath from Japan Jean-Paul Casanova 
Caudina intermedia, anew species of sea cucumber from the South China Sea (Echinodermata: 
Holothuroidea: Molpadiida) Yulin Liao and David L. Pawson 
Hyporhamphus meeki, a new species of halfbeak (Teleostei: Hemiramphidae) from the Atlantic 
and Gulf coasts of the United States Heidi M. Banford and Bruce B. Collette 
A new species of Oedipina (Amphibia: Caudata: Plethodontidae) from northern Honduras 
James R. McCranie, Larry David Wilson, and Kenneth L. Williams 
Upper Triassic reptile footprints and a coelacanth fish scale from the Culpeper Basin, Virginia 
Robert E. Weems and Peter G. Kimmel 
A new intergeneric wood warbler hybrid (Parula americana x Dendroica coronata) (Aves: 
Fringillidae) Gary R. Graves 
Karyotypic characteristics of Sorex tundrensis Merriam (Mammalia: Soricidae), a Nearctic 
species of the S. araneus-group V.R. Rausch and R. L. Rausch 


207 
221 
225 


237 


243 
201 
296 


305 


315 
S25 


332 
339 


346 


353 
359 


366 
369 
385 
390 
402 


410 


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PROC. BIOL. SOC. WASH. 
/ 106(3), 1993, pp. 417-428 


A NEW SPECIES OF A-GIANT:, THOMASOMYS 
(MAMMALIA: MURIDAE: SIGMODONTINAE) FROM 
THE ANDES OF NORTHCENTRAL PERU 


Mariella Leo L. and Alfred L. Gardner 


Abstract.—We describe Thomasomys apeco, new species, from the upper 
montane forest zone of the eastern slope of the Andes of northcentral Pert 
(department of San Martin, Rio Abiseo National Park). This impressive mam- 
mal is the largest known living thomasomyine rodent, exceeded in size of skull 
and dentition only by the extinct Megaoryzomys curioi from the Galapagos 


Islands, Ecuador. 


Resumen. —Describimos Thomasomys apeco, nueva especie, de la vertiente 
oriental de los andes al nordeste del Peru (departamento de San Martin, Parque 
Nacional Rio Abiseo). Este llamativo animal es el mas grande de los thoma- 
sominos vivientes, siendo superado en tamano de craneo y denticion solo por 
Megaoryzomys curioi de las Islas Galapagos, Ecuador. 


In 1987, the Peruvian Association for the 
Conservation of Nature (APECO), with the 
financial assistance of the David and Lucile 
Packard Foundation, began a four-year fau- 
nal survey and inventory of the Rio Abiseo 
National Park located in western departa- 
mento San Martin, Pert (Fig. 1). The park, 
centered at approximately 07°45’S, 77°15'W, 
covers 2745 square kilometers encompass- 
ing the major part of the Rio Abiseo wa- 
tershed on the eastern (Amazonian) slope 
of the Andes and drains into the Rio Hualla- 
ga. The elevational range is from near 1000 
to more than 4000 m and includes seven 
identified life zones according to the Hold- 
ridge Life Zone classification (Tosi 1960) as 
modified by Young & Leon (1988). Surveys 
during the first year of study were at higher 
elevations (from about 3000 to 3600 m) in 
Tropical Alpine Zone and Tropical Mon- 
tane Rain Forest (Paramo Pluvial Alpino 
and Bosque Pluvial Montano Tropical, re- 
spectively; Tosi 1960). These wet paramo 
and elfin forest habitats produced a variety 
of small mammals including a previously 
undescribed giant thomasomyine rodent 
that may be known as: 


Thomasomys apeco, new species 


Holotype.— Adult male, Museo de His- 
toria Natural of the Universidad Nacional 
Mayor de San Marcos (MUSM) 7197, from 
Valle de Los Chochos, ca. 25 km NE Pataz, 
3280 m, Parque Nacional Rio Abiseo, San 
Martin, Peru. Collected by Mariella Leo L. 
on 27 Jul 1987, original number MLL 055. 
The holotype is a well-made skin with skull 
and mandibles (latter separated at symphy- 
sis) in good condition, except that tip of 
right paroccipital process is missing (bro- 
ken). 

Paratypes.— Eight paratypes (skins with 
skulls and mandibles): MUSM 7199 female 
from the type locality; MUSM 7196 female 
(with body in fluid), MUSM 7203 female, 
MUSM 7204 female, MUSM 7202 female, 
MUSM 7201 male, and MUSM 7198 fe- 
male from Pampa del Cuy, ca. 24 km NE 
of Pataz, 3260-3380 m; and MUSM 7200 
male from Puerta del Monte, ca. 26 km (at 
60°) from Pataz, 3250 m. 

Distribution. —Known only from the vi- 
cinity of the type locality and the neigh- 
boring Pampa del Cuy Valley to the south. 


418 


PARQUE NACIONAL; 


os 


CHAGUAL ® 
ee 


Se 


Fig. 1. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


S. 
a 


ms” 
lag 


J 
Hua 


-. ood 


ee 


.wsRio 


Departmental Boundary 
Park Boundary 

Major Rivers 

Nearby Communities 
Survey Area 


0 10 Km 


Map of Rio Abiseo National Park showing location of area where inventories were conducted. The 


western boundary of park corresponds to border between departments of La Libertad and San Martin. 


Puerta del Monte is in the lower Pampa del 
Cuy Valley. 

Etymology.—The name is based on the 
acronym APECO (Asociacion Peruana para 
la Conservacion de la Naturaleza), the Pe- 
ruvian Association for the Conservation of 
Nature responsible for conducting the fau- 
nal inventory of the Rio Abiseo National 
Park. The species epithet apeco 1s treated as 
a noun in apposition. The singular genitive 
endings -/ or -ae are not appropriate because 
these endings are for modern personal 
names. 

Diagnosis. —Thomasomys apeco, endem- 
ic to the northeastern Andes of Peru, is the 
largest living thomasomyine and, along with 
Kunsia tomentosus (Lichtenstein, 1830), 
Nectomys squamipes (Brants, 1827), and the 
extinct Megaoryzomys curioi (Niethammer, 


1964) and Megalomys spp., is among the 
largest Sigmodontinae known (Gylden- 
stolpe 1932, Steadman & Ray 1982, Nowak 
1991). The combination of large size, long 
black and white tail, and long unwebbed 
hind feet distinguish this species from other 
sympatric Sigmodontinae and related tho- 
masomyines. Dental features that distin- 
guish this species from other large Tho- 
masomys include the especially well- 
developed anteroloph and parastyle on M1 
(dental terminology follows Reig 1977; also 
see Carleton & Musser 1989), connection of 
anteroloph to anterior mure independent of 
anterolabial conule and its connecting loph, 
well-developed posteroloph of M1 and M2 
whose labial extension (beyond metaloph) 
persists in well-worn teeth, decidedly an- 
terior-posterior orientation of metalophids 


VOLUME 106, NUMBER 3 


419 


Table 1.—Measurements of Thomasomys apeco new species, from Parque Nacional Rio Abiseo, San Martin, 
Pera. Linear measurements are in millimeters and mass in grams; values are the mean followed by range in 


parentheses. Age-class categories defined in text. 


Age 1 Age 2 Age 3 Age 4 
Measurement (n= 1, 4) (n= 6,42 + 28) (n = 1, 2) (n = 1, 2) 
Length 
Total 469 504.3 560 558 
(469-548) 
Tail 299 303.3 324 320 
(279-329) 
Hind foot (cu) 56 53.8 54 35 
(50-59) 
Ear (from notch) 31 28.7 31 31 
(27-31) 
Skull 45.2 48.1 50.5 51.0 
(44.3-50.9) 
Condylobasal 42.6 45.1 47.7 47.8 
(41.9-47.7) 
Nasal 16.3 17.8 18.9 18.7 
(16.3-19.1) 
Palatilar 20.1 223 22.6 
Post palatal 15.5 18.3 18.7 
Incisive foramina 9.6 10.6 rts 11.4 
(9.4—11.7) 
Maxillary toothrow (alveolar) 10.1 9.8 10.2 10.0 
(9.4—10.1) 
Breadth 
Zygomatic 24.8 255 240 oT 
(23.6—27.4) 
Mastoidal broken 17.8 18.6 18.2 
(17.2-18.2) 
Postorbital a3 5:2 5.0 32 
(4.9-5.4) 
Palatal (postdental constriction) 7.9 7.6 6.9 
Rostral 8.8 8.7 9.9 9.7 
(7.8-9.8) 
Mass 190 225.5 258 300 
(164-335) 


and entolophids (and correspondingly long 
meso- and posteroflexids) in the first two 
lower molars (a feature shared with TJ. au- 
reus [Tomes, 1860]), crescent-shaped pos- 
terolophid on m1 extending to lingual mar- 
gin of tooth, and m3 similar in form to m2 
because of large, well-developed entoconid. 
Also distinctive is the thick, well-developed 
jugal and the relatively deep notch between 


the lacrimal and the zygomatic ramus of the 
maxilla. 

Measurements of the holotype.—Linear 
measurements are in millimeters and mass 
(weight), in grams: Total length, 498.0; tail, 
300.0; hind foot, 50.0; ear, 27.0; greatest 
length of skull, 50.2; condyloincisive length, 
45.9; palatilar length, 22.4: postpalatal 
length, 17.8; incisive foramen length, 11.7; 


420 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 2.—Measurements of Thomasomys apeco new species, Thomasomys aureus complex, and Megaory- 
zomys curioi. Age classes of T. apeco and T. aureus complex are combined. Measurements of Megaoryzomys 
curioi from Steadman & Ray (1982:6). Linear measurements are in millimeters and mass in grams; values are 
the mean followed by range (in parentheses) and sample size. See ““Additional Specimens Examined” for sources 
of 7. aureus. 


Measurements T. “aureus” T. apeco M. curioi 
Length 
Total 372.4 514.3 
(335-421) 10 (469-560) 9 
Tail 218.6 307.0 
(199-248) 10 (279-329) 9 
Hind foot (cu) 36.7 54.0 
(33-41) 10 (50-59) 9 
Ear (from notch) 232 29.4 
(21-24) 10 (27-31) 9 
Skull 38.2 48.3 55:6 
(34.6-41.8) 12 (44.3-51.0) 9 
Condylobasal Seo 45.4 54.5 
(31.9-40.1) 13 (41.9-47.8) 9 (53.3-55.6) 2 
Condyloincisive 339 
(30.7-38.6) 13 
Palatilar 16.0 233 
(13.9-18.4) 13 (24.6-26.5) 3 
Postpalatal 14.0 
(12.5-16.5) 13 
Incisive foramina 8.1 10.6 10.8 
(7.1-9.1) 13 (9.4-11.7)9 (9.6—11.7) 5 
Nasal 14.4 17.9 
(12.4—-15.8) 12 (16.3-19.1) 9 
Rostrum 14.2 
(12.4—-16.0) 12 
Maxillary toothrow Tes 10:9 
(6.8—7.8) 13 (10.0-11.5) 5 
Maxillary toothrow (alveolar) TES 9.9 Lid 
(7.0-8.1) 13 (9.4—10.2) 9 (10.9-12.3) 13 
Mandibular toothrow 7.5 120 
(6.8-8.0) 13 (11.5-12.7) 9 
Breadth 
Zygomatic 20.1 25.8 34.3 
(18.6-21.4) 13 (23.6—27.4) 9 (33.8-34.8) 2 
Braincase 14.8 19.0 
(13.9-15.5) 13 (18.7-19.2) 3 
Mastoidal 14.8 17.9 
(13.9-15.9) 12 (17.2-18.6) 8 
Postorbital 4.9 Sey 8.2 
(4.3-5.3) 13 (4.9-5.4) 9 (7.3-8.6) 4 
across molars (M2—M2) ile 
(7.3-8.5) 13 
Palate (postdental constriction) ~ 6.0 
(5.4-6.8) 13 
Rostral 6.8 8.9 | i) 
(6.1—7.2) 13 (7.8-9.9) 9 (11.0-12.3) 3 
Zygomatic plate 355 | 8.1 
(2.5-3.7) 13 (6.9-8.7) 6 


VOLUME 106, NUMBER 3 


Table 2.—Continued. 


421 


Measurements T. “aureus” T. apeco M. curioi 
Depth 
Braincase 1:2 
(10.2-11.9) 13 
Mass 91.8 23520 
(58-136) 9 (164-335) 9 


nasal length, 19.4; zygomatic breadth, 25.7; 
braincase breadth (above zygoma), 25.7; 
mastoidal breadth, 18.1; interorbital con- 
striction, 5.4; breadth across molars (M2- 
M2), 9.9; palatal breadth (post dental), 7.1; 
rostral breadth, 8.8; rostral length, 19.3; zy- 
gomatic plate breadth, 4.3; mesopterygoid 
fossa breadth, 3.5; maxillary tooth row 
length, 9.5; alveolar maxillary tooth row 
length, 9.7; mandibular tooth row length, 
9.9; alveolar mandibular tooth row length, 
9.5; mandibular length; 28.2; mass, 249.0. 
Additional measurements. —See Tables 1 
and 2 for additional measurements. 
Description. —Dorsal fur long (under fur 
as long as 22 mm middorsally with guard 
hairs reaching 30 mm) and broadly dark 
gray based (between Slate-Gray and Slate 
Color; capitalized color terms from Ridg- 
way 1912), slightly darker subterminally, 
and broadly tipped with paler. Guard hairs 
black, conspicuous, and imparting a coarse 
quality to the fur. Dorsum bright with color 
varying between individuals from Raw Si- 
enna to Ochraceous-Tawny and Tawny Ol- 
ive streaked with black (guard hairs). Color 
darkest middorsally and becoming gradu- 
ally clearer (less streaked with black) over 
shoulders, sides, and flanks where color 
grades into Cinnamon or Cinnamon-Buff, 
becoming Ochraceous-Buff on venter. One 
age-class-1 male (MUSM 7200) with short- 
er and harsher pelage from nape to middle 
of back that may be remnant of juvenile 
pelage; otherwise colored as in adults. Ven- 
tral coloration brightest along sides where 
it may encroach toward ventral midline, and 


on band across chest and along thoracic 
midline. Extent of Ochraceous-Buff color- 
ation on venter variable among individuals, 
most extensive on MUSM 7199 where 
nearly entire ventral surface is uniformly 
colored; somewhat less extensive on MUSM 
7200 where coloration is strongest on chest 
(both as a pectoral band and along thoracic 
midline); and least extensive on MUSM 
7196 where coloration is brightest only on 
middle of chest (thoracic midline) and in 
perianal region. The main color differences 
between specimens seem related to the ex- 
tent of yellow pigment in hair tips. The nar- 
rower the terminal color band, the paler the 
color (approaches Pale Buff) and the more 
conspicuous the basal gray color of the fur. 
Fur on throat dark gray based, tipped with 
buff to whitish, and usually paler than re- 
mainder of venter. Fur on chin (below lower 
incisors) white to base. Eye ring somewhat 
darker than remainder of head, which oth- 
erwise is colored like the dorsum. Ears 
clothed inside and out with short dark brown 
to blackish hair and lack evidence of a pale 
rim. Although fur behind and below ears 
clearer (less black-lined) than remainder of 
head, post or subauricular spots character- 
istic of many thomasomyines absent. An- 
kles completely encircled with blackish hair 
(near Fuscous Black), with the same color 
extending over outer upper surface of hind 
feet and base of toes to claws. Claws, digital 
bristles, and inner (and sometimes outer) 
dorsal surface of metatarsals conspicuously 
paler than remainder of hind foot. Outer 
edge of wrist and metacarpals whitish with 


422 


a few long white tactile hairs; otherwise, 
forefeet dark with paler claws and digital 
bristles. Tail longer than head and body (57 
to 60% of total length) and weakly penicil- 
late. Color on basal two-thirds to three-fifths 
of tail dark brown to blackish with terminal 
one-third to two-fifths white. Vibrissae long 
(some exceeding 70 mm), a few either uni- 
formly pale or dark brown throughout, but 
most dark brown with pale tips. Hind feet 
long; plantar surface naked to ankle. Hallux 
shortest with base of claw well short of first 
phalanx on digit IJ. Middle three toes of 
hind foot nearly of equal length with digit 
III only slightly longer than digits II and IV; 
tip of claw of digit V extends to base of claw 
of digit IV. 

Skull large and robust with flaring zygo- 
matic arches and narrow postorbital con- 
striction (Fig. 2). Nasals long, subparallel 
above rostral capsules, and terminate at 
nearly same level as posterior extensions of 
premaxillae. Margins of postorbital con- 
striction elevated (but not beaded) and de- 
fine the pronounced depression (groove) on 
frontals along midline of skull. Dorsal pro- 
file low and sloping forward along nasals 
over rostrum. Squamosal roots of zygoma 
massive and flaring with posterolateral ridge 
converging toward lambdoidal crests in a 
nearly straight line giving braincase a 
V-shaped lateral profile when viewed from 
above. Auditory bullae relatively large, al- 
though not extending below plane of pter- 
ygoid processes. Mastoid not fenestrate. 
Parapterygoid fossae equal to or narrower 
than mesopterygoid fossa. Alisphenoid strut 
present, although its development variable 
(missing on one side in one individual). 
Sphenofrontal foramen and associated 
squamosal-alisphenoid groove (see Carle- 
ton & Musser 1989:fig. 20) on lateral floor 
of braincase absent. 

Upper incisors orthodont; upper and low- 
er incisors normal for genus. Upper molars 
robust, always longer (anteriorly-posteri- 
orly) than wide, and forming a graded series 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


(Fig. 3). First upper molar with well-devel- 
oped, comparatively broad anteriomedial 
flexus, which opens anteriorly onto a shal- 
low basin rimmed anteriorly by a low ridge 
bearing a variable number of accessory cus- 
pules and lophules (dental terminology fol- 
lows Reig 1977). Protoloph short and weak- 
ly developed or absent. Anteroloph and 
anterostyle well developed with the latter 
expanded anteriorly-posteriorly at margin 
of tooth. Anteroloph, anterior mure, and 
lophs connecting anterolingual and anter- 
olabial conules to anterior mure appear to 
arise from a common juncture in unworn 
teeth (anteromedian flexus, anteroflexus, 
protoflexus, and anteromedial bend of par- 
aflexus converging), although anterolingual 
loph becomes more isolated in worn den- 
tition. Posterior end of anteromedial flexus 
(anterior internal fold of Hershkovitz 1944) 
isolated as a small, shallow enamel island 
in some individuals. Anteroloph well de- 
veloped and continuous from anterior mure 
to labial margin in M2 and M3. A low an- 
terior cingulum forms anterior margin of 
protoflexus in M2 and M3. Paraloph in all 
three upper molars joins median mure be- 
tween junctures of protoloph and mesoloph. 
A paralophule also extending from para- 
cone to mesoloph (connection may be in- 
complete in M3), resulting in isolation of 
median portion of mesoflexus as an enamel 
island. Well-developed mesoloph extending 
to labial margin in all three molars. Meta- 
loph joining posteroloph at approximately 
mid-length in M1 and M2 in unworn den- 
tition (connection appears closer to labial 
margin in worn teeth because of attrition of 
posteroloph). The broad hypoflexus may 
contain low accessory cuspules on lingual 
margin. Enteroloph either absent or, at best, 
visible in unworn teeth only as a lateral 
swelling from the median mure in M1 and 
M2. The small metacone on M3, although 
visible in unworn teeth, becoming more dif- 
ficult to locate as tooth wears. 

Lower molars also form a graded series; 


VOLUME 106, NUMBER 3 423 


Fig. 2. Dorsal, ventral, and lateral views of cranium, and lateral view of mandible of holotype of Thomasomys 
apeco, new species, MUSM 7197 (original number MLL 055). Vertical bar equals 10 mm. 


424 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


% 
~ 
at 
‘ae 
» 
* 
red 
Ss 
a 
$ 


Fig. 3. Left maxillary and mandibular toothrows of Thomasomys apeco, new species, and T. aureus (Tomes, 
1860): T. apeco, MUSM 7198, a maxillary and b mandibular; T. aureus, MLL 312, c maxillary and d mandibular. 


Vertical bar equals 5 mm. 


VOLUME 106, NUMBER 3 


however, m3 is similar to m2 in length and 
development of the entoconid, although 
narrower posteriorly. The prominent and 
deep anteromedial flexid separates high, but 
relatively narrow, lingual and labial conu- 
lids (each about one-half size of metaconid). 
The short and weakly-developed protolo- 
phid inconspicuous; in its place, the anter- 
olabial margin of m1 enclosed by a prom- 
inent anterolabial cingulum extending 
posteriorly from outer surface of anterola- 
bial conulid to protoconid base. Anterola- 
bial cingulum in m2 and m3 fused to pro- 
toconid and, with moderate tooth wear, 
enclosing protoflexid as a comma-shaped 
island. Metalophid continuous with lateral 
extension of anterolophid in all three lower 
molars, reflected by long mesoflexid sepa- 
rating metacone and median mure. The nar- 
row anteroflexid extends to anterior mure 
(evident in all but well-worn teeth). The 
shallow metaflexid partly confluent with an- 
teroflexid. Mesostylid fused to posterolat- 
eral margin of metaconid in each molar and, 
with relatively little wear, connection iso- 
lates mesoflexid as internal island. Proto- 
lophid narrower than protoconid and di- 
rected slightly anteriorly as it joins median 
mure. In m1, both entolophid and entolo- 
phulid join mesolophid, reflected by long 
posteroflexid separating entoconid from 
median mure. In m2 and m3, entolophid 
joins median mure at level of anteriormost 
connection of hypolophid and median mure. 
Entolophulid extending to mesolophid re- 
sulting in short, narrow entoflexid whose 
medial extension becomes a small enamel 
island lost relatively early in wear, at which 
time entolophid assumes a quite broad ap- 
pearance. Entolophulid of m1, evident only 
in unworn teeth, isolating medial entoflexid 
as small enamel island (internal entoflexid) 
lost early in wear, at which time evidence 
of entolophulid is lost. Despite original dou- 
ble connection, resultant entolophid nar- 
rower than entoconid. The relatively broad 
hypoflexid extending to near midline in m1 


425 


and m2, and approximately two-thirds 
width of tooth in m3. In all three lower 
molars, hypoflexid may contain one or more 
small cuspulids at outer margin. A shallow 
flexid indents posterolabial side of each mo- 
lar at juncture of posterolophid and hypo- 
lophid. Distal end of posterolophid in m1 
and m2 lies close to entoconid and, with 
relatively little wear, appears fused to en- 
toconid isolating posteroflexid as an island. 
In m3, posteroflexid appears enclosed only 
in well-worn teeth. Lower molars with de- 
cidedly anterior orientation of metalophids 
and entolophids (and correspondingly long 
meso- and posteroflexids) in ml and m2 
along with the relatively well-developed en- 
toconid in m3 and correspondingly shallow 
hypoflexid. 

Age categories based on teeth wear are 
defined as follows: Age 0, all cheek teeth not 
fully erupted and functional; age 1, all teeth 
erupted and functional, lingual cusps show- 
ing relatively little wear; age 2, lingual cusps 
worn, but not flat, occlusal surfaces showing 
some wear on all cusps; age 3, all lingual 
cusps on M2 and M3 worn flat, labial cusps 
showing wear; age 4, all cusps worn, occlusal 
surface of M3 flat, some major dental fea- 
tures may be obliterated. 

Comparisons.—Among known species, 
T. apeco needs comparison with only Me- 
gaoryzomys curioi and the species of large 
Thomasomys comprising the 7. aureus 
complex. Megaoryzomys curioi 1s larger (see 
Table 2) with relatively square molars and 
a shallow notch between lacrimal and zy- 
gomatic ramus of maxilla (see Steadman & 
Ray 1982:figs. 7D and 9A-lower). Dental 
characteristics (e.g., paraloph joins para- 
cone to median mure; compare A and B in 
Steadman & Ray 1982:fig. 4) place M. curioi 
well apart from other known species of larg- 
er thomasomyines. Although 7. apeco ap- 
proaches M. curioi in size, dental features 
suggest closer affinity with species of the 7. 
aureus complex. The only major dental dif- 
ferences we have found that separate 7. ape- 


426 


co from specimens of 7. aureus from Peru, 
including Rio Abiseo National Park, are the 
more medial orientation of the paraloph in 
M1 and M2, more prominent anterior mure 
in M2, relatively larger and more widely 
separated anterior conulids in ml, more 
rounded posteroloph in ml and m2, and 
larger entoconid on m3 with a correspond- 
ingly shorter hypoflexid that result in sim- 
ilar appearance of m2 and m3 (Fig. 3). Cra- 
nial features other than larger size (Table 2) 
that can be used to distinguish 7. apeco from 
T. aureus are the shallower zygomatic notch, 
much deeper notch between lacrimal and 
zygomatic ramus of maxilla, and much 
thicker jugal. The larger and thicker jugal 
in T. apeco results in a greater distance be- 
tween distal ends of maxillary and squa- 
mosal contribution to the zygomatic arch 
(distance exceeds greatest depth of zygo- 
matic arch at its midpoint). The jugal is 
conspicuously thin in 7. aureus and the gap 
between ends of maxilla and squamosal filled 
by the jugal is less than the greatest depth 
of the zygomatic arch at its midpoint. Ex- 
ternally, 7. apeco is much larger and more 
robust than 7. aureus and, although pro- 
portionally about the same length (from 56 
to 61% of total length), the tail is white dis- 
tally for nearly half its length in contrast to 
being uniformly dark in 7. aureus. 
Additional specimens examined. —Speci- 
mens we examined are deposited either in 
the collections of the Museo de Historia 
Natural, Universidad Nacional Mayor de 
San Marcos, Lima, Pera (MUSM); the Na- 
tional Museum of Natural History, Wash- 
ington, D.C., U.S.A. (USNM) [division of 
mammals,], USNM-P [paleobiology]); or are 
indicated by field numbers (MLL [Mariella 
Leo L.] and MRR [Monica Romo R.]). The 
majority of the latter will be deposited in 
the MUSM, which commonly has been re- 
ferred to in the literature as the Museo de 
Historia Natural “Javier Prado.” 
Thomasomys aureus complex: Colombia: 
Cundinamarca, Bogota, USNM 251957, 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


topotype of Thomasomys princeps (Tho- 
mas, 1900); Cundinamarca, Choachi, 
USNM 251956; Cundinamarca, Laguna 
Vergon [= Laguna del Verjon], USNM 
251976. ECUADOR: Pichincha, Rio Con- 
dor Huachana, 3.45 km NE of Lloa, USNM 
513588, 513589. PERU: San Martin, Rio 
Montecristo, ca. 28 km NE of Pataz, MLL 
312; San Martin, Puerta del Monte, ca. 26 
km (at 60°) from Pataz, MLL 249; San Mar- 
tin, Las Palmas, ca 32 km NE of Pataz, 
MRR 579 and 594, MLL 340 and 343; San 
Martin, Las Papayas, ca. 32 km NE of Pa- 
taz, MLL 390; Junin, Rio Palca, 15 km W 
of San Ramon, USNM 507265 and 507266; 
Cuzco, Torontoy, 3260 m, USNM 194818; 
Cuzco, Tocopoque [= Tocopogueyu], Oc- 
cobamba Valley, 3000 m, USNM 194826- 
194828. Megaoryzomys curioi (Nietham- 
mer, 1964). ECUADOR: Islas Galapagos, 
Isla Santa Cruz, USNM-P 284204 (15 mo- 
lars), USNM-P 284213 (mandible with m2 
and m3), 284276 (right M1), USNM-P 
284283 (right mandible); USNM-P 284287 
(right m1), USNM-P 284343 (right man- 
dible with all molars), USNM-P 284346 
(right mandible with m1). 

Remarks.—One female was pregnant in 
August 1987 with a single embryo (CR = 
5.0 mm); the only female caught the follow- 
ing July showed well-developed mammae. 
Age-class-2 males had well-developed tes- 
tes in July and August (length = 23.0 mm), 
while the age-class-1 male had small testes 
in July (4.6 mm). 

A total of ten specimens were caught of 
which one was released. The holotype and 
the age-class-1 female collected at Los Cho- 
chos were caught in live traps, one at the 
end of a log and the other on a bank above 
a small stream, on the forest floor in an 
isolated patch of elfin forest. The age-class-4 
female was caught at Pampa del Cuy in an 
unbaited conibear trap set in a runway lead- 
ing up from a rivulet of water through grass 
on a slope in wet pampa habitat over a hun- 
dred meters from forest. Three were taken 


VOLUME 106, NUMBER 3 


in unbaited live traps placed in a small shal- 
low stream bordered by a few small trees in 
otherwise pampas habitat. Traps (three 
**Tomahawk’’) were placed to intercept any 
animals moving along the stream. The 
Puerto del Monte specimen was captured 
within continuous forest. Three additional 
specimen were trapped in pampas habitat 
associated with a small stream bordered by 
bushes and scattered small trees. One of 
these was dusted with fluorescent powder 
and released. Subsequent trailing of this an- 
imal showed that it had climbed a small tree 
and followed a branch to a bank above the 
stream where originally caught. 

Stomach morphology of MUSM 7196 
(age-class-4 female) shows similarity to that 
described and illustrated by Carleton (1973: 
14) for a T. aureus from Peru in contrast to 
the morphology he described for other spe- 
cies of Thomasomys. The stomach of our 
T. apeco was not sufficiently distended to 
permit a detailed comparison; nevertheless, 
while extension of cornified epithelium into 
the antrum is extensive, it is not as extreme 
as in 7. aureus and does not separate the 
glandular epithelium into two zones. 


Acknowledgments 


The inventory of Parque Nacional Rio 
Abiseo was supported by the Abiseo River 
National Park Research Project from the 
University of Colorado, the David and Lu- 
cile Packard Foundation, and the Pew 
Charitable Trust. Museum work for Leo was 
supported by the Smithsonian International 
Fellowship Program and the Peruvian As- 
sociation for Conservation of Nature. We 
gratefully acknowledge the many persons 
who were part of the inventory team, es- 
pecially Daniel Latorre and Monica Romo 
who were directly involved in the collection 
of these specimens. We also thank person- 
nel from the Parque Nacional Rio Abiseo 
for their help at several stages of the project, 
as well as the Direccion General Forestal y 


427 


de Fauna for authorizing and facilitating re- 
search in the park. We are indebted to 
George C. Steyskal for advice on the form 
of the species name and to Charles O. Han- 
dley, Jr., Division of Mammals, National 
Museum of Natural History, for sharing his 
knowledge of the genus Thomasomys and 
under whose direction the Smithsonian Fel- 
lowship was given. We greatly appreciate 
the assistance of Clayton E. Ray and Robert 
W. Purdy of the Department of Paleobiol- 
ogy, National Museum of Natural History, 
in examining material representing Mega- 
oryzomys curiol. 


Literature Cited 


Carleton, M. D. 1973. A survey of gross stomach 
morphology in New World Cricetinae (Roden- 
tia, Muroidea), with comments on functional 
interpretations.— Miscellaneous Publications, 
Museum of Zoology, University of Michigan 
146:1-43. 

——, & G.G. Musser. 1989. Systematic studies of 
oryzomyine rodents (Muridae, Sigmodontinae): 
a synopsis of Microryzomys.—Bulletin of the 
American Museum of Natural History 191:1- 
83. 

Gyldenstolpe, N. 1932. A manual of Neotropical sig- 
modont rodents.—Kungliga Svenska Veten- 
skapsakademiens Handlingar, Stockholm, Se- 
ries 4, 11(3):1-164, plates 1-18. 

Hershkovitz, P. 1944. A systematic review of the 
Neotropical water rats of the genus Nectomys 
(Cricetinae).— Miscellaneous Publications Mu- 
seum of Zoology, University of Michigan 58:1- 
101, 4 pls., 1 map (folded). 

Nowak, R.M. 1991. Walker’s mammals of the world. 
Fifth Edition. The Johns Hopkins University 
Press, Baltimore, 2:xii + 643-1629. 

Reig, O. A. 1977. A proposed unified nomenclature 
for the enamelled components of the molar teeth 
of the Cricetidae (Rodentia).—Journal of Zo- 
ology, London 181:227-241. 

Ridgway, R. 1912. Color standards and color no- 
menclature. Washington, D.C., iv + 43 pp., 53 
pls. 

Steadman, D., & C. Ray. 1982. The relationships of 
Megaoryzomys curioi, an extinct cricetine ro- 
dent (Muroidea: Muridae) from the Galapagos 
Islands, Ecuador.—Smithsonian Contributions 
to Paleobiology 51:1-—23. 

Tosi, J. A., Jr. 1960. Zonas de vida natural en el Peru. 


428 


Instituto Interamericano de Ciencias Agricolas 
de la OEA Zona Andina, Boletin Técnico No. 
5: Wil Gee? EL ispp: 

Young, K., & B. Leon. 1988. Vegetacion de la Zona 
Alta del Parque Nacional Rio Abiseo, San Mar- 
tin.— Revista Forestal del Peru 15(1):3—20. 


(MLL) Asociacion Peruana para la Con- 
servacion de la Naturaleza, Parque José de 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Acosta 187, Magdalena, Lima 17, Peru; 
(ALG) Biological Survey Field Station, Na- 
tional Ecology Research Center, U.S. Fish 
and Wildlife Service, National Museum of 
Natural History, Washington, D.C. 20560, 
US.A. 


PROC. BIOL. SOC. WASH. 
106(3), 1993, pp. 429-435 


ZOOGEOGRAPHY AND GEOGRAPHIC VARIATION OF 
ATLAPETES RUFINUCHA (AVES: EMBERIZINAE), 
INCLUDING A DISTINCTIVE NEW 
SUBSPECIES, IN SOUTHERN 
PERU AND BOLIVIA 


J. V. Remsen, Jr. 


Abstract. —A distinctive new subspecies, Atlapetes rufinucha terborghi (Em- 
berizinae), is described from the isolated Cordillera Vilcabamba, a spur range 
of the Andes in Dpto. Cuzco, Peru. This population is isolated from the nearest 
populations of A. rufinucha by more than 200 km, and intervening areas with 
suitable habitat are inhabited by another species, A. schistaceus. The new taxon 
is greener breasted than any other subspecies of A. rufinucha. The four sub- 
species found in southern Peru and Bolivia represent four discrete phenotypes 
with respect to plumage. Geographic variation in size in the southern Andes 


does not support Bergmann’s Rule. 


The Cordillera Vilcabamba, Dpto. Cuz- 
co, Peru, is a mountain range isolated from 
the main chain of the Andes by deep river 
valleys with tropical, non-montane habitats 
(Terborgh 1971, Haffer 1974). Although 
specimens of birds collected there in the late 
1960’s by John Weske and John Terborgh 
have yet to be analyzed in a systematic man- 
ner, some endemic taxa have been or are 
being described: Schizoeaca vilcabambae 
(Vaurie et al. 1972, Remsen 1981), Crani- 
oleuca marcapatae weskei (Remsen 1984), 
Ochthoeca fumicolor subsp. nov. (P. Hosey, 
in litt.), and Coeligena violifer subsp. nov. 
(J. Weske and J. P. O’Neill, pers. comm.). 

While examining specimens of Andean 
Atlapetes for a project on their patterns of 
distribution (Remsen & W. S. Graves 1994), 
I found five specimens of A. rufinucha col- 
lected by Weske and Terborgh in the Vil- 
cabamba that represent an important range 
extension for this species. Furthermore, 
these specimens differ distinctly from any 
other population of A. rufinucha, so much 
so that even with just five specimens, it is 
clear that they represent a new taxon, which 
may be known as: 


Atlapetes rufinucha terborghi, 
new subspecies 


Holotype.— American Museum of Natu- 
ral History (AMNH) #820436; mist-netted 
by John S. Weske and John W. Terborgh 
on 22 Jul 1967 in the Cordillera Vilcabam- 
ba, 2630 m, Departamento Cuzco, Peru, 
12°37'S, 73°33'W. The specimen, prepared 
by Weske (#1334), is a female in breeding 
condition (ovary and oviduct much en- 
larged, largest ovum 8 mm; brood patch 
present) with a completely pneumatized 
skull and little fat. 

Description. —Capitalized color names are 
from Ridgway (1912). Crown closest to Ha- 
zel, becoming slightly paler (Cinnamon-Ru- 
fous) on hind-crown and nape, with some 
Cinnamon Rufous feathers extending to ex- 
treme upper back. Rest of back, upper sides 
of wings, and tail black, obscurely suffused 
with olivaceous tones on back and on outer 
webs of secondaries. Upper tail coverts Ol- 
ivaceous Black. Lores, broad superciliary, 
face, and auriculars black, contiguous with 
black of dorsum at neck, but contrasting 
with the darker back. Chin and throat Light 


430 


Greenish Yellow. Faint, broken malar stripe 
dark olive. Breast closest to Javel Green. 
Center of belly like throat, blending to broad, 
darker, Olive Green flanks and slightly paler 
undertail coverts. Rather than uniformly 
colored, the underparts have faint hints of 
obscure streaks throughout; the breast is not 
sharply demarcated from the paler throat 
and belly. Undersides of remiges and rec- 
trices Fuscous. Soft part colors recorded by 
Weske: iris rich brown; bill black; legs dark 
brown. Measurements: wing chord 70.2 mm; 
exposed culmen 18.5 mm; tail 74.2 mm; 
and tarsus 23.9 mm. 

Diagnosis. —Atlapetes r. terborghi is 
greener and darker ventrally over-all than 
any other subspecies of A. rufinucha. It can 
be distinguished from the nearest popula- 
tion of A. rufinucha, A. r. melanolaemus 
(which A. r. terborghi resembles in lacking 
a loral spot and black feathers on fore- 
crown), by its yellow-green throat and malar 
area, which is variably clouded with black 
in A. r. melanolaemus. In A. r. terborghi, 
the breast, although greenish and darker than 
the throat or belly, lacks the variable amount 
of irregular blackish scalloping that char- 
acterizes melanolaemus. The nominate 
subspecies of A. rufinucha, from dptos. La 
Paz and Cochabamba, Bolivia, (which is 


more similar in plumage to other subspecies . 


of A. rufinucha throughout the Andes than 
either A. r. terborghi or A. r. melanolaemus) 
differs from A. r. terborghi in the following 
ways: (1) A. r. terborghi lacks the yellowish 
loral spot of the nominate subspecies; this 
area 1s black in A. r. terborghi; (2) in A. r. 
terborghi, the chestnut of the crown extends 
to the bill, whereas in the nominate sub- 
species the feathers closest to the bill on the 
forehead are black; (3) the nominate form 
has a conspicuous black malar stripe, 
whereas A. r. terborghi has only a faint trace 
of a darker malar stripe; (4) the underparts 
of the nominate subspecies are a bright yel- 
low, whereas those of A. r. terborghi are 
greener; even in the two most yellow spec- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


imens of A. r. terborghi, the yellow is paler 
and more washed out than in the nominate 
subspecies. The back and uppertail coverts 
of A. r. terborghi are slightly paler than those 
of A. r. melanolaemus or the nominate sub- 
species; the black of the face of A. r. ter- 
borghi contrasts with the paler back, where- 
as there is little or no contrast between the 
face and back of A. r. melanolaemus or the 
nominate subspecies. In these respects, A. 
r. terborghi resembles distant A. r. carrikeri 
(but A. r. carrikeri is much paler-backed, 
paler-breasted, and smaller in size); such 
mosaic distribution of characters is a prom- 
inent feature of geographic variation in A. 
rufinucha (Paynter 1978). 

Distribution. —Cordillera Vilcabamba, 
Dpto. Cuzco, Peru. 

Paratypes.—Four other specimens were 
also mist-netted in the Cordillera Vilca- 
bamba by Weske and Terborgh and _ pre- 


pared by Weske: (1) AMNH 820438, 2620 


m, 12°37’S, 73°33’W, male, 18 Jul 1967; (2) 
AMNH 820437, 2620 m, 12°37’S, 73°33'W, 
male, 30 Jul 1967; (3) AMNH 820609, 3300 
m, 12°36'S, 73°30’W, female, 17 Jul 1968; 
and (4) AMNH 820633, 3500 m, 1[2°36’S, 
73°29'W, female, 31 Jul 1968. The two males 
are darker below than the holotype, with 
the underparts more heavily suffused with 
green; the two females are paler than the 
holotype, with brighter yellow throats and 
bellies. The holotype, a female, was selected 
partly because it is intermediate in ventral 
coloration, although closer to the two males. 
One female paratype (820609) has a trace 
of the yellow loral spot and a few black 
feathers on the forecrown, characters-more 
fully developed in other subspecies. Soft part 
colors were not recorded on any of the para- 
types, but in the dried study skins, the bills, 
tarsi, and toes appeared identical in color 
to those of the holotype. Measurements of 
the paratypes and holotype. are in Table 1. 

Etymology. —It is a pleasure to name this 
distinctive taxon endemic to the Cordillera 
Vilcabamba for the person who engineered 


VOLUME 106, NUMBER 3 


and conducted its ornithological explora- 
tion, Dr. John W. Terborgh. The survey of 
the Vilcabamba by Terborgh and John 
Weske represents the most rigorous and well- 
executed inventory of any area of the Andes. 
The name is particularly appropriate be- 
cause patterns of distribution of brush- 
finches in the Andes provide evidence for 
the influence of interspecific competition on 
their zoogeography (Remsen & Graves 
1994), a major theme of Terborgh’s (1971) 
research in the Vilcabamba. 


Natural History 


The only information available comes 
from the specimen labels. Four specimens 
were mist-netted in humid, montane cloud- 
forest and elfin forest, from 2620 to 3300 
m, and the fifth was mist-netted in “‘mixture 
of tall grassland and elfin forest on crest of 
mountain range” at 3500 m; Weske (1972) 
gave the elevational limits as 2520 to 3520 
m. All five specimens are adults: for four of 
the five, skull pneumatization was recorded 
as ““complete,”’ and for the one with no skull 
notation, the testes were highly enlarged 
(largest 11 mm). Both males had enlarged 
testes and cloacal protuberances. The ho- 
lotype female was also in breeding condi- 
tion, with the largest ovum 8 mm, but for 
the other two females, gonad information 
was recorded as “ovary 5 X 4 mm (not 
enlarged)” and “ovary 6 mm (not en- 
larged).”’ Thus, at least some individuals 
seemed to be breeding in July, which is dur- 
ing the driest part of the year in the southern 
Andes (Weske 1972, Fjeldsa & Krabbe 
1990). 

If A. r. terborghi is like other subspecies 
of A. rufinucha, then it should be acommon, 
conspicuous species that favors forest edge 
rather than interior; and it should forage 
actively, often in mixed-species flocks, from 
near ground to the subcanopy while search- 
ing foliage of trees and shrubs, and epi- 
phyte-covered as well as bare branches 
(Remsen 1985, Fjeldsa & Krabbe 1990). 


431 


Table 1.—Measurements (in mm) of holotype and 
paratypes of At/apetes rufinucha terborghi. 


Exposed 

Wing Tail Tarsus culmen 

Specimen (AMNH #; sex) chord length length length 
820436; 2 (holotype) 70.2 74.2 23.9 18.5 
820609; ¢ Ae 79 5253) 17.9 
820633; 2 Ge nI4.2., 247 18.4 
820437; 3 TALS IG6:3- ~ 25:0 18.6 
820438; 3 iss Tt. ~~ 25.0 19.4 


Geographic Variation 


Atlapetes r. terborghi is the northernmost 
of four distinctive subspecies distributed 
from about 18°S in northern Dpto. Santa 
Cruz, Bolivia, north to ca. 12°S in Dpto. 
Cuzco, Peru. Paynter (1978) summarized 
plumage features of the three previously de- 
scribed subspecies (the nominate subspe- 
cies, A. r. carrikeri, and A. r. melanolae- 
mus), each of which represents a discrete, 
strongly marked unit (Graves 1985). The 
only signs of intergradation between any of 
the subspecies are as follows. A specimen 
(Louisiana State University Museum of 
Natural Science, hereafter LSUMZ, 96808) 
in a series of 55 of the nominate subspecies 
from Cotapata, Chuspipata, and Sacramen- 
to Alto, Dpto. La Paz, has black scalloping 
on the breast and an enlarged black malar 
area, thereby approaching melanolaemus; 
whether this represents intergradation or in- 
dividual variation within the nominate form 
cannot be determined. 

Specimens from the El Choro area, Prov. 
Ayopaya, northern Dpto. Cochabamba, are 
anomalous in their variable crown color. 
Most show the nearly typical cinnamon-ru- 
fous crown of the nominate subspecies, but 
several are paler to varying degrees. For ex- 
ample, some (e.g., LSUMZ 36861) are no- 
tably but not greatly paler, some are paler 
still (e.g., Academy of Natural Sciences, 
Philadelphia 134927; Field Museum of 
Natural History 217844), and two (ANSP 
134928, FMNH 217837) are very pale, al- 


432 


most as pale as A. r. baroni of the Rio Ma- 
ranon valley of Peru. 

Atlapetes rufinucha is one of many (but 
not all; see Graves 1991, Kratter 1993) An- 
dean bird species and superspecies that show 
geographic variation in body size that con- 
tradicts Bergmann’s “Rule” (Remsen 1984; 
Remsen et al. 1991; R. Brumfield, pers. 
comm; D. Wiedenfeld, pers. comm.). Over 
the limited latitudinal range of the southern 
populations of A. rufinucha a non-Berg- 
mannian pattern is evident. To illustrate 
geographic variation in size, I used wing 
length as an index of body size. For 30 male 
A. rufinucha specimens with body mass data 
from Dpto. Puno, Peru, to Dpto. Santa Cruz, 
Bolivia, wing length is significantly corre- 
lated with cube root of body mass (Kendall’s 
Tau corrected for ties = 0.27, P = 0.04), in 
spite of the large potential measurement er- 
ror associated with body mass caused by use 
of different scales by different workers at 
different localities on birds that had often 
been in mist nets for varying periods. How- 
ever, for 24 females from the same area, 
wing length is not significantly correlated 
with cube root of body mass (Kendall’s Tau 
corrected for ties = 0.07, P = 0.66). For 
males, wing length decreases significantly 
with increasing latitude (Kendall’s Tau cor- 
rected for ties = 0.14; P = 0.03; Fig. 1). For 
females, wing length also decreases with in- 
creasing latitude, but the relationship is not 
Statistically significant (Kendall’s Tau cor- 
rected for ties = 0.14, P = 0.10; Fig. 1). For 
both males and females, the relationship of 
wing length to latitude may not be linear; 
only increased sample sizes from southern 
Peru and northern Bolivia will determine 
whether the apparent trough at 14—-15°S and 
consequent bimodal distribution is real or 
an artifact of low sample size. Elevation is 
not significantly correlated with wing length 
in either sex (Kendall’s Tau corrected for 
ties = 0.075 for males, 0.14 for females, P 
= 0.27, 0.09, respectively) or with latitude 
(Kendall’s Tau corrected for ties = 0.013 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


for males, —0.062 for females, P = 0.84, 
0.46, respectively). 

Interpretation of latitudinal gradients in 
body size is complex when the populations 
are not in genetic contact (Graves 1991), as 
is the case in the four taxa of A. rufinucha. 
In two of the four taxa, sample sizes and 
latitudinal ranges are large enough to ana- 
lyze trends within a taxon. In A. r. rufinucha, 
both males and females show a “Bergman- 
nian” trend, but the relationship is not sta- 
tistically significant (Kendall’s Tau correct- 
ed for ties = —0.115 for males, —0.155 for 
females, P = 0.15, both sexes). In A. r. me- 
lanolaemus, both males and females show 
a “non-Bergmannian” trend, but the rela- 
tionship is statistically significant only for 
females (Kendall’s Tau corrected for ties = 
0.085 for males, 0.50 for females, P = 0.63, 
0.007, respectively). 


Zoogeography 


Recent fieldwork in Peru by the Field Mu- 
seum of Natural History has extended the 
distribution of A. r. melanolaemus north in 
Dpto. Cuzco to 13°13’S, where John W. 
Fitzpatrick and David Willard collected 
specimens at Pillahuata, 2510 m, in the Rio 
Cosnipata valley, near the northern limit of 
the Cordillera de Carabaya. This locality is 
about 75 km north of the Marcapata region 
of southeastern Dpto. Cuzco, the previous 
northern limit for A. r. melanolaemus. Oth- 
er areas of humid montane forest in Dpto. 
Cuzco between the Cosnipata valley and the 
Cordillera Vilcabamba are inhabited by a 
different species, Atlapetes schistaceus. 
Remsen & Graves (1994) found that A. ru- 
finucha and A. schistaceus replace each oth- 
er throughout their extensive Andean dis- 
tributions; they proposed that these two 
species were either close relatives and com- 
petitors or, conversely, that they were yel- 
low and gray representatives of the same 
lineage. In either case, A. r. terborghi and A. 
r. melanolaemus are separated from each 


VOLUME 106, NUMBER 3 433 
80 
8 1 i 
! 
is oe Pie , e | @ 
e | 7 e- @ @ ® ; 
ace ® e_. 
Beh Ge Bitoniy : anima 
aren 18 — rat oO O : 
~— = iF @ u Si 1 
: al = | oo : 
S ree Hy PO aE ! 
z | Ea E a : 
Ll ore I 
2 3 | : 
= : " ! ! 
3 : Oo : : 
| : 
carrikeri i rufinucha ! melanolaemus | terborghi 
! | ! 
@ males 
CL) females 
50 
-19 -18 -17 -16 -15 -14 -13 -12 
S. LATITUDE 
Fig. 1. Relationship of wing length and latitude (degrees S Lat.) for four subspecies of Atlapetes rufinucha in 


southern Peru and northern Bolivia. Latitudes, and elevations for localities not recorded on the specimen labels, 
were taken from Stephens & Traylor (1983) and Paynter (1992). Vertical dashed lines represent approximate 
boundaries between four subspecies of region. Diagonal lines represent regression lines of wing length on latitude 
for males (upper line; y = 82.1 + 0.72x, r? = 0.11) and females (lower line; y = 76.2 + 0.60x, r? = 0.13); lines 
presented only to illustrate general trends, not formal statistical analyses, because data are largely inappropriate 


for parametric statistics. 


other by more than 200 km, and so A. r. 
terborghi is yet another isolated population 
of A. rufinucha (Paynter 1978). The gap, 
between the Urubamba-Concebidayoc val- 
ley east to at least the Rio Vilcanota valley, 
is inhabited by A. schistaceus canigenis. 
Which taxon of Atlapetes, if any, inhabits 
the region from there east to the Rio Yan- 
atili valley and Rio Yavero valley, the 
northwestern limit of A. r. melanolaemus, 
is unknown. 

Recent fieldwork in Peru by the Museum 
of Natural Science, Louisiana State Uni- 
versity, has extended the distribution of A. 
r. melanolaemus south to near the Bolivian 


border, where L. C. Binford and T. S. Schu- 
lenberg collected specimens in Dpto. Puno 
at Valcon, 3000 m, 14°26’S, and Abra de 
Marucunca, 2000 m, 14°14’S. The southern 
limit of A. r. melanolaemus is unknown but 
is probably somewhere in northern Dpto. 
La Paz, possibly the north side of the Rio 
Mapiri canyon, another region virtually 
unexplored by ornithologists. 

Weske (1972) listed one locality record 
for Atlapetes tricolor in the Vilcabamba, at 
2100 m, below the lower limit of A. r. ter- 
borghi. If A. tricolor occurs at lower eleva- 
tions than A. rufinucha in the Vilcabamba, 
then this would represent a similar pattern 


434 


in elevational distribution to that in the 
Western Andes of Colombia and Ecuador, 
where the two species replace each other at 
about 2000 m (Remsen & Graves 1994). In 
central Peru, from central Dpto. Cuzco north 
to Dpto. Huanuco, A. tricolor is replaced at 
higher elevations by A. schistaceus. 

Additional recent fieldwork in Peru by the 
Museum of Natural Science, Louisiana State 
University, has extended the distribution of 
A. r. rufinucha southeastward from its pre- 
vious southern limit in Prov. Chapare, Dpto. 
Cochabamba, Bolivia, into Prov. Carrasco, 
where C. Gregory Schmitt and Donna C. 
Schmitt collected a specimen at Quebrada 
Majon, 6.6 km northwest of Lopez Men- 
doza, 3150 m (17°32'S, 65°22'W). This 
specimen is indistinguishable from speci- 
mens from Prov. Chapare. 

A specimen (LSUMZ 38472) collected by 
F. Steinbach at San Mateo, extreme eastern 
Prov. Carrasco, Dpto. Cochabamba, near 
the Dpto. Santa Cruz border, represents A. 
r. carrikeri, formerly known only from Dpto. 
Santa Cruz. The specimen matches the type 
specimen of A. r. carrikeri at ANSP (M. B. 
Robbins, in litt.). 


Acknowledgments 


I thank Franc¢ios Vuilleumier and his cu- 
ratorial staff (American Museum of Natural 
History) for loaning the specimens of the 
new taxon. I am grateful to the staffs of the 
Academy of Natural Sciences at Philadel- 
phia, the American Museum of Natural 
History, and the Field Museum of Natural 
History for access to their collections and 
hospitality. Mark B. Robbins compared 
LSUMZ specimens of A. r. carrikeri to the 
type specimen at ANSP. T. S. Schulenberg, 
Scott M. Lanyon, and David Willard com- 
pared crown colors of A. r. rufinucha from 
El Choro to that of A. r. baroni and loaned 
specimens of that taxon from the Field Mu- 
seum of Natural History. Steven W. Cardiff, 
Gary R. Graves, Manuel Marin A., Thomas 
A. Munroe, John P. O’Neill, Raymond A. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Paynter, Jr., and David A. Wiedenfeld made 
many helpful comments on the manuscript. 


Literature Cited 


Fjelda, J., & N. Krabbe. 1990. Birds of the high An- 
des. Zoological Museum, University of Copen- 
hagen, Denmark, 876 pp. 

Graves, G. R. 1985. Elevational correlates of speci- 
ation and intraspecific geographic variation in 

plumage of Andean forest birds.— Auk 102:556— 

579. 

. 1991. Bergmann’s rule near the equator: lat- 

itudinal clines in body size of an Andean pas- 

serine bird. — Proceedings of the National Acad- 

emy of Sciences 88:2322—2325. 

Haffer, J. 1974. Avian speciation in tropical South 
America.— Nuttall Ornithological Club Publi- 
cation no. 14, 390 pp. 

Kratter, A. W. 1993. Geographic variation in the 
Yellow-billed Cacique (Amblycercus holoseri- 
ceus), a partial bamboo specialist.— Condor (in 
press). 

Paynter, R. A., Jr. 1978. Biology and evolution of 

the avian genus A¢lapetes (Emberizinae).—Bul- 

letin of the Museum of Comparative Zoology 

148:323-369. 

1992. Ornithological gazetteer of Bolivia. 
Second edition. Museum of Comparative Zo- 
ology, Cambridge, Massachusetts, 185 pp. 
Remsen, J. V., Jr. 1981. A new subspecies of Schi- 
zoeaca harterti (Aves: Furnariidae), with com- 
ments on the taxonomy of Schizoeaca.—Pro- 
ceedings of the Biological Society of Washington 
94:1068-1075. 

1984. Geographic variation, zoogeography, 
and possible rapid evolution in some Crani- 
oleuca spinetails.— Wilson Bulletin 96:515—523. 
. 1985. Community organization and ecology 
of birds of high elevation humid forest of the 
Bolivian Andes. Pp. 733-756 in P. A. Buckley 
et al., ed., Neotropical Ornithology. Ornitho- 
logical Monographs No. 36. 

—-, & W.S. Graves. 1994. Distribution patterns 
and zoogeography of Atlapetes brush-finches 
(Emberizinae) of the Andes. — Auk (in press). 

— , O. Rocha O., C. G. Schmitt, & D. C. Schmitt. 
1991. Zoogeography and geographic variation 
of Platyrinchus mystaceus in Bolivia and Peru, 
and the Circum-Amazonian distribution pat- 
tern.—Omnitologia Neotropical 2:77-83. 

Ridgway, R. 1912. Color standards and color no- 
menclature. Published by the author. Washing- 
ton, D.C. 

Stephens, L., & M. A. Traylor, Jr. 1983. Ornitholog- 
ical gazetteer of Peru. Museum of Comparative 
Zoology, Cambridge, Massachusetts, 271 pp. 


VOLUME 106, NUMBER 3 435 


Weske, J. S. 1972. The distribution of the avifauna 
in the Apurimac Valley of Peru with respect to 
environmental gradients, habitats, and related 
species.—Unpubl. Ph.D. dissertation, Univer- 
sity of Oklahoma, 137 pp. 


Terborgh. J. 1971. Distribution on environmental 
gradients: theory and a preliminary interpreta- 
tion of distributional patterns in the avifauna of 
the Cordillera Vilcabamba, Peru.—Ecology 52: 


23-40. 
Vaurie, C., J.S. Weske, & J. W. Terborgh. 1972. Tax- 
onomy of Schizoeaca fuliginosa (Furnariidae), Museum of Natural Science, Louisiana 


with description of two new subspecies. —Bul- i ; ay 
letin of the British Ornithologists’ Club 92:142- State University, Baton Rouge, Louisiana 
144. 70803, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(3), 1993, pp. 436-441 


A NEW HYBRID MANAKIN 
(DIXIPHIA PIPRA X PIPRA FILICAUDA) 
(AVES: PIPRIDAE) FROM THE ANDEAN 

FOOTHILLS OF EASTERN ECUADOR 


Gary R. Graves 


Abstract. —A new intergeneric hybrid manakin (Dixiphia pipra xX pipra fili- 
cauda) is described from the Andean foothills of eastern Ecuador. The adult 
male specimen represents the first record of hybridization among species be- 
longing to different clades of the traditional but polyphyletic genus Pipra (Prum 
1990, 1992). The collecting locality, Sarayacu (1°44’S, 77°29'W), apparently 
lies in a narrow zone of elevational overlap between the two parental species. 
The hybrid is nearly intermediate between the parental species in size, plumage 


pattern, and color. 


The few documented cases of hybridiza- 
tion 1n neotropical manakins (Pipridae) have 
been between species within the same su- 
perspecies, or between species in different 
genera (Parkes 1961, Haffer 1967, Parsons 
et al. 1993). Prum’s (1990, 1992) recent re- 
vision of the family indicated that the genus 
Pipra, as currently recognized (e.g., Sibley 
& Monroe 1990), is polyphyletic, composed 
of three monophyletic clades (Pipra, Dixi- 
pDhia, Lepidothrix). Hybridization among 
species belonging to different “‘Pipra’”’ lin- 
eages is unknown and the subject of this 
paper. 

An enigmatic manakin was collected by 
M. Olalla on 21 August 1951, at Sarayacu 
(1°44’S, 77°29'W;, Paynter & Traylor 1977), 
Provincia de Pastaza, Ecuador. Not clearly 
assignable to any described species, the 
specimen (Museo Ecuatoriano de Ciencias 
Naturales no. 2748) had been variously 
identified on the tag by taxonomists as a 
hybrid (i.e., Pipra filicauda x Chiroxiphia 
sp., P. filicauda x Heterocercus sp., P. fili- 
cauda x P. erythrocephala). Analyses of 
plumage characters and external morphol- 
ogy revealed that none of these hypotheses 
was correct: the specimen represents a hy- 
brid between Dixiphia pipra (white-crowned 


manakin) and P. filicauda (wire-tailed man- 
akin). 


Materials and Methods 


Sexed as a male, the specimen appears 
fully mature as judged by its glossy black 
mantle, wings, and tail. I compared it with 
series of all species of manakins in the Mu- 
seo Ecuatoriano de Ciencias Naturales and 
the National Museum of Natural History, 
Smithsonian Institution. For the purpose of 
the hybrid diagnosis (see Graves 1990), I 
considered all species of manakins (n = 16) 
that occur in western Amazonia and the ad- 
jacent Andean foothills of Colombia, Ec- 
uador, and Peru north of the Rio Maranon 
as potential parental species. Measurements 
of wing chord, length of central rectrices (= 
tail), and bill length from anterior edge of 
nostril were taken with digital calipers. Col- 
or comparisons were made under examo- 
lites (Macbeth Corp.). 


Results 


Plumage characters.—Determination of 
parentage was facilitated by the boldly pat- 
terned plumage of the hybrid (Fig. 1) and 
by the small number of potential parental 


\~R V r 
\ ' Vv 
oS Q’ 6b 
we Y si 


gs4-z 
hel rs ce 445 e4 
ctor. oeenee: 


“at aité »- Bona’ 


Fig. 1. Ventral, lateral, and dorsal views of adult males of Pipra filicauda (bottom), a presumed Dixiphia 
pipra x P. filicauda hybrid, and Dixiphia pipra (top). 


438 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
ras, 
O 
/ \ 
D. pipra Je 
@4 
CY ® 
Vo ae 
65 
O O05 
_ \@ 
ee © O 
P.. .tilicauda 
60 ® 0% 
Deane. 
P. erythrocephala 
S 
& 
HS 
15 ZU) 25 30 35 AO 
TAIL 


Fig. 2. Bivariate plot of wing and tail lengths (mm) of Dixiphia pipra, Pipra filicauda, P. erythrocephala, and 
a presumed D. pipra x P. filicauda hybrid (filled circle). 


species. Of the many possible combinations 
of parental species, only one (Pipra filicauda 
x Dixiphia pipra) could combine to pro- 
duce the diagnostic characters of the hybrid: 
(1) golden yellow crown and hindneck; (2) 
dark facial patch; (3) mottled yellow and 
black underparts; (4) glossy black tail, wings, 
and mantle; and (5) shallowly forked tail 
(see Appendix). 

The distribution of carotenoid pigments 
in the plumage of the hybrid is similar to 
manakins in the Pipra aureola superspecies, 
including P. filicauda. This fact and the 
forked tail of the hybrid suggest that P. fil- 
icauda is one of its parental species. Because 
P. filicauda has an unmarked yellow face, 
the dark facial patch of the hybrid must 
have been inherited from the other parental 
species. The dark facial patch of the hybrid 


is formed by feathers of the loral and sub- 
ocular areas and the auriculars. Several spe- 
cies of manakins in western Amazonia have 
black auriculars. Xenopipo atronitens and 
Heterocercus spp. can be eliminated from 
consideration because they possess struc- 
tural modifications of the remiges and rec- 
trices, respectively, that are absent in the 
hybrid. Likewise, the plumage of the hybrid 
lacks the distinctive plumage characters, 
even in traces, of several other potential pa- 
rental species: Lepidothrix isidorei (blue 
rump patch); Chiroxiphia pareola (blue 
mantle), and Masius chrysopterus (frontal 
tuft, yellow webs of remiges and rectrices). 
Two remaining species with black auricu- 
lars have black body plumage and contrast- 
ing crowns: D. pipra (white crown) and L. 
coronata (blue crown). Although the inher- 


VOLUME 106, NUMBER 3 


439 


Table 1.—Ranges and means (+one standard deviation) of measurements (mm) of adult males of Dixiphia 
pipra, Pipra filicauda, and P. erythrocephala from eastern Ecuador, and the hybrid, D. pipra x P. filicauda. 


Characters 
(n) Wing Tail Bill length 
D. pipra 10 64.5-69.2 25.1-28.1 6.4-7.2 
GT2 + 1t.32 26.8 + 1.02 6:7 3: 8:25 
P. filicauda 10 61.6-66.0 33.0-34.3 7.0-8.0 
63:3 = 1.33 34.3 + 1.52 7.4 + 0.26 
P. erythrocephala 10 55.9-62.3 17.6—21.2 5.9-6.8 
FoF 21D 19.42.19) 6.2 + 0.29 
Hybrid 1 67.7 29.7 7.1 


itance of structural colors is poorly under- 
stood, it is probable that a hybrid of L. cor- 
onata would exhibit at least traces of blue 
on the crown. I interpreted this absence as 
evidence that L. coronata was not a parent 
of the hybrid. Other combinations of spe- 
cies either lack the range of plumage pattern 
elements expressed in the hybrid or possess 
distinctive characters not found in the hy- 
brid. For example, the specimen in question 
could not represent a hybrid of P. filicauda 
and P. erythrocephala because neither spe- 
cies has black auriculars. Thus, a hybrid be- 
tween D. pipra and P. filicauda is the only 
remaining possibility. 

External morphology. —The parental hy- 
pothesis derived from plumage pattern and 
color was supported by a bivariate scatter- 
plot of wing and tail length (Fig. 2). Because 
the specimen had once been identified as a 
hybrid of P. filicauda and P. erythrocephala, 
the latter species has been included in Table 
1 for comparison. Under the assumptions 
used here (Graves 1990), if the specimen 
represents a hybrid of D. pipra and P. fili- 
cauda, then the mensural dimensions of the 
specimen should fall within the range of 
measurements of the parental species. This 
expectation was confirmed. Wing length of 
the hybrid was greater than that of either P. 
filicauda or P. erythrocephala, indicating that 
the hybrid is not a product of this combi- 
nation of species. Otherwise, the hybrid 
would be required to exhibit both atavism 


(hybrid expresses characters not found in 
either parental species) and morphological 
luxuriance (hybrid is larger than either pa- 
rental species). Neither phenomenon has 
been convincingly demonstrated in hybrids 
of passerine birds. 

Geography. — The collection locality, Sar- 
ayacu, lies at the base of the Andes on the 
Rio Bobonaza, a primary tributary of the 
Rio Pastaza. The elevation of this site was 
not recorded on the specimen label; the air- 
strip at Sarayacu lies between the 300 and 
600 m contour lines, approximately 40 km 
upslope from the 300 m contour interval 
and within 10 km of the 600 m contour 
interval (1:1,000,000 Mapa Fisico, Instituto 
Geographico Militar, Quito, 1991). Pipra 
filicauda is widespread in the lowlands of 
eastern Ecuador (recorded to 500 m ele- 
vation in Colombia, Hilty & Brown 1986). 
However, D. pipra has not been reported in 
Ecuador below 400 m elevation (M. B. Rob- 
bins, pers. comm.). Thus, the hybrid was 
apparently collected in a narrow zone of 
overlap between the two species. 


Discussion 


The hybrid exhibits a blend of parental 
plumage patterns rather than a mosaic of 
plumage elements. The one exception is the 
peculiar dark facial patch of the hybrid. The 
auricular area of both parental species is 
concolor with the throat and breast. Under 


440 


magnification, the auriculars and subocular 
feathers of the hybrid appear grizzled yellow 
and black. At arm’s length, however, this 
region contrasts with the adjacent yellow 
plumage of the throat and crown. Inheri- 
tance of a dark facial patch has also been 
noted previously in another manakin hy- 
brid, Pipra aureola Xx Heterocercus linteatus 
(Parkes 1961). These observations suggest 
that pigmentation of the facial region in Pi- 
pra manakins is controlled independently 
of the crown and throat. 

A surprising variety of carotenoid pig- 
ments has been isolated from manakins. 
Brush (1969) extracted the keto-carote- 
noids, canthaxanthin and astaxanthin, from 
crown feathers of Pipra aureola, which is 
closely related to P. filicauda, a parent of 
the hybrid described here. In a later analysis 
employing high-performance liquid chro- 
matography, Hudon et al. (1989) isolated at 
least nine carotenoid pigments, including 
xanthophylls and lutein, from the red crown 
feathers of P. rubrocapilla. These data sug- 
gest that the crown feathers of P. filicauda 
may be colored by a similar diversity of 
pigments. In contrast, the tips of the crown 
feathers are unpigmented in the other pa- 
rental species, Pipra pipra. The yellowish- 
orange crown of the hybrid manakin prob- 
ably represents a dilution of pigments pres- 
ent in P. filicauda. In an analogous case, 
Brush (1970) isolated a single carotenoid 
pigment from the rump feathers of the tan- 
agers, Ramphocelus flammigerus (bright 
scarlet) and R. icteronotus (lemon yellow) 
and their hybrids. He hypothesized that the 
“orange” rump color in hybrids was due 
simply to quantitative differences in the 
amount of the pigment present. In mana- 
kins, the expression of “‘red’’ may also be 
dosage dependent. If so, the intermediate 
crown color of the hybrid may reflect genetic 
heterozygosity. An alternate hypothesis is 
that the red pigments of P. filicauda are ab- 
sent in the hybrid leaving only yellow and 
yellowish-orange pigments in its crown 
feathers. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Acknowledgments 


I thank Miguel Moreno and Marco Ja- 
come for loaning the specimen and for per- 
mission to examine their collections in Qui- 
to, Mark Robbins, Academy of Natural 
Science, Philadelphia, for the loan of com- 
parative material, Alan Brush for discussion 
of pigments, and Richard Banks, Ralph 
Browning, Town Peterson, and Van Rem- 
sen for comments on the manuscript. Trav- 
el was supported by a grant from the Bio- 
diversity Program and the Department of 
Vertebrate Zoology of the National Muse- 
um of Natural History, Smithsonian Insti- 
tution. I thank Victor Krantz for photo- 
graphs. 


Literature Cited 


Brush, A. H. 1969. On the nature of “Cotingin.” — 

Condor 71:47-48. 

1970. Pigments in hybrid, variant and me- 
lanic tanagers (birds). — Comparative Biochem- 
istry and Physiology 36:785-793. 
Graves,G.R. 1990. Systematics of the “green-throat- 

ed sunangels”’ (Aves: Trochilidae): valid taxa or 
hybrids?— Proceedings of the Biological Society 
of Washington 103:6-25. 

Haffer, J. 1967. Speciation in Colombian forest birds 
west of the Andes.— American Museum Novi- 
tates 2294:1-57. 

Hilty, S. L., & W. L. Brown. 1986. A guide to the 
birds of Colombia. Princeton University Press, 
Princeton, New Jersey, 836 pp. 

Hudon, J., A. P. Capparella, & A. H. Brush. 1989. 
Plumage pigment differences in manakins of the 
Pipra erythrocephala superspecies.—Auk 106: 
34-41. 

Parkes, K.C. 1961. Intergeneric hybrids in the family 
Pipridae.— Condor 63:345-350. 

Parsons, T. J., S. L. Olson, & M. J. Braun. 1993. 
Unidirectional spread of secondary sexual 
plumage traits across an avian hybrid zone.— 
Science 260:1643-1646. 

Paynter, R. A., Jr., & M. A. Traylor, Jr. 1977. Or- 
nithological gazetteer of Ecuador. Museum of 
Comparative Zoology, Harvard University, 
Cambridge, Massachusetts, 151 pp. 

Prum, R. O. 1990. Phylogenetic analysis of the evo- 

lution of display behavior in the Neotropical 

manakins (Aves: Pipridae). — Ethology 84:202— 

231. 

. 1992. Syringeal morphology, phylogeny, and 


VOLUME 106, NUMBER 3 


evolution of the Neotropical manakins (Aves: 
Pipridae).— American Museum Notitates 3043: 
1-65. 

Sibley, C. G., & B. L. Monroe, Jr. 1990. Distribution 
and taxonomy of birds of the world. Yale Uni- 
versity Press, New Haven, Connecticut, 1111 
pp. 


Department of Vertebrate Zoology, Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. 20560, 
U.S.A. 


Appendix 


Comparative descriptions of the hybrid, Dixiphia 
pipra X Pipra filicauda, and its parental species (males 
in definitive plumage) (see Fig. 1). 

The forecrown of filicauda is yellow; feathers of the 
crown, hindneck, and upper mantle are pale yellow 
broadly tipped with lustrous orangish-red. In pipra, the 
forecrown, crown and upper hindneck are white; the 
basal fifth of the crown feathers is light gray to medium 
gray. In the hybrid, the forecrown, crown, hindneck, 
and scattered feathers of the upper mantle are golden 
yellow; feathers of the crown have yellowish-orange 
tips and gray bases. 

The mantle, lower back, rump, wings, and tail of 
filicauda, pipra, and the hybrid are glossy black; yellow 
feathers occur along the bend of the wing in filicauda 
(traces of yellow in the hybrid). Inner webs of the inner 
remiges of filicauda possess large white spots concealed 


441 


in the folded wing; remiges of pipra are uniformly black; 
those of the hybrid are black with a narrow pale border 
near the base of the inner web of the secondaries. 

The tail of filicauda is uniquely modified among the 
Piprinae; the barbless rachi of the three outer pairs of 
rectrices extend 30 to 60 mm beyond the tips of the 
tapered vanes. In pipra the tail is slightly rounded; 
feather tips are broad. The tail of the hybrid is shallowly 
forked (11 mm) and intermediate in shape. The central 
rectrices are broadly rounded, the outer rectrices are 
tapered; rachi do not extend beyond the vanes. 

The lores, eye ring, and auriculars are yellow in /fi- 
icauda. In pipra the lores and auriculars are black; the 
white of the crown extends ventrally to the eye ring. 
The lores, subocular area, and auriculars of the hybrid 
are grizzled yellow and black producing a distinctive 
face patch. 

In filicauda the underparts from the throat to the 
vent are yellow; the undertail coverts are black; tibial 
feathers are grayish-black with some yellowish bar- 
bules; underwing coverts are yellowish white. The un- 
derparts including underwing coverts are black in pi- 
pra. The underparts of the hybrid are yellow mottled 
with black. Scattered deposits of melanin can be seen 
under magnification (30 x) in feathers that appear to 
be entirely yellow; other feathers are visibly mottled, 
and some near the sides of the breast are predominately 
black. 

The bills of filicauda, pipra, and the hybrid are black, 
whitish along the tomia (in dried skins). Feet and legs 
of filicauda, pipra, and the hybrid are dark blackish- 
brown (in dried skins). 


PROC. BIOL. SOC. WASH. 
106(3), 1993, pp. 442-449 


A NEW SPECIES OF AQUATIC BUFO 
(ANURA: BUFONIDAE) FROM CLOUD FORESTS IN 
THE SERRANIA DE SIBERIA, BOLIVIA 


Michael B. Harvey and Eric N. Smith 


Abstract.—A new species of the Bufo veraguensis group is described from 
cloud forests in the Serrania de Siberia, Departamento de Cochabamba, Bolivia. 
Full webbing of the feet distinguishes the new taxon from any other New World 
species of Bufo. Morphology and behavior of this species suggest that it is an 
aquatic form of Bufo, the first reported from the Western Hemisphere. Cranial 
crest development increases with body size in Bufo quechua, an undescribed 
species of Bufo, and B. veraguensis. Presence of indistinct, rather than prom- 
inent, parietal crests in the holotype of B. echinoides may be due to incomplete 
development of the crests. Thus, the sole reported characteristic distinguishing 
B. echinoides from B. quechua is not valid. We conclude that B. echinoides is 
a junior synonym of B. quechua. 

Resumen. —Se describe una nueva especie de sapo de la Serrania de Siberia, 
en los Andes de Cochabamba, Bolivia. Pertenece al grupo B. veraguensis y se 
distingue de los demas miembros de éste por tener la membrana interdigital 
basal en las manos y abarcando el 100% en los pies, carecer de las crestas 
supraorbitales, tener crestas parietales lisas, carecer de timpano, poseer glan- 
dulas parotoideas ovoides mas largas que anchas, tener una linea dorsolateral 
de tubérculos, tener el vientre con algunos tubérculos lisos y agrandados (no 
conicos), poseer el primer dedo de la mano mas largo que el segundo. La nueva 
especie es aparentemente la primera especie acuatica de Bufo del hemisferio 
occidental. Diez espécimenes incluyendo dos paratipos de B. quechua y el 
holotipo de B. echinoides no tienen una diferencia morfologica significativa 
entre si. La supuesta ausencia de crestas parietales originalmente usada para 
diagnosticar a B. echinoides muestra ser el resultado de desarrollo incompleto. 
Proponemos la sinonimizacion de B. echinoides bajo B. quechua. 


Toads currently assigned to the Bufo ver- 
aguensis group occur throughout forested 
slopes of the Andes in Peru and Bolivia. 
Although most toads of the B. veraguensis 
group are allopatric (Duellman & Schulte 
1992), four species referable to this group 
occur sympatrically in cloud forests in the 
departments of Cochabamba and Santa 
Cruz, Bolivia. In addition to the widespread 
B. veraguensis and poorly known B. que- 
chua, two other species in these cloud for- 
ests are undescribed. Herein, we describe 
the most distinctive of these two toads, the 


first apparently aquatic species of Bufo from 
the Western Hemisphere. Thereafter, we 
discuss the validity of B. echinoides, a re- 
cently described (Reynolds & Foster 1992) 
taxon from Cochabamba referred to the B. 
veraguensis group. 


Methods 


Field work in the Serrania de Siberia was 
from 5 December 1991 to 15 February 1992. 
We recorded locality information, eleva- 
tion, air temperature, time of day, and mis- 


VOLUME 106, NUMBER 3 


Fig. 1. 


Adult male Bufo amboroensis (MNK 953); 


cellaneous ecological and behavioral notes 
with each specimen. 

In addition to newly collected specimens 
reported here, we also examined the holo- 
type of Bufo echinoides, nine specimens of 
B. quechua including two paratypes, and 17 
specimens of B. veraguensis (Appendix). 
Measurements were made with a dial cali- 
per under a dissecting scope to the nearest 
0.1 mm (or to the nearest mm for specimens 
of B. veraguensis and juvenile specimens of 
B. quechua). Webbing formulae follow those 
of Savage & Heyer (1967) as modified by 
Myers & Duellman (1982); terminologies 
for cranial crests and color descriptions fol- 
low those of Cei (1980) and Smithe (1975), 
respectively. Morphometric characteristics 
examined are snout vent length (SVL), head 
length (HL), head width (HW), interorbital 
distance (ID), dorsal eyelid width (EW), eye 
diameter (ED), eye-nostril distance (EN), 
paratoid width (PW), paratoid length (PL), 
tibia length (TL), and foot length (FL). Col- 
lection abbreviations refer to the Carnegie 
Museum of Natural History (CM); Museo 
de Historia Natural ““Noel Kempff Merca- 
do,’ Santa Cruz, Bolivia (MNK); Univer- 


443 


SVL = 37.1 mm. 


sity of Michigan, Museum of Zoology 
(UMMZ); United States National Museum 
of Natural History (USNM); and the Uni- 
versity of Texas at Arlington Collection of 
Vertebrates (UTA). 


Description of New Species 


Bufo amboroensis, new species 
Fig. 1 


Holotype.—Museo de Historia Natural 
“Noel Kempff Mercado,”’ Santa Cruz, Bo- 
livia, (MNK) AM-953, adult male, collected 
8 Jan 1992 by Michael B. Harvey and Eric 
N. Smith from a small stream 12.7 km by 
road E of El Enpalne along road to Khara 
Huasi, Provincia Carrasco, Estado Cocha- 
bamba, Bolivia, 2150 meters. 

Paratype. —UTA A-39337, an adult male 
collected with the holotype. 

Diagnosis.— Bufo amboroensis may be 
distinguished from all other species of Bufo 
in the Western Hemisphere by its large, 
completely webbed feet. It may further be 
distinguished from all other Bolivian spe- 
cies of Bufo by the combination of (1) su- 
praorbital crests absent (2) parietal crests 


444 


weak (3) external tympanum absent (4) lat- 
eral row of conspicuously enlarged tubercles 
present on body (5) some ventral body tu- 
bercles slightly enlarged and elevated (not 
conical), but venter mostly smoothly areo- 
late (6) hands webbed basally (7) first finger 
longer than second. 

All four species of Bufo occuring in cloud 
forests of the Serrania de Siberia lack a tym- 
panum and have extensively webbed feet. 
However, webbing does not extend over the 
tips of the toes in any of these, except B. 
amboroensis. The venter of B. amboroensis 
is relatively smooth and completely lacks 
conical tubercles, while numerous conical 
tubercles cover the venters of B. quechua, 
B. veraguensis, and an undescribed species 
of Bufo. In addition, B. veraguensis and B. 
quechua have much longer fingers than does 
B. amboroensis. Bufo amboroensis also lacks 
supraorbital crests, which further distin- 
guishes it from B. veraguensis and the un- 
described species of Bufo. 

Description of holotype.—Body robust; 
head slightly wider than long (HW:HL 1.07); 
head length occupying 32% of SVL; snout 
subacuminate in dorsal view, rounded and 
slightly sloping in profile (Fig. 2); supraor- 
bital crests absent; parietal crests weak; nos- 
trils not protuberant, directed laterally; can- 
thus rostralis rounded; loreal slightly 
concave; interorbital distance greater than 
dorsal eyelid width (ID:EW = 1.10) and 
considerably greater than eye-nostril dis- 
tance (ID:EN = 1.50), but slightly less than 
eye diameter (ID:ED = 0.93); lips rounded 
with V-shaped notch at symphysis of upper 
jaw; rostral keel present. Mostly smooth 
paratoids ovoid, longer than wide (PL:PW 
= 1.14), with some diffuse tubercles later- 
ally; paratoid separated from caudal margin 
of dorsal eyelid by glandular postorbital crest 
in contact with paratoid. 

Arms short and slightly robust; fingers 
relatively short (Fig. 3); relative lengths of 
fingers 3 > 1 > 4 > 2; webbing fleshy and 
tuberculate, extending as fringe on lateral 
edges of digits; hands webbed basally; web- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


bing formula I2'4-2'2112-31I13-3IV; nuptial 
excrescence extensive, covering medial sur- 
face of slightly enlarged prepollex and ex- 
tending onto dorsal surface of digit 1, also 
covering dorsal surface of digit 2, and me- 
dial surfaces of digit 2 and distal one-half 
of 3; low palmer tubercles large and ovoid, 
twice size of ovoid, elevated pollical tuber- 
cles; low subarticular tubercles round, paired 
or bifid on third and fourth fingers, simple 
on others; slightly raised supernumery tu- 
bercles smaller than subarticular tubercles. 

Legs long and slender, robust; foot length 
shorter than tibial length (TL:FL = 1.01); 
relative lengths of toes 4 > 5=3 >2> 1; 
toes completely webbed; broad fringe ex- 
tending distally from center of inner meta- 
tarsal tubercle to tip of digit one, broad fringe 
also along lateral border of digit five; web- 
bing enclosing tips of each digit; webbing 
formula IO-OII0-OITIO-OIVO-OV; tarsal fold 
present as row of tubercles; inner metatarsal 
tubercle elongate, twice as long as ovoid 
outer metatarsal tubercle; indistinct subar- 
ticular tubercles round and paired, bifid, or 
simple, apparently without regularity; in- 
distinct supernumery tubercles smaller than 
subarticular tubercles. 

External tympanum absent; choanae 
small, ovoid, and widely separated; elon- 
gate, ovoid tongue twice as long as wide, 
rounded posteriorly, free along one-third of 
its posterior length; vocal slits absent; skin 
of dorsal body, head, and limbs covered in 
rounded, spinous tubercles, evenly spaced, 
forming conspicuously enlarged, lateral row; 
smaller conical tubercles forming subman- 
dibular and labial rows; gular and ventral 
body surfaces mostly smooth with very few 
low tubercles, more on limbs; venter areo- 
late. 

Color in preservative (alcohol after buf- 
fered formalin): Dorsum olive to glaucous; 
tubercles olive gray, bordered or not in 
blackish neutral gray; smoke gray vertebral 
stripe extending from posterior tip of coccyx 
to interorbital region; two interorbital sub- 
triangular bars, the first olive, the second 


VOLUME 106, NUMBER 3 445 


Fig. 2. Dorsal and lateral views of the head of Bufo amboroensis (MNK 953); HL = 12.0 mm. 


446 


> 4 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Palmar and plantar hand and foot of Bufo amboroensis (UTA A-39337); scale equals 5 mm. 


blackish neutral brown; broken blackish 
neutral gray stripe just ventral to lateral row 
of tubercles; smoke gray venter grading to 
flesh color at throat; limbs olive to glaucous 
with blackish neutral gray bands dorsally. 
Variation.— The paratype differs from the 
holotype in relatively few characteristics. 
The foot is longer than the tibia (TL:FL = 


0.90 vs. 1.01), a tarsal fold is present as a 
row of enlarged tubercles on the left but 
absent on the right tarsus, a dorsolateral row 
of enlarged tubercles is present though con- 
siderably less conspicuous, and the para- 
toids are relatively much longer (PL:PW = 
1.75 vs. 1.14). 

The dorsum of the paratype is brownish 


VOLUME 106, NUMBER 3 


olive to glaucous with smoke gray tubercles, 
the tubercles are not edged in darker pig- 
ment as in the holotype, a vertebral stripe 
is absent, the dorsal surface of the limbs is 
brownish olive with diffuse blackish neutral 
gray bands, and the venter is glaucous with 
some flesh color on the throat. 

Measurements: Measurements of the ho- 
lotype are followed by those of the paratype 

in parentheses. SVL 37.1 (38.5); HL 12.0 
(12.3); HW 12.8 (12.1); ID 4.2 (4.1); EW 
3.8 (3.7); EN 2.8 (3.0); ED 4.5 (4.2); PL 4.2 
(5.6); PW 3.7 (3.2); TL 16.0 (16.3); FL 15.9 
(18.1). 

Comment. — Based on external character- 
istics, we tentatively assign Bufo amboroen- 
sis to the B. veraguensis group sensu Duell- 
man & Schulte (1992). With other members 
of the B. veraguensis group, B. amboroensis 
shares extensive webbing of the feet, re- 
duced cranial crests, a lateral row of tuber- 
cles on the body, absence of an external 
tympanum, and the first finger longer than 
the second. However, B. amboroensis dif- 
fers from all other species in this group by 
having relatively large tubercles on the dor- 
sum. 

Duellman & Schulte (1992) considered 
the monophyly of the B. veraguensis group 
highly suspect, a view supported by the oc- 
currence in sympatry of four species refer- 
able to this group in the Serrania de Siberia. 

Distribution and ecology.— Bufo ambor- 
oensis is known only from the type locality, 
a northeast facing slope overlooking the Rio 
Chua Khocha, 12.7 km NW of El Enpalne, 
Provincia de Carrasco, Departamento de 
Cochabamba: 17°50'31”S, 64°45'18’W. 
Slopes in the area are covered in cloud for- 
est, parts of which have been altered by se- 
lective logging of the largest trees, probably 
more than 20 years ago. 

Both specimens were found together at 
1340 hr on a warm sunny day (air temper- 
ature 20°C) at 2150 m. Both were motion- 
less on the bottom of a clear stream (water 
temperature 15°C) about a meter deep and 
two meters wide and in a stretch of rela- 


447 


tively little current where the stream 
emerged from cloud forest and formed a 
pool before passing under a dirt road. We 
approached the stream slowly and did not 
see either specimen jump into the water, 
suggesting that the toads were found in an 
undisturbed state. 

Extensive webbing of the feet and loca- 
tion of the toads on the bottom of a stream 
suggests an aquatic existence. Other than its 
obvious utility for swimming, the webbing 
may also play a role in respiration by in- 
creasing the surface area of skin similar to 
the flaps of skin present on some perma- 
nently aquatic, high elevation anurans such 
as Telmatobius culeus (Hutchinson et al. 
1976). 

Etymology.—The specific epithet is an 
adjective for the wildlife preserve in which 
Bufo amboroensis occurs: Parque Nacional 
Amboro. 


Taxonomic Status of Bufo echinoides 


Bufo echinoides was described recently 
(Reynolds & Foster 1992) based on a single 
male specimen (SVL 26.5 mm) from the 
Chapare region of Cochabamba. The au- 
thors assigned this species to the B. vera- 
guensis group and remarked that it differed 
from B. quechua Gallardo, 1961 by lacking 
cranial crests. However, both in and among 
three species of the B. veraguensis group that 
we examined, this characteristic varies with 
body size. 

Cranial crests are barely visible in an adult 
male (SVL = 31.7 mm) representing an un- 
described species of the Bufo veraguensis 
group from the Serrania de Siberia, and not 
visible in either of two juveniles of this un- 
described species we collected (SVL = 19.0, 
13.5 mm). However, a complete comple- 
ment of cranial crests are well developed in 
the largest specimens, a male (SVL = 41.6 
mm) and female (SVL = 53.6), of this spe- 
cies. Similarly, in B. veraguensis we exam- 
ined (n = 17), cranial crests are absent in 
specimens smaller than 31 mm SVL (” = 


448 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Morphometric characteristics of Bufo quechua, and B. echinoides. Abbreviations are listed in text. 


SVL HW:HL 

Bufo quechua 

CM 4223 54.8 1.16 

CM 4224 33,5 0.89 

UMMZ 68166 19.3 0.86 

UMMZ 76075 43.0 Ibe | 

UMMZ 89414 44.2 107 

UMMZ 172542 50.6 las 

UTA A-39338 39.8 1.16 
Bufo echinoides 

USNM 257799 26.5 1.10 


11) and increase in prominence with in- 
creasing size. These observations suggest 
that this character should be used with cau- 
tion when diagnosing species in the B. ver- 
aguensis group or when using keys employ- 
ing cranial crests as diagnostic characters 
such as that provided by Duellman & 
Schulte (1992). 

We were unable to find any significant 
morphological differences between the ho- 
lotype of Bufo echinoides and specimens of 
B. quechua. Some cranial crests (postorbi- 
tal, canthal, orbitotympanic) are well de- 
veloped in B. echinoides and all specimens 
of B. quechua examined. Parietal crests were 
not visible in juvenile specimens of B. que- 
chua (SVL = 15-19 mm; n = 4) but were 
visible in larger specimens (SVL = 33.5— 
50.6 mm; n = 6). The holotype of B. echi- 
noides is intermediate in size relative to these 
two groups. Contrary to the report by Reyn- 
olds & Foster (1992:87), we find that pari- 
etal crests, though poorly developed, are 
visible in the holotype of B. echinoides, with 
the left parietal crest being slightly more 
pronounced than the right. 

Morphometric ratios of Bufo echinoides 
(Table 1) lie within the range of ratios for 
specimens of B. quechua or are only slightly 
lower (PL:PW) than the ratios of a male 
paratype (CM 4224). Similarly, character- 
istics of the arms, legs, hands, and feet, in- 
cluding webbing formulae, relative lengths 


ID:EW 


ID:ED ID:EN PL:PW TL:FL 
07, 1.43 LJ 0.88 
0.93 1.86 1.50 1.03 
1.00 1.33 1.70 0.96 
LESS 1.67 Jie} 0.85 
19 eo 2.30 0.95 
1232 eral 1.58 0.97 
0:92 1:67 192 0.88 
1.06 1.42 1.48 1.02 


of digits, and condition of tubercles, are not 
significantly different in eight specimens in- 
cluded in Table 1. All eight specimens also 
possess the characteristic conical, spinous 
tubercles that cover both the dorsum and 
venter and also form an enlarged row on 
the dorsolateral aspect of the body. In both 
B. echinoides and B. quechua, morphology 
of the tongue and choanae are also very sim- 
ilar. Differences or similarities in color pat- 
tern could not be assessed because the colors 
of most specimens of B. quechua had faded. 

The type series of Bufo quechua was col- 
lected at Incachaca, Department of Cocha- 
bamba, at 2500 m. Although we were un- 
able to find this locality on any maps of 
Bolivia, it presumably lies within the ““Yun- 
gas of the Department of Cochabamba” 
(Gallardo 1961:6), the same region and ap- 
proximate elevation (456 meters higher) 
where B. echinoides was collected. Reynolds 
& Foster (1992) did not report collecting 
any specimens of B. quechua, nor did they 
mention that specimens of this taxon were 
compared directly with the holotype of B. 
echinoides. Because Bufo echinoides lacks 
any significant characteristics that distin- 
guish it from paratypes of B. quechua, and 
the two taxa come from about the same 
geographic area and elevation, we propose 
that Bufo echinoides Reynolds & Foster, 
1992 is a junior synonym of Bufo quechua 
Gallardo, 1961. 


VOLUME 106, NUMBER 3 


Acknowledgments 


We thank A. G. Kluge, C. J. McCoy, and 
R. P. Reynolds for loan of specimens ex- 
amined in this study. We thank J. A. Camp- 
bell for reviewing the manuscript and for 
the enormous amount of assistance and 
guidance he has provided throughout our 
studies on Bolivian reptiles and amphibi- 
ans. Without the support and guidance of 
J. A. Campbell and E. D. Brodie Jr. our 
research in Bolivia would have been im- 
possible. Many people in Bolivia aided us 
in the field and during the process of ob- 
taining a permit for export of the specimens. 
We especially wish to acknowledge logisti- 
cal support afforded us by H. Justiniano and 
A. Castillo of the Fundacion Amigos de la 
Naturaleza. We thank members of the Bo- 
livian Consejo Nacional, as well as I. Pi- 
naya, Dpto. Vida Silvestre, MACA; and M. 
Avalos, Dpto. Vida Silvestre, UTD-CDF 
SC, for granting us permission to export 
specimens and for their patience and kind 
understanding throughout the permit pro- 
cess. We acknowledge the kind assistance 
during the permit process afforded us by P. 
Bettella, T. Centurion, and N. Vacas of the 
Museo Noel Kempff Mercado, as well as E. 
Forno, of Fondo Nacional Para El Medio 
Ambiente, and P. Ergeta, of the Coleccion 
Nacional de Fauna. We also express our 
thanks to our good friends Barbara and Ian 
Phillips of the El Refugio project for their 
encouragement and logistical support. Fi- 
nally, we thank H. Centeno and K. Casta- 
neda for their logistical and editorial assis- 
tance. 


Literature Cited 


Cei, J. M. 1980. Amphibians of Argentina.—Moni- 
tore Zoologico Italiano, New Series 2:1-609. 

Duellman, W. E., & R. Schulte. 1992. Description of 
a new species of Bufo from northern Peru with 
comments on phenetic groups of South Amer- 


449 


ican toads (Anura: Bufonidae).—Copeia 1992: 
162-172. 

Gallardo, J. M. 1961. Three new toads from South 
America: Bufo manicorensis, Bufo spinulosus al- 
tiperuvianus and Bufo quechua.—Breviora 141: 
1-8. 

Hutchinson, V. H., H. B. Haines, & G. Engbretson. 
1976. Aquatic life at high altitude: respiratory 
adaptations in the Lake Titicaca frog, Te/ma- 
tobius culeus.— Respiration Physiology 27:115-— 
129. 

Myers, C. W., & W. E. Duellman. 1982. A new spe- 
cies of Hyla from Cerro Colorado, and other 
tree frog records and geographical notes from 
western Panama.—American Museum Novi- 
tates 2752:1-32. 

Reynolds, R. P., & M. S. Foster. 1992. Four new 
species of frogs and one new species of snake 
from the Chapare region of Bolivia, with notes 
on other species. — Herpetological Monographs 
6:83-104. 

Savage, J. M., & W. R. Heyer. 1967. Variation and 
distribution in the tree-frog genus Phyllomedusa 
in Costa Rica, Central America.—Beitrage zur 
Neotropischen Fauna 5:111-131. 

Smithe, F. B. 1975. Naturalist’s color guide. The 
American Museum of Natural History, New 
York, 16 pp. 


Department of Biology, UTA Box 19498, 
The University of Texas at Arlington, Ar- 
lington, Texas 76019-0498, U.S.A. 


Appendix 
Specimens Examined 


Specimens examined are followed in parentheses by 
their SVL in mm. Bufo amboroensis MNK 953 (37.1) 
holotype, UTA A-39337 (38.5) paratype, B. echinoides 
USNM 257799 (26.5) holotype, B. gquechua CM 4223 
(54.8) paratype, CM 4224 (33.5) paratype, UMMZ 
68163 (3 specimens, 15—18 mm), UMMZ 68166 (19.3), 
UMMZ 76075 (43.0), UMMZ 89414 (44.2), UMMZ 
172542 (50.6), UTA A-39338 (39.8), B. sp. MNK 950 
(41.6), MNK 951 (19.0), MNK 952 (13.5), UTA 
A-39335 (53.6), UTA A-39336 (31.7), B. veraguensis 
MNK 954 (33), MNK 955 (38), MNK 956 (45), MNK 
957 (22), MNK 958 (21), MNK 959 (23), MNK 960 
(32), UTA A-39296 (30), UTA A-39309 (31), UTA 
A-39322 (48), UTA A-39324 (20), UTA A-39325 (20), 
UTA A-39327 (22), UTA A-39331 (18), UTA A-39332 
(17), UTA A-39333 (45), UTA A-39334 (20). 


PROC. BIOL. SOC. WASH. 
106(3), 1993, pp. 450-454 


REVISION OF YLASOIA SPEISER, 1920 
(INSECTA: DIPTERA: BOMBYLITDAE: LOMATIINAE) 


Marcia Souto Couri and Carlos José Einicker Lamas 


Abstract. — Ylasoia Speiser, 1920 is redescribed and the known neotropical 
species, are revised, based on examination of the types. Y. abbreviata (Wie- 
demann), 1830 and Y. caloptera (Macquart), 1834 are synonymized with Y. 
pegasus (Wiedemann), 1828, the type species of the genus. 


Speiser (1920) erected the genus Ylasoia 
to include three south neotropical species: 
Y. abbreviata (Wiedemann), 1830, Y. cal- 
optera (Macquart), 1834 and Y. pegasus 
(Wiedemann), 1828. The generic name was 
proposed as an anagram of the Australian 
genus Alysoia Rondani, 1863, one of the 
genera segregated from Comptosia Mac- 
quart, 1840 by Rondani (1863), based 
mainly on wing venation—number of sub- 
marginal cells and cross veins (both genera 
have two sub-marginal cells). Edwards 
(1930) stated that too much importance was 
placed on this character and that the genera 
previously included in Comptosia should be 
reunited. Speiser (1920) nevertheless char- 
acterized Y/asoia on other characters, com- 
pared it with A/lysoia, and mentioned that 
these two genera were probably closely re- 
lated. 

Bowden (1971), Hull (1973), and Even- 
huis (1980) have discussed the classification 
of Comptosia group. The first and the last 
papers treated the Australian genera. 

Hull (1973) gave a detailed description of 
Y/asoia and stated that the genus “‘seems to 
be nearer the Australian genus Oncodosia 
Edwards, than to the South American gen- 
era like Lyophlaeba Rondani.” He exam- 
ined a series of Y. pegasus from Brazil and 
according to him, two of the three Ylasoia 
species may perhaps be subspecies. 

Since the original description, no more 
species have been included or described in 
this genus. 


Painter & Painter (1974) made notes and 
redescriptions on the types of the three spe- 
cles. 

The study of a material of Y/asoia spp. 
deposited at Museu Nacional (Rio de Ja- 
neiro) collection, showed that some specific 
characters cited in literature vary greatly 
when a long series is examined. 

Examination of the types and direct com- 
parisons with material at hand enabled a 
revision of the species and redescription of 
the genus. 


Ylasoia Speiser 


Ylasoia Speiser, 1920:213-214.—Hull. 
1973:307, 356.— Painter & Painter, 1974: 
107.—Painter et al., 1978:29. 


Type species. —Anthrax pegasus Wiede- 
mann, 1828 (orig. desig.) 

Redescription. —Length, 12-17 mm. 
Head: Eyes holoptic in male, separated at 
anterior ocellar level by a distance about 
twice ocellar diameter; face convex, slightly 
pronounced in profile; scape rectangular, 3— 
4 times the length of pedicel, with dark 
brown pile, pedicel globular, short, with pile 
as in scape; flagellomere long, bare, 5—6 times 
the length of pedicel; palpus broad basally, 
1-segmented; proboscis dark brown, short, 
not extended beyond oral margin. Thorax: 
Scutum velvety black, with two brown lon- 
gitudinal stripes anteriorly extended to 0.75 
of it, bare dorsally, and with black bristles 
laterally; scutellum velvety black, with some 


VOLUME 106, NUMBER 3 


black lateral bristles; pleura light brown, ka- 
tepisternum with bristles in upper third; 
anepimeron with pile in apical third of in- 
ferior half; meron bare; metepimeron with 
pile in the upper extremity near posterior 
spiracle; halter dark brown, long; knob yel- 
low at dorsal apex and totally yellow ven- 
trally. Legs: Brown with black bristles and 
few black scales; hind femur at anteroven- 
tral surface with 4—S bristles at apical third; 
claws dark brown; pulvilli 0.65 from the 
length. of claws. Wing: Brown with an an- 
terior white and posterior hyaline sub-me- 
dian transverse band, other white or hyaline 
variable areas in cells; anal and axillary cells 
with a big hyaline area, (Some males with 
darker marks) (Fig. 1); 2 sub-marginal cells 
({R; and R,); 3rd posterior cell (2nd M,) 
narrowed marginally for a variable distance; 
fringe of wing brown, longer and darker at 
anterior margin, specially near base. Ab- 
domen: Long, as wide as thorax; tergites 
concolorous with thorax; sparse pile dor- 
sally and more evident laterally. 

Female: Similar to male, except for eye 
distance, which is about 3 times ocellar di- 
ameter. 


Ylasoia pegasus (Wiedemann) 


Anthrax pegasus Wiedemann, 1828:298.— 
Walker, 1849:264.—Speiser, 1920:214.— 
Hull, 1973:356.— Painter & Painter, 1974: 
42: 

Anthrax abbreviata Wiedemann, 1830: 
637.—Osten-Sacken, 1887:140.—Al- 
drich, 1905:228.—Speiser, 1920:215.— 
Painter & Painter, 1974:109. 

Lomatia caloptera Macquart, 1834:612.— 
Speiser, 1920:215. 

Comptosia caloptera Macquart, 1834:412.— 
Painter & Painter, 1974:110. 

Ylasoia pegasus Speiser, 1920:214.—Hull, 
1973:358, figs. 148, 367, 659, 670, 985, 
986, 987.—Painter et al., 1978:29. 

Ylasoia abbreviata Speiser, 1920:215.— 
Painter & Painter, 1974:109-110.— 
Painter et al., 1978:29. 


451 


Ylasoia caloptera Speiser, 1920:215.— 
Painter & Painter, 1974:110-112.— 
Painter et al., 1978:29. 

Ylasoia pegasa Painter & Painter, 1974:112- 
aS. 


Holotype. —(Fig. 2). Female: A. pegasus 
m./ Mus. rog. Berol. [handwritten label]. 
Anthrax pegasus Wiedemann. Holotype 
[rose label]. Pegasus/ coll. Wiedem. (Fig. 3). 
Deposited at Naturhistorishes Museum, 
Wien. The specimen lacks both flagello- 
meres, both fore legs, left middle leg and 
the tarsi of the hind leg. 

Diagnosis. — Wing with the white/hyaline 
transverse band reaching posterior margin 
or not; hyaline areas at anal and axillary 
cells with variable extensions, sometimes 
faint, specially in some males, elongated hy- 
aline area distal half of second basal cell 
present or not; other hyaline areas variable. 
(Figs. 1, 2, 5 and 7). Male genitalia (Fig. 4) 
in lateral view with basistylus linear ovate; 
dististylus long, hooklike apically; aedeagus 
sclerotized, large in base; epandrium tri- 
angular; cerci long. 

Material examined. —(deposited at Mu- 
seu Nacional, Rio de Janeiro): BRAZIL. 
Minas Gerais: Pocos de Caldas, Morro S. 
Domingos, 1 2, 15 Sep 1968, J. Becker, O. 
Roppa e O. Leoncini; Morro do Ferro, 3 4, 
29-30 Mar 1964, 3 éand 1 2, 27 Jan 1965, 
1 éand 3 2, 22 Mar 1966, 1 6, 24 Mar 1966, 
J. Becker, O. Roppa e O. Leoncini; Cam- 
buquira, 2 2, Dec 1933, A. Marques, | 2, 
Feb 1942, H. S. Lopes, Cambuquira, 1 4, 
Feb 1941, Lopes & Gomes; Serra do Caraca, 
Santa Barbara, 1 6, Feb 1976, H. S. Lopes. 
Sao Paulo: Ypiranga, 2 2 e 2 6, Mar 1936, 
Santos, Leme, Fazenda Graminha, | 2, Al- 
ceu e Santos; Rio Grande do Sul: Pelotas, 
1 9, Mar 1957, C. Biezanko (MNRJ); Rio 
de Janeiro: Friburgo, 2 6 and 2 2, Feb 1933, 
C.F.M.L., 2 6, Feb 1932, Prof. M.-L.; Nova 
Friburgo (900M), 2 6, Jan 1946, Wigod., 
Itatiaia, 1.2; 18 Feb 1933, I 4, Feb 1959, 
W. Zikan; Itaguay, Serra da Caveira 600M, 
1 6 and 1 9, 25 Feb 1948, W. Zikan; Pe- 


452 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


co 
Coil... Wiedetie 


as 1 ee 


L727 203 Pidcl 
sf 
Fic cop bx A 


Calo 


oe 
Det. R. H. Pain 


@HoLoTYfE 2 
3 Ce mptosca 


caloptera Macquart 
det. J.E. Chainey, 1987. 


0,5 


Figs. 1-8. 1-4, Ylasoia pegasus, 1, Wing of holotype; 2, holotype; 3, labels of holotype; 4, male genitalia; 
5-6, Ylasoia abbreviata, 5, holotype; 6, labels of holotype; 7-8, Ylasoia caloptera, 7, holotype; 8, labels of 
holotype. 


VOLUME 106, NUMBER 3 


tropolis, Le Vallon, Alt. Mosella, 1 4, 24 
Jan—23 Feb 1958, D. Albuquerque. 

Comments. — Although some characters, 
such as the position of crossvein r-m in re- 
lation with discal cell; length of ending of 
first posterior, anal and third posterior cells 
and width of axillary and anal cells, men- 
tioned by Painter & Painter (1974) as di- 
agnostic ones were observed in the types of 
the three species, considerable variation is 
evident when a series is examined. In ad- 
dition, the examined specimens show a 
combination of other characters mentioned 
as distinct, especially the patterns of wing 
coloration. 

Hull’s (1973:495, fig. 367) illustration of 
Y. pegasus wing shows also another pattern 
of coloration, differing from the holotype. 
On the other hand, the male genitalia as 
illustrated in fig. 985 (p. 568) is very similar 
in all material we dissected, and there is no 
reason to treat them as separate species. 

The holotypes of Y. abbreviata and Y. 
caloptera (Fig. 5 and Fig. 7), here synony- 
mized with Y. pegasus, were also examined: 


Ylasoia abbreviata: Male. “Brasil V. Olf’ 
[Brasil, von Olfers] / abbreviata Wied. 
[handwritten labels]. Typus [red label]. 
Zool. Mus. Berlin. 1612. (Fig. 6). Depos- 
ited at Berlin Museum. The specimen is 
in good condition except abdominal seg- 
ments beyond the sixth are missing. Wie- 
demann (1830) incorrectly specifies Mex- 
ico as the type locality. 

Ylasoia caloptera: Male. Comptosia / fas- 
cipennis / 6 Macq. [handwritten labels]. 
Brazil ? / ex Bigot coll. / BM 1960 - 539. 
? HOLOTYPE ? / Comptosia / caloptera 
Macquart / det. J. E. Chainey, 1987 
[handwritten labels]. Y/asoia / caloptera 
/(Mq.)/ Det. R. H. Painter [19]60 [hand- 
written labels; this label with left wing 
glued to corner]. Type ? [red circle label]. 
Holotype ? [red circle label]. (Fig. 8). De- 
posited at The Natural History Museum, 
London. The specimen lacks its head, the 
fore and middle pair of legs, and part of 
the thorax has been eaten. 


453 


Acknowledgments 


Weare very grateful to Dra. R. Contreras- 
Lichtenberg (Naturhistorishes Museum, 
Wien), Dr. H. Schumann (Museum fur Na- 
turkunde der Humboldt—Universitat, Ber- 
lin) and Dr. J. Chainey (The Natural His- 
tory Museum, London) for the loan of type 
material, to Dr. Neal Evenhuis (Bishop Mu- 
seum, Honolulu) for the review of the 
manuscript and also to Dr. S. Fragoso (Mu- 
seu Nacional, Rio de Janeiro) for the pho- 
tographs. 


Literature Cited 


Aldrich, J. M. 1905. Catalogue of North American 
Diptera.—Smithsonian Miscellaneous Collec- 
tions, XLVI:679 pp. 

Bowden, J. 1971. Notes on some Australian Bom- 
byliidae in the Zoological Museum, Copenha- 
gen (Insecta, Diptera).—Steenstrupia 1:295-—307. 

Edwards, F. W. 1930. Bombyliidae, Nemestrinidae, 
Cyrtidae, in British Museum (Natural History), 
Diptera of Patagonia and South Chile 5(2):162- 
197, London. 

Evenhuis, N.L. 1980. Studies in Pacific Bombyliidae 
(Diptera) V. Notes on the Comptosia group of 
the Australian region, with Key to genera and 
descriptions of new genus and three new spe- 
cies.— Pacific Insects 21(4):328-334. 

Hull, F. M. 1973. Beeflies of the world. The genera 
of the family Bombyliidae. — United States Na- 
tional Museum Bulletin 286:1-687. 

Macquart, J. 1834. Histoire naturelle des Insectes. 
Diptéres. Tome premier. Diptera 1:578 pp., in 
N. E. Roret, ed., Collection des Suites a Buffon. 
Paris. 

Osten-Sacken, C. R. 1887. Diptera, Vol. I. Pp. 129- 
160, 161-176, 177-208, 209-216, pl. 3, in F. 
D. Godman & O. Salvin, eds., Biologia Centrali 
Americana, Zoologia-Insecta-Diptera 1:378 pp., 
6 pls. London. 

Painter, R. H., & Painter, E.M. 1974. Notes on, and 
redescriptions of, types of South American 
Bombyliidae (Diptera) in European and United 
States Museums.— Research Publications Kan- 
sas State University Experiment Station 168:1- 
eee DB 


,&J.Hall. 1978. Family Bombyliidae: 
1-92. A catalogue of Diptera of the Americas 
south of the United States. Departamento de 
Zoologia, Universidade de Sao Paulo. 

Rondani, A. C. 1863. Diptera exotica revisa et an- 
notata.— Modena, 99 pp. 

Speiser, P. 1920. Zur Kenntnis der Diptera Orthor- 


454 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


rhapha Brachycera.—Zoologische Jahrbucher 
(Abteilung fur Systematik) 43:195-220. 
Walker, F. 1849. List of the specimens of dipterous 
insects in the collection of the British Museum 
2:23 1-484; 4:689-1172. London. 
Wiedemann, C. R. W. 1828. Aussereuropaische 
zweifliigelige Insekten 1:xxxi1 + 608 pp., 7 pls. 
. 1830. Aussereuropaische zweifliigelige Insek- 
ten 2:XII + 684 pp., 5 pls. 


Departamento de Entomologia, Museu 
Nacional, Universidade Federal do Rio de 
Janeiro, Quinta da Boa Vista, Sao Cristo- 
vao, Rio de Janeiro, RJ. CEP 20.940-040 
Brasil. 


PROC. BIOL. SOC. WASH. 
106(3), 1993, pp. 455-466 


NEW RECORDS OF ENTOCYTHERID OSTRACODS 
INFESTING BURROWING AND CAVE-DWELLING 
CRAYFISHES, WITH DESCRIPTIONS OF 
TWO NEW SPECIES 


Horton H. Hobbs, Jr., and Daniel J. Peters 


Abstract.—The ranges, including several scores of new localities, of 15 en- 
tocytherid ostracods infesting burrowing and cave-dwelling crayfishes collected 
in the southeastern United States (most from Kentucky, Tennessee, and West 
Virginia) are summarized. The genera represented are Ascetocythere, Cymo- 
cythere, Dactylocythere, Donnaldsoncythere, Lordocythere, Phymocythere, and 
Uncinocythere. The new Dactylocythere cryptoteresis and Phymocythere lophota 
are described from Upshur and Wirt counties, West Virginia, respectively. 


This study of entocytherids associated 
with burrowing and cave-dwelling crayfish- 
es was prompted by the receipt of some 50 
lots of ostracods from Raymond F. Jezeri- 
nac and G. Whitney Stocker, of The Ohio 
State University at Newark. The specimens 
were retrieved by them from crayfish col- 
lections made from burrows and caves in 
Kentucky and West Virginia. Augmenting 
these lots are a large number of samples 
provided us previously by Raymond W. 
Bouchard, of the Academy of Natural Sci- 
ences of Philadelphia. This report consists 
of summaries of the distributions of the 15 
entocytherid species that were obtained in 
the collections made by Jezerinac and 
Stocker; included also are new locality re- 
cords provided by Bouchard and others. 
Two of the species included are previously 
undescribed, Dactylocythere cryptoteresis 
and Phymocythere lophota, both from West 
Virginia. 

As has been pointed out in many previous 
entocytherid studies, most collections have 
been made by students of crayfishes and 
usually all of the specimens collected in a 
locality were preserved together in a single 
container. Consequently only in instances 
in which a single crayfish species was col- 
lected at a locality can one be certain as to 


the host of the ostracods found in the con- 
tainer. Thus all of the crayfishes listed under 
‘“Hosts’’ below should be considered “‘pos- 
sible hosts.”’ Definitely established associ- 
ations with hosts are marked by asterisks 
(7): 

The synonymies presented include cita- 
tions to the original description, and update 
the synonymies and references cited by Hart 
& Hart (1974) or Hobbs & Peters (1977). 
Among the abbreviations used in listing the 
localities are: Ck = creek, R = river, US 
Hwy = U.S. Highways, Rte = State High- 
ways or Routes, Co. Rd = County Roads, 
cos = counties. In citing the hosts, subge- 
neric names are omitted as are citations to 
their authors and dates, all of which are list- 
ed in a recent checklist of American cray- 
fishes (Hobbs 1989). 


Ascetocythere myxoides Hobbs & Hart 


Ascetocythere myxoides Hobbs & Hart, 
1966:45-46, figs. 21-23.—Hart & Hart, 
1974:42-43, pl. VIII, figs. 1-3, pl. X_LVII. 


Previously known range. — Reported from 
the Cheat and Potomac basins from only 2 
localities in Randolph County, West Vir- 
ginia (type locality), and Prince Georges 


456 


County, Maryland. Subsequent collecting in 
the latter locality failed to disclose the pres- 
ence of either the entocytherid or its host, 
and inasmuch as neither has been reported 
to occur in the piedmont or coastal plain 
section of Maryland and neighboring states, 
the latter locality must be questioned. 

New locality records. —West Virginia: 
Grant Co., 4.1 airmi (6.6 airkm) SW of Stre- 
by on Nat. Forest Rd, 18 Aug 1985, R. F. 
Jezerinac, on Cambarus b. bartonii & C. 
monongalensis. Preston Co., Ditch 1.0 mi 
(1.6 km) N of Brandonville, 10 Sep 1984, 
G. W. Stocker, on C. dubius. Pocahontas 
Co., Roadside ditch and seep 1.0 mi (1.6 
km) S of Thornwood on US Hwy 250, 23 
Jun 1987, GWS, RFJ, on C. monongalensis 
and C. bartonii carinirostris. Randolph Co., 
E shore of Shavers Fk of Cheat River in 
Monongahela Nat. Forest, 28 Jul 1969, C. 
Adler, S. Arnold, on C. monongalensis. 
Pennsylvania: Greene Co., 1.8 airmi (2.9 
airkm) NE of Crabapple, 15 Sep 1984, GWS, 
V. Stocker, on C. monongalensis. 

Hosts: Cambarus b. bartonii, C. b. cari- 
nirostris, C. dubius*, and C. monongalen- 
sis*. This entocytherid is probably restrict- 
ed to the last two species. 

Drainage systems.—Cheat and Guyan- 
dotte basins (to Ohio and Mississippi) and 
perhaps the Potomac basin. 


Ascetocythere riopeli Hobbs & Walton 


Ascetocythere riopeli Hobbs & Walton, 1976: 
393-395, fig. la-d. 


Previously known range.—The Cumber- 
land Basin from only 2 localities in Letcher 
and Pike counties, Kentucky. 

New locality records. —Kentucky: Wolfe 
Co., Roadside ditch 2 mi (3.2 km) S of Rog- 
ers on Big Andy Ridge Rd, off Rte 715, 10 
Oct 1987, GWS, RFJ, et al., on C. dubius. 
Breathitt Co., Seep 3.0 mi (4.8 km) NNE of 
Camp Lewis on Rte 30, 16 Apr 1988, GWS, 
RFJ, et al., on C. dubius. 

Host: Cambarus buntingi, C. distans, C. 
dubius*, C. robustus, and Orconectes rusti- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


cus. As pointed out by Hobbs and Walton, 
1976:396, “it is highly probable that this 
ostracod is confined to [the burrowing] C. 
dubius.” 

Drainage systems. — Big Sandy, Cumber- 
land, Kentucky, and Licking basins (to Ohio 
and Mississippi). 


Ascetocythere sclera Hobbs & Hart 


Ascetocythere sclera Hobbs & Hart, 1966: 
42-43, figs. 15, 16.—Hart & Hart, 1974: 
40-41, pl. VIII, figs. 8-13, pl. XLVII.— 
Hobbs & McClure, 1983:777.—Hobbs & 
Peters, 1991:71-72. 


Previously known range. —Big Sandy, 
Clinch, Elk, Guyandotte, Kanawha, and Po- 
tomac basins in Buchanan, Dickinson, 
Highland, Russell, and Tazewell counties, 
Virginia; and Clay, McDowell, Raleigh (type 
locality), and Wyoming counties, West Vir- 
ginia. 

New locality records. —West Virginia: 
Boone Co., Seep, 4.3 mi (6.9 km) N of Kop- 
perston on Rte 85, 5 Jul 1988, GWS, RFJ. 
Kanawha Co., Kanawha State Forest camp- 
ground, 5 mi (8.5 km) W of Marmet, 8 Oct 
1988, GWS, RFJ. Wyoming Co., Burrows 
along spring in Twin Falls State Park, 7.8 
mi (12.5 km) E of Pineville, 6 Jul 1988, 
GWS, RFJ. Ditch 1 mi (1.6 km) N of 
McGraws on Co. Rd 5, 29 Jul 1988, GWS, 
T. Jones. Cambarus dubius served as the 
host in all of these localities. 

Host.—It is probably restricted to Cam- 
barus dubius. 

Drainage systems. — Big Sandy, Coal, Elk, 
Guyandotte, Kanawha basins (to the Ohio 
and Mississippi); Clinch Basin (to the Ten- 
nessee, Ohio, and Mississippi); and Poto- 
mac Basin. 


Cymocythere gonia Hobbs & Hart 


Cymocythere gonia Hobbs & Hart, 1966: 
51, figs. 36, 37.—Hart & Hart, 1974:44— 
45, pl. IX, figs. 12-14, pl. XLVI.— Hobbs 
& Walton, 1975:15. 


VOLUME 106, NUMBER 3 


Previously known range. — Holston Basin 
from 2 localities in Grainger (type locality) 
and Loudon counties, Tennessee. 

New locality records. —Alabama: Cle- 
burne Co., trib to Henry Ck off Co. Rd 66 
(T15S, R1OE, Sec 26N), 21 Apr 1973, R. 
W. Bouchard, J. D. Way, on C. halli and C. 
striatus. Marshall Co., Trib to Big Spring 
Ck at Co. Rd 12 (T9S, R2E, Sec 28SW), 20 
Apr 1973, RWB, JDW, on C. striatus, Pro- 
cambarus a. acutus, and O. erichsonianus. 
Shelby Co., Peavine Ck in Oak Mt. St. Park 
(T20S, R2W, Sec 8), 24 Mar 1974, RWB, 
J. W. Bouchard, on C. striatus, C. acan- 
thura, and P. lophotus. Georgia: Catoosa Co., 
Hurricane Ck above jct with Peters Ck off 
Rte 151, 24 Apr 1968, E. T. Hall, Jr., H. 
H. Hobbs, Jr., on C. extraneus, C. girardi- 
anus, C. striatus, and P. lophotus. Tennes- 
see: Anderson Co., Seepage area on SE side 
of Poplar Ck at Rte 61, 13-16 Mar 1972, 
D. A. Etnier, F. L. Oakberg, on C. deweesae. 
Blount Co., Pitner Ck off Co. Rd 2427, NW 
of Ellejoy, 4 May 1970, D. Walker, on C. 
bartonii cavatus, C. longirostris, O. erich- 
sonianus. Swamp on US Hwy 129 approx 
2.5 mi (4 km) N of Rte 72, 27 Apr 1971, 
DAE, on C. acanthura, C. striatus. Tem- 
porary pond at Co. Rd 2423, NE of Mid- 
way, 30 Apr 1970, RWB, on C. acanthura, 
C. striatus, C. longirostris. Cocke Co., Bur- 
rows between Del Rio and Harmony Grove 
on St Rte 107, 2 March 1972, RWB, FLO, 
DAE, C. Saylor, J. P. Dewees, on C. acan- 
thura. Grainger Co., ditch, 1 mi (1.6 km) N 
of Bean Station on unnumbered Co. Rd, 23 
Jul 1987, GWS, RFJ, D. Chrisman, on 
Cambarus acanthura and C. striatus. Sevier 
Co., Ditch, 3 mi (4.8 km) E of Boyds Creek 
on Rte 338, 31 Mar 1986, GWS, RFJ, on 
C. diogenes. Cove Ck at Co. Rd 2422, SW 
of Pigeon Forge, 22 Apr 1969, RWB, on C. 
b. bartonii, C. longirostris, O. erichsonianus 
and O. forceps. 

Hosts.—Cambarus acanthura*, C. bar- 
tonii bartonii, C. b. cavatus, C. halli, C. di- 
ogenes*, C. longirostris, C. striatus, C. dew- 
eesae.*, Orconectes erichsonianus, O. 


457 


forceps, O. spinosus, Procambarus a. acutus, 
and P. lophotus. 

Drainage systems. —Holston, French 
Broad, and Tennessee basins (to Ohio and 
Mississippi) and Cahaba and Tallapoosa 
basins (to Alabama and Mobile rivers). 


Dactylocythere coloholca Hobbs & Hobbs 


Dactylocythere coloholca Hobbs & Hobbs, 
1970:7, fig. 2.—Hart & Hart, 1974:53- 
54, pl. III, figs. 1-5, pl. XLVIII. 


Previously known range. —A single local- 
ity in the Cumberland Basin in Whitley 
County, Kentucky; no subsequent report of 
it has appeared in the literature. 

New locality records. — Kentucky: Breath- 
itt Co., Ditch 1.0 mi (1.6 km) S of Co. line 
on Rte 30, 16 Apr 1988, GWS, RFJ, M. 
Allen, on C. dubius. Morgan Co., Ditch 2.5 
mi ENE of Hazel Green on unnamed Rd 
off Rte 203, 10 Oct 1987, GWS, RFJ, D. 
Chrisman, P. Matesich, on C. dubius. Wolfe 
Co., Ditch in Koomer Ridge National For- 
est campground, campsite No 4, 4.2 mi (6.7 
km) ESE of Slade, 10 Oct 1987, GWS, RFJ, 
et al., on C. dubius. Ditch along Big Andy 
Ridge Rd off Rte 715, 2.0 mi (3.2 km) S of 
Rogers, 10 Oct 1987, GWS, RFJ, DC, PM, 
on C. dubius. Tennessee: Morgan Co., 11.6 
mi (18.6 km) E of Grimsley on Co. Rd., 5 
Jul 1969, P. C. Holt, V. F. Holt, on C. cri- 
nipes and C. sphenoides. Fentress Co., 
Frizsche Ck, about 0.1 mi (0.2 km) E of 
Allardt on unmarked road, 10 Jul 1969, 
PCH, VFH, on C. distans. Campbell Branch, 
0.4 mi (0.7 km) NW of jct Rte 52 on un- 
marked road, 10 Jul 1969, PCH, VFH, C. 
distans. Virginia: Lee Co., Wallin Ck. 0.5 
mi (0.8 km) W of Scott Co. line on US Hwy 
58, 15 Nov 1970, RWB, JDW, on C. lon- 
girostris, C. (C.) sp., Orconectes erichsoni- 
anus. West Virginia: Upshur Co., Burrows 
at jct of Rte 20 and Co. Rd 40/2, 0.2 mi 
(3.2 km) N of Arlington, 26 May 1989, 
GWS, RFJ, on C. diogenes. 


Hosts. —Cambarus crinipes, C. di- 


458 


ogenes*, C. dubius*, C. distans*, C. longi- 
rostris, C. sphenoides*, C. sp., and Orco- 
nectes erichsonianus. 

Drainage systems.—Cumberland, Ken- 
tucky, Little Kanawha, Powell, and Ten- 
nessee basins (to Ohio and Mississippi). 

Remarks. —Specimens collected in the lo- 
calities cited above in Morgan and Wolfe 
counties, Kentucky, are somewhat smaller 
than those reported from the type locality by 
Hobbs & Hobbs (1970): ranging from 420 
to 460 (X = 437) wm in length and 231-259 
(¥ = 241) um in height. In addition, the 
subangular posteroventral margin of the 
shell is sometimes more rounded; of the 3 
teeth on the preaxial border of the clasping 
apparatus, only the most proximal is well 
developed; and the accessory groove in some 
of the specimens almost reaches the level 
of the dorsal extremity of the spermatic loop. 


Dactylocythere crawfordi Hart 


Dactylocythere crawfordi Hart, 1965:255, 
figs. 1, 2.—Hart & Hart, 1974:55, pl. XIII, 
figs. 10-13, pl. XLIV.—Hobbs & Mc- 
Clure, 1983:776.— Hobbs & Peters, 1989: 
527, 199T-Gy, Fi. 


Previously known range. —Great Miami, 
Little Miami, Muskingum, Ohio, Scioto, and 
White basins in Decatur, Marion, and White 
counties, Indiana; Auglaise, Clinton, Frank- 
lin, Jackson, Licking, and Logan (type lo- 
cality) counties, Ohio; and Mason County, 
West Virginia. 

New locality records.—(The host was 
Cambarus diogenes unless otherwise not- 
ed.) Kentucky: Christian Co., Creek in Per- 
ryville State Park off Rte 109, Apr 1969, J. 
E. Pugh, D. J. Peters, HHH. Taylor Co., 
Ditch 1.0 mi (1.6 km) SW of Mannsville on 
Rte 70, 25 Mar 1987, GWS, RFJ, on Cam- 
barus diogenes and C. ortmanni. Ohio: Erie 
Co., Ditch, 3.2 mi (5.7 km) SE of Castalia, 
9 Oct 1983, J. Norrocky; Miller Rd, *4 mi 
(1 km) E of Rte 99, 29 Oct 1983, JN; Ditch 
on Wahl Rd just E of White’s Landing, 15 
Mar 1984, JN. Jackson Co., 2.8 mi (4.5 km) 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


NNE of Jackson, 1 May 1983, GWS, RFJ, 
R. F. Thoma. Ottawa Co., 8.1 mi (12 km) 
NE of Clinton, 10 Oct 1983, JN, on Orco- 
nectes rusticus. Perry Co., ditch 2.75 mi (4.4 
km) WSW of Corning, 3 Sep 1983, JN. Pond 
bank 8 mi (12.8 km) SE of New Lexington, 
3 Sep 1983, JN. Sandusky Co., W side of 
White’s Landing, Sec 4, between Rts 277 & 
283, 11 Dec 1983, JN, on F. fodiens. 1 mi 
(1.6 km) S of White’s Landing, 22 Nov 1983, 
JN. Ditch 2 mi (3.2 km) NNW of Vickery, 
20 Feb 1984, JN, on F. fodiens. 

Hosts.—Cambarus diogenes*, C. laevis, 
C. ortmanni, F. fodiens*, Orconectes s. san- 
bornii, and O. rusticus*. 

Drainage systems. — White Basin (to Wa- 
bash and Mississippi); Little Basin (to Cum- 
berland, Tennessee, Ohio, and Mississippi); 
Great Miami, Little Miami, Scioto, Rac- 
coon, Licking-Muskingum, Hocking, Ka- 
nawha, and Green basins (to Ohio and Mis- 
sissippi); and Lake Erie Basin. 


Dactylocythere crena Hobbs & Walton 


Dactylocythere crena Hobbs & Walton, 
1975:14, figs. 2a—f. 


Previously known range.—Known from 
only the type locality in the French Broad 
Basin, in Loudon County, Tennessee. 

New locality records. —Through an over- 
sight, Hobbs and Walton did not cite the 
two following localities from which they had 
specimens: Tennessee: Blount Co., Banks of 
little Tennessee River between Harrison 
Branch and Tallasee at Rte 72 and US Hwy 
129, 26 Apr 1970, DAE, on C. striatus. 
Temporary pond at Co. Rd 2423, NE of 
Midway, 30 Apr 1970, RWB, on C. acan- 
thura, C. striatus, C. longirostris. An addi- 
tional record was recently obtained in Se- 
vier Co., 3 mi E of Boyds Creek on Rte 338, 
31 Mar 1986, GWS, RFJ, on C. diogenes. 

Hosts. —Cambarus acanthura, C. di- 
ogenes*, C. longirostris, and C. striatus*. 

Drainage systems.—French Broad and 
Little Tennessee basins (to Tennessee, Ohio, 
and Mississippi). 


VOLUME 106, NUMBER 3 


1 


459 


.05 mm 


Fig. 1. a—c, Dactylocythere cryptoteresis, n. sp.; d, e, Phymocythere lophota, n. sp. (a, e, Copulatory complex 
of paratypic male; b, d, Shell of holotypic male; c, Shell of allotypic female). 


Dactylocythere cryptoteresis, new species 
Fig. la—c 


Diagnosis. — Male with eye pigmented and 
located about 0.2 shell length from anterior 
margin. Shell (Fig. 1b) ovate with greatest 
height slightly posterior to midlength where 
1.3 times height at levels of eye. Margin 
entire, lacking emarginations and promi- 
nences and nowhere angular. Submarginal 
setae absent dorsally and more abundant 
anterodorsally and posteroventrally. Shell 
length of males 413-448 (¥ = 425, n = 4) 
um; shell height 224-238 (¥ = 229, n = 4) 
pum. 

Copulatory complex (Fig. 1a) with arched 
peniferum gently rounded ventrally and 
meeting cephalic margin in acute angle. Ac- 
cessory groove reaching or almost reaching 
dorsal extremity of spermatic loop. Clasp- 
ing apparatus with horizontal and vertical 
rami disposed at angle of about 70 degrees, 
subequal in length but thickening and merg- 
ing imperceptibly in area of junction. Dorsal 
ramus entire, lacking shoulder on cephalic 
margin, comparatively thick, and weakly 
sinuous. Horizontal ramus with gently 
curved, entire postaxial margin; preaxial 
margin irregular and bearing 2 reduced 


(sometimes almost indescernible) teeth; apex 
of ramus with 4 acute, reflexed subapical 
denticles. Finger guard rather obscure but 
massive, short, and apparently unsclero- 
tized. Dorsal and ventral fingers unremark- 
able. 

Triunguis female. —Triunguis female with 
pigmented eye located 0.2 shell length from 
anterior margin. Shell (Fig. 1c) ovate with 
slight concavity ventrally just anterior to 
midlength; greatest height short distance 
posterior to midlength where almost 1.4 
times height at level of eye. Margins entire 
and nowhere angular. Submarginal setae as 
in male. Shell length 420-441 (¥ = 431, n 
= 7) wm; shell height 259-280 (X = 270, n 
= 7) wm. (Unfortunately several of the fe- 
males became fragmented in remounting the 
specimens. ) 

Genital complex consisting of bulbous 
tuberculiform lobe situated posterodorsally 
and projecting cephaloventrally; lacking 
J-shaped rod and amiculum. Slender tub- 
uliform pendant, which frequently previ- 
ously identified (perhaps mistakenly) as part 
of female genitalia, lying immediately an- 
terior to lobe. 

Type locality. —Crayfish burrows in ditch 


460 


at junction of Rte 20 and Co. Rd 40/2, 0.2 
mi (3.2 km) north of Arlington, Upshur Co., 
West Virginia. This locality, sampled on 26 
May 1989 by G. W. Stocker and R. F. Jez- 
erinac, is in the Little Kanawha River basin. 

Disposition of types. —The holotypic male 
and allotypic female are deposited in the 
National Museum of Natural History 
(Smithsonian Institution), USNM 260072 
and 260073, respectively. Paratypic males 
are in the collection of H. H. Hobbs III, 
Wittenberg University, and the Smithson- 
ian Institution. 

Host.—Cambarus diogenes. 

Entocytherid associates. —Dactylocythere 
coloholca. 

Relationships. —Dactylocythere crypto- 
teresis seems to have its closest affinities 
with those members of the genus in which 
the females lack an amiculum and is more 
similar to D. coloholca than to the others. 
The rounded posteroventral margin of the 
shell, the thickened junction of the rami of 
the clasping apparatus of the male, the re- 
duced size of the teeth on the preaxial bor- 
der of the horizontal ramus, and an acces- 
sory groove reaching dorsally to about the 
level of the dorsal extremity of the sper- 
matic loop will distinguish this ostracod 
from its relatives. 

Etymology.—G. crypto = hidden + ter- 
esis = guard; alluding to the difficulty in 
discerning the limits of the finger guard in 
males of this species; noun in apposition. 


Dactylocythere daphnioides (Hobbs) 


Entocythere daphnioides Hobbs, 1955:325, 
figs. 1-9. 

Dactylocythere daphnioides.—WHart, 1962: 
130.— Hobbs, Holt, & Walton, 1967:42.— 
Hart & Hart, 1974:56, pl. XIV, figs. 1-5, 
pl. XLVIII.—Hobbs & Peters, 1977:27, 
29-30. "41, 50, 525. 57> 72. lee TO oko: 
324, 327-329. 


Previously known range.—In describing 
this ostracod, Hobbs (1955) reported ma- 
terial that covered most of the currently 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


known range. Hobbs, Holt, & Walton (1967: 
42) cited a few new localities and described 
its range as extending “‘from the Watauga 
drainage system in Avery and Watauga 
Counties, N.C.; the New River system from 
Alleghany and Ashe Counties, N.C., to Po- 
cahontas County, W. Va.; and the Pound 
drainage system in Dickerson County, Va.” 
In their monograph, Hart & Hart (1974:56) 
added a number of new localities among 
which are four (those from Kentucky, Mis- 
souril, Clay and Fentress counties, Tennes- 
see) in need of confirmation. The most re- 
cent additions to the range was presented 
by Hobbs & Peters (1977:28, 72) who cited 
32 “localities in the Mountain and upper 
Piedmont provinces [of North Carolina] in 
the headwaters of the Pee Dee [Yadkin], 
Catawba, Little Tennessee, French Broad, 
Nolichucky, Watauga, and New rivers.” Ex- 
cluding the questioned localities mentioned 
above, this entocytherid ranges through the 
upper Pee Dee, and Catawba basins of North 
Carolina, the Tennessee River basin above 
Walden Gorge, headwaters of the Big Sandy 
River in Virginia, and throughout much of 
the Kanawha Basin. 

New localities. —Because it has not been 
reported previously from subterranean wa- 
ters, we cite the three following localities: 
West Virginia: Greenbrier Co., General Da- 
vis Cave (37°45'20’N, 80°33'15”W), 9 Sep 
1989, GWS etal., on C. nerterius. McClungs 
Cave (37°52'52’N, 80°23'24”W) 9 Sep 1989, 
GWS, D. Hemmerly, TJ, on C. nerterius. 
Pocahontas Co., Cave Creek Cave 
(38°12'12”N, 80°08’40’W), 20 Jul 1989, 
GWN, RFJ, TJ, on C. bartonii carinirostris. 

Hosts. —Cambarus acuminatus, C. as- 
perimanus, C. b. bartonii*, C. b. cariniros- 
tris*; C. chasmodactylus*;: C.dubiEae 
longirostris, C. longulus, C. nerterius*, C. 
reburrus, C. robustus*, C. sciotensis*, C. ve- 
teranus, Orconectes s. sanbornii* and O. s. 
erismophorous*. 

Drainage systems.—Scattered localities 
in headwater tributaries of the Catawba and 
Little Tennessee rivers in North Carolina 


VOLUME 106, NUMBER 3 


and Tennessee northward, in tributaries of 
the Tennessee, Pee Dee, Big Sandy, and New 
rivers, northward to the Greenbrier River 
in Pocahotas County and Little Kanawha 
Basin in Wirt County, West Virginia. 


Dactylocythere macroholca Hobbs & Hobbs 


Dactylocythere macroholca Hobbs & Hobbs, 
1970:9, fig. 3.—Hart & Hart, 1974:62, pl. 
XVI, figs. 6-10, pl. XLIX.—Hobbs & 
Walton, 1977:606, 609.—Hobbs & Pe- 
ters, 1989:326, 327-329; 1991:64, 69. 


Previously known range.—Known from 
only 9 localities in the Barren, Cumberland, 
Kentucky, and Licking basins in Allen, Bath, 
Madison, and Mason counties, Kentucky; 
and Fentress, Hawkins, and Pickett coun- 
ties, Tennessee. 

New localities. —Seventeen new localities 
have come to our attention in this study. 
Indiana: Randolph Co., Ditch 1.4 airmi (2.2 
airkm) SSE of Lynx on Co. Rd 700s, 1 May 
1986, GWS, RFJ, on Cambarus diogenes. 
Kentucky: Cumberland Co., Roadside ditch 
at intersection of Rts 912 & 704, 24 Mar 
1987, GWS, RFJ, DH, on C. striatus and 
C. (Jugicambarus) sp. Grayson Co., Bear Ck 
at Grayson Springs, 7.4 mi (11.8 km) N of 
Peoria on Rte 226, 11 Apr 1973, J. E. Pugh, 
G. B. Hobbs, HHH, on C. tenebrosus. War- 
ren Co., Stream 4.4 mi (7 km) SE of Butler 
Co. line on US Hwy 231, 11 Apr 1973, JEP, 
GWH, HHH, on C. graysoni, C. tenebrosus, 
C. diogenes, and O. putnami. Stream 4.4 mi 
(7 km) SE of Butler Co. line on US Hwy 
231, on C. diogenes, C. striatus, C. tenebro- 
sus, and O. putnami. Tennessee: Cannon 
Co., Brawley’s Fk off Co. Rd 4323 S of Cur- 
lee, 28 Mar 1971, RWB, JDW, on C. gray- 
soni, C. sp., and O. placidus. Clay Co., Hur- 
ricane Ck at Rte 52 in Oak Grove, 24 Dec 
1968, RWB, W. C. Starnes, on C. graysoni, 
C. tenebrosus, O. compressus, and O. put- 
nami. Big Trace Ck in Hermitage Springs 
at Rte 52, 11 Mar 1968, RWB, WCS, on C. 
graysoni, C. rusticiformis, O. compressus, 
and O. putnami. Hurricane Ck at Rte 52, 


461 


Oak Grove, 20 Mar 1972, RWB, JDW, on 
C. graysoni, C. tenebrosus, O. compressus, 
and O. putnami. Davidson Co., Sevenmile 
Ck at Co. Rd 6158 in Oglesby, 27 Mar 1971, 
RWB, JDW, on C. graysoni, C. tenebrosus, 
O. shoupi, and O. sp. DeKalb Co., Dry Ck 
at Co. Rd 4360 off US Hwy 70, S of Dow- 
elltown, 9 Nov 1968, RWB, WCS, on C. 
friaufi, C. graysoni, C. tenebrosus, and O. 
placidus. Hawkins Co., Approx 3 mi (4.8 
km) S of Kyle’s Ford on Rte 70, 25 Sep 
1971, RWB, DAE, FLO, CS, on C. dubius. 
Lawrence Co., Little Shoal Ck in Davey 
Crockett St Park off US Hwy 64, 27 Oct 
1973, RWB, JWB, on C. girardianus, C. 
graysoni, C. (Hiaticambarus) sp., O. spi- 
nosus, and O. forceps. Macon Co., Stream 
about 300 m E of Sumner Co. line at Rte 
52, 24 Dec 1968, RWB, WCS, on C. gray- 
soni, O. compressus, and O. putnami. Put- 
nam Co., Falling Water River off US Hwy 
7ON, NW of Rocky Point, 30 Jul 1969, 
RWB, R. Sayrs, A. Gnilka, on C. graysoni, 
C. rusticiformis, C. tenebrosus, and O. pla- 
cidus. Smith Co., Trib of Snow Ck in Elm- 
wood off US Hwy 70, 23 Mar 1971, RWB, 
JDW, on C. friaufi, C. graysoni, C. tenebro- 
sus, and O. sp. Sumner Co., Bledsoe Ck at 
US Hwy 231-31E and Rte 6, N of Boze, 24 
Mar 1971, RWB, JDW, on C. graysoni, C. 
tenebrosus, O. placidus, and O. sp. Caney 
Fork Ck at Rte 52, E of Portland, 17 Aug 
1969, RWB, on C. graysoni, C. tenebrosus, 
O. compressus, O. placidus, and O. sp. 

Hosts. —Cambarus batchi*, C. diogenes*, 
C. dubius*, C. friaufi, C. girardianus, C. 
graysoni, C. laevis*, C. rusticiformis, C. 
striatus, C. tenebrosus*, C. (Hiaticambarus) 
sp., C. (Jugicambarus) sp., C. sp., Orco- 
nectes compressus, O. forceps, O. placidus, 
O. putnami, O. shoupi, O. spinosus, and O. 
sp. 

Drainage systems.—The range includes 
segments of the following river basins: Ten- 
nessee (including the Holston), Cumber- 
land, Barren-Green, Kentucky, Licking, and 
Whitewater (to Ohio and Mississippi riv- 
ers). 


462 


Dactylocythere myura Hobbs & Walton 


Dactylocythere myura Hobbs & Walton, 
1970:859, figs. 2e, f, 3e, f, h.—Hart & 
Hart, 1974:64, pl. XVII, figs. 11-14, pl. 
XLIX. 


Previously known ranges. —Only 2 local- 
ities in the South Fork of the Holston River 
basin in Smith and Washington counties, 
Virginia. 

New locality.—Tennessee: Sullivan Co., 
Roadside ditch 11.9 mi SW of Tennessee- 
Virginia line on US Hwy 11, 13 Sep 1969, 
RWB, on C. dubius. Virginia: Washington 
Co., Along Garrett Ck, 1.5 mi (2.4 km) S 
of Holston on Co Rd 11, 9 Aug 1984, GWS, 
RFJ, on C. dubius. Ditch 1.2 mi E of Bow- 
den on US Hwy 33, 10 Apr 1986, GWS, 
RFJ, on C. dubius. 

Host. —Cambarus dubius. 

Drainage systems.—South Fork of Hol- 
ston River Basin (to Tennessee River). 


Dactylocythere prionata (Hart & Hobbs) 


Entocythere prionata Hart & Hobbs, 1961: 
174, figs. 15-17. 

Dactylocythere prionata Hart, 1962:130.— 
Hart & Hart, 1966:5; 1974:66, pl. X VIII, 
figs. 11-13, pl. XLIX. 


Previously known range. —Caves and 
springs in the Barren, Cumberland, and 
Kentucky basins of Jackson, Pulaski, and 
Warren counties, Kentucky. 

New locality records. —Kentucky: Pulaski 
Co., Stream flowing into Sloan’s Valley 
Cave, 5 Apr 1969, JEP, DJP, HHH, on C. 
tenebrosus. Taylor Co., 1 mi (1.6 km) SW 
of Mannsville on Rte 70, 25 Mar 1987, 
GWS, RFJ, on C. diogenes and C. ortmanni. 
Wayne Co., 3 caves: 2 at Rte 92 in Elk Spring 
Valley between Rte 776 and Oil Valley, and 
another at Rte 92, NW of Coopersville, 9 
Apr 1971, RWB, on C. tenebrosus. Oldham- 
Trimble cos., Pattons Ck, 2 mi (3.2 km) NW 
of Sligo, 19 Apr 1980, J. A. Thoma, RFJ, 
M. McCluskey, on C. diogenes, C. ortman- 
ni, C. ornatus, and O. rusticus. Tennessee: 
Clay Co., Hurricane Ck on Rte 52 at Oak 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Grove, 20 Mar 1972, RWB, on C. graysoni, 
C. tenebrosus, O. compressus, and O. put- 
nami. Scott Co., Marsh at Kentucky-Ten- 
nessee state line on US Hwy 27, 9 Apr 1971, 
RWB, DAE, on C. diogenes, and C. striatus. 
Hosts. — Cambarus diogenes, C. graysoni, 
C. ornatus, C. ortmanni, C. striatus, C. te- 
nebrosus*, Orconectes australis packardi*, 
O. compressus, O. putnami, and O. rusticus. 
Drainage systems. —Cumberland, Bar- 
ren-Green, Kentucky and Ohio basins. 


Donnaldsoncythere donnaldsonensis (Kliie) 


Entocythere donnaldsonensis Klie, 1931: 
334, figs. 1-9. 

Donnaldsoncythere donnaldsonensis. — Hart, 
1962:131.—Hart & Hart, 1974:78-79, pl. 
XXIII, fig. 6, pl. L._—Hobbs & Walton, 
1976:396, 399, 403; 1977:603, 606, 609, 
612.—Hobbs & Peters, 1977:22, 24-25, 
30, 33, 38, 41, 44, 50, 52, 55, 57, 69 fig. 
21; 1982:300. 307, 308, 311, 312 (fig. 7); 
1989:325, 326, 327-328; 1991:67, 69, 70— 
72, 73, 74.—Hobbs & McClure, 1983:772, 
716, 777, 778. 

Donnaldsoncythere hiwasseensis. —Hobbs 
& Walton, 1975:10, 12, 13, 18, 19.—Pe- 
ters, 1975:111, 5, 7-8, 10, 14, 19-20, 22, 
25-31, 33-34, 46. 


[For a complete synonymy, see Hobbs & 
Peters (1977:43-44); only references to lo- 
calities that have been recorded subsequent 
to those listed by Hart & Hart (1974) are 
included here.] 

This ostracod occurs so commonly (and 
on such a wide range of ecologically diverse 
crayfishes) throughout its known range—ac- 
cording to Hobbs & Peters 1977:44, “‘north- 
ern Georgia to Indiana and Maine”’—that 
there seems little reason to cite new local- 
ities that do not extend the currently rec- 
ognized limits of its distribution. New re- 
cords that have come to our attention 
include those in Kentucky: Breathitt, El- 
liott, Magoffin, Morgan, and Wolfe cos, on 
C. dubius; Virginia: Washington Co., on C. 
dubius; and West Virginia: Boone, Braxton, 
Greenbrier, Kanawha, Mercer, Pocahontas, 


VOLUME 106, NUMBER 3 


Randolph, Wirt, and Wyoming cos, on C. 
carinirostris, C. dubius, C. monongalensis, 
and C. nerterius. 

Hosts. —For reasons pointed out above, 
listing the large number of hosts here seems 
a bit excessive. One might anticipate that it 
probably infests all crayfish species occur- 
ring within its range. 

Drainage systems.— Atlantic Basin: Sa- 
vannah, Santee, Pee Dee, Roanoke, James, 
York, Potomac, Susquehanna, Delaware, 
Hudson, and St. Francis; Gulf of Mexico 
Basin: Coosa, Tennessee (all tributaries 
above and including the Sequatchie and in 
the Elk and Duck), Cumberland (wide- 
spread), Ohio (from headwaters and south- 
ern tributaries to Whitewater basin in In- 
diana). St. Lawrence Basin: Lake Erie basin 
eastward to northern Maine. 


Lordocythere petersi Hobbs & Hobbs 


Lordocythere petersi Hobbs & Hobbs, 1970: 
11, 16, fig. 9a-d.—Hart & Hart, 1974: 
103, pl. XXX, figs. 4—7, pl. LIT. —Hobbs 
& Peters, 1977:58—-59, fig. 30; 1991:72. 


Previously known range. — Four localities 
in the Cumberland, Emory, and Hiwassee 
basins in Whitley Co., Kentucky (type lo- 
cality), Cherokee Co., North Carolina, and 
Morgan and Scott counties, Tennessee. 

New locality records. —Kentucky: Knox 
Co., G. R. Hampton Elementary School 
grounds on Rte 11 in Barbourville, 1 Apr 
1986, GWS, RFJ, on C. diogenes. 

Hosts. —Cambarus acanthura*, C. di- 
ogenes*, C. dubius, C. nodosus*, and C. 
sphenoides. 

Drainage systems. —Cumberland, Emo- 
ry, and Hiwassee basins (to Tennessee and 
Mississippi rivers). 


Phymocythere lophota, new species 
Fig. Id, e 


Diagnosis. — Male with eye pigmented and 
located slightly less than 0.2 shell length from 
anterior margin. Shell (Fig. 1d) subovate, 
shallowly excavate ventrally anterior to 
midlength; greatest height about 0.6 shell 


463 


length from anterior margin where 1.4 times 
height at level of eye. Margins entire, lack- 
ing emarginations and prominences. Sub- 
marginal setae absent dorsally but rather 
evenly distributed along other borders. Shell 
length 399-406 (X = 404, n = 3) um; shell 
height 217-224 (X¥ = 219, n = 3) um. 

Copulatory complex (Fig. le) with peni- 
ferum arched posteriorly, swollen ventrally 
and with ventral emargination resulting in 
bilobed appearance, swollen anteroventral 
area with crest and produced anteriorly in 
subacute, sclerotized, beaklike prominence 
disposed anterodorsally. Long inverted 
U-shaped penis situated in swollen area, its 
basal part situated in posterior lobe and 
apex, which directed anteroventrally, lying 
in anterior lobe. Clasping apparatus 
L-shaped with vertical ramus tapering and 
joining horizontal ramus in gentle curve 
rather than at angle; latter ramus increasing 
in size distally and bearing 4 small reflexed 
subapical denticles; both rami otherwise 
unadorned. Finger guard tapering from 
swollen base but slightly compressed and 
troughlike apically. Ventral finger strongly 
curved caudally at about 100 degrees at end 
of basal two-fifths; dorsal finger unremark- 
able. 

Triunguis female. — Unknown. 

Type locality.—Standingstone Creek at 
bridge on Co. Rd 3, 2.8 mi (4.3 km) NE of 
Cherry, 4.1 mi (6.6 km) ESE of Elizabeth, 
Wirt Co., West Virginia. This creek is in the 
Little Kanawha River drainage. The spec- 
imens were collected by G. W. Stocker and 
R. F. Jezerinac on 7 Oct 1988. 

Disposition of types. —The holotypic male 
is deposited in the National Museum of 
Natural History (Smithsonian Institution), 
USNM 260074. Paratypes are in the col- 
lection of H. H. Hobbs III, Wittenberg Uni- 
versity and the Smithsonian Institution. 

Host. —Cambarus monongalensis. 

Entocytherid associates. —Donnaldson- 
cythere donnaldsonensis. 

Range and specimens examined. —Known 
only from a single collection, consisting of 
4 males, made in the type locality. 


464 


Relationships. —This is the second spe- 
cies to be assigned to the genus Phymocy- 
there. It differs from Ph. phyma in possess- 
ing an anterodorsally directed acute 
prominence on the swollen, crested, bilobed 
ventral part of the peniferum. 

Etymology. —G. lophos = crest; lophotos 
= crested (/ophotus-a-um) adj.; alluding to 
the crest and acute prominence on the an- 
teroventral part of the peniferum. 


Phymocythere phyma (Hobbs & Walton) 


Entocythere phyma Hobbs & Walton, 1962: 
42, figs. 10-13. 

Cymocythere phyma. — Hart, 1962:129. 

Phymocythere phyma. —Hobbs & Hart, 
1966:48-49.— Hobbs & Walton, 1966:7; 
Hobbs, Holt, & Walton, 1967:46.— Wal- 
ton & Hobbs, 1971:88.—Hart & Hart, 
1974:110, pl. XX XII, figs. 7-9, pl. LIT. — 
Hobbs & McClure, 1983:777. 


Previously known range. —Twelve local- 
ities in the Big Sandy, Cheat, James, New 
(Kanawha), Potomac, and Tygart basins in 
Craig, Giles, and Rockingham counties, 
Virginia; and Greenbrier, McDowell, Pen- 
dleton, Randolph, and Summers counties, 
West Virginia. 

New localities. —West Virginia: Green- 
brier Co., U.S. 219 Cave, 7 Jul 1989, GWS, 
RFJ, on Cambarus nerterius. General Davis 
Cave, 1 mi (1.6 km) NW of Fort Spring, 9 
Sep 1989, GWS, DH, TJ, S. Van Luik, on 
C. nerterius. Wades Cave 3.2 mi S of Max- 
welton, 9 Sep 1988, GWS, DH, TJ, on C. 
b. carinirostris. Monroe Co., Steeles Cave 
(37°33'52”N, 80°33'00"W), 8 Sep 1989, 
GWS, DH, TJ, on C. b. carinirostris and O. 
virilis. McDowell Co., Panther Ck, 5 mi (8 
km) S of Panther, 19 Jun 1981, RFT, on C. 
dubius, C. sciotensis, Cambarus sp., and Or- 
conectes sp. 

Hosts. —Cambarus b. bartonii*, C. b. car- 
inirostris*, C. dubius, C. nerterius*, C. ro- 
bustus, C. sciotensis, C. sp., Orconectes spi- 
nosus, O. virilis and O. sp. 

Drainage systems.—Big Sandy, Cheat, 
Greenbrier, New, and Tygart basins (to 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Ohio), James, and Potomac basins (to At- 
lantic Ocean). 


Uncinocythere zancla Hobbs & Walton 


Uncinocythere zancla Hobbs & Walton, 
1963:456-457, figs. 1-3. Hart & Hart, 
1966:8; 1974:140-141, pl. XX XVIIL, figs. 
10-12, pl. LVIl.—Hobbs & Walton, 1976: 
397(?); 1977:606. 


Previously known range. —Hart and Hart 
(1974:141) cited the localities (29, however, 
the three records from North Carolina and 
Georgia were based on misidentifications) 
and hosts (14, however, three must be de- 
leted for the reason just given) known for 
this entocytherid at that time. Subsequently 
11 more localities, involving 11 additional 
hosts, became known. Of these, we have 
been unable to confirm the locality cited by 
Hobbs & Walton (1977) reporting this os- 
tracod from Sevier Co., Tennessee, on 
Cambarus carolinus; inasmuch as this lo- 
cality is somewhat removed from the other 
known localities, and the host is not oth- 
erwise known to harbor this ostracod, nei- 
ther the locality nor the host appears in the 
summary below. The record in Carter Co., 
Kentucky, cited by Hobbs and Peters (1989: 
326, 327, 328, 329) was based on the mis- 
identification of specimens of U. simondsi 
(Hobbs & Walton, 1960). Until now, the 
species was known to be widespread in the 
Cumberland, Duck, and Elk basins, and a 
few localities were recorded in the north- 
ward flowing segment of the Tennessee Riv- 
er. Localities had been established in Adair, 
Allen, Hardin, and Hart counties, Ken- 
tucky; and Cannon, Davidson, DeKalb, 
Dickson, Fentress, Franklin, Hickman, 
Humphreys, Lawrence, Lincoln, Marshall, 
Maury, Perry, Pickett, Putnam, Rutherford, 
Wayne, Williamson, and Wilson counties, 
Tennessee. As pointed out by Hobbs & Pe- 
ters (1977:63) the published records for 
North Carolina were based on misidentifi- 
cations. 

New localities. —More than 100 addition- 
al localities are now known from Adair, Al- 


VOLUME 106, NUMBER 3 


len, Grant, Grayson, Hardin, Hart, Logan, 
Madison, Taylor, and Warren counties, 
Kentucky; and Bedford, Clay, Cannon, 
Cheatham, Clay, Coffee, Davidson, De- 
Kalb, Dickson, Fentress, Giles, Hickman, 
Houston, Humphreys, Lawrence, Lewis, 
Lincoln, Macon, Marshall, Maury, Mont- 
gomery, Overton, Pickett, Putnam, Rob- 
ertson, Rutherford, Smith, Stewart, Sum- 
ner, Trousdale, Wayne, Williamson, and 
Wilson counties, Tennessee. 

Hosts.—Barbicambarus cornutus, Cam- 
barus bartonii bartonii, C. b. cavatus, C. 
brachydactylus, C. carolinus, C. crinipes, C. 
cumberlandensis, C. diogenes, C. dubius, C. 
friaufi, C. gentryi, C. girardianus, C. gray- 
soni, C. ortmanni, C. robustus, C. rustici- 
formis, C. striatus*, C. tenebrosus, Orco- 
nectes barrenensis, O. compressus, O. 
erichsonianus, O. forceps, O. i. inermis*, O. 
mirus, O. placidus*, O. putnami*, O. rhoad- 
esi, O. rusticus, O. shoupi, O. spinosus, and 
O. sp. 

Drainage systems. —Tennessee Basin be- 
tween mouth of Sequatchie River and 
northward flowing segment (including the 
Elk, and Duck watersheds), and Cumber- 
land (including the Harpeth), and Green 
river systems. 


Acknowledgments 


We extend our thanks to Raymond F. 
Jezerinac and G. Whitney Stocker, as we do 
to Raymond W. Bouchard, for making 
available to us most of the material studied 
and to other collectors cited among the new 
locality records. We are also grateful to the 
three just named for furnishing us with the 
identifications of the hosts and for com- 
ments on the manuscript. For their con- 
structive criticisms of this work, we extend 
appreciation to C. W. Hart, Jr., of the 
Smithsonian Institution, and H. H. Hobbs 
III, of Wittenberg University. 


Literature Cited 


Hart, C. W. 1962. A revision of the ostracods of the 
family Entocytheridae.— Proceedings of the 


465 


Academy of Natural Sciences of Philadelphia 

114(3):121-147. 

1965. New entocytherid ostracods and dis- 
tribution records for five midwestern states. — 
Transactions of the American Microscopical 
Society 84:255-259. 

, & D. G. Hart. 1966. Four new entocytherid 

ostracods from Kentucky, with notes on the 

troglobitic Sagittocythere barri.—Notulae Na- 
turae of the Academy of Natural Sciences of 

Philadelphia 388:1-10. 

—., & H. H. Hobbs, Jr. 1961. Eight new trog- 
lobitic ostracods of the genus Entocythere (Crus- 
tacea, Ostracoda) from the eastern United 
States. — Proceedings of the Academy of Natural 
Sciences of Philadelphia 113(8):173-185. 

Hart, D. G., & C. W. Hart, Jr. 1974. The ostracod 
family Entocytheridae.—Academy of Natural 
Sciences of Philadelphia Monograph 18:ix + 
239 pages. 

Hobbs, H. H., Jr. 1955. Ostracods of the genus En- 

tocythere from the New River system of North 

Carolina, Virginia, and West Virginia. —Trans- 

actions of the American Microscopical Society 

74(4):325-333. 

1989. An illustrated checklist of the Amer- 
ican crayfishes (Decapoda: Astacidae, Cambar- 
idae, and Parastacidae).—Smithsonian Contri- 
butions to Zoology 480:1i1 + 236 pages. 

, & C. W. Hart, Jr. 1966. On the entocytherid 

ostracod genera Ascetocythere, Plectocythere, 

Phymocythere (gen. nov.), and Cymocythere, with 

descriptions of new species. — Proceedings of the 

Academy of Natural Sciences of Philadelphia 

118(2):35-61. 

, P. C. Holt, & M. Walton. 1967. The cray- 

fishes and their epizootic ostracod and bran- 

chiobdellid associates of the Mountain Lake, 

Virginia, region.—Proceedings of the United 

States National Museum 123(3602):1-84. 

& H. H. Hobbs III. 1970. New entocytherid 
ostracods with a key to the genera of the sub- 
family Entocytherinae.—Smithsonian Contri- 

butions to Zoology 47:1-19. 

,& A.C. McClure. 1983. Onasmall collection 

of entocytherid ostracods with the descriptions 

of three new species.— Proceedings of the Bio- 

logical Society of Washington 96:770-779. 

, & D. J. Peters. 1977. The entocytherid os- 

tracods of North Carolina.—Smithsonian Con- 

tributions to Zoology 247:iv + 75 pages. 

——, & 1982. The entocytherid ostracod 

fauna of northern Georgia. — Proceedings of the 

Biological Society of Washington 95:297-318. 

,& . 1989. New records of entocytherid 

ostracods infesting burrowing crayfishes, with 

the description of a new species, Ascetocythere 


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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


stockeri. — Proceedings of the Biological Society 
of Washington 102:324-330. 


——., & 


1991. Additional records of en- 


tocytherid ostracods infesting burrowing cray- 


fishes, with 
Proceedings 


descriptions of five new species. — 
of the Biological Society of Wash- 


ington 104:64-75. 


—., & M. Walton. 1960. Three new ostracods of 


the genus Entocythere from the Hiwassee drain- 
age system in Georgia and Tennessee. — Journal 
of the Tennessee Academy of Science 35:1 7-23. 


eavemeeny -) 
Entocythere 


1962. New ostracods of the genus 
from the Mr. Lake region, Virgin- 


ia.— Virginia Journal of Science 132:42-48. 


=== 


. 1963. Three new ostracods (Ostra- 


coda, Entocytheridae) from the Duck River 


drainage in 


Tennessee.— American Midland 


Naturalist 69:456-461. 


, & 


1966. A new genus and six new 


species of entocytherid ostracods (Ostracoda, 
Entocytheridae).— Proceedings of the United 
States National Museum 119(3542):1-12. 


——., & 


. 1970. New entocytherid ostracods 
from Tennessee and Virginia.— Proceedings of 


the Biological Society of Washington 82(68):85 1— 


864. 
eee 


. 1975. New entocytherid ostracods 


from Tennessee with a key to the species of the 
genus Ascetocythere.— Proceedings of the Bio- 
logical Society of Washington 88(2):5—20. 


Bese see 


. 1976. New entocytherid ostracods 


from Kentucky and Tennessee. — Proceedings of 
the Biological Society of Washington 89(33):393- 
404. 

——,, & . 1977. New entocytherid ostracods 
of the genus Dactylocythere.— Proceedings of the 
Biological Society of Washington 90:600-614. 

Klie, W. 1931. Campagne spéologique de C. Bolivar 
et R. Jeannel dans l’Amerique du Nord (1928). 
3. Crustacés Ostracodes.—Biospeologica: Ar- 
chives de Zoologie Expérimentale et Générale 
71(3):333-344. 

Peters, D. J. 1975. The entocytherid ostracod fauna 
of the James and York River basins with a de- 
scription of a new member of the genus Ento- 
cythere. — Virginia Polytechnic Institute and State 
University, Research Division Bulletin 937111 + 
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Walton, M., & H. H. Hobbs, Jr. 1971. The distri- 
bution of certain entocytherid ostracods on their 
crayfish hosts. — Proceedings of the Academy of 
Natural Sciences, 123(4):87-103. 


(HHH) Department of Invertebrate Zo- 
ology, National Museum of Natural His- 
tory, Smithsonian Institution, Washington, 
D.C. 20560, U.S.A.; (DJP) York High 
School, 9300 George Washington Highway, 
Yorktown, Virginia 23692, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(3), 1993, pp. 467-489 


SCOPALATUM VORAX (ESTERLY, 1911) AND 
SCOLECITHRICELLA LOBOPHORA PARK, 1970, 
CALANOID COPEPODS (SCOLECITRICHIDAE) 


ASSOCIATED WITH A PELAGIC 
TUNICATE IN MONTEREY BAY 


Frank D. Ferrari and Deborah K. Steinberg 


Abstract. —The last five copepodid stages of Scopalatum vorax and the last 
three of Scolecithricella lobophora are described from specimens collected in 
association with the pelagic tunicate Bathochordaeus sp. A non-feeding, stage 
six nauplius of the former species has three anterior naupliar appendages, five 
posterior appendage buds, and no mouth. The addition of setae during cope- 
podid development to the uniramal appendages of S. vorax suggests that distal 
segments of maxilla 2 are exopodal while those of the maxilliped are endopodal; 
addition of setae on antenna 1 are endopodal in pattern. The developmental 
pattern of the maxilliped of S. vorax suggests that it consists of a coxa, a basis 
with a distomedial lobe, and five endopodal segments. 


During the last three years, scientists at 
the University of California, Santa Cruz and 
Monterey Bay Aquarium Research Institute 
(MBARI) have undertaken a study of ma- 
rine snow in Monterey Bay (Pilskaln et al. 
1991, Silver et al. 1991). One of us (DKS) 
is studying the ecological role of metazoans 
associated with large mucus structures which 
contribute significantly to marine snow in 
Monterey Bay. These mucus structures, 
which are on the order of tens of centimeters 
in diameter, are derived from the house and 
filtering apparatus of a midwater larvacean, 
Bathochordaeus sp. (Barham 1979, Galt 
1979, Hamner & Robison 1992). 

We have found several copepods associ- 
ated with the house and filtering apparatus 
of the larvacean. These include the poeci- 
lostomatoids Oncaea conifera and O. sim- 
ilis, an harpacticoid, Microsetella rosea, and 
Metridia pacifica, a calanoid common in the 
pelagic waters. One of the most abundant 
calanoid associated with larvacean houses 
is a large, bright-red scolecitrichid, Scopa- 
latum vorax (Esterly 1911), which previ- 
ously was known from a single adult female. 


Scolecithricella lobophora Park, 1970, an- 
other rare scolecitrichid also was collected 
around the houses. 

Adult females and several other cope- 
podid stages of both species are described 
from specimens collected in the field. A fifth 
copepodid of Scottocalanus thomasi is the 
only juvenile scolecitrichid described (Sew- 
ell 1929). A nauplius of Scopalatum vorax 
which hatched from an egg produced by one 
of several females in culture is described; 
this is the first report of a scolecitrichid nau- 
plius. We also analyse the developmental 
patterns of setal addition to the first six ap- 
pendages of Scopalatum vorax, and from 
these patterns develop hypotheses about the 
identity and homologies of antenna 1, max- 
illa 2, and the maxilliped. 

The family Scolecitricidae initially was 
established as a subfamily, Scolecithrichina, 
of the Calanidae by Giesbrecht (1892). Sars 
(1902) used the now widely-accepted name, 
Scolecithricidae; Bowman & Abele (1982) 
changed the family name to Scolecitrichi- 
dae, using the correct latin transliteration of 
the genitive of the Greek word for “‘thrix.” 


468 


Bradford (1973) recently redefined the fam- 
ily and its genera. Roe (1975) established 
Scopalatum for the “‘Amallophora’’ altera 
group of scolecitrichids defined by Bradford 
(1973); that group included Esterly’s Sco- 
lecithrix vorax. Esterly’s species has not been 
recorded since its initial description. Sars 
(1902) established Scolecithricella. S. lo- 
bophora Park, 1970 has been recorded once, 
as Amallothrix lobophora by Roe (1975). 


Methods 


Copepods were collected with houses of 
Bathochordaeus sp. in Monterey Bay in wa- 
ter deeper than 1000 m over a submarine 
canyon (36°42’N, 122°02’W) on seven dif- 
ferent occasions between 8 October 1989 
and 20 December 1991. Depths of samples 
ranged from 198-310 m in temperatures of 
7.6°-9.0°C and salinities 33.95-—34.07%o. 
Samples were collected during the day using 
a remotely-operated submersible, the Ven- 
tana. Two different types of samplers on the 
submersible were used to collect the lar- 
vacean houses and associated copepods. The 
“‘detritus sampler,’ from Harbor Branch 
Oceanographic Institution, is a 7.5 liter 
plexiglass cylinder with an opening/closing 
lid at either end. The “‘suction sampler”’ is 
a vacuum system; samples are drawn 
through a vacuum nozzle and deposited onto 
a rotating carousel containing canisters, each 
of which is fitted with a 165 wm mesh net 
and cod end. Larvacean houses were fixed 
with buffered 4% formaldehyde and their 
associated copepods subsequently were sep- 
arated. In a few cases, living specimens of 
Scopalatum vorax were pipetted from house 
samples before fixation and placed in 0.2 
um filtered seawater in a dark, 10°C cold 
room. These copepods were fed a mixture 
of the algae Dunaliella tertiolecta, Isochrysis 
galbana, and Thalassiosira weissflogii. One 
nauplius hatched from several eggs pro- 
duced by a female in culture. 

Specimens were preserved in the labo- 
ratory in 0.5% propylene phenoxytol/4.5% 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


proplylene glycol/95.0% water. They were 
clared in steps through 50.0% lactic acid/ 
50.0% water to 100% lactic acid, and stained 
by adding a solution of chlorazol black E 
dissolved in 70.0% ethanol/30.0% water. 

The naupliar stage is presumed to be the 
sixth and is abbreviated N6; second through 
sixth copepodid stages are CII to CVI. Tho- 
racic and abdominal somites are numbered 
according to their relative developmental 
age as interpreted from data of Hulsemann 
(1991). The first and oldest thoracic somite 
bears the maxilliped and is fused with the 
cephalon. The youngest is the seventh; it is 
the only thoracic somite without an ap- 
pendage. In adult calanoids it is the first 
somite of the urosome, and in adult females 
it is fused to the second abdominal somite. 
The first, and oldest, abdominal somite is 
the last; it bears the caudal rami. The youn- 
gest is immediately anterior to the oldest, 
and the remaining abdominal somites in- 
crease in age anteriorly. 

Appendages are Al = antennule; A2 = 
antenna; Mn = mandible; Mx1 = maxillule; 
Mx2 = maxilla; Mxp = maxilliped; ap- 
pendages on thoracic somites are P1—5; cau- 
dal ramus = CR. Designations of appendage 
segments generally follow Huys & Boxshall 
(1991) except for Mx2 and Mxp; exopods 
= Re; endopods = Ri; medial lobes of a 
segment = li, lateral lobe = le. Terminal 
segments of Mx2 are exopodal. Mxp has at 
most 5 endopodal segments. 

Ramal segments on P1—4 are numbered 
by their developmental age (see Hulsemann 
1991, Ferrari & Ambler 1992, for a discus- 
sion of the age of these segments) and not 
proximal-to-distal as is the usual case for 
copepod descriptions. The distal-most seg- 
ment of a ramus is the first segment. The 
second segment is immediately distal to the 
basipod. If present, the third segment is im- 
mediately proximal to the distal (or first) 
segment of a 3-segmented ramus. For a 
3-segmented ramus, the proximal segment 
is the second segment, the middle segment 
is the third segment, and the distal segment 


VOLUME 106, NUMBER 3 


is the first segment. Thus, developmentally 
homologous segments are given the same 
number in this system. The number of setae 
recorded for the segments follows this same 
scheme. 

Armament elements of appendages are 
setae. Examples of the quality of setae are 
illustrated. Three setae and one aesthetasc 
on a segment of Al are designated 3+ 1; if 
these elements are broken, one number is 
given for setae plus aesthetascs. Bradford’s 
setae are those modified setae on Re of Mx2 
and syncoxa of Mxp; their diversity and tax- 
onomic value were described by Bradford 
(1973). Breaking planes are annular regions 
on a seta where the cuticular wall is thinner 
(Von Vaupel Klein 1982:112). Setules are 
epicuticular extensions of a seta and den- 
ticles are epicuticular extensions of an ap- 
pendage segment; spinules, epicuticular ex- 
tensions of a somite, are not found on these 
species. Groups of denticles are distally po- 
larized if their tips point distally; they are 
radially polarized if their longitudinal axes 
appear to diverge from a central point. Von 
Vaupel Klein’s organ on P1 (the appendage 
of thoracic somite 2) consists of the curved 
basipodal seta and tubercle with denticles 
on the endopodal segment; its taxonomic 
value was described by Von Vaupel Klein 
(1972). 


Results 


Scopalatum vorax (Esterly, 1911) 
Figs. 1-9 


Scolecithrix vorax.—Esterly, 1911:327-328, 
figs. 15, 21, 29, 45, 68, 93, 96, 99. 

?Amallophora smithae. —Grice, 1962:205— 
206, pit. 15, figs. 12—22 (see remarks). 


CVI female. — Length of 8 specimens 2.64, 
2266, 2.68, 2.70 (2), 2.75 (2); 2.93 mm; av- 
erage Pr length/Ur length = 4.6; average Pr 
length/Pr depth = 2.5. 

Pr (Fig. 1A): 4 segments; Ist a complex 
of 5 cephalic somites plus thoracic somites 
1 and 2; thoracic somites 3 and 4 simple 


469 


and articulated; fourth segment a complex 
of thoracic somites 5 and 6. 

Ur (Fig. 1B): 4 segments; Ist a genital 
complex of thoracic somite 7 and abdom- 
inal somite 2 (Fig. 1C); abdominal somites 
3, 4, 1 articulated. 

Rostrum (Fig. 1H): 2 long filaments; ar- 
mament of labrum and paragnath as in Fig. 
1E. 

Al: (Fig. 2A—-E) 23 articulated segments 
Witthowos lle 2 2b adele 12 4-1 
etl le alee Me 1 2. 
2, 7+1 setae + aesthetascs; lst segment with 
2 rows of small denticles. 

A2 (Fig. 3A, B): Coxa with 1 seta and a 
row of long denticles; basis with 2 setae. Re 
7-segmented with 0, 1, 1, 1, 1, 1, 4 setae 
with thick, dense setules above breaking 
plane and few scattered setules below; ter- 
minal 3 setae 1.5 times length of medial 
ones. Ri 2-segmented with 3, 15 (7 terminal, 
8 subterminal) setae. 

Mn (Fig. 3C, D): Coxa with a row of thin 
and a row of thick denticles; basis with 2 
setae. Re 5-segmented with 1, 1, 1, 1, 3 
setae, each with breaking plane. Ri 
2-segmented with 1, 9 setae. 

Mx! (Fig. 3E, F): Le with 9 setae, largest 
7 with breaking plane. Re 1-segmented with 
8 setae; baseoendopod with medial sets of 
5 and 3, and 6 terminal setae. Li 2 and 3 
with 2 and 4 setae. Li | with 9 apical and 
2 posterior setae; denticles on anterior and 
posterior surfaces. 

Mx2 (Fig. 3G, H, I): Li 1-4 of coxa each 
with 3 setae and posterior denticles; li of 
basis with 4 setae. Re indistinctly segment- 
ed with 8 Bradford’s setae; 1 thick with long 
apical setules, 4 thin with short apical se- 
tules, and 3 thin with tiny apical setules. 

Mxp (Fig. 4A, B): Coxa with 7 (1 Brad- 
ford’s) setae, basis with 5 (2 on a subter- 
minal medial lobe). Ri with 4, 4, 3, 4 (1 
lateral), 4 setae. 3 areas of denticles on coxa 
and a longitudinal row of denticles on basis. 

P1 (Fig. 4C): Coxa with medial denticles, 
and basis with medial and lateral denticles. 
Re 3-segmented with 5, 1, 2 setae; segments 


470 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Scopalatum vorax, CVI female: A, animal, lateral; B, Ur, lateral; C, genital segment with attached 
spermatophore, lateral; D, CR, dorsal; E, labrum and labium, lateral; F, P5, posterior; G, unusual PS, posterior; 
H, rostrum, lateral. Line 1 = 1.0 mm for A; line 2 = 0.1 mm for B—D; line 3 = 0.1 mm for E-G. 


VOLUME 106, NUMBER 3 471 


Fig. 2. Scopalatum vorax, CVI female: A, free segments 1—8 of Al; B, free segments 9-14 of Al; C, free 
segments 15-18 of Al; D, free segments 19-23 of Al; E, teminal setae of last segment of Al; F, P2, posterior, 
arrow to Re3 anterior; G, P3, posterior; H, P4, posterior. Line = 0.2 mm; letters on proximal-most segments 
of A-D. 


472 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Scopalatum vorax, CV1 female: A, A2; B, examples of lateral setae on subterminal (left) and apical 
setae on terminal (right) segment of A2; C, gnathobase of Mn; D, basis, Re, and Ri of Mn; E, Mx1, anterior; 
F, li of Mx1 with posterior setae darkened; G, coxa of Mx2; H, basis of Mx2; I, Re of Mx2. Line = 0.2 mm. 


VOLUME 106, NUMBER 3 473 


Fig. 4. Scopalatum vorax, CVI female: A, Mxp, anterior; B, basis of Mxp, posterior; C, Pl, anterior. CV 
female: D, Mxp li of basis and Ri. Line = 0.2 mm. 


474 


2 and 3 with medial denticles. Ri 
l-segmented with 5 setae; Von Vaupel 
Klein’s organ with 13-15 denticles begin- 
ning medially as a linear set of 3-4 short 
denticles and continuing laterally as longer 
denticles scattered over surface of tubercle. 
Breaking plane on inner setae of both rami. 

P2 (Fig. 2F): Coxa with medial seta and 
denticles. Re 3-segmented with 8, 2, 2 setae; 
posterior face of segments 1 and 3 and an- 
terior face of segment 1 with distally polar- 
ized denticles. Ri 2-segmented with 5, | se- 
tae; posterior face of segment 1 with 3 sets 
of distally polarized denticles. Breaking 
plane on inner setae of both rami. 

P3 (Fig. 2G): Coxa with medial seta and 
3 sets of denticles, 2 medial and | lateral. 
Re 3-segmented with 8, 2, 2 setae; posterior 
face of segments 1 and 3 with distally po- 
larized denticles; segment 3 with a poste- 
rior, cuticular ridge. Ri 3-segmented with 
5, 1, 1 setae; posterior face of segment 1 
with 2 sets of radially polarized denticles; 
segment 3 with a set of radially polarized 
denticles. Breaking plane on inner setae of 
both rami. 

P4 (Fig. 2H): Coxa with medial seta. Re 
3-segmented with 8, 2, 2 setae; posterior 
face of segment 3 with distally polarized 
denticles; segments 1 and 3 with epicutic- 
ular ridges. Ri 3-segmented with 5, 1, 1 se- 
tae; anterior face of segment 1 with 2 sets 
of distally polarized denticles. Breaking 
plane on inner setae of both rami. 

P5 (Fig. 1F, G): Coxa of both left and 
right legs fused to interpodal coupler. Ar- 
ticulating segment a baseoexopod, usually 
with 2 apical setae; a specimen with 3 setae 
on left PS. 

CR (Fig. 1D): 4 large terminal setae, a 
smaller seta on a distomedial, ventral lobe, 
and a very small seta on distolateral, dorsal 
lobe. 

Spermatophore (Fig. 1C): A simple, prox- 
imally narrow sac; on 1 female it is placed 
directly over the copulatory pore. 

CV female. — Differs from CVI female as 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


follows: length of 4 specimens 2.02, 2.13, 
2.23, 2.55 mm; average Pr length/Ur length 
= 4,3. 

Pr (Fig. 5A): 5 segments; lst a complex 
of 5 cephalic somites plus thoracic somite 
1; thoracic somite 2 fused ventrolaterally to 
anterior complex; thoracic somites 3 and 4 
articulated; 5th segment a complex of tho- 
racic somites 5 and 6. 

Ur (Fig. 5A): 4 segments; thoracic somite 
7 and abdominal somites 2, 3, 1 articulated. 

A2 (Fig. 5C): Ri terminal segment with 
14 setae (7 terminal, 7 subterminal). 

Mn (Fig. 5D): Ri2 with 8 setae. 

Mxp (Fig. 4D): Ri with 3, 3, 2, 3 (1 lat- 
eral), 4 setae. 

P5 (Fig. SE): Basis and Re 1 separate. 

CV male. — Differs from CV female as fol- 
lows: length of 6 specimens 2.17, 2.25, 2.30, 
2.40, 2.42, 2.63 mm; average Pr length/Ur 
length = 4.4. 

Pr (Fig. 5B): 6 segments; thoracic somites 
5 and 6 articulated. 

P5 (Fig. 5F): Coxa and basis separate; Re 
2-segmented with 2, 1 setae; Ri 1-segmented 
with | seta. 

CIV male. — Differs from CV male as fol- 
lows: length of 2 specimens 1.69, 1.59 mm; 
average Pr length/Ur length = 4.1; average 
Pr length/Pr depth = 2.8. 

Ur (Fig. 6A): 3 segments; thoracic somite 
7 and abdominal somites 2, 1 articulated. 

Al (Fig. 6B): 23 segments; proximal 8 
segments with 3, 3+1, 141, 1, lies 
2+1 setae + aesthetascs; proximal segment 
with 1 row of denticles. 

Mx! (Fig. 6C): Baseoendopod with me- 
dial sets of 4 and 3, and 5 terminal setae. 

Mxp (Fig. 6D): Ri with 3, 2, 1, 2 (1 lat- 
eral), 4 setae. 

P1 (Fig. 6E): Re 2-segmented with 7, 1 
setae; segment 2 with medial denticles. 

P2 (Fig. 6F): Re 2-segmented with 9, 2 
setae; posterior face of segment 1 with prox- 
imal set of large and distal area of small 
distally polarized denticles. 

P3 (Fig. 6G): Re 2-segmented with 9, 2 


VOLUME 106, NUMBER 3 


475 


Fig. 5. Scopalatum vorax, CV female: A, animal, lateral; C, A2, tip of Ri; D, Mn, tip of Ri; E, PS, posterior. 
CV male: B, Th4-6 lateral; F, PS, posterior. Line 1 = 1.0 mm for A, B; line 2 = 0.1 mm for C, D, E; line 3 = 


0.1 mm for F. 


setae; posterior face of segment | with prox- 
imal set of large and distal area of small 
distally polarized denticles. Ri 2-segmented 
with 6, | setae; posterior face of segment 1 
with 3 sets of radially polarized denticles. 
P4 (Fig. 6H): Re 2-segmented with 9, 2 
setae; posterior face of segment 1 without 
distally polarized denticles; segment 1, 3 
without cuticular ridge. Ri 2-segmented with 


6, 1 setae; anterior face of segment | without 
distally polarized denticles. 

P5 (Fig. 61): Re 1-segmented with 2 setae; 
Ri 1-segmented without seta. 

CIV female. —Differs from CIV male as 
follows: length of 2 specimens 1.78, 1.65 
mm. 

P5: (Fig. 6J) Re 1-segmented with | seta. 

CITT. —Differs from CIV male as follows: 


476 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 6. Scopalatum vorax, CIV male: A, animal, lateral; B, proximal 8 segments of Al; C, Ri of Mx1; D, 
end of basis and Ri of Mxp; E, Re of P1; F, Re of P2; G, Re (left) and Ri (right) of P3; H, Re (left) and Ri 
(right) of P4; I, P5. CIV female: J, P5. Line 1 = 1.0 mm for A; line 2 = 0.1 mm for B-J. 


VOLUME 106, NUMBER 3 


length of 3 specimens 1.29, 1.30, 1.41 mm; 
average Pr length/Ur length = 3.9. 

Ur (Fig. 7A): 2 segments; thoracic somite 
7 and abdominal somite | articulated. 

Al (Fig. 7B—D): 22 articulated segments 
meg 4 0.1.2... 1 ft, 2. 1, 1, I, 
1, 2, 1, 1, 2, 2, 8 setae plus aesthetascs to- 
gether. 

Mn (Fig. 7E): Ri2 with 6 apical setae. 

Mxl (Fig. 7F, G): Le with 8 setae. Re 
l-segmented with 7 setae; baseoendopod 
with medial sets of 4 and 2, and 5 terminal 
setae. Lil with 1 apical seta reduced in size. 

Mxp (Fig. 7H): 6 segmented; Ri 
4-segmented with 1, 1, 2 (1 lateral), 4 setae. 

P3 (Fig. 71): Re 2-segmented with 7, 2 
setae; posterior face of segment 1 without 
proximal set of large distally polarized den- 
ticles. 

P4 (Fig. 7J): Re 1-segmented with 7 setae; 
without distally polarized denticles. Ri 
l-segmented with 5 setae; without distally 
polarized denticles. 

P5: not apparent. 

CIT. — Differs from CIII as follows: length 
of 4 specimens 0.93, 0.97, 0.99, 1.09 mm; 
average Pr length/Ur length = 3.3. 

Pr (Fig. 8A): 5 segments; thoracic somite 
2 fused ventrolaterally and laterally to an- 
terior complex. 

Ur (Fig. 8A): 2 segments; thoracic somite 
6 and abdominal somite | articulated. 

Al (Fig. 8B—E): 16 articulated segments 
mam. 1-0, 2,0; 1, 0, 1, 2,45 1, 2, 2, 
8 setae plus aesthetascs together. 

A2 (Fig. 8F): Ri terminal segment with 
10 (6 terminal, 4 subterminal) setae. 

Mn (Fig. 8G): Ri2 with 5 setae. 

Mx1 (Fig. 8H): Le with 6 setae. Re 
l-segmented with 6 setae; baseoendopod 
with medial sets of 3 and 2, and 5 terminal 
setae. Lil with 7 apical and 2 posterior se- 
tae. 

Mxp (Fig. 81): 5 segments with 7 (1 Brad- 
ford’s) setae on coxa, 4 (2 on a subterminal 
medial lobe) on basis. Ri 3-segmented with 
1, 1, 4 setae. 

P1: basis with 1 medial seta. 


477 


P2 (Fig. 8J): Re 2-segmented with 7, 1 
setae; posterior face of segment 1 with dis- 
tally polarized denticles. Ri 2-segmented 
with 5, 1 setae; posterior face of segment 1 
with 2 sets of distally polarized denticles. 

P3 (Fig. 8K): Coxa unarmed. Re 
1-segmented with 7 setae; without distally 
polarized denticles. Ri 1-segmented with 6 
setae; without polarized denticles. 

P4 (Fig. 8L): A bilobed bud on posterior 
edge of thoracic somite 5; outer lobe with 
2 setae, inner lobe with 1 seta; lobes and 
setae point dorsally. 

N6 (Fig. 9A). — Length of 1 specimen 0.57 
mm. Without mouth; ventrally a cuticular 
ridge between A2 and Mn. No caudal setae 
apparent. 

A (Fig. 9B): 2-segmented; segment 2 with 
3 lateral, 3 terminal setae and denticles. 

A2 (Fig. 9C): 1 basal segment. Re indis- 
tinctly 1l-segmented with 5 setae. Ri 
l-segmented with 3 apical setae. 

Mn (Fig. 9D): Indistinctly segmented with 
inner lobe on basal area. Re with 5 setae, 
Ri with 3. 

Mx! (Fig. 9A): A bilobed bud. 

Mx2 (Fig. 9A): A unilobed bud. 

Mxp (Fig. 9A): A unilobed bud. 

P1 (Fig. 9A): A unilobed bud. 

P2 (Fig. 9A): A unilobed bud. 

Remarks. —Esterly (1911) described Sco- 
palatum vorax (as Scolecithrix vorax) from 
a single female specimen 1.6 mm collected 
in a vertical net tow from 310 fm (558 m) 
off San Diego; the location of that specimen 
is unknown. Our specimens agree in general 
with Esterly’s description and illustrations 
of the appendages, except that our speci- 
mens are larger and thoracic somites 5 and 
6 are fused. 

Of the remaining nominal species (Roe 
1975) in the genus, S. dubia (T. Scott, 1894), 
S. farrani Roe, 1975, S. gibbera Roe, 1975, 
and S. smithae (Grice, 1962), only S. smi- 
thae has been reported from the Pacific 
Ocean. It was described as Amallophora 
smithae by Grice (1962) from one adult fe- 
male, 1.40 mm, collected from the equa- 


478 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 7. Scopalatum vorax, CIII: A, animal, lateral; B, free segments 1-11 of Al; C, free segments 12-17 of 
Al; D, free segments 18-22 of Al; E, Mn, tip of Ri; F, Mx1; G, Mx1, armament of lil; H, end of basis and Ri 
of Mxp; I, Rel of P3; J, P4. Line 1 = 1.0 mm for A; line 2 = 0.1 mm for B-J; letters on proximal-most segments 


of B—D. 


torial Pacific Ocean southeast of Hawaii. 
The description of that specimen agrees in 
most character states with ours, including 
the fusion of thoracic somites 5 and 6, and 
the armament of most appendages; how- 
ever, PS of that specimens has only one ter- 
minal seta. The dissected type specimen is 


mounted on a slide and consists of 10 sub- 
mounts with 1 Al, 2 A2; 1 Mn; 2 Mx]; 2 
Mx2; 2 Mxp; nothing; 2 P2 complete with 
coupler; 2 P3 missing left Re2-3; 2 P4 miss- 
ing right Re2-3 and Ri2-3; nothing. 

The type localities of S. farrani and S. 
gibbera are in the Atlantic Ocean. P5 of the 


VOLUME 106, NUMBER 3 479 


Fig. 8. Scopalatum vorax, Cll: A, animal, lateral; B, free segments 1-6 of Al; C, free segments 7-11 of Al; 
D, free segments 12-14 of Al; E, free segments 15-16 of Al; F, A2, tip of Ri; G, Ri of Mn; H, Mx1; I, basis 
and Ri of Mxp; J, Re (right) and Ri (left) of P2; K, P3; L, right P4 (dorsal is up). Line 1 = 0.3 mm for A; line 
2 = 0.1 mm for B-L; letters on proximal-most segments of B—D. 


480 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 9. Scopalatum vorax, N6: A, animal, ventral; B, Al; C, A2; D, Mn. Line 1 = 0.2 mm for A; line 2 = 
0.1 mm for B—-D. 


VOLUME 106, NUMBER 3 


former has two free segments and an un- 
armed mandibular gnathobase; the latter 
species has a characteristic dorsal protu- 
berance on the middle of the cephalosome. 
S. dubia is known only from male speci- 
mens. 

Our specimens exhibit an overlap in body 
length among different copepodid stages. For 
example the length of one CV female (2.65 
mm) is within the length range of CVI fe- 
males (2.64—2.93 mm). From these few ob- 
servations we do not believe that variation 
in length of adult copepodids is known well 
enough to provide definitive information for 
separating our specimens from S. vorax or 
S. smithae. One of our adult females exhib- 
ited asymmetry in the number of terminal 
setae on P5, suggesting that PS armament 
may not provide definitive information for 
separating species. Based on present speci- 
mens and our limited knowledge of the ge- 
nus we believe that our specimens should 
be assigned to S. vorax, and that S. smithae 
may be conspecific with it. 

Larger collections of co-occurring adult 
females and males of Scopalatum will pro- 
vide opportunities for careful descriptions 
of the male P5 and should resolve the status 
of these species. 

Identity and homologies. —In addition to 
variation in size and number of setae on PS, 
the most easily detected variation among 
our specimens of S. vorax is the number of 
polarized denticles on rami of P2-4. For ex- 
ample, denticle numbers on left and right 
endopodal segments | and 3 of P3 from 3 
specimens are given in Table 1. The differ- 
ence in counts between left and right ap- 
pendages and among specimens suggests that 
the numbers of denticles may not have spe- 
cific taxonomic value. 

The position of surface denticles on P2-4 
during copepodid development, however, 
does provide information about the growth 
of the appendage cuticle. Denticles initially 
appear on the posterior surface of segment 
1 of leg 3 at CIII. We believe that these 
denticles are homologous to those on the 


481 


Table 1.— Variation in numbers of polarized den- 
ticles on the left and right endopod of P3 for three 
specimens (#1-—#3) of Scopalatum vorax. Number to 
the left of semicolon is the count for the set on the 
subterminal segment; two numbers to the right of the 
semicolon are the counts for two sets (separated by a 
comma) on the terminal segment. 


Left Right 
#1 5; 4, 3 Sey Aten 
#2 7; 4, 4 6; 6, 4 
#3 7, 6,4 $214.3 


3rd segment of P3 at CV. Their develop- 
ment suggests that growth of the cuticular 
exoskeleton of that leg occurs at the same 
time, and along the same mediolateral axis, 
as the formation of the setae on that seg- 
ment. These findings support data of Ferrari 
& Ambler (1992) who used formation ho- 
mology to show that a new inner seta and 
outer seta often are added to the proximal 
border of the distal segment of P3 of Dioith- 
ona oculata during the molt prior to the 
formation of the new segment that will bear 
those setae. 

Table 2 shows copepodid stages at which 
setae are added to either ramus of the bira- 
mus A2, Mn and Mx1, and to the uniramus 
Al, Mx2 and Mxp. Armament of exopods 
of the biramus appendages is complete by 
CII while endopods continue to add setae 
often through CVI. Al and Mxp continue 
to add setae late in development while Mx2 
does not. These results suggest that the ter- 
minal segments of Mx2 are exopodal, not 
endopodal as has been suggested by Gies- 
brecht (1894) and Huys & Boxshall (1991), 
and the terminal segments of Mxp are en- 
dopodal. The developmental pattern of se- 
tal addition to Al is similar to that of an 
endopod. 

Table 3 gives two interpretations of the 
number of setae on the terminal and 4 sub- 
terminal endopodal segments, and on the 
basal lobe of the Mxp. This latter structure 
has recently been called an endopodal seg- 
ment by Huys & Boxshall (1991). During 


482 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 2.—Stages of development of Scopalatum vorax at which setae are added (A) to rami of A2, Mn, Mx1, 


and to Al, Mx2, Mxpd. 


A2 Mn 
Al Re Ri Re 
cil = oe = a 
CII A A 
GIY, A 
CV A 
CVI A 


copepodid development, setae are added to 
the 4 subterminal endopodal segments but 
there is no change in number of setae on the 
lobe of the basis. These data and Von Vau- 
pel Klein (1982) suggest that this latter 
structure is a true lobe of the basis, and not 
subterminal, endopodal segment fused to 
the basis. 

Based on its developmental pattern, we 
have interpreted P5 of the adult female as 
the coxa of both left and right legs which 
are fused to the interpodal coupler; the ar- 
ticulating segment is the basis fused to ex- 
opod 1. Post-maxillipedal appendages usu- 
ally are transformed from a bud to an 


Table 3.— Two interpretations of the number of setae 
on the distal inner lobe of the basis (L) and on the five 
endopodal segments (numbered proximally to distally) 
of the maxilliped of Scopalatum vorax for stages CII— 
CVI. A—when new segments are armed with only one 
seta; B— when new segments are added proximally to 
the distal-most segment. a = segment presumed ab- 
sent. 


1 Ist 2nd 3rd 4th Sth 
A 
CII yD 1 a a 1 4 
CIll 2 1 1 a De 4 
CIV y 3 y 1 2 4 
GV! 2 3 3 2 3 4 
CVI Dy 4 4 3 4 4 
B 
CII 2 1 1 a a 4 
CIlIl 2 1 1 2 a 4 
CIV D 3 D 1 2 4 
CV iD 3 3 Ms 3 4 
CVI 2 4 4 3 4 4 


lod 
r 
Oo 
ee 


Mxpd 


> +P | 
>P>p | 
>> P>p | 


appendage having a basis articulating with 
the coxa and l-segmented rami (Ferrari 
1988, Ferrari & Ambler 1992). For PS of 
calanoids this transformation usually oc- 
curs during the molt to CIV (Ferrari 1988). 
P5 of CIV male (Fig. 61) exhibits this trans- 
formed morphology. Its exopod has 2 setae 
and its endopod is unarmed. P5 of CIV fe- 
males (Fig. 6J) is similar but has one ramal 
segment which, we believe, is an exopod 
because it is armed with a seta. P5 of CV 
female (Fig. SE) is morphologically simular; 
the unarmed basis articulates with the coxa 
and with a 1-segmented exopod now armed 
with 2 setae. An alternate explanation 1s that 
the basis has fused to the coxa and the ex- 
opod has gained an unarmed, second seg- 
ment; CV male has a 2-segmented exopod 
but the second segment has an outer seta 
(Fig. 5F). For CVI female the basis of P5 
has fused with the exopodal segment, while 
the fused coxa and coupler remain. In the 
alternate developmental pattern, the artic- 
ulating segment would incorporate two or 
three fused exopodal segments. We prefer 
the first developmental pattern and its in- 
terpretation of the articulating segment, a 
basis fused to exopod 1. 

Behavior and ecology. — Houses of shal- 
lower living larvaceans are known to serve 
as surface habitat for a variety of zooplank- 
ton and can be used as a food source for 
some copepods (Alldredge 1972, 1976; Oht- 
suka & Kubo 1991). S. vorax could be seen 
clearly with the submersible’s video camera 
moving around the inner food collecting fil- 
ter of larvacean houses and less often near 


VOLUME 106, NUMBER 3 


its outer house or “‘sheet”” (Hamner & Ro- 
bison 1992). Preliminary analyses of the gut 
contents of S. vorax show similarities be- 
tween food items, diatom and crustacean 
skeletal parts, and diatom and crustaceans 
associated with the houses. These data sug- 
gest that the community of organisms as- 
sociated with larvacean houses is being uti- 
lized as a food source by a S. vorax. 

The presence of lobes of Mx1, Mx2, Mxp, 
P1, and P2, suggests that our nauplius is a 
stage 6 calanoid nauplius. Absence of a 
mouth opening, labrum, paragnaths and 
gnathobase on A2 suggests that this nau- 
plius cannot feed. If the rules of Izawa (1987) 
for a reduced number of naupliar stages can 
be extended to calanoids, we expect S. vorax 
to develop through six naupliar stages. We 
do not expect these lecithotrophic nauplii 
to occur in shallow depths. 


Scolecithricella lobophora Park, 1970 
Figs. 10-13 


Scolecithricella lobophora. — Park, 1970:511, 
515, figs. 188-201. not Amallothrix lo- 
bophora. —Roe, 1975:329, fig. 17 (see re- 
marks). 


CVI female. —length of 4 specimens 1.26, 
1.52, 1.75, 1.85 mm; average Pr length/Ur 
length = 3.7. 

Pr (Fig. 10A): 4 segments; Ist a complex 
of 5 cephalic somites plus thoracic somites 
1 and 2; thoracic somites 3 and 4 simple 
and articulated; fourth segment a complex 
of thoracic somites 5 and 6. 

Ur (Fig. 10B): 4 segments; Ist a genital 
complex of thoracic somite 7 and abdom- 
inal somite 2; abdominal somites 3, 4, 1 
articulated. 

Rostrum (Fig. 10A): 2 short points. 

Al (Fig. 11B—E): 22 articulated segments 
mane, Ft 21 2. 21, 24 1 2 +7, 4, 1, 
Paes 1 1, ty te 2 EE OT 1S Oo 744 
setae + aesthetascs; segments 1, 8-11 dis- 
tally with circumferential denticles. 

A2: Coxa with | seta and row of long 
denticles; basis with 2 setae. Re 7-segmented 


483 


with 0, 1, 1, 1, 1, 1, 4 setae with breaking 
planes. Ri 2-segmented with 2, 15 (7 ter- 
minal, 8 subterminal) setae and lateral den- 
ticles. 

Mn (Fig. 10C, D): Coxa with 2 rows of 
thin and 1| row of thick denticles; basis with 
2 setae; denticles medially and laterally. Re 
4-segmented with 1, 1, 1, 3 setae. Ri 
2-segmented with 1, 9 setae. 

Mx! (Fig. 10E, F): Le with 9 setae. Re 
l-segmented with 8 setae; baseoendopod 
with medial sets of 5 and 3, and 6 terminal 
setae. Li 2 and 3 with 2 and 4 setae. Li 1 
with 9 apical and 2 posterior setae; denticles 
on anterior and posterior surfaces. 

Mx2: Li 1-4 of coxa each with 3 setae and 
posterior denticles; li of basis with 4 setae. 
Re indistinctly segmented with 8 Bradford’s 
setae: 5 thin with short apical setules, and 
3 thin with tiny apical setules. 

Mxp: Coxa with 6 (1 Bradford’s) setae, 
basis with 5 (2 ona subterminal medial lobe). 
Ri 5-segmented with 4, 4, 3, 4 (1 lateral), 4 
setae; 3 areas of denticles on coxa and a 
longitudinal row of denticles on basis. 

P1: Coxa and basis with medial denticles. 
Re 3-segmented with 5, 0, 2 setae; segment 
2 with medial denticles. Ri 1-segmented with 
5 setae; Von Vaupel Klein’s organ with 8— 
10 denticles below crest of tubercle. Break- 
ing plane on inner setae of both rami. 

P2 (Fig. 12A): Coxa with medial seta; ba- 
sis with distolateral denticles. Re 
3-segmented with 8, 2, 2 setae; posterior 
face of segments 1 and 3 with distally po- 
larized denticles. Ri 2-segmented with 5, 1 
setae; posterior face of segment | with 3 sets 
of radially polarized denticles. Breaking 
plane on inner setae of both rami. 

P3 (Fig. 12B, C): Coxa with medial seta 
and 2 medial sets of denticles; basis with 
distolateral set of denticles. Re 3-segmented 
with 8, 2, 2 setae; posterior face of segments 
1 and 3 with distally polarized denticles; 
anterior face of segment 2 with distally po- 
larized denticles. Ri 3-segmented with 5, 1, 
1 setae; posterior face of segment 3 with a 
set of radially polarized denticles and of seg- 


484 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 10. Scolecithricella lobophora, CV1 female: A, animal, lateral; B, Ur, lateral; C, gnathobase of Mn, 
anterior; D, basis, Re and Ri of Mn, posterior; E, Mx1; F, lil of Mx1. Line 1 = 0.2 mm for A; line 2 = 0.2 
mm for B; line 3 = 0.1 mm for C-F. 


ment | with 2 sets of radially polarized den- and basis with distolateral denticles. Re 
ticles. Breaking plane on inner setae of both 3-segmented with 8, 2, 2 setae; posterior 
rami. face of segment 1 with distally polarized 

P4 (Fig. 12D, E): Coxa with medial seta denticles. Ri 3-segmented with 5, 1, 1 setae; 


VOLUME 106, NUMBER 3 485 


Fig. 11. Scolecithricella lobophora, CV1 female: A, CR, ventral; B, free segments 1-10 of Al; C, free segments 
11-14 of Al; D, free segments 15-18 of Al; E, free segments 19-22 of Al. Line 1 = 0.2 mm for A; line 2 = 
0.1 mm for B-E; letters on proximal-most segments of D-G. 


486 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 12. Scolecithricella lobophora, CV\ female: A, P2, posterior; B, P3, posterior; C, Re2 of P3, anterior; 
D, P4, posterior; E, Ril, 3 of P4, anterior; F, P5, posterior. Line = 0.1 mm. 


anterior face of segments 1 and 3 with dis- legs fused to interpodal coupler; basis sep- 


tally polarized denticles. Breaking plane on arate. Re 1-segmented with 2 apical setae. 
inner setae of both rami. CR (Fig. 11A): 4 large terminal setae, and 


P5 (Fig. 12F): Coxa of both left and right a smaller seta on dorsal and on lateral sur- 


VOLUME 106, NUMBER 3 487 


Fig. 13. Scolecithricella lobophora, CV female: A, Th 5-6 and Ur; B, Ri2 of A2; C, Ri2 of Mn; D, baseoen- 
dopod of Mx1; E, Ri of Mxp; F, PS. CIV female: G, Th 5-6 and Ur; H, Ri2 of A2; I, Ri2 of Mn; J, baseoendopod 
of Mx1; K, Ri of Mxp; L, Re of Pl; M, Re of P2; N, Re (left) and Ri (right) of P3; O, Re (left) and Ri (right) 
of P4; P, P5. Line 1 = 0.2 mm for A, B; line 2 = 0.1 mm for C-P. 


faces; medially and dorsally with denticles. Ur (Fig. 13A): 4 segments; thoracic so- 

Terminal setae each with breaking plane. mite 7 and abdominal somites 2, 3, | ar- 
CV female. —differs from CVI female as __ ticulated. 

follows: length of 1 specimen 1.52 mm; Pr A2 (Fig. 13B): Ri2 with 14 setae (7 ter- 

length/Ur length = 4.1. minal, 7 subterminal). 


488 


Mn (Fig. 13C): Ri2 with 8 setae. 

Mx1 (Fig. 13D): Baseoendopod with 5 
terminal setae. 

Mxp (Fig. 13E): Ri with 3; 3,2, 3 (1 lat- 
eral), 4 setae. 

P5 (Fig. 13F): Rel terminal seta shorter 
than lateral. 

CIV female. —differs from CV female as 
follows: length of 1 specimen 1.26 mm; Pr 
length/Ur length = 3.4. 

Ur: (Fig. 13G) of 3 segments; thoracic 
somite 7 and abdominal somite 2 and 1 
articulated. 

Al: 22 segments; proximal 8 segments 
with 3, 35 12riG. io, ter 1 ea 2 aesetac 
+ aesthestascs. 

A2: (Fig. 13H) Ri2 with 13 setae (7 ter- 
minal, 6 subterminal). 

Mn (Fig. 131): Ri2 with 7 setae. 

Mx! (Fig. 13J): Baseoendopod with me- 
dial sets of 4 proximal and 3 distal setae. 

Mxp (Fig. 13K): Ri with 2, 2, 1, 2 (1 lat- 
eral), 4 setae. 

P1 (Fig. 13L): Re 2-segmented with 7, 0 
setae; segments 1 and 3 with medial den- 
ticles. 

P2 (Fig. 13M): Re 2-segmented with 9, 2 
setae; posterior face of segment 1 with dis- 
tally polarized denticles. 

P3 (Fig. 13N): Re 2-segmented with 9, 2 
setae; posterior face of segment 1 with dis- 
tally polarized denticles. Ri 2-segmented 
with 6, 1 setae; posterior face of segment 1 
with 3 sets of large and | set of small po- 
larized denticles. 

P4 (Fig. 130): Re 2-segmented with 9, 1 
setae; posterior face of segment 3 without 
distally polarized denticles. Ri 2-segmented 
with 6, 1 setae; anterior face of segment 1 
with distally polarized denticles. 

P5 (Fig. 13P): Re indistinctly separate 
from basis with 1 seta. 

Remarks. — Scolecithricella lobophora was 
described by Park (1970) from a single adult 
female collected in the southern Gulf of 
Mexico. Our specimens differ from Park’s 
description and illustration of 7 terminal 
setae on an articulating endopod of Mxl1 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


and a lobe on the dorsodistal edge of the 
cephalosome. However, the undescribed 
Mx1 of the type specimen has only 5 ter- 
minal setae. Although our specimens do not 
possess a lobe on the cephalosome, we have 
chosen to refer them to S. /obophora. 

Roe (1975) described males and females 
of Amallothrix lobophora (Park, 1970) with 
3 posterior setae on endite | of Mxl. We 
have not observed variation in this char- 
acter among our specimens and believe that 
Roe’s specimens are not conspecific with S. 
lobophora. 


Acknowledgments 


We thank the crew of the R/V Pt. Lobos 
and the pilots of Ventana for their help col- 
lecting specimens. Dr. Janet Bradford- 
Grieve, Dr. Janet Reid, and an anonymous 
reviewer carefully reviewed the manuscript. 
This work was supported in part by the 
Monterey Bay Aquarium Research Insti- 
tute, and by NSF OCE-9012340 to Mary 
Silver and NSF OCE-9015602 to C. Pil- 
skaln. 


Literature Cited 


Alldredge, A. 1972. Abandoned larvacean houses: a 
unique food source in the pelagic environ- 
ment.—Science 117:885-887. 

1976. Discarded appendicularian houses as 
sources of food, surface habitats, and particulate 
organic matter in planktonic environments. — 
Limnology and Oceanography 24:14—23. 
Barham, E. 1979. Giant larvacean houses: observa- 

tions from deep submersibles.—Science 205: 
1129-1131. 

Bowman, T., & L. Abele. 1982. Classification of the 
recent Crustacea. Pp. 1-27 in D. E. Bliss, ed., 
The biology of Crustacea 1:xx + 319. 

Bradford, J. 1973. Revision of family and some ge- 
neric definitions in the Phaennidae and Scole- 
cithricidae (Copepoda: Calanoida).— New Zea- 
land Journal of Marine and Freshwater Research 
7:133-153. 

Esterly, C. 1911. Third report on the Copepoda of 
the San Diego region. — University of California 
Publications in Zoology 6:313-352. 

Ferrari, F. 1988. Developmental patterns in numbers 
of ramal segments of copepod post-maxillipedal 
legs. —Crustaceana 54:256-293. 


VOLUME 106, NUMBER 3 


—., &J. Ambler. 1992. Nauplii and copepodids 
of the cyclopoid copepod Dioithona oculata (Oi- 
thonidae) from a mangrove cay in Belize.— Pro- 
ceedings of the Biological Society of Washington 
105:275-298. 

Galt,C. 1979. First records ofa giant pelagic tunicate, 
Bathochordaeus charon (Urochordata, Larva- 
cea), from the eastern Pacific Ocean, with notes 
on its biology.— Fishery Bulletin 77:514-519. 

Giesbrecht, W. 1892. Systematik und Faunistik des 

pelagischen Copepoden des Golfes von Neapel 

und der angrenzenden Meeres-abschnitte. — 

Fauna und Flora des Golfes von Neapel und der 

angrenzenden Meeres-abschnitte 19:1-831 + 54 

pits. 

. 1893. Mittheilungen iiber Copepoden 1-6.— 

Mittheilungen aus der Zoologischen Station zu 

Neapel 11:56-106. 

Grice, G. 1962. Calanoid copepods from equatorial 
waters of the Pacific Ocean.—Fishery Bulletin 
61:167-246. 

Hamner, W., & B. Robison. 1992. In situ observa- 
tions of giant appendicularians in Monterey 
Bay.— Deep-Sea Research 39:1299-1313. 

Hulsemann, K. 1991. The copepodid stages of Dre- 
panopus forcipatus Giesbrecht, with notes on the 
genus and a comparison to other members of 
the family Clausocalanidae (Copepoda Calan- 
oida).— Helgolander Meeresuntersuchungen, 45: 
199-224. 

Huys, R., & G. Boxshall. 1991. Copepod evolution. 
The Ray Society, London, 648 pp. 

Izawa, K. 1987. Studies on the phylogenetic impli- 
cations of ontogenetic features in the poecilo- 
stome nauplii (Copepoda: Cyclopoida).— Pub- 
lications of the Seto Marine Biological 
Laboratory 32:151-217. 

Ohtsuka, S., & N. Kubo. 1991. Larvaceans and their 
houses as important food for some pelagic co- 
pepods.— Bulletin of the Plankton Society of Ja- 
pan, Special Volume, 535-551. 

Park, T. 1970. Calanoid copepods from the Carib- 
bean Sea and Gulf of Mexico. 2. New species 
and new records from plankton samples. — Bul- 
letin of Marine Sciences 20:472-546. 

Pilskaln, C., M. Silver, D. Davis, K. Murphy, B. Grit- 
ton, S. Lowder, & L. Lewis. 1991. A quanti- 


489 


tative study of marine aggregates in the mid- 
water column using specialized ROV instru- 
mentation.—Oceans °91 Proceedings 2:1175-— 
1182. 

Roe, H. 1975. Some new and rare species of calanoid 
copepods from the northeastern Atlantic. — Bul- 
letin of the British Museum (Natural History), 
Zoology 28:297-372. 

Sars, G. 1902. Copepoda Calanoida, part V & VI 
Scolecithricidae, Diaixiidae, Stephidae, Thary- 
bidae, Pseudocyclopiidae. Pp. 49-72 + plts 34— 
48 in An account of the Crustacea of Norway, 
with short descriptions and figures of all of the 
species, 4:144 + 96 plts. The Bergen Museum, 
Bergen. 

Scott, T. 1894. Report on the Entomostraca from the 
Gulf of Guinea, collected by John Rattray, 
GSc.—Transactions of the Linnean Society of 
London (ser. 2, Zoology) 6:1-161 + plts 1-15. 

Sewell, R. 1929. The Copepoda of Indian Seas. Cal- 
anoida.— Memoires of the Indian Museum 10: 
1-221. 

Silver, M., C. Pilskaln, & D. Steinberg. 1991. The 
biologists’ view of sediment trap collections: 
problems of marine snow and living organisms. 
Pp. 76—93 in H. Wassman, A. Heiskaven, & O. 
Linidahl, eds., Sediment trap studies in nordic 
countries, Symposium Proceedings. 

Vaupel Klein, J. von. 1982. A taxonomic review of 

the genus Euchirella Giesbrecht, 1888 (Cope- 

poda, Calanoida). II. The type-species Euchi- 
rella messinensis (Claus, 1863). A. The female 

of f. typica.—Zoologische Verhandelingen, 198: 

1-131 + 23 plts. 

. 1972. Anew character with systematic value 

in Euchirella (Copepoda, Calanoida).—Zoolo- 

gische Mededelingen 47:497-512 + 6 plts. 


(FDF) Invertebrate Zoology, National 
Museum of Natural History, Smithsonian 
Institution, Washington, D.C. 20560, 
U.S.A.; (DKS) Division of Natural Sci- 
ences, University of California, Santa Cruz, 
California 95064, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(3), 1993, pp. 490-496 


A NEW SPECIES OF BENTHANA BUDDE-LUND 
FROM BRAZILIAN CAVES 
(CRUSTACEA; ISOPODA; ONISCOIDEA) 


Idalina Maria Brasil Lima and Cristiana Silveira Serejo 


Abstract. — Benthana iporangensis, a new species of terrestrial isopod inhab- 
iting caves of Sao Paulo, Brasil, is described, figured and compared with the 
closest species: Benthana longipenis and Benthana santosi. The position of the 
‘“‘noduli laterales”’ is presented for the three species here studied. 


Many studies (Jackson 1926; Van Name 
1936; Verhoeff 1941; Verhoeff 1951; Gru- 
ner 1955; Lemos de Castro 1958, 1985) have 
contributed to the knowledge of the genus 
Benthana Budde-Lund, 1908. 

Lemos de Castro (1958b) reviewed the 
genus, described seven new species and pro- 
vided a detailed redescription and a key for 
all hitherto known species. He described two 
additional new species, B. moreirai and B. 
dimorpha (1985). 

The genus Benthana Budde-Lund, 1908 
resembles Ctenoscia Verhoeff, 1917, Ben- 
thanoides Lemos de Castro, 1965 and Ben- 
thanops Barnard, 1932, in having inner 
ctenate teeth on outer lobe of maxillula. 

Benthana is easily distinguished by the 
presence of a sharp curved tooth on the lat- 
eral margin of male first exopodite of pleo- 
pod. 

A new species of the genus Benthana is 
here described based on specimens collect- 
ed in Brazilian caves. A diagnosis of the 
genus is also presented. 


Benthana Budde-Lund, 1908 


Diagnosis.—Cephalon frontal line not 
visible, lateral lobes small, suprantennal line 
very prominent. Eyes small with numerous 
ommatidea. Mandibles with three penicilli 
on the right appendage and two on the left. 
Outer lobe of the maxillula with four simple 
outer spines, five large ctenate ones and a 
small and simple one. Endite of maxilliped 


with an anterior and a posterior spine and 
several short teeth at distal margin. Pleon 
abruptly narrower than pereion. Exopodites 
of the first male pleopod provided with a 
sharp, curved tooth on posterior margin. 
Endopodites of second male pleopod with 
a distal extremity very slender and long. 
Telson triangular, with sides straight or 
slightly sinuous, extremity subacute. Both 
rami of the uropods inserted at same level 


-or very close, always posterior to tip of tel- 


son. In some species sexual dimorphism in 
pereopods 1-3 and 7 and in the proportions 
of uropods exopodites and endopodites, 
longer in male than in female. 

Type species.—Benthana picta (Brandt, 
1833). 


Benthana iporangensis, new species 
Figs. 1-4 


Type material. —Museu Nacional collec- 
tion. Iporanga, Sao Paulo— Aguas Quentes 
Caves: 2 6¢6 and 3 99, Eleonora Trajano leg., 
November 1983, 1 6 Holotype MN 239.MI; 
1 6 and 3 92 Paratypes MN 240.MI; 1 4, L. 
Ishibe leg., August 1978, Paratype MN 
241.MI—Areias de Cima Caves: | 6, E. Tra- 
jano leg., November 1983, Paratype MN 
242.MI; 1 6, E. Trajano leg., September 
1985, Paratype MN 243.MI—Areias de 
Baixo Caves: | 9, E. Trajano leg., February 
1985, Paratype MN 244.MI. 

Description. — Adult male: 6.4-8.6 mm. 
Body subconvex 3.3 times as long as broad. 


VOLUME 106, NUMBER 3 491 


A 


0.1mm 


01mm 


Fig. 1. Benthana iporangensis new species. Male. A: maxilla; B: dorsal view; C: right mandible; D: maxilliped; 
E: maxillule; F: cephalon, dorsal view. 


492 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Benthana iporangensis new species. Male. A: pereopod 1; B: pereopod 2; C: pereopod 3; D: pereopod 
7. (Figs. to same scale: B and D.) 


Body surface smooth, without pigmenta- with the lateral parts concave. Eyes small 
tion. Head big ('4 of the total body length), with about 18 ommatidia without pigmen- 
as wide as long. Lateral lobes small. Su- tation. Antennula surpassing the supranten- 
prantennal line distinctly marked, sinous, nal line. Antennae reaching the end of fourth 


VOLUME 106, NUMBER 3 493 


Fig. 3. Benthana iporangensis new species. Female. A: pereopod 1; B: pereopod 2; C: pereopod 7; Male. D: 
antenne. (Figs. to same scale: A and B.) 


494 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ee 
= 
e 
nl 
=) 


0.5 


it iv Vv VI vil 


| 


on 
(@) 
x 


i iv v vi Vil 


lH VV vi vn 


| uo mw Vv ViVi 


Fig. 4. Benthana iporangensis new species. Male. A: pleopod 2; B: telson and uropods; €: pleopod 1; Noduli 
laterales: (b/c coordinates). D: B. longipenis Lemos de Castro, 1958; E: B. santosi Lemos de Castro, 1958; F: 
B. iporangensis n. sp.—(d/c coordinates). G: B. longipenis Lemos de Castro, 1958; H: B. santosi Lemos de 


Castro, 1958; I: B. iporangensis n. sp. 


VOLUME 106, NUMBER 3 


thoracic segment when well drawn back. 
Flagellum as long as fifth joint, triarticulate, 
setose; its second joint smaller than first and 
third ones, which are subequal. Mandibles 
with molar penicil consisting of numerous 
fringed setae. Outer branch of the maxillula 
with four conic teeth; the first one is the 
longest and the second is the smallest; inner 
branch with six teeth; the fifth one is simple 
and smaller than the others that are ctenate. 
Maxilla with the inner lobe rounded and 
very setose; the outer one has the superior 
margin a little concave in the middle. 

Pereon slightly convex; all pereonal epi- 
mera a few detached, visible only on ventral 
view. Anterolateral angles of pereonal seg- 
ments 1-3 project forward; segments 5-7 
with posterior angles well projected back- 
ward, the seventh reaches the anterior mar- 
gin of the third pleonite. The fourth segment 
has the posterior and anterior angles almost 
straight. ““Noduli laterales’? short, ap- 
proaching to the posterior margin in a de- 
creasing way from pleonites 1-7 (Fig. 4F, 
b/c coordinate); the fourth pleonite with the 
““nodulus lateralis’’ distinctly more distant 
from the lateral margin than the others (Fig. 
41, d/c coordinate). 

Pleon narrow, a few convex laterally, neu- 
pleurons of segments 3-5 downwards di- 
rected, its extremities distinctly separate 
when seen from above. Exopodite of male 
first pleopod with distal margin round with 
a few short setae; lateral margin very con- 
cave with a sharp dentiform extension. En- 
dopodite short and thick with a row of min- 
ute teeth on medial margin. Endopodite of 
the second male pleopod narrow with a long 
capillary extremity; exopodite without se- 
tae. Telson twice as wide as long; lateral 
Margins straight; tip of telson proeminate 
and subacute. Uropods shorter than pleon. 
Protopodite exceeding beyond the telson tip, 
with a furrow on lateral margin. Exopodites 
and endopodites inserted in the same level, 
both furrowed laterally, on the outer and 
inner sides. Setae are present on medial 
margins. 


495 


Sexual dimorphism: Male pereopods 1- 
3 with merus and carpus more densely se- 
tose. Ischium of female pereopod 7 with the 
distal part provided with only a spine. The 
ratio between the length exopodite and en- 
dopodite of uropods smaller in males than 
in females. 

Length of the largest specimen male: 8.6 
mm; female: 10.4 mm. 

Etymology. —The specific name refers to 
the locality where the specimens were col- 
lected. 

Remarks. —Examination of comparative 
material of Benthana picta(Brandt, 1833) 
taken from different states of Brazil, Rio de 
Janeiro (95 specimens), Sao Paulo (110 
specimens), Minas Gerais (15 specimens), 
Parana (3 specimens), Brasilia (25 speci- 
mens) shows that it is readly distinguished 
from the new species by having: a. a visible 
depression in the distal dorsal part of pleo- 
telson; b. the ratio of the uropod endopod/ 
exopod length that is 1/1 in B. picta and 2/3 
in B. iporangensis. 

Benthana iporangensis closely resembles 
Benthana longipenis Lemos de Castro, 1958 
and Benthana santosi Lemos de Castro, 
1958 by the similarities of telson and uro- 
pods, but the uropod endopod is compar- 
atively shorter than in these two species. 

Benthana iporangensis differs from Ben- 
thana longipenis and Benthana santosi by 
having: a) head as wide as large with rela- 
tively small eyes; b) second antennae con- 
siderably shorter; c) maxillula with the outer 
spine not ctenate and medium sized: d) 
maxillipeds with different numbers of spines 
on endite and palp; e) first male pleopods 
by the position of setae, shape of the den- 
tiform expansion of the exopodite and by 
length and width of endopodite; f) second 
male pleopods by the exopodite without se- 
tae and by the different shape of endopodite; 
it also differs in the position of the “‘noduli 
laterales.”’ 

The three related species have been re- 
corded from the same southern part of Bra- 


496 


zil (Sao Paulo, Rio de Janeiro and Minas 
Gerais). 


Acknowledgments 


We are thankful to Dr. Eleonora Trajano 
(Departamento de Zoologia, Instituto de 
Biociéncias, Universidade de Sao Paulo) 
who gave the material for study. 


Literature Cited 


Barnard, K.H. 1932. Contributions to the crustacean 
fauna of South Africa. 11. Terrestrial Isopoda. — 
Annals of the South African Museum, Cape 
Town 30:179-388. 

Brandt, J. F. 1833. Conspectus monographie Crus- 
taceorum Oniscodorum Latreilii.— Bulletin de 
la Societé Impériale des Naturalistes de Moscou 
6:171-193. 

Budde-Lund,G. 1908. Isopoda von MadagasKar und 
Ostrafika mit Diagnosen verwandter Arten. Pp. 
263-308, tafn. XII-X VII in A. Voeltzkow, Reise 
in Ostrafika in den Jahren 1903-1905. Wissen- 


schaftliche Ergebenisse. Stuttgart, Band 2, Heft 


4. 

Gruner, H. E. 1955. Die Gattung Benthana Budde- 
Lund, 1908 (Isopoda, Oniscoidea).—Zoolo- 
gischer Jahrbiicher., Jena, Band 83, Heft 6:441- 
452. 

Jackson, H.G. 1926. Woodlice from Spain and Por- 
tugal, with an account of Benthana, a sub-genus 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


of Philoscia-Crustacea.— Proceedings of the 
Zoological Society of London 1:183-201. 

Lemos de Castro, A. 1958a. Benthanoscia longicau- 

data, a new genus and species of terrestrial iso- 

pod of the family Oniscidae (Isopoda Oniscoi- 
dea).— American Museum Novitates, New York 

1884:1-7. 

1958b. Revisao do género Benthana Budde- 

Lund, 1908 (Isopoda Oniscidea).— Archivos do 

Museu Nacional, Rio de Janeiro 46:85-118. 

. 1985. Duasespécies novas brasileiras de Ben- 

thana Budde-Lund, 1908 (Isopoda, Oniscoidea, 

Philosciidae).— Revista Brasileira de Biologia, 

Rio de Janeiro 45(3):241-—247. 

Van Name, W. G. 1936. The American Land and 
Fresh-Water isopod Crustacea— Bulletin of the 
American Museum of Natural History 71:1—535. 

Verhoeff, K. W. 1917. Zur Kenntnis der Atmung und 

der Atmungsorgane der Isopoda Oniscoidea. 

Ueber Isopoden. 20Aufsatz. — Biologischer Zen- 

tralblat Erlangen & Leipzig 37:133-127. 

1941. Zur Kenntnis sudamerikanischer On- 

iscoideen. — Zoologischer Anzeiger 133:114—126. 

1951. Land-Isopoden. Pp. 69-76 in E. Tit- 

schack, Beitrage zur Fauna Perus, 2. 


(IMBL & CSS) Departamento de Inver- 
tebrados, Museu Nacional— Universidade 
Federal do Rio de Janeiro— Quinta da Boa 
Vista, Sao Cristovao— Rio de Janeiro, C.E.P. 
20940-040. 


PROC. BIOL. SOC. WASH. 
106(3), 1993, pp. 497-507 


LINCA PINITA, A NEW PHOXOCEPHALID GENUS 
AND SPECIES (CRUSTACEA: AMPHIPODA) FROM 
THE ARGENTINE CONTINENTAL SHELF 


Gloria M. Alonso de Pina 


Abstract.—Linca pinita, a new genus and species belonging to the family 
Phoxocephalidae, is described from the benthos of the Argentine continental 
shelf, southwest Atlantic, based on a unique specimen, a female, collected 
during benthos dredging carried out by the R/V Shinkai Maru. It is closely 
related to species in the Birubiinae and Brolginae groups with which it shares 
many characters. Although the new taxon is convergent with the Brolginae, it 


is best placed in the Birubiinae. 


The unique specimen studied herein was 
collected by the R/V Shinkai Maru, through 
sampling of benthic communities on the Ar- 
gentine continental shelf, during four cruis- 
es (IV, V, X and XI) carried out in 1978 
and 1979. The material was obtained by 
means of a Picard dredge, the surface sed- 
iment was represented primarily by sand 
and the grain size was less than 2 mm. 

The description of the new genus and spe- 
cies follows the style of Barnard & Drum- 
mond (1978) who used a standardized form 
established for identification of phoxoce- 
phalids. The entire amphipod was mea- 
sured drawing a lateral parabolic line from 
the tip of the rostrum to the posterior ex- 
treme of the urosome. 


Linca, new genus 


Diagnosis. —Female only, male un- 
known. Eyes not apparent in preserved 
specimens. Antennae 1-2, flagella not re- 
duced. Antenna 1, article 2 elongate, ventral 
setae proximal and midventral. Antenna 2, 
article 1 simple, without any process; spines 
on article 4 in 3 rows. Molar reduced to a 
small protrusion, pillow-shape, bearing 2 
short and 1 elongate spines, and pubescent, 
palpar hump small. Maxilla 1, palp biarti- 
culate; inner plate with 4 setae. Maxilliped, 


apex of palp article 3 not protuberant, dactyl 
elongate, apical segment distinct, medium. 

Gnathopods subchelate, small, but dis- 
similar, gnathopod 2 slightly larger than 
gnathopod 1. Gnathopod 1, article 5 longer 
than in gnathopod 2, with posterior margin 
free, not hidden by articles 4 or 6; hand of 
gnathopods 1-2 respectively ovate and 
slightly broadened, poorly setose anteriorly. 
Pereopod 5, article 2 of broad form, not 
tapering distally. 

Uropod 1, peduncle without ventral spike- 
like spine, without enlarged displaced spine; 
inner ramus with 1 row of marginal spines. 
Uropods 1-2, rami lacking spination along 
their dorsal margin. Uropod 3, article 2 of 
outer ramus carrying 2 elongate setae. Tel- 
son without special dorsal and lateral spines 
or setae. Epimerae 1-2 lacking long poste- 
rior setae. Urosomite 3 without dorsal hook 
or special process. 

Description. — Rostrum fully developed, 
not constricted. Prebucal parts ordinary; 
right mandibular incisor with 4 teeth; right 
lacinia mobilis divided into two parts; man- 
dibular palp, article 1 short, palp medium, 
apex of article 3 oblique, article 2 without 
outer setae. Maxilla 1, outer plate with 11 
spines, one spine specially thickened. Max- 
illa 2, outer and inner plates moderately spi- 


498 


nose. Maxilliped, inner plate with 1 main 
spine, of reduced setosity. 

Gills present on coxae 2—7. Coxae 2-4 
without special anterodorsal humps. Per- 
eopods 3-4, article 5 bearing setae proxi- 
mally on posterior edge; article 6 with all 
posterior spines thick and stiff, midapical 
spine or seta absent. Pereopods 5-6, article 
2 setae absent posteriorly; articles 4—5 nar- 
row. Pereopod 7 of normal size, article 2 
naked ventrally, without facial setae; article 
3 not especially enlarged; dactyl normal. 

Epimeron 2 with 2 medial setae above 
ventral ridge. Epimeron 3 bearing 4 long 
setae. Urosomite | without large lateral fa- 
cial spines, bearing 2 ventral and midven- 
tral crescents or bundles of setae. 

Uropod 1, peduncle normally elongate, 
with dorsolateral spine situated apically, 
medial spines widely spread. Uropod 2, pe- 
duncle with only 1 medial spine distally. 
Uropods 1-2, peduncles with faint comb on 
their distal corners. Uropod 3, peduncle 
lacking extra subapical setae or spines. Tel- 
son, each lobe with 2 apical spines plus small 
plumose seta. 

Type species. — Linca pinita, new species. 

Etymology.—The generic name is de- 
rived from the nickname of Lin-Calel 
meaning “white meat’ in the Indian lan- 
guage for a region of Argentina. The specific 
name refers to the familiar diminuitive of 
the surname of my husband. 

Relationships. —The new genus has many 
similarities with the Birubiinae and Brol- 
ginae generic groups. It seems difficult to 
separate from the Birubiinae, but it has three 
spines on the molar with one of them very 
enlarged, and the apices of the peduncles of 
uropods 1-2 bear combs very faint. This 
genus differs from the Brolginae group in 
the elongate article 2 of antenna 1 and the 
normal as opposed to reduced size of article 
5 of antenna 2. 

Linca differs from Birubius Barnard & 
Drummond, 1978, in those characters men- 
tioned above for the birubiins, namely: 
number of spines on the molar and combed 
corners of peduncles of uropods 1-2. They 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


can be separated also by the single thick 
apical spine on the inner plate of maxilliped, 
by the ventral setae on peduncle of uropod 
1, and by the setation of urosomite 1 which 
in Birubius is generally naked or with sparse 
apicoventral setae or spines near base of 
uropod 1. 

The new genus resembles the brolgin ge- 
nus Fuegiphoxus Barnard, 1980, in the en- 
larged spine of the molar, in possesing a 
main spine only on the inner plate of the 
maxillipeds, in the presence of a midapical 
spine or seta on article 6 of pereopods 3-4, 
and in the combed peduncular apices of uro- 
pods 1-2. They differ in the elongate article 
2 of antenna 1, in the midventrally dis- 
possed main setae on article 4 of antenna 1 
(which in Fuegiphoxus is confined apically), 
in the presence of pubescence on the molars, 
and in that article 5 of gnathopod 2 is not 
shortened in the female as in Fuegiphoxus. 

Linca also resembles the brolgin genus 
Eyakia Barnard, 1979, in the enlarged spine 
of the molars, the elongate article 2 of an- 
tenna 1, and the presence of pubescence on 
the molars, but it differs from that genus in 
having additional proximal spines on article 
4 of antenna 2, one main spine (not two) on 
the inner plate of the maxillipeds, an un- 
tapered article 2 of pereopod 5, the pedun- 
cular apices of uropods 1—2 combed (they 
usually are in the Brolginae, but not in 
Eyakia), and the absence of a midapical 
spine or seta on article 6 of pereopods 3-4. 

The characters that separate the subfam- 
ilies Birubiinae and Brolginae are not com- 
pletely defined. The mandibular molars have 
little significance compared to the strong 
weighting of the length and setation of ar- 
ticle 2 of antenna 1 in all the Phoxoce- 
phalidae subfamilies. The main character 
bringing Linca close to the Brolginae is the 
presence of three molar spines, but it is best 
placed with the Birubiinae because of the 
elongate article 2 of antenna 1, and the com- 
bination of spination placement on article 
2 of antenna 1, the size of article 5 of an- 
tenna 2, and the setation on the inner plate 
of maxilla 1. 


VOLUME 106, NUMBER 3 


B 1ooy 


499 


(wy 


Figs. 1-4. Linca pinita, new genus, new species: 1, antenna 1; 2, right mandible; 3, left lacinia mobilis; 4, 


mandibular palp. Scales: A, Figs. 1, 4; B, Figs. 2, 3. 


Linca pinita, new species 
Figs. 1-32 


Description. —Head about 16% of the to- 
tal body length, greatest width about 43% 
of length; rostrum unconstricted, broad, 
elongate, exceeding middle of article 2 of 
antenna 1. Eyes not apparent in preserved 


specimens. Antenna | (Fig. 1), article 1 of 
peduncle about 1.5 times as long as wide, 
about 2.4 times as wide as article 2, ventral 
margin with 3 plumose setae, dorsal apex 
weakly produced with 1 small seta; article 
2 about 0.8 times as long as article 1, with 
midventral row of 6 setae and 8 shorter 
proximal setae, primary flagellum with 12 


500 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


200y 


Figs. 5,6. Linca pinita, new genus, new species: 5, peduncle of antenna 2; 6, flagellum of antenna 2. 


articles, about 0.6 times as long as peduncle, 
articles 2-9 each bearing | short aesthetasc; 
accessory flagellum with 10 articles. Anten- 
na 2 (Figs. 5, 6), article 1 without cone-like 
process; article 3 with thin anterodorsal 
spine and 2 small submarginal setae; spine 
formula of article 4 = 3-3-7, dorsal margin 


with 3 notches bearing 6 setae, 6 setae and 
1 spine and 3 setae and | spine (distal to 
proximal), ventral margin with about 11 
groups of 1-3 long to medium setae, 1 long 
ventral spine distally; article 5 about 0.7 
times as long as article 4, with single facial 
spine, dorsal margin bearing | set of distal 


VOLUME 106, NUMBER 3 


A__200Yy 


13 


B__500yY 


Figs. 7-13. Linca pinita, new genus, new species: 7, maxilla 1; 8, maxilla 2; 9, maxilliped; 10-12, coxae 1- 


3; 13, coxa 5. Scales: A, Figs. 7-9; B, Figs. 10-13. 


medium to short setae, ventral margin with 
8 sets of 1 long seta each, 3 ventrodistal long 
to medium spines set facially; flagellum 
about 0.8 times as long as articles 4—5 of 
peduncle combined, with 13 articles. 
Mandibles weakly extended near palpar 


attachment, right incisor with 4 teeth, left 
incisor composed of two divided branches: 
one of them with | tooth and the other one 
with 2 teeth; right lacinia mobilis bifid, dis- 
tal branch shorter than proximal, flabellate, 
tridentate, proximal branch simple, point- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 14-17. 
pereopod 3. Scales: A, Figs. 14-16; B, Fig. 17. 


ed, left lacinia mobilis (Fig. 3) with 4 teeth, 
middle teeth shortened; right rakers 17 plus 
1-2 reduced in size, left rakers 15 plus 1-2 
much smaller; molars composed of bulbous 
protrusions, each with 1 long serrate and 2 
short spines, molars without plume but pu- 


Linca pinita, new genus, new species: 14, coxa 4; 15, 16, gnathopods 1-2; 17, article 6 of 


besent (Fig. 2); palp (Fig. 4), article 1 short, 
article 2 carrying 1 long inner apical seta 
and 4 other short inner setae, article 3 about 
1.2 times as long as article 2, oblique apex 
with about 10 spine-setae, basofacial for- 
mula = 4—2. Maxilla 1 (Fig. 7), inner plate 


VOLUME 106, NUMBER 3 503 


Figs. 18-21. Linca pinita, new genus, new species: 18, pereopod 3; 19, article 6 of pereopod 4; 20, pereopod 
4; 21, epimeron 1. Scales: A, Figs. 18, 20, 21; B, Fig. 19. 


thin, bearing | long apial plumose seta, 1 submarginal setae. Maxilla 2 (Fig. 8), inner 
shorter apical plumose seta medially and 2 _ plate shorter than outer, with 4 short medial 
much shorter lateral simple setae apically; setae, outer not broader than inner, with 5 
outer plate with 11 spines; palp article 2 apical setae on lateral edge. Maxilliped (Fig. 
with 7 apical medial marginal spines and 6 9), inner plate with | large thick apical spine, 


504 


Figs. 22-25. 
uropod 2. 


4 submarginal setae apically and 4 medial 
setae; outer plate with 14 medial and apical 
spines and 1 apical seta on lateral edge; palp 
article 1 without lateral setae distally, article 
2 with 2 groups of 1 and 4 apical setae on 
lateral edge, medial margin of article 2 mod- 


WN, 


WQXQ A 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


\ Ss : 


ig <4 
J 
z. 


WSS 


500 Y 


Linca pinita, new genus, new species: 22, pereopod 5; 23, pereopod 7; 24, epimeron 2; 25, 


erately setose, article 3 with 6 facial setae, 
3 lateral setae, article 4 bearing 2 accessory 
small setae, with nail medium in length. 
Coxa 1 (Fig. 10) not expanded apically, 
anterior margin almost straight; main ven- 
tral setae of coxae 1—4 = 23-19-19-21, pos- 


VOLUME 106, NUMBER 3 


505 


Figs. 26-29. Linca pinita, new genus, new species: 26, pereopod 6; 27, article 6 of pereopod 6; 28, epimeron 


3; 29, uropod 1. 


teromost seta of coxa | longest, of coxae 2- 
3 slightly shorter (Figs. 11, 12), of coxa 4 
very short; anterior and posterior margins 
of coxa 4 (Fig. 14) divergent, posterior mar- 
gin very oblique, posterodorsal corner 
rounded, posterodorsal margin short, con- 


cave, width-length ratio of coxa 4 = 149: 
176. Gnathopods generally ordinary (Figs. 
15, 16), gnathopod 2 weakly enlarged, width 
ratios of articles 5—6 of gnathopods 1-2 = 
41:47 and 41:53, length ratios = 90:88 and 
78:90; palmar humps ordinary, palms 


506 


Figs. 30, 31. JLinca pinita, new genus, new species: 
30, uropod 3; 31, telson. 


strongly oblique; article 5 of gnathopod 1 
elongate, ovate, posterior margin flat, article 
5 of gnathopod 2 elongate, ovate, posterior 
margin rounded. Pereopods 3-4 (Figs. 17- 
20) similar, but article 5 of pereopod 4 
slightly stouter than that of pereopod 3, sub- 
marginal setae formula on article 4 of both 
pereopods = 9 and 10, parallel to apex, setae 
formula on article 5 = 9 and 13, longest 
spine on article 5 extending to apex of article 
6, article 5 without proximal spines poste- 
riorly, spine formula of article 6 = 8 plus 8 
and 8 plus 10 but without middistal seta, 
clump of apical spines thin and long; ac- 
clivity on inner margin of dactyls of per- 
eopods 3-4 represented by slit in which is 
inserted a short seta. Coxa 5 (Fig. 13), 6 and 
7 posteroventral seta formula = 10-41. 
Pereopods 5-6, articles 4—5 narrow, facial 
spine rows sparse, facial ridge formula of 
article 2 on pereopods 5—7 = Q-1-1. Width 
ratios of articles 2, 4, 5, 6 of pereopod 5 = 
82:57:44:27, of pereopod 6 = 159:72:49:27, 
of pereopod 7 = 184:53:45:23, length ratios 
of pereopod 5 = 169:80:82:90, of pereopod 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


6 = 204:116:106:108, of pereopod 7 = 231: 
67:57:78; article 2 of pereopod 7 reaching 
apex of article 4, posterior margin with 7 
medium serrations, medial apex of article 
6 apparently with scarcely comb setae, bear- 
ing 6-7 digital processes (Figs. 22, 23, 26, 
2H: 

Epimeron | (Fig. 21), posteroventral cor- 
ner rounded, posterior margin weakly con- 
vex, with 2 widely spaced very short setae, 
anteroventral margin with 10 medium se- 
tae, ventral face with about 16 long setae; 
epimeron 2 (Fig. 24) quadrate, posteroven- 
tral corner rounded, posterior margin with 
1 small seta, about 14 facial setae in irreg- 
ular row; epimeron 3 (Fig. 28), posteroven- 
tral corner rounded, posterior margin with 
4 setae, each inserted in an indentation of 
the edge, ventral margin with 5 spread setae 
in posterior half. 

Urosomite | with lateral seta at base of 
uropod 1, brush of setae ventral to uropod 
1 and brush of midventral setae, articula- 
tion line complete, urosomites smooth. 
Uropod 1 (Fig. 29), peduncle with 7 baso- 
facial setae, ventral cluster of 7 setae, 1 lat- 
eral spine distally, 7 marginal spines me- 
dially and 1 distal spine on lateral edge; inner 
and outer rami with 3 dorsal spines. Uropod 
2 (Fig. 25), peduncle with about 9-10 dorsal 
spines, basalmost thin and elongate, 1 me- 
dium apical spine medially; outer ramus 
with 1 dorsal spine, inner with 2 dorso- 
medial spines. Uropods 1-2, corners of pe- 
duncles with fine apical comb-like setae on 
lateral margin (only seen with high power); 
rami with articulate but tightly fixed ter- 
minal spines. Uropod 3 (Fig. 30), peduncle 
with 8 ventral spines, dorsally with 1 lateral 
spine and 1 medial spine and seta; rami of 
different length, inner extending to M. 87 
on article 1 of outer ramus, apex with 2 
setae, medial and lateral margin with 0 and 
1 sesta; article 2 of outer ramus short, 0.12, 
bearing 2 long setae; | apical seta on medial 
margin of article 1, lateral margin with up- 
ward slope, carrying 4 spines, apex with 3 


spines laterally. Telson (Fig. 31), length- 


VOLUME 106, NUMBER 3 


1laooy 


507 


Fig. 32. Linca pinita, new genus, new species: lateral view. 


width ratio = 60:53, almost fully cleft, each 
apex of medium width, rounded, lateral lobe 
broad, shallow, bearing short lateral and long 
medial spines separated by short plumose 
seta, midlateral plumose setae different in 
length, largest of usual size. 

Body smooth (Fig. 32), yellow in alcohol. 

Holotype. —Museo Argentino de Ciencias 
Naturales No. 33444, female, 11.87 mm. 

Type locality. —Argentine continental 
shelf, Shinkai Maru IV, benthic Sta 92: 
50°30’S, 62°31'W, 159 m depth, 14 Aug 
1978, donator Dr. R. O. Bastida. 


Acknowledgments 


I am very grateful to the late Dr. J. Lau- 
rens Barnard for his invaluable suggestions 
with this group of phoxocephalids. 


Literature Cited 


Barnard, J. L. 1979. Revision of American species 
of the marine Amphipod Genus Paraphoxus 
(Gammaridea: Phoxocephalidae).— Proceed- 
ings of the Biological Society of Washington 92: 
368-379. 

. 1980. Two new Phoxocephalid Genera, Fue- 
giphoxus and Phoxorgia, from Magellanic South 
America (Amphipoda: Crustacea).—Proceed- 
ings of the Biological Society of Washington 93: 
849-874. 

—., & M. M. Drummond. 1978. Gammaridean 
Amphipoda of Australia. Part III: The Phoxo- 
cephalidae. —Smithsonian Contributions to Zo- 
ology 245:1-551. 


Museo Argentino de Ciencias Naturales 
““Bernardino Rivadavia,” Division Inver- 
tebrados, Avda. Angel Gallardo 470, (1405) 
Buenos Aires, Argentina. 


PROC. BIOL. SOC. WASH. 
106(3), 1993, pp. 508-513 


THREE NEW SPECIES OF 
STRENGERIANA FROM COLOMBIA 
(CRUSTACEA: DECAPODA: PSEUDOTHELPHUSIDAE) 


Martha R. Campos and Gilberto Rodriguez 


Abstract.—Three new pseudothelphusid crabs, Strengeriana cajaensis, S. 
maniformis, and S. flagellata, are described from the Central Cordillera of 
Colombia. The last species displays a rudimentary flagellum on the exopod of 
the third maxilliped. This peculiar morphology reinforces the hypothesis that 
there exists a close phylogenetic relationship between the species of Strenger- 
jana from northern South America and the species of Epilobocera from the 


West Indies. 


The number of known species of Stren- 
geriana, a group of small, primitive fresh- 
water crabs from northern Colombia, has 
rapidly increased in recent years. The sys- 
tematics and biogeography of the genus have 
been recently reviewed by Rodriguez & 
Campos (1989). Explorations in the Central 
Cordillera of Colombia have revealed the 
presence of three new species, which are de- 
scribed in the present contribution. One of 
these species possesses a rudimentary palp 
in the exopod of the third maxilliped. 

The material is deposited at the Museo 
de Historia Natural, Instituto de Ciencias 
Naturales, Universidad Nacional de Co- 
lombia, Bogota (ICN-MHN). Other abbre- 
viations used are cb. for carpace breadth, 
and cl. for carapace length. 


Tribe Strengerianini Rodriguez, 1982 
Genus Strengeriana Pretzmann, 1971 
Strengeriana cajaensis, new species 
Fig. 1A—F 


Material.—Municipio Cajamarca, creek 
by the side of the Central Highway, 1560 
m above sea level, Tolima Department, Co- 
lombia; 3 Aug 1988; R. Sanchez; 1 male 
holotype, cb. 20.6 mm, cl. 13.2 mm (ICN- 
MHN No CR 0939). 

Description. —The carapace is narrow (cb/ 
cl = 1.57). The cervical grooves are straight 


and deep, reaching the lateral margins. The 
antero-lateral margins have a depression 
behind the orbits followed by a few indis- 
tinct papillae and a second shallow depres- 
sion at the level of the cervical groove; the 
rest of the margins have approximately 12 
ill-defined papillae, which are regularly 
spaced. The postfrontal lobes are small, oval 
shaped and low. The median groove is ab- 
sent. The surface of the carapace between 
the front and the postfrontal lobes is in- 
clined anteriorly and towards the mid-line. 
The upper border of the front is rounded, 
without conspicuous tubercles, and slightly 
bilobed in dorsal view. The lower margin 
is thin, and slightly sinuous in frontal view. 
The surface of the front between the upper 
and lower borders is low. The surface of the 
carapace is smooth and polished; the limits 
between the regions of the carapace are in- 
distinct. 

The palm of the larger (right) chela is 
strongly inflated; the fingers gape moder- 
ately when closed. The walking legs are slen- 
der, but not unusually elongated, the largest 
being those of the second and third pair, 
which are also of the same length (total length 
approximately 1.28 the breadth of cara- 
pace); the merus in the third pair is 3.5 times 
longer than wide. The exopod of the third 
maxilliped is 0.97 times the length of the 


VOLUME 106, NUMBER 3 


509 


Fig. 1. Strengeriana cajaensis, new species, holotype. A—D, Left first gonopod: A, Total view, caudal; B, 
Same, detail of apex, lateral view; C, Same, cephalic view; D, Same, mesial view. E, Third maxilliped, left; F, 
Aperture of efferent channel, left. (bs, basal spine; cl, cephalic lobe; cs, cephalic spine; ll, lateral lobe; mel, mesial 


lobe; ml, marginal lobe; mp, mesial plate.) 


lateral margin of the ischium of the endog- 
nath. The orifice of the efferent branchial 
channel is almost closed by a spine at the 
jugal angle and by the production of the 
lateral lobe of the epistome. 

The male first gonopod is short and stocky 
in caudal view, with a wide expansion placed 
crosswise to the apical portion, where the 
slit-like genital pore is located, and a con- 
spicuous rounded bulge on the lateral side. 
The apical expansion consists of a lateral 
twisting of the marginal lobe (Fig. 1A, B, C, 
ml), a rounded tooth-like lateral lobe (Fig. 
1A-C, Il) which ends in an acuminate tip, 
and a widening of the caudal surface (Fig. 
1A, mp) which gives a hammer-like ap- 
pearance to the appendage. In cephalic view 
the apex presents (1) a bifid mesial lobe (Fig. 
1C, D, mel) enveloped basally by the mesial 


plate (Fig. 1A, C, D, mp), and with a small 
basal spine (Fig. 1C, bs); and (2) a large 
conical cephalic lobe (Fig. 1C, D, cl) with a 
hooked cephalic spine (Fig. 1C, D, cs) on 
its distal surface. In addition to the strong 
caudal setae, the appendage has numerous 
strong spinules on the lateral bulge which 
extend to the lateral surface, the mesial bor- 
der has more slenderer spines, and the distal 
border of the lateral lobe has small spinules. 

Color.—In the holotype specimen pre- 
served in alcohol, the dorsal surface of the 
carapace and pereiopods is uniformly dark 
brown. The ventral surface is cream col- 
ored. 

Etymology. —The specific name refers to 
an abbreviation of Cajamarca, Colombia, 
where the type locality is situated. 

Remarks.—This species can be clearly 


510 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ih cane eis 


mp 


SSS 
Aa 


Fig. 2. Strengeriana maniformis, new species, holotype (except F—I). A-E, Holotype, first left gonopod: A, 
Total view, caudal; B, Same, detail of apex, cephalic view; C, Same, mesial view; D, Third maxilliped, left; E, 
Aperture of efferent channel, left; F, G: immature specimen, left gonopod (F, Detail of apex, cephalic view; G, 
Same, mesial view). Strengeriana flagellata, new species, holotype: H, Third maxilliped, left; I, Aperture of 


VOLUME 106, NUMBER 3 


distinguished from any other in the genus 
by the hammer-shaped first gonopod. The 
slit-like genital pore, with small spinules over 
the caudal surface, and the folded mesial 
plate (Fig. 1A, C, D, mp) are characters it 
shares with other species of the genus. Fur- 
thermore, a number of homologies exist be- 
tween the first gonopod of S. cajaensis and 
Strengeriana chaparralensis Campos & 
Rodriguez, 1984. The bifid mesial lobe (Fig. 
1C, D, mel) has the same appearance and 
direction in both species, but it is shorter in 
S. cajaensis; the basal spine of this lobe (Fig. 
1C, bs) is similar in both species. The large 
conical cephalic lobe (Fig. 1C, D, cl) of S. 
cajaensis is homologous with the spiniform 
cephalic lobe (Rodriguez & Campos 1989, 
Fig. 5, t) of S. chaparralensis; the cephalic 
spine (Fig. 1C, D, cs) is present in both. The 
large lateral bulge, covered with spines, of 
S. cajaensis is rudimentary in S. chapar- 
ralensis. 4 


Strengeriana maniformis, new species 
Fig. 2A—G 


Material. — Municipio Cajamarca, creek 
by the side of the Central Highway, 1560 
m above sea level, Tolima Department, Co- 
lombia; 3 Aug 1988; R. Sanchez; 1 male 
holotype, cb. 32.6 mm, cl. 21.2 mm, 1 ju- 
venile (ICN-MHN No CR 0938). 

Description. —The carapace is narrow (cb/ 
cl = 1.54). The cervical grooves are slightly 
arcuate, thin on the distal half, deep and 
wide proximally, ending far from the lateral 
margins. The antero-lateral margins have a 
depression behind the orbits followed by 5- 
6 papillae and a series of approximately 10 
regularly-spaced small teeth. The postfron- 
tal lobes are small, oval shaped and low, 
marked on their anterior margins by the 
rugosities of the carapace. The median 


— 


S11 


groove is absent. The surface of the carapace 
between the front and the postfrontal lobes 
is moderately inclined anteriorly, and to- 
wards the middle. The upper border of the 
front is well defined, slightly convex in dor- 
sal view, V-shaped in frontal view, marked 
with a row of conspicuous, well defined tu- 
bercles, and interrupted at the middle by a 
deep notch. The lower margin is strongly 
sinuous in frontal view. The surface of the 
front between the upper and lower borders 
is low. The surface of the carapace is smooth, 
covered by small papillae not visible to the 
naked eye; the regions are strongly de- 
marked. 

The palm of the larger chela is moderately 
inflated; the fingers do not gape. The walk- 
ing legs are slender, but not unusually elon- 
gated, the largest being those of the third 
pair (total length approximately 1.13 the 
breadth of carapace); the merus in this pair 
is 2.64 times longer than wide. The exopod 
of the third maxilliped overreaches the lat- 
eral margin of the ischium of the endognath. . 
The orifice of the efferent branchial channel 
is almost closed by a spine at the jugal angle 
and by the production of the lateral lobe of 
the epistome. 

The male first gonopods are short and 
stocky, with a long mesial process (Fig. 
2A, mp) extending back to the middle of 
the appendage; the marginal lobe (Fig. 2A, 
ml) is simple, with a short ridge on its lateral 
surface; the mesial lobe (Fig. 2B, mel) forms 
with the cephalic lobe (Fig. 2B, cl) a long 
slit where the genital pore is located; the 
cephalic lobe bears a strong conical cephalic 
process (Fig. 2A, B, cp) and a strong prox- 
imal cephalic spine (Fig. 2A, cs). The long 
mesial process is oblong, with a strong distal 
spine (Fig. 2A, B, sl), followed proximally 
by a series of spinules (Fig. 2A, B, sp) which 


efferent channel, left. (f, rudimentary flagellum; cl, cephalic lobe; cp, cephalic process; cs, cephalic spine; ml, 
marginal lobe; mp, mesial process; mel, mesial lobe; sl, distal spine of the mesial process; s2, lateral spine of 


the mesial process; sp, spinules of the mesial process.) 


512 


progressively diminish in size, and a slender 
spine Fig. 2A, s2), located near the middle 
of the lateral surface of the process. In ad- 
dition to the strong caudal setae and the 
small spines of the genital pore, the gonopod 
bears scattered long spines on the lateral 
side, small spinules over the distal border 
of the cephalic lobe and a few tiny setae on 
the mesial side. 

Color.—The carapace and pereiopods of 
specimens preserved in alcohol are entirely 
covered by a characteristic mottled pattern 
of light brown over a dark brown back- 
ground. 

Etymology.—The specific name is from 
the Latin ““manus,”’ in reference to the hand- 
shape mesial process of the gonopods, and 
“‘formis,”” shape. 

Remarks.—The species is closely allied 
to Strengeriana huilensis Rodriguez & 
Campos, 1989, but both species can be eas- 
ily distinguished by the characteristic shape 
of the mesial lobe of the first gonopod. 


Strengeriana flagellata, new species 
Fig. 2H, I 


Material. — Patio Bonito, Municipio Co- 
corna, 5O km SE of Medellin, creek by the 
side of the road, 1600 m above sea level, 
Antioquia Department, Colombia; 26 Jul 
1989; G. Susatama; 1 male holotype, cb. 
18.7 mm, cl. 12.3 mm (ICN-MHN No CR 
1194). 

Description. —The carapace is narrow (cb/ 
cl = 1.52). The cervical grooves are shallow 
and straight, ending far from the lateral 
margin. The antero-lateral margin has a de- 
pression behind the orbit followed by a se- 
ries of approximately 5 papillae and 15 small 
teeth which become indistinct posteriorly. 
The postfrontal lobes are small, oval shaped 
and low, obsolescent, not clearly delimited 
anteriorly. The median groove is absent. The 
surface of the carapace in front of the post- 
frontal lobes is moderately inclined ante- 
riorly, and towards the middle. The upper 
border of the front is well defined, bilobed 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


in dorsal view, excavated in frontal view, 
marked with a row of 8 conspicuous, well 
defined tubercles at each side, interrupted 
at the middle by a shallow notch. The lower 
Margin is sinuous in frontal view. The sur- 
face of the front between the upper and low- 
er borders is high. The surface of the cara- 
pace is smooth, covered by small papillae 
not visible to the naked eye; the limits be- 
tween the regions of the carapace are indis- 
tinct. 

The palm of the larger chela is strongly 
inflated; the fingers do not gape. The walk- 
ing legs are slender, but not unusually elon- 
gated, the largest being those of the third 
pair (total length approximately 1.05 the 
breadth of carapace); the merus in this pair 
is 3 times longer than wide. The exopod of 
the third maxilliped overreaches the lateral 
margin of the ischium of exognath, and is 
provided with a rudimentary flagellum. The 
orifice of the efferent branchial channel is 
almost closed by a spine at the jugal angle 
and by the production of the lateral lobe of 
the epistome. 

The male first gonopods are strongly 
arched in the caudocephalic plane. The apex 
is formed by three distinct lobes; the mesial 
and marginal lobes are rounded, cup-shaped, 
with their borders strongly demarked; the 
border of the mesial lobe is strongly bent 
over at their cephalic and caudal ends; the 
cephalic lobe is very wide in lateral view, 
strongly sinuous in cephalic view, with its 
distal margin partially fused to the field of 
spines. There is a small finger-like projec- 
tion located subapically on the mesial sur- 
face. The internal cavity of the mesial lobe 
is densely covered by long spines; there is 
a patch of shorter spines on the external 
surface of the cephalic lobe. 

Color.—In the holotype specimen pre- 
served in alcohol, the dorsal surface of the 
carapace and pereiopods is uniformly light 
brown. The ventral surface is cream col- 
ored. 

Etymology.—The specific name is from 
the Latin “‘flagellum,” a whip, and refers to 


VOLUME 106, NUMBER 3 


the rudimentary flagellum in the exopod of 
the third maxilliped. 

Remarks. — The holotype specimen is not 
fully mature, but even at this stage some 
characters of the first male gonopods are 
discernible. These appendages resemble 
those of Strengeriana fuhrmanni (Zimmer, 
1912), but they are conspicuously expanded 
in the caudo-cephalic plane, and the caudal 
lobe thus formed is strongly prominent and 
sinuous. The species can be clearly distin- 
guished from any other in the genus by the 
unusual morphology of the exopod of the 
third maxilliped which overreaches the lat- 
eral margin of the ischium of the endognath, 
and possesses a rudimentary flagellum (Fig. 
2H, f). This flagellum is absent in all species 
of Pseudothelphusidae, except for some 
species of Epilobocera from the West Indies. 
The unusually long exopod of the species of 
the South American Strengeriana and the 
West Indian Epilobocera has been inter- 
preted by Rodriguez (1986) as a proof of the 
phylogenetic affinity of both genera. The 
presence of a rudimentary flagellum on the 
exopod of the third maxilliped in this spe- 
cies strongly reinforces this point of view. 


Literature Cited 


Campos, M. R., & G. Rodriguez. 1984. New species 
of freshwater crabs (Crustacea: Decapoda: Pseu- 


513 


dothelphusidae) from Colombia.— Proceedings 
of the Biological Society of Washington 97:538— 
543. 

Pretzmann, G. 1971. Fortschritte in der Klassifizie- 
rung der Pseudothelphusidae. — Sitzungsberich- 
ten der Osterreich Akademie der Wissenschaf- 
ten, Mathematisch-Naturwissenschaftliche 
Klasse (1)179(1-4):14-20. 

Rodriguez, G. 1982. Les Crabes d’eau douce d’Ame- 

rique. Famille des Pseudothelphusidae. — Faune 

Tropicale 22:1—223. 

1982. Centers of distribution of Neotropical 
fresh-water crabs. Jn R. H. Gore & K. L. Heck, 
eds., Biogeography of the Crustacea. —Crusta- 
cean Issues 4:51-67. 

—., & M.R. Campos. 1989. Cladistic relation- 
ships of freshwater crabs of the tribe Strenger- 
ianini (Decapoda: Pseudothelphusidae) from the 
northern Andes, with comments on their bio- 
geography and descriptions of new species.— 
Journal of Crustacean Biology 9:141-156. 

Zimmer, C. 1912. Beitrag zur Kentniss der Sisswas- 
ser dekapoden Kolumbiens. Jn O. Fuhrmann & 
E. Mayor, eds., Voyage d’exploration scienti- 
fique en Colombie.— Mémoires de la Societé 
néuchateloise des Sciences naturelles 5:1-8a 


(MRC) Universidad Nacional, Instituto 
de Ciencias Naturales, Apartado Aéreo 
53416, Bogota, Colombia; (GR) Instituto 
Venezolano de Investigaciones Cientificas, 
Apartado 21827, Caracas 1020A, Venezue- 
la. 


PROC. BIOL. SOC. WASH. 
106(3), 1993, pp. 514-522 


FURTHER REMARKS ON THE IDENTITY OF 
SUDANONAUTES ORTHOSTYLIS BOTT, 1955, 
(CRUSTACEA: DECAPODA: POTAMOIDEA: POTAMONAUTIDAE) 
WITH COMPARISONS WITH OTHER SPECIES 
FROM NIGERIA AND CAMEROON 


Neil Cumberlidge 


Abstract. —Sudanonautes orthostylis Bott, 1955 a freshwater crab from West 
Africa is redescribed exclusively from type material. The species is now rec- 
ognized by a combination of characters of the carapace, chelipeds, and gono- 
pods, which are illustrated. It is compared to other species of Sudanonautes 
Bott, 1955, and to the other genera of freshwater crabs occurring in West Africa. 
The species is restricted to the tropical rain forest zone of southwest Cameroon, 


West Africa. 


The rain forest region of southeast Ni- 
geria and southern Cameroon is home to a 
large number of species of freshwater crabs 
of the family Potamonautidae Bott, 1970 
(Bott 1955, 1959, 1964; Monod 1977, 1980; 
Cumberlidge 1989, 1991, 1993a, b; Cumber- 
lidge & Clark 1992). These crabs belong to 
either Potamonautes MacLeay, 1838, Su- 
danonautes Bott, 1955, or Potamonemus 
Cumberlidge & Clark, 1992. Those species 
which possess one or more side spines on 
the carapace and an elongated, flagellum- 
like terminal segment of gonopod 2 belong 
to Potamonautes. Several other species of 
small-bodied freshwater crabs have a car- 
apace lacking side spines and a second gon- 
opod with a short terminal segment. Those 
with a third maxilliped whose exopod lacks 
a flagellum belong to Potamonemus, while 
crabs which possess this flagellum belong to 
Sudanonautes. 

However, identification of the similar- 
looking members of Sudanonautes found in 
the rain forest zone of Nigeria and Came- 
roon is hampered by the lack of a reliable 
taxonomic key to the species. The current 
literature (Bott 1955, 1964; Monod 1980; 
Cumberlidge 1989, 1993a) records five spe- 
cies of Sudanonautes found in this region: 


S. pelii (Herklots, 1861), S. aubryi (H. Milne- 
Edwards, 1853), S. africanus (A. Milne-Ed- 
wards, 1869), S. orthostylis Bott, 1955, and 
S. granulatus (Balss, 1929). 

Sudanonautes orthostylis was first de- 
scribed by Bott (1955) as a subspecies of S. 
decazei. That work included photographs of 
the carapace and chelipeds of the holotype 
from Cameroon and a sketch of the terminal 
segment of gonopod 1. In a later work, Bott 
(1964) considered S. (S.) d. decazei as a ju- 
nior synonym of S. pelii without discussion 
of the subspecies. Sudanonautes pelii (Herk- 
lots, 1861) and S. decazei (A. Milne-Ed- 
wards, 1886) are synonymous taxa (Cum- 
berlidge 1989), but S. orthostylis does not 
appear to be closely related to S. pelii, as 
this study will show. As such, there is a need 
to establish the identity and affinities of S. 
(S.) decazei orthostylis Bott, 1955. 

This anomaly prompted Cumberlidge 
(1989) to redescribe S. orthostylis Bott, 1955, 
based on a large male specimen from the 
Oban Hills, Nigeria, as well as numerous 
additional material from that country. 
Cumberllidge (1989) ascribed differences 
between specimens of S. orthostylis from 
different localities in Nigeria to intraspecific 
variation. Subsequent examination of this 


VOLUME 106, NUMBER 3 


Nigerian material in the light of recent find- 
ings (Cumberlidge & Clark 1992; Cumber- 
lidge 1993a, 1993b) indicates that part of 
this material should be removed from S. 
orthostylis and reassigned to Sudanonautes 
granulatus (Balss, 1929), and part to a spe- 
cies of Potamonemus. 

These developments have made it nec- 
essary to clarify the taxonomy of S. ortho- 
stylis by describing the species exclusively 
from the holotype and paratypes from Cam- 
eroon. The taxonomically important char- 
acters of the carapace, chelipeds, mandible, 
third maxilliped, and gonopods are illus- 
trated. The species is compared to other 
species of Sudanonautes Bott, 1955, and to 
the other genera of freshwater crabs occur- 
ring in West Africa. 


Methods 


The type series of S. orthostylis Bott, 1955 
was examined during a visit to the Zoolo- 
gische Museum der Humboldt-Universitat, 
Berlin, Germany (ZMB), and the holotype 
was subsequently loaned. Two other para- 
types were examined during a visit to the 
Natur-Museum und Forschungs-Institut 
Senckenberg, Frankfurt am Main, Germany 
(SMF). The type specimen of S. pelii was 
loaned from the Nationaal Naturhistorisch 
Museum, Leiden, The Netherlands (NNM). 
The type specimens of S. africanus and S. 
aubryi were examined in the Museum Na- 
tional d’Histoire Naturelle, Paris, France 
(MNHN). Large series of these latter two 
species from other collections were also ex- 
amined. Four dimensions of the carapace, 
carapace length, carapace width, carapace 
height, and front width, were recorded from 
each specimen using digital calipers (Table 
1, Fig. 3a). The relative proportions of the 
latter three measurements (adjusted for body 
size, CL) of S. orthostylis were calculated 
(Fig. 3b). These proportions were also cal- 
culated for the series of the other three spe- 
cies of Sudanonautes. One-factor ANOVA 
repeated measures analysis was used to test 


515 


for significant differences between the mean 
of the carapace proportions of S. orthostylis 
and the three closely related species (Table 
2). The right mandible and the left first and 
second gonopods were removed from the 
specimens in order to describe these struc- 
tures from different views. The length of the 
propodus of the right and left chelipeds of 
males and females was measured longitu- 
dinally along the ventral margin. 

Abbreviations. —CW = carapace width at 
widest point; CL, carapace length, measured 
along median line; CH, carapace height, 
maximum depth of cephalothorax; FW, 
front width, width of front measured along 
anterior margin. 


Sudanonautes orthostylis Bott, 1955 
(Figs. 1-3, Tables 1, 2) 


Sudanonautes (Sudanonautes) decazei or- 
thostylis Bott, 1955:301, fig. 63a, b; pl. 
29, fig. 2a-d. 

Sudanonautes (Sudanonautes) pelii pellti 
Bott, 1964:32 (part) (not Cancer (Thel- 
phusa) pelii Herklots, 1861). 

Sudanonautes orthostylis, Cumberlidge, 
1989:230 (part), figs. la-g, 2a-—. 


Material. —Holotype: male (CW 25.8 
mm), Bipindihof, Cameroon, collected by 
Zenker, ZMB 11093. Paratypes: 8 males, 3 
females, 3 juveniles; ZMB 11093, SMF 
2439. 

Type locality.—Bipindihof, Cameroon. 
This locality is most likely Bipindi (3°08'N, 
10°30’E). 

Diagnosis. —Terminal segment of gono- 
pod | straight for most of its length, curving 
sharply outward just before tip; completely 
lacking longitudinal groove; subterminal 
segment of gonopod | slim (Fig. 2f-h); ter- 
minal segment of gonopod 2 extremely short 
(Fig. 21). Major cheliped of adult male lon- 
ger, higher than minor cheliped (Fig. 2a, b, 
Fig. la). Dactylus of major cheliped of adult 
male broad, flat; dactylus, pollex enclosing 
long narrow interspace along their cutting 


516 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Sudanonautes orthostylis from Cameroon, West Africa. Carapace dimensions (mm) and proportions 
relative to body size (CL) of the type series. (M = male, F = female, ad = adult, juv = juvenile, n = 14.) 


Sex Cw CE CH FW CW/CL CH/CL FW/CL 
Holotype, ZMB 11093 
M 25.8 Neal 8.5 7.8 1.43 0.47 0.43 
Paratypes, ZMB 11093 
M 27.8 1726 9.5 8.8 1.58 0.54 0.50 
M 21,4 18.6 8.9 7.9 1.47 0.48 0.42 
M Jee | 165 8.9 “RE 1.44 0.54 0.47 
M 22.8 [>5 8.0 Lez [47 0.52 0.46 
M Pas heir fs 4 72 6.9 1.43 0.47 0.45 
M pS it {5:7 2 q2 1.38 0.46 0.46 
M 18.2 12.7 6.1 6.3 1.43 0.48 0.50 
F (ad) 221 16.0 8.7 TO 1.38 0.54 0.44 
F (uv) 19.2 13.0 | 6.6 1.48 0.55 0.51 
F (uv) 16.7 11.6 5.6 5.6 1.44 0.48 0.48 
F (juv) 3:3 9.7 4.6 4.6 1:37 0.47 0.47 
Paratypes, SMF 2439 
M 23.0 16.5 8.5 hss) 1.39 O52 0.45 
F (ad) 28.0 18.5 10.0 9.5 LS 0.54 0.51 


edges when fingers closed (Fig. 2a). Carpus 
of cheliped with 2 large pointed teeth, sec- 
ond smaller than first (Fig. 2c). Carapace 
distinctly convex, about half carapace length 
(Table 2); carapace, anterolateral margin, 
lower margin of orbit, postfrontal crest (i.e., 
fused epigastric, postorbital crests), smooth 
(Fig. la, b). Small intermediate tooth on 
anterolateral margin between exo-orbital 
and epibranchial teeth (Fig. 1b); vertical 
suture on flank meeting anterolateral mar- 
gin at epibranchial tooth (Fig. 1b). Small 
species, mature at CW 22.0 mm. 
Description of holotype — Carapace (Fig. 
la, b).—Ovoid, widest in anterior third 
(CW/CL = 1.43), relatively high, with max- 
imum height in anterior region (CH/CL = 
0.47). Anterior margin of front straight, 
curving under, front relatively narrow, about 
one-third carapace width (FW/CW = 0.30). 
Surface of carapace smooth with no deep 
grooves. Postfrontal crest consisting of fused 
epigastric, postorbital crests, smooth, end- 
ing before meeting anterolateral margins; 
mid-groove broad, shallow. Exo-orbital 
tooth blunt, low, intermediate tooth pres- 


ent, small low, epibranchial tooth small, low. 
Anterolateral margin of carapace smooth. 
Posterolateral margin curving inward, con- 
tinuous with anterolateral margin. Posterior 
margin about two-thirds as wide as carapace 
width. 

Each flank with 2 sutures, 1 longitudinal, 
1 vertical, dividing flank into 3 parts (Fig. 
1b). Longitudinal suture dividing suborbit- 
al, subhepatic regions from pterygostomial 
region, beginning at respiratory opening and 
curving backward across flank. Short ver- 
tical suture dividing suborbital region from 
subhepatic region (Fig. 1b); suture begin- 
ning just beneath epibranchial tooth, curv- 
ing down to meet longitudinal flank groove, 
marked by row of small rounded teeth. 
Groove between sternal segments 2 and 3 
complete; groove between sternal segments 
3 and 4 consisting of 2 small notches at sides 
of sternum (Fig. 1d). Third maxillipeds fill- 
ing entire oral field, except for transversely 
oval efferent respiratory openings at supe- 
rior lateral corners. Flagellum on exopod of 
third maxilliped (Fig. 1c). Ishium of third 
maxilliped smooth, with faint vertical 


VOLUME 106, NUMBER 3 


ed 


Fig. 1. 


Sudanonautes orthostylis, adult male from Bipindihof, Cameroon (CW 25.8 mm), ZMB 11093. a, 


whole animal, dorsal aspect; b, cephalothorax, frontal aspect; c, abdomen; d, left third maxilliped and detail of 
sternum. Scale bar equals 10.00 mm (a, b, d), and 5.0 mm (c). 


groove (Fig. Ic). Mandibular palp 
2-segmented; terminal segment single, un- 
divided, small hard, flap at junction be- 
tween segments (Fig. 2d, e). First 5 segments 
of male abdomen broad, short, tapering in- 
ward; last 2 segments long, narrow, last seg- 
ment rounded at distal margin (Fig. Ic). 
Chelipeds (Figs. 1a, 2a—c). — Unequal, right 


longer (23.5 mm), higher (12.0 mm) than 
left (18.0 mm, 7.0 mm respectively). Dac- 
tylus of right cheliped broad, flattened, fin- 
gers enclosing long narrow interspace when 
closed, palm of propodus swollen. Fingers 
of right cheliped with series of small pointed 
teeth along length. Anterior dorsal margin 
of merus of right and left chelipeds with 


518 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Sudanonautes orthostylis, adult male from Bipindihof, Cameroon (CW 25.8 mm), ZMB 11093; a, 
right cheliped, frontal view; b, left cheliped, frontal view; c, detail of carpus and merus of right cheliped, dorsal 
view; d, right mandible anterior view; e, right mandible posterior view; f, left gonopod 1, caudal view; g, distal 
portion of left gonopod 1 turned to show medial margin; h, left gonopod 1, cephalic view; 1, left gonopod 2, 
caudal view. Scale bar equals 10.00 mm (a-c), and 2.0 mm (d, e, fH). 


VOLUME 106, NUMBER 3 


Carapace Dimension (mm) 


CW = - 1.87 + 1.57 CL, r = 0.98 
CH = - 0.54 + 0.54 CL, r = 0.94 


FW= 0.69 + 0.42 CW, r = 0.93 


0 
8 cee Pee he OAS} ae 20 
Carapace Length (mm) 


519 


1.6 


1.4 


1.0 


CW/CL = 1.31 + 0.009 CL, r = 0.282 
0.87 CH/CL = 0.45 + 0.004 CL, r = 0.410 
FW/CL = 0.52 - 0.004 CL, r = 0.325 


Carapace Proportion 


0.6 


0.4 
8 10 «6120—~COA CUTE C18 20 


Carapace Length (mm) 


Fig. 3. Comparisons of 14 specimens of Sudanonautes orthostylis from Bipindihof, Cameroon. a, dimensions 
of the carapace (CW, CH, FW) compared to body size (CL), r values (all at df= 13) indicating highly significant 
correlation (P < 0.001) between size classes. b, relative proportions of the carapace (CW/CL, CH/CL and FW/ 
CL) compared to body size (CL), r values (all at df= 13) indicate no significant correlation (P > 0.01) between 


size classes. 


rows of small pointed teeth, largest close to 
distal end. Carpus of cheliped with 2 large 
pointed teeth on inner margin, second 
smaller than first. Left cheliped similar to 
right, but smaller in all respects. Walking 
legs (pereiopods 2-5) slender, P4 longest, P5 
shortest. Dactyli of P2—5 tapering to point, 
each bearing rows of downward-pointing 
sharp bristles; dactylus of P5 shortest of the 
4 legs. 

Gonopods.—Terminal segment of gono- 
pod | straight for most of its length, curving 
sharply outward just before tip; lacking lon- 
gitudinal groove; lateral margins fringed by 
sparse bristles; subterminal segment gono- 
pod | slim narrowest at junction between 


segments, widest at basal end (Fig. 2f-h). 
Caudal face of subterminal segment form- 
ing raised triangular flap extending halfway 
across segment, flap tapering diagonally to 
point at junction with terminal segment, 
forming roof of chamber for gonopod 2; ce- 
phalic face of subterminal segment narrow, 
forming lower floor of chamber for gono- 
pod 2. 

Gonopod 2 (Fig. 21) shorter than gonopod 
1 (reaching only junction between last 2 seg- 
ments of gonopod 1). Terminal segment 
gonopod 2 extremely short, only ,, as long 
as subterminal segment; terminal segment 
with pointed tip. Subterminal segment gon- 
opod 2 widest at base, tapering gradually 


520 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 2.—Means (+SD) of ratio of carapace width (CW), carapace height (CH), and front width (FW), to 
body size (CL) of Sudanonautes orthostylis compared to four closely related species of Sudanonautes from 


Cameroon and Nigeria, (n = 14 in all cases). 


CW/CL 

X+SD 
Sudanonautes orthostylis 1.44 +0.1 
Sudanonautes pelii 13722208 
Sudanonautes africanus L377 = Ost 
Sudanonautes aubryi? 527i Oe 
Sudanonautes granulatus ft 40> O51 


CH/CL FW/CL 
X+SD X¥+SD 
0.50 + 0.03 O:47 3310.03 
0.49 + 0.02 0.407 + 0.01 
0.407 + 0.1 0.402 + 0.1 
0.61 20:1 0.387 + 0.1 
O22 == 081 0.392 + 0.1 


4 Proportion significantly different from that of S. orthostylis at 95% confidence limits. 


> Adult specimens only, all from Nigeria. 


inward along length, forming long, thin, 
pointed, upright process which supports 
short terminal segment. 

Adult female paratype (CW 22.0 mm, Ta- 
bles 1, 2).—Anterolateral margin behind 
epibranchial tooth bearing row of small, 
rounded low teeth. Right, left chelipeds same 
proportions as male of same size, unequal 
in both length (20.0 mm, 13.0 mm), height 
(9.0 mm, 5.0 mm). Mature female abdomen 
very wide reaching coxae of pereiopods 2- 
5. Segments of female abdomen becoming 
gradually longer distally, first, fifth becom- 
ing gradually wider, abdomen being widest 
at line separating fourth, fifth segments. 
Sixth segment, telson together forming near 
semicircle. 

Juvenile and pubertal size classes (Fig. 3a, 
b, Tables 1, 2).—Sexual maturity judged by 
development of female abdomen: abdomen 
of mature females overlapping bases of cox- 
ae of walking legs; pleopods broad, hair- 
fringed. Pubertal molt, from pubertal stage 
to sexual maturity, occurring after CW = 
22.0 mm. Dimensions of the carapace vary- 
ing with age (Fig. 3a). Relative proportions 
of carapace (width, CW/CL, height, CH/CL, 
width of frontal margin, FW/CL) of juve- 
nile, pubescent S. orthostylis not signifi- 
cantly different from adults (Fig. 3b). 

Size. — Measurements given in Table 1. 

Distribution. —Sudanonautes_ orthostylis 
is known only from the rivers and streams 
of the rain forests of south west Cameroon. 


Discussion 


A number of species of freshwater crabs 
from the forested regions of Cameroon and 
Nigeria bear a superficial resemblance to the 
holotype of S. orthostylis from Cameroon. 
Included in this group are S. pelii, S. aubryi, 
S. africanus, S. granulatus, and the three 
species of Potamonemus (Cumberlidge & 
Clark 1992, Cumberlidge 1993b). These taxa 
can be distinguished from S. orthostylis as 
follows. 

Comparison of S. orthostylis from Cam- 
eroon with the lectotype of S. pelii from 
Elmina, Ghana revealed the following dif- 
ferences: (1) the terminal segment of gon- 
opod 1 of S. orthostylis is straight, curving 
sharply at its tip, while that of S. pelii is 
curved evenly along its entire length; (2) the 
dactylus of the major cheliped of S. ortho- 
stylis is broad and flat, while that of S. pelii 
is narrow and slightly arched; (3) the post- 
frontal crest of S. orthostylis almost meets 
the anterolateral margin close to the epi- 
branchial tooth, while that of S. pe/ii meets 
this margin behind the epibranchial tooth; 
(4) the carapace and frontal margin of S. 
orthostylis are significantly wider than those 
of S. pelii (Table 2); and (5) S. orthostylis is 
a much smaller species maturing at CW 22.0 
mm, compared to maturity at CW 45.0 mm 
in the larger S. pelii. 

A smooth rounded carapace is a character 
shared by both S. orthostylis and S. aubryi. 


VOLUME 106, NUMBER 3 


The two taxa may be distinguished as fol- 
lows: (1) the terminal segment of gonopod 
1 of S. orthostylis lacks a groove, is straight, 
and curves sharply outward at the tip, while 
that of S. aubryi bears a longitudinal groove 
and curves along its entire length; and (2) 
the carapace of S. orthostylis is significantly 
flatter and less widened, and the frontal 
margin is wider than those proportions of 
S. aubryi (Table 2). 

Sudanonautes orthostylis may be distin- 
guished from S. africanus, the type species 
of the genus as follows: (1) gonopod 1 of S. 
orthostylis is straight, curving only at the 
tip, while that of S. africanus curves evenly 
along its length; (2) the dactylus of the major 
cheliped of S. orthostylis is broad and flat 
and the propodus lacks large teeth, while 
the dactylus of S. africanus is slightly arched 
and the propodus possesses a diagnostic 
broad, flat tooth; (3) the carapace of S. or- 
thostylis is smooth, while that of S. african- 
us has a warty texture in the posterior re- 
gion, and distinct granulations in the anterior 
corners; (4) the carapace ofS. orthostylis is 
significantly higher and wider, and the fron- 
tal margin is wider than in S. africanus (Ta- 
ble 2); and (5) S. africanus is a much larger 
species than S. orthostylis maturing at CW 
70.0—90.0 mm, rather than around CW 22.0 
mm for S. orthostylis. Specimens of S. or- 
thostylis of CW 25.0 mm or more show 
greatly enlarged chelipeds (male) and broad- 
ened abdomen (female), whereas the che- 
lipeds and abdomen of S. africanus of this 
size are small and undeveloped (i.e., juve- 
niles). 

Sudanonautes orthostylis may be distin- 
guished from S. granulatus (Balss, 1929) 
[(fide Cumberlidge 1933a)] as follows: (1) 
the proximal two-thirds of the terminal seg- 
ment of gonopod 1 of S. orthostylis is 
straight, lacks a groove, and curves sharply 
outward at the tip, whereas that of S. gran- 
ulatus curves along its entire length, and the 
gonopod bears a longitudinal groove; (2) the 
vertical suture on the flank of S. orthostylis 


521 


originates at the epibranchial tooth, where- 
as that of S. granulatus originates at the 
intermediate tooth; (3) the dactylus of the 
major cheliped of the adult male of S. or- 
thostylis is broad and flat, but not arched, 
whereas that of S. granulatus is narrow and 
dramatically arched; (4) the major cheliped 
of adult S. granulatus is longer than the car- 
apace width, whereas that of S. orthostylis 
is shorter than the carapace width; and (5) 
the carapace and the frontal margin of S. 
orthostylis are significantly wider than in S. 
granulatus (Table 2). 

The short terminal segment of gonopod 
2 of the species of Potamonemus resembles 
that of S. orthostylis but the lack of a fla- 
gellum on the exopod of the third maxilli- 
ped of Potamonemus and the absence of a 
well-defined intermediate tooth on the an- 
terolateral margin of the carapace clearly 
separate the 2 genera (Cumberlidge & Clark 
1992, Cumberlidge 1993b). 


Acknowledgments 


I am very grateful to Dr. H. E. Gruner of 
the Zoologische Museum of the Humboldt- 
Universitat, Berlin for his helpful cooper- 
ation during a visit, and for loaning the ho- 
lotype of S. orthostylis. I also thank the staff 
at the Natur-Museum und Forschungs-In- 
stitut Senckenberg, Frankfurt am Main, 
Germany for their help during a visit. Pro- 
fessor Dr. L. B. Holthuis of the Nationaal 
Naturhistorisch Museum, Leiden, The 
Netherlands is thanked for the loan of the 
type specimen of S. pelii. Thanks also to 
Associate Editor Dr. Rafael Lemaitre and 
two reviewers for their helpful comments 
on the manuscript. I am especially grateful 
to artist Anne C. Martin of Northern Mich- 
igan University, U.S.A, for all of the illus- 
trations used in this paper. Part of this work 
was supported by a Faculty Grant from 
Northern Michigan University, Marquette, 
Michigan, U.S.A. 


522 


Literature Cited 


Balss, H. 1929. Potamonidae au Cameroon. Jn: Con- 
tribution a l’étude de la faune du Cameroun. — 
Faune Colonies Frangaises 3:1 15-129. 

Bott, R. 1955. Die Siisswasserkrabben von Afrika 

(Crust., Decap.) und ihre Stammesgeschichte. — 

Annales du Musee Royal du Congo Belge, (Ter- 

vuren, Belgique,) C. Zoologie 1 (3, 3):209-352. 

1959. Potamoniden aus West-Afrika.— Bul- 
letin de l’Institut Fondamental D’Afrique Noire, 
Série A 21 (3):994-1008. 

1964. Decapoden aus Angola unter beson- 
derer Beriicksichtigung der Potamoniden (Crust. 
Decap.) und einem Anhang: Die Typen von 
Thelphusa pelii Herklots 1861.—Publicacoes 
Culturais da Companhia de Diamantes de An- 
gola, Lisboa, 69:23-24. 

1970. Betrachtungen liber die Entwicklungs- 
geschichte und Verbreitung der Sitsswasser- 
Krabben nach der Sammlung des Naturhisto- 
rischen Museums in Genf/Schweiz.— Revue 
Suisse de Zoologie 77(2), 24:327-344. 
Cumberlidge, N. 1989. Redescription of Sudanon- 
autes orthostylis (Bott, 1955), a freshwater crab 
from Nigeria, Cameroon and Ghana (Decapo- 
da, Potamonautidae), with notes on its ecolo- 
gy.—Crustaceana 56(3):230-245. 

. 1991. Sudanonautes kagoroensis, a new spe- 

cies of freshwater crab (Decapoda: Potamoidea: 

Potamonautidae) from Nigeria.— Canadian 

Journal of Zoology 69:1938-1944. 

. 1993a. Redescription of Sudanonautes gran- 

ulatus (Balss, 1929) (Potamoidea, Potamonau- 

tidae) from West Africa.—Journal of Crusta- 
cean Biology (in press). 

1993b. Two new species of Potamonemus 
Cumberlidge & Clark, 1992 (Brachyura, Pota- 
moidea, Potamonautidae) from the rain forests 
of West Africa.— Journal of Crustacean Biology 
13:571-584. 

——, & P. Clark. 1992. A new genus and species 
of freshwater crab from Cameroon, West Africa 
(Crustacea, Brachyura, Potamoidea, Potamon- 
autidae).—Bulletin of the British Museum of 
Natural History (Zoology), London 58(2):149- 
156. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Herklots, J. A. 1861. Symbolae carcinologicae. Etudes 
sur la classe des Crustacés Pp. 1—43, Leiden. 
Macleay, W. S. 1838. Illustrations of the zoology of 
South Africa; being a portion of the objects of 
natural history chiefly collected during an ex- 
pedition into the interior of South Africa, under 
the direction of Dr. Andrew Smith, in the years 
1834, 1835, and 1836; fitted out by “The Cape 
of Good Hope Association for Exploring Cen- 
tral Africa.” Jn A. Smith, Illustrations of the 
zoology of South Africa; consisting chiefly of 
figures and descriptions of the objects of natural 
history collected during an expedition into the 
interior of South Africa, in the years 1834, 1835, 
and 1836; fitted out by “The Cape of Good 
Hope Association for Exploring Central Afri- 
ca.’’, (Invertebrates). London, 75 pp, 4 pls. 
Milne-Edwards, H. 1853. Observations sur les affin- 
itiés zoologiques et la classification naturelle des 
Crustacés.—Annales des Sciences Naturelles, 
Zoologie, Paris, Série 3, 10:163-228. 
Milne-Edwards, A. 1869. Révision du genre Thel- 
phusa et description de quelques especes nou- 
velles faisant partie de la collection du Mu- 
séum.—Nouvelles Archives du Muséum 
d’Histoire naturelle, Paris, 5:161-191. 

1886. La description de quelques Crustacés 
du genre Thelphusa recueillis par M. de Brazza 
dans les régions du Congo. — Bulletin de la So- 
ciété Philomathique de Paris, Série 7, 10:148- 
leaks 
Monod, T. 1977. Sur quelques crustacés Décapodes 
africaines (Sahel, Soudan).—Bulletin de Mu- 
séum National d’ Histoire naturelle, Série 3, 500: 
1201-1236. 

1980. Décapodes. Pp. 369-389 in J.-R. Du- 
rand and C. Léveque, eds., Flore et Faune Aqua- 
tiques de l’Afrique Sahelo-Soudanienne. Edi- 
tions de l’Office de la Recherche Scientifique et 
Technique Outre-Mer Collection Initiations— 
Documentations Techniques No. 44(1), Paris, 
389 pp. 


Department of Biology, Northern Mich- 
igan University, Marquette, Michigan 
49855, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(3), 1993, pp. 523-531 


THREE GENERA REMOVED FROM THE 
SYNONYMY OF PINNOTHERES BOSC, 1802 
(BRACHYURA: PINNOTHERIDAE) 


Raymond B. Manning 


Abstract. — Three genera, Arcotheres Burger, 1895, Holothuriophilus Nauck, 
1880, and Zaops Rathbun, 1900, are removed from the synonymy of Pin- 
notheres Bosc, 1802. Arcotheres contains only its type species from the Indo- 
West Pacific, A. palaensis (Burger, 1895). Holothuriophilus is represented by 
two species from the eastern Pacific, its type species H. trapeziformis Nauck, 
1880 and H. pacificus (Poeppig, 1836), a senior synonym of H. silvestrii (Nobili, 
1901), aspecies previously assigned to Pinnaxodes Heller, 1865. Zaops contains 
a western Atlantic species, Z. ostreum (Say, 1817), the American oyster pea 
crab; Z. ostreum is an adult and a senior synonym of the type species of Zaops, 


Pinnotheres depressum Say, 1817. 


Ongoing studies on some Atlantic species 
of Pinnotheres prompted me to review the 
status of its synonyms. As a result of this 
review, I believe that three genera synony- 
mized with Pinnotheres Bosc, 1802 by 
Rathbun (1918), Schmitt et al. (1973), and 
Manning & Holthuis (1981), e.g., Arcotheres 
Burger, 1895, Holothuriophilus Nauck, 
1880, and Zaops Rathbun, 1990, should be 
recognized as separate genera. 

Five other pinnotherid genera have been 
recognized since the summary of pinnothe- 
rid taxa was published by Schmitt et al. 
(1973), as follows: 

Indopinnixa Manning & Morton, 1987 
[type species Indopinnixa sipunculana 
Manning & Morton, 1987] is a member of 
the Pinnothereliinae and is not considered 
further here; all of the other newly recog- 
nized genera listed here are members of the 
subfamily Pinnotherinae. 

Calyptraeotheres Campos, 1990 [type 
species Fabia granti Glassell, 1933] differs 
from Pinnotheres in having a two-segment- 
ed mandibular palp. 

Clypeasterophilus Campos & Griffith, 
1990 [type species Dissodactylus rugatus 


Bouvier, 1917] resembles Dissodactylus 
Smith, 1870 and differs from Pinnotheres 
in having the dactyli of the walking legs bifid 
as well as in having the three segments of 
the mandibular palp placed end-to-end. 

Limotheres Holthuis, 1975 [type species 
Limotheres nasutus Holthuis, 1975] also dif- 
fers from Pinnotheres in having the seg- 
ments of the third maxilliped placed end- 
to-end; it further differs in having a 
projecting, triangular rostrum and three 
longitudinal postfrontal grooves on the car- 
apace. 

Tumidotheres Campos, 1989a [type spe- 
cies Pinnotheres margarita Smith, 1869; the 
genus also includes 7. maculatus (Say, 
1818)] differs from Pinnotheres in having 
(a) the dactyli of the walking legs dissimilar 
and unequal, that of the fifth leg much the 
longest, and (b) a spatulate dactylus on the 
palp of the third maxilliped, inserted near 
midlength of the propodus and not extend- 
ing beyond the propodus. As pointed out 
by Campos (1989a:693) the dactylus of the 
third maxilliped in the type species of Pin- 
notheres, the European P. pisum (Linnaeus, 
1767), is styliform and inserted basally on 


524 


the ventral margin of the propodus; the dac- 
tyli of the walking legs in P. pisum are sim- 
ilar and subequal. 

One genus previously considered to be a 
pinnotherid has been removed from the 
family. Mortensenella Rathbun, 1909 [type 
species Mortensenella forceps Rathbun, 
1909] was transferred from the Pinnother- 
idae to the subfamily Camptandriinae of the 
Ocypodidae by Harminto & Ng (1991). 

The genus Orthotheres Sakai, 1969 [type 
species Orthotheres turboe Sakai, 1969], in 
which the segments of the palp of the third 
maxilliped are placed end-to-end, was re- 
viewed by Campos (1989b); it contains some 
species previously assigned to Fabia and 
Pinnotheres. 

Judging from the accounts in Davidson 
(1968) of Cryptophrys concharum Rathbun, 
1893, the type species of Cryptophrys Rath- 
bun, 1893, and of Fabia subquadrata Dana, 
1851, the type species of Fabia Dana, 1851, 
I believe that there may be grounds to con- 
sider these two genera as distinct, an action 
that is beyond the scope of this paper. Cryp- 
tophrys and Fabia are now regarded as syn- 
onyms (Schmitt et al. 1973:22). 

The diagnoses given below will distin- 
guish each genus from Pinnotheres s.s. In 
the diagnoses, pereopods are indicated by 
their abbreviations, e.g., Pl to P5 (P1 is the 
cheliped, P5 the last leg or the fourth walk- 
ing leg). In the legends, measurements are 
given as carapace length < carapace width, 
in millimeters. USNM is an acronym for 
the National Museum of Natural History, 
Smithsonian Institution, Washington. 


Arcotheres Burger, 1895 
Fig. 1 


Arcotheres Burger, 1895:361. 


Type species. —Pinnotheres palaensis 
Burger, 1895, by subsequent designation by 
Rathbun (1918:62). Gender masculine. 

Diagnosis. —Carapace subhexagonal. 
Third maxilliped with ischium and merus 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


indistinguishably fused; exopod with fla- 
gellum; palp 3-segmented; propodus much 
longer than carpus; spatulate dactylus in- 
serted proximally on ventral margin of 
propodus, not extending to apex of propo- 
dus. Dactyli of walking legs dissimilar, those 
of the third (P4) and fourth (P5) walking 
legs longer than and different from dactyli 
of first (P2) and second (P3) walking legs in 
both sexes. Abdomen of 7 segments in each 
SEX. 

Remarks.—Arcotheres differs from Pin- 
notheres in having the dactyli of the last two 
walking legs longer than and different from 
the dactyli of the first two, and in having a 
spatulate rather than styliform dactylus on 
the third maxilliped. 

Known only from the type species from 
the Indo-West Pacific. In pelecypods. 


Holothuriophilus Nauck, 1880 
Figs. 2-3 


Holothuriophilus Nauck, 1880:66. Name no. 
319 on Official List of Generic Names in 


Zoology. 


Type species. —Holothuriophilus trapezi- 
formis Nauck, 1880, by original designation 
and monotypy. Gender masculine. 

Diagnosis. —Carapace broader than long, 
subrectangular. Third maxilliped with is- 
chium and merus indistinguishably fused; 
exopod with flagellum; palp 3-segmented; 
propodus shorter than carpus, conical; spat- 
ulate dactylus articulated basally on prop- 
odus, extending beyond end of propodus. 
Dactyli of walking legs similar and sub- 
equal, short. Abdomen of 7 segments in both 
SEXeS. 

Remarks.—I consider Holothuriophilus 
to be a valid genus distinct from Pinnax- 
odes. It differs (a) in carapace shape, with 
the greatest width of the carapace anterior 
to the midlength in Holothuriophilus, pos- 
terior to the midlength in Pinnaxodes; (b) 
in the proportions of the walking legs which 
are short and stout, with very short dactyli 


VOLUME 106, NUMBER 3 


Eyres 


525 


/ 


yy, 


Fig. 1. Arcotheres palaensis Birger, spent female, 7.8 x 11.1 mm, Philippines, USNM 256948. a, Dorsal 
view (walking legs of left side omitted; extent of abdomen indicated on both sides); b, Third maxilliped. 


in Holothuriophilus, slender and elongate in Pinnaxodes whereas these segments are 
with long dactyli in Pinnaxodes; and (c) in __indistinguishably fused in Holothuriophilus. 
the structure of the third maxilliped, which Members of Pinnaxodes inhabit echinoids, 
has a suture between the ischium and merus' whereas species of Holothuriophilus inhabit 


526 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Holothuriophilus pacificus (Poeppig), spent female, 11.5 x 14.2 mm, San Juan Bay, Peru, USNM 
256986. a, Dorsal view (walking legs of left side omitted); b, Third maxilliped, inner aspect; c, Same, outer 
aspect. 


VOLUME 106, NUMBER 3 


527 


Fig. 3. Species of Holothuriophilus. a, H. pacificus (Poeppig), San Vicente, Chile [from Garth (1957:fig. 10A), 
as Pinnaxodes silvestrii (Nobili]; b, H. pacificus (Poeppig), Talcahuano, Chile [from Poeppig (1836:pl. 4, fig. IID), 
as Leucosia pacifica]; c, Holothuriophilus trapeziformis Nauck, Mazatlan, Mexico [from Birger (1895:pl. 9, fig. 


26)]. 


holothurians (see summary of Chilean pin- 
notherid hosts in Garth 1957:92). 

The status of the other species listed in 
Pinnaxodes by Schmitt et al. (1973) should 
be re-examined, a task beyond the scope of 
this work. 

In my opinion, Holothuriophilus trape- 
ziformis Nauck, 1880 is congeneric with 
Pinnaxodes silvestrii (Nobili, 1901) and its 
junior synonym Pinnaxodes meinerti Rath- 
bun, 1904; Garth (1957:88) synonymized 


the latter two species. Nauck’s species may 
be the northern counterpart of H. silvestrii. 

I believe that Leucosia pacifica Poeppig, 
1836, from Talcahuano, Chile, which was 
considered by Rathbun (1937:183, foot- 
note) to be a pinnotherid and was listed as 
incertae sedis by Garth (1957:91) and 
Schmitt et al. (1973:137), is a species of 
Holothuriophilus. Poeppig’s figure of his 
species shows a crab with the carapace shape 
of Holothuriophilus, broader than long, with 


528 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Zaops ostreum (Say), adult female [from Williams (1984:fig. 354), as Pinnotheres ostreum Say]. 


its greatest width anteriorly, and the short 
dactyli characteristic of members of that ge- 
nus. The third maxilliped shown by Poeppig 
is extremely stylized and does not resemble 
that of members of either Holothuriophilus 
or Pinnaxodes. | consider Leucosia pacifica 
Poeppig, 1836 to be a senior synonym of 
Pinnaxodes silvestrii (Nobili, 1901). That 
species should be known as Holothuriophi- 
lus pacificus (Poeppig, 1836). 

The figures of H. pacificus given by Poep- 
pig (as Leucosia pacifica) and Garth (as Pin- 
naxodes silvestrii) are reproduced here in 
Fig. 3, along with the figure of H. trapezi- 
formis published by Birger. 

Known from the eastern Pacific. In hol- 
othurians. 


Zaops Rathbun, 1900 
Figs. 4—5 


Zaops Rathbun, 1900:588, 590. 


Type species. —Pinnotheres depressum 
Say, 1817, a subjective junior synonym and 
juvenile of Pinnotheres ostreum Say, 1817 


(see Williams 1984:445), by original des- 
ignation and monotypy. Gender masculine. 

Diagnosis. —Carapace subhexagonal. 
Third maxilliped with ischium and merus 
indistinguishably fused; exopod with fla- 
gellum; propodus much longer than carpus; 
dactylus minute, inserted near midlength of 
ventral margin of propodus. Dactyli of 
walking legs dissimilar and unequal, that of 
second walking leg (P3) much the longest in 
adult females; propodus of first walking leg 
(P2) dilated distally in females. Abdomen 
of 7 segments in each sex. 

Remarks. —The walking legs of the type 
species of Zaops are quite distinctive, with 
the club-shaped propodus on the first walk- 
ing leg (P2) and the long dactylus on the 
second walking leg (P3). Zaops ostreum 
shares the distally dilated propodus of the 
first walking leg with the eastern Pacific Pin- 
notheres clavapedatus Glassell, 1935 [? = 
Pinnotheres lithodomi Smith, 1870], but 
Glassell’s species differs in having expanded 
propodi on the first two walking legs (P2, 
P3) and a long dactylus on the third walking 
leg (P4) rather than on the second (P3). 


VOLUME 106, NUMBER 3 


Fig. 5. 


529 


Zaops ostreum (Say), spent female, 10.5 x 12.1 mm, market in New York, USNM 4991. a, Third 


maxilliped; b, Cheliped; c, Second pereopod; d, Third pereopod; e, Fourth pereopod; f, Fifth pereopod. 


Zaops 18 monotypic. Zaops ostreum 1s 
known from localities between Massachu- 
setts and Brazil in the western Atlantic (Wil- 
liams 1984). In bivalve mollusks, especially 
the oyster, Crassostrea virginica (Gmelin); 
possibly in worm tubes (Williams 1984). 


Acknowledgments 


Studies on systematics of pinnotherids are 
supported by the Smithsonian Marine Sta- 
tion at Link Port, Florida. This is contri- 
bution number 313 from that station. The 
figures were prepared by Lilly King Man- 
ning. 


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Schmitt, W. L., J. C. McCain, & E.S. Davidson. 1973. 
Fam. Pinnotheridae. Brachyura I: Decapoda I. 
In H.-E. Gruner & L. B. Holthuis, eds., Crus- 
taceorum Catalogus. W. Junk, Den Haag 3:1- 
160. 

Smith, S. I. 1869. Pinnotheres margarita Smith, sp. 

nov. P. 245 (footnote) in A. E. Verrill, On the 

parasitic habits of Crustacea.—American Nat- 

uralist 3:239-250. 

1870. Ocypodoidea. Notes on North Amer- 


VOLUME 106, NUMBER 3 


ican Crustacea, I.—Transactions of the Con- 
necticut Academy of Arts and Sciences 2:1 13- 
176, pls. 2-5. 

Williams, A. B. 1984. Shrimps, lobsters and crabs of 
the Atlantic coast of the eastern United States, 
Maine to Florida. Smithsonian Institution Press, 
Washington, 550 pp. 


531 


Department of Invertebrate Zoology, Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. 20560, 
U:S.A. 


PROC. BIOL. SOC. WASH. 
106(3), 1993, pp. 532-544 


A NEW SUBGENUS AND SPECIES OF CRAYFISH 
(DECAPODA: CAMBARIDAE) OF THE GENUS 
CAMBARUS, WITH AN AMENDED 
DESCRIPTION OF THE SUBGENUS 
LACUNICAMBARUS 


Raymond F. Jezerinac 


Abstract. —A new subgenus of crayfish, Tubericambarus, and a new species, 
Cambarus (Tubericambarus) thomai, are described. The new species is most 
closely related to C. (T.) acanthura, new combination, but differs from it in 
that the distomedian spine on the mesial ramus of the uropod does not over- 
reach the rounded margin of the ramus, and the merus always has a well 
developed spiniform tubercle on the distroventral articular rim. The species 
occurs in Kentucky, Ohio, Pennsylvania, Tennessee, and West Virginia. The 
definition of the subgenus Lacunicambarus is amended and distributional maps 


of the subgenera are included. 


As early as 1885, Faxon (p. 72) noted that 
what is currently called Cambarus (Lacuni- 
cambarus) diogenes is a species complex 
consisting of at least three forms: an eastern 
form found on the Atlantic Costal Plain, a 
western form extending from the western 
side of the Appalachian Mountains west- 
ward to the eastern slopes of the Rocky 
Mountains and from the Gulf Coast north- 
ward to southern Canada, and a variety des- 
ignated by him as Cambarus Diogenes var. 
Ludovicianus, from the environs of New 
Orleans, Louisiana. Faxon’s variety was lat- 
er elevated to subspecific rank by Hay (1899: 
959). Marlow (1960:248) attempted to clar- 
ify the taxonomy of this group, but his ma- 
jor contribution was providing further ev- 
idence for recognizing C. d. ludovicianus as 
a valid subspecies. In 1969, Hobbs revised 
the genus Cambarus by dividing it into 10 
subgenera, provided diagnoses for the sub- 
genera, and listed the species belonging to 
each of them. Cambarus diogenes and re- 
lated forms were placed in the subgenus La- 
cunicambarus. Two additional taxa of the 
complex were described, C. (L.) miltus Fitz- 


patrick, 1978, and C. (L.) acanthura Hobbs, 
1981. In 1989, Hobbs (p. 26) raised C. (L.) 
d. ludovicianus to specific rank and repeated 
the statement he made in 1974 (p. 20) that 
“This [the C. (L.) diogenes group] is a spe- 
cies complex that needs considerable atten- 
tone 

After studying the complex for the last 12 
years, I have concluded that the complex 
consists of two subgenera Lacunicambarus 
and Tubericambarus, new subgenus, and at 
least five additional species or subspecies. 
The subgenus Lacunicambarus is amended, 
the new subgenus is defined, and one new 
species 1s described herein. 


Lacunicambarus Hobbs, 1969, amended 


Diagnosis. —Eyes reduced and pigment- 
ed. Antennae not heavily fringed on mesial 
border. Rostrum with margins moderately 
thickened, usually without spines or tuber- 
cles. Postorbital and cervical spines absent. 
Suborbital angle prominent and often acute 
to subacute. Branchiostegal spine reduced 
to small tubercle or absent. Areola oblit- 


VOLUME 106, NUMBER 3 


A B 
Fig. 1. 


533 


a 


‘ 


G D 


Cambarus (Lacunicambarus) diogenes: A, dorsal view right chela; B, ventral view right chela, (SPT 


= 3 subpalmar tubercles). Cambarus (Tubericambarus) acanthura: C, dorsal view right chela; D, ventral view 


right chela. 


erated or linear along much of its length, 
constituting 37-45% (X = 42%) of total 
length of carapace, never bearing more than 
2 punctations in narrowest part. Chela (Fig. 
1A) moderately robust with dactyl length/ 
palm length ratio greater than 1.9 on first 
form males; dorsomesial surface of palm of 
chela with 2 well developed rows of tuber- 
cles usually numbering 6-8 each, third row 
running to knob at base of the dactyl, and 
additional scattered tubercles present be- 
tween second and third rows; dorsomesial 
surface mostly punctate laterally; lateral 
margin of fixed finger of chela subcostate, 
with punctations but never bearing row of 
spines; fingers gaping and with moderately 
well defined dorsomedian longitudinal ridg- 
es; proximal opposable margin of dacty] dis- 
tinctly concave; inconspicuous tuft of setae 
sometimes present at mesial base of fixed 
fingers, dorsolateral base slightly impressed; 
subpalmar tubercles (Fig. 1B) 1-3. Medial 
spine on mesial ramus of uropod never 
overreaching caudal margin of ramus. Form 
I male with coxa of fourth pereiopod lacking 
large ventral setiferous pit on caudomesial 


boss; first pleopods contiguous at base, dis- 
tal portion of shaft straight; terminal ele- 
ments consisting of (1) short, broad, blade- 
like, distally truncate or rounded central 
projection (rarely with subapical notch) re- 
curved at about 90° to shaft, (2) swollen 
mesial process variously shaped and di- 
rected, and frequently bearing 1-4 small 
tuberculiform prominences apically, and (3) 
often rudimentary caudal knob at caudo- 
lateral base of central projection. 

Females with annulus ventralis subsym- 
metrical, slightly movable; first pleopod 
present and reaching cephalically beyond 
caudal margin of annulus. 

Type species.—Cambarus (Lacunicam- 
barus) diogenes Girard, 1852:88. 

Species. —Cambarus (Lacunicambarus) 
diogenes Girard, 1852:88, C. (Lacunicam- 
barus) ludovicianus Faxon, 1884:144, and 
C. (Lacunicambarus) miltus Fitzpatrick, 
1978:749. 

Range. — Disjunct (Fig. 2). Along the At- 
lantic Costal Plain from Maryland to East- 
ern Georgia; from Western Georgia to East- 
ern Texas, northward from Louisiana to 


sere 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Geographic range of the subgenera Lacunicambarus: A, range of C. (L.) diogenes; B, range of C. (L.) 


miltus; C, range of C. (L.) ludovicianus. 


Canada, as far west as Denver, Colorado, 
and as far east as western New York via the 
north shore of Lake Erie. 


Tubericambarus, new subgenus 


Diagnosis. —Eyes reduced and pigment- 
ed. Antennae not heavily fringed on mesial 
border. Rostrum with margins moderately 
thickened without spines or tubercles 
(sometimes present on small juveniles). 
Postorbital and cervical spines absent. Sub- 
orbital angle prominent and often acute to 
subacute. Branchiostegal spine reduced to 
small tubercle or usually absent. Areola usu- 
ally obliterated or linear, and constituting 
40-47% (X = 43%) of total length of cara- 
pace, never bearing more than | punctation 
in narrowest part. Chela (Fig. 1C) moder- 
ately robust with dactyl length/palm length 
ratio less than 1.8 on first form males; me- 
sial 3 to %4 of dorsal palmar surface of chela 
studded with small tubercles; dorsomesial 
surface tuberculate, punctate laterally; lat- 


eral margin of fixed finger of chela smooth 
or costate with punctations but never bear- 
ing row of spines; fingers slightly gaping with 
well defined dorsomedian longitudinal ridg- 
es; proximal opposable margin of dactyl 
concave; tufts of setae at mesial base of fixed 
fingers usually absent, dorsolateral base with 
moderate impression; subpalmar tubercle 
(Fig. 1D) usually absent, occasionally 1 
present. Median spine on mesial ramus of 
uropod reaching or overreaching caudal 
margin of ramus. Form I male with coxa of 
fourth pereiopod lacking large ventral se- 
tiferous pit on caudomesial boss; first ple- 
opods contiguous at base, distal portion of 
shaft straight; terminal elements consisting 
of (1) short, broad, blade-like, distally 
rounded central projection (rarely with sub- 
apical notch) recurved at about 90° to shaft, 
(2) swollen mesial process variously shaped 
and directed, bearing 1 small tuberculiform 
prominence apically, and (3) lacking caudal 
knob at caudolateral base of central projec- 
tion. 


VOLUME 106, NUMBER 3 


535 


Fig. 3 Geographic range of the subgenus Tubericambarus: A, range of C. (T.) acanthura, B, range of C. (T.) 


thomai; C, range of C. (T.) sp. A. 


Females with annulus ventralis subsym- 
metrical, slightly movable; first pleopod 
present and reaching cephalically beyond 
caudal margin of annulus. 

Type species. —Cambarus (Tubericam- 
barus) acanthura Hobbs, 1981:215, new 
combination. 

Species. —Cambarus (Tubericambarus) 
acanthura Hobbs, 1981:215, and C. (Tube- 
ricambarus) thomai, new species. C. (Tube- 
ricambarus) sp. A. 

Gender. — Masculine. 

Etymology. —Tuber- (L. tuberosus = full 
of lumps or protuberances) combined with 
Cambarus, in reference to the tubercles cov- 
ering a significant portion of the palm of the 
chela. 

Range. —The Gulf Costal Plain of Florida 
(Fig. 3), Georgia, and Mississippi north- 
ward up the Appalachian Plateau to Lake 
Erie, Southern Michigan, and west to Cen- 
tral Illinois and Eastern Missouri. Appar- 
ently absent in the Blue Grass Region of 
Kentucky. 


Hobbs’ (1972:108) taxonomic key should 
be modified as follows: 


12(10) 


13(12) 


Dactyl of chela lacking broad 
concavity on basal '2 of oppos- 
able margin (fig. 90e); first pleo- 
pod with central projection 
distinctly longer than cephalo- 
caudal diameter of shaft at base 
of projection (fig. 92b-d) .... 


.. Depressicambarus Hobbs, 1969: 


Dactyl of chela with broad con- 
cavity on basal '2 of opposable 
margin (fig. 90d); first pleopod 
with central projection equal in 
length to, or shorter than, ceph- 
alocaudal diameter of shaft at 
base of projection (fig. 92a) .. 
Mesial and dorsomesial surface 
of palm with 2 distinct rows of 
tubercles, third row extending 
to knob at base of dactyl, ad- 
ditional tubercles between sec- 


112 


13 


536 


ond and third rows; dactyl 
length greater than 1.9 times 
palm length; 1-3 subpalmar tu- 
bercles usually present ...... 
... Lacunicambarus Hobbs, 1969: 
127 
Mesial and dorsomesial %4 to 4 
surface of palm studded with 
small tubercles; dactyl length 
less than 1.8 times palm length; 
subpalmar tubercles 1 or usu- 
ally _absent- So f344 el ey 
. Tubericambarus, new subgenus 


Cambarus (Tubericambarus) thomai, 
new species 
Fig. 4, Table 1 


Cambarus diogenes Girard, 1852:88 [in 
part].— Williamson, 1899:48. [in part].— 
Ortmann, 1905a:398 [in part], 1905b:123 
[in part].—Newcombe, 1929:286.— 
Rhoades, 1944a:146 [in part], 1944b:98 
[in part].— Marlow, 1960:231 [in part]. 

Cambarus diogenes diogenes. —Hay, 1899: 
959 [in part].— Marlow, 1960:233. 

Cambarus (Bartonius) diogenes. —Ort- 
mann, 1906:402 [western part]. — Turner, 
1926:168 [in part]. 

Cambarus (Lacunicambarus) diogenes di- 
ogenes.—Hobbs, 1969:110 [in part].— 
Bouchard, 1972:56 [in part], 1975:595 [in 
part].— Hobbs, 1974:20; [in part].— Law- 
ton, 1979:47.—Thoma & Jezerinac, 1982: 
136.—Jezerinac & Thoma, 1984:123 
[eastern form].—Jezerinac 1985:7 [east- 
ern form]. 

Cambarus (Lacunicambarus) diogenes.— 
Jezerinac, 1985:7 [eastern].—Jezerinac 
and Stocker, 1989:2; 1990:8.—Hobbs, 
1989:24 [in part]. 

Cambarus (Lacunicambarus) sp. A.—Jez- 
erinac, 1986:178 (eastern Ohio). 


Diagnosis. — Pigmented; eyes slightly re- 
duced. Rostrum usually straight or some- 
times gently decurved in lateral view, mar- 
gins converging, slightly thickened, without 
marginal spines or tubercles, lacking me- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


dian carina, shallowly excavated. Carapace 
laterally compressed, without cervical spines 
or tubercles. Branchiostegal tubercles very 
small or absent. Suborbital angle acute. 
Postorbital ridges weak, never ending in 
spines or tubercles. Areola usually obliter- 
ated, constituting, in adults, 39.8-42.5% (X 
= 42.3%) of entire length of carapace, and, 
if open, with room for only 1 row of punc- 
tations in narrowest part. Antennal scale 2.5— 
2.8 times as long as wide, broadest at about 
midlength. Mesial 4 surface of palm of che- 
la (Fig. 1C) with distinct to adpressed tu- 
bercles, mesial row consisting of 6-8. No 
tufts of elongate setae at base of propodus. 
Opposable margin of dactyl weakly incised. 
Ratio of palm length to dactyl length av- 
eraging 1.6. Dorsomedian longitudinal ridg- 
es strong. Dorsolateral impression at base 
of propodus moderate to strong. Ventral 
surface of chela with 1, or usually without, 
subpalmar tubercle (Fig. 1D). Ventral sur- 
face of carpus with spiniform tubercle on 
distal articular rim. Mesial ramus of uropod 
with distomedian spine reaching caudal 
margin, but never extending beyond. First 
pleopods of form I male contiguous at base, 
with convexity near midlength of cephalic 
surface; terminal elements consisting of (1) 
short, non-tapering, distally truncate central 
projection, and (2) conically shaped mesial 
process, both directed caudally at angle 
slightly greater than 90°. Hooks on ischium 
of third pereiopods only. Female with an- 
nulus ventralis elliptical, slightly longer than 
broad, and rather deeply embedded in ster- 
num. 

Holotype male, Form I.— Body subovate 
(Fig. 4A, J), laterally compressed. Abdomen 
narrower than cephalothorax (12.5 and 17.7 
mm); maximum width of carapace greater 
than depth at caudodorsal margin of cer- 
vical groove (17.7 and 16.8 mm). Areola 
closed with no punctations in narrowest part; 
length comprising 43.1% of total length of 
carapace. Rostrum with convergent, slightly 
thickened, margins; acumen not distinctly 
delimited basally, anterior tip upturned and 


VOLUME 106, NUMBER 3 Si 


“ 


Kd “h \ 
bra 


HOT 
Rs | z 


ye 


Fig. 4. Cambarus (Tubericambarus) thomai, new species. All from holotype male, Form I, except C, E, from 
morphotype male, form II and I, from allotype female: A, lateral view of carapace; B, C, mesial view of first 
pleopods; D. caudal view of first pair of pleopods; E, F, lateral view of first pleopod; G, antennal scale; H, 
epistome; I, annulus ventralis; J, dorsal view of carapace; K, proximal podomeres of third, fourth, and fifth 
pereiopods; L, dorsal view of distal podomeres of cheliped. (See Table 1 for precise measurements.) 


538 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Measurements (mm) of Cambarus (Tube- 
ricambarus) thomai, new species. 


Morpho- 
Character Holotype Allotype type 
Carapace 
Height 16.0 17.0 14.6 
Width WT 17.3 13.9 
Length 39.4 40.0 322 
Areola 
Length 16.8 16.9 13.4 
Rostrum 
Width at eyes 4.2 4.2 3.8 
Length 7.6 8.4 6.4 
Length to anterior 
postorbital ridges 6.3 6.4 5.2 
Postorbital ridge 
Width 7.8 ZZ 5.9 
Chela—right 
Length of lateral 
margin of palm 31.1 28.6 Dope 
Length of mesial 
margin of palm 11.6 10.6 3 
Width of palm 14.5 13.2 10.3 
Length of dactyl 18.2 16.9 13.7 
Thickness of palm 92 8.9 6:5 
Abdomen 
Length 40.0 44.3 Se, 
Width 25 18.1 a7 
Gonopod 
Length 9.6 7.8 
Antennal scale 
Length D3 5.4 5.0 
Width Da 1:9 1.8 


reaching base of ultimate podomere of an- 
tennular peduncle; upper surface of rostrum 
concave with no punctations other than 
usual submarginal ones. Subrostral ridge 
weak but evident in dorsal aspect along bas- 
al *3 of rostrum. Postorbital ridge weak, 
grooved dorsolaterally, and ending cephal- 
ically without spine or corneous tubercle. 
Suborbital angle very prominent; bran- 
chiostegal spine represented by small tu- 
bercle. Cervical spine absent. Hepatic and 
branchiostegal regions with granules. Re- 
mainder of carapace punctate dorsally and 


granulate laterally. Abdomen subequal in 
length to carapace, pleura short, subtrun- 
cate, rounded caudoventrally. Cephalic sec- 
tion of telson with 2 spines on left (3 right) 
caudolateral corner. Proximal podomere of 
uropod with weak distal spine on mesial 
lobe; mesial ramus of uropod with promi- 
nent median rib ending distally in strong 
distomedian spine not overreaching margin 
of ramus, laterodistal spine of ramus also 
strong. 

Cephalomedian lobe of epistome (Fig. 4H) 
short and subtriangular with uniform mar- 
gins, ventral surface rather flat; main body 
with shallow fovea; epistomal zygoma 
arched. Ventral surface of proximal podo- 
mere of antennular peduncle with small 
acute spine at base of distal third. Antennal 
peduncle without spines; antennal scale (Fig. 
4G) 2.5 times as long as broad, broadest 
slightly proximal to midlength, mesial bor- 
der forming gentle arc; distal spine strong, 
reaching distal extremity of antennular pe- 
duncle. Mesial half of ventral surface of is- 
chium of third maxilliped studded with ir- 
regular rows of long, stiff setae; submarginal 
lateral row on podomere consisting of much 
smaller flexible ones; distolateral angle not 
acute. 

Length of right chela (Fig. 4L) 78.9% that 
of carapace; width 46.3% of length; palm 
length 37.0% of chela length; dactyl length 
1.6 times palm length. Dorsomesial 4 sur- 
face of palm studded with tubercles, me- 
sialmost row composed of 6 (left 7) tuber- 
cles, dorsolateral half punctate, punctations 
deep and large in vicinity of dorsolateral 
base of fixed finger; lateral surface of palm 
and fixed finger subcostate; ventral surface 
of palm punctate, with small corneous tu- 
bercle on articular rim opposite base of dac- 
tyl; no subpalmar tubercle (Fig. 1D). Both 
fingers of chela with well defined submedian 
ridges dorsally and ventrally; opposable 
margin of fixed finger with row of 6 tubercles 
(fourth from base enlarged) along proximal 
?/; of finger and additional large one on lower 
level at base of distal fourth. Opposable 


VOLUME 106, NUMBER 3 


margin of dactyl with row of 9 tubercles, 
(first and fourth from base larger) along 
proximal *%; single row of minute denticles 
extending distally from fifth (sixth on left) 
tubercle; mesial surface of dactyl with row 
of 4 adpressed tubercles (7 left) basally giv- 
ing way to punctations distally. Dorsome- 
dian longitudinal ridges on both fingers well 
developed. Moderate dorsolateral impres- 
sion at base of fixed finger. 

Carpus of cheliped (Fig. 4L) with distinct 
furrow dorsally; dorsomesial surface with 
row of 8 (left 7) low tubercles; dorsolateral 
surface with sparse punctations; mesial sur- 
face with 1 large spiniform tubercle and 3 
additional small ones; ventral surface with 
1 spiniform tubercle on distal articular rim. 
Merus with 2 premarginal tubercles dorsal- 
ly, ventrolateral row of 4 (2 reduced on left) 
tubercles, and ventromesial row of 12 (11 
left); podomere otherwise smooth. Ventral 
ridge of ischium with 4 small tubercles. Is- 
chium of third pereiopod (Fig. 4K) with 
simple hook extending proximally over ba- 
sioischial articulation, not opposed by tu- 
bercles on basis. Coxa of fourth pereiopod 
(Fig. 4K) with vertically disposed cau- 
domesial boss; that of fifth pereiopod (Fig. 
4K) lacking boss, its ventral membrane 
bearing oblique row of small sclerites armed 
with stiff setae. 

First pleopods contiguous at base (Fig. 
4D), reaching coxa of third pereiopod; cen- 
tral projection (Fig. 4B, F) short, not taper- 
ing, lacking subapical notch, rounded api- 
cally, and not extending beyond mesial 
process; mesial process conical, tapering, and 
directed essentially caudolaterally. Both ter- 
minal elements bent caudally at angle slight- 
ly greater than 90°; caudal knob absent. 

Allotype female.—Excluding secondary 
sexual characteristics, differing from holo- 
type in following respects: areola length 
42.1% of total length of carapace; cephalo- 
median lobe of epistome with thickened 
margins; antennal scale 2.8 times as long as 
broad; right chela 71.5% of carapace length; 
opposable margin of dactyl with row of 8 


539 


tubercles (7 left), only first tubercle enlarged; 
mesial surface of dactyl with 7 (6 left) squa- 
mous tubercles; merus with 4 (3 left) pre- 
marginal tubercles dorsally, ventrolateral 
row of 3 tubercles, and ventromesial row of 
11. 

Annulus ventralis (Fig. 41) deeply em- 
bedded in V-shaped sternum, subcircular in 
outline, with narrow median longitudinal 
furrow in cephalic half ending in central de- 
pression; tongue extending caudosinistrally 
across caudal side of depression, disap- 
pearing beneath thickened caudosinistral 
wall; sinus reverse S-shaped and tilted sinis- 
trally at almost 90° ending under caudal wall 
slightly dextral to median line. Postannular 
sclerite oval. First pleopod reaching mid- 
length of annulus when abdomen flexed. 

Morphotypic male, Form II.—Differing 
from holotype in following respects: areola 
length 41.6% of carapace length; antennal 
scale 2.8 times as long as broad; right chela 
68.6% of carapace length; palm length 33.0% 
of chela length; opposable margin of right 
fixed finger without enlarged tubercle (third 
enlarged on left); tip of right fixed finger 
slightly damaged; opposable margin of dac- 
tyl with first and second tubercles enlarged 
(third on left); merus with 3 premarginal 
tubercles dorsally, ventrolateral row of 10 
tubercles and ventromesial row of 9; central 
projection of first pleopod (Fig. 4C, E) non- 
corneous and blunt. 

Type locality. —A roadside ditch on the 
property of the Union Elementary School 
at the intersection of State Route (St Rte) 
79 and County Road (Co Rd) 18, Section 
22, Perry Township, Coshocton County, 
Ohio, (2.1 air km NW of West Carlisle; 5.6 
air km SSE of New Guilford), [40°12’45’N, 
82°07'50” W]. The specimens were dug from 
burrows without chimneys in a ditch having 
permanently flowing water from a spring. 
The surrounding vegetation was grass (a 
lawn). The collection was made on 12 July 
1989 and consisted of 8 4II and 10 2. Some 
of the males were kept alive in the labora- 
tory until they molted which occurred be- 


540 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


tween 11-17 September. The growth incre- 
ments (carapace length) were 0.8 to 1.0 mm 
per individual. 

Disposition of the types.—The holotype, 
allotype, and morphotype are in the collec- 
tion of the National Museum of Natural 
History, Smithsonian Institution, Washing- 
ton, D.C. (USNM 260068, 260069, and 
260070, respectively). Paratypes (7 4I and 
9 9) are in The Ohio State University at 
Newark Crayfish Museum. Specimens from 
localities other than the type locality are 
excluded from the type series. 

Range and specimens examined. —I\ have 
examined 434 specimens of which 91 were 
(Form I) males, 133 were (Form 2) males, 
and 210 were females, from 151 localities. 
Since the number of collections is large, only 
one collection from each county is cited. 
The entire list, however, may be obtained 
from the author or The Ohio State Univer- 
sity at Newark library. Most of the collec- 
tions were made in roadside ditches and 
seeps. Unless stated otherwise, the collec- 
tors were R. F. Jezerinac (RFJ) and G. W. 
Stocker (GWS)—Coll 1, RFJ, GWS, and D. 
Chrisman (DC)—Coll 2, and RFJ, GWS, 
and T. Jones (TJ)—Coll 3. 

KENTUCKY-Bell Co: Fourth intersec- 
tion (inters) E of 15th Street in Middlesboro, 
1 Apr 1986, Coll 1, 3 2. Boone Co: St Rte 
20, 0.3 km (0.2 mi) W of Interstate (1) 275, 
19 Mar 1987, Coll 1, 1 2. Boyd Co: St Rte 
757, 3.2 km (2.0 mi) W of Whites Creek 
Road (Rd), 1 Sep 1986, Coll 1, Dave Hile 
(DH), 2 2. Carter Co: St Rte 1, 0.8 km (0.5 
mi) S of County (Co) Rd 1496, 1 Sep 1986, 
Coll 1, DH, 3 2. Estill Co: Inters St Rte 52 
and Co Rd 1, 2 Apr 1986, Coll 1, 3 2. Grant 
Co: St Rte 22, 1.0 km (0.6 mi) W of Co Rd 
36, 16 May 1989, RFJ, 1 I, 1 2. Greenup 
Co: St Rte 1, 0.2 km (0.1 mi) S of Co Rd 
1459, 6 Aug 1984, Coll 1, 1 6II, 1 2. Knox 
Co: G. R. Hampton Elementary School, St 
Rte 11 in Barborville, 1 Apr 1986, Coll 1, 
4 61, 8 2. Laurel Co: Sublimity Elementary 
School in London, 2 Apr 1986, Coll 1, 1 2. 
Lawrence Co: Roe Creek Rd just W of Unit- 


ed States Route (U.S. Rte) 23, 1 Sep 1986, 
Coll 1, DH, 1 ¢II, 1 2. Morgan Co: Inters 
U.S. Rte 460 and Co Rd 1000, 16 Apr 1988, 
Coll 1, M. Allen, 3 2. Oldham Co: Pattons 
Creek Rd, 3.2 km (2.0 mi) W of U.S. Rte 
42, 19 Apr 1980, RFJ, J. Thoma, M. 
McCluskey (MM), 2 2—2 2 ovig. Powell Co: 
Inters Main Street and Wells Street in Clay 
City, 2 Apr 1986, Coll 1, 1 2. Taylor Co: St 
Rte 70, 1.6 km (1.0 mi) SW of St Rte 337, 
25 Mar 1987, Coll 1, 1 9. Whitley Co: Inters 
St Rte 1277 and U.S. Rte 25W, 25 Mar 
1985, Coll 1, 1 2. 

OHIO-Adams Co: St Rte 41, 0.2 km (0.1 
mi) N of Township (Twp) Rd 125, 13 Jun 
1983, Coll 1, 3 6II, 3 2. Carroll Co: Co Rd 
20, 1.6 km (1.0 mi) N of St Rte 542, 2 Jun 
1984, Coll 1, D. M. Williams (DMW), 1 ¢I. 
Clinton Co: Inters St Rte 380 and Twp Rd 
260, 12 Oct 1985, Coll 2, 1 2. Coshocton 
Co: Type Locality, 12 Jul 1987, Coll 1, R. 
F. Thoma, N. Gillombardo, Z. Thoma, 8 
éII, 10 2. Crawford Co: Co Rd 12, 1.6 km 
(1.0 mi) NE of Twp Rd 117, 18 May 1984, 
Coll 1, 1 éI, 1 2—ovig. Erie Co: E edge of 
Crystal Rock, 22 Aug 1976, J. Norrocky, 1 
2. Fairfield Co: Co Rd 69, 0.8 km (0.5 m1) 
N of Revenge, 1 Aug 1982, MM, RFJ, 1 2. 
Franklin Co: Twp Rd 5, 0.3 km (0.2 mi) E 
of Co Rd 107, 1 Mar 1983, D. Rice (DR), 
G. Phiney (GP), 1 6I. Gallia Co: Co Rd 50 
1.6 km (1.0 mi) NW of St Rte 790, 8 Jun 
1984, Coll 2, 2 2—1 2 with young. Greene 
Co: Co Rd 22, 1.6 km (1.0 mi) Sto@spRee 
35, 12 May 1984, Coll 1, 3 61, 1 4Il, 1 9— 
ovig. Hardin Co: Co Rd 22, 0.2 km (0.1 mi) 
E of Ramshorn Rd, 14 Apr 1985, Coll 1, 
R. J. Jezerinac, 1 4641. Highland Co: Twp Rd 
124, 0.8 km (0.5 mi) S of St Rte 506, 31 
Mar 1984, Coll 1, DMW, 1 I, 1 2. Hocking 
Co: Co Rd 11, 0.5 km (0.3 mi) E of Twp 
Rd 237, 23 Jun 1984, V. Stocker (VS), GWS, 
1 611. Huron Co: Co Rd 167, 0.8 km (0.5 
mi) W of St Rte 60, 8 Aug 1982, RFJ, 1 2. 
Jackson Co: St Rte 93, 0.3 km (0.2 mi) S of 
Co Rd 36, 1 May 1983, Coll 1, RFT, 1 41, 
1 6II, 3 2. Lawrence Co: Co Rd 5, 0.3 km 
(0.2 mi) S of Twp Rd 198, 20 Apr 1986, 


VOLUME 106, NUMBER 3 


Coll 2, 1 4II, 2 2, 1 2—ovig. Licking Co: St 
Rte 586, 2.4 km (1.5 mi) NW of St Rte 16, 
19 Mar 1983, RFT, RFJ, 3 éI. Logan Co: 
Twp Rd 127, 0.2 km (0.1 mi) W of Twp Rd 
129, 22 Aug 1982, RFJ, 1 ¢ Il, 1 9. Ma- 
honing Co: 1.4 km (0.9 mi) NW of Sebring, 
7.5 km (4.7 mi) NE of Alliance, 20 Oct 1979, 
RFT, 1 °. Marion Co: St Rte 98, 1.3 km 
(0.8 mi) S of St Rte 95, 16 May 1982, K. 
Matesich, RFJ, 1 41. Madison Co: U.S. Rte 
42, 0.2 km (0.1 mi) N of Co Rd 145, 10 Jul 
1982, RFJ, 1 sII. Medina Co: Twp Rd 94, 
0.2 km (0.1 mi) W of Co Rd 59, 6 Aug 1982, 
RFJ, 1 °. Meigs Co: St Rte 124, 1.8 km (1.1 
mi) E of St Rte 246, 7 Aug 1985, Coll 1, 1 
61, 1 611. Monroe Co: St Rte 7 at mile post 
5.5, 9 July 1983, GWS, 1 °. Morrow Co: 
Twp Rd 124, 0.8 km (0.5 mi) W of St Rte 
61, 28 Apr 1984, J. Frenton, GWS, 2 I. 
Muskingum Co: St Rte 146, 0.3 km (0.2 m1) 
W of Chandlersville, 19 May 1985, K. Ba- 
Kee) < Perry Co: St Rte 37, 0.2 km (0.1 
mi) E of Co Rd 23, DR, GP, 4 61. Pickaway 
Co: Co Rd 280, 1.6 km (1.0 mi) E of Twp 
aoe, 2 Oct 1983, Coll 1, 1 oI, 2 9. Pike 
Co: Co Rd 65A, 0.3 km (0.2) E of Co Rd 
68, 5 May 1984, Coll 1, RFT, 4 2. Seneca 
Co: St Rte 53, 0.8 km (0.5 mi) S of Twp Rd 
92, 19 May 1985, Coll 1, 1 41, 1 9. Summit 
Co: Co Rd 253, 11 Aug 1983, Ohio Envi- 
ronmental Protection Agency, 1 éII. Tus- 
carawas Co: Co Rd 69, 2.1 km (1.3 mi) E 
of St Rte 250, 2 Jun 1984, Coll 1, DMW, 
2 61, 3 2, 1 2—ovig. Vinton Co: St Rte 278, 
3.2 km (2.0 mi) N of St Rte 667, 14 Mar 
1983, DR, GP, 1 4I. Washington Co: St Rte 
7, 0.2 km (0.1 mi) E of Co Rd 46, 7 Aug 
1985. Coll 1, 1 9. 
PENNSYLVANIA-Beaver Co: Rd S of 
Rural Road (RR) 04048, 1 Jul 1985, VS, 
Gws. 2 2. Butler Co: RR 10113, 0.5 km 
(0.3 mi) S of St Rte 422, 27 Jun 1984, VS, 
GWS, 1 II. Fayette Co: Inters RR 26022 
and St Rte 819, 18 Jun 1984, Coll 1, 1 41, 
1 II. Greene Co: RR 3009, 2.1 km (1.3 mi) 
W of St Rte 19, 26 May 1984, GWS, 1 9— 
ovig. Washington Co: S of I-70 exit, (2.4 air 
km S of Denningsville), 19 Jul 1984, VS, 


541 


GWS, 1 2. Westmoreland Co: RR 64015 
just N of Twp Line, (3.8 km SE of Milligan), 
28 Jun 1984, VS, GWS, 1 4Il, 4 &. 

TENNESSEE-Monroe Co: Inters U.S. 
Rte 128 and St Rte 33 at Monroe-Loudon 
County Line, 31 Mar 1986, Coll 1, 1 41, 1 
2. Sevier Co: St Rte 338, 4.8 km (3.0 mi) E 
of Boyds Creek Rd, 31 Mar 1986, Coll 1, 1 
OH: 148. 

WEST VIRGINIA-Barbour Co: Arden 
Rd: 1.3" km; (0/8emi) E-of U.S. Rte 119.8 
Apr 1986, Coll 1, 1 I. Braxton Co: U.S. 
Rte 19, 6.2 km (3.8 mi) E of St Rte 5, 25 
May 1985, Coll 2, 1 41. Cabell Co: Guyan 
Creek Rd, 4.0 km (2.5 mi) NE of St Rte 2, 
19 Oct 1985, Coll 2, 1 41. Dodridge Co: St 
Rte 18, 0.8 km (0.5 mi) S of Co Rd 66, 14 
Apr 1986, Coll 1, 2 2—1 2 ovig. Gilmer Co: 
Rd to Cedar Run State Park, (2.9 air km 
SSW of Glenville), 25 May 1985, Coll 2, 1 
éII, 2 2. Harrison Co: St Rte 20, 2.6 km (1.6 
mi) E of Dola, 24 Aug 1984, VS, GWS, 1 
2. Kanawha Co: Inters St Rte 25 and Co Rd 
25/13, 9 Apr 1988, GWS, 1 2. Lincoln Co: 
Co Rd 40, 1.6 km (1.0 m1) S of St Rte 3, 
21 Jun 1989, Coll 1, 1 2. Logan Co: Co Rd 
5! 0:Seknr (0S mi)" Sof Co Rd 328" Jul 
1988, GWS, TJ, 1 41, 1 ¢II. Lewis Co: Inters 
Co Rd 10 and Co Rd 10/8, 8 Sep 1988, Coll 
1, 1 2. Marion Co: Inters U.S. Rte 250 and 
Co Rd 8, 12 Aug 1988, Coll 3, 1 @II, 3 2. 
Mason Co: U.S. Rte 33, 0.8 km (0.5 mi) S 
of Graham Station, 6 Aug 1985, Coll 1, 1 
éI. Putnam Co: Co Rd 5, 0.6 km (0.4 mi) 
SW of U.S. Rte 35, 1 Oct 1988, Coll 3, 1 2. 
Ritchie Co: Co Rd 50/34, 1.8 km (1.1 m1) 
NE of U.S. Rte 50, 17 Jul 1988, Coll 3, 1 
2. Roane Co: St Rte 27, 2.4 km (1.5 mi) W 
of St Rte 29, 11 Sep 1988, Coll 3, 1 2. Taylor 
Co: Co Rd 3, 2.4 km (1.5 mi) N of St Rte 
76, 11 Aug 1988, Coll 3, 1 II, 1 2. Tyler 
Co: Conaway Run Lake State Park camp- 
ground, 13 Apr 1986, Coll 1, 1 2. Upsur Co: 
St Rte 20, 1.9 km (1.2 mi) S of St Rte 4, 25 
May 1985, Coll 2, 2 II. Wayne Co: U.S. 
Rte 52, 1.4 km (0.9 mi) W of St Rte 35, 20 
Oct 1985, Coll 2, 4 4II, 4 2. Wood Co: Co 
Rd 11, 2.9 km (1.8 mi) N of St Rte 68, Coll 


542 


Table 2.—Seasonal data of C. (T.) thomai, new spe- 
cies. 


Second 
form 
males 


First 
form 


males *Females Sex ratio 


Month (number) (number) (number) (M:F) erous 
February 2 
March 26 7 15 @24) 1 
April 19 3 38 (leis7) 5 
May 10 11 32 (let) 10 
June fi 19 35 (1:1.4) Z 
July Ps 14 24 CieT.5) 
August 10 26 34 CUSIEt) 
September 8 7 20 (1:1.3) 
October 3 5 et (1:1.4) 


@ Numbers include ovigerous females. 


1, 1 61. Wyoming Co: St Rte 971, 3.7 km 
(2.3 mi) SW of St Rte 10, 6 Jul 1988, Coll 
L 1 Gl hg: 

Color notes. —In the central portion of the 
species range, the color pattern is rather uni- 
form and consists of a dark brownish-olive 
carapace and abdomen. The tips of the fin- 
gers, lateral margin of the Chela, and dorsal 
knob on the chela at the base of the dactyl 
is orangish to reddish. The rostral margins 
are cream. The undersurface is cream to 
white. Red bands are never present on the 
rim of the articulation joints and caudal 
margins of the abdominal tergites. 

The color patterns at the western and 
southern periphery of the range is more 
variable. In southwestern Ohio (Highland 
County), the carapace tends to be chestnut- 
brown with the abdomen being darker than 
the thoracic portion of the carapace. A 
darker brown band is present on the ante- 
rior flank of the cervical groove. The lateral 
margin of the finger, the palmer tubercles, 
tubercles on the dorsomesial surface of the 
merus, and the mesial spine on the merus 
are orangish. The dorsal knobs on the distal 
rim of the palm of chela at the base of the 
dactyl and the knob on the ventral rim are 
reddish. The undersurface is cream to white. 
In northern Kentucky and southern West 
Virginia, the basic body color tends to be 
emerald green with additional structures 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


colored like those in southwestern Ohio. 
Two specimens were collected from Clinton 
County, Ohio, (one specimen was de- 
stroyed) the basic body color of which was 
speckled emerald green with orange and red 
as described above. Two individuals, one 
collected in Medina County, Ohio, and the 
other from Marion County, West Virginia, 
were blue, apparently lacking red chromato- 
phores. 

Variations. — As expected in a species that 
occupies a wide geographical area, minor 
variations occur in most body structures and 
body proportions. The material was ex- 
amined for clinal variations and characters 
that might be restricted to local populations, 
but none was found. However, those spec- 
imens collected in Tennessee tend to have 
a more deeply excavate rostrum. In this spe- 
cies, there appears to be more variation in 
color than in body structures. Additional 
meristic and morphometric data (simple 
descriptive statistics, ratios, and regression 
analysis) are available from the author or 
the library at The Ohio State University at 
Newark. 

Size.—The largest specimen examined 
was a female with a carapace length of 53.8 
mm from Mason County, West Virginia. 
The largest Form I male measured 51.9 mm 
and the smallest 26.6 mm. For measure- 
ments see Table 1. 

Life-history notes.—Ovigerous females 
were collected on 31 March; 14, 19, and 20 
April; 1, 4, 7, 12, 18, and 26 May; and 2 
June. Females with young attached were 
captured on 27 May and 8 and 11 June. 
Form I males were found from 1 March 
through 18 June and from 19 July to 19 
October. Additional seasonal data are pre- 
sented in Table 2. 

Crayfish associates. —Collected with C. 
(T.) thomai at one or more sites in Ohio 
were C. (C.) b. cavatus Hay (1902), C. (C.) 
ortmanni Williamson (1907), C. (C.) sci- 
otensis Rhoades (1944b), C. (P.) robustus 
Girard (1852). In the other states, its as- 
sociates included C. (C.) b. cavatus, C. (C.) 


VOLUME 106, NUMBER 3 


b. carinirostris Hay (1914), C. (J.) monon- 
galensis Ortmann (1905a), C. (J.) dubius 
Faxon (1884), C. (P.) robustus, and C. (C.) 
sclotensis. 

Relationships. —Cambarus (T.) thomai is 
most closely related to C. (T.) acanthura but 
differs from the latter in that the distome- 
dian spine on the mesial ramus of the uro- 
pod does not overreach the distal margin of 
the ramus and the ventral surface of the 
carpus has a spiniform tubercle on the distal 
articular rim. Sometimes the spiniform tu- 
bercle is observed on C. (7.) acanthura but 
it is never well developed. 

Etymology.—I take pleasure in naming 
this crayfish in honor of Roger Francis 
Thoma of the Ohio Environmental Protec- 
tion Agency, a student of crayfishes and a 
friend. 


Acknowledgments 


I thank all individuals who helped collect 
the specimens used in this study, especially 
G. W. Stocker, D. Chrisman, and T. Jones. 
Special thanks are extended to Dr. H. H. 
Hobbs, Jr., the Smithsonian Institution, Dr. 
H. H. Hobbs, III, Wittenberg University, 
and Dr. J. F. Fitzpatrick, Jr., the University 
of South Alabama, and two anonymous re- 
viewers for their comments concerning the 
manuscript. Dave Dennis and Susan Hen- 
drix of The Ohio State University prepared 
Fig. 4 and funds for them were provided by 
The Ohio State University, Department of 
Zoology. Partial funding for publication was 
provided by the Professional Standards 
Committee of The Ohio State University at 
Newark. 


Literature Cited 


Bouchard, R. W. 1972. A contribution to the knowl- 
edge of Tennessee crayfishes. Unpublished Ph.D. 
dissertation, University of Tennessee, Knox- 
ville, 113 pp. 

1975. Geography and ecology of crayfishes 
of the Cumberland Plateau and Cumberland 
Mountains, Kentucky, Virginia, Tennessee, 
Georgia and Alabama. Part II. The genera Fal- 


_—_————— 


543 


licambarus and Cambarus. Pp. 585-605 in J. 
W. Avault, Jr., ed., Freshwater crayfish II. Lou- 
isiana State University Division of Continuing 
Education, Baton Rouge, 676 pp. 

Faxon, W. 1884. Descriptions of new species of Cam- 
barus, to which is added a synonymical list of 
the known species of Cambarus and Astacus. — 
Proceedings of the American Academy of Arts 
and Sciences 20:107-158. 

. 1885. A revision of the Astacidae. Part I. The 

genera Cambarus and Astacus. —Memoirs of the 

Museum of Comparative Zoology at Harvard 

College 10(4):v + 186. 

Fitzpatrick, J. F., Jr. 1978. A new burrowing crawfish 
of the genus Cambarus from southwestern Al- 
abama (Decapoda: Cambaridae).— Proceedings 
of the Biological Society of Washington 90:367- 
374. 

Girard, C. 1852. A revision of the North American 
Astaci, with observations on their habits and 
geographic distribution.—Proceedings of the 
Academy of Natural Science of Philadelphia 
6:87-91. 

Hay, W. P. 1899. Synopsis of North American in- 

vertebrates, VI: The Astacidae of North Amer- 

ica.—American Naturalist 33(396):957-966. 

1902. Observations on the crustacean fauna 
of Nickajack Cave, Tennessee. — Proceedings of 
the United States National Museum 25(1292): 
417-439. 

1914. Cambarus bartonii carinirostris Hay. 
Pp. 384—385 in Water Faxon, ed., Notes on the 
crayfishes in the United States National Mu- 
seum and Museum of Comparative Zoology, 
with descriptions of new species and subspecies 
to which is appended a catalogue of the known 
species and subspecies.— Memoirs of the Mu- 
seum of Comparative Zoology at Harvard Col- 
lege 40(8):349-427. 

Hobbs, H. H., Jr. 1969. On the distribution and phy- 

logeny of the crayfish genus Cambarus. Pp. 93- 

178 in P. C. Holt, R. L. Hoffman, & C. W. Hart, 

Jr., eds., The distributional history of the biota 

of the southern Appalachians. Part I: Inverte- 

brates. Research Monograph 1, Virginia Poly- 
technical Institute, Blacksburg, 295 pp. 

1972. Crayfishes (Astacidae) of North and 
Middle America in biota of freshwater ecosys- 
tems, identification manual. Water Pollution 
Control Research Series, United States Envi- 
ronmental Protection Agency, Washington D.C. 
9:1-173. 

1974. A checklist of the North and Middle 
American crayfishes (Decapoda: Astacidae and 
Cambaridae).—Smithsonian Contributions to 
Zoology 166:161 pp. 

1981. The crayfishes (Decapoda: Cambari- 


544 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


dae) of Georgia.—Smithsonian Contributions 

to Zoology 318:549 pp. 

1989. An illustrated checklist of the Amer- 
ican crayfishes (Decapoda: Astacidae, Cambari- 
dae, and Parastacidae).—Smithsonian Contri- 
butions to Zoology 488:236 pp. 

Jezerinac, R. F. 1985. On the two forms of Cambarus 

(Lacunicambarus) diogenes (Decapoda: Cam- 

baridae) in Ohio.—Ohio Journal of Science 

85(2):7 (Abstract). 

1986. Endangered and threatened crayfishes 
(Decapoda: Cambaridae) of Ohio.— Ohio Jour- 
nal of Science 86(4):177—180. 

—., & G. W. Stocker. 1989. Distributions of the 
primary burrowing crayfishes of the genera Fal- 
licambarus and Cambarus (Decapoda: Cam- 
baridae) in West Virginia. — Ohio Journal of Sci- 
ence 89(2):2—3 (Abstract). 

——., & 1990. Distributions of the cray- 
fishes (Decapoda: Cambaridae) of West Virgin- 
ia. Part II. The genera Cambarus and Fallicam- 
barus. —Proceedings of the West Virginia 
Academy of Science 62(1):7—8 (Abstract). 

—., & R. F. Thoma. 1984. An illustrated key to 
Ohio Cambarus and Fallicambarus (Decapoda: 
Cambaridae), with comments and a new sub- 
species record.—Ohio Journal of Science 84(3): 
120-125. 

Lawton, S. M. 1979. A taxonomic and distributional 
study of the crayfishes (Decapoda: Cambaridae) 
of West Virginia with diagnostic keys to species 
of the genera Cambarus and Orconectes. Un- 
published M.S. Thesis, Marshall University, 
Huntington, West Virginia, 107 pp. 

Marlow, G. 1960. The subspecies of Cambarus di- 
ogenes.—American Midland Naturalist 64(1): 
229-250. 

Newcombe, C. L. 1929. The crayfishes of West Vir- 
ginia.—Ohio Journal of Science 29:276-288. 

Ortmann, A. E. 1905a. The crawfishes of western 


Pennsylvania.— Annals of the Carnegie Muse- 

um 3(2):387—406. 

1905b. The mutual affinities of the species 
of the genus Cambarus, and their dispersal over 
the United States.— Proceedings of the Ameri- 
can Philosophical Society 44:91-136. 

1906. The crawfishes of the state of Penn- 
sylvania.—Memoirs of the Carnegie Museum 
2(10):343-523. 

Osburn, R. C., & E. B. Williamson. 1898. The cray- 
fishes of Ohio. — Sixth Annual Report, The Ohio 
State Academy of Science, 1898:11. 

Rhodes, R. R. 1944a. The crayfishes of Kentucky, 

with notes on variation, distribution and de- 

scription of new species and subspecies.— 

American Midland Naturalist 31:111-149. 

1944b. Further studies on distribution and 
taxonomy of Ohio crayfishes, and the descrip- 
tion of a new subspecies. — Ohio Journal of Sci- 

ence 44:95-99. 

Thoma, R. F., & R. F. Jezerinac. 1982. New distri- 
butional records from crayfishes (Cambarus and 
Fallicambarus) from Ohio, including a new sub- 
species record.— Ohio Journal of Science 82(3): 
136-138. 

Turner, C. L. 1926. The crayfishes of Ohio.—Ohio 
Biological Survey 13:145-195. 

Williamson, E. B. 1899. Notes on Ohio Astacidae. — 

Seventh Annual Report of the Ohio Academy 

of Science 1899:47-48. 

1906[1907]. Notes on the crayfish of Wells 
County, Indiana, with descriptions of new spe- 
cies. Pp. 749-763, plate 35 in Thirty-first An- 
nual Report of the Department of Geology and 
Natural Resources, Indiana, 772 pp. 


Department of Zoology, The Ohio State 
University at Newark, 1179 University 
Drive, Newark, Ohio 43055, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(3), 1993, pp. 545-553 


TWO NEW RECORDS OF THE GENUS HEPTACARPUS 
(CRUSTACEA: DECAPODA: HIPPOLYTIDAE) 
FROM JAPANESE WATERS 


Tomoyuki Komai 


Abstract.—Two species of the hippolytid genus Heptacarpus, H. maxillipes 
(Rathbun, 1902) and H. moseri (Rathbun, 1902), both of which have been 
known mainly from the eastern Pacific Ocean including the Bering Sea, are 
recorded from Japan for the first time on the basis of specimens collected from 
the Pacific coast of northern Japan. The identification was confirmed by com- 
parison with American specimens. For each species, a synonymy and detailed 
description, including the color pattern of fresh material, are given. 


The hippolytid genus Heptacarpus 
Holmes now contains 33 species, which are 
all restricted to the North Pacific Ocean. 
Hayashi (1992) reported 11 species from 
Japan and its adjacent waters, all from lit- 
toral or sublittoral zones. 

Recent collections from the Pacific coast 
of northern Japan revealed the presence of 
two bathyal species of Heptacarpus, H. 
maxillipes (Rathbun, 1902) and H. moseri 
(Rathbun, 1902). These two species have 
been reported from the eastern Pacific in- 
cluding the Bering Sea (Rathbun 1902, 1904; 
Butler 1980), and H. maxillipes has been 
reported near Kurile Islands (Birshtein & 
Zarenkov 1970). Since descriptions of these 
two species by the previous authors are in- 
adequate, I describe them herein in detail. 

The following abbreviations are used in 
the text: CL, postorbital carapace length; 
HUMZ, Laboratory of Marine Zoology, 
Faculty of Fisheries, Hokkaido University; 
USNM, National Museum of Natural His- 
tory, Smithsonian Institution, Washington, 
D.C. 


Heptacarpus maxillipes (Rathbun, 1902) 
Figs. 1A, 2, 3 


Spirontocaris maxillipes Rathbun, 1902: 
898.—Rathbun, 1904:92, fig. 40. 


Heptacarpus maxillipes.—Holthuis, 1947: 
12 (list).—Birshtein & Zarenkov, 1970: 
420.—Hayashi, 1979:12, tab. 1.—Hay- 
ashi, 1992:110, tab. 32. 


Material examined. —HUMZ-C 936, East 
of Cape Erimo, 42°02.3’N, 144°07.9’E, 915— 
970 m, 6 Sep 1989, otter trawl (T/S Oshoro- 
Maru), 1 female (5.5 mm CL), coll. T. Ko- 
mai; HUMZ-C 1003, off Fukushima Pref., 
37°07.5'N, 142°41.2’E, 412 m, 1 Jun 1989, 
otter trawl (R/V Tanshu-Maru), 2 females 
(6.8 mm CL), coll. O. Yamamura. 

Comparative material. —-USNM 183004, 
Aleutian Islands, 2 males (5.1, 5.2 mm CL), 
1 female (7.0 mm: CL). 

Description. —Integument thin, smooth. 
Rostrum (Fig. 2A) slightly curved upward, 
reaching or slightly overreaching distal end 
of antennular peduncle, 0.86—0.89 times as 
long as carapace, with ventral blade rather 
deep and dorsal blade poorly developed; lat- 
eral carina axis-like, sharply ridged proxi- 
mally; apex sharply pointed; dorsal margin 
with 6 teeth over entire length, including 2 
on carapace posterior to orbital margin, 
posteriormost tooth situated at anterior 4 
of carapace; ventral margin convex, with 4 
small teeth on distal %5. Carapace (Figs. 1A, 
2A, B) with suborbital lobe rounded from 
dorsal aspect, falling far short of antennal 


546 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


FSS 


0 
yan 
ee 
aa 
rs 
ae 
qo 


Fig. 1, 


A, Heptacarpus maxillipes (Rathbun, 1902), HUMZ-C 1003, female, 6.8 mm CL, entire animal; B, 


Heptacarpus moseri (Rathbun, 1902), HUMZ-C 1056, female, 9.6 mm CL, entire animal. 


tooth; pterygostomian tooth small; postros- 
tral median carina becoming obscure just 
behind posteriormost median tooth. 

Abdominal somites (Figs. 1A, 2C) with- 
out carinae or dorsal teeth, not strongly ge- 
niculate. Pleuron of fourth somite with weak 
tooth at posteroventral angle. Sixth somite 
0.7 times as long as carapace and 2.1-2.9 
times as long as proximal depth. Telson (Fig. 
2D) 0.9 times as long as carapace, 3.4 times 
as long as proximal width, lateral margins 
of anterior %4 subparallel, remaining * ta- 
pering to convex posterior margin; dorsal 
surface slightly convex, armed with 3 pairs 
of dorsolateral spines and 3 pairs on pos- 
terior margin. 


Cornea (Fig. 2A, B) fairly large, strongly 
dilated, without ocellus. 

Antennule (Fig. 2A, B) with peduncle 
reaching at least to distal 4 of scaphocerite, 
proximal segment longer than distal 2 seg- 
ments combined, without teeth on distal 
margin; stylocerite sharp, falling slightly 
short of distal margin of proximal segment; 
intermediate segment longer than wide, with 
strong dorsolateral tooth; distal segment also 
longer than wide, with dorsodistal tooth. 
Outer flagellum with proximal aestetascs 
bearing portion composed of 10 or 11 thick- 
ened articles. Inner flagellum slender, slight- 
ly longer than outer flagellum. 

Antenna (Fig. 2A, B) with carpocerite 


VOLUME 106, NUMBER 3 547 


Cry 14 PE 


AM 


i] 


Fig. 2. Heptacarpus maxillipes (Rathbun, 1902). HUMZ-C 936, female, 5.5 mm CL. A, anterior carapace, 
rostrum, and cephalic appendages in left aspect; B, same in dorsal aspect; C, posterior three abdominal somites 
in left aspect; D, telson in dorsal aspect; E, left third maxilliped; F, left first pereopod; G, same, chela in extensor 
aspect; H, left second pereopod; I, same, chela in extensor aspect; J, left third pereopod; K, same, dactylus in 
lateral aspect. 


548 


0.5 mm 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Heptacarpus maxillipes (Rathbun, 1902). USNM 183004, male, 5.2 mm CL. A, anterior carapace, 
rostrum, and cephalic appendages in lateral aspect, eye removed; B, endopod of first pleopod in dorsal aspect; 
C, appendix interna and appendix masculina of second pleopod in dorsomesial aspect. 


reaching distal % of scaphocerite. Scapho- 
cerite 0.80—0.82 times as long as carapace, 
3.2 times as long as wide; lateral margin 
almost straight, distolateral tooth falling 
somewhat short of blade. 

Mouthparts typical of genus. Third max- 
illiped (Fig. 2E) long and relatively thin, 
overreaching scaphocerite by length of dis- 
tal “% of ultimate segment; ultimate segment 
3.3 times as long as carpus, mesial surface 
with numerous clusters of stiff setae, and 
tipped with 5 or 6 corneous spines; coxa 
with epipod; exopod absent. 

Pereopods without epipods. First pereo- 
pod (Fig. 2F) fairly stout, overreaching dis- 
tal end of scaphocerite by length of distal '4 
of chela; dactylus (Fig. 2G) 0.7 times as long 
as palm; palm 2.7 times as long as wide; 
carpus about half length of chela; merus 6.1 
times as deep as long, unarmed. Second pe- 
reopod (Fig. 2H) somewhat longer than first 
pereopod, slender, overreaching scaphocer- 
ite by length of chela and distal three articles 
of carpus; dactylus and fixed finger (Fig. 21) 
thickly setose distally, dactylus about half 
length of palm; carpus composed of 7 ar- 
ticles, second and third articles 1.6 times 
and 2.3 times as long as first article respec- 
tively. Third to fifth pereopods relatively 
long, almost similar morphologically. Third 
pereopod (Fig. 2J) overreaching scaphoce- 
rite by length of dactylus and almost all of 


propodus; dactylus (Fig. 2K) 0.2 times as 
long as propodus, bifid terminally, with 5 
or 6 accessory spinules over entire length 
of flexor margin; propodus with 2 rows of 
spinules on flexor margin; carpus 0.47 times 
as long as propodus; merus with 3 lateral 
spines in distal half. Fourth pereopod over- 
reaching scaphocerite by length of dactylus 
and distal ¥%4 of propodus; merus with 1 or 
2 lateral spines in distal 4. Fifth pereopod 
overreaching scaphocerite by length of dac- 
tylus and distal half of propodus; merus with 
1 subterminal spine. 

First pleopod with endopod convention- 
al. Uropod (Fig. 1A) with endopod slightly 
overreaching tip of telson; exopod slightly 
longer than endopod, lateral margin nearly 
straight. 

Coloration.—In fresh condition, entire 
animal transparent with red dots over entire 
body, devoid of distinct band or pattern. 
Distal spines of third maxilliped, tip of fin- 
gers of first chela and tip of dactylus of last 
three pairs of pereopods dark brown. 

Biological note.—The three specimens 
examined were collected from muddy bot- 
tom of the continental slope at a depth rang- 
ing from 412 to 970 m. The female speci- 
mens collected off Fukushima have well- 
developed ovaries which are visible through 
the carapace. 

Remarks. —Birshtein & Zarenkov (1970) 


VOLUME 106, NUMBER 3 


recorded for the first time Heptacarpus 
maxillipes from the Western Pacific (off Ku- 
rile Islands), but they did not give descrip- 
tion or figures of their specimens. I com- 
pared the present three female specimens 
with one female and two male specimens 
from the Aleutian Islands deposited at the 
USNM, and I could not find any significant 
differences between the Japanese and Aleu- 
tian specimens. Rathbun (1904) gave the 
proportional length of scaphocerite to the 
carapace as “‘about two-thirds,”’ but the ra- 
tio varies considerably, 0.63-0.81 in the 
Aleutian specimens, and 0.80—0.82 in the 
Japanese specimens. Further, Rathbun 
(1902, 1904) described the proportional 
length of rostrum to the carapace as “‘about 
four-fifths,” but the ratio of the Aleutian 
specimens examined here is 0.88 in the fe- 
male and 0.63 and 0.69 in two males; the 
ratios of the two Japanese specimens in 
which the rostra are intact are 0.89 and 0.94 
respectively. The apparently short rostrum 
in the male (Fig. 3A) may be ascribed to the 
sexual difference, since sexual dimorphism 
in length of the rostrum has been reported 
in other hippolytids (e.g., Greve 1963, Hay- 
ashi 1979, Criales 1992). 

Other than the proportionate length of the 
rostrum, the males differ from the females 
in the following points: the rostrum is much 
shallower than in the female (Fig. 3A); the 
outer and inner antennular flagella are 
somewhat longer than in the female (Fig. 
3A); the endopod of the first pleopod (Fig. 
3B) is slightly longer than half of the exopod 
and tapers gradually to a simple tip; the 
distal portion bears adhesive hooks; the me- 
sial margin is fringed with minute bristles; 
the appendix interna of the second pleopod 
(Fig. 3C) is more slender, and more strongly 
curved than in the female; and the appendix 
masculina (Fig. 3C) is about two-thirds of 
the appendix interna, and is armed with 
about 16 long spines. 

Heptacarpus maxillipes belongs to the 
group having epipods only on the maxilli- 
peds, and three Japanese species, 1.e., H. 
camtchaticus (Stimpson, 1860), H. genicu- 


549 


latus (Stimpson, 1860), and H. pandaloides 
(Stimpson, 1860), have been assigned to this 
group (Hayashi 1979, 1992). Heptacarpus 
maxillipes, however, is distinguished im- 
mediately from them by the moderately 
stout body, the rostrum shorter than the 
carapace, the eye lacking an ocellus, the 
scaphocerite distinctly shorter than the car- 
apace, and the long pereopods. 

In addition, Wicksten (1990) presented a 
key to the hippolytid shrimp of the Eastern 
Pacific Ocean, but she omitted H. maxil- 
lipes. Heptacarpus maxillipes is inserted into 
Wicksten’s key as follows: 


56. Scaphocerite shorter than cara- 


TE CGh le ea ee er 56a 
Scaphocerite as long as or longer 
MiangCatapaGe 4.) % 2... on a7 


56a. Rostrum longer than carapace; 
telson shorter than sixth abdom- 
inal somite; third maxilliped not 
overreaching scaphocerite ..... 
H. decorus (Rathbun) 
Rostrum shorter than carapace; 
telson longer than sixth abdomi- 
nal somite; third maxilliped over- 
reaching scaphocerite 
ae 3, a ea H. maxillipes (Rathbun) 


Sey witai-= ce wel @ im 


Distribution. —Heptacarpus maxillipes 
was recorded from the Aleutian Islands at 
depths of 418 to 1144 m by Rathbun (1904), 
and was recorded subsequently from the Pa- 
cific side of Kurile Islands at depths of 280 
to 1580 m by Birshtein & Zarenkov (1970). 
The present report corroborates the occur- 
rence of this poorly known species in the 
Western Pacific Ocean, and extends further 
its range to the west and south. 


Heptacarpus moseri (Rathbun, 1902) 
Figs. 1B, 4 


Spirontocaris gaimardii.—Rathbun, 1899: 
556 (in part). 

Spirontocaris moseri Rathbun, 1902:897.— 
Rathbun, 1904:91, fig. 39.—? Hart, 1930: 
104. 

Eualus moseri. —Makarov, 1941:126. 


550 


Heptacarpus moseri.—Holthuis, 1947:21 
(list). —Birshtein & Vinogradov, 1953: 
216.—Hayashi, 1979:12, tab. 1.— Butler, 
1980:223, pl. 6A.—Wicksten, 1990:595 
(key).— Hayashi, 1992:110, tab. 32. 


Material examined. —HUMZ-C 1056, off 
Hiroo, Hokkaido, 42°11.45’E, 143°41.6’E, 
325 m, 30 May 1990, otter trawl (R/V Tan- 
shu-Maru), 1 female (9.6 mm CL), coll. T. 
Komai. 

Comparative material. -USNM 183092, 
southwest of East Cape, Attu Island, 
52°55'40N, 173°26’E, 247 m (Albatross Sta- 
tion 1906: 4784), 5 females (7.3-8.7 mm 
CL). 

Description. —Integument thin, smooth. 
Rostrum (Fig. 4A) (injured on distal part) 
horizontal in general, reaching distal end of 
scaphocerite, 1.16 times as long as carapace, 
with deep ventral blade and poorly devel- 
oped dorsal blade; lateral carina axis-like, 
proximal half sharply ridged; apex sharply 
pointed; dorsal margin with 7 teeth over 
entire length, including 2 on carapace pos- 
terior to orbital margin, posteriormost tooth 
situated at about anterior 3 of carapace; 
ventral margin convex, with 5 small teeth 
on about distal half. Carapace (Figs. 1B, 4A, 
B) with suborbital lobe rounded from dorsal 
aspect, falling far short of antennal tooth; 
pterygostomian tooth rather strong; post- 
rostral median carina becoming obscure just 
behind posteriormost median tooth. 

Abdominal somites (Fig. 1B) without ca- 
rinae or dorsal tooth, not strongly genicu- 
late. Pleuron of fourth somite with weak 
tooth at posteroventral angle. Sixth somite 
0.7 times as long as carapace and 1.9 times 
as long as proximal depth. Telson (Fig. 4C) 
0.9 times as long as carapace, 4.2 times as 
long as proximal width, lateral margins of 
anterior 3 subparallel, remaining 7% taper- 
ing to convex posterior margin; dorsal sur- 
face slightly convex, armed with 5 pairs of 
dorsolateral spines and 3 pairs on posterior 
margin. 

Cornea of eye (Fig. 4A, B, D, E) large, 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


strongly dilated; ocellar sinuosity distinct, 
but not forming true ocellus; mesial face of 
ocular peduncle considerably swollen prox- 
imally. 

Antennule (Fig. 4A, B) with peduncle 
reaching middle of scaphocerite; proximal 
segment much longer than distal 2 segments 
combined, without teeth on distal margin; 
stylocerite sharp, reaching beyond middle 
of intermediate segment of peduncle; inter- 
mediate segment longer than wide, with 
strong dorsolateral tooth; distal segment as 
long as wide, with strong dorsodistal tooth. 
Outer flagellum with proximal aestetascs 
bearing portion composed of about 20 
thickened articles. Inner flagellum slender, 
about twice length of outer flagellum. 

Antenna (Fig. 4A, B) with carpocerite 
falling slightly short of middle of scapho- 
cerite. Scaphocerite 0.94 times as long as 
carapace, 4.0 times as long as wide; lateral 
margin almost straight, distolateral tooth 
falling somewhat short of blade. 

Mouthparts typical of genus. Third max- 
illiped (Fig. 4F) stout, overreaching scapho- 
cerite by length of distal half of ultimate 
segment; ultimate segment 2.9 times as long 
as carpus, mesial surface with numerous 
clusters of stiff setae, and tipped with 4 or 
5 corneous spines; coxa with epipod; exo- 
pod absent. 

Pereopods with epipods on first pair. First 
pereopod (Fig. 4G) stout, falling slightly 
short of scaphocerite; dactylus (Fig. 4H) half 
length of palm, palm 3.0 times as long as 
wide; carpus 0.6 times as long as chela; me- 
rus 4.4 times as long as deep, unarmed. Sec- 
ond pereopod (Fig. 41) much longer than 
first pereopod, slender, overreaching scaph- 
ocerite by length of chela and distal 3 arti- 
cles of carpus; dactylus and fixed finger (Fig. 
4J) thickly setose distally, dactylus 0.8 times 
as long as palm; carpus composed of 7 ar- 
ticles, second and third articles 1.3 and 2.3 
times as long as first article respectively. 
Third to fifth pereopods relatively long, al- 
most similar morphologically. Third pereo- 
pod (Fig. 4K) overreaching scaphocerite by 


VOLUME 106, NUMBER 3 554 


Uv 4 
SUI LL Vy md 


Yy y Wy, YYES D 
so 


I ) 


x 
J 


Fig. 4. Heptacarpus moseri (Rathbun, 1902). HUMZ-C 1056, female, 9.6 mm CL. A, anterior carapace, 
rostrum, and cephalic appendages in lateral aspect; B, same in dorsal aspect; C, telson in dorsal aspect; D, eye 
in dorsal aspect; E, same in mesial aspect; F, left third maxilliped; G, left first pereopod; H, same, chela in 
extensor aspect; I, left second pereopod; J, same in extensor aspect; K, left third pereopod; L, same, dactylus in 
lateral aspect. 


552 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


length of dactylus and distal half of prop- 
odus; dactylus (Fig. 4L) 0.19 times as long 
as propodus, bifid terminally, with 4 or 5 
accessory spinules over entire length of flex- 
or margin; propodus with 2 rows of spinules 
on flexor margin; carpus 0.36 times as long 
as propodus; merus with 3 lateral spines in 
distal half. Fourth pereopod overreaching 
scaphocerite by length of dactylus and distal 
’3 of propodus; dactylus 0.18 times as long 
as propodus, with 5 or 6 accessory spinules; 
carpus 0.39 times as long as propodus; me- 
rus with 3 lateral spines. Fifth pereopod 
overreaching scaphocerite by length of dac- 
tylus; dactylus 0.19 times as long as prop- 
odus, with 5 accessory spinules; carpus 0.36 
times as long as propodus; merus with 3 
lateral spines. 

First pleopod with endopod convention- 
al. Uropod with endopod (Fig. 1B) distinct- 
ly overreaching tip of telson; exopod slightly 
longer than endopod, lateral margin nearly 
straight. 

Coloration.—In life, body entirely red- 
dish with translucent background. Abdo- 
men having wide dark red band on each 
somite. Third maxilliped white except me- 
sial surface of distal two segments colored 
with dark red. First and second pereopods 
nearly colorless. Last three pairs of pereo- 
pods with ischium and merus bearing red 
bands on white background; distal three seg- 
ments dark red. Each protopodite of pleo- 
pods having white spot on lateral side; uro- 
pod red in distal half, exopod with two fine 
white spots at distal portion and base of 
distolateral spine respectively. 

Biological note.—The present specimen 
was trawled on a muddy bottom at a depth 
of 325 m, where the temperature was 
Bag tet 

Remarks.—This single female specimen 
was compared with the specimens identified 
as H. moseri from off Attu Island, Aleutian 
Islands, and deposited at the USNM. No 
significant difference was observed. AI- 
though Rathbun (1904) gave the propor- 


tionate length of scaphocerite to the carapace 
as “‘five-sevenths” (=ca. 0.71) measured 
along the lateral margin, the ratio is 0.88- 
0.90 (0.89-0.94 in total length) in all the 
examined specimens. Dr. F. A. Chace, Jr. 
of the USNM informed me that a propor- 
tion ratio of the scaphocerite was actually 
0.73 in the illustrated specimen from the 
type lot of H. moseri. It is suggested that the 
ratio varies considerably from 0.73 to 0.94 
in H. moseri. Butler (1980) gave the pro- 
portionate length of rostrum to the carapace 
as 0.8—1.0, and that of dactylus to the prop- 
odus as 0.24 in the third pereopod and 0.23 
in the fourth pereopod. In the specimens 
examined here, the propotion ratio of the 
rostrum to the carapace is 1.0—1.2, and those 
of dactylus to the propodus of the pereopods 
0.19-0.20 in both the third and fourth per- 
eopods. These differences are so identical or 
so slight that they could be within the range 
of variation of the species. Further, Butler 
(1980) indicated the presence of a ridge on 
the inner margin (=face?) of the ocular pe- 
duncle. In the specimens examined here, no 
ridge-like structure is apparent, but a swell- 
ing is observed on the inner face of the oc- 
ular peduncle. Butler’s term “ridge”? may 
indicate this swelling. 

Rathbun (1904) stated that “Specimens 
from the two southernmost localities show 
a tendency to differ from the types,”’ though 
she combined them then. The identity of 
the southern population still remains un- 
settled. Hart’s (1930) record of H. moseri 
from the intertidal zone of Vancouver Is- 
land still remains somewhat in doubt, since 
the color pattern of the living animals de- 
scribed by her differs noticeably from that 
described by Butler (1980:224) and that of 
the present specimen. It is likely that her 
specimens belong to the closely related spe- 
cies H. littoralis Butler, 1980, which is dis- 
tributed along the American coast from Ba- 
ranof Island, Alaska to Seattle, Washington, 
at depths of 0-18 m (Wicksten 1990). 

In the Western Pacific Ocean, H. moseri 


VOLUME 106, NUMBER 3 


is the only species belonging to the group 
having epipods only on the first pereopod 
in addition to the maxillipeds. 
Distribution. —H. moseri has been known 
from the Bering Sea to Destruction Island, 
Washington (Wicksten 1990) and has a very 
wide bathymetric range from the intertidal 
zone to a depth of 1100 m (Butler 1980). 
The present specimen represents the first 
record of this species from the Western Pa- 
cific Ocean, as well as from Japanese waters. 


Acknowledgments 


I thank Drs. F. A. Chace, Jr., B. Kensley, 
and R. Lemaitre of the Smithsonian Insti- 
tution, K.-I. Hayashi of Shimonoseki Uni- 
versity of Fisheries, M. K. Wicksten of Tex- 
as A & M University, K. Amaoka of 
Hokkaido University, and two anonymous 
reviewers for thoughtfully reading the ear- 
lier draft of this manuscript and for their 
many valuable comments. I also thank Dr. 
Y. Sakurai and O. Yamamura of Hokkaido 
University for providing part of the mate- 
rial, Dr. Kensley and Ms. J. Clark of the 
Smithsonian Institution for arranging the 
loan of comparative material from Ameri- 
can waters, and Dr. Chace, Jr. for re-ex- 
amination of illustrated specimen from the 
type lot of H. moseri. The cooperation and 
assistance given by the staff on board the 
R/V Tanshu-Maru and T/S Oshoro-Maru 
are also acknowledged. 


Literature Cited 


Birshtein, Y. A., & L. G. Vinogradov. 1953. Novie 
dannie po faune desyatinogikh rakiibraznikh 
(Decapoda) Beringova Morya. [New data on the 
decapod crustacean fauna in the Bering Sea.]— 
Zoologicheskii Zhurnal 32(2):215-228. 

, & N. A. Zarenkov. 1970. O donnikh desy- 
atinogikh rokoobraznikh (Crustacea, Decapo- 
da) roiona Kurile-Kamchatskogo zeroba. [Bot- 
tom decapods (Crustacea, Decapoda) of the 
Kurile-Kamchatka trench area.].— Trudy Insti- 
tuta Okeanologii 86, Akademiya Nauk SSSR: 
420-426. 


553 


Butler, T. H. 1980. Shrimps of the Pacific coast of 
Canada.—Canadian Bulletin of Fishery and 
Aquatic Science 202:1—280. 

Criales, M.M. 1992. Redescription of the hippolytid 
shrimp Trachycaris rugosa (Bate) (Crustacea: 
Decapoda: Caridea) from the Western Atlantic, 
with notes on sexual dimorphism.—Proceed- 
ings of the Biological Society of Washington 105: 
562-570. 

Greve, L. 1963. The genera Spirontocaris, Lebbeus, 
Eualus and Thoralus in Norwegian waters (Crust. 
Dec.).—Sarsia 11:29-42. 

Hart, J. F. L. 1930. Some decapods from the south- 
eastern shores of Vancouver Island.— Canadian 
Field Naturalist 44(5):101-109. 

Hayashi, K. 1979. Studies on hippolytid shrimps from 

Japan— VII. The genus Heptacarpus Holmes. — 

The Journal of the Shimonoseki University of 

Fisheries 28:1 1-32. 

. 1992. Prawns, shrimps and lobsters from Ja- 

pan (64). Family Hippolytidae-Genera Alope & 

Heptacarpus.— Aquabiology 79:108-112. (in 

Japanese) 

Holthius, L. B. 1947. The Decapoda of the Siboga 
Expedition Part IX. The Hippolytidae and 
Rhynchocinetidae collected by the Siboga and 
Snellius expeditions with remarks on other spe- 
cies. —Siboga Expeditie 39(a8):1—100. 

Makarov, V. V. 1941. Fauna decapoda Beringova 1 
Chukotskogo Morei. [The decapod Crustacea of 
the Bering and Chukchi Seas.].—Issledovaniya 
Darinevoctochnikh Morei SSSR 1:111-163. 

Rathbun, M. J. 1899. List of Crustacea known to 

occur on and near Pribilof Islands. Pp. 555—557 

in D. S. Jordan, ed., The fur seals and fur seal 

islands of the north Pacific Ocean, Part 3. U.S. 

Treasury document 2017, Washington, D.C. 

1902. Descriptions of new decapod crusta- 
ceans from the west coast of North America. — 

Proceedings of the United States National Mu- 

seum 24:885-—905. 

. 1904. Decapod crustaceans of the northwest 

coast of North America. — Harriman Alaska Ex- 

pedition, Series 10:1-—210. 

Wicksten, M. K. 1990. Key to the hippolytid shrimp 
of the Eastern Pacific Ocean.— Fishery Bulletin 
U.S., 88:587-598. 


Laboratory of Marine Zoology, Faculty 
of Fisheries, Hokkaido University, Minato- 
machi 3-1-1, Hakodate, Hokkaido 041, Ja- 
pan; present address, Natural History Mu- 
seum and Institute, Chiba, 955-2 Aoba-cho, 
Chuo-ku, Chiba 260, Japan. 


PROC. BIOL. SOC. WASH. 
106(3), 1993, pp. 554-565 


TWO NEW HERMIT CRABS 
(CRUSTACEA: DECAPODA: PAGURIDAE) 
FROM THE CARIBBEAN SEA 


Rafael Lemaitre and Néstor H. Campos 


Abstract. —Two new species of hermit crabs are described from the southern 
Caribbean, Pylopagurus pattiae, living in empty polychaete tubes, and Xylo- 
pagurus tayrona, living in pieces of wood. Xylopagurus tayrona is the second 
known species of Xylopagurus A. Milne Edwards, 1880, in the western Atlantic, 
and the third of the genus. Both species are fully illustrated. 


During studies of the marine environ- 
ments of the Caribbean coast of Colombia, 
several interesting hermit crab specimens 
were collected in the vicinity of Santa Mar- 
ta. Thirty-seven of these specimens were 
found living in empty polychaete tubes, and 
represent an undescribed species of the ge- 
nus Pylopagurus A. Milne Edwards & Bou- 
vier, 1891 (sensu McLaughlin 1981). An- 
other specimen, a large ovigerous female 
found living in a hollow piece of wood, 
clearly belonged in the genus Xylopagurus 
A. Milne Edwards, 1880, but differed from 
the only known western Atlantic species of 
this genus, X. rectus A. Milne Edwards, 
1880, in several significant characters. In 
order to ascertain the identity of our spec- 
imen we compared it with the type material 
of A. Milne Edwards’ taxon deposited at the 
Museum of Comparative Zoology, Harvard 
University (MCZ 4097, 4098), and con- 
cluded that the Santa Marta specimen rep- 
resented an undescribed species of Xylo- 
pagurus. Subsequently, while examining 
supplemental, unreported material identi- 
fied as X. rectus from the University of Mi- 
ami collections, we discovered that in ad- 
dition to X. rectus, the material contained 
numerous specimens conspecific with our 
Santa Marta specimen. The two new spe- 
cies, named P. pattiae, and X. tayrona, are 
described and illustrated. 

The material used for this study remains 


deposited in: Instituto de Ciencias Natur- 
ales, Museo de Historia Nacional, Univer- 
sidad Nacional de Colombia (ICN-MHN- 
CR); Instituto de Investigaciones Marinas 
de Punta de Betin, Santa Marta (INVE- 
MAR-CRU); Rosenstiel School of Marine 
and Atmospheric Sciences, University of 
Miami (UMML); and National Museum of 
Natural History, Smithsonian Institution, 
Washington, D.C. (USNM). The following 
abbreviations or symbols are used: O, M/V 
Oregon; P, R/V John Elliott Pillsbury; SL, 
shield length, measured from the tip of the 
rostrum to the midpoint of the posterior 
margin of the shield; and 2°, ovigerous fe- 
males. 


Pylopagurus pattiae, new species 
Figs. 1, 2 


Holotype.—é (SL 2.0 mm), Bahia de 
Chengue, Parque Nacional Natural Tay- 
rona, north of Santa Marta, Colombia, 
dredged in 20-40 m, 3 Dec 1988, coll. N. 
H. Campos, USNM 259412. 

Paratypes. —Parque Nacional Natural 
Tayrona, north of Santa Marta, Colombia: 
2 6 (SL 1.7, 1.8 mm), Bahia de Chengue, 
dredged in 20-40 m, 3 Dec 1988, coll. N. 
H. Campos, USNM 251896.—2 4 (SL 1.5, 
1.7 mm), 2 9? (SL 1.7, 1.8 mm), Bahia de 
Cinto, dredged in 20-40 m, 30 Nov 1989, 
coll. N. H. Campos, USNM 251897.—16 6 


VOLUME 106, NUMBER 3 


(SL 1.1-2.0 mm), 4 2 (SL 1.5—1.9 mm), 10 
92 (SL 1.6-1.9 mm), Bahia de Concha, 
dredged in 40-60 m, 3 Dec 1988, coll. N. 
H. Campos, INVEMAR-CRU-1202, ICN- 
MHN. 

Description. —Shield (Fig. 1a) distinctly 
longer than broad; dorsal surface glabrous, 
evenly convex, with few scattered short se- 
tae; anterior margin between rostrum and 
lateral projections concave. Rostrum tri- 
angular, acute or bluntly rounded. Lateral 
projections broadly triangular, rounded. 
Anterolateral margin sloping. 

Ocular peduncles (Fig. 1a, b) stout, com- 
bined length of peduncle and cornea more 
than half length of shield. Cornea moder- 
ately dilated, pigmented. Dorsal surface na- 
ked or with few scattered short setae. Ocular 
acicles triangular, terminating acutely and 
with small submarginal spine; separated ba- 
sally by slightly more than basal width of 1 
acicle. 

Antennular peduncle slightly overreach- 
ing distal margin of cornea; segments naked 
or with few scattered short setae. Basal seg- 
ment with small spine on lateral face dis- 
tally. Dorsal flagellum long, about same 
length as that of ultimate and penultimate 
segments combined. 

Antennal peduncle (Fig. 1a, c) slightly ex- 
ceeding distal margin of cornea, supernu- 
merary segment present. Fifth segment slen- 
der, with row of setae on mesial margin. 
Fourth segment with setae on distomesial 
margin. Third segment with small spine 
(usually curved dorsally) on ventrodistal an- 
gle. Second segment with dorsolateral distal 
angle produced and terminating in strong 
spine, bearing few setae on lateral face dis- 
tally; dorsomesial distal angle with small 
spine. First segment with spine at ventrodis- 
tal angle. Acicle exceeding distal margin of 
fourth segment, nearly straight, slender, and 
terminating in strong spine, bearing few se- 
tae distally. Flagellum short, reaching to 
about distal margin of carpus of right che- 
liped, articles usually with pair of short se- 
tae, equal or less than 1 article in length. 


555 


Sternite of third maxillipeds with small 
spine on each side of midline. 

Right cheliped (Fig. 1d, e) with chela about 
twice as long as broad; fingers dorsoven- 
trally flattened, terminating in small cor- 
neous claw, cutting edges formed by un- 
equal calcareous teeth. Chela with ovate 
region on dorsal surface forming an oper- 
culum delimited by raised ridge on palm, 
lateral margin of fixed finger, and mesial 
margin of dactyl. Operculum with surface 
flat, naked, margins finely crenulate. Palm 
with scattered small tubercles and short se- 
tae on posterior portion of dorsal face and 
lateral and mesial faces; ventral face smooth, 
naked, or at most with scattered minute tu- 
bercles and short setae. Carpus longer than 
broad, distal margin often with small spine 
mesially; dorsal margin with row of small 
tubercles or protuberances bearing short se- 
tae; lateral and mesial faces with scattered 
small tubercles and setae; ventral face 
smooth or with scattered small tubercles and 
short setae. Merus more than 2 times as long 
as broad, triangular in crossection, distal 
margin often with small spine laterally; dor- 
sal margin with few minute protuberances 
bearing short setae; ventral face smooth or 
with scattered small tubercles and short se- 
tae. Ischium with row of long setae on ven- 
tromesial margin. 

Left cheliped (Fig. 1/) reaching to about 
distomesial angle of palm of extended right 
chela. Fingers with tufts of setae on distal 
half of dorsal and ventral faces; cutting edge 
of dactyl with distal row of small, fused, 
corneous spinules; cutting edge of fixed fin- 
ger formed of small calcareous teeth. Palm 
longer than broad, broadest between dis- 
tomesial angle and lateral margin; dorso- 
lateral margin with finely dentate, low ridge 
on distal 74; dorsomesial margin with sev- 
eral minute tubercles; dorsal surface also 
with several minute tubercles proximally; 
ventral face smooth, with scattered short 
setae. Carpus with 4 blunt to sharp spines 
on dorsodistal margin; dorsal surface with 
row of setae on each side of midline; mesial, 


556 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


hs ey Bay 


Fig. 1. Pylopagurus pattiae, new species. Holotype, USNM 259412: a, shield and cephalic appendages; 5b, 
right eye, lateral view; c, left antennal peduncle, lateral view; d, right cheliped, dorsal view; e, right chela, lateral 
view; f, left chela, dorsal view; g, second left pereopod, lateral view; h, third left pereopod, lateral view. Scales 
equal 1 mm (a, b, d-h), and 5 mm (c). (Stippled areas in a, b, and ovals shown on chela in d, indicate color 
pattern.) 


VOLUME 106, NUMBER 3 


lateral and ventral faces smooth, or at most 
with scattered setae. Merus triangular in 
crossection, with row of bristles on dorso- 
distal margin; dorsal margin with row of 
setae; mesial, lateral and ventral faces 
smooth, or at most with scattered setae. Is- 
chium with row of long setae on ventrome- 
sial margin. 

Ambulatory legs (second and third pereo- 
pods; Fig. lg, /) similar from right to left, 
usually not exceeding extended right che- 
liped, or at most overreaching right cheliped 
by length of claws of dactyls; mesial and 
lateral faces of segments naked. Dactyls 
about same length as propodi, terminating 
in long, slender corneous claw; dorsal mar- 
gin with row of long setae and bristles; ven- 
tral margin armed with 5-8 long, slender 
corneous spines (including spine at base of 
claw). Propodi with row of setae on dorsal 
margin, and dorsodistal row of bristles; ven- 
tral margin with row of long, slender cor- 
neous spines and long setae, and row of bris- 
tles on ventrodistal angle. Carpi with small 
dorsodistal spine, dorsal row of setae and 
few bristles; ventral margin with 1 or 2 long 
bristles. Meri and ischia with row of setae 
and some bristles on dorsal and ventral 
margins. 

Sternite of third pereopods (Fig. 2b) with 
anterior lobe subovate, setose. Sternite of 
fourth pereopods with cluster of capsulate 
setae. Sternite of fifth pereopods with pair 
of capsulate setae. 

Fourth pereopods with inconspicuous 
preungual process at base of claw. 

Abdomen straight (Fig. 2a), tightly fitting 
into tube housing. Uropods and telson sym- 
metrical. Telson (Fig. 2c) with lateral mar- 
gins of posterior lobes evenly rounded (of- 
ten nearly sub-semicircular), each ending 
posteriorly in strong spine; posterior lobes 
separated by shallow median cleft, with ter- 
minal margins oblique and each armed with 
2-3 unequal sharp or blunt spines. 

Females with 15-40 eggs about 0.4 mm 
in maximum width. 

Color (Fig. la, b, d).—In life, general col- 


SST 


oration orange. Ocular peduncles with light 
orange band at about midline. Cutting edges 
of fingers orange. Operculum of right chela 
with three longitudinally ovate white spots 
with orange outlines, as follows: two me- 
dially on palm and base of fixed finger re- 
spectively, and one medially on base of dac- 
tyl and extending slightly on palm. Capsulate 
setae on sternite of fourth pereopods yellow. 

Habitat. — Found living in corneous, semi- 
transparent tubes (Fig. 2a) built by poly- 
chaetes of the genus Hyalinoecia, family 
Onyphidae. The tubes inhabited by the 
specimens have one opening slightly larger 
than the other, and range in length from 25 
to 28.5 mm. The tube openings range in 
diameter from 1.6 to 1.9 mm (larger open- 
ings), and from 1.1 to 1.3 mm (smaller 
openings). 

Distribution. —So far known only from the 
Bays of Chengue and Cinto, north of Santa 
Marta, on the Caribbean coast of Colombia; 
20-60 m. 

Etymology.—The specific name is de- 
rived from “Pat,” a friendly version of Dr. 
Patsy A. McLaughlin’s first name, and is 
given in recognition of her many contri- 
butions to hermit crab systematics in gen- 
eral and the Pylopagurus-group of species 
in particular. 

Remarks. —This species most closely re- 
sembles Pylopagurus discoidalis (A. Milne 
Edwards, 1880), a species distributed from 
the Lesser Antilles to North Carolina (Pro- 
venzano 1963). A. Milne Edwards & Bou- 
vier (1893:79) and recently Williams (1984: 
227), have suggested that the large chela of 
P. discoidalis is initially elongate, but be- 
comes more discoidal with age. Pylopagurus 
pattiae differs from P. discoidalis most sig- 
nificantly in the shape and armature of the 
dactyls, propodi and carpi of the ambula- 
tory legs. In P. pattiae these segments all are 
moderately long and narrow (lateral view), 
whereas they all are short and broad in P. 
discoidalis. Although the ventral margins of 
the dactyls each carry a row of corneous 
spines in both species, the dorsal margins 


558 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


a 
b,c 


Fig. 2. Pylopagurus pattiae, new species. a, ovigerous female (SL 1.8 mm) shown inside polychaete tube 
housing, USNM 251897. b, c, holotype, USNM 259412: b, sternum; c, telson, dorsal view. Scales equal 5 mm 
(a), and 1 mm (8, c). 


VOLUME 106, NUMBER 3 


of the propodi and carpi of P. discoidalis 
are each armed with one or two rows of 
spinules or small spines. With the exception 
of the dorsodistal carpal spines, these sur- 
faces are unarmed in P. pattiae. The ocular 
peduncles of P. discoidalis are nearly twice 
as broad at the base of the cornea as the 
peduncular base, and the corneae are 
strongly dilated. In P. pattiae the ocular pe- 
duncles are nearly uniform in breadth over 
their entire length and the corneae are only 
slightly dilated. The general shape of the 
telson in both species is similar; however, 
the new species has only two or three spines 
on the terminal margins, whereas P. dis- 
coidalis has several more. The color pat- 
terns of the right cheliped in particular, also 
differ. This chela in P. discoidalis is basically 
white with a band of red or reddish-orange 
on the dactyl and fixed finger, and irregular 
bands on the palm. The extent of pigmen- 
tation on the palm is variable. In the new 
species, the base color of the chela is orange 
with ovate patches of white rimmed with 
orange on the palm, fixed finger and dactyl. 
Pylopagurus discoidalis most commonly oc- 
cupies shells of Dentalium spp., whereas P. 
pattiae has only been found living in cor- 
neous, semi-transparent tubes built by poly- 
chaetes of the family Onyphidae (Hyaii- 
noecia sp.). 


Xylopagurus tayrona, new species 
Figs. 3-5 


Holotype. —22 (SL 11.9 mm), Punta 
Gayra, Bahia de Santa Marta, Colombia, 
118 m, from trap and living inside of a bam- 
boo-like piece of wood, Feb 1984, coll. Oc- 
tavio Galvis, USNM 251902. 

Paratypes.—1 2 (SL 15.5 mm), 12°27'N, 
69°51'W, 0-5648, 229 m, 2 Oct 1965, 
UMML 32:4996.—26 6 (SL 3.0-12.3 mm), 
15 2? (SL 2.5-5.8 mm), 1 92 (SL 9.0 mm), 
11°22'N, 73°44'W, P-783, 174 m, 31 Jul 
1968, USNM 265141, 265142.—1 6 (SL 
14.4 mm), 11°6.7'N, 74°30'W, O-4844, 183 
m, 17 May 1964, UMML 32:4995.—1 ?@(SL 


559 


13.0 mm), 11°5.2’N, 64°45'W, P-722, 91 m, 
21 Jul 1968, USNM 265143. 

Description. —Shield (Fig. 3a) distinctly 
longer (about 1.4 times) than wide, evenly 
convex. Dorsal surface glabrous except for 
few scattered minute pits and tufts of setae; 
with short, low dorsal ridge posterior to ros- 
trum, and small, shallow depressed region 
posterior to each lateral projection. Ros- 
trum subtriangular, slightly exceeding lat- 
eral projections. Anterior margin nearly 
straight. Lateral projections broadly sub- 
triangular. Anterolateral margins sloping, 
with distinct indentation on distolateral an- 
gle. Accessory calcified portions of carapace 
partially fused to shield (see Fig. 3a). Bran- 
chiostegite with narrow calcified portion ad- 
jacent and parallel to /inea anomurica; an- 
terior margin rounded, setose. Posterior 
carapace with dorsal surface well calcified 
medially. 

Ocular peduncles stout, combined length 
of peduncle and cornea about half length of 
shield; cornea dilated. Ocular acicles (Fig. 
3a, b) developed as broad ovate plate armed 
marginally with 13-14 small spines (10-12 
in small specimens, SL <= 3.0 mm); sepa- 
rated basally by about basal width of 1 aci- 
cle. 

Third maxilliped with ischium (Fig. 3e) 
having well developed crista dentata and 
strong accessory tooth. 

Antennular peduncle reaching to about 
distal margin of cornea. Ultimate segment 
about 2.4 to 2.8 times as long as broad. 
Ultimate and penultimate segments sub- 
equal in length, with scattered setae. Basal 
segment with ventrodistal spine, and setose 
lobe on lateral face proximally. Dorsal fla- 
gellum long, about twice as long as ultimate 
segment; ventral flagellum usually with 12 
articles. 

Antennal peduncle (Fig. 3a, c, d) not ex- 
ceeding distal margin of cornea, supernu- 
merary segment present. Fifth segment with 
long setae on lateral margin. Fourth seg- 
ment with scattered setae dorsally, and short 
transverse row of long setae on ventral face 


560 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Xylopagurus tayrona, new species. Holotype, USNM 251902: a, shield (arrows indicate accessory 
calcified portions of carapace) and cephalic appendages; b, ocular acicles, dorsal view; c, right antennal peduncle, 
lateral view; d, right antennal acicle, dorsal view; e, basis, ischium, and merus of left third maxilliped, external 
view; f, right cheliped (tip of fixed finger reconstructed); g, merus and carpus of same, mesial view; h, left cheliped. 
Scales equal 5 mm (a, g, h, f), 2 mm (b), 3 mm (¢, e), and 2 mm (d). 


VOLUME 106, NUMBER 3 


distally. Third segment with blunt ventro- 
distal angle. Second segment with distola- 
teral angle developed as very broad subrec- 
tangular projection armed with 12-15 small 
spines on distal margin (8 spines in small 
specimens SL <= 3.0 mm); mesial margin 
with dorsodistal angle armed with 7-8 small 
spines (4 in small specimens SL = 3.0 mm). 
First segment unarmed, with tuft of setae 
on lateral face. Acicles short, broad, straight, 
terminating in fan-like distal margin armed 
with 14—16 small spines (elongate and with 
4 small spines in small specimens SL < 3.0 
mm). Flagellum reaching to about mid- 
length of right chela, densely setose, each 
article with up to 10 short and long setae. 
Chelipeds markedly dissimilar. Right 
cheliped (Fig. 3/) strong, massive. Fingers 
each terminating in blunt calcareous claw. 
Dactyl with small tubercles on dorsal sur- 
face, and row of tufts of setae parallel to 
cutting edge; mesial margin with tufts of 
setae and small tubercles; cutting edge prox- 
imally with 1 massive, molar-like calcare- 
ous tooth about half as long as dactyl, and 
distally with 8 small calcareous teeth; ven- 
tromesial margin with row of tubercles in- 
terspersed with tufts of setae; ventral surface 
smooth except for scattered tufts of setae. 
Fixed finger slightly overreaching dactyl; 
cutting edge proximally with 2 large molar- 
like calcareous teeth, and distally with 4 
small teeth; ventromesial margin with row 
of tubercles interspersed with tufts of setae. 
Palm about 1.2 to 1.4 times as long as wide; 
spiniform dorsomesial projection about %4 
as long as dactyl, with tubercles and spines 
on dorsal and mesial faces; dorsal surface 
armed on distal 7s with numerous tubercles 
and spines (largest on medial region; tuber- 
cles and spines less numerous in small spec- 
imens SL < 5.0 mm), smooth and with scat- 
tered tufts of setae proximally; dorsomesial 
and dorsolateral margins each delimited by 
irregular row of small tubercles or spines; 
mesial face with numerous small tubercles 
and spines on distal '4; lateral face convex, 
with numerous tufts of setae; ventral face 


561 


smooth except for scattered small tubercles 
and tufts of setae. Carpus about as long as 
wide, triangular in cross-section; dorsal 
margin distinctly delimited (usually elevat- 
ed), projecting slightly anteriorly over palm, 
and armed with series of transverse rows of 
small tubercles usually bearing setae at bas- 
es; dorsolateral and dorsomesial faces with 
transverse rows of long bristles; ventral face 
smooth, with scattered tufts of setae. Merus 
about as long as wide, triangular in cross 
section, unarmed, with scattered setae. 

Left cheliped (Fig. 3h) slender, short, 
reaching to about distal margin of carpus of 
right cheliped. Fingers longer than palm, 
terminating in corneous claws, opposing 
faces of fingers concave; cutting edges each 
with row of minute, fused corneous teeth; 
dorsal and ventral surfaces with numerous 
tufts of setae. Palm smooth except for scat- 
tered setae. Carpus and merus subequal in 
length, unarmed, with scattered short setae. 

Ambulatory legs (Fig. 4a—d) similar from 
right to left. Dactyls each terminating in cor- 
neous claw, with cluster of short bristles near 
base of claw on lateral and mesial faces: 
ventral margin nearly straight, usually armed 
with 30 or more short and long corneous 
spines (small specimens SL <= 3.0 mm with 
as few as 15 spinules); with row of setae 
dorsally, and 3-4 short oblique rows of setae 
dorsolaterally and dorsomesially. Ischia, 
meri, carpi, and propodi unarmed except 
for row of 4—6 minute spinules on ventral 
margin of propodus; segments with setae or 
tufts of setae on dorsal and ventral margins; 
lateral and mesial faces of segments with 
scattered setae or with longitudinal rows of 
tufts of short setae. Sternite of third pereo- 
pods (Fig. 4g) narrow, anterior lobe naked, 
flat, sloping down anteriorly. 

Fourth pereopod (Fig. 4e) with dactyl 
subtriangular, terminating in corneous claw 
curved laterad, with tufts of setae on dor- 
solateral and ventrolateral margins, and 
longitudinal row of strong corneous spi- 
nules directed slightly laterad. Propodus 
strongly produced ventrally; propodal rasp 


562 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


‘ +++ a, b,g 
je 
a SS 


Fig. 4. Xylopagurus tayrona, new species. Holotype, USNM 251902: a, second right pereopod, lateral view; 
b, third right pereopod, lateral view; c, dactyl of right third pereopod, lateral view; d, tip of dactyl of left third 
pereopod, lateral view; e, right fourth pereopod, lateral view; f right fifth pereopod, lateral view; g, sternum, 
ventral view. Scales equal 5 mm (a, b, g), 2 mm (c), 1 mm (d@), 2 mm (e¢, f). 


well developed, occupying subovate area inform of grid. Merus and carpus unarmed, 
covering nearly half of lateral face of seg- with scattered setae. 

ment and formed of numerous minute scales Fifth pereopod (Fig. 4/) subchelate, with 
densely packed and arranged in straightrows propodus bearing long setae on dorsal and 


VOLUME 106, NUMBER 3 


ventral margin; rasp formed of conical or 
lanceolate scales. Merus and carpus un- 
armed, with scattered setae. 

Tergite of sixth abdominal somite strong- 
ly calcified, operculate (Fig. 5a), with raised 
crenulate margins bearing evenly spaced 
tufts of long setae (more separated on an- 
terior margin). Dorsal surface divided into 
anterior and posterior regions by broad 
transverse depression; anterior region oc- 
cupying about *%4 surface of tergite; posterior 
region directed slightly upwards (planes of 
surface of anterior and posterior regions 
forming obtuse angle). Anterior and pos- 
terior regions separated laterally by narrow 
notch. Anterior region nearly flat, covered 
with numerous small, shallow, rounded pits, 
and often scattered, irregularly-shaped, short 
grooves; occasionally with pair of small me- 
dian tubercles near center of operculum. 
Posterior region covered with numerous 
small, shallow, rounded pits; with median 
longitudinal furrow not reaching posterior 
margin of tergite, and usually with 1 or 2 
small tubercles on each side near postero- 
lateral angle; with broad, depressed, trian- 
gular region basally encompassing nearly 
entire posterior margin, and with blunt apex 
in advance of anteriormost point of median 
furrow. 

Telson (Fig. 55, c, d) not visible in dorsal 
view of abdomen, more than 2 times as 
broad as long. Lateral margins with tuft of 
short setae near distolateral angle. Dorsal 
surface with 3 depressions (2 deep lateral, 
and 1 shallow median). Posterior margin 
divided into very broadly rounded lobes by 
shallow (sometimes inconspicuous) cleft, 
and armed with small, mostly blunt spines 
with corneous tips. 

Uropods (Fig. 5b) symmetrical. Protopod 
robust, dorsal face flattened; posterodorsal 
margin corneous, proximally with broad tri- 
angular and subrectangular teeth, and row 
of tufts of long setae; ventrodistal angle pro- 
duced into prominent blunt spine with row 
of setae laterally. Rasp of endopod and ex- 


563 


opod formed of strong conical and lanceo- 
late scales; endopod with small denticles on 
anterior margin; rasp of exopod occupying 
almost entire surface of lateral face, scales 
stronger and more spaced proximally. 

Male with paired first and second pleo- 
pods modified as gonopods; first pleopod 
1-segmented, curved anteriorly, with long 
dense setae distally; second pleopod 
3-segmented, distal segment directed ante- 
riorly and long setae distally. Female with 
biramous unpaired left second to fourth 
pleopods; eggs about 1.8 mm in diameter. 

Etymology.—The specific name is given 
in honor of the Tayrona indians, whose de- 
scendants still populate the coastal moun- 
tain range of the Sierra Nevada de Santa 
Marta. 

Distribution.—Southern Carribean Sea, 
from Venezuela and Colombia; 91 to 229 m. 

Remarks. —Xylopagurus tayrona can be 
separated most readily from the only other 
known western Atlantic species of the ge- 
nus, X. rectus, by the armature of the dactyls 
of the ambulatory legs, and the shape of the 
operculate tergite of the sixth abdominal so- 
mite. In X. tayrona the dactyls of the am- 
bulatory legs are armed with 30 or more 
corneous spines, whereas in X. rectus there 
are at most six spines. In X. tayrona, the 
tergite of the sixth abdominal somite is 
nearly flat on the anterior half, and the lat- 
eral margin of the anterior region is sepa- 
rated from that of the posterior region by a 
narrow notch. In contrast, the tergite of the 
sixth abdominal somite of X. rectus is very 
uneven on the anterior region, with irregular 
ridges, pits, and a distinct median longitu- 
dinal groove with raised margins; and the 
lateral margin of the anterior region is sep- 
arated from that of the posterior region by 
a broad rounded notch. 

Xylopagurus tayrona most closely resem- 
bles the eastern Pacific X. cancellarius Wal- 
ton, 1950. Similarities between the two spe- 
cies are evident in the massive right cheliped 
with a triangular carpus, the development 


564 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


/ 


i / wu 
\ 


e 7 


~ 


_ Sy LE 

“i 
b \ = Ly) 
—s 


fp 3 


Rips 5: 


NN Vw / 
ae Vil Bs 


ee wn S 
Bory NM \ \| NY vy IOS 


| WG 
aS LOD 
1 AB 


d 


Xylopagurus tayrona, new species. Holotype, USNM 251902: a, operculate tergite of sixth abdominal 


somite, dorsal view (figure by Molly K. Ryan); }, left uropod and margin of tergite of sixth abdominal somite, 
lateral view (left side of figure is posterior region); c, posterior portion of tergite of sixth abdominal somite and 
telson, ventral view; d, telson. Scales equal 5 mm (a), 3 mm (b), 5 mm (c), and 2 mm (d). 


of a fan-like terminal margin on the anten- 
nal acicles, the dense setation of the anten- 
nal flagella, and the armature of the ventral 
margin of the dactyls of the ambulatory legs. 
However, X. tayrona lacks the two pairs of 
prominent central, conical protuberances 
present on the tergite of the sixth abdominal 
somite of XY. cancellarius. 


Acknowledgments 


We wish to thank O. Galvis and other 
personnel from INVEMAR for helping to 
recover the specimen of Xylopagurus tay- 
rona. The help of M. M. Criales, J. Garcia- 
Gomez, N. Voss, and A. B. Johnston in 
arranging loans of specimens from MCZ and 


VOLUME 106, NUMBER 3 


UMM, is gratefully acknowledged. Molly 
K. Ryan prepared Fig. 5a. This study was 
possible, in part, thanks to a Short-Term 
Visitor travel grant awarded to one us (NHC) 
by the Office of Fellowships and Grants, 
Smithsonian Institution, during the sum- 
mer of 1992. Thanks are also extended to 
P. A. McLaughlin, for comments on an ear- 
lier version of the manuscript; and K. Fau- 
chald and J. J. A. Laverde, for the name of 
the polychaete. 


Literature Cited 


Milne Edwards, A. 1880. Reports on the results of 
dredging, under the supervision of Alexander 
Agassiz, in the Gulf of Mexico, and in the Ca- 
ribbean Sea, 1877,’78,’79, by the United States 
Coast Survey Steamer “Blake’’, Lieut.-Com- 
mander C. D. Sigsbee, U.S.N., and Commander 
J. R. Bartlett, U.S.N., Commanding. 8. Etudes 

préliminaires sur les crustacés.— Bulletin of the 

Museum of Comparative Zoology, Harvard 8(1): 

1-68. 

,& E. L. Bouvier. 1891. Observations généra- 
les sur les paguriens recueillis dans la mer des 
Antilles et le Golfe du Méxique, par le Blake et 
le Hassler, sous la direction de M. Alexandre 
Agassiz.— Bulletin de la Société Philomatique 
de Paris (8)3:102-110. 

——., & 1893. Reports of the results of 
dredging under the supervision of Alexander 
Agassiz, in the Gulf of Mexico (1877-78), in the 
Caribbean Sea (1878-79) and along the Atlantic 


565 


coast of the United States (1880), by the U. S. 
Coast Survey Steamer “Blake’’, Lieut.-Com S. 
D. Sigsbee, U.S.N., and Commander J. R. Bart- 
lett, U.S.N., Commanding. 33. Description des 
Crustacés de la Famille des Paguriens recueillis 
pendant l’expédition.— Memoirs of the Muse- 
um of Comparative Zoology at Harvard College 
14(3):1-172 + pls. 1-12. 

McLaughlin, P. A. 1981. Revision of Pylopagurus 
and Tomopagurus (Crustacea: Decapoda: Pa- 
guridae), with the descriptions of new genera 
and species: Part I. Ten new genera of the Pa- 
guridae and a redescription of Tomopagurus A. 
Milne Edwards and Bouvier.— Bulletin of Ma- 
rine Science 31(1):1-30. 

Provenzano, A. J., Jr. 1963. Pylopagurus discoidalis 
(A. Milne Edwards, 1880) (Decapoda, Ano- 
mura) found off North Carolina (U.S.A.), a 
northern record for the genus. — Crustaceana 5(3): 
239-240. 

Walton, B. C. 1950. Some new and rare Pacific pa- 
gurids.—Journal of the Washington Academy 
of Sciences 40(6):188-193. 

Williams, A. B. 1984. Shrimps, lobsters, and crabs 
of the Atlantic coast of the eastern United States, 
Maine to Florida. Smithsonian Institution Press, 
Washington, D.C., 550 pp. 


(RL) Department of Invertebrate Zoolo- 
gy, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 
20560, U.S.A.; (NHC) Instituto de Ciencias 
Naturales, Universidad Nacional de Co- 
lombia, INVEMAR, A. A. 1016, Santa 
Marta, Colombia. 


PROC. BIOL. SOC. WASH. 
106(3), 1993, pp. 566-581 


STOMATOPOD CRUSTACEA FROM TOBAGO, 
WEST INDIES 


Marilyn Schotte and Raymond B. Manning 


Abstract. — Eleven shore species of stomatopods are reported from localities 
around Tobago; only two species, Gonodactylus oerstedii Hansen and G. bredini 
Manning, had been recorded previously from the island. Gonodactylus carib- 
baeus and Nannosquilla tobagoensis are described as new. Lysiosquilla bimi- 
niensis Bigelow is referred to the new genus Bigelowina, and Lysiosquilla flori- 
densis Manning is removed from the synonymy of L. digueti Coutiére and is 
transferred to the new genus Alachosquilla; both of these species had been 


referred to Acanthosquilla. 


Until now, only two stomatopods, Gon- 
odactylus bredini Manning and G. oerstedii 
Hansen had been reported from Tobago 
(Manning 1969). A search of records in the 
stomatopod collection of the National Mu- 
seum of Natural History yielded no new or 
unpublished records from Tobago other than 
those mentioned by Manning (1969). These 
species, along with Pseudosquilla ciliata, re- 
ported below, are the most common and 
widespread shore stomatopods in the north- 
western Atlantic. 

During a survey in 1990 of invertebrates 
and fish in shore habitats off Tobago in the 
southeastern Caribbean Sea, nine species of 
stomatopod crustaceans were collected by 
one of us (MS) and other survey partici- 
pants; two of the species proved to be new. 
Nearly all were collected at rotenone poison 
stations in coral habitats in depths between 
11 and 13-28 meters. Subsequently, Rich- 
ard Heard, Gulf Coast Research Labora- 
tory, provided us with material of four spe- 
cies that he collected on Tobago in 1992. 
Two species that he collected were not rep- 
resented in our collections from 1990. Thus 
the collections reported here provide a five- 
fold increase in the number of species of 
stomatopods known from Tobago. The col- 
lection also supplied valuable information 
on habitat and color in life of the species. 


Since Manning’s (1969) monograph on 
stomatopod crustaceans from the western 
Atlantic, a number of papers have cited 
range extensions for some members of the 
group into the Gulf of Mexico and southern 
Florida, e.g., Camp (1973) and Gore & 
Becker (1975, 1976). Studies by Adkison et 
al. (1983), Adkison & Hopkins (1984), Camp 
(1971), Camp & Manning (1982, 1986), 
Fausto-Filho & Lemos de Castro (1973), 
Hernandez Aguilera & Hermoso Salazar 
(1988), Manning (1970, 1979), and Man- 
ning & Hart (1981), added 20 new species 
of stomatopods to the East American fauna. 
Only six of the 20 were discovered in the 
Caribbean Sea. GOmez & Ortiz (1985) re- 
ported that 18 species occurred in Cuban 
waters; five of these, e.g., Acanthosquilla 
biminiensis (Bigelow), Gonodactylus bredi- 
ni, G. oerstedii, Lysiosquilla glabriuscula 
(Lamarck), and Pseudosquilla ciliata (Fa- 
bricius), are reported here from Tobago. A 
total of 22 stomatopod species were re- 
ported from the Caribbean coast of Colom- 
bia by Werding & Miller (1990), nine of 
which were new records for that area. Mark- 
ham & Donath-Hernandez (1990) and 
Markham et al. (1990) recently recorded five 
species from Quintana Roo, Mexico; four 
of these five also occur at Tobago. 


VOLUME 106, NUMBER 3 


The 20 East American species described 
since 1969 are: 


Gonodactyloidea 


Gonodactylidae: 
Gonodactylus lightbourni Manning & 
Hart, 1981, Bermuda. 
Gonodactylus moraisi Fausto-Filho & Le- 
mos de Castro, 1973, Brazil. 
Gonodactylus petilus Manning, 1970, 
southwestern Caribbean Sea. 
Pseudosquillidae: 
Parsquilla boschii Manning, 1970, Argen- 
tina. 


Lysiosquilloidea 


Coronididae: 

Acoridon manningi Adkison, Heard, & 

Hopkins, 1983, Gulf of Mexico. 
Nannosquillidae: 

Mexisquilla horologii (Camp, 1971), Gulf 
of Mexico. 

Nannosquilla adkisoni Camp & Manning, 
1982, Gulf of Mexico. 

Nannosquilla baliops Camp & Manning, 
1982, Florida. 

Nannosquilla candidensis Hernandez 
Aguilera & Hermoso Salazar, 1988, 
Mexico. 

Nannosquilla carolinensis Manning, 1970, 
North Carolina. 

Nannosquilla dacostai Manning, 1970, 
Brazil. 

Nannosquilla disca Camp & Manning, 
1986, Gulf of Mexico. 

Nannosquilla heardi Camp & Manning, 
1982, Gulf of Mexico. 

Nannosquilla taguensis Camp & Man- 
ning, 1982, U.S. Virgin Islands. 

Nannosquilla vasquezi Manning, 1979, 
Panama. 

Nannosquilla virginalis Camp & Man- 
ning, 1986, U.S. Virgin Islands. 

Nannosquilla whitingi Camp & Manning, 
1982, Florida. 

Nannosquilla yucatanica Camp & Man- 
ning, 1986, Mexico. 


567 


Tetrasquillidae: 
Tectasquilla lutzae Adkison & Hopkins, 
1984, Gulf of Mexico. 


Squilloidea 


Squillidae: 
Squilla decimdentata Manning, 1970, 
southwestern Caribbean Sea. 


Abbreviations used below include: ft, feet; 
JTW, acronym for J. T. Williams’ field sta- 
tions; leg., collector; m, meters; mm, mil- 
limeters; sta, station; TL, total length, mea- 
sured on the midline; TOB, acronym for 
Schotte’s field stations. All measurements 
are in millimeters. 

In the species accounts below, “‘Habitat” 
refers to habitat on Tobago. In the sections 
on ‘Material,’ the number in parentheses 
after the number of specimens is total length. 

All of the material reported here is de- 
posited in the collections of the National 
Museum of Natural History, Smithsonian 
Institution, Washington, D.C. (USNM). 


Systematic Account 


Order Stomatopoda 
Superfamily Gonodactyloidea 
Family Gonodactylidae 
Genus Gonodactylus Berthold, 1827 
Gonodactylus bredini Manning, 1969 


Gonodactylus bredini Dingle, 1969:108 
[January]. 

Gonodactylus bredini Manning, 1969:315, 
figs. 87, 88 [April]. 


Material.—Sta TOB-16, east of North 
Paint,.21.mz 1.625). 

Color in life.—Body color yellow-pink; 
posterior margins of carapace and all so- 
mites with small light brown patches dot- 
ting entire margin; carinae on telson with 
similar patches; medial brown patch of pig- 
ment at articulation of rostral plate and car- 
apace; meral spot white, flanked by two 
small patches of dark brown; base of dactyl 
bright purple-pink; proximal end of prop- 


568 


odus with faint patches of brown; posterior 
margin of carpus olive-green. 

Size.—Male (1), TL 25 mm. Manning 
(1969) studied specimens as large as 75 mm. 

Habitat.— Rock wall to rubble flat at 21 m. 

Remarks.—The specific name bredini was 
published twice in 1969. Under the present 
(third) edition of the International Code of 
Zoological Nomenclature, the specific name 
bredini Dingle, published three months be- 
fore bredini Manning, has to be considered 
a nomen nudum. Dingle’s use of the name, 
in an account of the behavior of larvae, is 
not accompanied by a description or words 
that distinguish his taxon from others and 
lacks a bibliographic reference to such a 
statement. 

Gonodactylus bredini is one of the two 
most common shore species of Gonodac- 
tylus in the Caribbean, the other being G. 
oerstedii. In the field the two may be distin- 
guished at once by the color of the meral 
spot in live specimens, white in G. bredini, 
reddish to purple in G. oerstedii. 

One male was recorded from Tobago by 
Manning (1969); it lacked data on habitat 
and locality. 

Distribution. —Bermuda, North Carolina, 
and Florida to Curagao; shore to 55 m. Camp 
(1973) reported specimens from the Gulf of 
Mexico in depths to 73 m. 


Gonodactylus caribbaeus, new species 
Fig. 1 


Material. —Sta TOB-12, off Little Tobago 
Island, 18 m: 1 6(30.5), 1 29(31.5), paratypes 
USNM 252678.—Sta TOB-13, cove near 
Speyside, 11 m: 2 66 (27.5—33), paratypes, 
USNM 252679.—Sta TOB-21, Buccoo 
Reef, 14 m: 1 ¢ (31), holotype, USNM 
252677.—Sta TOB-40, St. Giles Island, 6- 
11 m: 1 6 (33), paratype, USNM 252680. 

Diagnosis. — Rostral plate as long as broad, 
obtusely rounded laterally, anterior margins 
sloping to slender median spine. Ocular 
scales small, separate, erect. Anterior four 
abdominal somites unarmed posterolater- 
ally, fifth with sharp posterolateral spines; 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


abdomen lacking distinct black chromato- 
phores. Sixth abdominal somite with 6 ca- 
rinae, each armed posteriorly, submedians 
and intermediates inflated. Abdominal 
width-carapace length indices 790-880 
(mean 830). Telson slightly longer than 
broad, of oerstedii-type, with dorsal tuber- 
cles on carinae. Carinae of telson well-de- 
fined except for low accessory intermedi- 
ates, not strongly inflated in males. Median 
carina broad, terminating in single tubercle. 
Accessory median carinae broad, with 1-3 
tubercles dorsally. Knob rounded, with 2- 
4 tubercles. Anterior submedian carinae 
with 2 dorsal tubercles. Submedian margin- 
al teeth slender, with 3-6 dorsal tubercles, 
inner margins lined with denticles, movable 
apices present. Intermediate marginal teeth 
sharp, with 0-1 dorsal tubercles. Interme- 
diate denticles sharp, inner with 1-2 dorsal 
tubercles. Lateral carinae ending in blunt 
tooth. Uropodal exopod with 11-12 mov- 
able spines, distalmost largest. 

Color in life. —Body pale yellow with thin 
blue bands at posterior margin of carapace 
and all somites; carinae on sixth abdominal 
somite and telson tinged with blue, green 
on median and submedian carinae; dactyl 
lavender-pink, propodus blue with olive- 
green and orange distally; carpus blue, ol- 
ive-green distally; merus pale yellow, olive 
with dark blue tinges distally; meral spot 
white, flanked by two dark brown spots. 

Size.— Males (5), TL 27.5-33 mm; fe- 
male (1), TL 31.5 mm. Other measurements 
of male holotype, TL 31 mm: rostral plate 
length 2.3, width 2.1; carapace length 7.3; 
fifth abdominal somite width 6.2; telson 
length 5.3, width 4.9. 

Habitat. —On Tobago, sublittoral on 
rough substrates, in depths between 6-11 m 
and 35 m, including at 14, 18, and 21 m, 
on bottoms with coral heads; covered with 
coral and sea whips; rock wall to rubble flat; 
sloping fore reef; and rock, rubble, and live 
and dead coral. 

Remarks. —Gonodactylus caribbaeus re- 
sembles G. curacaoensis and differs from all 
other American species in having the fifth 


VOLUME 106, NUMBER 3 


569 


Fig. 1. 


Gonodactylus caribbaeus, new species, male holotype, TL 31 mm. A, Sixth abdominal somite, telson, 


and right uropod; B, Rostral plate; C, Fourth and fifth abdominal somites, lateral view of left side; D, Submedian 
and intermediate teeth of telson, right side; E, Posterior two abdominal somites, telson, and left uropod, lateral 


view. 


abdominal somite armed with a spine at its 
posterolateral angle. It resembles both G. 
spinulosus Schmitt, 1924 and G. minutus 
Manning, 1969 in having the dorsal carinae 
of the telson ornamented with tubercles, but 
both of these species have rounded, un- 
armed posterolateral angles on the fifth ab- 
dominal somite. This new species can be 


distinguished from G. spinulosus in the field 
by its lack of the dark patches of pigment 
on the sixth thoracic and first abdominal 
somites that are characteristic of the latter 
species. 

The telson of the specimen from sta TOB- 
40 is damaged; it lacks the left marginal 
tooth. 


570 


Etymology. —Named for the general area 
in which it was found, the Caribbean Sea. 
Distribution.—Known only from Toba- 


go. 


Gonodactylus curacaoensis Schmitt, 1924 


Gonodactylus oerstedii var. curacaoensis 
Schmitt, 1924a:80, pl. 8, fig. 6. 

Gonodactylus curacaoensis. —Manning, 
1969:307, fig. 85. 


Material.—Sta TOB-6, off Speyside, 11 
m: 1 2 (62).—Sta TOB-12, off Little Tobago 
Island, 18 m: 1 6 (27).—Sta TOB-16, east 
of North Point, 21 m: 1 6 (42).—Sta TOB- 
17, northeast of Charlotteville, 12 m: 2 4é 
(35, 57).—Sta TOB-21, Buccoo Reef, 14 m: 
1 6(21), 2 92 (24, 29).—Sta TOB-38, Buccoo 
Reef, 14 m: 1 6 (39).—Sta TOB-44, The 
Sisters, 18-26 m: 3 92 (21, 28, 50). 

Color in life. —Body color variable: pale 
yellow, pinkish or pale to dark blue-green, 
especially in larger males; often with thin 
band of blue-green at posterior margin of 
all somites. Dactyl and distal part of prop- 
odus pale to bright purple-pink; rest of 
propodus, carpus, and articulation of merus 
variously outlined in blue; meral spot pale 
or white (sometimes with faint orange blush 
in large males) and flanked by two dark pur- 
ple-brown spots. 

Size. —Males (6), TL 21-57 mm; females 
(6), TL 21-62 mm. Manning (1969) re- 
ported males as large as 59 mm and females 
as large as 72 mm. 

Habitat.—Taken in a variety of sublit- 
toral coralline habitats in depths between 
11 and 18—26 m: in coral heads, 18 m; from 
rock wall to rubble flat, crevices in 21 m; 
coral-encrusted rocks and ledges, 12 m; 
sloping fore reef in 14 m, and outer reef 
slope, coral with sand pockets in 14 m; ver- 
tical rock wall and slope, dead coral and 
rocks, 18—26 m. 

Remarks. —This is the first record of this 
species from Tobago. 

Distribution. —Bahamas and southern 
Florida to Colombia; shore and shallow 
sublittoral to 38 m, usually on coral reefs. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Gonodactylus oerstedii Hansen, 1895 


Gonodactylus Oerstedii Hansen, 1895:65, 
footnote [part]. 

Gonodactylus oerstedii.—Manning, 1969: 
325, fig. 89. 


Material. —Sta TOB-21, Buccoo Reef, 14 
m: 2 92 (38, 58).—Heard sta 3, Lover’s 
Beach, shore to about 2 m: 3 juvenile ¢ (9- 
16).— Heard sta 5, Sandy Bay, shore to 1.5 
m: 15 66 (9.5-52), 19 9° (11-43), 3 juveniles 
(9), some early larvae.— Heard sta. 9, Petit 
Trou, 1-1.5 m: 1 2 (50). 

Color in life. —Female: body color yellow 
or pale blue-green with overall speckling of 
dark chromatophores including merus; 
multiple blue and green bands on sixth ab- 
dominal somite and telson; dactyl bright 
pink at base, light blue distally; meral spot 
bright pink or pale purple-blue persisting in 
preservation, flanked by two dark blue or 
blue-brown spots. In preservative, female 
with numerous persistent black spots on 
body, in bands on claw. 

Size. — Males (17), TL 9-52 mm; females 
(21), TL 11-58 mm; juveniles (3), TL 9 mm. 
Manning (1969) studied males with total 
lengths of up to 76 mm and females as large 
as 68 mm. | 

Habitat. —Shallow lagoon with turtle 
grass, bottom coarse sand to fine coralline 
silt, 1—-1.5 m; shore to about 2 m on pro- 
tected beach area; from washing of live rock 
taken between shore and 1.5 m; and from 
sloping fore reef at a depth of 14 m. 

Remarks.—This species was recorded 
from Pigeon Point and Buccoo Reef, To- 
bago by Manning (1969). 

Distribution. — Bermuda, Bahamas, 
southern Florida and throughout the Carib- 
bean; shore and shallow sublittoral. 


Gonodactylus spinulosus Schmitt, 1924 


Gonodactylus oerstedii var. spinulosus 
Schmitt, 1924b:96, pl. 5, fig. 5. 

Gonodactylus spinulosus. —Manning, 1969: 
299, fig. 83. 


Material.—Sta TOB-6, off Speyside, 11 


VOLUME 106, NUMBER 3 


m: 1 ?(21).—Sta TOB-21, Buccoo Reef, 14 
m: 1 2(36).—Sta TOB-36, east side of North 
Point, 5—12 m: 1 2 (20).—Sta TOB-39, Lon- 
don Bridge Rock, 13-28 m: 1 6 (22).—Sta 
TOB-40, Saint Giles Island, 6-11 m: 1 ¢ 
(24).—Sta TOB-42, Bloody Bay, 0-3 m: 1 
6 (18), 2 juveniles (7, 10.5). 

Color in life. —Body color and chromato- 
phore pattern as described in Manning 
(1969:303); meral spot without distinctive 
pigment, sometimes flanked by one or two 
blue spots. 

Size. —Males (3), TL 18-24 mm; females 
(3), TL 20-36 mm; juveniles (2), TL 7 and 
10.5 mm. Manning’s (1969) material in- 
cluded males to TL 48 mm, females to TL 
54 mm. 

Habitat. —On Tobago, adults were taken 
in sublittoral habitats on rough bottoms in 
depths between 5—12 and 13-28 m on slop- 
ing fore reef; vertical wall with rock, coral, 
and rubble, 5-12 m; vertical rock wall to 
ledge, 13-28 m; in rock and rubble with live 
and dead coral, 6—1 1 m; and on vertical rock 
wall and slope with dead coral and rocks, 
18—26 m. Juveniles were taken in shallower 
habitats, in O—3 m on sand, algae and rocks. 

Remarks. —This species has not been re- 
corded previously from Tobago. 

Distribution. —Bermuda and southern 
Florida to Tobago; shore and shallow sub- 
littoral, on coral reefs. 


Family Pseudosquillidae 
Pseudosquilla ciliata (Fabricius, 1787) 


Squilla ciliata Fabricius, 1787:333. 
Pseudosquilla ciliata. —Manning, 1969:264, 
fig. 74. 


Material. —Sta TOB-21, Buccoo Reef, 14 
m: 1 6 (43).—Heard sta 3, Lover’s Beach, 
shore to about 2 m: 1 postlarva (19).— Heard 
sta 9, Petit Trou, 1-1.5 m: 1 2 (43). 

Color in life. —Body pale pink-white with 
diffuse pink mottling overall including dac- 
tyl, propodus, and merus; dark lateral spots 
on first thoracic and first abdominal so- 
mites; two dark spots at base of telson; spines 
and carinae of sixth abdominal segment and 


a7 1 


telson banded with bright pink; meral spot 
without distinctive pigment, flanked by two 
pink-purple patches. 

Size. — Male (1), TL 43 mm; female (1), 
TL 43 mm; postlarva (1), TL 19 mm. Spec- 
imens reported by Manning (1969) included 
males to 80 mm long, females to 89 mm, 
and postlarvae ranging from 18 to 24 mm. 

Habitat. —Shore to about 2 m on a beach 
and shallow reef area; in a shallow lagoon 
with turtle grass, bottom from coarse sand 
to fine coralline silt, in 1-1.5 m; and sub- 
littoral, on sloping fore reef at a depth of 
14 m. 

Remarks. —This species has not been re- 
ported previously from Tobago. 

Distribution. —Widely distributed in all 
tropical oceans except the eastern Pacific. 
In the western Atlantic it occurs from Ber- 
muda, the Bahamas, and Florida, south- 
ward to Brazil; shore and shallow sublit- 
toral. 


Superfamily Lysiosquilloidea 
Family Lysiosquillidae 
Lysiosquilla glabriuscula (Lamarck, 1818) 
Fig. 2 


Squilla glabriuscula Lamarck, 1818:188. 
Lysiosquilla glabriuscula. —Manning, 1969: 
34, figs. 5c-d, 6. 


Material. —Sta TOB-16: Tobago, east of 
North Point, 21 m: 1 2 postlarva (29). 

Color in life. —Body yellow-white; anten- 
nal scale, propodus and merus of claw with 
median spotted areas; carapace with three 
bands of brown pigment, posteriormost dif- 
fuse; thoracic somites with diffuse brown 
band at posterior margin; abdominal so- 
mites with diffuse, broken bands anteriorly 
and dark, thinner bands at posterior margin, 
usually interrupted at midline; telson with 
three large dark spots; endopod and exopod 
dark, without pigment near margins. 

Size. —Postlarval female (1), TL 29 mm. 
Adults of this species exceed 200 mm in 
total length (Manning 1969). 

Habitat. —Sublittoral, rock wall to rubble 
flat, crevices, at a depth of 21 m. 


572 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


B 


Fig. 2. Lysiosquilla glabriuscula (Lamarck), female postlarva, TL 29 mm. A, Anterior part of carapace and 
frontal appendages; B, Sixth abdominal somite, telson and left uropod. 


Remarks. —This species has not been re- 
corded previously from Tobago, although 
that is well within its known range. 

We take this opportunity to provide a fig- 
ure of the postlarva, which differs from the 
adult in having distinct submedian denti- 
cles. 

This species apparently replaces L. sca- 
bricauda (Lamarck, 1818) in reef habitats; 
L. scabricauda is the common species of the 
genus in other shore habitats in the western 
Atlantic. 

Distribution. —South Carolina and Flor- 
ida to Brazil; shore and shallow sublittoral, 
near coral reefs. 


Family Nannosquillidae 
Alachosquilla, new genus 


Diagnosis. —Size small, TL less than 50 
mm. Cornea subglobular. Antennal pedun- 
cle with 2 papillae, 1 mesial, 1 ventral. Ros- 
tral plate rectangular, with 3 apical projec- 
tions. Mandibular palp present or absent. 
Five epipods present. Claw lacking distal 
ventral spine on ischium. Sixth abdominal 
somite with posterolateral spines; posterior 
margin unarmed ventrally. Telson with 5 
spines dorsally, 4 pairs of fixed primary 
marginal teeth, submedians, intermediates, 
laterals, and marginals, and 1 pair of mov- 
able submedian teeth; ventral surface with- 


out spinules. Spines of uropodal exopod 
sharp. 

Type species. —Lysiosquilla floridensis 
Manning, 1962, by present designation. 

Derivation of name.—From the Greek, 
alachos, bedfellow, in combination with the 
generic name Squilla, alluding to the asso- 
ciation of members of this genus with bal- 
anoglossid worms. Coutiére (1905) record- 
ed the association of A. digueti with a 
polychaete and a balanoglossid. Holthuis 
(1967:25) noted that all specimens of A. vi- 
cina then known were taken with balano- 
glossids, and Rodrigues (1971:209), in the 
original account of Callianassa guassutinga, 
recorded the occurrence of A. floridensis with 
balanoglossids in Brazil. 

Included species. —Two from the Amer- 
icas, Alachosquilla digueti (Coutiére, 1905: 
174), new combination, from the eastern 
Pacific and Alachosquilla floridensis (Man- 
ning, 1962), new combination, from the 
western Atlantic. It also includes Alacho- 
squilla vicina (Nobili, 1904:229), new com- 
bination, from the Red Sea, the Gulf of 
Aden, and the Philippines in the Indo-West 
Pacific (Nobili 1904, Kemp 1915, Holthuis 
1967). 

Remarks.—The rectangular rostral plate 
with three anterior projections and the 
subglobular cornea will distinguish mem- 
bers of this genus from members of Acan- 


VOLUME 106, NUMBER 3 


313 


Fig. 3. Alachosquilla floridensis (Manning), male, TL 33 mm. A, Carapace and frontal appendages; B, Sixth 


abdominal somite, telson and uropods. 


thosquilla sensu stricto in which the rostral 
plate is triangular, with a single apical spine, 
and the cornea is bilobed (see Chopra 1939: 
fig. 8). 

Members of this genus are unique in the 
Nannosquillidae in having the presence of 
the mandibular palp variable. In some spec- 
imens it is present on one side, absent on 
the other. 


Alachosquilla floridensis (Manning, 1962), 
new combination 
Fig. 3 


Lysiosquilla floridensis Manning, 1962a: 
221; 1969:67, fig. 16. 


Material.—Sta TOB-39, Saint Giles Is- 
land, 13-28 m: 1 ¢(30).—Sta TOB-40, Lon- 
don Bridge Rock, 6-11 m: 1 4 (33). 

Color in life. —Body cream-colored; car- 
apace speckled with brown chromato- 
phores, posterolateral angles outlined with 
dark semi-circles; diffuse speckled bands 
plus dark brown bands on posterior margins 
on all body segments; sixth abdominal so- 
mite with dark posterolateral spots and col- 
or bands on lateral margins; submedian 
dorsal spines with two dark basal spots; en- 


dopod dark, diffuse brown pigment on basal 
segment of exopod. 

Size.—Males (2), TL 30 and 33 mm. 
Manning (1969) studied males 45-48 mm 
long. 

Habitat. —Sublittoral, in 6-11 and 13-28 
m; from rock, rubble, with live and dead 
coral, 6—11 m; and vertical rock wall to ledge, 
13-28 m. This species, which burrows in 
level bottom substrates, certainly was taken 
from sand on the ledge rather than on the 
wall itself. 

Remarks. —Manning (1974:105) synon- 
ymized the western Atlantic Lysiosquilla 
floridensis Manning, 1962 with L. digueti 
(Coutiére, 1905) from the eastern Pacific; 
both were then in Acanthosquilla, where they 
were placed by Manning (1963). We now 
believe that they should be recognized as 
distinct species. In all specimens of A. /flor- 
idensis available for study, the median black 
spot on the telson is distinctly divided into 
two spots, whereas all specimens of A. di- 
gueti that we have examined have a single 
black spot on the telson. 

In addition to the material from Florida 
and Brazil recorded by Manning (1969), 
there are specimens in the national collec- 


574 


tions from Lake Worth Inlet, Florida 
(USNM 256863, 2 TL 49 mm), from Virgin 
Gorda, U.S. Virgin Islands (USNM 170214, 
3 6, TL 36, 38, and 44 mm), Isla Margarita, 
Venezuela (USNM 170214, 2, TL 14 mm), 
and Bahia, Brazil (USNM 150780, 6, TL 33 
mm). 

The mandibular palp is absent in both 
specimens from Tobago. 

Distribution. —Southern Florida to Bra- 
zil; shore. 


Bigelowina, new genus 


Diagnosis. —Size small, TL less than 50 
mm in adults. Cornea subglobular. Anten- 
nal peduncle with 2 papillae, 1 mesial, 1 
ventral. Rostral plate quadrangular, with 
apical spine. Mandibular palp present. Five 
epipods present. Claw lacking distal ventral 
spine on ischium. Sixth abdominal somite 
with posterolateral spines, posterior margin 
unarmed ventrally. Telson with 5 submar- 
ginal spines dorsally, marginal armature 
comprising 2 pairs of primary marginal 
teeth, laterals and marginals; ventral surface 
unarmed. Spines of uropodal exopod sharp. 

Type species.—Lysiosquilla biminiensis 
Bigelow, 1893. 

Included species. —Two, the amphi- 
American Bigelowina biminiensis (Bigelow, 
1893), new combination, and the eastern 
Atlantic Bigelowina septemspinosa (Miers, 
1881:368), new combination (see Manning 
1977 for an account of this species). 

Etymology.—Named for Robert Payne 
Bigelow (10 July 1863-6 September 1955), 
zoologist and educator, who was the Amer- 
ican pioneer in studies on the systematics 
of stomatopods. The gender is feminine. 

According to his biography on p. 396 of 
The National Cyclopaedia of American Bi- 
ography, vol. 46, R. P. Bigelow was edu- 
cated at Harvard and Johns Hopkins Uni- 
versity. He joined the Massachusetts 
Institute of Technology in 1893 as an in- 
structor in biology and retired as professor 
of zoology in 1933. He led the Johns Hop- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


kins zoological expeditions to Jamaica in 
1891 and 1893. In 1892 he was joined by 
E. A. Andrews and T. H. Morgan on an 
expedition to Bimini, a trip chronicled in 
Andrews, Bigelow & Morgan (1945), during 
which they drank from the “Fountain of 
Youth” (p. 339). They may well have sam- 
pled waters from the fountain of youth, for 
as they noted in 1945, their combined ages 
then totalled nearly 250 years. L. B. Hol- 
thuis (in litt., 9 Aug 1992) remarked that he 
purchased Bigelow’s stomatopod library for 
$35 in 1952 and met Bigelow in 1953. 

Bigelow’s published work on stomato- 
pods (1891, 1893a, 1893b, 1894, 1901, 
1926, 1931, 1941), though not voluminous, 
established a firm foundation for subse- 
quent work on the systematics of American 
members of the group. 

Remarks. —Members of Bigelowina, which 
until now have been placed in Acanthosquil- 
la Manning, 1963, resemble species of 
Acanthosquilla in having a submarginal, fan- 
shaped row of five dorsal spines on the tel- 
son. Species of Bigelowina can be distin- 
guished from those of Acanthosquilla by 
their rectangular rather than triangular ros- 
tral plate and by their subglobular cornea; 
the cornea is bilobed in species of Acan- 
thosquilla sensu stricto. The characteristic 
rostral plate and cornea of Acanthosquilla 
are well-illustrated in Chopra (1939:fig. 8). 

Members of Bigelowina differ from species 
of Alachosquilla in having a single rostral 
spine and in having four rather than two 
pairs of primary marginal spines on the tel- 
son. 

Many of the species now placed in Acan- 
thosquilla will be assigned to other genera 
in a revision of the stomatopod genera now 
in preparation. 


Bigelowina biminiensis (Bigelow, 1893), 
new combination 
Fig. 4 


Lysiosquilla biminiensis Bigelow, 1893b: 
Os 


VOLUME 106, NUMBER 3 


Fig. 4. Bigelowina biminiensis (Bigelow), female, TL 45 mm. A, Carapace and frontal appendages; B, Sixth 


abdominal somite, telson and uropods. 


Acanthosquilla biminiensis. —Manning, 
1969:63, figs. 14, 15. 


Material. — Heard sta 4, Bloody Bay, shore 
to about 2 m: 1 2 (45). 

Color. —Color in life not recorded. In pre- 
servative, posterolateral angle of carapace 
ringed with black semicircle containing 
dusky circle in center; thoracic somite 8 with 
short black line on each side of posterior 
margin; sixth abdominal somite with short, 
dark line posterolaterally, flanked anteriorly 
by dusky circle. 

Size.—Female (1), TL 45 mm. Males 
measuring 16-62 mm and females 21-50 
mm long have been recorded in the litera- 
ture (Manning 1969, 1974). 

Habitat.—On sand bottom with some 
rocks between beach and 2 m depth. 

Remarks. — Additional material of this 
species added to the national collections 
since Manning’s (1969) monograph has pro- 
vided a much clearer picture of the distri- 
bution of this species. There is material from 
off South Carolina in 37 m (USNM 174488), 
off Georgia in 14, 34 and 46 m (USNM 
128350, 174484, 174486, and 174487), and 


off northeastern Florida in 15 m (USNM 
174485). 

Distribution. —Bahamas and South Car- 
olina to Brazil in the western Atlantic and 
from Panama in the eastern Pacific (Man- 
ning 1974); shore and shallow sublittoral to 
a depth of 46 meters. 


Genus Nannosquilla Manning, 1963 
Nannosquilla tobagoensis, new species 
Fig. 5 


Material. —Sta TOB-21, Buccoo Reef, 14 
m: 1 6 (20), holotype, USNM 252681. 

Diagnosis. —Eye small, extending to end 
of second segment of antennular peduncle. 
Cornea subglobular, width 0.8 times rostral 
plate length. Ocular scales with bases fused 
medially, apices distinct, rounded. Anterior 
margin of ophthalmic somite produced into 
blunt median projection, ventral spine also 
present. 

Antennular peduncle short, less than half 
as long as carapace. Upper flagellum with 
15-16 articles, longer lower flagellum as long 
as upper, with 16-17 articles, shorter lower 
flagellum with 8 articles. Antennular pro- 


576 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 5. 
ages; B, Claw; C, Right uropod, ventral view; D, Telson, ventral view; E, Telson, posterior view; F, Posterior 
part of sixth abdominal somite and telson. 


cesses visible as anteriorly-directed spines 
projecting beyond sides of rostral plate, 
barely overreaching anterolateral corners of 
plate. 

Antennal peduncle falling short of ante- 
rior margin of eye. Flagellum with 11-12 
articles. Antennal scale short, extending 
about to middle of distal segment of anten- 
nal peduncle. 

Rostral plate subquadrangular, length 0.8 


Nannosquilla tobagoensis, new species, male holotype, TL 20 mm. A, Carapace and frontal append- 


times width, apex falling short of midlength 
of eyestalks. Lateral margins of plate par- 
allel, convex. Anterolateral corners forming 
rounded right angles. Anterior margin shal- 
lowly biconcave, low, obtuse median pro- 
jection not extending beyond anterolateral 
corners of plate. 

Mandibular palp absent. Four epipods 
present. 

Dactylus of claw with 8 teeth. Proximal 


VOLUME 106, NUMBER 3 


notch on outer margin of dactylus flanked 
by subacute proximal lobe and more round- 
ed, larger distal lobe. Carpus with angled 
projection distally on upper margin. 

Basal segments of walking legs unarmed. 

Sixth abdominal somite with acute pos- 
terolateral angles. 

Telson width about twice median length, 
with broad, obtuse median projection, sep- 
arated from lateral projections by shallow 
submedian concavities. Marginal armature 
consisting of, on each side of midline, 6 
submedian denticles, a movable submedian 
tooth near the posterior margin, and 7 lat- 
eral teeth. 

Uropodal exopod with 5 movable spines 
on outer margin of proximal segment, distal 
2 spatulate; inner margin of proximal seg- 
ment with 2 stiff setae. Outer spine of basal 
prolongation of uropod the longer. 

Color in life. —Body color off-white with 
sparse, scattered brown chromatophores on 
all somites except sixth abdominal somite, 
densest pigment on fifth abdominal somite. 

Size. — Male (1), TL 20 mm. Other mea- 
surements of unique male holotype: cara- 
pace length 3.5; rostral plate length 1.0, 
width 1.4; telson length 1.4, width 2.8. 

Habitat. —Sublittoral, in 14 m on sloping 
fore reef. 

Remarks. —This is the twenty-sixth spe- 
cies of the genus, all but one of which occur 
only in the Americas; six species are known 
from the Caribbean (Camp & Manning 
1982, 1986; Manning 1970, 1979). This ge- 
nus has not been recorded previously from 
the Trinidad and Tobago region. 

This species resembles N. virginalis Camp 
& Manning, 1986, differing from it in hav- 
ing a more quadrate rostral plate with a low- 
er anterior projection, longer antennular 
processes that extend beyond the anterolat- 
eral corners of the rostral plate, a shorter 
antennal peduncle that does not extend be- 
yond the end of the eye, and eight rather 
than seven teeth on the claw. 

Etymology. —Named for the type locali- 
ty. 


S77 


Distribution. —Known only from the type 
locality. 


Superfamily Squilloidea 
Family Squillidae 
Alima alba (Bigelow, 1893) 


Squilla alba Bigelow, 1893:103. 

Alima hyalina. —Manning, 1962b:496; 
1969:128, figs. 37, 38, 39a. [Not Alima 
hyalina Leach, 1817, a larva.] 

Alima neptuni.—Manning & Lewinsohn, 
1986:13, 15 [not Cancer neptuni Linnae- 
us, 1768, a larva]. 


Material. —Heard sta 10A, Buccoo Reef, 
about 2 m: | 6 (46). 

Size. —One male only collected, TL 46 
mm. 

Habitat. —Sand patch in back reef area in 
a depth of about 2 m. 

Remarks. —Manning (1962) identified a 
pelagic larva originally described from the 
Gulf of Guinea, Alima hyalina Leach, 1817 
as the larva of the species then known as 
Squilla alba Bigelow, 1893. Subsequently, 
Manning reported two different species of 
Alima from localities in the western Atlantic 
(1969) and the eastern Atlantic, A. hyalina 
(Leach) and A. hieroglyphica (Kemp, 1911). 
In 1986 Manning & Lewinsohn identified 
A. hyalina with a larval form named by Lin- 
naeus. 

We question the wisdom of using names 
based on larval forms for adult species un- 
less the adult can be raised from larvae from 
a known parent, especially in stomatopods 
in which there are so many larval forms. 
The alima larva reported by Manning (1962) 
that metamorphosed into a postlarva that 
could be identified with Squilla alba cer- 
tainly resembled the larva named by Leach, 
but with two adults of the same genus oc- 
curring in the area from which Leach’s larva 
was taken, the Gulf of Guinea, there is no 
way to positively identify his larva with ei- 
ther of the two adults. For the same reason 
the name used by Linnaeus, based on a larva 
from the central Atlantic, cannot be used 


578 


for the species. We believe it is best to use 
the name based on the adult for the species, 
that used by Bigelow in 1893. 

There is an adult female, TL 42 mm, of 
this species in the national collections taken 
by D. L. Felder and colleagues from a tidal 
flat exposed at low water at Peanut Island 
in Lake Worth, Palm Beach County, Flor- 
ida, on August 1987. 

This species has not been reported pre- 
viously from Tobago. 


Acknowledgments 


This study is a component of a compre- 
hensive survey of the fauna of Tobago being 
carried out for the Tobago House of Assem- 
bly by J. David Hardy, whose support dur- 
ing the collecting expedition we appreciate. 
We would also like to acknowledge the lo- 
gistical help from the Marine Fisheries Sec- 
tion of Tobago Fisheries in Scarborough. 
Richard Heard kindly shared collections 
with us. Manning’s studies on the system- 
atics of stomatopods are supported by the 
Smithsonian Marine Station at Link Port, 
Florida. This is contribution number 312 
from that station. 


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Department of Invertebrate Zoology, Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. 20560, 
U.S.A. 


Appendix—Station Data 


A. Station Data and Species List for Collections 
Made by M. Schotte and Others, 1990 


TOB-6 (= JTW 90-4): Tobago; off Speyside (1 1°18'N, 
60°32'W), leeward side of Little Tobago Island, 35 ft 
(11 m), rotenone, 6 Sep 1990, leg. D. Johnson, et al.: 
Gonodactylus curacaoensis, G. spinulosus. 

TOB-12 (= JTW 90-5): Tobago, off Little Tobago 
Island (11°18’N, 60°30’W), coral heads, depth 60 ft (18 
m), rotenone, 7 Sep 1990, leg. D. Johnson, et al.: Gon- 
odactylus caribbaeus, new species, G. curacaoensis. 

TOB-13: Tobago, cove near Speyside (11°18'N, 
60°32’W), large rock covered with coral and sea whips, 
depth 35 ft(11 m), rotenone, 7 Sep 1990, leg. M. Schotte, 
et al.: Gonodactylus caribbaeus, new species. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


TOB-16: Tobago, east of North Point (11°20’N, 
60°33’W), rock wall to rubble flat, crevices, depth 70 
ft (21 m), rotenone, 8 Sep 1990, leg. D. Johnson, et 
al.: Gonodactylus bredini, G. curacaoensis, Lysiosquilla 
glabriuscula. 

TOB-17: Tobago, northeast of Charlotteville 
(11°19'N, 60°33’W), coral-encrusted rocks and ledges, 
depth 40 ft (12 m), rotenone, 8 Sep 1990, leg. M. Schotte, 
et al.: Gonodactylus curacaoensis. 

TOB-21: Tobago, Buccoo Reef (11°11'N, 60°49’W), 
sloping force reef, depth 45 ft (14 m), rotenone, 10 Sep 
1990, leg. M. Schotte, et al.: Gonodactylus caribbaeus, 
new species, G. curacaoensis, G. oerstedii, G. spinu- 
losus, Nannosquilla tobagoensis, new species, Pseu- 
dosquilla ciliata. 

TOB-36 (= JTW 90-9): Tobago, Man O’War Bay, 
east side of North Point (11°20’N, 60°33’W), vertical 
wall with rock, coral, and rubble, depth 15—40 ft (5— 
12 m), rotenone, 8 Sep 1990, M. Schotte, et al.: Gon- 
odactylus spinulosus. 

TOB-38 (= JTW 90-10): Tobago, Buccoo Reef 
(11°11'N, 60°49’W), outer reef slope, coral with sand 
pockets, depth 45 ft (14 m), rotenone, 10 Sep 1990, 
leg. M. Schotte, et al.: Gonodactylus curacaoensis. 

TOB-39 (= JTW 90-14): Tobago, St. Giles Island, 
London Bridge Rock (11°21'N, 60°32’W), vertical rock 
wall to ledge, depth 40-80 ft (13—28 m), rotenone, 12 
Sep 1990, leg. J. T. Williams, et al.: Alachosquilla flor- 
idensis, Gonodactylus spinulosus. 

TOB-40 (= JTW 90-15): Tobago, Saint Giles Island 
(11°21’N, 60°31'W), rock, rubble, live and dead coral, 
depth 20-35 ft (6-11 m), 12 Sep 1990, rotenone, leg. 
J. T. Williams, et al.: Alachosquilla floridensis, Gono- 
dactylus caribbaeus, new species, Gonodactylus spi- 
nulosus. 

TOB-42 (= JTW 90-16): Tobago, Bloody Bay 
(11°18'W, 60°38’W), mouth of Bloody Bay River, sand, 
algae, and rocks, depth 0-10 ft (0-3 m), rotenone, 13 
Sep 1990, leg. J. T. Williams, et al.: Gonodactylus spi- 
nulosus. 

TOB-44: Tobago, The Sisters (11°20’N, 60°39’W), 
vertical rock wall and slope, dead coral and rocks, depth 
60-85 ft (18-26 m), rotenone, 14 Sep 1990, leg. J. T. 
Williams, et al.: Gonodactylus curacaoensis. 


B. Station Data and Species List for Collections 
Made by R. Heard, 1992 and 1993 


Heard Sta. 3: Tobago, Lover’s Beach, protected beach 
and shallow reef area on northwest corner of Man O’War 
Bay (11°19’N, 60°34’W), shore to a depth of about 2 
m, 6 Apr 1992: Gonodactylus oerstedii, Pseudosquilla 
ciliata. 

Heard Sta 4: Tobago, Bloody Bay (11°18’N, 60°38’W), 
sand bottom with some rocks, beach to a depth ofabout 
2 m, 4 Apr 1992: Bigelowina biminiensis. 

Heard Sta 5: Tobago, Sandy Bay (= Milford Bay, 


VOLUME 106, NUMBER 3 


11°09’N, 60°50’W), near Pigeon Point, west end of is- 
land, protected sand beach bordered by Buccoo Reef, 
from shore to a depth of 1.5 m, washing of live rock, 
7 Apr 1992: Gonodactylus oerstedii. 

Heard Sta 9: Tobago, Petit Trou (= Lowlands La- 
goon, 11°08’N, 60°47'W), a shallow lagoon with rich 


581 


patches of turtle grass, bottom varied from coarse sand 
to fine coralline silt, depth 1-1.5 m, 7 Apr 1992: Gon- 
odactylus oerstedii, Pseudosquilla ciliata. 

Heard Sta 10: Tobago, Buccoo Reef, sand patch in 
back reef area, depth about 2 m, 11 Jun 1993: Alima 
alba. 


PROC. BIOL. SOC. WASH. 
106(3), 1993, pp. 582-586 


A NEW SPECIES OF SPHAERODORIDAE 
(ANNELIDA: POLYCHAETA) FROM 
SOUTHERN CALIFORNIA 


Jerry D. Kudenov 


Abstract. —Sphaerodoropsis sexantenella, new species, 1s described from 
Southern California. It is most closely related to both S. oculata Fauchald and 
S. pycnos Fauchald in having 10—11 rows of dorsal macrotubercles arrayed in 
a zig-zag pattern, and may differ from these and all other known congeners in 
having characteristic retractable accessory papillae on inferior lateral prosto- 


mial antennae. 


Fauchald (1974:257-289) reviewed 
sphaerodorid polychaetes from world areas, 
redefined genera, and in all, recognized nine 
genera on the basis of characters that gen- 
erally had not been applied in systematically 
consistent ways previously. In contrast, Pet- 
tibone (1982) recognized only four genera. 
This apparent discrepancy reflects the fact 
that Pettibone relied on traditional char- 
acters, which are fewer in number than those 
used by Fauchald (1974), to define sphae- 
rodorid genera. However, Pettibone’s 
scheme may result in polytypic genera, and 
at least one genus (Ephesiopsis Hartman and 
Fauchald) is excluded from her overview of 
the family. Fauchald’s system has been used 
almost exclusively by most workers since 
1974. It is used in the present study in an 
attempt to maintain monotypic definitions 
of genera and to facilitate species compar- 
isons. This seems particularly appropriate 
since Perkins (1987) moved the genus Levi- 
dorum to a new family, Levidoridae, and 
Kudenov (1987) described the new genus, 
Amacrodorum, that is strongly isolated 
morphologically in lacking macrotubercles. 
Differences between Fauchald’s (1974, 1977) 
and Pettibone’s (1982) schemes need to be 
more thoroughly addressed 

The present material derives from the on- 
going California Phase II Monitoring Pro- 
gram (CAMP) of the Minerals Management 


Service (MMS) as part of their Outer Con- 
tinental Shelf Environmental Studies Pro- 
gram, and represents part of Science Ap- 
plications International Corporation’s 
(SAIC) overall effort to produce a taxonom- 
ic atlas of the macroinvertebrate fauna of 
the Santa Maria Basin and the western Santa 
Barbara Channel. 

Type materials are deposited in the Na- 
tional Museum of Natural History, Smith- 
sonian Institution (USNM); both types and 
non-types in the Natural History Museum 
of Los Angeles County (LACM). 


Sphaerodoropsis sexantennella, 
new species 
Figs. 1-2 


Sphaerodoropsis species A. Kudenov, 1992: 
4-379, fig. 4. 122A—-K. 


Material examined. -MMS CAMP 
Phase II, Sta. PJ-1, rep. 2, 34°55.79'N; 
120°49.91'W, 145 m, 5 specimens (LACM- 
AHF POLY); Sta., PJ-1, rep. 3; sames3 
(LACM-AHF POLY); Sta. PJ-7, rep. 1, 
34°55.79'N, 120°48.60'W, 123 m, holotype 
(USNM 157606), 7 paratypes (LACM-AHF 
POLY 1626; USNM 157607); Sta. PJ-7, rep. 
2, same, 3 paratypes (LACM-AHF POLY 
1627); Sta. PJ-8, rep. 1, 34°S63:87 RE 
120°49.91’W, 142 m, 3 (LACM-AHF 


VOLUME 106, NUMBER 3 


POLY); Sta. PJ-8, rep. 2, same, 3 (LACM- 
AHF POLY); Sta. PJ-8, rep. 4, same, 2 
(LACM-AHF POLY); Sta. PJ-9, rep. 1 
B4255-T9LNe 120°51-23'°W, 169 m; 3 
(LACM-AHF POLY); Sta. PJ-9, rep. 2, 
same, 1 (LACM-AHF POLY); Sta. PJ-9, 
rep. 3, same, 1 (LACM-AHF POLY); Sta. 
PJ-10, rep. 1, 34°53.63'N, 120°49.91'W, 147 
m, 2 (LACM-AHF POLY); Sta. PJ-10, rep. 
3, same, 3 (LACM-AHF POLY); Sta. PJ- 
Misep: 2, 34°57.95'N,; 120°49.91'W, 136 
m, 2 (LACM-AHF POLY); Sta. PJ-11, rep. 
3, same, 1 (LACM-AHF POLY). 

Description. —Holotype measuring about 
1 mm long, 0.2 mm wide excluding setae, 
with 14 setigerous segments. Specimens 
having 12-14 setigerous segments oviger- 
ous. Prostomium with a short globular me- 
dian antenna plus two pairs of short digitate 
to globular lateral antennae (Fig. 1 A—E). Su- 
perior lateral antennae smaller, shorter than 
inferior lateral antennae. Superior lateral 
antennae apparently lacking accessory pa- 
pillae; inferior lateral antennae each with 
two retractable accessory papillae on me- 
dian basal surfaces (Fig. 1D, E). These nip- 
ple-shaped, surrounded by a circlet of six 
cirriform appendages (Fig. 1E). Two pairs 
of additional accessory papillae on body wall 
near midline, between superior and inferior 
lateral antennae (Figs. 1E, 2A, B). Pair large 
medial eyes present, deeply embedded in 
body wall, on a line behind median antenna 
at level of setiger 1 (Fig. 1D). Peristomial 
cirri short, papilliform (Fig. 1B—D). 

Parapodia each with broad, bluntly con- 
ical acicular and erect digitiform presetal 
lobes; postsetal lobes absent (Fig. 1F, G). 
Parapodial papillae numbering three, in- 
cluding one each on anterior parapodial sur- 
faces after setiger 2, and one each on prox- 
imal superior and inferior edges of all 
parapodia (Fig. 1G); papillae otherwise ab- 
sent from anterior surfaces of first 1—2 se- 
tigers (Fig. 1F). Ventral cirri large, digiti- 
form, not projecting beyond tip of acicular 
lobes (Fig. 1F, G). 

Macrotubercles arrayed in ten or eleven 


583 


rows, forming a zig-zag pattern on dorsum, 
sessile, lacking terminal papillae. Papillae 
short, blunt, present dorsally between rows 
of macrotubercles; present ventrally in six 
staggered rows (Fig. 1H). 

Setae entirely composite, of one kind, 
generally numbering six per fascicle (Fig. 
1F, I-K); shafts inflated, sometimes with an 
indistinct subdistal spur (Fig. 11), forming 
terminal sockets for long, falcate blades; 
blades with indistinctly serrated cutting 
margins and sharp recurved tips (Fig. 1K), 
varying up to two times longer than the 
shortest blade in setiger 1 (Fig. 1C), becom- 
ing nearly equal in length within a fascicle 
thereafter. 

Anus terminal, with pair of dorsal anal 
papillae and longer midventral anal cirrus 
(Fig. 1L, M). 

Remarks.— Sphaerodoropsis sexanten- 
nella is unusual in having characteristic and 
retractable accessory papillae on both the 
inferior lateral antennae and on the body 
wall between the inferior and superior lat- 
eral antennae (Fig. 2; note that Fig. 2A isa 
composite illustration of USNM 157606- 
157607). Such a feature has apparently not 
been noted previously in the genus Sphae- 
rodoropsis, and strongly isolates S. sexan- 
tennella from other described species in this 
taxon. However, this trait is difficult to ob- 
serve in S. sexantennella since, 1) all spec- 
imens tend to be about 1 mm long, 2) the 
general size of all prostomial structures is 
strongly reduced when compared to other 
known species of Sphaerodoropsis, 3) the 
prostomium is strongly retracted in all spec- 
imens except the holotype and one paratype 
(both illustrated), and 4) accessory papillae 
were detected only in the holotype. This 
species would have been described as lack- 
ing accessory papillae, had they not been 
protracted in the holotype; both their total 
number and distribution are here consid- 
ered to be tentative pending the acquisition 
of additional specimens (Fig. 2B). It is sug- 
gested that the types of other sphaerodorid 
taxa be examined for the presence of re- 


584 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Sphaerodoropsis sexantennella, new species: A-C, paratype (USNM 157607); D-N, holotype (USNM 
157606). A, Anterior end and setiger 1, dorsal view; B, Anterior end, ventral view; C, Anterior end and setiger 
1, ventral view; D, Anterior end, dorsal view; E, Anterior end, ventral view; F, Right setiger 2, anterior view; 
G, Right setiger 8, anterior view; H, Ventrum, setigers 4-6, ventral view, arrow points anteriorly; I, Pygidium, 
dorsal view; J, Pygidium, ventral view; K-L, Composite falcigers; M, Serrated blade of composite seta, oblique 


VOLUME 106, NUMBER 3 


585 


mAn 


Fig. 2. Sphaerodoropsis sexantennella, new species: A-B, paratype (USNM 157607) and holotype (USNM 
157606). A, Composite of anterior end showing spatial relationships between left accessory papillae, inferior 
lateral antennae and body wall, ventral view. B, Schematic showing distribution of accessory papillae (solid 
circles) on the body wall and inferior lateral antennae in relation to the median antenna and superior lateral 
antennae. Abbreviations given in legend of Fig. 1. Scale = .05 mm: A. 


tractable accessory papillae on and around 
the prostomial antennae. 

Sphaerodoropsis sexantennella is most 
closely related to S. oculata Fauchald, 1974, 
and S. pyncos Fauchald, 1974, both of which 
were described originally from Antarctica. 
Sphaerodoropsis sexantennella and S. ocu- 
lata have ten or eleven rows while S. pycnos 
has eleven rows of dorsal macrotubercles 
arrayed in a zig-zag pattern. The first two 
species have well-developed eyes, and gen- 
erally similar parapodia, although all para- 
podial structures of S. sexantennella are 
much more stout than those of S. oculata; 
S. pycnos lacks eyes, and has foliaceous 
rather than digitiform presetal lobes. Sphae- 
rodoropsis sexantennella differs from S. 
oculata in having short prostomial antennae 
rather than long ones; papilliform peristo- 
mial cirri instead of digitiform structures; 
parapodial papillae on anterior, not poste- 
rior parapodial surfaces; blades of compos- 


— 


ite setae that are both longer and serrated; 
and ventral papillae arrayed in 6 orderly 
rows that do not cover the ventrum; and 
may differ in having 3 pairs of retractable 
accessory papillae only on inferior lateral 
prostomial antennae instead of having non- 
retractable papillae on all prostomial anten- 
nae (Fauchald 1974:fig. 19). 

Etymology. —The epithet, sexantennella, 
derives from the following Latin terms, in- 
cluding the prefix sex-, meaning six or six- 
fold; antenn, the root for the New Latin 
term antenna or feeler, and the suffix -e// 
added to the noun stem to form a dimin- 
utive. It refers to the small cirriform ap- 
pendages present on each of the nipple- 
shaped accessory papillae on inferior lateral 
antennae, and on the body wall between the 
superior and inferior lateral antennae. 

Type locality. —CAMP Phase II Sta. PJ- 
7, 34°55.79’'N, 120°48.60'W. 


view. Abbreviations: ap, accessory papilla; ilAn, inferior lateral antenna; mAn, median antenna; mt, macro- 
tubercle; PC, peristomial cirrus; slAn, superior lateral antenna. Scale 1 = 0.01 mm: A, B, D-G; Scale 2 = 0.05 
mm: C, H; Scale 3 = 0.05 mm: I, J; Scale 4 = 0.01 mm: K—-M. 


586 


Distribution. —Southern California, in 
depths of 123-169 m. 


Acknowledgments 


I am indebted to James A. Blake, SAIC, 
for making these materials available for 
study, and for reviewing the manuscript, 
which was also improved by comments from 
two anonymous reviewers. This study is 
based on work funded by MMS Contract 
No. 14-35-0001-30484 to Science Appli- 
cations International Corporation, Woods 
Hole, Massachusetts. 


Literature Cited 


Fauchald, K. 1974. Sphaerodoridae (Polychaeta: Er- 
rantia) from world-wide areas. — Journal of Nat- 
ural History, London 8:257-289. 

. 1977. The Polychaete worms. Definitions and 
keys to Orders, Families and Genera.— Los An- 
geles County Museum of Natural History, Sci- 
ence Series 28:1-190. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Kudenov, J. 1987. Four species of Sphaerodoridae 
(Annelida: Polychaeta) including one new genus 
and three new species from Alaska. —Proceed- 
ings of the Biological Society of Washington 
100(4):917-926. 
1992. Family Sphaerodoridae Malmgren 
1867. Pp. 371-383 in J. A. Blake, ed., Taxo- 
nomic atlas of the benthic fauna of the Santa 
Maria Basin and Western Santa Barbara Chan- 
nel. Vol. 4. Annelida Part 1. U.S. Department 
of the Interior, Minerals Management Service. 
Perkins, T. H. 1987. Levidoridae (Polychaeta), new 
family, with remarks on two new species of Levi- 
dorum from Florida.— Bulletin of the Biological 
Society of Washington 7:162-168. 

Pettibone, M. 1982. Polychaeta. Pp. 3-43 in S. P. 
Parker, ed., Synopsis and classification of living 
organisms. Vol. 2. McGraw-Hill. 


Department of Biological Sciences, Uni- 
versity of Alaska Anchorage, 3211 Provi- 
dence Drive, Anchorage, Alaska 99508, 
U.S Age 


PROC. BIOL. SOC. WASH. 
106(3), 1993, pp. 587-590 


A NEW MARINE SPECIES OF SMITHSONIDRILUS 
(OLIGOCHAETA: TUBIFICIDAE) FROM 
THE FLORIDA KEYS 


Christer Erséus 


Abstract. —Smithsonidrilus exspectatus, new species, is described from barely 
subtidal sediments at Pigeon Key (near Marathon) and Bahia Honda, southern 
Florida. It appears phylogenetically intermediate between Smithsonidrilus — 
hummelincki and the most apomorphic group within the genus (Smithsonidrilus 
marinus, Smithsonidrilus involutus, Smithsonidrilus westoni, and Smithsoni- 
drilus multiglandularis); all these species show a northwest Atlantic distribu- 
tion. The new species has the same kind of spermatheca as, but a less elaborate 
copulatory sac (pseudopenis) than, those of the species in the S. marinus group. 
The dilated, heavily muscular ejaculatory duct appears to be an autapomorphy 


of S. exspectatus. 


The marine tubificid genus Smithsoni- 
drilus Brinkhurst, 1966, was revised by Er- 
séus (1990), who presented an hypothesis 
of the phylogenetic relationships between 
all species of the genus, based on a parsi- 
mony analysis of morphological characters. 
According to the most parsimonious clado- 
gram (Erséus 1990: fig. 31), a group of spe- 
cies form a highly apomorphic, monophy- 
letic, group within Smithsonidrilus, defined 
by at least two synapomorphies, (1) the un- 
paired ejaculatory duct, and (2) the modi- 
fication of the prostatic pads into atrial di- 
verticula. These species, Smithsonidrilus 
marinus Brinkhurst, 1966, Smithsonidrilus 
involutus Erséus, 1990, Smithsonidrilus 
westoni Erséus, 1982, Smithsonidrilus mul- 
tiglandularis Erséus, 1990, and Smithsoni- 
drilus hummelincki (Righi & Kanner, 1979), 
are all northwest Atlantic (largely Carib- 
bean) taxa. 

During an ongoing study of the distri- 
bution of marine Tubificidae in southern 
Florida, by Milligan and Erséus, an addi- 
tional member of this group was found. The 
species, Smithsonidrilus exspectatus, new 
species, is described in the present paper. 

The material was collected by M. R. Mil- 


ligan and C. Erséus at barely subtidal lo- 
calities in the Florida Keys, southern Flor- 
ida. The specimens were sorted live under 
a dissecting microscope from elutriated sed- 
iment samples, fixed in Bouin’s fluid, and 
later stained with paracarmine and mount- 
ed whole in Canada balsam. The type ma- 
terial has been deposited in the United States 
National Museum of Natural History 
(USNM), Washington, D.C., and the Swed- 
ish Museum of Natural History (SMNH), 
Stockholm. 


Smithsonidrilus exspectatus, new species 
Fig. 1 


Holotype. -USNM 
mounted specimen. 

Type locality. —Off small beach, NE cor- 
ner of Pigeon Key (W of Marathon), Florida 
Keys, 0.1 m, coarse sand with gravel and 
pebbles (4 May 1990). 

Paratypes. -USNM 160305, 160306, 
three specimens from type locality. SMNH 
Type coll. 4533, two specimens from close 
to rocks, W end of Bahia Honda (N side), 
beach in Bahia Honda State Recreation 
Area, Florida Keys, 0.5 m, medium to coarse 


160304, whole- 


588 


| 25um | 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


100 um | 


Fig. 1. 


Smithsonidrilus exspectatus, new species. A, Seta; B, Lateral view of spermatheca and male genitalia 


in segments X—XI. Abbreviations: aa, atrial ampulla; ad, atrial duct; ado, atrial duct of other side; d, diverticulum 
(modified prostatic pad); ed, ejaculatory duct; pr, prostate gland; pp, pseudopenis (copulatory sac); sa, sper- 
mathecal ampulla; sd, spermathecal duct; sdi, spermathecal diverticulum; sf, sperm funnel; sv, spermathecal 


vestibule; vd, vas deferens. 


sand with cobbles and gravel (3 May 1990); 
SMNH Type coll. 4534, one specimen from 
at foot of dock, N side of Pigeon Key (W of 
Marathon), Florida Keys, 0.6 m, heteroge- 
neous medium to coarse sand (4 May 1990). 

Etymology.—Named exspectatus (Latin 
for ‘expected’) indicating that the species is 
morphologically intermediate between oth- 
er known species of the genus (see Discus- 
sion below), as if it could have been pre- 
dicted to exist. 

Description. —Length (7 complete speci- 
mens) 10.4-16.5 mm, 54-87 segments. 


Width at XI 0.41-0.55 mm. Prostomium 
pointed or rounded triangular. Clitellum ex- 
tending over about *2X—AXII. Setae (Fig. 
1A) bifid, with upper tooth much thinner 
and shorter than lower. Setae 45-80 um long, 
3-5.5 wm thick at node, two or three (oc- 
casionally only one) per bundle anteriorly, 
totally absent from XI, two (occasionally 
only one) per bundle thereafter. Male pore 
unpaired, located mid-ventrally and pos- 
terior to middle of XI. Spermathecal pore 
unpaired, mid-ventral, near middle of X. 
Pharyngeal glands in (III) IV—V. Oesopha- 


VOLUME 106, NUMBER 3 


geal diverticula, in IX, slender. Male geni- 
talia (Fig. 1B) complex, paired for most 
parts; but ejaculatory duct and pseudopenis 
(copulatory sac) unpaired. Sperm funnel 
conspicuous and deep. Vas deferens thin- 
walled, ciliated, 25-35 um wide, about as 
long as atrial ampulla, but not clearly set off 
from latter. Atrial ampulla about 100-175 
um long, entally dilated, 25-55 wm wide; 
cilia inside not observed (see Remarks). Ec- 
tal part of atrial ampulla narrow, but bear- 
ing oval, heavily granulated diverticulum 
(=modified prostatic pad), up to 50-60 um 
long; however, diverticulum appearing 
poorly developed in some specimens. Large, 
lobed prostate gland attached to this diver- 
ticulum. Atrial ampulla ectally terminating 
in slender, non-granulated, atrial duct, 115- 
140 um long, 13-19 um wide. Atrial ducts 
of both sides joining each other while en- 
tering conspicuous, unpaired ejaculatory 
duct. Ejaculatory duct 100-165 um long, 
45-90 um wide, with heavily muscular, of- 
ten folded, walls; muscle layer maximally 
5-15 wm thick. Ejaculatory duct entering 
subapical, posterior part of compact copu- 
latory sac. This sac, acting as an eversible 
pseudopenis, 75-80 um deep, 65-75 um 
wide, with complex, folded walls [and in at 
least one paratype, bearing a small copu- 
latory gland]. Spermatheca (Fig. 1B) un- 
paired, consisting of (1) an inconspicuous 
vestibule, (2) a short duct, about 60-90 um 
long, 28-38 um wide, (3) a large, thin-walled, 
generally somewhat oval, ampulla, about 
200-280 um long, 80-140 wm wide, and (4) 
a filiform diverticulum, about 260-390 um 
long, 35-45 um wide, attached to inner end 
of ampulla. Sperm in random mass 
throughout spermatheca; mass denser in di- 
verticulum than in ampulla. 

Remarks.—In the whole-mounted spec- 
imens, cilia could not be observed in the 
atrial ampullae; cilia occur there in all con- 
geners. Possibly, the atrial ciliation is re- 
duced in S. exspectatus. 

Distribution and habitat.—Known only 


589 
Q 
Ww 
: D 
= w 
sonniodar? eh is 
rs) 2 xe) 
Ey estas pide at to 222 
® 9 x 8 = = 
= ® DD c = je) 
= = = iS B 
S = = w Ss ® 
= a) € i & = 


9 (state 2) 
6 (state 2) 


11 (state 2) 
8? 


5 (state 2) 
3 


Fig. 2. Smithsonidrilus exspectatus, new species, 
inserted in the most parsimonious cladogram of the 
(most apomorphic) species of Smithsonidrilus (after 
Erséus 1990:fig. 31). Numbers of character and char- 
acter states refer to table VI in Erséus (1990): character 
3, prostatic pads modified into atrial diverticula; 5 (state 
2), ectal part of atrial ducts modified into slender, un- 
paired ejaculatory duct; 6 (state 2), copulatory sac large, 
much folded; 8, copulatory glands present; 9 (state 2), 
spermathecal vestibule large, with folded walls; 10, 
spermathecal glands (associated with vestibule) pres- 
ent; 11 (state 2), spermathecal ampulla bipartite, bear- 
ing filiform diverticulum. 


from the Florida Keys (Atlantic coast of 
southern Florida). Barely subtidal (known 
from 0.1—0.6 m depth), medium to coarse 
sand. 


Discussion 


Smithsonidrilus exspectatus is morpho- 
logically intermediate between S. humme- 
lincki and the group consisting of S. mul- 
tiglandularis, S. marinus, S. involutus and 


590 


S. westoni. Its copulatory (pseudopenial) sac 
is more bulbous and complex than the one 
of S. hummelincki, but not as elaborate as 
those of the others. The other species (but 
not S. hummelincki) all have copulatory 
glands (one or more) associated with their 
pseudopenes; in a single specimen of S. 
exspectatus a small copulatory gland, sim- 
ilar in size and shape to the one of S. in- 
volutus (see Erséus 1990:fig. 36E), was not- 
ed. 

The spermatheca of S. exspectatus is sim- 
ilar to those of S. marinus, S. involutus, S. 
westoni and S. multiglandularis, particular- 
ly with regard to the filiform diverticulum 
(Fig. 1B, sdi), but its vestibule (sv) is not as 
developed as in the four other species. In S. 
hummelincki, the spermatheca lacks a di- 
verticulum. 

If S. exspectatus is inserted in an already 
published cladogram (Erséus 1990:fig. 31; 
see Fig. 2), it will intervene between S. hum- 
melincki and S. multiglandularis, sharing 
with the latter state 2 of character 11 (sper- 
mathecal ampulla with filiform diverticu- 
lum), but not state 2 of characters 6 (cop- 
ulatory sac large, much folded) and 9 
(spermathecal vestibule large, with folded 
walls) (numbers referring to Erséus 1990: 
table VI). As indicated above, it may also 
share character 8 (copulatory gland) with 
the most apomorphic species of Smithson- 
idrilus. 

The paired parts of the atrial ducts of S. 
exspectatus are narrow and not granulated, 
and thus they resemble those of S. hum- 
melincki more than those of most other con- 
geners. Whether this resemblance is syna- 
pomorphic or convergent is uncertain. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


The dilated, heavily muscular ejaculatory 
duct appears autapomorphic for S. exspec- 
tatus. A possible reduction of the ciliation 
in the atrial ampullae (Remarks above) 
would also be unique, but this feature needs 
confirmation on sectioned material. 


Acknowledgments 


I am indebted to Mr. Michael R. Milligan 
(Mote Marine Laboratory, Sarasota, Flori- 
da), for technical assistance, both in the field 
and with the preparation of specimens, and 
for critically reviewing a draft of the manu- 
script; to Ms. Christine Hammar (SMNBH), 
for lettering Fig. 1; and to the Swedish Nat- 
ural Science Research Council, for financial 
support. 


Literature Cited 


Brinkhurst, R. O. 1966. A contribution to the sys- 
tematics of the marine Tubificidae (Annelida, 
Oligochaeta).— Biological Bulletin 130:297-303. 

Erséus,C. 1982. Revision of the marine genus Smith- 

sonidrilus Brinkhurst (Oligochaeta, Tubifici- 

dae). —Sarsia 67:47—-54. 

. 1990. The marine Tubificidae (Oligochaeta) 

of the barrier reef ecosystems at Carrie Bow Cay, 

Belize, and other parts of the Caribbean Sea, 

with descriptions of twenty-seven new species 

and revision of Heterodrilus, Thalassodrilides 

and Smithsonidrilus. — Zoologica Scripta 19:243— 

303. 

Righi, G., & E. Kanner. 1979. Marine Oligochaeta 
(Tubificidae and Enchytraeidae) from the Ca- 
ribbean Sea.—Studies of the Fauna of Curacao 
and other Caribbean Islands 58:44—-68. 


Department of Invertebrate Zoology, 
Swedish Museum of Natural History, Box 
50007, S-104 05 Stockholm, Sweden. 


PROC. BIOL. SOC. WASH. 
106(3), 1993, pp. 591-601 


TWO NEW SPECIES OF PHASCOLION 
(SIPUNCULA: PHASCOLIONIDAE) FROM TROPICAL 
AND SUBTROPICAL WATERS OF THE 
CENTRAL WESTERN ATLANTIC 


Mary E. Rice 


Abstract. —Phascolion (Isomya) gerardi, a new species of Sipuncula, is de- 
scribed from littoral waters of the Bahamas, Belize, and the Yucatan coast of 
Mexico where it inhabits rubble associated with coral reefs. A new interstitial 
species, Phascolion (Phascolion) psammophilum, is described from depths of 
15-19 m off the coast of east central Florida. It is the first sipunculan, docu- 
mented as an adult rather than juvenile, to be reported as a member of the 
meiofaunal community. Observations on development, resulting from labo- 
ratory spawnings, reveal a typical lecithotrophic trochophore and a short-lived 
lecithotrophic pelagosphera larva that transforms into a crawling juvenile at 4 
days of age. Station data for these collections are presented in an appendix at 


the end of the paper. 


In collections of sipunculans from the wa- 
ters of the southeastern coast of Florida, the 
Bahamas, and the Caribbean, two new spe- 
cies of Phascolion have been discovered. 
These collections were made for a system- 
atic survey, currently in progress, of the 
phylum of marine worms, Sipuncula, in the 
western central Atlantic Ocean. Although a 
number of authors have described localized 
collections from these waters (cf. Fischer 
1922; ten Broeke 1925, Murina 1967a, 
1967b; Rice 1975) a comprehensive survey 
of the systematics and distribution of si- 
punculans from south Florida, the Baha- 
mas, and the Caribbean has not been un- 
dertaken. 

The most recent classification of the phy- 
lum Sipuncula is that of Gibbs and Cutler 
(1987) who defined two classes, four orders 
and six families. In a separate treatment of 
the family Phascolionidae, Cutler & Cutler 
(1985) recognized two genera, Phascolion 
and Onchnesoma. In the smaller genus 
Onchnesoma only 4 species are listed, 
whereas in Phascolion 5 subgenera and 23 


species are distinguished. The present paper 
adds two species to the genus Phascolion. 


Family Phascolionidae Cutler & Gibbs, 
1985 
Genus Phascolion Théel, 1875 
Subgenus Jsomya Cutler & Cutler, 1985 
Phascolion gerardi, new species 
Fig. 1 


Material examined. — Bahamas: Berry Is- 
lands (Rice, Sta. 209), 1 specimen, Belize: 
South Water Cay and Curlew Bank (Rice 
Sta. 263, 264), 7 specimens. Mexico: Puerto 
Morelos reef (Rice, Sta. 265, 268), 4 spec- 
imens. 

Holotype: USNM 160243. 

Paratypes: USNM 160244-160248. 

Diagnosis. — Total body length averaging 
17 mm. Thick trunk sometimes cylindrical, 
but more often spherical in living speci- 
mens. Thin, narrow introvert one to two 
times the length of trunk. Prominent pa- 
pillae, mammillate or elongate and cone- 
shaped, distributed closely over the entire 


592 


trunk, largest and most elongate at anterior 
and posterior extremities; larger papillae of- 
ten with multiple tips. Less conspicuous, 
sharply tapered papillae over introvert. Up 
to 24 digitiform tentacles surround mouth. 
Nuchal organ forms corrugated band sur- 
rounding dorsal half of introvert at base of 
tentacular crown. Bulbous expansion of in- 
trovert beneath tentacles bears numerous, 
irregularly arranged simple, curved hooks. 
Ventral retractor muscle divides in mid- 
trunk, each branch attaching to body wall 
on either side of ventral nerve cord in pos- 
teriormost trunk. Single dorsal retractor 
muscle attaches posteriorly at same level as 
ventrals. Dorsal and ventral muscles sepa- 
rate at anterior extremity of introvert, re- 
maining attached by mesenteries to esoph- 
agus for length of introvert. Intestine looped, 
attached to body wall by fixing muscles. 
Long rectum, ending at anus on mid-intro- 
vert. Single right nephridium opening in an- 
terior trunk. Gonad at base of ventral re- 
tractor muscles. 

Description.—Of the 12 specimens on 
which this description is based, the tentacles 
were extended in 10 and the entire introvert 
was retracted in the others. The digitiform 
tentacles, having grooves on the inner sur- 
faces leading to the central mouth, num- 
bered 9 to 24 (Fig. la—c). The total body 
length (introvert plus trunk) ranged from 
approximately 8 to 30 mm; the introvert to 
trunk ratio ranged from 1.0 to nearly 3. 
Larger specimens, as a rule, had more ten- 
tacles than smaller. Depending on the state 
of contraction of the specimen, the region 
of the introvert just posterior to the tentac- 
ular crown may be expanded into a bulbous 
shape. The nuchal organ is a distinctive dor- 
sal band that extends half way around the 
base of the tentacles, becoming narrower 
laterally (Fig. 1c). It is distinguished by the 
numerous irregular folds or longitudinal 
corrugations. The “‘neck’’ or the area ante- 
rior to the bulbous expansion is smooth and 
its posterior border in many specimens ap- 
pears as an undulating fold. Covering the 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


surface of the bulb are scattered hooks. The 
hooks are brown, curved, and simple, av- 
eraging 30 um in basal width and 20 wm in 
height (Fig. 1d). One of the most distinctive 
external features of this species is the form, 
prominence and abundance of the papillae 
(Fig. la, e, f). They cover the entire trunk 
and are commonly largest at the anterior 
trunk or base of the introvert, and next larg- 
est at the posterior extremity. The largest 
papillae are elongate, often cone-shaped with 
two to three pores opening on apical pro- 
trusions; others have a tapered or rounded 
apex and a single pore. Small, sharply ta- 
pered papillae are closely distributed over 
the introvert. The anus is situated dorsally 
in the mid-introvert (Fig. la, g). Internally 
the retractor muscle column separates in the 
anterior introvert into a ventral and a dorsal 
retractor of similar diameter (Fig. 1g). The 
esophagus is enclosed between the two mus- 
cles, to which it is attached by mesenteries, 
for the length of the introvert, separating 
from the dorsal mesenteric connection 
slightly above the ventral separation. The 
ventral retractor muscle splits in mid-trunk 
into two muscles which attach on either side 
of the ventral nerve cord close to the pos- 
terior extremity. A single dorsal muscle at- 
taches posteriorly at about the same level 
as the ventrals. In all specimens dissected 
(6/12) the posterior end of the contractile 
vessel appears as a prominent bulge along 
the esophagus, where the latter presumably 
merges with the intestine. At this level a 
prominent fixing muscle attaches the esoph- 
agus to the dorsal body wall in the mid- 
trunk. The intestine forms several loose 
loops which are attached to the body wall 
by numerous other fixing muscles. The in- 
testine sometimes forms a short spiral of 2 
or 3 coils posteriorly. A long rectum extends 
to the anal opening in the mid-introvert. A 
single right nephridium opens in the ante- 
rior trunk and is attached by mesenteries to 
the body wall for most of its length. A nod- 
ule occurring on the left side at the same 
level as the right nephridiopore may rep- 


VOLUME 106, NUMBER 3 593 


Tessie 


g) 
2 


Fig. 1. Illustration of Phascolion gerardi, new species. a. External anatomy of the holotype. Note anus (An) 
in the mid-introvert region. b. Apical view of tentacles surrounding central mouth. c. Dorsal view of head 
showing nuchal organ (NO) and eyespots (E). d. Introvert hook. e. Enlargement of papillae from anterior trunk. 
f. Englargement of papillae from posterior trunk. Note multiple tips. g. Internal anatomy. Abbreviations. — An, 
anus; CV, contractile vessel; DRM, dorsal retractor muscle; E, eyespot; Es, esophagus; FM, fixing muscle; G, 
gonad; In, intestine; N, nephridium; NO, nuchal organ; R, rectum; RN, rudimentary nephridium; SB, saccular 
body; VNC, ventral nerve cord; VRM, ventral retractor muscle. (Illustrator: Charissa Baker) 


594 


resent a rudimentary left nephridium. The 
gonadal fringe at the bases of the ventral 
retractor muscles has well developed lob- 
ules and in some specimens extends as a 
strand between the two muscles posterior 
to the end of the ventral nerve cord. 
Remarks.—Phascolion gerardi is placed 
in the subgenus Jsomya recently created by 
Cutler & Cutler (1985) to include those spe- 
cies in which dorsal and ventral retractors 
are approximately equal in diameter and in 
which the esophagus departs from the re- 
tractor column posterior to the separation 
of the muscles. In Phascolion gerardi the 
retractor muscle column divides into dorsal 
and ventral muscles in the anterior intro- 
vert; these two muscles, of similar diameter, 
are attached by mesenteries to the esopha- 
gus, which lies between them, for the length 
of the introvert. In the anterior trunk or 
posterior introvert the mesenteric connec- 
tion ends and the muscles become separate 
from each other and from the esophagus. 
A distinguishing feature of Phascolion 
gerardi is the arrangement and prominence 
of the papillae. In this character, it most 
closely resembles Phascolion tuberculosum 
Théel, 1875, which also is in the subgenus 
Isomya. Papillae cover the trunk and are 
largest at its posterior and anterior extrem- 
ities. The largest papillae are elongate and 
may have two to three pores on apical pro- 
trusions, whereas those in the mid-body are 
usually mammillate and rounded, with a 
single pore. Holdfasts, common in many 
members of this genus, are lacking in P. 
gerardi and, according to Théel’s descrip- 
tion (Théel, 1905), also in P. tuberculosum. 
However, Cutler & Cutler (1985), in a re- 
evaluation of this character, describe pa- 
pillae in the midsection of the latter species 
as flattened spheres, resembling holdfasts 
without chitinization. Such a condition is 
not apparent in P. gerardi. Other differences 
include the shape of tentacles and position 
of the anus. The tentacles of P. tuberculo- 
sum are short and rounded and the anus 
opens in the anterior trunk, whereas the ten- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


tacles of P. gerardi are more elongate and 
digitiform and the anus opens midway along 
the introvert. 

Unlike most species of Phascolion, P. ge- 
rardi does not inhabit shells or construct 
tubes, nor does it possess the modified pap- 
illary holdfasts usually associated with such 
habitats. Specimens at all stations were 
found in cavities in coralline limestone. This 
species differs from many other rock-dwell- 
ing sipunculans in that it does not appear 
to create the burrow in which it lives, but 
rather inhabits pre-existing holes in the 
rocks. When the rock is fractured, the spec- 
imens, usually contracted and spherical, roll 
out from their exposed burrows. 

Nothing is known about reproductive bi- 
ology in this species, but on one occasion 
spawned eggs were observed. A specimen 
from the Bahamas, Sta. 209, kept alive in 
the laboratory, spawned eggs 12 days after 
collection. The eggs were spherical, opaque 
and white and enclosed by a thin egg en- 
velope. The average diameter (” = 50) was 
257 um with a range of 252—263 um. 

Etymology.—This species is named for 
the Honorable Sumner Gerard in appreci- 
ation of his gracious and generous help in 
our collecting efforts. It was on a cruise 
aboard his research vessel, Morning Watch, 
in the Berry Islands, Bahamas, that the first 
specimen of this species was discovered. 


Family Phascolionidae Cutler & Gibbs, 
1985 
Genus Phascolion Théel, 1875 
Subgenus Phascolion (sensu stricto) 
Théel, 1875 
Phascolion psammophilum, new species 
Figs. 2-4. 


Holotype: USNM 160249. 

Paratypes: USNM 160250-160253. 

Material examined.—Florida: off Fort 
Pierce, central east coast (Rice, Stas. 162, 
230, 230A, 236, 237, 254, 255, 255A, 256), 
221 specimens. 

Diagnosis.—An interstitial sipunculan, 


VOLUME 106, NUMBER 3 


averaging 4 mm in length. Introvert ap- 
proximately twice the length of trunk. Ten- 
tacular crown having two to eight elongate 
tentacles, followed by bulbous expansion 
bearing 1—4 irregular rings of curved hooks. 
Rounded papillae over surface of trunk; 
smaller and more pointed papillae on in- 
trovert. Anus on posterior 20% of introvert. 
Retractor muscle column divides in ante- 
riormost trunk into thick dorsal retractor 
and thin ventral retractor, both of which 
attach to body wall near posterior end of 
trunk. Ventral retractor splits in posterior 
extremity before attachment. Esophagus re- 
mains attached for short distance to ventral 
retractor before continuing into loosely 
looped intestine. Single right nephridium. 
Pair of saccular bodies on either side of ven- 
tral nerve cord near level of nephridiopore. 

Description.—Unlike most sipunculans 
in which the concave curvature of the body 
is ventral, this species manifests a dorsal 
concave curvature. The introvert is elongate 
and slender, whereas the trunk, depending 
on its state of contraction, is thickened and 
has a rounded posterior end. The anus opens 
on the introvert at a point about 20% of the 
total length of the introvert from its base. 
Measurements of total body lengths of 65 
specimens from 10 stations, with introverts 
and tentacles extended, averaged 4 mm 
(+1.0) with minimum and maximum 
lengths of 1.5—7 mm; the average ratio of 
introvert to trunk was 1.7:1 (1.0:1—2.4:1) 
The width varied depending on the state of 
contraction, but the maximum body width 
of the largest specimens was 0.5 mm. The 
elongate introvert terminates in a tentacular 
crown with central mouth and 2-8 tentacles 
(Fig. 2a—e). In smaller specimens, under 2 
mm, the number of tentacles is from two to 
four, whereas in those 3 mm and over the 
number may range from three to eight. As 
seen in the smallest specimens, the single 
pair of tentacles is dorsal, whereas in those 
specimens with four tentacles there is, in 
addition, a ventral pair (Fig. 2b, c). In spec- 
imens with six tentacles, there are four dor- 


395 


sals and two ventrals and in those with eight 
tentacles there are two additional ventrals 
(Fig. 2d, e). On the dorsal side of the ten- 
tacular crown at the terminal extremity of 
the animal, there is a flattened pad, revealed 
by SEM to be heavily ciliated, that is pre- 
sumed to be the nuchal organ (Fig. 3a, b). 
SEM also shows the inner surfaces of the 
tentacles to be grooved and ciliated (Fig. 
3b). Posterior to the tentacles, a short neck 
region is followed by a bulbous expansion 
of the anterior introvert which bears from 
two to four irregular rows of large curved 
hooks, the largest of which are 30 um in 
height and 30 um in basal width. In smaller 
specimens there may be only one row. The 
hooks are brownish in color, strongly curved 
and bluntly pointed (Fig. 2f). Scattered 
among the hooks are small dome-shaped 
papillae (Fig. 3c). The papillae on the re- 
mainder of the introvert are more pointed 
and prominent. Scanning electron micro- 
graphs reveal a central, apical pore through 
which ciliary extensions sometimes pro- 
trude. Larger, more rounded papillae cover 
the surface of the trunk, being most con- 
centrated in the anterior trunk (Fig. 3d). 
Apical regions of papillae often appear to 
be differentiated as knobs or caps (Fig. 3c, 
d). Papillary caps are present on some of the 
trunk papillae, but in those without caps or 
apical elevations two to three central pores 
are occasionally visible. Demarcation be- 
tween introvert and trunk is usually indi- 
cated by a constriction and in SEMs by a 
decrease in the circular undulations of the 
cuticle of the trunk. Holdfast papillae, pres- 
ent in most species of Phascolion, are ab- 
sent. 

Internally, the undivided retractor mus- 
cle column extends for the length of the in- 
trovert and separates in the anterior third 
of the trunk, usually close to the base of the 
introvert, into dorsal and ventral branches. 
These two branches attach to the body wall 
near the posterior extremity at approxi- 
mately the same level (Fig. 2g). The thick- 
ness of the dorsal muscle is two to four times 


596 


Fig, 2. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


L Laker 


Illustration of Phascolion psammophilum, new species. a. External anatomy. b-e. Ontogenetic se- 


quence of tentacle development from two to eight tentacles. Apical view; dorsal side at top. f. Introvert hook. 
g. Internal anatomy. Abbreviations. —See Fig. 1. (Illustrator: Charissa Baker) 


greater than that of the ventral. The ventral 
muscle usually splits near its point of at- 
tachment, straddling the end of the thick 
nerve cord. Gonadal tissue extends across 
the base of the ventral muscles. The esoph- 
agus is attached to the anterior ventral re- 


tractor muscle for a short distance posterior 
to its separation from the dorsal retractor. 
The swollen posterior end of the contractile 
vessel is apparent along the esophagus near 
its position of separation from the muscle. 
The intestine is not coiled, but is loosely 


VOLUME 106, NUMBER 3 


Fig. 3. 


\ 
Ry Tie ae 


Ld 


Scanning electron micrographs of Phascolion psammophilum, new species. a. Whole animal. Left 


side is dorsal. Scale bar, 150 um. b. Lateral view of anterior introvert showing 6 tentacles, dorsal nuchal organ 
(arrow), and hooks on bulbous introvert. Scale bar, 150 wm. c. Papillae from anterior introvert. Note central 
pores. Scale bar, 10 um. d. Papillae from mid-trunk. Some have papillary cap. Scale bar, 10 um. 


looped and attached to the body wall by a 
few fine fixing muscles. A caecum is present 
at the end of the intestine, and a long rectum 
extends into the introvert where it opens in 
a dorsal anus. A single right nephridium, 


free from the body wall for most of its length, 
attaches at the level of the nephridiopore in 
the anterior one-third of the trunk. Slightly 
anterior to this attachment is a pair of sac- 
ciform bodies on either side of the ventral 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Developmental stages of Phascolion psammophilium, new species. a. Gastrula. b. Trochophore. c. 


Pelagosphera larva, 112 days. d. Juvenile, 442 days. Abbreviations.—P, prototrochal cilia; M, metatrochal cilia; 
H, hook. Scale bars, 50 um. 


nerve cord, the right member of the pair pores. The thick, prominent nerve cord ends 
being lateral to the nephridium. Histologi- posteriorly just above the attachment of the 
cal sections demonstrate that these bodies _ ventral retractor muscle. 

are glandular and open to the exterior by Reproduction and development. —The 


VOLUME 106, NUMBER 3 


presence of mature gametes in the coelomic 
fluid is evidence that these specimens are 
adults rather than juveniles, as might be 
suggested by their small size. The popula- 
tion is dioecious. Spawnings of eggs and 
sperm have been observed in the laboratory 
in the months of April, May, August and 
December. 

Recently spawned, unfertilized eggs are 
slightly elongate, averaging 124 x 111 wm 
(n = 40), and enclosed by relatively thin egg 
envelopes that, as is true for all sipunculan 
eggs, are perforated by fine pores (Fig. 4a). 
Within 24 hours after fertilization a typical 
trochophore, having a wide band of pro- 
totrochal cilia, apical tuft, and dorsal eye- 
spots, develops within the egg envelope (Fig. 
4b). By 36 hours the trochophore has trans- 
formed into an elongated pelagosphera lar- 
va with a well-developed band of metatro- 
chal cilia (Fig. 4c). The larva swims near the 
bottom of the laboratory container or moves 
along on the bottom in a geometrid pattern 
of locomotion, similar to that of an inch- 
worm. Within 4 days the pelagosphera larva 
metamorphoses into a juvenile worm with 
fully formed gut. The presence of numerous 
yolk granules in the coelomic cavity sug- 
gests that the larva is lecithotrophic. The 
young juvenile has the shape characteristic 
of the adult, with elongate retractable intro- 
vert and anterior hooks (Fig. 4d). 

Remarks.—Previous to the collections 
and observations of Phascolion psammo- 
philum reported in this paper, sipunculans 
have been generally considered as only tem- 
porary inhabitants of the meiofaunal com- 
munity, represented by numerous uniden- 
tifiable juveniles. However, as demonstrated 
by the above observations on gametes and 
reproduction, this species, in its adult stage, 
is an integral member of the meiofaunal 
community. It has been previously referred 
to as an unidentified interstitial species (Rice 
1988, 1993). Only one other sipunculan, As- 
pidosiphon exiguus, has been reported as a 
member of the interstitial community (Ed- 
monds 1974, 1982). Although specimens 


599 


were presumed to be adults, gametes were 
not observed. 

Phascolion psammophilum was discov- 
ered during studies of sediments on the con- 
tinental shelf off the central east coast of 
Florida. It occurs at depths from 15 to 19 
meters in sediments characterized as me- 
dium to coarse sand and shell hash. Unlike 
most species of this genus, P. psammophi- 
lum does not occupy shells or tubes, but is 
found moving among the sand grains. 

The arrangement and fusion of retractor 
muscles and their relation to the esophagus 
are characteristic of the subgenus Phasco- 
lion, as defined by Cutler and Gibbs (1985). 
In two other species of this genus, P. hupferi 
and P. gerardi, the anus opens on the intro- 
vert; however, these species differ in many 
other characters and are placed in two other 
subgenera, respectively, Lesenka and Iso- 
mya. 

Etymology.—The name psammophilum 
refers to the sand habitat in which this spe- 
cies is found. (Gr. psammos, sand; Gr. phi- 
los, having affinity for.) 


Acknowledgments 


I am grateful to Julianne Piraino for her 
many efforts on behalf of this project, in- 
cluding collecting specimens, operating the 
SEM, and preparing the photographic plates. 
Special thanks go to Hugh Reichardt for 
collection of specimens, maintenance of de- 
velopmental cultures, and, along with Wil- 
liam Lee, operating the boats for offshore 
collections. Specimens of Phascolion psam- 
mophilum were sorted and generously con- 
tributed by Robert P. Higgins, along with 
Jon Norenburg, Edward Ruppert and Sher- 
ry Reed. Charissa Baker is respectfully ac- 
knowledged for rendering the illustrations, 
Figs. 1 and 2. John Pilger is thanked for 
assistance in collecting specimens from the 
Caribbean. The Honorable Sumner Gerard 
is acknowledged with genuine gratitude for 
support and encouragement during this pro- 
ject and for providing his ship Morning 


600 


Watch for collections in the Bahamas. 
Smithsonian Marine Station at Link Port 
Contribution #327. Caribbean Coral Reef 
Ecosystems Program Contribution #388. 


Literature Cited 


Cutler, E. B., & N. J. Cutler. 1985. A revision of the 
genera Phascolion Théel, and Onchnesoma Ko- 
ren and Danielssen (Sipuncula).— Proceedings 
of the Biological Society of Washington 98:809- 
850. 

—., & P.E. Gibbs. 1985. A phylogenetic analysis 
of higher taxa in the phylum Sipuncula.—Sys- 
tematic Zoology 34:162-173. 

Edmonds, S. J. 1974. A new species of Sipuncula 

(Aspidosiphon exiguus n.sp.) belonging to the 

interstitial fauna of marine beaches collected by 

Mr. L. Botosaneanu during the second Cuban- 

Rumanian Biospeleological Expedition to Cuba 

1973.—International Journal of Speleology 

6:187-192. 

1982. A sipunculan reported to be “‘intersti- 
tial’? from the Netherland Antilles.— Bijdragen 
tot de Dierkunde 52:228-230. 

Fischer, W. 1922. Westindische Gephyreen.—Zoolo- 
gischen Anzeiger 55:10-18. 

Gibbs, P. E., & E. B. Cutler. 1987. A classification 
of the phylum Sipuncula.— Bulletin of the Brit- 
ish Museum (Natural History) Zoology 52:43- 
58. 

Murina, V. V. 1967a. Report ofthe sipunculid worms 

from the sub-littoral zone of Cuba and the Mex- 

ican Gulf.—Zoologicheskiy Zhurnal 46:1329- 

1339. (In Russian). 

. 1967b. On the sipunculid fauna of the littoral 

of Cuba. — Zoologicheskiy Zhurnal 46:35—46. (In 

Russian). 

Rice, M.E. 1975. Survey ofthe Sipuncula of the coral 

and beachrock communities of the Caribbean 

Sea. Pp. 35-49 in M. E. Rice & M. Todorovic, 

eds., Proceedings of the International Sympo- 

sium on the Biology of the Sipuncula and 

Echiura, Vol. I, Naucno Delo, Belgrade, 355 pp. 

1988. Sipuncula, Chapter 32. Pp. 355-356 
in R. P. Higgins & H. Thiel, eds., Introduction 
to the study of meiofauna. Smithsonian Insti- 
tution Press, Washington, D.C. and London, 488 
pp 


1993. Sipuncula, Chapter 7. In F. W. Har- 
rison and M. E. Rice, eds., Microscopic anat- 
omy of invertebrates, Volume 12: Onychopho- 
ra, Chilopoda, and Lesser Protostomata. Wiley- 
Liss, New York (in press). 

ten Broeke, A. 1925. Westindische Sipunculiden und 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Echiuriden.—Bijdragen tot de Dierkunde, 24: 
81-96. 
Théel, H. 1875. Recherches sur le Phascolion (Phas- 
colosoma) strombi (Mont.).—Bihang Kungl. 
Svenska Vetenskaps-Akademens Handlingar 
3:1-7. 

1905. Northern and Arctic invertebrates in 
the collection of the Swedish State Museum. I. 
Sipunculids.—Bihang Kungl. Svenska Veten- 
skaps-Akademens Handlingar 39:1—130. 


Smithsonian Marine Station at Link Port, 
5612 Old Dixie Highway, Fort Pierce, Flor- 
ida 34946, U.S.A. 


Appendix: Station Data 


Station numbers listed below are from the station 
records of Mary E. Rice for her collections in Florida 
and the Caribbean. 


Station 162—27°31.0'N, 80°08.3'’W, 8.7 miles E from 
Fort Pierce, FL, collected by pipe dredge on Aquarius 
Cruise 7 at 15 m depth, 9 May 1978 

Station 209—Frazer’s Hog Cay, Berry Islands, Baha- 
mas, Morning Watch Cruise, collected by hand at 1 
m depth, 3 March 1982, in calcareous sandstone and 
coralline limestone 

Station 230—27°29.9'’N, 80°11.4’W, 5.6 miles E from 
Fort Pierce, FL, collected by anchor dredge on Tur- 
siops Cruise 24 at 15 m depth, 6 Dec 1982, in coarse 
sand with shell fragments 

Station 230A—27°30.9'N, 80°12.2’W, 5.4 miles NE 
from Fort Pierce, FL, collected by anchor dredge on 
Tursiops Cruise 25 at 15 m depth, 31 Jan 1983, in 
coarse sand with shell fragments 

Station 236—27°30.5'N, 80°12.0’W, 5.3 miles NE of 
Fort Pierce, FL, collected by sled dredge on Snook 
Cruise 80 at 15 m depth, 24 Feb 1983, in crushed 
shell and fine quartz sand 

Station 237 —27°30.3’N, 80°12.0’W, 5.3 miles E of Fort 
Pierce, FL, by sled dredge on Snook Cruise 80 at 15 
m depth, 24 Feb 1983, in crushed shell and fine 
quartz sand 

Station 254—27°33.3'N, 80°10.8’W, 7.8 miles NE of 
Fort Pierce, FL, collected by anchor dredge on Tur- 
siops Cruise 30 at 19 m depth, 6 Dec 1983, in me- 
dium sand and shell hash 

Station 255—27°32.3'N, 80°10.6’W, 7.3 miles NE of 
Fort Pierce, FL, collected by anchor dredge on Tur- 
siops Cruise 30 at 18 m depth, 6 Dec 1983, in sand 
and shell hash 

Station 255A—27°32.3'N, 80°10.7'N, 7.3 miles NE of 
Fort Pierce, FL, collected by anchor dredge on Tur- 
siops Cruise 31 at 15 m depth, 6 Mar 1984, in sand 
and shell hash 

Station 256—27°31.2’N, 80°10.1’W, 7.2 miles NE of 


VOLUME 106, NUMBER 3 


Fort Pierce, FL, collected by anchor dredge on Tur- 
siops Cruise 30 at 17 m depth, 6 Dec 1983, in shell 
hash 

Station 263—north of South Water Cay, west of To- 
bacco Reef, Belize, collected by sled dredge at 1 m 
depth, 26 June 1985, in rubble on sandy backreef 
area 

Station 264—Curlew Bank, Belize, collected by hand 
in less than 1 m depth, 26 June 1985, in coral rubble 


601 


Station 265— Puerto Morelos, Quintana Roo, Mexico, 
collected by hand while snorkeling at 1 m depth, 3 
July 1985, in coral rubble taken from reef crest area 

Station 268—Cabeza Reef, Puerto Morelos, Quintana 
Roo, Mexico, collected by hand while scuba diving 
at 9 m depth, 5 July 1985, in coral rubble from reef 


PROC. BIOL. SOC. WASH. 
106(3), 1993, pp. 602-605 


AN UNUSUAL SQUID PARALARVA (CEPHALOPODA) WITH 


TENTACULAR PHOTOPHORES 
Deborah L. Loffler and Michael Vecchione 


Abstract.— We describe an unusual squid paralarva from the eastern North 
Pacific. This paralarva has characters indicating it might be a gonatid, but if 
so it is the first gonatid species known to have photophores on its tentacles. 
Structures that might be photophores were also located on the interior of the 
mantle. One other gonatid species, Gonatus pyros, is known to have photo- 
phores but these are located only on the eyes. Gonatus pyros spawns in the 
eastern North Pacific, and its early paralarva has not been described. We there- 
fore tested the hypothesis that our squid may be an ontogenetic stage of G. 
pyros. Tentacles of young G. pyros were examined histologically to see if pho- 
tophore tissue might be embedded in the tentacles, but the results were incon- 
clusive. We also examined early stages of eight other gonatid species. None of 
these had either tentacular or internal photophores. We therefore describe this 


very unusual specimen, but its identity remains uncertain. 


During examination of a collection of 
cephalopods from the eastern North Pacific 
we found a paralarva with characters of the 
squid family Gonatidae, but also with what 
appeared to be a single photophore on each 
tentacle and two more photophores on the 
interior wall of the mantle just anterior to 
the bases of the fins. Only one gonatid spe- 
cies, Gonatus pyros Young, 1972, is known 
to possess photophores, and in that species 
a large photophore is located on the ventral 
surface of each eye (Young 1972). The pres- 
ence of photophores in locations other than 
the eyes on this paralarva is therefore note- 
worthy. 

Few small gonatid paralarvae (<10 mm 
mantle length, ML) have been described, 
and little is known about them. Kubodera 
& Jefferts (1984), in a study of distribution 
and abundance of the early life stages of 
North Pacific squids, found only three Go- 
natus pyros specimens <10 mm ML. Be- 
cause G. pyros was omitted from descrip- 
tions and keys to early paralarvae <10 mm 
ML by Kubodera & Okutani (1981) and 
Okutani & Clarke (1992), we hypothesized 


that our specimen may be an early stage of 
that species. If G. pyros were shown histo- 
logically to have photophore tissue embed- 
ded in its tentacles then we could argue that 
G. pyros does develop tentacular photo- 
phores early in life and that our specimen 
could be that species. 


Materials and Methods 


The specimen (USNM 817795; ca. 5 mm 
ML) was collected in the eastern North Pa- 
cific, off southern California at 34°N, 121°W 
on 13 November 1969. Sampling time was 
1730-1800 and depth was approximately 
20S: m 

We examined the possibility that the 
specimen was a Gonatus pyros because this 
is the only photophore-bearing gonatid 
known, and it is found in abundance in the 
sampled area (Kubodera & Okutani 1981). 
Examination of gross tentacular morphol- 
ogy of the smallest G. pyros specimens in 
the collections of the National Museum of 
Natural History revealed no photophores 
but a damaged area was consistently found 


VOLUME 106, NUMBER 3 


at approximately the same location as the 
photophores on our paralarva. The tenta- 
cles of three G. pyros, (USNM 727486, ML 
32 mm, 17 mm, and 16 mm) were examined 
histologically for embedded photophores. 
These specimens had been fixed in formalin 
and preserved in 45% isopropanol. Histo- 
logical preparation included ethanol dehy- 
dration, tissue embedding in paraffin, and 
frontal sectioning at 6 wm thickness. He- 
matoxylin and eosin stains were used and 
the sections were examined using light mi- 
croscopy. Other gonatids from the National 
Museum of Natural History were examined 
for the presence of photophores. These were 
the smallest specimens in the collection for 
each species listed below and included: Ber- 
ryteuthis anonychus (USNM 575631, ML 
68 mm), Gonatopsis borealis (USNM 
813458, ML 33 mm and 28 mm), Gonatus 
onyx (USNM 730044, ML 30 mm and 
USNM 730043, ML 17 mm), Gonatus ma- 
dokai (USNM 884252, ML 15 mm and 15 
mm), Gonatus fabricii (USNM 884254, ML 
18 mm and 19 mm), Gonatus berryi (USNM 
729867, ML 12 mm and 12 mm), Berry- 
teuthis magister (USNM 814633, ML 17 
mm and 21 mm), Gonatus antarcticus 
(USNM 884253, ML 42 mm). 


Results 


The specimen was identified to the family 
Gonatidae using a paralarval key to the 
cephalopod families (Sweeney et al. 1992). 
Identification was based on the following 
characters: arms with four rows of suckers, 
robust tentacles with minute suckers along 
most of the tentacular stalk and distinctive 
tentacular clubs developing >4 rows of 
minute suckers. 

General morphology (Fig. 1).— Mantle: 
Inverted but bell-shaped, thin, muscular; fins 
small, membranous; Mantle Width esti- 
mated to be ca. 60% Mantle Length (ML). 
Funnel-locking cartilage straight. Two small 
spheres, possibly photophores, located on 
interior wall of mantle just anterior to bases 


603 


SS [ee ee eee mn cow? 


Fig. 1. Ventral view of gonatid paralarva. Mantle 
shape estimated from inverted mantle. Scale bar = 2 
mm. 


604 


1mm 


Fig. 2. Ventral view of (left to right) arm IV, ten- 
tacle, and arm III on right side. Scale bar = 1 mm. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


of the fins, connected to a smaller centrally 
located sphere. 

Head: Squarish in shape, slightly narrow- 
er than mantle opening; large anterolateral 
eyes. No photophore anlagen on ventral 
surface of eyes. 

Arms: Stout, muscular. Arm formula 
I=II=III>IV. Suckers tetraserial on arms 
I, If and III. No suckers on arms IV. 

Tentacles (Fig. 2): Stout, muscular, ca. four 
times longer than longest arms, about 3.5 
mm long; smaller suckers cover most of oral 
surface of stalk, in two staggered rows prox- 
imally, increasing to five rows on distal stalk; 
club developing on distal ’s with numerous 
minute suckers in seven rows. Large pho- 
tophore on aboral surface ca. % of tentacle 
length from base. Photophore diameter 
about equal to tentacle diameter. Photo- 
phore round and distinctly darker in shade 
than tentacle. Photophore roughly spherical 
and partially embedded in tentacular mus- 
cle. 

Digestive gland: Oblong with ink sac em- 
bedded in ventral surface. 

When the specimen was stained with 
Methylene Blue, all tissues stained except 
the structures we presume to be photo- 
phores. Histological examination of G. py- 
ros tentacles failed to reveal photophore tis- 
sue. No photophores, either tentacular or 
internal, were found on any of the other 
gonatids examined. Photophores on the 
paralarva were not sectioned because of the 
uniqueness of the specimen. 


Discussion 


The specimen has familial characters to 
classify it as a probable gonatid squid. These 
characters include: arms with four rows of 
suckers, straight funnel-locking cartilage and 
tentacular clubs with >4 rows of minute 
suckers. Other squid families have species 
with tentacular and/or internal photo- 
phores. The Lycoteuthidae have both. But 
no family other than Gonatidae is known 
to have the sucker patterns of this specimen. 


VOLUME 106, NUMBER 3 


Its size of 5S mm ML places it in the size 
range least known for gonatids. The pres- 
ence of tentacular (and perhaps internal) 
photophores, however, is without prece- 
dence in the family. 

Gonatus pyros is the only species in the 
family Gonatidae that has been described 
to have photophores. The photophores of 
G. pyros are, however, in a different location 
than the ones on our specimen. Gonatus 
pyros has a large oval photophore on the 
ventral surface of each eye (Young 1972) 
whereas our gonatid paralarva lacks ocular 
photophores, but has one large round pho- 
tophore on the aboral surface of each ten- 
tacular stalk and possibly a complex of pho- 
tophores in the posterior mantle cavity. The 
location where the specimen was collected 
is in the general region where larger para- 
larvae of G. pyros have been found (Ku- 
bodera & Jefferts 1984). Gonatus pyros 
specimens >10 mm ML were found to be 
frequent and abundant off the Oregon and 
Washington coasts between 40-—SO°N and 
140-120°W (Kubodera & Jefferts 1984). 

Our histological examination was incon- 
clusive. Our inability to locate photophore 
tissue does not mean that it was not there 
at an earlier stage. Absorption of paralarval 
photophores has been suspected in the om- 
mastrephid squid Doscidicus gigas (C. F. C. 
Roper, pers. comm.). The gonatid pho- 
tophore tissue could have been completely 
resorbed by 16 mm ML. If so, this could 
explain the apparent weak spot in the ten- 
tacles of the small G. pyros we examined. 
In other words, we failed to support the hy- 
pothesis that our paralarva is an early stage 
of G. pyros but we do not feel that it has 
been refuted conclusively. Because the his- 
tological examination of G. pyros failed to 
indicate embedded or resorbed photo- 
phores, we have no evidence that our par- 
alarva is that species. Examination of other 
gonatid species did not reveal any charac- 
ters to link our paralarva with any of those 


605 


species. However, the squid treated in this 
paper, although unidentified, remains note- 
worthy in that it appears to be a gonatid 
squid with photophores on the tentacles. 


Acknowledgments 


We thank Clyde F. E. Roper, Smithson- 
ian Institution, for his comments and en- 
couragement. Support was provided by the 
Registry of Tumors and Lower Animals in 
the National Museum of Natural History 
for the histological work performed in this 
investigation. Keiko Hiratsuka Moore of the 
NMFS Systematics Laboratory inked the il- 
lustrations. We appreciate reviews of the 
manuscript by Bruce B. Collette, Michael J. 
Sweeney, Clyde F. E. Roper, Tsunemi Ku- 
bodera and Joseph B. Slowinski. 


Literature Cited 


Kubodera, T., & K. Jefferts. 1984. Distribution and 
abundance of the early life stages of squid, pri- 
marily Gonatidae (Cephalopoda, Oegopsida), in 
the northern Pacific. — Bulletin National Science 
Museum, (Tokyo), Ser. A, 10(4):165-193. 

—, & T. Okutani. 1981. The systematics and 
identification of larval cephalopods from the 
northern North Pacific.— Research Institute of 
North Pacific Fisheries, Hokkaido University, 
Special volume, 131-159. 

Okutani, T., & M. R. Clarke. 1992. Family Gona- 
tidae Hoyle, 1886. In M. J. Sweeney, C. F. E. 
Roper, K. Mangold, M. R. Clarke, & S. v. Bolet- 
zky, eds., ““Larval” and juvenile cephalopods: 
a manual for their identification. —Smithsonian 
Contributions to Zoology 513:139-156. 

Sweeney, M. J., C. F. E. Roper, K. Mangold, M. R. 
Clarke, & S. v. Boletzky, eds., 1992. “‘Larval”’ 
and juvenile cephalopods: a manual for their 
identification.—Smithsonian Contributions to 
Zoology 513:1—282. 

Young, R. E. 1972. The systematics and areal dis- 
tribution of pelagic cephalopods from the seas 
off southern California.—Smithsonian Contri- 
butions to Zoology 97:1-159. 


NMFS Systematics Laboratory, National 
Museum of Natural History, Washington, 
D.C. 20560, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(3), 1993, pp. 606-627 


A REVIEW OF THE GENUS CRITONIOPSIS IN 
CENTRAL AND SOUTH AMERICA 
(VERNONIEAE: ASTERACEAE) 


Harold Robinson 


Abstract.—The genus Critoniopsis Schultz-Bip. is expanded to contain 76 
species and includes the Central American species of Eremosis (DC.) Gleason. 
Additional species from South America are transferred to the genus including 
a number with opposite leaves such as Vernonia stellata of Brazil. Opposite- 
leaved Critoniopsis dorrii, C. palaciosii of Ecuador, and C. quillonensis of Peru, 
subopposite-leaved C. cotopaxensis of Ecuador, and alternate-leaved C. lewisii 
and C. steinbachii of Bolivia and C. uribei of Colombia are described as new. 


A group of neotropical Vernonieae hav- 
ing few florets in the heads and deciduous 
inner involucral bracts has been treated in 
various papers since 1900 under two dif- 
ferent names, Eremosis and Critoniopsis, 
sometimes at the sectional level under Ver- 
nonia Shreb. and at other times as distinct 
genera. The two names have been applied 
regionally (Gleason 1906, 1922; Cuatreca- 
sas 1956; Robinson 1980; Jones 1973), and 
their synonymy has often been casually as- 
sumed. A long overdue broader summary 
is offered here. Critoniopsis 1s treated at the 
generic level with Eremosis as a synonym. 
All the presently accepted members of the 
genus are listed, and all the species not pre- 
viously placed in Critoniopsis are here 
transferred to the genus. 

The first generic name for the group, Tur- 
pinia Lexarza (La Llave & Lexarza, 1824), 
was a later homonym. The oldest valid name 
for the group, Monosis sect. Eremosis DC. 
(1836), was established for three Mexican 
species of which Monosis salicifolia DC. has 
been treated as the lectotype (Gleason 1906, 
1922; Jones 1973). Eremosis was later raised 
to generic rank by Gleason (1906, 1922). 
Critoniopsis Schultz-Bip. was first described 
at the generic level in 1863 to include a 
single Colombian species, C. lindenii 
Schultz-Bip, but the name was subsequently 


used only at a sectional level under Ver- 
nonia by various authors including Baker 
(1873) for Brazilian species, Gleason (1923) 
for Bolivian species, and Cuatrecasas (1956) 
for northern Andean species. The genus Te- 
phrothamnus Schultz-Bip. (1863) was de- 
scribed at the same time as Critoniopsis, but 
was effectively reduced to synonymy by 
Badillo (1983) with the transfer of 7. par- 
adoxa Schultz-Bip. to Critoniopsis. 

The nomenclatural history has resulted in 
the name Eremosis having priority at the 
sectional level and Critoniopsis having pri- 
ority at the generic level. The various au- 
thors using Critoniopsis as a section of Ver- 
nonia did not consider Eremosis, and 
Gleason (1922), in his use of Eremosis as a 
genus, treated Critoniopsis as a doubtful 
synonym. Still, none of the authors provid- 
ed any differences on which separate sec- 
tions or genera could be based. In the rees- 
tablishment of Critoniopsis at the generic 
level by Robinson (1980), the possible syn- 
onymy of Eremosis was not a factor no- 
menclaturally and was not mentioned. 
Among recent authors, only Jones (1973) 
clearly stated that Critoniopsis and Ere- 
mosis were synonyms, and he used the ap- 
propriate name at the sectional level, Ver- 
nonia section Eremosis. 

The present paper accepts that the two 


VOLUME 106, NUMBER 3 


groups, Critoniopsis and Eremosis, are syn- 
onyms, and that the combined group de- 
serves generic status. There are some gen- 
eral differences in aspect of the 
Mesoamerican material, with its narrower 
heads and narrower involucral bracts, more 
often setuliferous achenes, slightly pointed 
hairs on the style branches, and the more 
often pubescent upper leaf surfaces. Nev- 
ertheless, intergradations and exceptions in 
the characters preclude recognition of the 
Mesoamerican material as a separate genus. 

Robinson (1980) emphasized the com- 
paratively close relationship of Critoniopsis 
to Piptocarpha R. Br., and Robinson et al. 
(1980) established the subtribe Piptocar- 
phinae for the three genera, Piptocarpha, 
Pollalesta H.B.K., and Critoniopsis. Of these 
three genera, Pollalesta is distinguished by 
the modified pappus of straps and scales 
rather than capillary bristles. The style base 
of the latter also seems to characteristically 
lack a sclerified basal node. Piptocarpha has 
traditionally been distinguished by having 
tails on the anthers, but the most significant 
difference is the sclerified and sharply point- 
ed nature of those tails. Some species pres- 
ently placed in Piptocarpha, like P. lusch- 
nathii Krasch. (Stifftia axillaris Barroso & 
G. da Vinha) have blunt-tipped sclerified 
tails. Basal tails are actually present in many 
species of Critoniopsis, but the cells below 
the polleniferous part of the anther are thin- 
walled, and the tip is often distinctly toothed. 
Two additional genera that have been more 
recently recognized as members of the Pip- 
tocarphinae, the opposite-leaved Josean- 
thus H. Robinson and the alternate-leaved 
Cuatrecasanthus H. Robinson, both have 
corollas that have lobes separated to the base 
of the limb at the level of insertion of the 
anther filaments. Thus the corollas have no 
throat. Huberopappus Pruski, Ekmania 
Gleason, and Gorceixia Baker were referred 
to the Piptocarphinae by Pruski (1992). The 
three genera all have a strongly coroniform 
pappus. The most recently recognized 
member of the subtribe, Dasyandantha H. 


607 


Robinson, has numerous hairs on the inner 
and outer surfaces of the corolla throat. 
The broad interpretation here of the ge- 
nus Critoniopsis includes almost all oppo- 
site-leaved Vernonieae of the western 
Hemisphere except those in the closely re- 
lated genus Joseanthus (Robinson, 1989). 
One exception is Lepidaploa canescens var. 
opposita (H. Robinson) H. Robinson (1990a) 
of northern Colombia. It should be noted 
that, although opposite leaves are a tenden- 
cy within Critoniopsis, few of the species of 
the genus having the characteristic seem to 
be immediate relatives of each other. The 
number of florets in the Critoniopsis head 
is usually less than 10, but it has previously 
been noted to be as high as 15 or 16 in C. 
pallida (Cuatrecasas 1956), and it is here 
extended to about 20 by the inclusion of 
Vernonia harlingii H. Robinson. The non- 
glanduliferous anther appendages with or- 
nate thickenings in their cell walls, which 
have been noted as a characteristic of the 
subtribe (Robinson 1992b), furnish a par- 
ticularly sharp delimitation of the Mexican 
members of Critoniopsis from species of 
Vernonanthura H. Robinson (1992a) of the 
subtribe Vernoninae in that area. Crito- 
niopsis proves to include all the species list- 
ed for Vernonia section Eremosis by Jones 
(1973) and Jones & Stutts (1981) along with 
two species not placed in the section by those 
authors, Vernonia autumnalis McVaugh and 
V. tequilana Jones & Stutts. Nevertheless, 
there are some southern species in this al- 
liance from Bolivia and Brazil that lack ob- 
vious thickenings in the cells of their anther 
appendages. Thickenings prove to be weak 
or lacking in other members of the Pipto- 
carphinae, such as Pollalesta. A few thin 
anther appendages with one gland at the 
base have been seen in a specimen of Pip- 
tocarpha triflora (Aubl.) Benn. ex Baker, but 
the gland is not on the appendage. Southern 
species of Critoniopsis having unthickened 
appendages can be distinguished from Ver- 
nonanthera by the deciduous inner invo- 
lucral bracts and by the many small glands 


608 


or small hairs on the corolla. Vernonan- 
thura also tends to have distinctive resin 
ducts filling the centers of the corolla lobes 
(Robinson 1992a) that are lacking in C7i- 
toniopsis. 

The genus Critoniopsis and its species are 
summarized as follows. 


Critoniopsis Schultz-Bip., Jahresber. Polli- 
chia 20/21:430. 1863. 
Type: Critoniopsis lindenii Schultz-Bip. 
Turpinia Lexarza in La Llave & Lexarza, 
Nov. Veg. Descr. 1:24. 1824 (non H.B.K., 
1807, nom. rej.; non Ventenat, 1807, nom. 
cons). Type: 7. tomentosa Lexarza. 
Monosis sect. Eremosis DC., Prodr. 5:77. 
1836. Lectotype: Monosis salicifolia DC. 
Tephrothamnus Schultz-Bip., Jahresber. 
Pollichia 20/21:431. 1863. Lectotype: T. 
paradoxus Schultz-Bip., chosen here. 
Vernonia sect. Critoniopsis (Schultz-Bip.) 
Benthy dé Hook.f. Gene P2230. 13873. 
Eremosis (DC.) Gleason, Bull. New York 
Bot. Gard. 4:227. 1906. 


Shrubs or trees to 13 m tall; stems and 
leaf undersurfaces densely pilosulous to to- 
mentose, rarely glabrous (C. glandulata); 
stems of alternate-leaved species often de- 
flected at nodes. Leaves simple, alternate or 
opposite, petiolate or rarely sessile (C. har- 
lingii, C. sagasteguii); blades often coria- 
ceous, broadly ovate to elliptical or obovate, 
base cuneate to slightly cordate, margins en- 
tire or remotely denticulate to serrulate dis- 
tally, apices obtuse to acuminate, upper sur- 
face glabrous to tomentose, surfaces 
sometimes with stellate hairs; venation pin- 
nate. Inflorescence terminal on leafy 
branches, usually pyramidally thyrsoid, with 
corymbose to subcymose branches. Heads 
homogamous discrete, sometimes crowded; 
involucral bracts slightly to strongly coria- 
ceous, subimbricate to imbricate in 4—6 se- 
ries, appressed, inner bracts easily decidu- 
ous, sometimes with strongly recurved basal 
margins, distal margins often split at ma- 
turity, rarely with a white marginal flange 
(C. harlingii); receptacle epaleaceous. Flo- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


rets mostly 2-11 (15 or 16 in C. pallida, ca. 
20 in C. harlingii); corollas regular, white 
to lavender, with distinct throat, lobes with 
numerous small glands outside, sometimes 
with short hairs; anther thecae without basal 
tails or with denticulate tails having thin- 
walled cells, apical anther appendage with- 
out glands, usually with ornate thickenings 
on cell walls (thin-walled in Brazilian and 
some Bolivian species); style with distinct 
broadened sclerified basal ring; style 
branches with blunt-tipped hairs. Achenes 
prismatic, usually with 8 veins, 3—8-ribbed, 
with many rounded idioblasts on surface 
usually in clusters, with or without setulae, 
raphids in walls subquadrate to short-ob- 
long; carpopodium shortly cylindrical to 
stopper-shaped, cells subquadrate in many 
series, with thickened cell walls; pappus bi- 
seriate, inner series elongate, of many bris- 
tles with broadened tips, short outer series 
often weak or nearly lacking. Pollen grains 
tricolporate, spinulose, type A. 

Chromosome numbers are known only 

from Mexican members of the genus: N = 
17, C. autumnalis (Jones 1979); N = 18, C. 
foliosa (Jones 1973 as V. steetzii); N = 17 
or 18, C. leiocarpa (Jones 1973); N = 19, 
V. obtusa (Jones 1973); N = 37 + 1, C. 
salicifolia (Jones 1973); N = 36, C. tomen- 
tosa (Jones 1973 as C. paniculata); N = 36- 
39, C. uniflora (Jones 1974). 

Previous combinations in Critoniopsis in- 
clude the type species (Schultz-Bip, 1863), 
the species treated by Cuatrecasas (1956) 
that were transferred by Robinson (1980), 
two Venezuelan species transferred by Badi- 
llo (1983), and two Andean species trans- 
ferred by Robinson (1990b). Two addition- 
al species were named from Venezuela by 
Badillo (1989). Species that have been placed 
in Vernonia sect. Critoniopsis that are here 
excluded from the genus include four in Ba- 
ker (1873) and three in Cuatrecasas (1956). 
The Brazilian species included by Baker, but 
excluded here, are Vernonia puberula Less., 
V. diffusa Less. and V. discolor Less., which 
belong to the genus Vernonanthura H. Rob- 


VOLUME 106, NUMBER 3 


inson (1992a), and V. serrata Less., which 
belongs to Dasyanthina H. Robinson. The 
three species included by Cuatrecasas, but 
excluded here, are Vernonia crassilanata 
Cuatr. of Colombia and Ecuador and V. tri- 
chotoma Gleason of Colombia, which are 
members of Joseanthus H. Robinson (1989), 
and V. vargasii Cuatr. of Peru, which is ap- 
parently a synonym of Vernonanthura pat- 
ens (H.B.K.) H. Robinson (1992a). The spe- 
cies concepts of Jones (1973) and Jones & 
Stutts (1981) are accepted here. 

The 76 species presently accepted in Cvi- 
toniopsis are as follows: 


Critoniopsis angusta 
(Gleason) H. Robinson, 
comb. nov. 


Eremosis angusta Gleason, N. Amer. F1. 33: 
98. 1922. 

Vernonia angusta (Gleason) Standl., Publ. 
Field. Mus. Nat. Hist., Bot. Ser. 11:276. 
1936. 


Distribution. —Guatemala. 


Critoniopsis autumnalis 
(McVaugh) H. Robinson, 
comb. nov. 


Vernonia autumnalis McVaugh, Contr. 
Univ. Michigan Herb. 9(4):477. 1972. 


Distribution. —Mexico. 

Unlike other members of the genus, the 
species is a short-lived perennial herb. It 
was excluded from the section Eremosis by 
Jones (1973), but the anther appendages lack 
glands and have thickenings on the cell walls 
as in other members of Critoniopsis. 


Critoniopsis baadii 
(McVaugh) H. Robinson, 
comb. nov. 


Vernonia salicifolia (DC.) Schultz-Bip. var. 
baadii McVaugh, Contr. Univ. Michigan 
Herb. 9:484. 1972. 


609 


Vernonia baadii (McVaugh) S. Jones, Brit- 
tonia 25:113. 1973. 


Distribution. —Mexico. 


Critoniopsis barbinervis 
(Schultz-Bip.) H. Robinson, 
comb. nov. 


Vernonia barbinervis Schultz-Bip. in Seem., 
Bot. Voy. Herald 297. 1856. 

Eremosis barbinervis (Schultz-Bip.) Glea- 
son, Bull. New York Bot. Gard. 4:232. 
1906. 


Distribution. —Mexico. 


Critoniopsis bitriflora 
(Cuatr.) H. Robinson, 
Phytologia 46:439. 1980. 


Vernonia bitriflora Cuatr., Bot. Jahrb. Syst. 
77:64. 1956. 


Distribution. —Colombia. 


Critoniopsis bogotana 
(Cuatr.) H. Robinson, 
Phytologia 46:439. 1980. 


Vernonia bogotana Cuatr., Bot. Jahrb. Syst. 
77:65. 1956. 

Vernonia calerana Cuatr., Not. Syst. Paris 
15(2):238. 1956. 


Distribution. —Colombia. 

The species is the most common of the 
genus in the area of Depto. Cundinamarca, 
but reports from outside of that area are not 
accepted here. 


Critoniopsis boliviana 
(Britton) H. Robinson, 
comb. nov. 


Vernonia boliviana Britton, Bull. Torrey Bot. 
Club. 18:332. 1891. 

Vernonia paucisquamata Rusby, Bull. New 
York Bot. Gard. 4:376. 1907. 


Distribution. —Ecuador, Bolivia. 
Many specimens are known from Bolivia, 


610 


and the species was recently collected in Ec- 
uador: Zamora-Chinchipe: Nangaritza 
Canton Ridge crest of Cordillera del Con- 
dor, above Pachicutza, on the disputed Peru- 
Ecuador border, Neill & Palacios 9523 (MO, 
QCNE, US). 


Critoniopsis brachystephana 
(Cuatr.) H. Robinson, 
Phytologia 46:439. 1980. 


Vernonia brachystephana Cuatr., Bot. Jahrb. 
Syst. 77:66. 1956. 


Distribution. —Colombia. 

The species seems to be distinguished 
from the closely related C. lindenii only by 
the lack of distinct, large squamae in the 
outer pappus. 


Critoniopsis cajamarcensis 
(H. Robinson) H. Robinson, 
comb. nov. 


Vernonia cajamarcensis H. Robinson, Phy- 
tologia 53:393. 1983. 


Distribution. —Peru, Bolivia. 


Critoniopsis cotopaxensis H. Robinson, 
Sp. NOv. 
(Fig. 1) 


Plantae arborescentes ad 13 m altae; caules 
subhexagonales dense patentiter rufo-to- 
mentosi. Folia alterna vel subopposita, peti- 
olis 2-4 cm longis dense rufo-hirtellis; lam- 
inae subcoriaceae ovatae vel oblongo-ovatae 
13-22 cm longae 6.5—10.0 cm latae base 
obtusae vel breviter acutae margine integrae 
apice leniter breviter anguste acuminatae 
supra planae vel vix rugulosae parce pilo- 
sulae glabrescentes subtus in nervis et ner- 
vulis dense prominulae sordide tomentellae 
et parce glanduliferae, pilis in nervis pri- 
mariis rufo-hirsutis, pilis in nervis secun- 
dariis et nervulis pallidis base stellatae ar- 
matis, pilis in areolis albis minute stellatis, 
nervis secundariis patentiter pinnatis. Inflo- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


rescentiae laxe pyramidaliter thyrsoideae in 
ramis primariis et ramulis alternis vel sub- 
oppositis late patentibus rufo-hirsutae vel 
patentiter tomentosae, pedunculis in fasci- 
culis plerumque 2-3 ca. 0-3 mm longis. Ca- 
pitula submatura ca. 7 mm alta 4-5 mm 
lata; bracteae involucri ca. 25 appresse im- 
bricatae ca. 3—4-seriatae ovatae vel anguste 
oblongae 1.5—4.5 mm longae 1.0-1.4 mm 
latae apice breviter acutae vel obtusae extus 
distincte vel in partibus superioribus leniter 
puberulae. Flores 8 vel 9; corollae albae? ca. 
4.2 mm longae, tubis immaturis ca. 1.3 mm 
longis, faucibus ca. 0.5 mm longis, lobis ca. 
2.5 mm longis extus ubique minute glan- 
duliferis; thecae antherarum ca. 1.7 mm 
longae base non caudatae; appendices an- 
therarum ca. 0.4 mm longae, cellulis dis- 
tincte reticulate ornatis; pili stylorum apice 
rotundati. Achenia immatura ca. 1 mm lon- 
ga glabra; setae pappi interiores ca. 4.2 mm 
longae distaliter sensim distincte latiores; 
setae exteriores immaturae indistincte. 
Grana pollinis in diametro ca. 40 um tri- 
colporata spinulosa. 

Type: ECUADOR: Cotopaxi: Carretera 
Latacunga—Pilalo—Quevado, 5—15 km al este 
de Pilalo, 00°55’S, 79°01'W, 2700-3350 m, 
bosque muy humedo Montano Bajo, bos- 
que disturbado, arbol de 13 m, capitulos 
color crema blanco, 22 May 1988, Ceron, 
Neill & Palacios 3804 (holotype US; isotype 
MO). 

Critoniopsis cotopaxensis is similar to C. 
palaciosii, described below, in the size and 
shape of its leaves and the stellate or basally 
armed hairs of the leaf undersurface, but the 
leaves of the latter seem rigorously opposite 
with secondary veins more ascending, the 
heads have about 6 florets, and the achenes 
have a pubescence of uniseriate setulae. The 
present species is similar to C. suaveolens 
in its general aspect and 8 or 9 florets in the 
heads, but the latter has smaller leaves with 
a rugose upper surface, contorted simple 
hairs on the lower surface, and a denser in- 
florescence. 


VOLUME 106, NUMBER 3 


UNITED STATES 


3145460 


NATIONAL HERBARIUM 


611 


ECUADOR 


ASTERACEAE 
Cael i -_ 
Crit op sis 


Prove COTOPAXI: 

Carretera Latacunga-Pilal0-Quevedo. 
5-15 km al este de Pilalo. 

Bosque muy humedo Nontano BajO-. 
Bosque disturbado. 

00°55'S 79°01'W 2700-3350 m 


Arbol de 13 m; capitulos color 
crema blanco. 


Carlos E. CerOnr 22 mayo 1988 
D. Neillr W. Palacios 3804 
MISSOURI BOTANICAL GARDEN H ERBARIUM (MO) 


Fig. 1. Holotype of Critoniopsis cotopaxensis H. Robinson, Cer6n, Neill & Palacios 3804 (US). 


612 


Critoniopsis cuatrecasasii H. Robinson, 
Phytologia 46:439. 1980. 


Distribution. —Colombia. 


Critoniopsis dorrii H. Robinson, sp. nov. 
(Fig. 2) 


Plantae arborescentes 2—6 m altae; caules 
brunnescentes superne dense appresse to- 
mentelli inferne subglabrescentes. Folia op- 
posita vel interdum alterna, petiolis 6-12 
mm longis; laminae coriaceae ellipticae 
plerumque 4—8 cm longae 1.8—3.8 cm latae 
base et apice breviter acutae margine inte- 
grae supra planae glabrae subtus in nervulis 
minute reticulate distincte prominulae dense 
pallide tomentellae, pilis congestis minute 
stellatis, nervis secundarlis utrinque ca. 8 
vel 9. Inflorescentiae in ramis foliatis ter- 
minales laxe pyramidaliter thyrsoideae ap- 
ice et in ramis rotundate corymbosae laxae, 
pedunculis 1-8 mm longis dense appresse 
stellate tomentellis. Capitula 10-12 mm alta 
3—4 mm lata; bracteae involucri ca. 40 ap- 
presse imbricatae ca. 5-seriatae late ovatae 
vel lineari-lanceolatae 1—S mm longae 1.0- 
1.3 mm latae apice breviter acutae extus 
parce appresse puberulae. Flores 9-12 in 
capitulo; corollae albae? ca. 7 mm longae 
parce minute glanduliferae, tubis infundi- 
bularibus ca. 5 mm longis, faucibus ca. 1 
mm longis, lobis ca. 2mm longis; thecae 
antherarum ca. 2 mm longae base vix vel 
non caudatae; appendices antherarum 0.45-— 
0.50 mm longae, parietibus cellularum dis- 
tincte reticulate ornatis; pili stylorum apice 
rotundati. Achenia ca. 3.8 mm longa glabra 
vel perpauce minute setulifera; setae pappi 
interiores ca. 5.5 mm longae distaliter sen- 
sim distincte latiores. Grana pollinis in dia- 
metro ca. 40 um tricolporata spinulosa. 

Type: ECUADOR: Azuay: Cuenca-—Sol- 
dados road (following N bank of Rio Ya- 
nuncay), 19-20 km W of San Joaquin 
(2°55’'S, 79°5'W), small tree 2—3 m high, 22 
Jun 1989, Dorr & Valdespino 6404 (holo- 
type US; isotypes F, NY). Paratype: EC- 
UADOR: Azuay: mountains above Say- 
ausid, 3000-3200 m, tree S—6 m high, corolla 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


white, 18 Mar 1974, Harling & Andersson 
12585 (GB). 

Critoniopsis dorrii resembles C. sodiroi (C. 
pichinchensis) of north-central Ecuador in 
habit, especially with the mostly opposite 
leaves. There is also a resemblance to C. 
pycnantha of southernmost Ecuador and 
northern Peru. The new species differs from 
the latter two by the looser inflorescence, 
the larger heads with more florets, and by 
the strongly prominulous dense reticulation 
of veinlets on the lower leaf surface. Leaves 
on the type series are strictly opposite, but 
those of the paratype are alternate on some 
branches. 


Critoniopsis duncanii 
(S. Jones) H. Robinson, 
comb. nov. 


Vernonia duncanii S. Jones, Brittonia 25: 
108. 1973. 


Distribution. —Mexico. 


Critoniopsis elbertiana 
(Cuatr.) H. Robinson, 
Phytologia 46:440. 1980. 


Vernonia elbertiana Cuatr., Bot. Jahrb. Syst. 
77:68. 1956. 


Distribution. —Columbia, Ecuador. 

The species is one of two noted by Cua- 
trecasas (1956) for the contracted bases and 
recurved “‘auriculate’”’ basal margins of the 
median and inner involucral bracts. The type 
and one more recent collection (King 6199, 
US) are from Depto. Putumayo in southern 
Colombia. Two other collections have been 
seen from adjacent Prov. Napo in Ecuador 
(Palacios 5360, 6416, MO, US). All speci- 
mens are from the eastern escarpment of 
the Andes. 


Critoniopsis floribunda 
(H.B.K.) H. Robinson, 
Phytologia 46:440. 1980. 


Vernonia floribunda H.B.K., Nov. Gen. et 
Sp., ed. fol. 4:30. 1818. 


VOLUME 106, NUMBER 3 


UNITED STATES 


3187575 


NATIONAL HERBARIUM 


Holotype 


THE NEW YORK BOTANICAL GARDEN 
PLANTS OF ECUADOR 


No. 6404 Compositae 
- ; f eo he 
Cr ilmiepsis Omrti iH Rela i) 


ECUADOR. Prov. Azuay. Cuenca-Soldados road 
(following the N bank of the Rio Yanuncay), 19-20 km 
W of San Joaquin (2°55'S;79°5'W). 


Small tree, 2-3 m tall. 


L. J. Dorr & I. Valdespino 22 June 1989 


Fig. 2. Holotype of Critoniopsis dorrii H. Robinson, Dorr & Valdespino 6404 (US). 


613 


614 


Vernonia affinis H.B.K., Nov, Gen. et Sp., 
ed. fol. 4:30. 1818. 


Distribution. —Ecuador, Peru. 

The type specimen has been ascribed, with 
some question, to Peru. Recent collections 
identified as the species are from Prov. Loja 
in southermost Ecuador (Ollgaard, Madsen 
& Christensen 74596, AAU, UCQ, US; 
Dalessandro 657, MO, US). 


Critoniopsis foliosa 
(Benth.) H. Robinson, 
comb. nov. 


Monosis foliosa Benth., Pl. Hartw. 19. 1839. 
Vernonia steetzii Schultz-Bip. in Seem, Bot. 
Voy. Herald 297. 1856. 
Vernonia Ssteetzii var. callilepis Schultz-Bip. 
in Seem., Bot. Voy. Herald 297. 1856. 
Vernonia foliosa (Benth.) Schultz-Bip., Jah- 
resber. Pollichia 18/19:161. 1891, non V. 
foliosa Gardn. 1846. 

Eremosis foliosa (Benth.) Gleason, Bull. New 
York Bot. Gard. 4:228. 1906. 

Eremosis steetzii (Schultz-Bip.) Gleason, 
Bull. New York Bot. Gard. 4:230. 1906. 

Vernonia mucronata Blake, Contr. Gray 
Herb. 523195 1917, 

Eremosis callilepis (Schultz-Bip.) Gleason, 
N. Amer. Fl. 33:98. 1922. 

Vernonia aristifera Blake, Contr. U.S. Natl. 
Herb. 23:1415. 1926. 


Distribution. — Mexico. 
Critoniopsis franciscana 


(Cuatr.) H. Robinson, 
Phytologia 46:440. 1980. 


Vernonia franciscana Cuatr., Bot. Jahrb. 
Syst. 77:69. 1956. 


Distribution.—Colombia. 
Critoniopsis glandulata 


(Cuatr.) H. Robinson, 
Phytologia 46:440. 1980. 


Vernonia glandulata Cuatr., Bot. Jahrb. Syst. 
11269: 1956: 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Distribution. —Colombia, Venezuela. 

The species is distinctive in the heads that 
are 12 mm high when mature and in the 
large coriaceous leaf blades, with rounded 
apices, glabrous undersurfaces, and the short 
basal decurrences with recurved margins. 
The basal decurrence of the blade is not as 
abruptly delimited as that of C. killipii. 


Critoniopsis harlingii 
(H. Robinson) H. Robinson, 
comb. nov. 


Vernonia harlingii H. Robinson, Phytologia 
44:66. 1979. 


Distribution. —Ecuador. 

The large, sessile, opposite leaves, the 
broad, whitish, dissected margins of the in- 
volucral bracts, and the heads with about 
20 florets are distinctive. 


Critoniopsis heydeana 
(Coult.) H. Robinson, 
comb. nov. 


Vernonia heydeana Coult., Bot. Gaz. 
(Crawfordsville) 20:42. 1895. 

Eremosis heydeana (Coult.) Gleason, Bull. 
New York Bot. Gard. 4:234. 1906. 


Distribution. —Mexico, Guatemala. 


Critoniopsis huairacajana 
(Hieron.) H. Robinson, 
Phytologia 46:440. 1980. 


Vernonia huairacajana Hieron., Bot. Jahrb. 
Syst. 19:43. 1894. 


Distribution. —Ecuador, Peru. 

The species was described from Prov. 
Azuay in southern Ecuador and has been 
collected in nearby Prov. Canar (Prieto P-76, 
NY, US; King & Almeda 7743, US). A spec- 
imen from Depto. Cajamarca in Peru is also 
identified as this species (Becker & Terrones 
2283, US). 


VOLUME 106, NUMBER 3 


Critoniopsis huilensis 
(Cuatr.) H. Robinson, 
Phytologia 46:440. 1980. 


Vernonia huilensis Cuatr., Bot. Jahrb. Syst. 
g7-71: 1956. 


Distribution. —Colombia. 


Critoniopsis jalcana 
(Cuatr.) H. Robinson, 
comb. nov. 


Vernonia jalcana Cuatr., Ann. Missouri Bot. 
Gard: 52:312. 1965. 


Distribution. — Peru. 


Critoniopsis jelskii 
(Hieron.) H. Robinson, 
Phytologia 46:440. 1980. 


Vernonia jelskii Hieron., Bot. Jahrb. Syst. 
36:459. 1905. 


Distribution. — Peru. 


Critoniopsis jubifera 
(Rusby) H. Robinson, 
comb. nov. 


Vernonia jubifera Rusby, Mem. Torrey Bot. 
Club 6:53. 1896. 

Vernonia conwayi Rusby, Bull. New York 
Bet soard. 8:125..1912. 


Distribution. — Bolivia. 


Critoniopsis killipii 
(Cuatr.) H. Robinson, 
Phytologia 46:440. 1980. 


Vernonia killipii Cuatr., Bot. Jahrb. Syst. 
ea-74,. 1956. 

Vernonia bogotana var. santanderensis 
Cuatr., Bot. Jahrb. Syst. 77:66. 1956. 


Distribution. —Colombia, Venezuela 
(Tachira). 

The species is easily distinguished by the 
base of the leaf blade that has a strongly 
recurved margin and an abrupt decurrence 


for about 1 cm on the petiole. Specimens in 


615 


addition to the type include one from Dep- 
to. Santander, Colombia (Killip & Smith 
18332, US) and one from Edo. Tachira in 
Venezuela (Steyermark & Dunsterville 
98656, US) that have been annotated as 
Vernonia bogotana var. santanderensis 
Cuatr. by Keeley. 


Critoniopsis leiocarpa 
(DC.) H. Robinson, 
comb. nov. 


Vernonia leiocarpa DC., Prodr. 5:34. 1836. 

Cacalia leiocarpa (DC.) Kuntze, Rev. Gen. 
Pi; 2:970. 1891. 

Eremosis leiocarpa (DC.) Gleason, Bull. 
New York Bot. Gard. 4:232. 1906. 

Eremosis melanocarpa Gleason, Bull. New 
York Bot. Gard. 4:232. 1906. 


Distribution. —Mexico, Guatemala, Be- 
lize, El Salvador, Honduras, Nicaragua. 


Critoniopsis lewisii H. Robinson, 
sp nov. 


Plantae fruticosae et subarborescentes vel 
leniter scandentes 3-4 m altae, ramulis nu- 
merosis; caules atro-brunnescentes subtere- 
tes irregulariter striati dense minute puberu- 
li. Folia alterna, petiolis 0.4—0.7 mm longis; 
laminae subcoriaceae ovatae vel anguste 
ovatae plerumque 4.0—6.5 cm longae 1.3- 
2.2 cm latae base breviter acutae margine 
perminute remote denticulatae apice acutae 
supra glanduliferae cetera glabrae in ner- 
vulis minime prominulis subtus dense glan- 
dulo-punctatae subglabrae in nervis major- 
ibus appresse puberulae in nervulis non 
prominulis dense brunnescentiter reticula- 
tae, nervis secundariis pinnatis utrinque 7 
vel 8 in angulis 50—75° patentibus. Inflores- 
centiae late rotundate corymbosae in ramis 
dense corymbosae in glomerulis capitulo- 
rum subsessiliorum terminatis, pedunculis 
0-2 mm longis dense breviter sordide pub- 
erulis vel subvelutinis. Capitula ca. 11 mm 
altae; bracteae involucri ca. 20 appresse im- 
bricatae ca. 4—-seriatae ovatae vel anguste 


616 


oblongae 1.5—7.0 mm longae 1.0-2.2 mm 
latae apice acutae extus sparse appresse pal- 
lide puberulis. Flores 3—5; corollae purpu- 
reae ca. 7 mm longae, tubis 3.5-4.0 mm 
longis, faucibus ca. 0.8 mm longis extus 
pauce glanduliferis, lobis ca. 3 mm longis 
ca. 0.5 mm latis extus glanduliferis distaliter 
densiores apice pauce pilulosis, pilis leniter 
T-formibus; thecae antherarum ca. 3 mm 
longae, caudis basilaribus 0.8 mm longis re- 
trorse denticulatis; appendices antherarum 
ca. 0.55 mm longae 0.27 mm latae, cellulis 
distincte leniter ornate annulatae; pili sty- 
lorum apice rotundati. Achenia ca. 3 mm 
longa 10-costata in sulcis glandulifera cetera 
glabra; setae papillae pluriseriatae pler- 
umque ca. 7 mm longae inferne scabridulae 
distaliter laeviores vix latiores, setae pappi 
breviores pauces irregulares. Grana pollinis 
in diametro ca. 35 wm tricolporata spinu- 
losa. 

Type: BOLIVIA: La Paz: Prov. Inquisivi: 
‘““Quebrada Jancha Kaihua,” along a ravine 
joining Rio Ocsalla ca. 3 km down river 
from Laguna Huara Huarani, 10 km N of 
Choquetanga, along upper edge of ravine 
cloud forest, Clethra, Hesperomeles, Wein- 
mannia, Saracha, Berberis, Gynoxys, Myr- 
ica are all common, 16°45’S, 67°17'W, 3400- 
3600 m, vine over small trees, inflorescence 
white, 3 Sep 1991, Marko Lewis 39696 (ho- 
lotype US; isotypes LPB, MO). Paratype: 
BOLIVIA: La Paz: Prov. Inquisivi: 
‘““Chachacomani,” slope SW of the Rio Oc- 
salla, a few hundred meters above river and 
3 km SE of its mouth 12 km NE of Cho- 
quetanga, 16°17’S, 67°17'W, 3300 m, forest 
edges, large shrub or small tree 3—4 m high, 
phyllaries deep black-red-purple, flowers 
white-purple, 18 Apr 1991, Marko Lewis 
38834 (LPB, MO, US). 

The new species has the aspect of Crito- 
niopsis quinqueflora of southeastern Brazil 
and C. weberbaueri of northern Peru. The 
Brazilian species differs most obviously by 
having a differentiated, short, scale-like out- 
er pappus series and a distinctly setuliferous 
achene. The Peruvian species is most close- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ly related, but it differs by the fewer branch- 
es, the generally larger leaves, the glands on 
the leaf undersurfaces that are very dense 
and almost touching, the more numerous 
and more prominent secondary veins, the 
more ascending branches of the inflores- 
cence, and the denser and longer pubescence 
of the involucre. 


Critoniopsis lindenii Schultz-Bip.., 
Jahresber. Pollichia 20/21:431. 1863. 


Vernonia lindenii (Schultz-Bip.) Cuatr., Bot. 
Jahrb. Syst. 57:72. 1956. 


Distribution. —Colombia. 

Specimens have been determined recent- 
ly from Depto. Antioquia (Uribe Uribe 
2071, US; Escobar & Velasquez 7534, US; 
Zarucchi, Betancur & Roldan 5273, MO, 
US) and Depto. Quindio (Gentry, Velez & 
Carvajal 65348, MO, US). 


Critoniopsis littoralis 
(Brandg.) H. Robinson, 
comb. nov. 


Vernonia littoralis Brandg., Erythea 7:3. 
1899. 

Eremosis littoralis (Brandg.) Gleason, North. 
Amer. P1:33:100. 1922 


Distribution. —Mexico. 


Critoniopsis macphersonii 
(S. Jones & Stutts) H. Robinson, 
comb. nov. 


Vernonia macphersonii S. Jones & Stutts, 
Brittonia 33:546. 1981. 


Distribution. —Mexico. 


Critoniopsis macvaughii 
(S. Jones) H. Robinson, 
comb. nov. 


Vernonia macvaughii S. Jones, Brittonia 25: 
105. 1973. 


Distribution. —Mexico. 


VOLUME 106, NUMBER 3 


Critoniopsis magdalenae 
(Barroso) H. Robinson, 
comb. nov. 


Vernonia magdalenae Barroso, Arq. Jard. 
Bot. Rio Janeiro 13:12. 1954. 


Distribution. — Brazil. 


| Critoniopsis meridensis 
(Badillo) Badillo, 
Ernstia 16:16. 1983. 


Vernonia meridensis Badillo, Rev. Fac. 
Agron. Univ. Central (Maracay). 9:87. 
1976. 


Distribution. — Venezuela. 


Critoniopsis mucida 
(Cuatr.) H. Robinson, 
Phytologia 46:440. 1980. 


Vernonia mucida Cuatr., Bot. Jahrb. Syst. 
THT2. 1956. 


Distribution. —Colombia. 


Critoniopsis occidentalis 
(Cuatr.) H. Robinson, 
Phytologia 46:440. 1980. 


Vernonia occidentalis Cuatr., Bot. Jahrb. 
Syst. 77:73. 1956. 


Distribution. —Colombia, Ecuador. 

The species was originally described from 
Depto. Valle in Colombia, but specimens 
have been seen recently from Depto. Narinfo 
(Gentry, Benavides & Keating 60522, MO, 
US; Beltran 44, US) and farther south in 
Ecuador in Prov. Carchi (Hoover et al. 2556, 
MO, US; Palacios & Rubio 7271), Prov. 
Napo (Palacios 5369, MO, US), and Prov. 
Pichincha (Jaramillo et al. 8055, 8068, 8089, 
MO, US; Zak & Jaramillo 2519, MO, US; 
Wak ti%6, 1239, 1251 US). 


Critoniopsis oolepis (Blake) H. Robinson, 
comb. nov. 


Vernonia oolepis Blake, Contr. Gray Herb. 
a22 20: 191. 


617 


Eremosis oolepis (Blake) Gleason, North. 
Amer. FI. 33:97. 1922. 


Distribution. — Mexico. 


Critoniopsis obtusa (Gleason) H. Robinson, 
comb. nov. 


Eremosis obtusa Gleason, N. Amer. FI. 33: 
99. 1922. 

Vernonia obtusa (Gleason) Blake, Contr. 
U.S. Natl. Herb. 23:1415. 1926. 


Distribution. — Mexico. 


Critoniopsis ovata (Gleason) H. Robinson, 
comb. nov. 


Eremosis ovata Gleason, Bull. Torrey Bot. 
Club 40:331. 1913, non Vernonia ovata 
Less., 1829. 

Vernonia gleasonii Blake, Contr. Gray Herb. 
52:17. 1917, non V. gleasonii Ekman, 
1914. 

Vernonia durangensis Blake, Contr. U.S. 
Natl. Herb. 22:587. 1924. 


Distribution. — Mexico. 


Critoniopsis palaciosii H. Robinson, 
sp. nov. 


(Fig. 3) 


Plantae dendroideae ad 10 m altae; caules 
atro-brunnescentes dense brunneo-velutini. 
Folia opposita, petiolis 2-3 cm longis; lam- 
inae coriaceae ovatae 13-18 cm longae 4— 
8 cm latae base obtusae margine integrae 
apice breviter acutae supra rugulosae glan- 
dulo-punctatae caetera glabrae in nervulis 
non prominulis subtus in nervulis minute 
reticulate prominulae dense sordide tomen- 
tellae, pilis contortis base stellate armatis, 
nervis secundarlis utrinque ca. 13 inferne 
patentiores. Inflorescentiae pyramidaliter 
thyrsoideae in ramis primarilis oppositae 
dense pallide tomentosae vel sublanatae, ra- 
mulis dense breviter sordide velutinis in 
glomerulis capitulorum subsessiliorum ter- 
minatis. Capitula 8-9 mm alta; bracteae in- 
volucri ca. 22 appresse imbricatae ca. 


618 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


S19 }euwypI{Ue. 


ECUADOR 
ASTERACEAE 
CHionioes iosii  -Rebini 
Critonicpsis palaciosii LOD IP Sa}, 
Hi : 
JTOUIY Pe 4 
IMBABURA 
Canton Cotacachi. 
Carretera Cotacachi - Apuela. 
Sitio Tabla Chuap, a1 km de Hacienda 


La Providencia. Bosque primario. 
78°25'W 00°25'N 3100 m 


SS, 


a 


UNITED STATES 
Arbol de 10 m de altura. 


Hojas coriaceas con el envés parduzco. 
Capitulos verdes. Ligulas blancas. 


3215780 


NATIONAL HERBARIUM i 4 abril 1990 
Walter Palacios & C. Iguago 4867 
MISSOURI BOTANICAL GARDEN HERBARIUM (MO) 


Fig. 3. Holotype of Critoniopsis palaciosii H. Robinson, Palacios & Iguago 4867 (US). 


VOLUME 106, NUMBER 3 


4-seriatae ovatae vel anguste oblongae 1.5- 
6.0 mm longae et 1.0-1.5 mm latae apice 
obtusae extus superne dense puberulae. Flo- 
res ca. 6; corollae albae? ca. 6.5 mm longae, 
tubis ca. 2 mm longis, faucibus 1.5 mm lon- 
gis, lobis ca. 2.5 mm longis extus ubique 
minute gladuliferis; thecae antherarum ca. 
1.5 mm longae base breviter caudatae; ap- 
pendices antherarum ca. 0.4 mm longae, 
cellulis distincte ornate annulatae; pili sty- 
lorum apice rotundati. Achenia ca. 2.2 mm 
longa pilifera, pilis uniseriatis; setae pappi 
ca. 4.5 mm longae superne sensim distincte 
lateriores; setae exteriores perbreves. Grana 
pollinis in diametro 35—40 um tricolporata 
spinulosa. | 

Type: ECUADOR: Imbabura: Canton 
Cotacachi: carretera Cotacachi—Apuela, si- 
tio Tabla Chuap, a 1 km de Hacienda La 
Providencia, 78°25’W, 00°25’N, 3100 m, 
bosque primario, arbol de 10 m altura, ho- 
jas coriaceas con el envés parduzco, capi- 
tulos verdes, ligulas blancas, 4 Apr 1990, 
W. Palacios & C. Iguago 4867 (holotype US, 
isotype MO). 

Critoniopsis palaciosii is notable for the 
opposite leaves with large, ovate blades and 
the dense and prominulous veinlets on the 
lower surface. The only other species with 
opposite leaves with large blades is C. har- 
lingii of southern Ecuador, but the latter has 
sessile leaves, distinctive dissected whitish 
margins on the involucral bracts, and about 
20 florets in its heads. 


Critoniopsis pallens 
(Schultz-Bip.) H. Robinson, 
comb. nov. 


Vernonia pallens Schultz-Bip., Jahresber. 
Pollichia 18/19:161. 1861. 

Eremosis pallens (Schultz-Bip.) Gleason, 
Bull. New York Bot. Gard. 4:228. 1906. 

Vernonia michoacana McVaugh, Contr. 
Univ. Michigan Herb. 9:482. 1972. 


Distribution. — Mexico. 


619 


Critoniopsis pallida 
(Cuatr.) H. Robinson, 
Phytologia 46:440. 1980. 


Vernonia pallida Cuatr., Bot. Jahrb. Syst. 
77:74. 1956. 


Distribution. —Colombia. 


Critoniopsis paradoxa 
(Schultz-Bip.) Badillo, 
Ernstia 16:16. 1983. 


Tephrothamnus paradoxus Schultz-Bip.., 
Jahresber. Pollichia 20/21:432. 1863. 
Piptocarpha venezuelensis Badillo, Bol. Soc. 
Venez. Cienc. Nat. 10:280. 1946. 

Piptocarpha paradoxa (Schultz-Bip.) Aris- 
teguieta, Fl. Venezuela, Compositae 10(1): 
54. 1964. 


Distribution. — Venezuela. 


Critoniopsis paucartambensis 
(Dillon) H. Robinson, 
comb. nov. 


Vernonia paucartambensis Dillon, Britton- 
ia 36:336. 1984. 


Distribution. — Peru. 

The species was described from the Dep- 
to. Cuzco, and an additional specimen has 
been seen from Depto. Puno (Boeke & Boeke 
3076, NY, US). 


Critoniopsis pendula 
(Cuatr.) H. Robinson, 
Phytologia 46:440. 1980. 


Vernonia pendula Cuatr., Bot. Jahrb. Syst. 
T7239 19 56: 


Distribution. —Columbia. 
Critoniopsis peruviana 


(Cuatr.) H. Robinson, 
comb. nov. 


Vernonia peruviana Cuatr., Bot. Jahrb. Syst. 
VALS S56: 


Distribution. — Peru. 


620 


Critoniopsis popayanensis 
(Cuatr.) H. Robinson, 
Phytologia 46:440. 1980. 


Vernonia popayanensis Cuatr., Bot. Jahrb. 
Syst. 77:77. 1936. 


Distribution. —Colombia. 

The leaf blades of the species have a 
slightly recurved basal margin and a basal 
decurrence reminiscent of C. killipii, but the 
decurrence is in no way as strong or abruptly 
limited. The type specimen was from Dep- 
to. Cauca in Colombia, and there are two 
recent collections from Depto. Antioquia 
(Croat 69888, MO, US; Betancur, Roldan 
& Castano 1133, HUA, US). 


Critoniopsis pugana 
(S. Jones & Stutts) H. Robinson, 
comb. nov. 


Vernonia pugana S. Jones & Stutts, Brit- 
tonia 33:544. 1981. 


Distribution. —Mexico. 


Critoniopsis pycnantha 
(Benth.) H. Robinson, 
Phytologia 46:441. 1980. 


Vernonia pycnantha Benth., Pl. Hartw. 134. 
1844. 

Tephrothamnus? pycnanthus (Benth.) 
Schultz-Bip., Jahresber. Pollichia 20/21: 
433. 1863. 


Distribution. —Ecuador. 

The Hartweg type specimen was from 
*“montibus Paccha”’ which is presumed here 
to be in Depto. Ancash, Peru rather than 
Colombia or Ecuador as sometimes stated. 
Material matching the type photograph has 
been seen from Prov. Loja in southernmost 
Ecuador (Ollgaard, Laegaard, Thomsen, 
Korning & Illum 58006, AAU, US; Madsen 
75524, AAU, US). Specimens identified as 
the species from Colombia, central Ecua- 
dor, and Bolivia are considered here to be- 
long to various other species, some of which 
may be unnamed. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Critoniopsis quillonensis H. Robinson, 
sp. nov. 


Plantae arborescentes ad 7 m altae; caules | 
teretes dense granulate flavo-tomentelli, pi- 
lis globuliformis subsessilis sparse perbrev- 
iter spinulosis. Folia opposita vel suboppo- 
sita, petiolis 0.2—0.7 cm longis; laminae vix 
coriaceae anguste ovatae vel ellipticae 4-10 
cm longae 0.8—3.0 cm latae base anguste 
acutae vel acutae margine subserrulatae ap- 
ice anguste acutae vel acuminatae supra gla- 
brae in nervulis minute pallide prominulae 
subtus in nervulis reticulatae distincte 
prominulae dense stellulate tomentellae, pi- 
lis breviter stellatis breviter stipitatis, nervis 
secundariis utrinque 8-11 in angulis ca. 45° 
patentibus. Inflorescentiae terminales late 
pyramidaliter paniculatae, ramulis corym- 
biformis, pedunculis 0-2 mm longis dense 
flave granulo-tomentellis. Capitula ca. 10 
mm alta; bracteae involucri ca. 25 breviter 
suborbiculares vel oblongo-ellipticae 1—4 
mm longae 1.0-1.5 mm latae apice rotun- 
datae vel breviter obtusae extus glabrae. 
Flores ca. 9; corollae albae? ca. 6.5 mm lon- 
gae extus in faucibus et in apicis lobarum 
glanduliferae, tubis ca. 2.5 mm longis, fau- 
cibus 0.7 mm longis, lobis ca. 3 mm longis 
base ca. 0.5 mm latis; thecae antherarum 
ca. 1.2 mm longae base vix appendiculatae; 
appendices antherarum ca. 0.4 mm longae; 
base stylorum abrupte nodulosi, pilis sty- 
lorum apice rotundatae. Achenia ca. 3 mm 
longa glabra 6 vel 7-costata; setae pappi al- 
bae ca. 4 mm longae distaliter sensim dis- 
tincte latiores; squamae exteriores ca. 0.5 
mm longae. Grana pollinis in diametro ca. 
45 wm. 

Type: PERU: Cajamarca: Prov. San Mi- 
guel: Cerro Quillon (Agua Blanca), 3150 m, 
5 Jul 1986, Mostacero, Alvitez, Leiva, Me- 
jia & Pelaez 1286 (holotype, US; isotypes, 
F, MO, NY). 

The species is known only from the type 
collection. Data on the label indicates a 
roadside and “‘Arbol de hasta 7 m de alto 
con capitulos blancos.” 

The species is another with opposite to 


VOLUME 106, NUMBER 3 


subopposite leaves. Such forms are more 
common in Colombia and Ecuador, and 
only the sessile-leaved C. sagasteguii has 
previously been described with opposite 
leaves from Peru. The new species is dis- 
tinctive in its narrow, almost lanceolate, leaf 
blades with narrowly cuneate bases and acute 
apices, and by the subserrulate margins of 
the leaf blades. 


Critoniopsis quinqueflora 
(Less.) H. Robinson, 
comb. nov. 


Vernonia quinqueflora Less., Linnaea 6:656. 
1831. 


Distribution. — Brazil. 


Critoniopsis sagasteguli 
(Dillon) H. Robinson, 
comb. nov. 


Vernonia sagasteguii Dillon, Brittonia 36: 
333. 1984. 


Distribution. —Peru. 

Dillon (1984) suggests that the species fits 
naturally within the group of Vernonia, now 
placed in Joseanthus, that have opposite 
leaves, spreading inner involucral bracts, 
and cylindrical basal corolla tubes ending 
abruptly at the bases of the deeply cut lobes. 
The original description and the one corolla 
illustrated in the head indicate that the co- 
rolla is typical of Critoniopsis, and that the 
lobes are not separate to the bases of the 
anther filaments. 


Critoniopsis salicifolia 
(DC.) H. Robinson, 
comb. nov. 


Monosis salicifolia DC., Prodr. 5:77. 1836. 

Cacalia salicifolia (DC.) Kuntze, Rev. Gen. 
Pl. 2:971. 1891. 

Eremosis salicifolia (DC.) Gleason, Bull. 
New York Bot. Gard. 4:321. 1906. 

Eremosis leiophylla Gleason, Bull. New 
York Bot. Gard. 4:231. 1906. 


621 


Vernonia leiophylla (Gleason) Blake, Contr. 
Gray Herb. 52:18. 1917. 


Distribution. —Mexico. 


Critoniopsis sevillana 
(Cuatr.) H. Robinson, 
Phytologia 46:441. 1980. 


Vernonia sevillana Cuatr., Bot. Jahrb. Syst. 
77:78. 1956. 


Distribution. —Ecuador. 

The species was originally described from 
Prov. Azuay in Ecuador and two collections 
have been seen from Prov. Loja (Dodson & 
Thein 1343, US; Madsen 85459, AAU, US). 


Critoniopsis shannonii 
(Coult.) H. Robinson, 
comb. nov. 


Vernonia shannonii Coult., Bot. Gaz. 
(Crawfordsville) 20:42. 1895. 

Eremosis shannonii (Coult.) Gleason, Bull. 
New York Bot. Gard. 4:234. 1906. 


Distribution. —Guatemala. 


Critoniopsis sodiroi 
(Hieron.) H. Robinson, 
Phytologia 69:105. 1990. 


Piptocarpha sodiroi Hieron. ex Sodiro, Bot. 
Jahrb. Syst. 29:2. 1900. 

Vernonia pichinchensis Cuatr., Bot. Jahrb. 
Syst. 77:76. 1956. 

Critoniopsis pichinchensis (Cuatr.) H. Rob- 
inson, Phytologia 46:440. 1980. 


Distribution. —Ecuador. 

The opposite-leaved Ecuadorian species 
is known mostly from Prov. Pichincha, but 
has also been collected in Prov. Bolivar (As- 
plund 8223, US; Zak & Jaramillo 2578, MO, 
US), Prov. Chimborazo (Zak & Jaramillo 
2863, 3661, MO, US), Prov. Cotopaxi 
(Holm-Nielsen & Andrade 18510, AAU, 
US; Neill, Palacios & Cer6n 8426, MO, US), 
and Prov. Imbabura (Moran, Vaca, Vallejo 
& Paisano 38, MO, US). 


Critoniopsis standleyi 
(Blake) H. Robinson, 
comb. nov. 


Vernonia standleyi Blake, J. Wash. Acad. 
Ser 1371432892 3: 


Distribution. —Guatemala, El Salvador, 
Honduras, Nicaragua. 


Critoniopsis steinbachii H. Robinson, 
sp. nov. 
(Fig. 4) 


Plantae frutescentes ad 3 m altae in ramis 
arcuatae; caules brunnescentes in nodis dis- 
tincte leniter deflecti superne dense albo- 
tomentosi. Folia alterna, petiolis 1.0-1.5 cm 
longis; laminae membranaceae late oblon- 
go-ovatae plerumque 12-24 cm longae 5—- 
11 cm latae base rotundatae leniter cordatae 
margine multo minute denticulatae apice 
sensim anguste acuminatae supra planae 
sparse vel dense interdum evanescentiter 
pilosulae subtus vix pallidoires plerumque 
in nervis et nervulis dense pilosulae vel sub- 
lanatae, nervis secondariis pinnatis ut- 
rinque ca. 9 in angulis 45°-55° patentibus, 
nervulis vix prominulis. Inflorescentiae in 
ramis terminalibus thysoideae alterne ra- 
mosae, ramis aliquantum dense corymbo- 
so-cymosis, pedunculis 0.3—2.5 mm longis 
dense pallide puberulis. Capitula 5-6 mm 
alta; bracteae involucri ca. 20 appresse im- 
bricatae ca. 4-seriatae oblongae 1.0—3.5 mm 
longae 0.5—1.0 mm latae apice breviter acu- 
tae extus distaliter dense puberulae. Flores 
6—8 in capitulo; corollae lavandulae ca. 4.0 
mm longae extus ubique minute glandul- 
ferae non piliferae, tubis basilaribus ca. 2 
mm longae, faucibus ca. 1 mm longis, lobis 
ca. 1.2 mm longis; thecae antherarum ca. 1 
mm longae base breviter tenuiter caudatae 
et denticulatae; appendices apicales anther- 
arum oblongae ca. 0.4 mm longae et 0.18 
mm latae, parietibus cellularum plerumque 
tenuibus. Achenia ca. 2.5 mm longa ca. 
8-costata patentiter setulifera; setae pappi 
exteriores persistentes ca. 0.4 mm longae; 
setae papp!i interiores facile deciduae ca. 3.5 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


mm longae distaliter sensim distincte later- 
iores. Grana pollinis in diametro ca. 30 um 
tricolporata spinulosa. 

Type: BOLIVIA: Santa Cruz: Prov. Ich- 
ilo: Parque Nacional Amboro, ca. 15 km 
(SE) up the Rio Surutt, moist tropical forest 
on lower montane slopes, sandstone, 
17°44’'S, 63°40’W, 700 m, arching shrub, 3 
m, corollas lavender, 30 Aug 1985, J. C. 
Solomon & S. Urcullo 14171 (holotype US; 
isotypes MO, NY). Paratypes: BOLIVIA: 
Santa Cruz: Cerro Hosana, 27 Aug 1917, 
Steinbach 34993 (NY, US), 34995 (NY). 

Specimens of Critoniopsis steinbachii have 
been either undetermined in herbaria or 
provisionally identified as C. yungasensis 
Britton. Relationship to the latter species 
seems closest, but the leaves of the latter are 
narrowly ovate with rounded bases, the up- 
per surface is slightly rugulose with slightly 
prominulous veinlets, the hairs of the un- 
dersurface are longer, and the inflorescence 
is more open and more spreading. 


Critoniopsis stellata 
(Spreng.) H. Robinson, 
comb. nov. 


Conyza stellata Spreng., Neue Entdeck. 
2:142. 1820. 

Vernonia oppositifolia Less., Linnaea 4:273. 
1829. 

Vernonia stellata (Spreng.) Blake, Contr. 
U.S. Natl. Herb. 22:587. 1924. 

Vernonanthura stellata (Spreng.) H. Rob- 
inson, Phytologia 72:74. 1992. 


Distribution. — Brazil. 

The species is mostly distinct by its large, 
opposite leaves. In spite of the name of the 
species, the hairs are not stellate. The recent 
transfer of the species to Vernonanthura was 
in error. 


Critoniopsis suaveolens 
(H.B.K.) H. Robinson, 
Phytologia 46:441. 1980. 


Vernonia suaveolens H.B.K., Nov. Gen. et 
Sp., ed fol. 4:30. 1818. 


VOLUME 106, NUMBER 3 623 


hu 


3 4 
uluult uuu 


- 


2 


wu 


hh 


centimet 


‘Fieldwork supported by the National Science Foundation 


BOLIVIA 


UNITED STATES a : it he ‘ 
3116536 


NATIONAL HERBARIUM 


J. 


C. Solomon S. Urcullo } 
MISSOURI BOTANICAL GARDEN HERBARIUM (MO) 


) 


Fig. 4. Holotype of Critoniopsis steinbachii H. Robinson, Solomon & Urcullo 14171 (US). 


624 


Distribution. —Ecuador. 

The type of the species is cited from Co- 
lombia, but all recent collections identified 
in this study are from Ecuador in Prov. Bo- 
livar (Zak & Jaramillo 2533, 2538, 2721, 
2740, MO, US) and Prov. Bolivar-Chim- 
borazo (Zak & Jaramillo 2802, MO, US). 


Critoniopsis tamana Badillo, 
Ernstia 53:10. 1989. 


Distribution. — Venezuela. 


Critoniopsis tequilana 
(S. Jones & Stutts) H. Robinson, 
comb. nov. 


Vernonia tequilana S. Jones & Stutts, Brit- 
tonia 33:544. 1981. 


Distribution. —Mexico. 


Critoniopsis tarchonanthifolia 
(DC.) H. Robinson, 
comb. nov. 


Monosis tarchonanthifolia DC., Prodr. 5:77. 
1836. 

Vernonia tarchonanthifolia (DC.) Schultz- 
Bip., Linnaea 20:507. 1847. 

Vernonia purpurascens Schultz-Bip. in 
Walp., Rep. Bot. Syst. 2:945. 1843. 

Oliganthes karwinskii Schultz-Bip., Lin- 
naea 20:505. 1847. 

Cacalia karwinskii (Schultz-Bip.) Kuntze, 
Rev. Gem. PE2:9702 189 

Eremosis tarchonanthifolia (DC.) Gleason, 
Bull. New York Bot. Gard. 4:230. 1906. 


Distribution. — Mexico. 


Critoniopsis tomentosa 
(La Llave & Lex.) H. Robinson, 
comb. nov. 


Turpinia tomentosa La Llave & Lex., Nov. 
Veg. Descr. 1:24. 1824, non Vernonia to- 
mentosa (Walt.) Ell., 1821. 

Vernonia paniculata DC., Prodr. 5:23. 1836. 

Monosis tomentosa (La Llave & Lex.) DC., 
Prodf’ 3:77. 1856. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Vernonia monosis Schultz-Bip., Linnaea 20: 
507. 1847. 

Cacalia tomentosa (La Llave & Lex.) 
Kuntze, Rev. Gen. Pl. 2:969. 1891. 

Cacalia paniculata (DC.) Kuntze, Rev. Gen. 
PL2:970. 139. 

Cacalia monosis (Schultz-Bip.) Kuntze, Rev. 
Gen. Pl. 2:970. 1891. 

Eremosis tomentosa (La Llave & Lex.) 
Gleason, Bull. New York Bot. Gard. 
4:229. 1906. 


Distribution. —Mexico. 


Critoniopsis triflosculosa 
(H.B.K.) H. Robinson, 
comb. nov. 


Vernonia triflosculosa H.B.K., Nov. Gen. et 
Sp., ed. fol. 4:40. 1818. 

Gymnanthemum congestum Cass., Dict. Sci. 
Nat. 20:1 10. 1821. 

Vernonia triantha Nees & Schauer, Linnaea 
19:714. 1847. 

Cacalia triflosculosa (H.B.K.) Kuntze, Rev. 
Genr PLi2:97 1 tse 

Cacalia triantha (Ness & Schauer) Kuntze, 
Rev. Gen. Pl. 2:971, 1891. 

Vernonia palmeri Rose, Contr. U.S. Natl. 
Herb 1: 101~ rs9or- 

Vernonia luxensis Coult., Bot. Gaz. (Craw- 
fordsville) 20:41. 1895. 

Vernonia dumeta Klatt., Bull. Soc. Roy. Bot. 
Belgique 35:277. 1896. 

Eremosis palmeri (Rose) Gleason, Bull. New 
York Bot. Gard. 4:233. 1906. 

Vernonia chacalana Blake, Contr. Gray 
Herbe52719S 1917: 


Distribution. —Mexico, Guatemala, El 
Salvador, Honduras, Nicaragua, Costa Rica, 
Panama. 


Critoniopsis tungurahuae 
(Benoist) H. Robinson, 
Phytologia 46:441. 1980. 


Vernonia tungurahuae Benoist, Bull. Soc. 
Bot. Fr. 83:804. 1936. 


Distribution. —Ecuador. 


VOLUME 106, NUMBER 3 


Critoniopsis turmalensis Badillo, 
Ernstia 53:12. 1989. 


Distribution. — Venezuela. 


Critoniopsis uniflora 
(Schultz-Bip.) H. Robinson, 
comb. nov. 


Vernonia uniflora Schultz-Bip., Linnaea 20: 
506. 1847. 

Eupatorium uniflorum Sess. & Moc., Pl. 
Nov. Hisp. 164. 1890, non Vernonia uni- 
flora Schultz-Bip. 

Cacalia uniflora (Schultz-Bip.) Kuntze, Rev. 
Gen. Pl. 2:968. 1891. 

Cacalia baptizanda Kuntze, Rev. Gen. PI. 
2:968. 1891. 


Distribution. —Mexico. 


Critoniopsis uniflosculosa 
(Cuatr.) H. Robinson, 
Phytologia 46:441. 1980. 


Vernonia uniflosculosa Cuatr., Bot. Jahrb. 
Syst. 77: 81. 1956. 


Distribution. —Colombia. 


Critoniopsis unguiculata 
(Cuatr.) H. Robinson, 
Phytologia 46:441. 1980. 


Vernonia unguiculata Cuatr., Bot. Jahrb. 
Syst. 77:80. 1956. 


Distribution. —Colombia. 

The species is similar to C. elbertiana in 
the recurved basal margins of its median 
involucral bracts, but it seems to be restrict- 
ed to the western or central cordilleras of 
Colombia, and the bases of its leaf blades 
seem more obtuse to nearly rounded. The 
type is from Depto. Valle, but specimens 
have been seen from Depto. Cauca (Luteyn, 
Luteyn & Morales 7445, NY, US) and Com. 
Choco (Silverstone-Sopkin 4493, US). 


Critoniopsis uribei H. Robinson, 
sp. nov. 


Plantae subarborescentes a 3 m altae in 
truncis robustae; caules leniter angulati 


625 


dense sordide tomentosi. Folia alterna, peti- 
olis 0.7—1.5 cm longis; laminae valde rigide 
coriaceae late oblongae plerumque 9-14 cm 
longae 4-9 cm latae base rotundatae mar- 
gine integrae in partibus anguste reflexae 
apice obtusae supra subnitidae minime 
prominule reticulo-venulosae minute glan- 
dulo-punctatae subtus dense brunneo-to- 
mentosae, nervis secundarlis pinnatis utrin- 
que ca. 7 late patentibus. Inflorescentiae 
terminales pyramidales in ramis dense cor- 
ymbiformes, bracteis in nodis alternis 2-4 
inferioribus foliiformibus 4—9 cm longis 1.5- 
4.0 cm latis, penduculis nullis. Capitula 15- 
17 mm alta cylindrica, bracteae involucri 
ca. 18 ovatae vel anguste ellipticae 2-10 mm 
longae 1.5—3.0 mm latae apice anguste ro- 
tundatae extus sparse vel dense tomentellae, 
marginis inferioribus in bracteis interiori- 
bus leniter reflexis. Flores ca. 4; corollae 
submaturae ca. 8 mm longae, tubis ca. 1.5 
mm longis, faucibus ca. 2 mm longis, lobis 
ca. 4.5 mm longis base ca. 0.8 mm latae 
extus distaliter glanduliferis; thecae anther- 
arum ca. 3.5 mm longae base distincte brev- 
iter truncatae caudatae; appendices anther- 
arum 0.4—0.5 mm longae induratae; basi 
stylorum immatura indistincte nodulosi, 
pilis stylorum multiseptatis apice obtusis vel 
rotundatis. Achenia submatura ca. 4 mm 
longa 8—10-costata glandulifera; setae pappi 
flavescentes 8-9 mm longae apice sensim 
distincte latiores, setae exteriores minutae 
indistinctae. Grana pollinis in diametro 50- 
55 um tricolporata spinulosa. 

Type: COLOMBIA: Boyaca: Cordillera 
Oriental, carretera de Duitama a Charala, 
mas alla del Paramo de La Rusia, 3000 m, 
12 Oct 1959, L. Uribe Uribe 3396 (holotype 
US). 

The species is known only from the type 


- specimen. Data with the type states “‘Ar- 


bolito pequeno, de 3 m de altura; tronco 
robusto; follaje oscuro: hojas en la haz de 
color verde muy subido; en al envés con 
tinte ceniciento o ferruginoso.” the only 
other members of the genus having heads 
as long are C. glandulata, which is almost 
wholly glabrous, and C. Auilensis, which has 


626 


narrower, toothed leaves with less pubes- 
cence. 


Critoniopsis ursicola 
(Cuatr.) H. Robinson, 
Phytologia 46:441. 1980. 


Vernonia ursicola Cuatr., Bot. Jahrb. Syst. 
77:82. 1956. 


Distribution. —Colombia. 


Critoniopsis weberbaueri 
(Hieron.) H. Robinson, 
comb. nov. 


Vanillosmopsis weberbaueri Hieron., Bot. 
Jahrb. Syst. 40:352. 1908. 

Vernonia ramospatana MacLeish, Syst. Bot. 
9:135. 1984. 


Distribution. — Peru. 

The species has a strong superficial re- 
semblance to C. quinqueflora of Brazil, but 
the two are not considered closely related. 
The anther appendages of the northern Pe- 
ruvian species have thickened cell walls, but 
the appendages of C. quinqueflora have thin 
walls, as in other southern species. 


Critoniopsis woytkowskii 
(S. Jones) H. Robinson, 
comb. nov. 


Vernonia woytkowskii S. Jones, Fieldiana, 
Bot. n.s. 5:29. 1980. 

Vernonia lambayequensis S. Jones, Fieldi- 
ana, Bot. n.s. 5:28. 1980. 


Distribution. — Peru. 

Isotypes have been seen of both V. woyt- 
kowskii and V. lambayequensis, and an ad- 
ditional specimen has been seen, Plowman, 
Sagastegui, Mostacero, Mejia & Pelaez 
14308 (NY, US), all from near Olmos, Dept. 
Lambayeque, Peru. Vernonia lambaye- 
quensis was originally stated to have pappus 
bristles ca. 9 mm long and corollas ca. 8 
mm long, but the isotype has pappus bristles 
only ca. 6 mm long and corollas only ca. 
6.5 mm long, scarcely longer than the mea- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


surements stated for V. woytkowskii. All the 
specimens have setuliferous achenes. The 
isotype of V. lambayequensis looks different 
primarily because the heads are less mature. 


Critoniopsis yamboyensis 
(Benoist) H. Robinson, 
comb. nov. 


Vernonia yamboyensis Benoist, Bull. Soc. 
Bot. France 83:804. 1936 [1937]. 


Distribution. — Ecuador. 

The species is unusual in the genus in 
having ovate rather than oblong or elliptical 
leaf blades. 


Critoniopsis yungasensis 
(Britton) H. Robinson, 
comb. nov. 


Vernonia yungasensis Britton, Bull. Torrey 
Bot. Club. 18:332. 1892. 


Distribution. — Bolivia. 

Co-types from the New York Botanical 
Garden have been examined and Rusby 
1732 stamped from Columbia College Her- 
barium is designated here as lectotype. 


Acknowledgments. 


Jose Cuatrecasas of the Smithsonian In- 
stitution and Santiago Diaz Piedrahita of 
the Universidad Nacional de Colombia, 
Bogota made helpful comments during the 
study. Photographs of holotypes were pre- 
pared by Victor E. Krantz, Staff Photogra- 
pher, National Museum of Natural History. 


Literature Cited. 


Badillo, V. M. 1983. Nuevas combinaciones 0 si- 

nonimia en Compositae de Venezuela. —Ernstia 

16:16. 

. 1989. Dos especies nuevas del genero C7i- 

toniopsis y otra del genero Vernonia (Compos- 

itae).—Ernstia 53:10-14. 

Baker, J. B. 1873. Compositae. I. Vernoniaceae. Jn 
C. F. P. Martius, ed., Flora brasiliensis 6(2):1— 
180. 

Cuatrecasas, J. 1956. Neue Vernonia-Arten und Syn- 


VOLUME 106, NUMBER 3 


opsis der andinen Arten der Sektion Critoniop- 
sis. — Botanische Jahrbicher fiir Systematik 77: 
52-84. 

Dillon, M. O. 1984. Two new species of Vernonia 
(Asteraceae: Vernonieae) from Peru.—Brittonia 
36:333-336. 

Gleason, H. A. 1906. A revision of the North Amer- 

ican Vernonieae. — Bulletin of the New York Bo- 

tanical Garden 4:144—243. 

1922. Vernonieae.—North American Flora 
33:47-101. 

1923. The Bolivian species of Vernonia. — 
American Journal of Botany 10:297-309. 
Jones, S. B. 1973. Revision of Vernonia section Er- 
emosis (Compositae) in North America. — Brit- 
tonia 25:86-115. 

1974. Vernonieae (Compositae) chromo- 
some numbers.— Bulletin of the Torrey Botan- 
ical Club 101:31-34. 

1979. Chromosome numbers of Vernonieae 
(Compositae).— Bulletin of the Torrey Botanical 
Club 106:79-84. 

— & J. G. Stutts. 1981. Three new species of 
Vernonia (Compositae: Vernonieae) from Mex- 
ico.—Brittonia 33:544—-546. 

La Llave, P. & J. Lexarza. 1824. Novorum vegeta- 
bilium descriptiones, fasc. 1. Mexico. viii + 32 
pp. 

Pruski, J. 1992. Compositae of the Guayana High- 
lands—VI. Huberopappus maigualidae (Ver- 
nonieae), a new genus and species from Vene- 
zuela.—Novon 2:19-25. 

Robinson, H. 1980. Re-establishment of the genus 
Critoniopsis (Vernonieae: Asteraceae).— Phy- 
tologia 46:437-442. 


627 


1989. Two new genera of Vernonieae (As- 
teraceae) from the northern Andes with dis- 
sected corolla limbs Cuatrecasanthus and Jo- 
seanthus. — Revista de la Academia Colombiana 
de Ciencias Exactas, Fisicas y Naturales 17:207-— 
23! 

1990a. Studies in the Lepidaploa Complex 
(Vernonieae: Asteraceae) VII. The genus Lepi- 
daploa.—Proceedings of the Biological Society 
of Washington 103:464-498. 

1990b. New combinations in the Asteraceae 
(Vernonieae, Heliantheae, Mutisieae).— Phyto- 
logia 69:105-107. 

1992a. A new genus Vernonanthura (Ver- 
nonieae, Asteraceae).— Phytologia 73:65—76. 

1992b. Notes on Lychnophorinae from Mi- 
nas Gerais, Brazil, a synopsis of Lychnophorop- 
sis Schultz-Bip., and the new genera Antere- 
manthus and Minasia  (Vernonieae: 
Asteraceae). — Proceedings of the Biological So- 
ciety of Washington 105:640-652. 

—., F. Bohlmann, & R. M. King. 1980. Che- 
mosystematic notes on the Asteraceae. III. Nat- 
ural subdivisions of the Vernonieae.—Phyto- 
logia 46:421-436. 

Schultz-Bipontinus, C. H. 1863. Geschichte der Gat- 
tung Lychnophora. —Jahresbericht der Pollichia 
20/21:329-439. 


Department of Botany, NHB-166, Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. 20560, 
WES-A: 


PROC. BIOL. SOC. WASH. 
106(3), 1993, p. 628 


BIOLOGICAL SOCIETY OF WASHINGTON 
120th Annual Meeting, 19 May 1993 | 


The meeting was called to order by Storrs Olson, President, at 12:30 p.m. in the Waldo 
Schmitt Room, National Museum of Natural History. 

Storrs briefly summarized this year’s activities: 

Chad Walter presented the Treasurer’s Report. Total income for the period of 1 January 
1992 to 31 December 1992 was $98,362, and total expenditures were $82,143. Expen- 
ditures included management costs of $8,770 paid to Allen Press for three quarters {issues 
1-3, Vol. 105}. Chad noted that the costs for issue 105-4 were not included in 1992 
expenditures, that follow-up of many unpaid FY91-—92 invoices were received in FY92, 
and that sales of back issues were unusually high. He thus cautioned that the net increase 
for the year of $16,219 was unusual and should not be expected annually. 

Storrs announced that the Council had voted to increase membership dues in a two- 
tiered fashion, as: Annual dues $25.00 (for USA and non-USA addresses), and; Library 
subscriptions $40.00 for USA and non-USA addresses. Non-USA members or subscribers 
may pay an additional $25.00 to receive the Proceedings by AirMail. 

Brian Robbins, Editor, then presented his report. The four issues of Volume 105 of the 
Proceedings were published on 12 March, 11 June, 15 October, and 18 December 1992, 
containing a total of 90 papers and 896 pages. There were 93 submissions in 1992, down 
from 101 in 1991, and as of 1 May 1993, submissions were running even with the total 
for the same period in 1992. There is no current backlog. Brian noted that the new Associate 
Editor for Vertebrate Zoology is Thomas A. Munroe, replacing G. David Johnson. 

Stephen Cairns (Council Member) presented the results of an analysis he conducted of 
the contents of the Proceedings by major taxonomic groupings. The impetus for this 
analysis was a perception on the part of some that there are an inordinate number of 
papers published on Crustacea. The results show that over the past ten years (Volumes 
96-105) the Proceedings has published papers on 30 of the 33 phyla with the following 
breakdown: 38.5% crustaceans; 35.5% other invertebrates; 23% vertebrates; 2.3% botany; 
0.5% general. Over the past six years the number of crustacean papers has increased at 
the expense of all other invertebrates. Contributions on vertebrates have remained re- 
markably constant at about 21%. 

The meeting was adjourned at 1:00 p.m. 


Respectfully submitted, 
G. David Johnson 
Secretary 


INFORMATION FOR CONTRIBUTORS 


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CONTENTS 


A new species of a giant Thomasomys (Mammalia: Muridae: Sigmodontinae) from the Andes 
of northcentral Peru Mariella Leo L. and Alfred L. Gardner 
Zoogeography and geographic variation of At/apetes rufinucha (Aves: Emberizinae), including 
a distinctive new subspecies, in southern Peru and Bolivia J. V. Remsen. Je 
A new hybrid manakin (Dixiphia pipra x Pipra filicauda) (Aves: Pipridae) from the Andean 
foothills of eastern Ecuador Gary R. Graves 
A new species of aquatic Bufo (Anura: Bufonidae) from cloud forests in the Serrania de Siberia, 
Bolivia. Michael B. Harvey and Eric N. Smith 
Revision of Y/asoia Speiser, 1920 (Insecta: Diptera: Bombyliidae: Lomatiinae) 
Marcia Souto Couri and Carlos José Einicker Lamas 
New records of entocytherid ostracods infesting burrowing and cave-dwelling crayfishes, with 
descriptions of two new species Horton H. Hobbs, Jr., and Daniel J. Peters 
Scopalatum vorax (Esterly, 1911) and Scolecithricella lobophora Park, 1970, calanoid copepods 
(Scolecitrichidae) associated with a pelagic tunicate in Monterey Bay 
Frank D. Ferrari and Deborah K. Steinberg 
A new species of Benthana Budde-Lund from Brazilian caves (Crustacea; Isopoda; Oniscoidea) 
Idalina Maria Brasil Lima and Cristiana Silveira Serejo 
Linca pinita, a new phoxocephalid genus and species (Crustacea: Amphipoda) from the Ar- 
gentine continental shelf Gloria M. Alonso de Pina 
Three new species of Strengeriana from Colombia (Crustacea: Decapoda: Pseudothelphusidae) 
Martha R. Campos and Gilberto Rodriguez 
Further remarks on the identity of Sudanonautes orthostylis Bott, 1955, (Crustacea: Decapoda: 
Potamoidea: Potamonautidae) with comparisons with other species from Nigeria and Cam- 


eroon Neil Cumberlidge 
Three genera removed from the synonymy of Pinnotheres Bosc, 1802 (Brachyura: Pinno- 
theridae) Raymond B. Manning 
A new subgenus and species of crayfish (Decapoda: Cambaridae) of the genus Cambarus, with 
an amended description of the subgenus Lacunicambarus Raymond F. Jezerinac 
Two new records of the genus Heptacarpus (Crustacea: Decapoda: Hippolytidae) from Japanese 
waters Tomoyuki Komai 


Two new hermit crabs (Crustacea: Decapoda: Paguridae) from the Caribbean Sea 
Rafael Lemaitre and Néstor H. Campos 
Stomatopod Crustacea from Tobago, West Indies 
Marilyn Schotte and Raymond B. Manning 
A new species of Sphaerodoridae (Annelida: Polychaeta) from southern California 
Jerry D. Kudenov 
A new marine species of Smithsonidrilus (Oligochaeta: Tubificidae) from the Florida Keys 
Christer Erséus 
Two new species of Phascolion (Sipuncula: Phascolionidae) from tropical and subtropical waters 
of the central western Atlantic Mary E. Rice 
An unusual squid paralarva (Cephalopoda) with tentacular photophores 
Deborah L. Loffler and Michael Vecchione 
A review of the genus Critoniopsis in Centrai and South America (Vernonieae: Asteraceae) 
Harold Robinson 
Biological Society of Washington: 120th Annual Meeting 


417 


429 


436 


442 


450 


455 


467 


490 


497 


508 


514 


523 


332 


545 


554 


566 


582 


587 


591 


602 


606 
628 


~ PROCEEDINGS 


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BIOLOGICAL SOCIETY 


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VOLUME 106 NUMBER 4 


17 DECEMBER 1993 


ISSN 0006-324X 


THE BIOLOGICAL SOCIETY OF WASHINGTON 


1992-1993 
Officers 
President: Storrs L. Olson Secretary: G. David Johnson 
President-elect: Janet W. Reid ‘Treasurer: T. Chad Walter 
Elected Council 

Stephen D. Cairns Jon L. Norenburg 

Richard C. Froeschner Lynne R. Parenti 

Alfred L. Gardner F. Christian Thompson 


Custodian of Publications: Austin B. Williams 


PROCEEDINGS 


Editor: C. Brian Robbins 


Associate Editors 


Classical Languages: George C. Steyskal Invertebrates: Jon L. Norenburg 
Frank D. Ferrari 

Plants: David B. Lellinger Rafael Lemaitre 

Insects: Wayne N. Mathis Vertebrates: Thomas A. Munroe 


Membership in the Society is open to anyone who wishes to join. There are no prerequisites. 
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Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biological 
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POSTMASTER: Send address changes to PROCEEDINGS OF THE BIOLOGICAL SOCIETY 
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PROC. BIOL. SOC. WASH. 
106(4), 1993, pp. 629-632 


A NEW SPONGE SPECIES, CERATOPSION CRUSTOSUM 
(DEMOSPONGIAE: RASPAILITDAE), FROM DEEP 
WATERS OF THE GULF OF MEXICO 


Belinda Alvarez and R. W. M. Van Soest 


Abstract.—A new sponge, Ceratopsion crustosum from deep waters (70-159 
m) of the Gulf of Mexico is described. It is the first record of a Ceratopsion 
from the Gulf, or indeed from Atlantic waters. The new species is a typical 
species of Raspailiidae with an axial, extra-axial, and ectosomal skeleton, and 
with all the diagnostic characters of Ceratopsion Strand, 1928, except for the 
occurrence of trichodragmata. The definition of Ceratopsion is expanded to 
include species having these microscleres. 


During a 5-year survey by the Mineral 
Management Service (United States De- 
partment of the Interior), off southwest 
Florida in the Gulf of Mexico, a large num- 
ber of sponges were collected and deposited 
at the National Museum of Natural History, 
Smithsonian Institution. The Axinellidae is 
one of the best represented families in this 
collection in terms of number of species and 
number of specimens of each species. The 
presence of several undescribed species, and 
the high intraspecific morphological varia- 
tion that has not been described previously, 
provided the incentive for a regional revi- 
sion of the Axinellidae (in preparation). 
Amongst the material selected for this re- 
vision, a new species of Ceratopsion Strand, 
1928 was found, dredged between 70 and 
159 m depth. This genus, formerly in the 
Axinellidae, is now placed in the Raspaili- 
idae, Order Poecilosclerida (Hooper 1991). 

Specimens were preserved in alcohol. 
Spicule slides and both thick and polished 
sections were prepared using the methods 
described by Riitzler (1978). Abbreviations 
used in the text are: USNM, National Mu- 
seum of Natural History, Smithsonian In- 
stitution (formerly United States National 
Museum), ZMA, Zoological Museum of 
Amsterdam. 


Family Raspailiidae Hentschel, 1923 
Genus Ceratopsion Strand, 1928 
Ceratopsion crustosum, new species 
Fig. 1 


Material examined. —Holotype: USNM 
42808, alcohol, 76 m, off Florida Keys, 
24°47'25’N, 83°51'09"W, coll. 25 April 
1981. Paratypes: USNM 42809 (alcohol), 
ZMA 10070, ZMA 10071 [alcohol, 76 m, 
off Florida Keys, 24°47'25’N, 83°51'09’W, 
coll. 25 Apr 1981]. 

Additional material.—USNM 41577, al- 
cohol, 70 m, Southwest Florida, off Cape 
Sable; .25716'53” Naw3°37'47'W,; colladd 
Nov 1980; USNM 42807, alcohol, 70 m, 
Southwest Florida, off Cape Sable, 
25°16'53’"N, 83°37'47’W, coll. 8 Feb 1982; 
USNM 42810, alcohol, 159 m, South- 
west Florida, off Naples, 25°44’'50’N, 
84°21'02”W, coll. 7 Feb 1982. 

Description. —Shape: One or more thin 
laminae on short peduncle, coalescing at 
some points, or folded, forming calycles 
(cups), with margins forked, undulating or 
uneven (Fig. 1a). 

Surface: Hispid from projecting long spic- 
ules. 

Consistency: Firm. 

Skeleton (Fig. 1b, c): Ectosomal, extra- 


630 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


t 
(ee aC ae Ad tig Sung eas Dae 
ct a om sf ke ae ay z . & ~~ Ae 


Fig. 1. Ceratopsion crustosum, new species. a, Holotype, USNM 42808; b, micrograph of longitudinal section 
of the skeleton; c, schematic drawing of a longitudinal section of the skeleton; d, micrographs of trichodragmata 
in the ectosome; e, choanosomal strongyles and modifications; f-g, ectosomal oxeas, h, ectosomal trichodragmata 
and raphides. Scales: a, 1 cm, b, 200 um, c, not to scale, d, 67 wm, e—-h, 100 um. 


VOLUME 106, NUMBER 4 


axial and axial skeleton differentiated. Ec- 
tosomal skeleton formed by continuous and 
compact dermal crust or palisade (100-350 
um) of oxeas, raphides and trichodragmata, 
that are generally scattered or sometimes 
arranged in sparse bundles, 100 wm thick 
approximately, oriented perpendicularly to 
surface; in places with round apertures of 
80-100 um diameter. Extra-axial skeleton 
generally obscured by leaf-shape habit of 
sponge; with single and sparse styloids, 
mostly broken, protruding from choano- 
some to surface through ectosome and areas 
with aspiculous sheets of spongin. Choano- 
somal skeleton axially condensed, com- 
pressed in cross section, with large stron- 
gyles and styloids oriented longitudinally in 
relation to axis. 

Choanocyte chambers: Circular to oval 
(35—48 um). 

Spicules (Fig. 1d—h; Table 1): Strongyles, 
typical or modified to styles and styloids 
(670-1300 um in length), some sinuous. Two 
size categories of oxeas (150-330 um and 
40-100 um in length); smaller ones bent at 
center or s-shaped. Raphides, some ar- 
ranged as trichodragamata (17.5—250 um in 
length). 

Discussion.—The newly described spe- 
cies shares characters typical of both Thri- 
nacophora Ridley & Dendy, 1886 and Cer- 
atopsion Strand, 1928. These genera, which 
have historically been placed within the Ax- 
inellidae, are considered by Hooper (1991) 
to belong to the Raspailiidae. Both genera 
according to Hooper are very similar in 
many characters including a specialized ec- 
tosomal skeleton, large extra-axial mega- 
scleres, a more-or-less radial arrangement 
of the extra-axial skeleton that protrudes a 
long way through the surface, and well de- 
veloped axial and extru-axial differentia- 
tion. The major differences between these 
genera are, a) the spicules in the axial skel- 
eton (long sinuous styles, strongyles or an- 
isoxeas in Ceratopsion; short, stout oxeas in 
Thrinacophora), b) the organization of the 
spicules in the axial skeleton (densely packed 


631 


Table 1.—Spicule dimensions for Ceratopsion crus- 
tosum. Measurements (in wm) are ranges of 25 spicules 
(or the number indicated in brackets) with means + 
standard deviation in parentheses. 


Holotype USNM 42808 


Strongyles 

Length 570-1300 (842.8 + 156.3) 

Width 10-25 (17.2 + 4.5) 
Oxeas I 

Length 150-290 (224.8 + 36.5) 

Width 2.5-7.5 (6:7 (1A) 
Oxeas II 

Length 42.5-100 (75:4 == 7229) 

Width 2.5—2.5 25/00) 
Trichodragmata 

Length 70-200 (154.6 + 38.0) [13] 

Width 5-10 (FASTA) [IS] 

Paratype USNM 42809 

Strongyles 

Length 670-1175 (948.4 + 129.9) 

Width 10-20 (i5i' 22328) 
Oxeas I 

Length 170-330 (260.4 + 38.0) 

Width 5-10 (73 =-14) 
Oxeas II 

Length 40-100 (2 Als h5,7) 

Width 2.5—2.5 (25,-=,. 0:0) 
Trichodragmata 

Length 17.5—250 (138:6+.70:3) 

Width 5-17.5 (9.2 st 27) 


in Ceratopsion; criss-cross in Thrinaco- 
phora), c) the presence of raphides and tri- 
chodragmata (absent in Ceratopsion; pres- 
ent in Thrinacophora). 

The new species was compared with Cer- 
atopsion ramosum Thiele, 1898 (fragment 
of type specimen from Berlin Museum No. 
957) and Thrinacophora funiformis Ridley 
& Dendy, 1886 and found to be similar in 
structure to both of them. The presence of 
raphides and trichodragmata in Thrina- 
cophora funiformis and the new species, but 
absent in C. ramosum and other species of 
Ceratopsion, could be a reason to allocate 
the new species to Thrinacophora. How- 
ever, the new species lacks the reticulate axis 


632 


of short, stout choanosomal oxeas, and criss- 
cross of axial spicules typical of Thrinacoph- 
ora. As the presence of raphides and tri- 
chodragmata is considered less important 
(because of wide-spread occurrence in many 
families and genera of the Demospongiae) 
than the architecture of the skeleton the new 
species is placed within Ceratopsion. The 
definition of Ceratopsion given by Hooper 
(1991) should therefore be emended to in- 
clude species with raphides and trichodrag- 
mata. 

Ceratopsion crustosum represents the first 
species of Ceratopsion recorded in Atlantic 
waters. Other species of Ceratopsion (see 
Hooper 1991:1328) have been reported from 
Japan, Indonesia, New Zealand, Australia, 
South Africa, and the Mediterranean. 

Etymology. — From the Latin crusta, crust 
referring to the dermal crust of oxeas that 
forms the ectosome. 


Acknowledgments 


Specimens were made available by the 
Mineral Management Service (formerly, 
Bureau of Land and Management, U.S. De- 
partment of Interior) through Dr. Kristian 
Fauchald, National Museum of Natural 
History (NMNH), Smithsonian Institution. 
Equipment and material was provided by 
the NMNH. Dr. Klaus Rutzler and Ms Kate 
Smith (NMNH), helped with the polished 
sections, photographic prints and provided 
constructive comments. Mr. Michael Car- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


penter (NMNH) assisted with photographic 
work. Dr. John Hooper, Queensland Mu- 
seum and Dr. Christopher Glasby, Austra- 
lian Biological Resource Studies, reviewed 
the manuscript and made important sug- 
gestions. 


Literature Cited 


Hentschel, E. 1923. Erste Unterabteilung der Meta- 
zoa. Parazoa. Einziger Stamm und einzige 
Klasse der ersten Unterabteilung: Porif- 
era=Schwamme.—Handbuch der Zoologie 
1:307-417. 

Hooper, J. N. A. 1991. Revision of the family Ras- 
pailiida (Porifera: Demospongiae), with descrip- 
tion of Australian species.—Invertebrate Tax- 
onomy 5:1179-1418. 

Ridley, S. O., & A. Dendy. 1886. Preliminary report 
on the Monoaxonida collected by H.M.S. 
“Challenger”. Part II.—The Annals and Mag- 
azine of Natural History, serie 5, 18:470-493. 

Ritzler, K. 1978. Sponges in coral reefs. Pp. 299- 
313 in D. R. Stoddart & R. E. Johannes, eds., 
Coral reefs: research methods. Monographs on 
Oceanographic Methodology, 5, Unesco, Paris. 

Strand, E. 1928. Miscellanea nomenclatoria zoolo- 
gica et palaentologica.—Archiv fir Naturge- 
schichte, Berlin, serie A, 92(8):30-—75. 

Thiele, J. 1898. Studien tiber pazifische Spongien. — 
Zoologica Heft. 24:1-72, pls. 1-8. 


(BA) Australian National University, Di- 
vision of Botany and Zoology, Canberra, 
ACT, 0200, Australia; (RWMS) Institute of 
Taxonomic Zoology, University of Am- 
sterdam, P.O. Box 4766-1009 at Amster- 
dam, The Netherlands. 


PROC. BIOL. SOC. WASH. 
106(4), 1993, pp. 633-644 


A NEW SPECIES OF FRESHWATER PLANARIAN FROM 
CHILE (PLATYHELMINTHES: TRICLADIDA), WITH 
A NOMENCLATURAL NOTE ON GIRARDIA FESTAE 
(BORELLI, 1898) 


Alejandro C. Curino and Néstor J. Cazzaniga 


Abstract. —A new species of triclad, Girardia canai (Paludicola, Dugesiidae), 
is described from the Mapocho River within the Region Metropolitana of Chile. 
The species has the following diagnostic characters: ventral testes; bursal canal 
forming a right angle at its opening into the genital atrium; epithelia of the 
atrium and bursal canal with intraepithelial nuclei; course of the sperm ducts 
very variable, generally asymmetric; the sperm ducts do not form intrapenial 
enlargements; instead, they change their histology abruptly, adopting a histo- 
logical structure identical to that of the ejaculatory duct; the union of these 
slender ducts occurs within the penis papilla; penis papilla notably long. The 
intrapenial structure of this species is of a new morphological type, and is 
considered a main attribute for defining the species. The aforementioned fea- 
tures distinguish the new species from all other Girardia species. Particular 
emphasis is placed in the discussion on the comparison with G. tigrina (Girard), 
G. rincona (Marcus), G. longistriata (Fuhrmann) and G. microbursalis (Hyman). 
The nomenclature of Girardia festae (Borelli, 1898) is reviewed on the grounds 


that the commonly used spelling G. festai is an unjustified emendation. 


Girardia Ball, 1974 (Paludicola: Dugesi- 
idae) is a Neotropical genus of freshwater 
planarians encompassing more than 30 de- 
scribed species which are for the most part 
very closely related to one another and show 
only small morphological differences. The 
phylogenetic position of this genus within 
the Dugesiidae family was investigated re- 
cently by de Vries & Sluys (1991). 

In 1986 planarians were collected from 
two streams, Rio Mapocho and Estero Re- 
Naca, near Vina del Mar, Chile, where there 
are previous records of Girardia chilla 
(Marcus, 1954) and G. sanchezi (Hyman, 
1959). We were particularly interested in G. 
sanchezi since Hyman (1959) suggested that 
it could be conspecific with G. anceps (Kenk, 
1930), for which we have already published 
a redescription (Cazzaniga & Curino 1987). 
However, the planarians we collected be- 
long to a new species of the genus Girardia, 
as described below. 


Material and methods.—The specimens 
from both localities were collected by the 
authors together with Dr. Patricio Sanchez 
on 26 October 1986. They were found under 
stones, mainly in backwater areas of the riv- 
ers. 

Before fixation in F.A.A. fluid (formal- 
dehyde-ethyl alcohol-acetic acid), the spec- 
imens were placed on ice in order to induce 
muscular relaxation. The material was se- 
rially cut at 7—9 um intervals into sagittal, 
transverse and frontal sections which were 
then stained in hematoxylin and eosin. 


Girardia candi, new species 
Figs. 1-9 


Diagnosis.—Girardia species with ven- 
tral testes; bursal canal forming a right angle 
at its opening into the genital atrium. Bursa 
copulatrix medium to large in size. Epithelia 
of the atrium and of the bursal canal with 


634 


intraepithelial nuclei. Course of sperm ducts 
very variable, generally asymmetric. No in- 
trapenial sperm vesicles. Instead, the sperm 
ducts change their histology abruptly, form- 
ing two slender ducts structurally identical 
to the ejaculatory duct with a tall epithelium 
of infranucleate cylindrical cells, pale cy- 
toplasm and a very narrow lumen. The ducts 
fuse within the penis papilla and the ejac- 
ulatory duct ends terminally. The penis pa- 
pilla is notably long. 

Distribution. —Type locality: Mapocho 
River, in the vicinity of the town of Tala- 
gante, Region Metropolitana, Chile. 

The other locality where they have been 
found is near the mouth of the Estero Rena- 
ca, to the south of Concon, V Region, Chile. 

Type material. —The type series includes 
19 sets of serial sections deposited as fol- 
lows. 

Museo Argentino de Ciencias Naturales 
‘““Bernardino Rivadavia’’ (MACN), Buenos 
Aires: Holotype: a set of sagittal serial sec- 
tions on seven slides (MACN #33438). 
Paratype #1: a set of transverse serial sec- 
tions on eight slides, specimen from the type 
locality (MACN #33439). 

American Museum of Natural History 
(AMNH), New York: Paratypes #2 and 3: 
a set of transverse serial sections on nine 
slides, specimen from the type locality 
(AMNH #1595); a set of sagittal serial sec- 
tions on 12 slides, specimen from Estero 
Renaca (AMNH #1596). 

Laboratorio de Ecologia Acuatica, Uni- 
versidad Nacional del Sur, Bahia Blanca: 
Paratypes #4 to 19: ten sets of sagittal serial 
sections, four sets of transverse serial sec- 
tions and one set of frontal serial sections. 
Thirteen specimens from the type locality 
and two from Estero Renaca. 

Description. —The fixed planarians are 9- 
10 mm in length and 2—4 mm in width. The 
head has the typical triangular shape of the 
genus Girardia, with visible but not con- 
spicuously large, rounded auricles. The 
ground pigmentation is light brown, with 
dark brown, almost black spots, the density 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


of which is very variable. In some speci- 
mens the spots are very abundant and de- 
limit a light longitudinal middorsal stripe; 
in others the spots are scarce and no stripe 
is apparent. The size and position of the eye 
cups conform to that described for all the 
species of Girardia from South America. 

The body wall has a cylindrical to cubic 
epithelium underlain by a weakly devel- 
oped subepidermal layer of circular muscles 
and a well-developed layer of longitudinal 
muscles; radial muscles were not discerned. 
The rhabdite-forming cells are located be- 
neath the muscular layers. The marginal ad- 
hesive glands form a ring round the ventral 
surface; they are very developed, strongly 
eosinophilous, and their secretion is gran- 
ular in appearance. 

The digestive system is typical for tri- 
clads. The main anterior branch of the gut 
extends briefly past the level of the ocelli, 
but does not extend anteriorly to the brain. 
The pharynx is of the dugesiid type. The 
macroscopic aspect of the pharynx was not 
observed in the live material. No epidermal 
or subepidermal pigmentary cells are ap- 
parent in the histological sections. 

The ovaries are ovoid in shape and lo- 
cated directly posterior to the brain branch- 
es, with a slight latero-medial displacement. 
The funnel-shaped oviducal tuba of each 
ovovitelline duct opens onto the lateral ex- 
ternal face of the respective ovary. The ovi- 
ducal tubae were filled with sperm in many 
of the specimens examined. 

The ovovitelline ducts have an infranu- 
cleate cubic epithelium surrounded by a cir- 
cular muscular layer. In transverse section, 
these ducts are surrounded by a layer of 
radially disposed nuclei, similar to that de- 
scribed for other Girardia and Schmidtea 
species (Marcus 1946, Schilt 1976). 

The ovovitelline ducts run dorsally along 
the nerve cords up to shortly before the gon- 
opore, where they curve upward and run 
toward the middle-sagittal plane; here they 
bend anteriorly and finally open into the 
bursal canal. In the majority of the speci- 


VOLUME 106, NUMBER 4 


BC 


SE 


635 


Za 
\a 


. 
1 


BO hs 


EDB MF G CG 


Figs. 1-2. Girardia canai, new species. Sagittal reconstructions of the copulatory apparatus. BC: bursa cop- 
ulatrix; BG: basophilous glands; BS: bursal canal; CG: cement glands; ED: ejaculatory duct; EDB: anterior 
branch of the ejaculatory duct; G: gonopore; MF: muscular fold; O: ovovitelloduct; S: sperm duct; SE: extrapenial 
seminal vesicle; SG: shell glands. Note that only the epithelia of S, EDB and ED are drawn. The diameter of S 
appears smaller than it in fact is, since the surrounding muscular coat is not depicted; EDB and ED do not have 


a muscular coat. 


mens examined the ducts open separately, 
but very closely together, into the postero- 
dorsal face of the bursal canal. In a few cases 
the openings are somewhat more separated. 
The posterior-most portion of the ovovi- 
telline ducts shows a change in histological 
structure, being lined with a normal, and 
not infranucleate, epithelium. 

The yolk glands have large polyhedrical 
cells with large nuclei. Their cytoplasm is 
eosinophilous and vacuolate. These glands 
occur mainly between and on the divertic- 


ula of the gut. In the majority of the spec- 
imens examined a variable number of these 
glands were observed to be present anteri- 
orly to the ovaries. 

The bursa copulatrix has a tall-celled ep- 
ithelium with basal nuclei (Figs. 3, 4), the 
cells being thicker at the apical end than at 
the base. These cells frequently have a vac- 
uole at their broader end, in the interior of 
which it is sometimes possible to observe 
masses of sperm in digestion. The size of 
the bursa copulatrix can vary greatly from 


636 


Figs. 3-5. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


= 


aig, 
‘ 
* 
ay 


Girardia canai, new species. 3-4, Sagittal sections of the holotype. 5, Sagittal section of paratype #4. 


VOLUME 106, NUMBER 4 


small to very large, sometimes filling the 
entire height of the body. No correlation 
was found between size and sperm content. 

The bursal canal originates from the cau- 
do-upper part of the bursa, running in a 
smooth curve over the penial complex. At 
its posterior end it bends downward to form 
a right angle and opens into the common 
atrium. The shell glands open into the bur- 
sal canal immediately below the opening of 
the ovovitelline ducts. These glands are 
strongly eosinophilous and very developed. 
The distal portion of the bursal canal is dis- 
placed to the left in the majority of the spec- 
imens examined, opening into the latero- 
dorsal portion of the common atrium; in 
only a few specimens is a sagittal course 
followed. The histological structure of the 
bursal canal (Fig. 7) shows a tall, cylindrical 
ciliate epithelium of nucleate cells, together 
with two muscle layers, one of which is sub- 
epidermal and circular and the other exter- 
nal and longitudinal. No other muscle layers 
or muscular additions are observed at the 
distal portion. 

The testes were ventral in all of the stud- 
ied specimens, extending from close behind 
the ovaries almost to the posterior end of 
the body. In only one of the specimens ex- 
amined was a testicular follicle present an- 
teriorly to one of the ovaries. As usual in 
the genus, the testes are at both sides of the 
nerve cords, with the exception of the phar- 
ynx and the genital complex zone, where 
they occur only on the outside. They are 
very abundant and show a continuous dis- 
tribution. 

The extrapenial seminal vesicles or false 
seminal vesicles are generally well devel- 
oped, forming two wide and sinuous tubes 
full of sperm originating at the level of the 
anterior half of the pharynx. 

Near the anterior face of the penis bulb 
or the origin of the papilla the extrapenial 
seminal vesicles abruptly become thinner 
and give rise to the sperm ducts. These ducts 
have a thin cavity and a cubic eosinophilous 
epithelium with normal intraepithelial nu- 


637 


clei and a well-developed circular muscle 
layer. The course of the ducts is extraordi- 
narily variable from one specimen to an- 
other and even from one side of the same 
organism to the other. Of the 16 completely 
mature individuals examined, 11 showed 
asymmetry in their course. There is no ev- 
idence to suggest that this variation is due 
to different states of contraction. Three main 
schematic models can be described for the 
course of the sperm ducts: 


1 The sperm duct runs upward and toward 
the bulb, curves both frontward and to- 
ward the middle of the body, penetrates 
into the bulb through the anterior-upper 
face and finally runs downward through 
the bulb (Fig. 1 on the right side of the 
specimen and Fig. 2 on the left side). This 
course clearly forms a loop similar to that 
described for Girardia anceps, though it 
never reaches such a dorsal position nor 
does it curve as far forward as it does in 
the latter species (Cazzaniga & Curino 
1987). 

2 The duct runs upward, curves toward the 
middle of the body, penetrates into the 
bulb through the lateral-upper face and 
then runs downward, the whole course 
running on the same transversal plane 
(Fig. 2 on the right side of the specimen), 
1.e., the ducts do not form a loop. 

3 The duct does not run upward, or does 
so only slightly and then goes directly to- 
ward the bulb, entering through the lat- 
eral-inferior face (Fig. | on the left side). 
This course can be straight or sinuous, 
forming an “‘S”’ on a horizontal plane. 


The asymmetry most frequently ob- 
served was a loop or type-1 course on one 
side and a type-3 course on the other side 
of the same specimen (Fig. 6). 

In all cases the sperm ducts run a long 
way into the interior of the bulb without 
becoming true intrapenial seminal vesicles, 
1.e., they do not form intrabulbar enlarge- 
ments. Instead, they transform into two 


638 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 6-9. Girardia canai, new species. 6, Transverse section of paratype #1. 7, Enlargement of the bursal 
canal of the holotype to show the presence of two muscle layers, the internal layer being circular. 8, Frontal 
section of the penis of paratype #5, illustrating the structure of the anterior branches of the ejaculatory duct 
uniting in the penis papilla. 9, Enlargement of figure 1. CM: circular musculature; EB: cephalic branch of the 
ejaculatory duct; ED: ejaculatory duct; LM: longitudinal musculature; S: sperm duct. 


slender ducts with the same histological 
structure as the ejaculatory duct (Figs. 8, 9). 

The course of the intrabulbar portion of 
the sperm duct varies greatly. When it runs 
upward after entering the bulb the histolog- 
ical transformation occurs in the upper-pos- 
terior zone of the latter; in one of the spec- 
imens examined, the duct runs first upward 
and then anteriorly, forming an intrabulbar 
loop. In others the duct runs downward and 
even in a lateral direction, so that the his- 
tological transformation of the duct takes 


place at some point between the posterior 
half of the bulb and the base of the papilla. 
In the type-3 course the duct ends with a 
right-angled curve towards the papilla and 
immediately changes its structure. 

The histological transformation of the 
sperm duct is abrupt. In the 16 mature spec- 
imens examined, the narrow lumen of the 
sperm ducts is reduced even further, the cu- 
bic epithelium replaced by a tall cylindrical 
epithelium of infranucleate cells with a pale 
cytoplasm (Fig. 8), and the circular muscle 


VOLUME 106, NUMBER 4 


layer disappearing, with the exception of a 
few isolated fibers. On the basis of purely 
topographical considerations these modi- 
fied intrapenial ducts would conventionally 
be called “‘sperm vesicles’; however, they 
have a very reduced lumen and upon unit- 
ing they give rise to the ejaculatory duct 
without undergoing any further modifica- 
tion in the histological structure. For these 
reasons we consider it more appropriate to 
denominate them ‘“‘anterior branches of the 
ejaculatory duct.” 

The union of these two narrow ducts oc- 
curs at a point located somewhere between 
the base and two thirds of the length of the 
papilla (Fig. 9). The ejaculatory duct gen- 
erally runs through the center of the papilla 
and its opening is terminal. 

The penis bulb is hemispherical and short. 
The penis papilla is very long and in almost 
all the specimens examined reaches the pos- 
terior wall of the common atrium or even 
the gonopore (Figs. 3, 5), making it difficult 
to differentiate the male atrium from the 
common one. The epithelium of the papilla 
is cubic and shows intraepithelial nuclei; it 
is underlain firstly by a circular layer of 
muscles and then by a much thinner layer 
of longitudinal ones. 

Basophilous glands extend from the pe- 
riphery of the bulb to the ejaculatory duct 
into which they open, their degree of de- 
velopment varying greatly from one indi- 
vidual to another. 

The epithelium of the atria has normal 
nuclei and varies from cubic to cylindrical. 
The atrial walls have a subepidermal cir- 
cular muscle layer that is more developed 
than the longitudinal one. In five of the nine 
sagittally sectioned specimens, a fold with 
circular muscle fibers was observed on the 
ventral wall of the male atrium (Fig. 5). 
There can be no ring-shaped diaphragm 
since a corresponding fold has never been 
observed on the dorsal wall of the atrium. 

The common atrium is surrounded by a 
ring of cement glands that open into it. They 
are very strongly eosinophilous, their secre- 


639 


tion is granular, and they extend a long way 
toward the posterior part of the body. 

In only one specimen was a posterior 
pocket of the common atrium observed 
similar to that described for G. tigrina and 
G. anceps (Ball 1971, Cazzaniga & Curino 
1987). In the rest of the material there was 
no evidence of evaginations in the posterior 
wall of the atrium. 

Etymology. —The word canai means 
“‘friend”’ in Mapuche language and was cho- 
sen as the name for the new species in grat- 
itude for the friendship accorded by Dr. Pa- 
tricio Sanchez of the Universidad Catolica 
de Santiago, who guided us in the search for 
the material. 


Discussion 


The species described here can clearly be 
assigned to the Girardia genus by virtue of 
its triangular head, the presence of two mus- 
cle layers in the bursal canal with the sub- 
epidermal one circular and with no ectal 
reinforcement, the absence of a diaphragm 
in the ejaculatory duct, and the numerous 
ventral testes which extend along the length 
of the body. 

The pigmentation of the outer pharyngeal 
wall is one of the autapomorphies proposed 
for defining Girardia (de Vries & Sluys 1991). 
This character has been described for many 
species, though it is quite variable intraspe- 
cifically, as demonstrated in G. rincona, G. 
festae or G. tigrina (du Bois Reymond-Mar- 
cus 1953; Marcus 1954, 1960; Kawakatsu 
et al. 1984; Ribas et al. 1989), and in the 
case of G. anceps the pharynx is fully de- 
pigmented (Cazzaniga & Curino 1987, Ka- 
wakatsu & Rovasio 1992). It is regrettable 
that no conclusive information on this point 
can be included for G. canai. 

De Vries & Sluys (1991) presumed a close 
affinity among Girardia Ball, 1974, Cura 
Strand, 1942 and Schmidtea Ball, 1974. The 
relationship between these three dugesiid 
genera rests on a single synapomorphy, 1.e., 
the presence of an angled bursal canal, in- 
stead of the plesiomorphic smoothly curved 


640 


bursal canal. Ball (1980) suggested that it is 
possible to distinguish two groups within 
the Girardia genus, each encompassing 
roughly half of the known species. He rests 
his argument heavily on the curvature, ei- 
ther smooth or angled, of the bursal canal 
at its opening into the genital atrium. It ap- 
pears that the course of the bursal canal, 
either smoothly curved or angled, is asso- 
ciated with the dorsal and ventral testes, 
respectively (Ball 1980). The unity of the 
genus and its affinities should thus be re- 
considered in future analyses of additional 
characters. 

With the exception of Beauchamp (1939), 
authors have accepted that the ventral or 
dorsal position of the testes is a useful tax- 
onomical character (Ball 1971). De Vries & 
Sluys (1991) stated that the male gonads are 
unsuitable for inferring phylogenetic rela- 
tionships between genera of Dugesiidae. Ball 
(1974) expressed a similar viewpoint, as- 
serting however that the position of the tes- 
tes may be useful for delimiting species 
groups. 

The new species belongs to the group pre- 
vailing in southern South America that has 
ventral testes and an angled bursal canal. 
For this reason the species with dorsal testes 
and for the most part smoothly curved bur- 
sal canals (Ball 1980; Kawakatsu et al. 1983, 
1992) will not be included in the discussion. 
Within this latter group is Girardia chilla 
(Marcus), known in central and southern 
Chile (Marcus 1954, Hyman 1959, Kawa- 
katsu et al. 1984), and which has an enor- 
mous cavity in the penial bulb making it 
immediately distinguishable from G. canai. 

Within the group of species with ventral 
testes, the conformation of the intrapenial 
ducts in the new species establishes a novel 
morphological model. In some species of 
Girardia such as G. rincona and G. longistri- 
ata the sperm ducts join almost immedi- 
ately beyond the separate entrance to the 
penial bulb, where they form the ejaculatory 
duct. In other species the normal configu- 
ration is the formation of a bifid, more or 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


less broad seminal vesicle, as in G. anceps, 
G. sanchezi, G. dorotocephala, G. tigrina, or 
the occurrence of a unique intrabulbar ves- 
icle from which the ejaculatory duct arises, 
as in G. festae, G. nonatoi and G. arndti. 
The new species is intermediate between the 
previously known forms without seminal 
vesicles and those in which the ejaculatory 
duct is short and limited to the penis papilla 
due to the formation of the intrapenial ves- 
icle: 

Girardia canai has no intrabulbar en- 
largements, but the union of the intrapenial 
ducts nevertheless occurs in the penis pa- 
pilla. This difference is an important argu- 
ment in favor of excluding the following 
species from the discussion: G. dimorpha 
(Bohmig) and G. sanchezi (Hyman), both 
from Chile, G. anceps (Kenk), G. arndti 
(Marcus), G. aurita (Kennell), G. doroto- 
cephala (Woodworth), G. festae (Borelli), G. 
guatemalensis (Mitchell & Kawakatsu), G. 
nonatoi (Marcus). 

Though G. tigrina (Girard) should also be 
excluded on the very same grounds, it is 
considered pertinent to include it in the dis- 
cussion since it is a very variable and wide- 
spread species that serves a useful purpose 
in placing the description of the new species 
in perspective. 

The diagnostic features of Girardia tigri- 
na have been summarized by Kenk (1972). 
Most morphological descriptions referring 
to material from various parts of the world 
where the species is either autochthonous 
or has been introduced conform to these 
characters (for example, Ball 1971; Kawa- 
katsu & Mitchell 1981; Ribas et al. 1989). 
Morphological variations mostly affect ex- 
ternal pigmentation patterns, pharynx pig- 
mentation, the occasional expansion of the 
intrabulbar sperm vesicles—uniting in a 
single cavity, or their contraction, appearing 
as tubular seminal vesicles, the position of 
the epithelial cell nuclei of the copulatory 
system, and some asymmetry in the course 
of the sperm ducts. Rather more stable char- 
acters are the short, conical penis papilla, 


VOLUME 106, NUMBER 4 


the sharp angle of the bursal canal at its 
opening into the genital atrium and the pres- 
ence of a well-developed blind diverticulum 
on the posterior wall of the common atrium 
(Ball 1971, Ribas et al. 1989). 

Girardia tigrina is a species found in South 
America, and the descriptions given by 
Marcus (1946) and Kawakatsu et al. (1992) 
of specimens found in Brazil and Uruguay, 
respectively, agree with the characters al- 
ready mentioned for the species. Other South 
American material with diverse character- 
istics also has been identified as G. tigrina 
but no explicit valuation of the described 
characters has been carried out. 

For example, the musculature that sur- 
rounds the bursal canal provides a character 
of phylogenetic importance (Ball 1974, Sluys 
& de Vries 1988, de Vries & Sluys 1991). 
Girardia has the plesiomorphic condition 
of only two muscle layers, the subepidermal 
one of which is made up of circular muscle, 
as shown in Fig. 7. The specimens from 
southern Brazil (Kawakatsu et al. 1981, 
1982, 1983, 1986) and Uruguay (Kawaka- 
tsu & Ponce de Leon 1990), however, are 
described as having a thin subepidermal 
layer of longitudinal muscle in the bursal 
canal and a variable number of muscle lay- 
ers exhibiting diverse degrees of develop- 
ment. 

Furthermore, the bursal canal of these 
south Brazilian planarians does not always 
form a sharp angle, some ectal reinforce- 
ments appear, the caudal diverticulum of 
the atrium wall is not always developed, the 
penis papilla is not always of a rounded con- 
ical form, and sometimes it is atypically long. 
All such divergences from the diagnostic 
characters of G. tigrina emphasize the need 
for a reassessment, since they would appear 
to separate some of the Brazilian planarians 
from tigrina or even from Girardia. 

We consider that G. candi, new species, 
has the greatest morphological affinity with 
the following usually accepted species of 
Girardia, all of which have ventral testes, 
angled bursal canal and no seminal vesicles: 


641 


G. rincona (Marcus), G. longistriata (Fihr- 
mann), and G. microbursalis (Hyman). 
Girardia rincona was originally described 
by Marcus (1954) for central Chile and re- 
described by Kawakatsu et al. (1984) for 
Peru. This is the only species for which an 
epithelium of the ejaculatory duct similar 
to that of G. canai has been described, al- 
though it differs significantly in a number 
of other characteristics. The bursa copu- 
latrix of G. rincona is even smaller than in 
the least developed cases of G. canai. Mar- 
cus (1954) and Kawakatsu et al. (1984) 
mention the presence of preovaric testicular 
follicles in G. rincona. In G. canai however, 
a testicular follicle was seen only once, ex- 
ceptionally, in this position, on one side of 
a specimen. Both the G. rincona material 
from Chile and that from Peru show the 
extrapenial seminal vesicles running up- 
ward to the anterior dorsal side of the penis 
bulb, while in G. canai this condition does 
not appear, not even in fully developed in- 
dividuals. In the new species the occurrence 
of sperm ducts that follow different, fre- 
quently asymmetric courses before entering 
the penial bulb is typical. In G. rincona the 
sperm ducts either unite in the penial bulb 
without giving rise to a bifid vesicle, as de- 
scribed by Marcus (1954), or give rise to a 
small bulbar cavity or seminal vesicle, as 
described by Kawakatsu et al. (1984), there- 
after continuing in the form ofa unique and 
narrow ejaculatory duct whose histology dif- 
fers from that of the sperm ducts. In G. 
canai, on the other hand, the sperm ducts 
run along about half the length of the penial 
bulb and then undergo an abrupt histolog- 
ical transformation, acquiring the same 
characteristics as the ejaculatory duct, be- 
fore uniting somewhere between the base of 
the papilla and a third of the way along to 
the opening. Kawakatsu et al. (1984) said 
that the penis papilla of the samples from 
Peru are accompanied by many eosinoph- 
ilous glands that do not appear in G. canai. 
Finally, the penis papilla in G. rincona is 
short and conical whereas that in the new 


642 


species is significantly longer, probably the 
longest ever described for this group of spe- 
cies. 

Girardia longistriata, from Colombia, has 
been briefly redescribed by Ball (1980). The 
greatest similarity is seen in the type of in- 
sunk epithelium that covers the ejaculatory 
duct, which is narrow and opens terminally 
but does not run centrally through the pa- 
pilla. There are however several differences 
in the Colombian species: the dorsum has 
a different color pattern, the penis papilla is 
conical and very short, and the sperm ducts 
unite within the bulb rather than in the pe- 
nis papilla as they do in G. canai. The initial 
course of the ovovitelline ducts is also dif- 
ferent. 

Girardia microbursalis from Connecticut, 
U.S.A., is known only through its original 
description (Hyman 1931), which lacks his- 
tological details and thus precludes an ac- 
curate comparison. From what can be seen 
it would appear to differ in the size of the 
bursa copulatrix (to which characteristic it 
Owes its specific name) and in the site where 
the sperm ducts or “‘anterior branches of the 
ejaculatory duct’? unite. Without a histo- 
logical description there is no reliable way 
of telling whether or not a transformation 
of the type described in the new Chilean 
species occurs. Hyman (1939) cited new lo- 
calities without giving any additional mor- 
phological data. Kenk (1974, 1989) sug- 
gested that it could be a synonym of G. 
tigrina, an opinion that was not sustained 
in the outcome of the ensuing discussion. 


A Note on the Nomenclature of 
Girardia festae 


A South American freshwater planarian, 
now under Girardia Ball, 1974, was de- 
scribed by Borelli (1898) as Planaria festae, 
based on the name of Dr. Enrico Festa. This 
original spelling is correct under article 31 
a(i) of the International Code of Zoological 
Nomenclature (International Trust for Zoo- 
logical Nomenclature 1985; example: podae 
from Nicolaus Poda). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Marcus (1946) suggested that “‘melhor 
seria festai” (festai would be better), and this 
comment was considered nomenclaturally 
valid by Kenk (1974). Du Bois Reymond- 
Marcus (1953) used the emendation for the 
explicit reason that festae “‘constitutes a lap- 
sus, as it was chosen in honor of Dr. Enrico 
Festa.’’ They thus changed the spelling to 
Dugesia festai on the grounds that Dr. Festa 
was male. This invalid emendation subse- 
quently became common usage. The origi- 
nal orthography was not in fact erroneous 
but the correct formation of the first de- 
clension genitive in Latin. 

Certainly, festai as an original spelling 
would also have been admissible (art. 31 
a(ii); example podai), but there is no valid 
reason for accepting an emendation to Bo- 
relli’s spelling of this species. Girardia festai 
(Marcus, 1946), based on an unjustified 
emendation, became a junior objective syn- 
onym of G. festae (Borelli, 1898). The fol- 
lowing references are, therefore, correct sub- 
sequent spellings: 

Euplanaria festae: Kenk (1930, explicit 
new generic combination), Beauchamp 
(1939). Dugesia festae: Hyman (1939, ex- 
plicit new generic combination); whereas 
those that follow are incorrect (art. 33 c): 

Dugesia festai: du Bois Reymond-Marcus 
(1953), Marcus (1960), Ball (1969, 1980), 
Mitchell & Kawakatsu (1972), Kawakatsu 
& Mitchell (1984a, 1984b), Kawakatsu et 
al. (1984), Kawakatsu & Rovasio (1992). 
Dugesia (Girardia) festai: Ball (1974, jus- 
tified subgeneric assignation), Kenk (1989). 


Acknowledgments 


Contribution #29 of the Laboratorio de 
Ecologia Acuatica, Departamento de Biolo- 
gia, Universidad Nacional del Sur. NJC is 
a Researcher of the Comision de Investi- 
gaciones Cientificas de la Provincia de Bue- 
nos Aires, Argentina. We are greatly in- 
debted to A. Boland for bringing us part of 
the bibliographical material from U.S.A., to 
M. Kawakatsu, for being so kind as to send 
us a large number of his papers (to which 


VOLUME 106, NUMBER 4 


for a variety of reasons we would otherwise 
have had no access) and in offering his help, 
to P. Hermann for providing the facilities 
of her laboratory, and to R. Delhey for his 
help in reading the German literature. This 
work was supported by a P.I.D. grant from 
the Consejo Nacional de Investigaciones 
Cientificas y Técnicas (CONICET), Argen- 
tina. 


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——,, J. Hauser, S. M. G. Friedrich, & O. de Souza 
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. 1960. Turbellaria from Curacao. — Studies on 

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Mitchell, R. W., & M. Kawakatsu. 1972. Freshwater 
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626. 

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Departamento de Biologia, Universidad 
Nacional del Sur, Peru 670, 8000 Bahia 
Blanca, Argentina. 


PROC. BIOL. SOC. WASH. 
106(4), 1993, pp. 645-660 


OCTOPUS ORNATUS GOULD, 1852 
(CEPHALOPODA: OCTOPODIDAE) IN AUSTRALIAN 
WATERS: MORPHOLOGY, DISTRIBUTION, AND 
LIFE HISTORY 


Mark D. Norman 


Abstract. — The morphology, distribution, and life history of the ““white-striped 
octopus,’ Octopus ornatus Gould, 1852, from Australian waters are reported. 
Information gathered on habitat preferences, activity patterns, foraging behav- 
ior and diet also are presented. Australian representatives of this species are 
described and compared with the neotype from Hawaiian waters. Prior reports 
of O. ornatus from Australia refer to a related species, Octopus aspilosomatis 
Norman, 1993a. Distributional records from Asia, the Indian Ocean and the 
South Pacific Ocean are presented. The known distribution of O. ornatus ex- 
tends from Easter Island and the Hawaiian islands, west to eastern Africa. 
Delineation of O. ornatus from related taxa is discussed. 


The “‘white-striped octopus,” Octopus or- 
natus Gould, 1852, is a large, nocturnally- 
active octopus found primarily in associa- 
tion with coral reefs throughout the tropical 
Indian and West Pacific Oceans. This spe- 
cies originally was described by Gould 
(1852) on the basis of specimens from the 
Hawaiian Islands collected on the Wilkes 
U.S. Exploring Expedition of 1838-1842. 
Original type material has never been traced 
and is presumed lost (Voss 1981). Voss 
(1981) redescribed the species and desig- 
nated a neotype from Oahu Island, Hawaii 
(93.7 mm ML male, USNM 730020). Voss 
also described additional specimens from 
Hawaii, the Marshall Islands and Kenya, 
and synonymized Callistoctopus arakawai 
Taki, 1964 from southern Japanese waters. 

Octopus ornatus was reported from Liz- 
ard Island at the northern end of the Great 
Barrier Reef, Australia by Roper & Hoch- 
berg (1987, 1988). Specimens and photo- 
graphs of the taxon referred to in these works 
have been re-examined and proved to be- 
long to a distinct species, Octopus aspilo- 
somatis Norman, 1993a. 


This paper presents the first record of the 
true O. ornatus from Australian waters. A 
detailed description of the Australian spec- 
imens and a comparison with the neotype 
are provided. Additional counts and indices 
from the neotype are provided beyond those 
included in Voss’s (1981) treatment. This 
species is reported for the first time from a 
number of new localities in the Pacific and 
Indian Oceans. Observations of live O. or- 
natus from Australia provide some infor- 
mation on habitat preferences, foraging be- 
havior, diet and activity patterns. 


Methods 


Nine live O. ornatus were encountered in 
the field in two visits to One Tree Island, 
Capricorn Bunker Group, southern Great 
Barrier Reef in October 1989 and Septem- 
ber 1990. Individual animals were observed 
and photographed in situ. Seven animals 
were collected and returned to aquaria where 
they were observed, described and photo- 
graphed. The seven specimens were killed 
in fresh water, fixed in 10% formalin for a 


646 


minimum of two weeks and preserved in 
70% ethanol, according to the techniques of 
Roper & Sweeney (1983). This material is 
now housed in the Museum of Victoria, 
Melbourne, Australia. Voss’s neotype of O. 
ornatus was examined in the National Mu- 
seum of Natural History, Smithsonian In- 
stitution, Washington, D.C. A total of 39 
additional preserved specimens were ex- 
amined in the collections of the Museum of 
Victoria, Melbourne, Australia (NMV), 
Australian Museum, Sydney, Australia 
(AMS), Santa Barbara Museum of Natural 
History (SBMNH), National Museum of 
Natural History, Smithsonian Institution, 
Washington, D.C. (USNM), California 
Academy of Sciences, San Francisco, Cali- 
fornia (CASIZ), British Museum (Natural 
History), London, England (BMNH) and 
Muséum National d’Histoire Naturelle, 
Paris, France (MNHN). 

Field collection at Lizard Island, northern 
Great Barrier Reef, in November 1989 en- 
abled observation and collection of O. as- 
pilosomatis Norman, 1993a, the species re- 
ferred to as O. ornatus by Roper & Hochberg 
(1987, 1988). Specimens from Lizard Island 
treated in these works were examined at the 
Smithsonian Institution, Washington, D.C. 
in June 1990. 

In the description and tables, measure- 
ments and indices follow Roper & Voss 
(1983:56) and Toll (1988). The following 
additional or modified indices and symbols 
also are employed: 

Arm Mantle Index (AMI): arm length as 
% of ML; Arm Width Index (AWI): arm 
width at widest point on stoutest arm, as % 
of ML; Free Funnel Index (FFI): length of 
free funnel portion as % of funnel length; 
Funnel Length Index (FLI): funnel length as 
% of ML; Funnel Organ Index (FOI): length 
of outer limb of funnel organ as % of medial 
limb length; Funnel Organ Length Index 
(FOLI): length of medial limb as % of funnel 
length; Gill Count (GC): number of gill la- 
mellae per demibranch not including the 
terminal lamella; Head Mantle Index (HMI): 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


head width as % of mantle width; Hecto- 
cotylized Arm Mantle Index (HAMI): length 
of hectocotylized arm as % of ML; Sper- 
matophore number (SpN): number of sper- 
matophores in Needham’s Sac; Stage of 
Maturity (StM): Immature (Imm: sex in- 
determinate or reproductive organs min- 
ute), Submature (S: reproductive organs dis- 
tinct but poorly developed) and Mature (M: 
developed spermatophores or eggs distinct); 
Sucker Count (SC): number of suckers on 
arm with highest sucker count. 


Family Octopodidae d’Orbigny, 1839 
Subfamily Octopodinae d’Orbigny, 1839 
Octopus ornatus Gould, 1852 
Figs. 1-5 


Octopus ornatus Gould, 1852:476, fig. 590, 
590a.— Gould, 1862:232.—Tryon, 1879: 
112, pl. 30, figs. 29-30.—Hoyle, 1886a: 
11; Hoyle, 1886b:220.—Robson, 1929: 
108.—Boone, 1938:357, pl. 151.—Adam, 
1941:9, 14.—Van Heukelem, 1966.— 
Voss, 1981:525, figs. 1-3, tables 1, 2.— 
Houck, 1982:152.—Nesis, 1982:74, plate 
(in 1987 translation).— Young, Harman 
& Hochberg, 1989:152, figs. 3, 5.— 
O’Shea, 1990:41, fig. 2. 

Polypus ornatus.—Berry, 1909:418; Berry, 
1914:294, pl. XLVI, figs. 1-2, textfig. 14. 

Callistoctopus arakawai Taki, 1964:292, pls. 
2-3, textfigs. 34-41.—Taki, 1965:324, 
textfig.; Taki, 1981:253.—Okutani, Ta- 
gawa & Horikawa, 1987:168, textfigs. 
66A, B. 

Octopus arakawai.—Dong, 1979:72, pl. 1, 
he. 2: 19ST 686, he: 123: 


Holotype. —Not extant. 

Holotype locality. —Sandwich Islands 
(=Hawaiian Islands). 

Neotype.—93.7 mm ML male (USNM 
730020), designated by Voss (1981). 

Neotype locality. — Pacific Ocean, Hawaii, 
Oahu Island, Black Point. 

Material examined. —See appendix. 

Diagnosis. —Medium to large nocturnal- 
ly-active species with distinctive color pat- 


VOLUME 106, NUMBER 4 


tern of buff or white longitudinal stripes on 
dorsal and lateral mantle, in distinct con- 
trast to a background of red brown or dark 
maroon red. Regular paired white spots on 
dorsal arm crown and along aboral surfaces 
of all arms. Interbrachial, lateral face of all 
dorsal suckers with large buff or white spot. 
Arms elongate (AMI 544.6-838.6), unequal 
in length, dorsal pair longest, grading to 
ventral pair shortest (AF 1.2.3.4). Webs 
shallow (WDI 5.3-11.3), dorsal web deepest 
(WF typically A.B.C.D.E). More than 300 
suckers per arm in submature and mature 
animals. Hectocotylized arm with 150-180 
suckers in submature to mature individuals. 
Large conical ligula in mature males (LLI 
4.3-5.7). Eggs small, capsule up to 3.5 mm 
long. Gills with 13-14 lamellae per demi- 
branch. 

Description. —Counts and indices for the 
neotype and Australian specimens are pre- 
sented in Tables | and 2. In the following 
text, the values for both the neotype and 
Australian material are given in brackets. 
The first number refers to the neotype, fol- 
lowed by the range and mean (underlined) 
in the Australian material. Voss (1981) re- 
ported that O. ornatus shows considerable 
variation in elongation of the mantle on 
preservation. A similar range was observed 
in the Australian material. 

Medium to large species with elongate 
arms (Figs. la, 4a); ML to at least 100 mm 
for males and 130 mm for females; TL to 
at least 1200 mm; weight to at least 1000 
g. Mantle shape variable, from ovoid to 
greatly elongated (Voss 1981, fig. lb-d: MWI 
76.6; 41.5-65.2-76.3). Mantle walls thick, 
muscular. Stylets present, poorly devel- 
oped. Pallial aperture moderately wide, ap- 
proximately half mantle width. Funnel 
broad-based and muscular (FLI 49.9; 37.0— 
47.9-59.7), free portion variable in length, 
approximately half funnel length (FFI 44.1; 
31.1-49.5-69.4). Funnel organ well devel- 
oped with broad limbs (Fig. 1b), outer limbs 
slightly shorter than median ones (FOI 85.6; 
76.4—82.1-—86.3). Funnel organ approxi- 


647 


mately 70% of funnel length (FOLI 80.3; 
62.1-70.6—77.5). 

Head slightly narrower than mantle (HWI 
52.7; 27.3-46.8-56.4, gravid 2: 37.3; HMI 
68.8; 65.2—76.1-87.6), neck distinct, slight- 
ly narrower than head. Eyes medium-sized, 
slightly pronounced. 

Arms long, typically 6-8 times mantle 
length (AMI 685.2; 544.6-688.1-838.6), 
moderately robust (AWI 23.9; 16.4-18.5- 
19.9), sub-cylindrical along length. Arm loss 
and regeneration evident at different levels 
along arms, on at least some arms in most 
specimens. Arms decrease in length and di- 
ameter from dorsal to ventral pairs, dorsal 
arms longest and most robust (AF 1.2.3.4). 
Suckers moderately large (SDI 14.0; 7.3- 
11.3-14.4), larger on dorsal arms; distinctly 
enlarged suckers absent in both sexes. Suck- 
ers moderately cylindrical with thick, mus- 
cular rims. Sucker rims finely scalloped, 
more pronounced in smaller distal suckers. 
More than 300 suckers on each intact arm 
in submature and mature specimens, fe- 
males with slightly higher sucker counts (SC 
neotype R2: 378; R4: 400; SC 6: 324—342- 
366,47 =3522SC 2: 3765382) 7 — 2). Webs 
shallow (WDI 7.8; 5.3-8.7—11.3), depths 
subequal, decreasing slightly from dorsal to 
ventral webs (WF typically A.B.C.D.E or A 
= B = C.D.E). Web margins extend as nar- 
row retractile membranes on dorso-lateral 
and ventro-lateral edges for at least 70% of 
length of all arms. 

Third right arm of males hectocotylized, 
short (HAMI 307.4; 253.9-310.2—353.6), 
approximately 60% length of opposite arm 
(OAI 84.7; 54.8-62.6-68.5). Ligula mod- 
erate sized in mature males (LLI 5.7; range 
for larger submature and mature Australian 
specimens: 4.3-4.9-5.6, n = 4), robust and 
cylindrical, tapering to blunt point (Fig. Ic). 
Ligula groove deep and usually closed. Cal- 
amus small but distinct (CLI 18.4; 13.6- 
15.3-18.4, n = 4). Spermatophore groove 
well developed, wide and thin with fine 
transverse ridges. Spermatophore guide dis- 
tinct, bordered by flattened papillae or digits 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


648 


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VOLUME 106, NUMBER 4 649 


Table 2.— Octopus ornatus Gould, 1852: Counts and indices for females from Australian waters. (S = sub- 
mature; M = mature; D = damaged; — = not recorded; ID = indistinct; * = mature ovarian eggs only; G = 


distended gravid female). 


Museum: NMV 
Reg. no.: F57919 
ML 173 
StM S 
aL 512 
MWI 65.8 
HWI 48.0 
HMI 129 
ALI:1 544.6 
2 469.6 
3 410.1 
4 388.1 
AWI 16.4 
SDIn oa 
SC 382 
WDI 8.9 
WF A.B. = C.D.E 
GC 14 
ELI* 
EWI* 
EN* 
FLI 41.7 
PEL 45.6 
FOI 83.5 
FOLI 62.1 


of skin. Approximately 160 suckers on hec- 
tocotylized arm (HASC 163; 152-164-172). 

Gills with 13-14 lamellae (holotype: 14) 
on both inner and outer demibranchs, plus 
terminal lamella. 

Digestive tract illustrated in Fig. 2a. Pos- 
terior salivary glands large (up to twice length 
of buccal mass), attached to lateral edges of 
crop diverticulum. Crop diverticulum well 
developed. Stomach bipartite. Caecum 
coiled to form 1.5 whorls, distinctly striat- 
ed. Digestive gland elongate, approximately 
conical in shape and bilobed at posterior 
end. Intestine long, moderately narrow with 
a reflexed bend approximately one third 
from proximal end. Walls of rectum mus- 
cular in distal quarter of intestinal tract. Ink 
sac well developed, partially embedded in 
ventral surface of digestive gland. Anal flaps 
present. 

Beaks illustrated in Fig. 2b—d. Upper beak 
with a short hooked rostrum and narrow 


NMV AMS 
F57922 C169234 
$3.5 13220 
S M 
738 1210 
68.7 73.2 
55.6 27:3 
80.9 3 1-3G 
756.9 838.6 
638.3 582.6 
540.1 35-0 
479.0 5)2:3 
18.1 — 
PAS) 11.5 
376 — 
8.5 8.9 
B.€ = DE C.B.D.E.A 
13 13 
7254 | 
0.5 
~35,000 
51.3 221 
46.2 48.4 
86.3 ID 
1333 ID 


hood (Fig. 2b). Lower beak with moderately 
sharp rostrum, narrow hood, widely spread 
wings and flared lateral walls separated in 
posterior half (Fig. 2c, d). Radula with seven 
teeth and two marginal plates in each trans- 
verse row (Fig. 4f, 1). Rhachidian tooth with 
2-3 lateral cusps, typically 3, on each side 
of large medial cone. Lateral cusps in sym- 
metrical seriation, migrating from lateral to 
medial position over 9-10 transverse rows. 

Male genitalia illustrated in Fig. 3. Penis 
in mature males of moderate size (PLI 9.4; 
in larger submature and mature Australian 
specimens 10.0-13.9-16.9, n = 5), genital 
aperture subterminal. Holotype contained 
damaged spermatophores. Spermatophores 
short (SpLI neotype: D; Australian mate- 
rial: 38.4—49.3, n = 4) and narrow (SpWI 
neotype: D; Australian material: 2.0—2.5, n 
= 4), apparently produced in low numbers 
(4 in Needham’s Sac of holotype; 8 and 9 
in two other males, NMV F57918, NMV 


650 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


a 
1 
' 
' 
i | 
i 
4 
" 
N 
il 
1 il 
il 
" 
il 
il 
i 11 
7 i) 
a i 
4 a 
i i 
: Fl 
: i : 
i LY - 
i H 
“ ae _ 
5 4) 
: #1 F 
R yf 4 b 
A giagd | 
i} . 2 ' 
PR BY Uk st 
5 a: 4 i 
fe a i { 4 U 
os i) Ek 3 7 cs 
e te Bad ff a EE ie 
S 25 Pat fe >. iki 
s 8 i oe 
elle : fi. 2 
-_ rE 
a 1 
Se Es 
“Fe ce 
5 by 
‘gg 
ZY 


a 


Fig. 1. Octopus ornatus Gould, 1852. a, Dorsal view of NMV F57920 (45.9 mm ML 3), Abbreviations DWS 


= dorsal white spots, HA = hectocotylized arm; b, Funnel organ of same specimen; c, Hectocotylus of NMV 
F57918 (104.2 mm ML 8). 


VOLUME 106, NUMBER 4 


651 


d 


Fig. 2. Octopus ornatus Gould, 1852. a, Digestive tract from NMV F57922 (83.5 mm ML 9), Abbreviations 
ASG = anterior salivary glands, BM = buccal mass, C = caecum, CD = crop diverticulum, CR = crop, DG = 
digestive gland, I = intestine, IS = ink sac, O = oesophagus, PSG = posterior salivary glands, R = rectum, S = 
stomach; b—d Beaks from NMV F57918 (104.2 mm ML 8), b, Upper beak, lateral view; c, Lower beak, lateral 


view; d, Lower beak, ventral view. 


F57924). Oral cap simple with long cap 
thread. Details of ejaculatory organ, cement 
body and sperm reservoir unclear in ma- 
terial examined due to poor preservation. 
Sperm reservoir approximately half total 
length (SpRI 44.0, 45.1). 

Only one gravid female encountered in 
specimens examined (132.0 mm ML; AMS 
169234). Mature ovarian eggs small, cap- 
sule length to 3.5 mm long (EgLI: 2.7; EgWI: 
0.5); produced in large numbers (approxi- 
mately 35,000 in ovary of above specimen). 
Oviducal glands partitioned into approxi- 
mately 20 radiating chambers in gravid fe- 
male and one submature female (1 2: 83.5 
mm ML, NMV 57922). 


Body coloration relatively fixed. Base col- 
or generally orange or red brown with dis- 
tinctive series of buff or white stripes and 
spots on arms, dorsal body and bases of 
dorsal suckers (Figs. la, 4a—e, g, h). Pair of 
small dorsal white spots are visible on dor- 
sal mantle when longitudinal stripes are 
suppressed (DWS in Fig. la; see also Fig. 
4d, g). In alarm display, base color becomes 
deep maroon red with pure white stripes 
and spots (Fig. 4a, b, g). During daylight 
hours the base color in “inactive”’ animals 
light pink or gray, iridocytes evident as ir- 
idescent green hue. 

Skin covered with small fixed patches 
separated by fine grooves to form an irreg- 


652 


SG2 


Fig. 3. Octopus ornatus Gould, 1852. Male repro- 
ductive tract from NMV F57918 (104.2 mm ML 8). 
Abbreviations A = appendix, D = diverticulum, MG 
= mucilaginous gland, NS = Needham’s sac, P = penis, 
SG1 = spermatophoric gland 1, SG2 = spermatophoric 
gland 2, T = testis, VD = vas deferens. 


ular reticulation over all dorsal surfaces (Fig. 
4d). Two moderate-sized papillae present 
over each eye (Fig. 4d). In alarm displays, 
longitudinal truncate flaps or ridges raised 
in centers of white stripes and spots (Fig. 
4c-e). 

Remarks. —The distinctive color pattern 
and sculpture of this species are typically 
visible in preserved specimens (Fig. 4h), en- 
abling easy identification. 

Sexual dimorphism is not marked in O. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ornatus. Based on the few mature specimens 
available, females appear to reach a larger 
size than males. Additional material should 
be examined to confirm these trends. 

Distribution.—In Australian waters, O. 
ornatus 1s reported from One Tree Island at 
the southern end of the Great Barrier Reef 
and on the New South Wales coast south to 
Sydney. 

The known distribution of this species is 
considerably expanded here. Octopus or- 
natus is reported for the first time from: 1) 
the South Pacific: Easter Island, Society Is- 
lands (Tahiti and Raiatea), Cook Islands 
(Raratonga) and Fiji Islands (Rotuma and 
Vatao Island); 2) the West Pacific: Philip- 
pines (Batan Island), New Caledonia (con- 
firmed from video footage of G. Boucher, 
MNHN); and 3) the Indian Ocean: Mada- 
gascar, Aldabra Atoll (West Island), Chagos 
Archipelago (Eagle Island), Reunion Island 
and the Seychelles. 

Octopus ornatus thus is widely distributed 
in tropical waters of the Indian and Western 
and Central Pacific oceans (Fig. 5). It is re- 
ported from Hawaii and Easter Island in the 
east, through the Pacific Islands to Asia and 
Australia, and into the Indian Ocean to East 
Africa. Taki (1964, 1965, 1981) reported 
this species from southern Japan under the 
name Callistoctopus arakawai. Okutani, 
Tagawa & Horikawa (1987) also used this 
name for O. ornatus from the Okinawa Is- 
lands, in tropical waters south of Japan. 
Nishimura (1992) contains an illustration 
of O. ornatus from the Bonin Islands, also 
in tropical waters south of Japan. 

O’Shea (1990) reported O. ornatus from 
New Zealand on the basis of a single female 
specimen. O’Shea illustrated the specimen 
(O’Shea 1990, fig. 2), which clearly is O. 
ornatus, providing the collection locality as 
“NZOI Station K23.”’ Additional locality 
details have since been obtained. This spec- 
imen was collected from the Tonga Islands 
(18°41.70'S, 173°57.40'’W) and, hence, does 
not constitute a record of this species from 
New Zealand waters. 


VOLUME 106, NUMBER 4 


Life history.—All nine O. ornatus en- 
countered in the field were found on One 
Tree Island at the southern end of the Great 
Barrier Reef. One Tree Island is a true coral 
island formed within a large shallow lagoon. 
All O. ornatus were found within the lagoon 
during night low tides between 1930 and 
0430 hr, active over coral rubble substra- 
tum in shallow water (0.2—2 m deep). Most 
specimens were foraging along the water’s 
edge adjacent to exposed intertidal reef flats. 
Several individuals were encountered in 
openings of large lairs. These lairs consisted 
of deep vertical holes excavated in coral 
rubble. One specimen retreated into its lair 
and blocked the entrance at several levels 
with large pieces of dead coral. No evidence 
of O. ornatus lairs could be found during 
daylight hours, the entrances sealed with 
rubble during the day. It is unknown wheth- 
er this species occupies permanent lairs or 
temporary/short-term refuges. 

Individuals foraged over coral rubble, ex- 
ploring burrows and holes with the dorsal 
arms, using tactile detection of prey. One 
specimen was caught carrying four alpheid 
shrimps on the proximal suckers of the lat- 
eral arms. A large captive individual of O. 
ornatus was observed to attack and com- 
mence devouring a small individual of an- 
other octopus species, Octopus alpheus Nor- 
man, 1993a. The intestine of another 
specimen (83.5 mm ML female, NMV 
F57922) contained the nearly intact upper 
and lower beak of an octopus, plus broken 
portions of other octopod beaks. No prey 
remains were found in middens around the 
entrance to lairs. 

Young et al. (1989) reported that the 
hatchlings of O. ornatus are planktonic. 
Nothing is known of courtship or brooding 
behavior, and insufficient mature speci- 
mens are available to determine whether 
any seasonality to spawning exists. 

Little information has previously been re- 
ported on the life history of O. ornatus. 
Houck (1982) reported that individuals of 
this species from Hawaii showed nocturnal 


653 


activity patterns, with maximum activity 
between 2100 and 0200 hr. Active O. or- 
natus were observed foraging over reef flats 
or on sand and gravel substrata during night 
low tides. Captive individuals were found 
to enter deep, sleep-like inactivity during 
daylight hours, burrowing under gravel or 
sand during these light periods. Houck 
(1982) also reported that O. ornatus readily 
accepted grapsid crabs and shrimp species. 


Discussion 


The majority of the counts and indices 
recorded from the neotype fell within or 
close to the range observed in the Australian 
material, often close to the mean. The only 
marked exception was the index of hecto- 
cotylized arm length over the length of the 
opposite arm (OAJ), which was much higher 
in the neotype than in the Australian ma- 
terial (OAI 84.7 versus 54.8-62.6-68.5 for 
the Australian material). The third left arm 
in the neotype was considerably shorter than 
in all other specimens examined, including 
Hawaiian material. The suckers on this are 
smaller in diameter than adjacent and op- 
posite arms suggesting that the arm has par- 
tially regenerated from earlier damage. An 
additional Hawaiian specimen (USNM 
214609) with intact arms had an OAI of 
58.1. In Australian specimens, there is no 
well-marked long notch in the cutting edge 
of the upper beak, as reported by Voss (1981) 
for Hawaiian material. 

Counts and indices were taken from ad- 
ditional specimens from Hawaii, Philip- 
pines, Fiji, Tahiti, Madagascar, Chagos Ar- 
chipelago, Cook Islands and Aldabra Atoll. 
All possessed typical body morphology of 
long unequal arms, short webs, 13-14 gill 
lamellae and the characteristic pattern of 
white stripes and truncate flat papillae. Males 
possessed 144-171 suckers on the hecto- 
cotylized arm. 

A number of cephalopod workers have 
recognized the distinct group of octopus 
species, often referred to as the Octopus 


654 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Octopus ornatus Gould, 1852. a, Photograph of live animal (NMV F57920: 45.9 mm ML 8); b-e, g 
Photographs of live animal (NMV F57918: 104.2 mm ML 8); b, Dorsal mantle and arm crown showing darkened 
alarm coloration, Photo R. Fenwick; c, Dorsal mantle and arm crown showing raised flaps in white bars, Photo 
R. Fenwick; d, Dorsal mantle showing patch and groove system and dorsal white spots (arrows); e, Lateral 
mantle and arm crown showing raised flaps in white bars, Photo R. Fenwick; f, Scanning EM photograph of 


VOLUME 106, NUMBER 4 


20°E 60°E 100°E 


cfg Oe ee 
LOSER eee 


140°E 
Fig. 5. Distribution of Octopus ornatus Gould, 1852. Symbols V = 


40°S 
180° 140°W 


type locality of O. ornatus, @ = type 


locality of Callistoctopus arakawai Taki, 1964, @ = material examined, A = published records. 


macropus group (Robson 1929, Adam 1941, 
Taki 1964, Voss 1981). These octopuses are 
characterized by elongate arms with dorsal 
arms always longest (arm formula typically 
1.2.3.4), high gill counts (10-14 per demi- 
branch), multicuspid radulae (rhachidian 
typically bears three lateral cusps on each 
side of a medial cone) and nocturnal activity 
patterns. Octopus ornatus exhibits all these 
characters and has been suggested to be a 
member of this group of species (Robson 
1929, Adam 1941, Voss 1981). 

Octopus ornatus often has been allied to 
the species from which this group receives 
its name, O. macropus Risso, 1826 (Gould 
1852, Berry 1914, Robson 1929, Voss 1981, 
Roper & Hochberg 1988). The description 
and distribution of O. macropus 3.s. cur- 
rently are being reviewed by Hochberg, 
Mangold and Norman, who consider this 
species to be restricted to temperate waters 
of the Mediterranean and the north west 
African coast. Considerable geographic and 
temperature boundaries separate the distri- 
butions of O. ornatus from that of O. macro- 
pus. The two species are easily distinguished 


on the basis of color patterns, gill lamellae 
counts (13-14 for O. ornatus vs. 11-12 for 
O. macropus) and hectocotylized arm suck- 
er counts (~160 for O. ornatus vs. ~110 
for O. macropus). 

The name, O. macropus Risso, 1826, reg- 
ularly has been applied to elongate Indo- 
West Pacific octopuses with dorsal arms 
longest and high gill lamellae counts (10- 
14 per demibranch) [Joubin 1894, 1898; 
Goodrich 1896; Hoyle 1904; Berry 1912, 
1914; Massy 1916; Wulker 1913, 1920; 
Odhner 1917; Robson 1926, 1929, 1932; 
Boone 1938; Adam 1939, 1942, 1946, 1954, 
1959, 1960, 1973; Rees & Stuckey 1954; 
Voss 1963; Nesis 1982 (plate in 1987 trans- 
lation); Roper, Sweeney, & Nauen 1984]. 
The restriction of the distribution of the true 
O. macropus to the Mediterranean and tem- 
perate Atlantic necessitates the need to crit- 
ically review the Indo-West Pacific mem- 
bers of the Octopus macropus group. It also 
makes available a number of species names 
from the Indo-West Pacific which previ- 
ously were synonymized with O. macropus 
(Robson 1929, Roper, Sweeney, & Nauen 


— 


radula from NMV F57918 (104.2 mm ML 4); g, Dorsal mantle showing alarm coloration and position of dorsal 
white spots (arrows); h, Dorsal mantle of preserved specimen showing characteristic color pattern (NMV F57918: 
104.2 mm ML 8); i, Scanning EM photograph of radula from NMV F57918 (104.2 mm ML 3) showing serial 
migration of lateral cusps on rhachidian tooth from lateral to medial position over 10-11 rows. 


656 


1984). The following nominal taxa show 
morphological and behavioral affinities with 
O. ornatus, yet are easily distinguished: 

Octopus lechenaultii d’Orbigny, 1826 (and 
its synonym O. cuvieri d’Orbigny, 1826) 
were both described from Pondicherry in 
southern India. The holotypes of both nom- 
inal species were examined in Paris (O. /e- 
chenaultii: 1 9: 61.8 mm ML, MNHN 4-12- 
972; O. cuvieri: 1 6: 42.9 mm ML, MNHN 
4-12-971). Octopus ornatus is distinct from 
this species with higher gill lamellae counts 
(13-14 for O. ornatus vs. 11-12 for O. leche- 
naultii) and more suckers on the hectoco- 
tylized arms (~ 160 for O. ornatus vs. 88 on 
the holotype of O. lechenaultii). 

Octopus luteus Sasaki, 1929 was de- 
scribed from two types lodged in the Hok- 
kaido Imperial University (110 mm ML 
male and 125 mm ML female), collected 
from the Pescadore Islands near Taiwan. 
Octopus ornatus 1s distinct from this species 
with longer arms (5.4—8.4 times ML in O. 
ornatus vs. 4—4.1 in O. luteus), more suckers 
on normal arms (>300 for O. ornatus vs. 
~200 for O. luteus) and on the hectocotyl- 
ized arm (~160 for O. ornatus vs. 88 on 
holotype of O. luteus), larger hectocotylus 
in mature males (4.3-5.7 for O. ornatus vs. 
3.7 for O. luteus) and more gill lamellae (13- 
14 for O. ornatus vs. 12 for O. luteus). 

Octopus nanhaiensis Dong, 1976 is known 
from a single male from Quangdong Prov- 
ince, China. It appears distinct from O. or- 
natus in possessing fewer gill lamellae (10 
vs. 13-14). Additional material and access 
to the type material is required to clarify the 
status of this little-known species. 

Octopus rapanui Voss, 1979 from Easter 
Island shows some similarities to O. orna- 
tus. Specimens of both species collected from 
Easter Island were examined in the MNHN 
collection. These species can be distin- 
guished easily by a number of characters. 
Octopus ornatus is distinct from this species 
having shallower webs [WDI 5.3-11.3 for 
O. ornatus vs. 18-23 for O. rapanui (Voss 
1979)], more suckers on the hectocotylized 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


arm [~160 for O. ornatus vs. ~105 for O. 
rapanui (MNHN specimens)], more gill la- 
mellae [13-14 for O. ornatus vs. 11-12 for 
O. rapanui (Voss 1979)]; and distinguishing 
color patterns (deep maroon red with white 
longitudinal stripes and spots for O. ornatus 
vs. white base color with fine purple brown 
chromatophores on dorsal surfaces for O. 
rapanul). 

Two species are known only from juve- 
nile specimens. Octopus taprobanensis Rob- 
son, 1926 was described from a single ju- 
venile of indeterminate sex (14.4 mm ML, 
BMNH 1925.11.23.2), collected from the 
Pearl Banks off Sri Lanka. It shows an arm 
formula of 1.2.3.4 and has 13-14 gill la- 
mellae. Octopus teuthoides Robson, 1929 
also was described from a juvenile of in- 
determinate sex (15.8 mm ML, BMNH 
1928.3.28.1), collected from Vanuatu. It also 
shows the 1.2.3.4 arm formula and has 11 
gill lamellae. Until the juvenile stages of 
members of the Octopus macropus group 
are resolved, it will not be possible to de- 
termine the status of these species. 

Rochebrune proposed a new genus and 
species, Eledonenta filholiana Rochebrune, 
1884, based on a male specimen from Fiji. 
Diagnostic characters for the genus were long 
arms with a single row of suckers. This per- 
ceived, single row of suckers led Roche- 
brune to place the genus in the Eledoninae 
Gray, 1849. The type specimen of this spe- 
cies was examined in Paris and is a poorly- 
fixed elongate specimen. The elongation of 
the arms appears to be an artefact of han- 
dling and/or fixation. There are two rows of 
suckers but arm elongation has spaced the 
suckers sufficiently for Rochebrune to in- 
terpret them as forming a zig-zag single row. 
Although poorly preserved, E. filholiana is 
clearly recognizable as a member of the Oc- 
topus macropus group. It has the typical arm 
formula of 1.2.3.4 and 10 gill lamellae. The 
status of this species needs resolution. Oc- 
topus ornatus is distinct from this species 
with higher gill counts (13-14 for O. ornatus 
vs. 10 for E. filholiana) and higher hecto- 


VOLUME 106, NUMBER 4 


cotylized arm sucker count (~160 for O. 
ornatus vs. 81 for holotype of E. filholiana). 

Voss (1981) synonymized Callistoctopus 
arakawai Taki, 1964 with O. ornatus. Other 
works have reported C. (or Octopus) araka- 
wai from Japanese and Chinese waters (Taki 
1964, 1965; Dong 1979; Taki 1981; Dong 
1987; Okutani, Tagawa & Horikawa 1987; 
Nishimura, 1992). Photographs, illustra- 
tions and descriptions provided in these 
works indicate the characteristic color pat- 
tern and morphology that identify these re- 
cords as O. ornatus. The reports of O. or- 
natus from southern Japan (Taki 1964, 
1965, 1981) are likely to be vagrant indi- 
viduals carried north to the warm bays and 
inlets of southern Japan during their plank- 
tonic phase by the warm Kuroshio Current. 
This species does not, however, appear per- 
manently established in the warm temper- 
ate waters of southern Japan. 

Recent research into tropical Australian 
octopuses (Norman, 1991, 1992, 1993a, 
1993b, 1993c) has resulted in the descrip- 
tion of four new species (O. alpheus, O. as- 
pilosomatis, O. dierythraeus and O. graptus, 
all Norman, 1993a). These octopuses ex- 
hibit characters typical of the Octopus mac- 
ropus group; all share an arm formula of 
1.2.3.4, moderate to high gill counts (10- 
14), multicuspid radula and nocturnal ac- 
tivity patterns. Full descriptions, and delin- 
eation of these species from O. ornatus, are 
presented in Norman (1993a). 

The distribution of O. ornatus is expand- 
ed to include the east coast of Australia and 
the South Pacific Ocean (Fig. 5). New rec- 
ords within the known range also are re- 
ported from the Indian Ocean, Asia and the 
North Pacific Ocean. Octopus ornatus ex- 
hibits a distribution typical of shallow-wa- 
ter, wide-ranging, tropical Indo-West Pa- 
cific marine taxa. As reported for other biota 
exhibiting this range (Briggs 1974), the dis- 
tribution of O. ornatus appears to be limited 
to waters within the 20°C winter isotherm. 

The specimens of O. ornatus collected 
from the warm temperate waters of New 


657 


South Wales probably result from plank- 
tonic juveniles carried south in the sporadic 
warm-water eddies of the East Australian 
Current. All were encountered on rocky 
shores of sheltered coasts, most in bays or 
inlets. Sporadic incursions of tropical spe- 
cies into temperate waters has been reported 
for many taxa (Bennett & Pope 1953, Wil- 
son & Allen 1987). Bennett & Pope (1953) 
suggested that inlets and bays along the New 
South Wales coast can act as warm water 
refuges for tropical species, enabling tropi- 
cal species to exist in such localities well 
south of their normal range. 


Acknowledgments 


This research forms part of a Ph.D. re- 
search program on octopods of the Great 
Barrier Reef and northern Australia, carried 
out in association with the Museum of Vic- 
toria and the University of Melbourne. Sin- 
cere thanks to F. G. Hochberg, M. Sweeney, 
Ce WoC. EE. Roper, T. Stranks and G. 
F. Watson for assistance with the manu- 
script and project; R. Fenwick and M. Tur- 
ner for field assistance; I. Loch, P. Colman 
and B. Rudman (AMS), J. Stanisic (QMB), 
C. F. E. Roper and M. Sweeney (USNM), 
T. Gosliner and E. Kool (CASIZ), F. G. 
Hochberg and H. Chaney (SBMNH), F. 
Naggs (BMNH) and R. and G. Boucher- 
Rodoni (MNHN) for assistance and access 
to collections; S. Hochberg for kind hospi- 
tality and support; D. Paul and C. Rowley 
for assistance with photographic plates. This 
research was made possible through support 
grants from the Museum of Victoria, Uni- 
versity of Melbourne, Victorian Institute of 
Marine Sciences, Australian Museum Post- 
graduate and Keith Sutherland Awards, 
Joyce W. Vickery Research Fund (Linnean 
Society of NSW), Ethel Mary Read Fund 
(Royal Zoological Society of New South 
Wales), Ian Potter Foundation, Hawalian 
Malacological Society, Royal Society of 
Victoria, Western Society of Malacologists, 
and the Harkness Fellowships, Common- 
wealth Fund. 


658 


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. 1993a. Four new species of the Octopus mac- 

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. 1993b. Systematics and biogeography of the 


VOLUME 106, NUMBER 4 


shallow-water octopuses (Cephalopoda: Octo- 

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659 


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Department of Invertebrate Zoology, 
Santa Barbara Museum of Natural History, 
2559 Puesta del Sol Rd., Santa Barbara, 
California 93105, U.S.A. 


Appendix.— Octopus ornatus Gould, 1852: 
material examined 


Neotype: HAWAII: 1 ¢: 93.7 mm ML, USNM 
730020, Oahu Island, Black Point (21°15'’N, 157°48'W), 
coll. S. Kempf, 11 Jan 1976. 

Australian material: QUEENSLAND: Capricorn 
Bunker Group, One Tree Island, 23°30’S, 152°05’E: 1 
3: 45.9 mm ML, NMV F57920, reef channel, 2 m, coll. 
M. Norman, 15 Oct 1989 (1930 hr); 1 2: 77.3 mm ML, 
NMV F57919, reef channel, 2 m, coll. M. Norman, 3 


660 


Sep 1990 (0430 hr, in mouth of lair, flushed with 
CuSO,); 1 6: 83.2 mm ML, NMV F57921, The Gutter, 
0.2 m, coll. M. Norman, 4 Sep 1990 (0400 hr); 1 2: 
83.5 mm ML, NMV F57922, The Gutter, 0.2 m, coll. 
M. Norman, 3 Sep 1990 (0215 hr); 1 6: 102.8 mm ML, 
NMV F57923, The Gutter, 0.2 m, coll. M. Norman, 
3 Sep 1990 (0330 hr); 1 46: 103.5 mm ML, NMV F57924, 
reef channel, 0.2 m, coll. M. Norman, 4 Sep 1990 (0430 
hr); 1 6: 104.2 mm ML, NMV F57918, The Gutter, 
0.2 m, coll. M. Norman, 3 Sep 1990 (0215 hr); NEW 
SOUTH WALES: 1 ¢: 75.7 mm ML, AMS C115738, 
north of Sydney, Newport (33°39’S, 151°19’E), inter- 
tidal on rock platform, coll. I. Bennett, 27 Jul 1962; 1 
6: 84.4 mm ML, AMS C156207, near Laurieton, Lake 
Cathie, 31°33’S, 152°5’E: coll. J: Ibbott= Mar-1977: “1 
6: 97.1, AMS C115737, Sydney, Manly, Fairy Bower 
(33°47'S, 151°17’E), 19 Jun 1962 (in fish net); 1 2: 132.0 
mm ML, AMS C169234, off Tuncurry (32°47’S, 
151°29’E), coll. J. C. Moore, 1965. 

Other material: VATAO ISLAND: 1?: 11.0 mm ML, 
USNM 817781, 19°48’S, 178°15'W, 4.6-9.1 m, coll. 
V. G. Springer, stn. 86, 14 Jun 1986 (poison station); 
FIJI: 12: 17.1 mm ML, USNM 817782, Rotuma, east 
end of Malaha, north coast, 12°30’S, 177°05’E, 0-1.8 
m, coll. V. G. Springer, ca. 15 May 1986 (rocky (lava) 
shore with some sand and coral rubble, rotenone sta- 
tion); 1?, 1 6: 17.1, 51.1 mm ML, USNM 817641, north 
east corner of Rotuma, just west of wharf, 12°30’S, 
177°05’E, 10.5 m, coll. V. G. Springer, 12 May 1986 
(isolated reef patch, mostly dead coral rocks, rotenone 
station); PHILIPPINES: 1?, 1 6: 22.5, 44.4 mm ML, 
USNM 817642, Batanes, Batan Island, White Beach 
past Mahatae, 20°24’45’”N, 121°55'02”E, 0-6 m, coll. 
D. Johnson, 22 Apr 1987 (surge channel at outer edge 
of reef, rotenone station); PALAU ISLANDS: 1 2: 22.9 
mm ML, CASIZ 031970, Angaur Island, Gangaraoi 
(6°55'N, 134°09’E), coll. De Witt, 22 Oct 1957; CHA- 
GOS ARCHIPELAGO: 3 2: 23.0-38.4 mm ML, USNM 
817640, Eagle Island, 6°10’45”S, 71°21'32”E, 0.5—1 m, 
coll. R. Winterbottom, 27 Nov 1979 (1015-1230 hr, 
tidal flats and channels, low tide, rotenone); 1 2: 87.0 
mm ML, BMNH unreg., Peros Banhos, Ile du Coin, 
off Jetty (5°18’S, 72°00’E), 6 m, coll. A. Shepherd, Joint 
Services Chagos Expedition 1978, Mar 1979 (Acc. No. 
2307); HAWAII: 1?: 26.7 mm ML, USNM 817783, 
Waianae (21°26’N, 158°11'W), coll. USBCF, Hono- 
lulu, 1 Jul 1951 (2100-2230 hr, attracted to night light); 
1 6: 27.9 mm ML, USNM 817784, Midway Island 
(28°12'N, 177°24'W), coll. USBCF, Honolulu, Jul or 
Aug 1955 (attracted to night light); 1 6: 31.8 mm ML, 
CASIZ 021564, Oahu, Honolulu Reef (21°30'N, 
158°00'W), coll. USS Albatross, 1902 (Berry voucher 
specimen SSB#179, CASIZ holotype no. 506); 1 2: 43.4 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


mm ML, CASIZ 034975, Leeward Islands, Laysan Is- 
land, west side on reef, 25°46.3’N, 171°44.7'W, coll. 
R. R. Harry, 27 Jun 1951; 1 2: 45.8 mm ML, CASIZ 
035015, east of Laysan Island, anchorage off Aro Reef, 
25°25:5'N, 170°41.5"W, coll. R. R? Harry, 5 Jul 1954: 
1 6: 73.8 mm ML, USNM 817787, 21°15'N, 157°44'W, 
Albatross surface tow net survey, serial no. 286, 2 Dec 
1891 (1700 hr, caught at surface; 1 6: 79.4 mm ML, 
CASIZ 034977, Oahu, Honolulu Fish Market (21°30'N, 
158°00'W), coll. D. S. Jordan and Joseph Grinnell (from 
Stanford collection); 1 4: 88.1 mm ML, USNM 214609, 
Oahu, Honolulu Market, 21°20’N, 157°55'W, Alba- 
tross cruise, between 1902 and 1914; REUNION IS- 
LAND, 1?: 26.9 mm ML, NMV F60139, (21°06’S, 
55°36’'E), coll. M. Jay, MNHN, 1991 (station N2); AL- 
DABRA ATOLL: 1 6: 30.8 mm ML, MNHN 4.7.923, 
Aldabra Island (10°30’S, 46°30’W), 1954; 2 6,29: 51.9- 
121.0 mm ML, USNM 817638, West Island, 9°22.8’S, 
46°12.4’E, 0.75 m, coll. H. A. Fehlmann, 14 Aug 1967 
(1700-1900 hr, on sand and coral rubble, rotenone); 
SOCIETY ISLANDS: 1 4, 2 9: 37.3-58.0 mm ML, 
NMV F30256, Tahiti, Teahupo Presquile de Taiarapu 
(17°51'S, 149°15’W), 0.1 m, coll. R. Boucher, 18 Mar 
and 1 Apr 1972 (rubble bottom in lagoon at night); 
unmeasured, USNM 575419, Raiatea, Uturoa, station 
77, (16°44’S, 151°25’W), 0.3 m, coll. H. A. Rehder, 28 
Apr 1957 (muddy sand flat east of church and school, 
swimming); unmeasured, USNM 576016, Tahiti, 
Taone (17°30’N, 149°30'W), coll. R. Sixberry; COOK 
ISLANDS: 1 6: 38.0 mm ML, CASIZ unreg., Ka’umata, 
Majaia, 21°54'30”S, 157°58’00”W, coll. Vanderbilt 
Foundation Cook Islands Expedition, 25 May 1958; 1 
6: 44.1 mm ML, CASIZ 034026, Manaia, 21°54’30’S, 
157°58'00’W, coll. D. S. Marshall; 1 9: 85.2 mm ML, 
USNM 817639, Raratonga, Arorangi (21°13’S, 
159°49’W), coll. G. Paulay, 14 Oct 1984 (on fringing 
reef, swimming at night); MADAGASCAR: 1 6: 67.2 
mm ML, SBMNH 64492, Libanona Beach, Taolanaro 
(Fort Dauphin), 25°02’S, 47°00’E, coll. Henry W. Cha- 
ney, 23 Mar 1990 (active on reef terrace amongst tide- 
pools and algae, intertidal at night); MARIANA IS- 
LANDS: 1 2: 72.5 mm ML, CASIZ 031971, Guam, 
0.5 mile SW of Agat Village, sand flat off north side of 
Bangi Point, 13°22'36”N, 144°38'53’E, coll. Fehlman, 
12 Oct 1958; EAST AFRICA: 1 6: 85.0 mm ML, USNM 
817778, Kenya, Mombasa Fish Market (4°04’S, 
39°40’E), coll. J. C. Miguel, 11 Feb 1979 (purchased 
at market); SEYCHELLE ISLANDS: 1 ¢: 92.1 mm 
ML, MNHN 4.4.880, (4°30’S, 55°30’E), rec. Dussu- 
mier, 1830; EASTER ISLAND: 1 ¢: 96.1 mm ML, 
MNHN 3.12.784, (27°05’S, 109°20’W); 1 2: 104.3 mm 
ML, MNHN 3.10.753, (27°05’S, 109°20’W). 


PROC. BIOL. SOC. WASH. 
106(4), 1993, pp. 661-665 


GRASSLEIA HYDROTHERMALIS, A NEW GENUS AND 
SPECIES OF AMPHARETIDAE (ANNELIDA: POLYCHAETA) 
FROM THE HYDROTHERMAL VENTS OFF THE 
OREGON COAST (U.S.A.), AT GORDA RIDGE 


Vivianne Solis-Weiss 


Abstract. —A new ampharetid genus (Grassleia hydrothermalis) from the hy- 
drothermal vents found off the coasts of Oregon is described. It differs from 
other genera in its unusual abdominal shape: very short (seven setigers) and 
constricted in the first. Its affinities with close related genera are discussed. 


As part of the biological material collect- 
ed on the June 1988 expedition to the Es- 
canaba Trough region of Gorda Ridge, off 
the coasts of Oregon and California, with 
the DSRV Alvin (Grassle & Petrecca 1992), 
J. F. Grassle found some ampharetids which 
he kindly provided me for study since we 
were examining the ampharetids from the 
Guaymas basin expedition of February 
1988. 

Gorda Ridge is a 300 km segment of vol- 
canically active oceanic ridge extending from 
the Blanco Fracture zone (43°00'N) to the 
Mendocino Fracture zone (40°20'N) and is 
included in the proclaimed U.S. Exclusive 
Economic Zone (Grassle & Petrecca 1992). 
The polychastes were found to dominate in 
all areas sampled, contributing, in the soft 
sediment from which the ampharetids were 
taken, to more than 80% of the fauna. 

The ampharetids collected there, in ex- 
cellent state of preservation, were found to 
belong to a new genus which is described 
below. 

The family Ampharetidae is well known 
in deep marine environments (Day 1967), 
and has been found also to colonize the hy- 
drothermal vents extensively (Desbruyeres 
et al. 1985), along with members of the 
closely related Alvinellidae (Desbruyeres & 
Laubier 1986). 


Materials and Methods 


Specimens were collected with the sub- 
mersible DSRV Alvin in its dive 2042 of 
June 12, 1988 (40°56.71'N, 127°29.31'W), 
in the Escanaba Trough region of Gorda 
Ridge, 70 nautical miles northward from 
the intersection of the ridge with the Men- 
docino Fracture Zone, within 200 nautical 
miles off the coasts of Oregon and Califor- 
nia, U.S.A. (Grassle & Petrecca 1992). 

The sample was obtained using a DSR V 
Alvin tube core as a scoop to shovel sedi- 
ment into a collection box of the DSRV 
Alvin basket. The amount of sediment was 
the equivalent of at least two steel box cores 
(225 cm? each) specifically designed for use 
by the DSRV Alvin (Grassle & Petrecca 
1992). 

At the surface, the material was rinsed 
Over sieves with mesh openings of 300 and 
63 um. Animals retained in the sieves were 
fixed in 10% formalin and stored in 80% 
ethanol (Grassle & Petrecca 1992). 

The types are deposited in the following 
collections: ICML—MEX (Instituto de Cien- 
cias del Mar y Limnologia, Universidad Na- 
cional Autonoma de México, Mexico); 
USNM (National Museum of Natural His- 
tory, Smithsonian Institution, Washington, 
D.C., U.S.A.); BMNH (The Natural History 


662 


Museum, London, England); AM (Austra- 
lian Museum, Sydney, Australia); HZM 
(Zoologisches Museum, Hamburg, Ger- 
many) and J. F. Grassle, personal collection, 
Rutgers University, New Jersey, U.S.A. 


Results 


Family Ampharetidae Malmgren, 1866 
Subfamily Ampharetinae Chamberlin 
Grassleia, new genus 


Type species.—Grassleia hydrothermalis, 
new species. 

Diagnosis. —Body short with fifteen tho- 
racic setigerous segments, of which last ten 
uncinigerous. Abdomen truncate, distinctly 
constricted in transition zone, seven ab- 
dominal uncinigerous segments present. 
Abdominal notopodia absent. Small paleae 
present. Four pairs of smooth branchiae 
present; branchiae fused at base. No mid- 
dorsal space between branchial groups. No- 
topodial and neuropodial cirri absent. Anus 
terminal. 

Etymology.—The genus is dedicated to 
Dr. J. Frederick Grassle in an attempt to 
show my sincere appreciation for all the un- 
selfish help and friendship provided over 
the past years in addition to providing the 
present material for study. 

Discussion. —-This genus differs from all 
the other genera in the ampharetids mainly 
by having a very short and truncate abdo- 
men, only seven setigers long, and by being 
distinctly constricted in the first abdominal 
segment. Only the genus Lysippides Hessle, 
1917 has eight abdominal segments, with 
the rest of the genera having up to 60 ab- 
dominal segments (Holthe 1986b). 


Grassleia hydrothermalis, new species 
Fig. 1A—D 


Material examined.—Escanaba Trough, 
Gorda Ridge, DSRV Alvin dive 2042, 12 
Jun 1988, 40°56.71'N, 127°29.31'W, depth 
3271) m: 

Holotype: USNM (157692), 21 para- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


types: (USNM 157693, 10; BMNH-ZB 
1993:18-19, 2; HZM P-21748, 2; ICML 
POH-68001, 7. Additional specimen de- 
posited: J. F. Grassle pers. collection, Rut- 
gers University, 2; AM, 3. 

Description. — The description is based on 
the holotype unless specified otherwise. All 
the specimens are complete. 

Holotype 9.6 mm long, 1.6 mm wide. 
Color in alcohol white, nearly translucent. 
Sizes of paratypes from 3.5 to 9.1 mm, with 
about half of them measuring around 7 mm. 

Prostomium slightly pointed, indistinctly 
trilobed with two longitudinal nuchal slits 
(Fig. 1A). No glandular ridges present. Oral 
tentacles not observed in any of the organ- 
isms collected. Segment 1 enlarged ventrally 
to form the lower lip. Buccal aperture large. 
Segment 2 distinct, achaetous (Fig. 1A). 

Thorax with fifteen setigers, the last ten 
being uncinigerous. Thorax measurements 
vary from 2.85 to 7.9 mm in length and 0.6 
to 1.6 mm in width, the holotype being the 
largest. Segment 3 laterally enlarged, over- 
lapping segment 2, with small, bristlelike 
paleae. Segments 3 and 4 fused laterally and 
hard to differentiate. Branchiae located on 
segment 4 (Fig. 1A). There are four pairs of 
branchiae united basally. No gap between 
the branchial groups. Each branchia has a 
branchiophore fused to form a basal mem- 
brane, and then emerges independently as 
a finely annulated rounded filament, the 
longest measuring about 3.4 mm in the ho- 
lotype (Fig. 1B). The length of the branchiae 
varies in direct relation to size, the longest 
per specimen varying from 1.6 to 4.6 mm 
in the paratypes. 

Notosetae start on segment 4. First no- 
topodial lobe reduced, setae arising close to 
the branchial base (Fig. 1A). 

The other notopodial lobes are elongate 
and remain the same size throughout the 
thorax. In some paratypes they are some- 
what larger towards the posterior thorax, 
and their setae are distinctly longer. They 
bear about seven to twelve unilimbate, 
slightly recurved and distally pointed cap- 


VOLUME 106, NUMBER 4 


illary setae of different lengths. The longest 
notoseta per specimen varies from 0.2 to 
0.7 mm (measuring 0.4 mm in the holo- 
type). 

The short, low-ridged neuropodial tho- 
racic lobes (uncinigerous tori) begin at se- 
tiger 6 and continue throughout the thorax. 
Thoracic uncini are located in single trans- 
verse rows, 44 to 50 per row. The number 
of uncini per setiger varies widely according 
to animal size: the holotype being the larg- 
est, has the largest number of uncini per 
setiger. Others can be as low as from 14 to 
17. Each uncinus bears 10 teeth in 4 trans- 
verse rows above a rounded basal prow. 

The abdomen, consisting of seven unci- 
nigerous segments, is unusually well set off 
from the thorax, nearly quadrangular when 
seen from above, narrower and separated 
from the former by a constriction in the first 
abdominal setiger (Fig. 1C). It measures 1.7 
mm long and 0.8 mm wide in the holotype 
and varies from 0.6 to 1.85 mm in length 
and 0.3 to 0.8 mm in width in the paratypes. 

The first abdominal torus is located in the 
constriction separating the thorax from the 
abdomen and its shape is intermediate be- 
tween the thoracic and abdominal tori, re- 
sembling more closely the thoracic tori, that 
is: not elongate, although it is a bit higher 
than the thoracic ones. The 6 following ab- 
dominal uncinigerous tori are rounded and 
elongate. Abdominal uncini are in single 
rows fringing the edge of each torus num- 
bering 26 to 28 per row in the holotype. 
Their number varies in direct relation to 
specimen size and can be as low as from 7 
to 9. Each uncinus bears 10 teeth arranged 
in 4 rows around a large single tooth and 
above a rounded basal prow (Fig. 1D). 

Abdominal notopodia, as well as noto- 
podial, neuropodial and anal cirri absent. 

Anus rounded and terminal, surrounded 
by 2 large and several small papillae. 

Tube: Only fragments of a tube could be 
seen on one of the specimens as a thin trans- 
lucent membrane almost completely cov- 
ered with fine sand grains and small debris. 


663 


Etymology. —The name refers to the hab- 
itat of the species, the hydrothermal vents. 

Biological notes.—The organisms were 
found only in soft sediments (never in hard 
substrates) at the base of active hydrother- 
mal mounds, or in the surroundings. They 
were located where hydrothermal fluid per- 
colates to the surface, and were found to 
dominate there, representing 37% of the 
fauna collected (Grassle & Petrecca 1992). 
They are, together with the majority of the 
polychaete fauna found in the area, surface 
deposit feeders. A detailed description of 
the fauna found at this site can be found in 
Grassle & Petrecca (1992). 

Distribution. —Known only from the 
Gorda Ridge. 


Discussion 


The species differs from other ampharetid 
species as indicated in the genus. 

Most of the ampharetid genera of both 
subfamilies: Melinninae and Ampharetinae 
have either 12 or 14 thoracic uncinigers 
(Fauchald 1977, Holthe 1986a). One genus 
has 15 (Weddellia Hartman 1967); 6 genera, 
in addition to the new genus, have 10, and 
2 genera have 9 uncinigers. Of the six genera 
with 10 uncinigers, 2, Melinnopsis and Mel- 
innopsides belong to the subfamily Melin- 
ninae; the other 4, plus the new genus are 
members of the subfamily Ampharetinae. 
These four genera include Decemunciger 
Zottoli 1982, Melinnata Hartman, 1965, 
Mexamage Fauchald, 1972 and Muggoides 
Hartman, 1965. 

Two genera, Melinnata and Muggoides, 
have 3 pairs of branchiae rather than 4, in 
addition to other important characters like 
a dorsal ridge across the dorsum on anterior 
thoracic segments and the last thoracic no- 
topodia elevated with modified notosetae in 
the case of Muggoides. The other 3 genera 
Decemunciger, Grassleia and Mexamage 
have 4 pairs of branchiae. The branchiae 
are arranged in a longitudinal series in Mex- 
amage and in transverse rows in Decemun- 
ciger and Grassleia. In Mexamage, the 


664 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


x) 
Kk) 


ds qqacsteg mentee. , 
Ail : te 
« = ee a hee 
: ws 2 : 
a ee , o. 
=i 


A qust 


’ 
1, 
os 
a 
‘\ 


gate ji 
Lad , ae 


ware 
\ 
Tu ay a 


MULL 


app & 
cy cece hued 

SM TTCCULL 

er 14 


Fig. 1. Grassleia hydrothermalis: A, Lateral view of entire worm; B, anterior end, dorsal view; C, posterior 
end, dorsal view; D, mid-abdominal uncinus, lateral view. A—C: holotype (USNM 157692) D: paratype (POH- 


68001). Abbreviations: I, II & III = segments 1, 2 & 3; p = prostomium. 


branchiae are each clearly related toa con- in all 4 specimens observed by Fauchald 
secutive segment; moreover, there isa wide (1972), so that no comparison can be made 
gap between the branchiae, in addition toa regarding this character. In Melinnata, the 
““complex branchial membrane.”’ Unfortu- number of abdominal segments is also un- 
nately, the abdominal segments were absent known, and in Muggoides there are 13 ab- 


VOLUME 106, NUMBER 4 


dominal segments. Decemunciger Zottoli, 
1982, consists of 13 thoracic segments and 
has 14 abdominal uncinigers. Grassleia con- 
sists of 15 thoracic setigers and has 7 ab- 
dominal uncinigers. 

It is interesting to note that all 5 genera 
of Melinninae come from deep waters (Fau- 
chald 1972, Hartman 1965, Zottoli 1982). 


Acknowledgments 


Thanks are due to Dr. K. Fauchald, cu- 
rator, USNM, Smithsonian Institution, for 
his careful review and many useful com- 
ments on the manuscript. Dr. Marian Pet- 
tibone, Professor Emeritus, USNM, Smith- 
sonian Institution, made useful comments 
on an earlier draft of the manuscript for 
which I am grateful. Dr. J. F. Grassle, in 
addition to revising the manuscript, pro- 
vided the material, for which I am deeply 
grateful. The editor and two anonymous re- 
viewers suggested improvements to the 
manuscript for which they are thanked. 


Literature Cited 


Day, J. W. 1967. A monograph of the Polychaeta of 
Southern Africa. Part 2, Sedentaria.— British 
Museum (Natural History) London, Publica- 
tions No. 656, xvii, pp. 459-878. 

Desbruyeres, D., F. Gaill, L. Laubier, & Y. Foulquet. 
1985. Polychaetous annelids from hydrother- 
mal vents ecosystems: an ecological overview. — 
Bulletin of the Biological Society of Washington 
6:103-116. 

—, & L. Laubier. 1986. Les Alvinellidae, une 
famille nouvelle d’annélides polychetes inféo- 
dées aux sources hydrothermales sous-marines: 


665 


systématique, biologie et écologie.— Canadian 
Journal of Zoology 64:2227-2245. 

Fauchald, K. 1972. Benthic polychaetous annelids 
from deep waters off western Mexico and ad- 
jacent areas in the eastern Pacific Ocean. — Allan 

Hancock Monographies on Marine Biology 7:1- 

ed pee 

. 1977. The Polychaete worms. Definitions and 

keys to the orders, families and genera. — Nat- 

ural History Museum of Los Angeles County 

Science series 28:1—-188. 

Grassle, J. F., & R. Petrecca. 1992. Soft-sediment 
hydrothermal vent communities of Escanaba 
Trough.— Bulletin of the United States Geolog- 
ical Survey (in press). 

Hartman, O. 1965. Deep water polychaetous anne- 

lids off New England to Bermuda and other 

North Atlantic areas.—Allan Hancock Foun- 

dation Occasional Papers 28:1—378. 

. 1967. Polychaetous annelids collected by the 

USNS Eltanin and Staten Island cruises, chiefly 

from Antarctic seas.—Allan Hancock Mono- 

graphies on Marine Biology 2:1—387. 

Hessle,C. 1917. Zur Kenntnis der Terebellomorphen 
Polychaeten.— Zoologiska bidrag fran Uppsala 
5:39-258. 

Holthe, T. 1986a. Polychaeta Terebellomorpha. Ma- 

rine invertebrates of Scandinavia vol. 7.—Uni- 

versitetsforlaget, Oslo, 192 pp. 
1986b. Evolution, systematics and distri- 
bution of the Polychaeta Terebellomorpha, with 

a catalogue of the taxa and a bibliography. — 

Gunneira 55:1-236. 

Zottoli, R. 1982. Two new genera of deep-sea poly- 
chaete worms of the family Ampharetidae and 
the role of one species in deep-sea ecosystems. — 
Proceeding of the Biological Society of Wash- 
ington 95:48—-57. 


Lab. de Ecologia Costera (Poliquetos), In- 
stituto de Ciencias del Mar y Limnologia, 
U.N.A.M., Apdo postal 70-305, Mexico, 
D.F., 04510, Mexico. 


PROC. BIOL. SOC. WASH. 
106(4), 1993, pp. 666-672 


A NEW SPECIES OF SCALE-WORM, HARMOTHOE 
COMMENSALIS (POLYCHAETA: POLYNOIDAE), FROM 
MANTLE CAVITIES OF TWO CHILEAN CLAMS 


N. Rozbaczylo and J. I. Canete 


Abstract. —A new species of polynoid, Harmothoe commensalis, is described 
from the mantle cavities of two subtidal clams, Gari solida and Semele solida, 
on the Chilean coast. About 50% of the clams investigated at Bahia La Herradu- 
ra, Coquimbo, contained single individuals of the new polynoid. This is the 
third documented record of commensal association between polynoid poly- 


chaetes and bivalve molluscs. 


Many polychaetes live in association with 
other animals. Polynoids, particularly, are 
commensals on or with numerous inverte- 
brates, such as sponges, cnidarians, mol- 
luscs, echinoderms, and other polychaetes 
(Pettibone 1953). Clark (1956) compiled a 
list of commensal polychaetes and their re- 
spective hosts that included thirty species 
from the Polynoidae and twenty species 
from other families. None of the polynoid 
species he mentioned had been found living 
in association with lamellibranch molluscs. 

Commensal polynoid polychaetes have 
been recorded from bivalve molluscs on two 
previous occasions. Pettibone (1984) de- 
scribed Branchipolynoe symmytilida, from 
mantle cavities of giant deep-sea mussels, 
Bathymodiolus thermophilus Kenk & Wil- 
son, 1985, at 2500 m in the Galapagos Rift 
vent area. A second species, B. seepensis, 
was also described by Pettibone (1986), from 
mantle cavities of mussels in the abyssal 
eastern Gulf of Mexico, near hypersaline 
seep-sites. 

During dissection of specimens of the 
subtidal clams Gari solida (Gray, 1828) and 
Semele solida (Gray, 1828) collected from 
Bahia La Herradura, at Coquimbo, poly- 
chaetes were found living in the mantle cav- 
ities (Fig. 1). Examination of the commensal 
revealed it to be an undescribed species of 
Polynoidae that we describe here as Har- 


mothoe commensalis, new species. This is 
the third record of commensalism between 
a polynoid and bivalve molluscs and is the 
first case reported from the Chilean south- 
east Pacific (Rozbaczylo 1985). 

Type specimens are deposited in the Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. 
(USNM)); Sala de Sistematica, Departamen- 
to de Ecologia, Pontificia Universidad Ca- 
tolica de Chile, Santiago (SSUC); and in the 
personal reference collections of both au- 
thors. 


Materials and Methods 


Clams were collected by diving at Bahia 
La Herradura, Coquimbo, Chile, at 9 and 
12 m depth from four 1 m? X 0.3 m deep 
samples in the north-west sector of the bay, 
at two sites characterized by different sed- 
iment granulometry composition of the sed- 
iments. Ninety-four bivalves (16 S. solida 
and 78 G. solida) were collected and ex- 
amined. Additionally, seven specimens of 
G. solida and three of S. solida were bought 
from fishermen at a cove in Tomé, Bahia 
de Concepcion, who said the clams had been 
collected between 10 and 15 m depth, near 
Tome. To select type specimens of the poly- 
noids, additional clams were collected at 10 
m depth at Bahia La Herradura, Coquimbo, 
on 24 Jan 1991. Scanning electron micro- 


VOLUME 106, NUMBER 4 


scopic (SEM) observations and photographs 
were executed by the first author with a JEOL 
JSM-25SII. Figures were prepared by means 
of a drawing tube on a Wild M-5 micro- 
scope. 


Harmothoe commensalis, new species 
Figs. 1—4 


Material examined. —Central Chile: Ba- 
hia La Herradura, Coquimbo, 29°58’S, 
71°22'W, 10 m, J. I. Canete, coll., from G. 
solida and S. solida, 24 Jan 1991, holotype 
(USNM 157690), 4 paratypes (USNM 
157691), and 2 specimens (SSUC 6700); 
Bahia La Herradura, from G. solida, 10—20 
m, J. I. Canete, coll., Dec 1986, 2 speci- 
mens; Bahia La Herradura, from S. solida, 
10-15 m, J. I. Canete, coll., 2 specimens. 
Tomé, Bahia de Concepcion, 36°37’S, 
72°57'W, from S. solida, 10-15 m, 20 Mar 
1989, 1 specimen (SSUC 6701). 

Description. —Length of adults 14-18 mm, 
width 3.5—4.2 mm, excluding setae, with 37— 
40 segments; body nearly linear, tapering 
slightly anteriorly and posteriorly (Fig. 2); 
oval in cross section. Elytra with two pig- 
mented zones of olivaceous to brownish 
color: larger medial and smaller lateral zones 
with colorless areas between, giving ap- 
pearance of a “heart’’ on each elytron, and 
continuous with colorless periphery (Fig. 3A, 
B). Body flesh-colored in live specimens, 
but changing according to sexual state of 
maturity. Elytra 15 pairs arranged on elytro- 
phores of segments 2, 4, 5, 7, alternate seg- 
ments to 23, 26, 29, and 32. Specimens un- 
der 10 mm in length with 14 pairs of elytra; 
these strongly imbricated, completely cov- 
ering dorsum, except for last 5—6 segments. 
Elytra variable in shape and size along body, 
becoming larger posteriorly; first pair of el- 
ytra circular, covering prostomium, follow- 
ing two pairs subreniform; and rest oval in 
shape. Elytral surface smooth with a few 
scattered microscopic tubercles (Fig. 3A); 
second and third pairs of elytra with small 
group of microtubercles near anterior curves 
of elytra (Figs. 3B, C, 4a—c); margins smooth, 


667 


lacking papillae. Elytrophores large, bul- 
bous. Dorsal cirri and dorsal tubercles on 
non-elytrigerous segments; dorsal tubercles 
conspicuous, nodular. Dorsal cirri with glo- 
bose cirrophores pigmented on anterior and 
posterior sides; styles fusiform, not extend- 
ing beyond tips of setae, pigmented in the 
lower half of its length, and covered with 
small pyriform papillae. Prostomium (Figs. 
3A, 4a) of light tan color, bilobed, slightly 
wider than long, with convex lateral mar- 
gins; with well developed cephalic peaks, 
diverging laterally from median antenna; 
median groove extending about half length 
of prostomium. Median antenna long, pap- 
illated, inserted on a large conspicuous ovoid 
ceratophore in anterior notch; lateral anten- 
nae inserted ventrally on small cerato- 
phores; styles short, basally globose, distally 
filiform, slightly pigmented and with small 
pyriform papillae. Palps up to twice length 
of prostomium, wider basally, tapering dis- 
tally to short digitiform tips; with short pa- 
pillae arranged in close-set longitudinal 
rows. Two pairs of black eyes in trapezoidal 
arrangement, anterior pair slightly larger lo- 
cated on lateral margin of middle of pro- 
stomium, posterior pair smaller displaced 
medially. First segment with tentaculo- 
phores lateral to prostomium, each with 
dorsal and ventral tentacular cirri similar to 
median antenna, with internal acicula and 
single curved notoseta on medial side. Sec- 
ond segment with first pair of large elytro- 
phores lateral to posterior half of prosto- 
mium, biramous parapodia and long ventral 
buccal cirri, similar to tentacular cirri, on 
neuropodia lateral to ventral mouth. An- 
terior end of evaginated pharynx with 18 
triangular papillae, nine dorsally and nine 
ventrally and two pairs of pointed jaws of 
golden to brown color; invaginated pharynx 
extending up to segment 11. Parapodia bi- 
ramous (Fig. 3D, E). Notopodium rounded 
lobe on anterodorsal face of neuropodium, 
extending into acicular lobe from which 
acicula projects; slightly pigmented on an- 
terior and posterior sides. Notosetae amber 


668 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. yl: 
mensalis, new species, “in situ.” 


colored, numerous (30-35), forming radi- 
ating bundle of four or five rows, increasing 
in length from dorsal to ventral rows (Fig. 
4d), longest notosetae not extending beyond 
tips of neuropodium. Notosetae slightly 
stouter than neurosetae, slightly curved, with 
unidentate blunt tips, with 33-34 transverse 
rows of spines (Figs. 3F, 4e). Neuropodial 
presetal aciculcar lobe diagonally truncate 
distally, with short stout digitiform supra- 
acicular process; neuroacicula only slightly 
stouter than notoacicula; postsetal lobe 
shorter, more or less rounded. Neurosetae 
amber-colored, from seven to nine supra- 
acicular and 39-40 subacicular. All neuro- 
setae similar (Fig. 4d); supraacicular ones 
longer, with 13-15 rows of spines; subacicu- 
lar ones shorter, decreasing in length from 
dorsal to ventral, bearing 9-13 rows of spines 
(Figs. 3G, 4f); all neurosetae with bidentate 
tips. Ventral cirri short, located at mid- 
length of neuropodia; styles basally globose 
and distally pointed; cirrophores and ven- 
tral cirri slightly pigmented, covered with 
very short scattered papillae. 


Specimen of clam Gari solida photographed immediately after dissection showing Harmothoe com- 


Pygidium small, with pair of long anal 
cirri, about as long as four or five posterior 
segments, with pigmented basal zones, and 
distally tapered, covered with small papil- 
lae. 

Geographical distribution. —Known only 
from the type locality and Bahia de Con- 
cepcion, Chile. 

The clams are distributed from Callao, 
Peru to Archipiélago de los Chonos, Chile 
(Osorio et al. 1979), therefore the geograph- 
ical distribution for H. commensalis could 
vary as other localities are investigated. 

Taxonomical remarks. —The genus Har- 
mothoe now contains in excess of 150 spe- 
cies (Hanley, pers. comm.). Many of these 
species are poorly described and illustrated 
and their distributions are poorly known. 
As a consequence, it is difficult to adequate- 
ly define the basis for new species in this 
genus. However, in this case, the unusual 
habit of commensalism with bivalve mol- 
luscs, the lack of evidence of free-living in- 
dividuals, and the lack of ornamentation on 
the elytra (unusual in the genus, but often 


VOLUME 106, NUMBER 4 


669 


Fig. 2. Harmothoe commensalis, new species, photographed alive after removal from the mantle cavity of 


the clam Gari solida. 


typical of commensals) provide strong 
grounds for the erection of a new species. 
Among the species of Harmothoe previ- 
ously described from Chilean and subant- 
arctic waters, only H. brevipalpa Bergstrom, 
1916, is morphologically similar to the new 
species but can be distinguished from it by 
the following characters. The elytral surface 
of H. brevipalpa is uniformly covered with 
conical, blunt microtubercles, except on the 
anterior and inner borders, while the elytral 
surface of H. commensalis has groups of 
microtubercles near the anterior curves of 
the elytra (Figs. 3B, 4b, c), the rest of the 
surfaces being almost smooth. In H. brevi- 
palpa, the posterior elytral borders have 
fringes of club-shaped papillae, while in H. 
commensalis the elytral borders lack papil- 
lae. In H. brevipalpa, the dorsal cirri are 
covered with papillae and the ventral cirri 
are smooth, while in H. commensalis both 
the dorsal and ventral cirri have papillae, 
although less numerous in the ventral ones. 
Also, the neuropodial acicular lobe of H. 
commensalis is less developed than in H. 
brevipalpa; and the numbers of notosetae 
(30-35) and neurosetae (46-49) are higher 


in H. commensalis, as compared to H. brevi- 
palpa: notosetae (15-20) and neurosetae 
(30-35). 

Etymology. —The name commensalis re- 
fers to the biological relationship between 
the new species and the clams. 

Ecological remarks. —In each of the four 
areas sampled, 22% to 78% of the clams 
contained a single commensal polynoid. The 
only case of more than one commensal per 
clam was two juvenile scaleworms. H. com- 
mensalis was not found free-living, sug- 
gesting it is an obligate commensal. In G. 
solida, 13 commensals were females (34%), 
19 were males (50%) and in six cases the 
sex could not be determined due to the ab- 
sence of gametes. In live animals, females 
were recognized by their reddish color, while 
males were cream-colored on the ventral 
surface. The lengths of the polychaetes 
ranged from 8.7 to 29.2 mm. There was no 
correlation between the lengths of the poly- 
noids and the clams; for instance, the largest 
clam (G. solida), at 82.4 mm long, contained 
a polynoid of only 13.5 mm in length. Poly- 
noids were not found in clams less than 60 
mm long, although samples included small- 


670 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


eer 
}, 
ton 
At Dir 
Lee Nive \ 
oF Raye | 
ee = Ms \ 
Li 
eee Hie : 
eos Eom Mr Wil “ily 
AS a = ul Oy w'(} 
Wa Be a ea A vj mr 
a waa { wp 
Ata Y Urea 
w =e Pin 
72 ae Ml] i \ { 
77 aa fy 
7 D We it! ; 
} { N " af 
wii my 
Unio 
Ih NUT 
> NW V IW ( 
‘ \ { y 
a ‘ inant nt 
Se ask lime { ‘ 
SSS SSS ; mn 
S SS Vasey pe 
~~ RS Let / 
S SS = PSN ae 
= S= =~ ~ : 
= SS aS 
= a7 ~ > =—_ 
= 
= 
Fig. 3. 


Harmothoe commensalis, new species (USNM 157691). A, anterior end in dorsal view; B, second 
left elytron; C, detail of microtubercles from anterior curved part of elytron; D, left elytrigerous parapodium, 


anterior view, acicula dotted; E, left cirrigerous parapodium, posterior view, acicula dotted; F, notoseta from 
parapodium 7; G, subacicular neuroseta, from parapodium 7. Scales = 1 mm for A, B, D, E; 0.02 mm for C, 
F, G. 


VOLUME 106, NUMBER 4 671 


/ 


q f | ey 


Ql 


Ne Ae ae 


o | 


i - 
* ait 
: c. 
‘ j pr 
v ~ 
7 4 ... . 4 *. 


Y 
| 


" 


i AS 
= 


rat ee 
Ay Ds 
+ 4a 4 
AA 6 n f 7 | 
Aa Weak | 
re) 


yr 
x 


‘4 


Sy 


Fig. 4. Harmothoe commensalis, new species, SEM micrographs: a, dorsal view of anterior end showing 
third pair of elytra, median and lateral antennae and tentacular cirri missing; b, left 3rd elytron showing 
microtubercles on anterior part; c, detail of microtubercles from anterior part of same; d, notosetae and neurosetae 
from anterior region of body; e, detail of notosetae; f, detail of neurosetae. Scales = 1 mm for a; 0.1 mm for 
b-f. 


672 


er clams. Other bivalve species of compa- 
rable size inhabiting the same area were also 
examined. The intertidal Protothaca taca 
(Molina, 1782) and the subtidal Ensis ma- 
cha (Molina, 1782), and Tagelus dombeii 
(Lamarck, 1818), were all without com- 
mensal polynoids. 

The presence of the polynoid has no ap- 
parent negative effect on the tissues of the 
clams. We do not know if the scale-worm 
shares the clams’ food particles or whether 
it feeds on pseudofeces accumulated in the 
mantle cavity of the bivalves. Thus, H. 
commensalis may be considered a com- 
mensal, as defined by Cheng (1967), because 
it derives physical shelter from the host, and 
is nourished on foods that are associated 
but are not a part of the host. 


Acknowledgments 


We thank Professor E. Gonzalez, pres- 
ently Academic Vice-Rector of the Univer- 
sidad Catolica del Norte, for his valuable 
logistics support for the collection of the 
samples and Veronica Flores for the prep- 
aration of photographic material. Drawings 
were done by Marcelo Bobadilla. Veronica 
Palma and Mariana Parra collaborated in 
the translation of descriptions published in 
German. We also acknowledge Dr. R. Al- 
bertini, Dean of the Facultad de Ciencias 
Biologicas, for the financial support for pub- 
lication of this paper. We especially thank 
Dr. Marian H. Pettibone of the National 
Museum of Natural History, Washington, 
D.C., for her valuable comments and sug- 
gestions on the manuscript. We are also 
grateful to Dr. R. Hanley for his valuable 
suggestions for improvement of the text. 


Literature Cited 


Bergstrom, E. 1916. Die Polynoiden des schwedisch- 
en Stidpolarexpedition 1901-—1903.—Zoologis- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ka Bidrag fran Uppsala 4:269-304, 2 figures, 
plates 2-5. 

Cheng, T.C. 1967. Marine molluscs as hosts for sym- 
bioses. Jn F. S. Russell, ed., Advances in marine 
biology, vol. 5. Academic Press, London, 424 
pp. 

Clark, R. B. 1956. Capitella capitata as a commensal, 
with a bibliography of parasitism and commen- 
salism in the polychaetes.—The Annals and 
Magazine of Natural History, ser. 12, 9(102): 
433-448. 

Gray, J. E. 1828. Spicilegia Zoologica, or original 
figures and short descriptions of new and unfi- 
gured animals. London 1:1-8, pls. 1-6. 

Kenk, V., & B. R. Wilson. 1985. A new mussel (Bi- 
valvia, Mytilidae) from hydrothermal vents in 
the Galapagos Rift zone. — Malacologia 26:253- 
Pid id 

Osorio, C., J. Atria, & S. Mann. 1979. Moluscos 
marinos de importancia economica en Chile. — 
Biologia Pesquera, Chile 11:3-47. 

Pettibone, M. H. 1953. Some scale-bearing poly- 

chaetes of Puget Sound and adjacent waters. 

University of Washington Press, 89 pp., pls. 1- 

40. 

1984. A new scale-worm commensal with 
deep-sea mussels on the Galapagos hydrother- 
mal vent (Polychaeta: Polynoidae).— Proceed- 
ings of the Biological Society of Washington 97: 
226-239. 

1986. A new scale-worm commensal with 
deep-sea mussels in the seep-sites at the Florida 
Escarpment in the eastern Gulf of Mexico (Poly- 
chaeta: Polynoidae: Branchipolynoidae).— Pro- 
ceedings of the Biological Society of Washington 
99:444-451. 

Rozbaczylo, N. 1985. Los Anélidos Poliquetos de 
Chile. Indice sinonimico y distribuciOn geo- 
grafica de especies. — Monografias Biologicas 3: 1— 
284. 


Departamento de Ecologia, Facultad de 
Ciencias Biologicas, P. Universidad Cato- 
lica de Chile, Casilla 114-D, Santiago, Chile; 
(JIC) Departamento de Oceanologia, Univ- 
ersidad de Concepcion, Casilla 2407, Con- 
cepcion, Chile. 


PROC. BIOL. SOC. WASH. 
106(4), 1993, pp. 673-677 


SCALISPINIGERA OCULATA HARTMAN, 1967 
(SCALIBREGMATIDAE: POLYCHAETA): SENIOR 
SYNONYM OF LACYDONIA ANTARCTICA 
(LACYDONIIDAE) HARTMANN-SCHRODER & 
ROSENFELDT, 1988 


Fredrik Pleijel and Kristian Fauchald 


Abstract. —Scalispinigera oculata Hartman, 1967, originally placed in the 
polychaete family Scalibregmatidae, is transferred to Lacydonia Marion & 
Bobretzky, 1875, in the family Lacydoniidae. Scalispinigera oculata is the type 
species of the genus, and Scalispinigera is consequently synonymized with 
Lacydonia. The second species described in the genus, S. cirrata Hartman & 
Fauchald, 1971, is likewise transferred to Lacydonia. Based on examination 
of the types Lacydonia antarctica Hartmann-Schroder & Rosenfeldt, 1988 is 
considered to be a junior synonym of Lacydonia oculata (Hartman, 1967), new 
combination. The relationship of L. oculata to L. mikrops Ehlers, 1913 is 


discussed. 


Hartman (1967) described the new genus 
and species Scalispinigera oculata from the 
Antarctic Peninsula, and, with a note that 
it did not conform to the conventional def- 
inition of the family, placed it in the family 
Scalibregmatidae. Examination of the ho- 
lotype of S. oculata indeed indicates that 
the species actually belongs to the genus La- 
cydonia Marion & Bobretzky, 1875, family 
Lacydoniidae Bergstrom, 1914. The posi- 
tion of S. oculata has been questioned ear- 
lier in studies by Kudenov & Blake (1978) 
and Blake (1981), in the former considered 
to be of uncertain position, in the latter 
questionably referred to the Hesionidae. 

The second species described in the ge- 
nus, S. cirrata Hartman & Fauchald, 1971, 
is likewise transferred to Lacydonia. 

Apart from L. oculata seven species are 
included in Lacydonia: 


L. miranda Marion & Bobretzky, 1875, from 
Mediterranean 
L. mikrops Ehlers, 1913, from Antarctic 


L. papillata Ushakov, 1958, from Kurile 
Trench, NW Pacific 

L. incognita Rullier, 1965, from West Af- 
rica 

L. cirrata Hartman & Fauchald, 1971, new 
combination, from U.S. east coast 

L. laureci Laubier, 1975, from Mediterra- 
nean 

L. antarctica Hartmann-Schroder & Rosen- 
feldt, 1988, from Antarctic 


Of these, L. antarctica is here considered 
a junior synonym of L. oculata. 

The redescription of Scalispinigera ocu- 
lata presented here is based completely on 
the holotype. Drawings were completed with 
a camera lucida. Institutions and museums 
are indicated by the following abbrevia- 
tions: CENTOB (Centre National de Tr 
d’Océanographique Biologique, Brest), 
USNM (National Museum of Natural His- 
tory, Smithsonian Institution, Washington, 
D.C.), ZMH (Universitat Hamburg, Zoolo- 
gisches Institut und Museum). 


674 


Family Lacydoniidae Bergstrom, 1914 
Genus Lacydonia Marion & Bobretzky, 
1875 


Lacydonia Marion & Bobretzky, 1875:57- 
61. 
Scalispinigera Hartman, 1967:134. 


Type species. —Lacydonia miranda Mar- 
ion & Bobretzky, 1875, by monotypy. 

Description. —Body-size small. Prosto- 
mium rounded, with four small, smooth 
frontal antennae. One pair of eyes or eyes 
lacking. Proboscis unknown. First visible 
segment achaetous, with one pair of small 
tentacular cirri. Parapodia and setae present 
from segment 2. Anteriormost parapodia 
uniramous, lacking dorsal setigerous lobes; 
following parapodia biramous with separate 
noto- and neuropodial setigerous lobes, 
aciculae and setae. All parapodia with small, 
inflated dorsal and ventral cirri. Notosetae 
capillaries, neurosetae composite spinigers. 
One pair of pygidial cirri. 


Lacydonia oculata (Hartman, 1967), 
new combination 
Fig. 1 


Scalispinigera oculata Hartman, 1967:134— 
135, pl. 41A-—C.—Kudenov & Blake, 
1978:428, 441.—Blake, 1981:1131, 1157. 
Lacydonia antarctica Hartmann-Schro- 
der & Rosenfeldt, 1988:36, figs. 11-13. 
New synonymy. 


Material examined. —Scalispinigera ocu- 
lata holotype (USNM 47326), Antarctic 
Peninsula, Anvers Island, Fort Lockroy, off 
Wiencke, 64°68'S, 63°30’W, shore, coll. W. 
Schmitt. Lacydonia antarctica holotype 
(ZMH 19098) and paratype (ZMH 19099), 
Antarctic Peninsula, King George Island, 
62°05.3’S, 57°39'W, 265 m, coll. U. Muh- 
lenhardt-Siegel; 1 specimen (ZMH P-20529), 
Antarctic, Elephant Island, 61°09.7'S, 
56°10.3'W, 290 m, coll. U. Mihlenhardt- 
Siegel, det. G. Hartmann-Schroder & P. Ro- 
senfeldt. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Description. —Holotype of S. oculata 
complete, 6 mm long and 0.8 mm wide (se- 
tiger 17, including parapodia but excluding 
setae) for ca. 45 setigers. Body dorso-ven- 
trally flattened, widest anteriorly, slowly ta- 
pering posteriorly. 

Prostomium rounded, slightly incised an- 
teriorly, about twice as wide as long (Fig. 
1A, B). Paired antennae small, papilliform, 
difficult to discern, ventral pair situated in 
small depressions (Fig. 1B). Pair of large 
eyes, anteriorly situated; lenses not ob- 
served. Small rounded median antenna sit- 
uated just posterior to eyes. Proboscis not 
observed. Segment 1 with one pair of small 
rounded tentacular cirri (Fig. 1A, B). Seg- 
ments 2—4 uniramous, without notopodial 
setigerous lobes, neuropodia with setigerous 
lobes bearing composite neurosetae. Para- 
podia following segment 4 biramous. Seg- 
ments biannulated with small intersegmen- 
tal areas dorsally and ventrally. Indistinct 
transverse ridges present medially on each 
segment (bands of cilia?). Notopodia from 
segment 2 with inflated dorsal cirrus, ovoid 
with pointed tips, inserted near notopodial 
base. Setigerous lobe conical, with single no- 
toacicula (Fig. 1C). Notosetae long, smooth 
capillaries. Neuropodium with conical se- 
tigerous lobe, single neuroacicula slightly 
stouter than notoacicula. Neurosetae com- 
posite spinigers, long and thin with single- 
toothed rostrum, blade serrated. Ventral 
cirrus similar to dorsal in size and shape, 
inserted subdistally on neuropodium. Py- 
gidium with two small inflated cirri and me- 
dian ventral papilla (Fig. 1D). 

Color: Eyes black. Body brownish yellow, 
darker brown pigment present laterally on 
each segment, extending both dorsally and 
ventrally. Most dorsal and ventral cirri with 
dark pigmented tips. 

Remarks. —WHartman’s interpretation of 
the characters of S. oculata was clearly in- 
fluenced by the fact that she considered the 
specimen to be a scalibregmatid. Contrary 
to the description above, she stated that bi- 
ramous parapodia were present from seg- 


VOLUME 106, NUMBER 4 675 


S RES 
SS = = - 
) aS = ae 
Ys 
S 
=e S} @, Li 
SS Boi/ZZ ee 
==o SSZE 
ZF Ze 
S| e= 4 
= — 
7 
Fig. 1. Lacydonia oculata, holotype (USNM 47326). A. Anterior end, dorsal view. B. Anterior end, ventral 


view. C. Left parapodium from ca, segment 20, posterior view. Only about half of full number of setae shown. 
D. Posterior end, ventral view. Setae omitted. Scale lines A-B 250 um, C 250 um, D 100 um. 


ment 2, that dorsal and ventral cirri were 
lacking, that the segments were triannulat- 
ed, and that the epithelium was weakly 
aerolated (probably segmental bands of cilia 
with adhering particles). 

Hartman’s species obviously is a lacy- 
doniid, and the type species, L. miranda, is 
illustrated here (Fig. 2) for comparison. As 
noted by Cantone (1973) the species is pro- 
vided with five antennae, rather than four; 
the median one was overlooked by Marion 
& Bobretzky (1875) in their original de- 
scription. While we see no obvious differ- 
ences between L. oculata and L. miranda, 
we would not synonymize the two species. 
We consider common Antarctic and Med- 
iterranean distributions of these species to 
be highly unlikely, and such a conclusion 
would have to be based on other material 
and methods. The intent was to place the 
species described by Hartman (1967) into 


the correct familial context, and a revision 
of the family Lacydoniidae is beyond the 
scope of this paper. 

Hartmann-Schroder & Rosenfeldt (1988), 
in their description of L. antarctica, only 
discussed two Lacydonia-species: L. mir- 
anda (which incorrectly was stated to have 
four rather than five antennae; observations 
based on examination of western Mediter- 
ranean specimens) and L. /aureci. Possibly 
they overlooked the existence of Ehlers’ spe- 
cies L. mikrops, which was also described 
from the Antarctic (Wilhelm II Land). Con- 
sidering Hartman’s mistake it is less sur- 
prising that Scalispinigera oculata was not 
discussed in their study, but comparing the 
type material of L. antarctica with that of 
S. oculata clearly shows them to be con- 
specific. Hartmann-Schroder & Rosen- 
feldt’s drawing (fig. 11) shows the median 
antenna as situated on a line between the 


676 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Lacydonia miranda, specimen from Italy, Sicily, Brucoli, 37°17'N, 15°11’E, 17 m (in FP’s collection). 
A. Anterior end, dorsal view. B. Anterior end, ventral view. C. Left parapodium from segment 10, posterior 
view. D. Posterior end, ventral view, setae omitted. All drawn to same scale; scale line 250 um. 


anterior rather than posterior sides of the 
eyes. This is due to the angle in which the 
specimen was drawn; examination of their 
material shows it to be situated more pos- 
terior. 

The original description and illustrations 
of S. cirrata show this species as well to 
agree with the generic definition of Lacy- 
donia, for which reason it is transferred to 
this genus. 

Of L. mikrops two syntypes (ZMH 
V-8549) and seven specimens from the 


Weddell Sea collected by EPOS 3 in 1989 
(CENTOB) were examined. Ehlers (1913) 
mentioned several specimens in his descrip- 
tion, but the one on which he based at least 
the major part of his description is unfor- 
tunately not one of the remaining syntypes. 
The two syntypes are in poor condition and 
probably the only remaining type material; 
for this reason we hesitantly allocate the 
Weddell specimens to L. mikrops. Never- 
theless, L. mikrops appears to differ from 
L. oculata in having much smaller eyes and 


VOLUME 106, NUMBER 4 


in having a prostomium as wide as long 
rather than much wider than long. 


Acknowledgments 


We wish to thank J. A. Blake, C. J. Glasby 
and M. H. Pettibone for valuable comments 
on the manuscript, G. Hartmann-Schroder 
(ZMH) and M. Segonzac (CENTOB) for loan 
of material, and P. Bouchet for admitting 
FP to the “Fifth Malacological Workshop” 
at Sicily, where part of the material was col- 
lected. Financial support for FP was pro- 
vided by the Swedish Natural Science Re- 
search Council (contracts 9555-306 and 
-307). 


Literature Cited 


Bergstrom, E. 1914. Zur Systematik der Polychae- 
tenfamilie der Phyllodociden.— Zoologiska Bi- 
drag fran Uppsala 3:37-224. 

Blake, J. A. 1981. The Scalibregmatidae (Annelida: 
Polychaeta) from South America and Antarctica 
collected chiefly during the cruises of the R/V 
Anton Brun, R/V Hero and USNS Eltanin. — 
Proceedings of the Biological Society of Wash- 
ington 94:1131-1162. 

Cantone, G. 1973. Una populazione di Lacydonia 
miranda Marion & Bobretzky (Annelida Poly- 
chaeta) con 5 antenne delle coste orientali della 
Sicilia.—Bollettino delle Sedute dell’Academia 
Gioenia di Scienze Naturali in Catania 12:237- 
240. 

Ehlers, E. 1913. Die Polychaeten-Sammlungen der 
deutschen Siidpolar-Expedition 1901-1903.— 
Deutsche Siidpolar-Expedition 13:397-598. 

Hartman, O. 1967. Polychaetous annelids collected 
by the USNS Eltanin and Staten Island cruises, 


677 


chiefly from Antarctic seas.—Allan Hancock 
Monographs in Marine Biology 2:1-387. 

Hartman, O., & K. Fauchald. 1971. Deep-water ben- 
thic polychaetes off New England to Bermuda 
and other North Atlantic areas.—Allan Han- 
cock Monographs in Marine Biology 6:1—327. 

Hartmann-Schroder, G., & P. Rosenfeldt. 1988. Die 
Polychaeten der “‘Polarstern’’-Reise ANT III/2 
in die Antarktis 1984. Teil 1: Euphrosinidae bis 
Chaetopteridae.— Mitteilungen aus dem Ham- 
burgischen zoologischen Museum und Institut 
85:25-72. 

Kudenov, J. D., & J. A. Blake. 1978. A review of the 
genera and species of the Scalibregmidae (Poly- 
chaeta) with descriptions of one new genus and 
three new species from Australia.—Journal of 
Natural History 12:427-444. 

Laubier, L. 1975. Lacydonia laureci sp. n., annélide 
polychéte nouvelle de 1’étage abyssal de Médi- 
terranée orientale.— Vie et Milieu 25:75-82. 

Marion, A. F., & N. Bobretzky. 1875. Etude des An- 
nélides du Golfe de Marseilles.—Annales des 
sciences naturelles 2:1-106. 

Rullier, F. 1965. Contribution a la faune des anné- 
lides polychétes du Dahomey et du Togo. —Ca- 
hiers océanographiques 3:5—66. 

Uschakov, P. V. 1958. Two new species of marine 
Polychaeta of the family Phyllodocidae from the 
abyssal depths of the Kuril-Kamchatka Trench 
[in Russian].—Trudy Instituta Okeanologii. 
Akademiya nauk SSSR 27:204—207. 


(FP) Swedish Museum of Natural His- 
tory, Box 50007, S-104 05 Stockholm, Swe- 
den, and (postal address): Tjarno Marine 
Biological Laboratory, Pl. 2781, S-452 96 
Stromstad, Sweden; (KF) Department of In- 
vertebrate Zoology, National Museum of 
Natural History, Smithsonian Institution, 
Washington, D.C. 20560, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(4), 1993, pp. 678-688 


POLY NOID POLYCHAETES ASSOCIATED WITH A 
WHALE SKELETON IN THE BATHYAL 
SANTA CATALINA BASIN 


Marian H. Pettibone 


Abstract. —A whale-fall community dominated by microbial mats of sulfur- 
oxidizing bacteria in the Santa Catalina Basin is inhabited by species similar 
to those associated with hydrothermal vents in the northeast Pacific. Polynoid 
polychaetes associated with the skeleton include: Bathykurila guaymasensis 
(Macellicephalinae); Peinaleopolynoe santacatalina, new species (Branchino- 
togluminae, emended); and Harmothoe craigsmithi, new species, and Subadyte 
mexicana (Harmothoinae). Both the holotype of Peinaleopolynoe sillardi Des- 
bruyéres & Laubier and additional specimens of Subadyte mexicana Fauchald, 
were examined and descriptions supplemented. 


Oceanographers aboard the deep sub- 
mergence research vessel (DSRV) A/vin dis- 
covered an intact, 20 m long skeleton of a 
blue or fin whale at a depth of 1240 m in 
the Santa Catalina Basin (33°14’N, 
118°30’W) during November, 1987. Sub- 
sequent collections revealed a characteristic 
“‘whale-fall” community associated with the 
skeleton (Smith et al. 1989, Allison et al. 
1991, Smith 1992). The skeleton was par- 
tially buried and the bone surfaces were cov- 
ered by a white “‘furry” microbial mat con- 
sisting of large filamentous sulfur-oxidizing 
bacteria (Beggiatoa sp.). The site was a sul- 
fide rich, reducing habitat similar to those 
associated with hydrothermal vents. In ad- 
dition, numerous invertebrate species, in- 
cluding large clams, mussels, limpets and 
snails, were associated with the bones in 
ways that were similar to those of hydro- 
thermal vent faunas of the Galapagos Rift, 
Guaymas Basin and Juan de Fuca Ridge. 
These species were unreported from the 
Santa Catalina Basin (Smith & Hamilton 
1983, Smith 1985, Kukert & Smith 1992). 
Additional Alvin dives to the whale-fall site 
were made in February 1991. 

Polynoid polychaetes collected on or 
within 0.5 m of the bones were sent to me 


for identification by Dr. Craig R. Smith, 
University of Hawaii, Manoa. Included are 
four species in three subfamilies: Bathyku- 
rila guaymasensis Pettibone, 1989 (Macel- 
licephalinae), originally described from the 
Guaymas Basin hydrothermal mounds; 
Peinaleopolynoe santacatalina, new species 
(Branchinotogluminae Pettibone, 1985a, 
emended); Harmothoe craigsmithi, new 
species, and Subadyte mexicana Fauchald, 
1972 (Harmothoninae). The subfamily 
Branchinotogluminae is emended to in- 
clude Peinaleopolynoe sillardi Desbruyéres 
& Laubier, 1988, obtained from artificially 
enriched substrates placed at a depth of 4800 
m in the northeast Atlantic off Spain. The 
original description of P. sillardi is supple- 
mented, based on an examination of the 
holotype kindly sent on loan from the Mu- 
séum National d’Histoire Naturelle, Paris 
(MNHNP). Harmothoe craigsmithi is sim- 
ilar to H. tenebricosa Moore, 1910, which 
was described from southern California in 
914-1463 m. Subadyte mexicana was orig- 
inally described from western Mexico in 
567-844 m. Its description is also supple- 
mented based on additional specimens from 
the Santa Catalina Basin and Channel Is- 
lands. 


VOLUME 106, NUMBER 4 


The larger polynoids, collected by the A/- 
vin, were part of the epifauna on the whale 
bones and designated by Bone Implant 
number (1 or 2) or Vertebra (V) number. 
The smaller polynoids, part of the infauna, 
were collected by Ekman cores (E), and sep- 
arated into subcores (A—D) and from 0-1, 
1-5, and 5-10 cm below the sediment sur- 
face. 

Types and additional specimens are de- 
posited in the collections of the Department 
of Invertebrate Zoology, National Museum 
of Natural History, Smithsonian Institution 
(USNM), the Natural History Museum of 
Los Angeles County (AHF-LACM), and 
with the donor of the collection, Dr. Craig 
R. Smith (CRS), of the Department of 
Oceanography, University of Hawaii at 
Manoa. 


Family Polynoidae 
Subfamily Macellicephalinae 
Hartmann-Schroder, 1971, 
emended Pettibone, 1976 
Genus Bathykurila Pettibone, 1976 
Bathykurila guaymasensis Pettibone, 1989 


Bathykurila guaymasensis Pettibone, 1989: 
159, figs. 1, 2. 


Material.—Santa Catalina Basin, Cali- 
fornia, 33°12’N, 118°30’W, 1240 m, Alvin 
Dives in Whale-fall site, in Feb 1991, buck- 
et washes: AD 2332, 20 Feb, Implant 1, 1 
specimen (USNM 157592), Implant 2, 4 
specimens (USNM 157593), 1 specimen 
(CRS); AD 2334, 22 Feb, Bone V21, 1 spec- 
imen (USNM 157594); AD 2336, 24 Feb, 
Bone V15, 5 specimens (USNM 157595). 

Description.—The specimens agree with 
those previously described from the vents 
in the Guaymas Basin in 2004—2020 m. The 
largest specimen measures 10 mm long, 8 
mm wide, with setae, with 15 segments, and 
7 pairs of elytrophores; smaller specimens 
are 1.5—-4 mm long, 1.5-—3 mm wide, with 
10-13 segments. Long ventral papillae on 
segment 11 (Fig. 1C, in Pettibone 1989) are 
present on 4 of the 12 specimens; 3 speci- 


679 


mens have small ventral papillae on seg- 
ments 10, 11, 12 (not reported previously). 

Remarks. —The specimens have numer- 
ous filamentous sulfur-bacteria attached 
(Beggiatoa sp.), characterized by their glid- 
ing motility and internal globular elemental 
sulfur, described and figured by Nelson et 
al. (1989). 

In their report on the fauna of the Santa 
Catalina Basin, Smith & Hamilton (1983: 
916) stated: “‘an undescribed macellicepha- 
lin polychaete made frequent excursions 
high above the sediment.”’ The specimen, 
collected by Dr. Kenneth L. Smith of the 
Scripps Institution of Oceanography, was 
sent to me for identification and described 
as anew macellicephalan species, Natopoly- 
noe kensmithi Pettibone (1985b:747). 


Subfamily Branchinotogluminae 
Pettibone, 1985a, emended 


The subfamily is emended to include 
Peinaleopolynoe sillardi Desbruyéres & 
Laubier, 1988 and P. santacatalina, new 
species. Instead of 10 pairs of elytra and 
elytrophores on segments 2, 4, 5, 7, 9, 11, 
13, 15, 17, 19, and dorsal cirri on the non- 
elytrigerous segments, including segments 
20 and 21, there may be 9 pairs of elytra, 
with the elytra and elytrophores lacking on 
segment 19, but also lacking dorsal cirri and 
thus not a typical cirrigerous segment, as in 
P. sillardi or 10 pairs of elytra with the ely- 
trophores and elytra extra small on segment 
19, as in P. santacatalina. Notopodial bracts 
are not present on some or all of the elytri- 
gerous segments, as in other members of the 
subfamily. Also the arborescent branchiae 
may begin on segment 2 and not on segment 
3, as indicated previously. 


Genus Peinaleopolynoe 
Desbruyéres & Laubier, 1988, emended 


Type species.—Peinaleopolynoe sillardi 
Desbruyéres & Laubier, 1988, by mono- 
typy. Gender feminine. 


680 


The genus includes the type species, P. 
sillardi, and a new species, P. santacatalina. 

Diagnosis. —Body short, with 21 seg- 
ments. Elytra and elytrophores numbering 
9 or 10 pairs, on segments 2, 4, 5, 7, 9, 11, 
13, 15, 17, and 19 or lacking on 19. Elytra 
large, subreniform, overlapping, covering 
dorsum, without tubercles or papillae. Dor- 
sal cirri with short cylindrical cirrophores 
and long distal styles; dorsal tubercles, in 
line with elytrophores, on non-elytrigerous 
segments. Arborescent branchiae attached 
on dorsal tubercles and bases of notopodia, 
beginning on segment 2 and continuing to 
near end of body. Prostomium bilobed, with 
triangular anterior lobes bearing minute 
frontal filaments, with median antenna in 
anterior notch and paired palps; without lat- 
eral antennae or eyes. First or tentacular 
segment not visible dorsally; tentaculo- 
phores lateral to prostomium, each with 
small acicular lobe on inner side, dorsal and 
ventral tentacular cirri, without setae. Sec- 
ond or buccal segment with first pair of elyt- 
rophores, biramous parapodia, and ventral 
or buccal cirri attached to basal parts of 
neuropodia, lateral to ventral mouth; styles 
longer than following ventral cirri. Para- 
podia biramous, with notopodia almost as 
long as neuropodia. Notopodia without no- 
topodial bracts, bulbous basally, extending 
distally into acicular processes; neuropodia 
with longer conical presetal lobes with acic- 
ular processes and shorter truncate postsetal 
lobes. Notosetae stouter than neurosetae, 
Straight, acicular, smooth or with lateral 
spines. Neurosetae long, slender, finely spi- 
nous, with slightly hooked tips. Pharynx with 
7 pairs of border papillae; jaws with lateral 
teeth. Ventral segmental papillae on seg- 
ments 12-15. Pygidium with pair of anal 
Cirri. 


Peinaleopolynoe sillardi 
Desbruyéres & Laubier 
Fig. 1 


Peinaleopolynoe sillardi Desbruyéres & 
Laubier, 1988:331, figs. 1, 2. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Material. —Northeast Atlantic Ocean, 
46°02’N, 16°40’W, 4800 m, from enriched 
module of colonization, 23 Jul 1985, ho- 
lotype (MNHNP UB631). 

Supplementary description.—The elytra 
and elytrophores are lacking on segment 19, 
thus there are 9 pairs of elytra, as originally 
indicated. However, segment 19 also lacks 
dorsal cirri, thus suggesting that it has sec- 
ondarily lost the characteristic structure. The 
large elytra are subreniform, thick, white, 
Opaque, without tubercles or papillae, ex- 
cept for some small posterior extensions, 
variable in size, shape, and number (Fig. 
1A). The notopodia and neuropodia of the 
biramous parapodia are subequal in size, 
both with projecting acicular processes; the 
notosetae are short to longer, not as long as 
the neurosetae (Fig. 1C). The stout acicular 
notosetae, without spines, taper to rounded 
tips (Fig. 1D). The slender neurosetae have 
slightly hooked bare tips, with double rows 
of spines along one border; the upper neu- 
rosetae are more slender than the middle 
ones (Fig. 1E). Segmental ventral papillae 
are present on segments 12-15; they are 
short and curved laterally (Fig. 1B). 


Peinaleopolynoe santacatalina, 
new species 
Fig. 2 


Material.—Santa Catalina Basin, Cali- 
fornia, 33°14’N, 118°30’W, 1240 m, Alvin 
Dives in Whale-fall site, in Feb 1991, buck- 
et washes: AD 2332, 20 Feb, Implant 1, 
paratype (USNM 157588), Implant 2, 2 
paratypes (USNM 157589); paratype (CRS); 
(AD 2334, 22 Feb, Bone V21, holotype 
(USNM 157587). 

Description. —Holotype (USNM 157587) 
22 mm long, 14 mm wide with setae, 21 
segments; paratype (USNM 157588) 20 mm 
long, 13 mm wide, 21 segments; 3 smaller 
paratypes (USNM 157589) 9-12 mm long, 
7-10 mm wide, 21 segments, last 2 very 
small. Dorsum with ciliated transverse 
bands, 2 per segment, extending onto bases 
of elytrophores and dorsal tubercles (Fig. 


VOLUME 106, NUMBER 4 


681 


D 


Fig. 1. 


a 
pA 
4 


= 


Peinaleopolynoe sillardi, holotype (MNHNP UB631): A, Right 3rd elytron from segment 5, with 


detail of posterior extensions; B, Ventral view of central part of left side of segments 12-15, showing ventral 
segmental papillae; C, Right elytrigerous parapodium from segment 5, anterior view; D, Tips of long and short 
notosetae from same; E, Upper and middle neurosetae from same, with detail of parts. Scales = 2.0 mm for A; 


2.0 mm for B; 1.0 mm for C; 0.1 mm for D, E. 


2A, B). Elytrophores large, bulbous, 10 pairs, 
oaseements 2.4.5, 7,9, 11, 13, 15, 17,19, 
smaller on segment 19 (Fig. 2A, B, F); elytra 
all missing, except for minute elytron on left 
19th elytrophore of one paratype. Dorsal 
tubercles and dorsal cirri on non-elytriger- 
ous segments; cirrophores rather long, cy- 
lindrical, on posterior sides of notopodia; 
styles long, filiform, extending beyond se- 
tae, shorter on segment 21 (Fig. 2B, G). 
Branchiae compact, arborescent, with nu- 
merous short bulbous terminal filaments, 
beginning on segment 2 and continuing to 
near posterior end, small on segment 20, 


lacking on segment 21; on elytrigerous seg- 
ments, branchiae forming single large groups 
between elytrophores and bases of noto- 
podia; on cirrigerous segments, branchiae 
in small groups attached to dorsal tubercles 
and larger groups near bases of notopodia 
(Fig. ZA, B, F,.G). 

Prostomium oval, deeply bilobed, form- 
ing triangular anterior lobes with delicate 
frontal filaments; median antenna with bul- 
bous ceratophore in anterior notch, style 
short, only slightly surpassing prostomium; 
without eyes; palps stout, tapering, about 
twice length of prostomium; tentaculo- 


682 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


J 


Fig. 2. Peinaleopolynoe santacatalina, A-C, F-I, holotype (USNM 157587); D, E, paratype (USNM 157589): 
A, Dorsal view of anterior end, left dorsal tentacular cirrus missing; B, Dorsal view of posterior end, including 
segments 17-21, styles of dorsal cirri of segment 20 and anal cirri missing; C, Ventral view of left side of segments 
12-15, showing ventral segmental papillae; D, Dorsal border papillae of pharynx; E, Jaw; F, Right elytrigerous 
parapodium from segment 9, anterior view, acicula dotted; G, Right cirrigerous parapodium from segment 10, 
posterior view; H, Long notoseta from same; I, Supraacicular neuroseta from same, with detail of parts; J, 
Subacicular neuroseta from same, with detail of parts. Scales = 1.0 mm for A, B; 2.0 mm for C; 0.2 mm for 
D; Evl-Ognm fork, G2 0.1,mm for H: I, 


VOLUME 106, NUMBER 4 


phores of segment | lateral to prostomium 
and palps, achaetous, each with small acic- 
ular process on medial side, long dorsal ten- 
tacular cirrus about as long as palp and 
slightly shorter ventral tentacular cirrus, and 
forming anterior and lateral lips of ventral 
mouth (Fig. 2A). Segment 2 or buccal seg- 
ment with first pair of large elytrophores, 
branchiae, biramous parapodia, and ventral 
buccal cirri inserted basally and extending 
beyond tips of neuropodia, with contribu- 
tions to posterior lip of ventral mouth (Fig. 
2A). Pharynx (dissected) with 7 pairs of dor- 
sal and ventral border papillae; hooked jaws 
with small and larger teeth on inner borders 
(Fig. 2D, E). 

Notopodia of biramous parapodia bul- 
bous basally, extending into long acicular 
processes on lower sides; neuropodia with 
subtriangular presetal lobes extending into 
long acicular processes, postsetal lobes 
shorter, truncate (Fig. 2F, G). Notosetae 
forming radiating bundles, short to long, al- 
most as long as neurosetae, stout, acicular, 
tapering to blunt tips, with double alter- 
nating rows of short spines on one side, 3- 
4 pairs on shorter notosetae and up to 11 
pairs on longer ones (Fig. 2F—H). Neuro- 
setae forming fan-shaped bundles, very nu- 
merous, slender, with slightly curved tips 
and double rows of spines; supraacicular 
neurosetae with more prominent spines than 
in subacicular neurosetae (Fig. 2F, G, I, J). 
Ventral cirri with small cirrophores on mid- 
dle of neuropodia, styles short, tapering, of- 
ten curved distally on posterior sides of neu- 
ropodia (Fig. 2F, G). Ventral segmental 
papillae 4 pairs, on segments 12-15, rather 
long, curved laterally (Fig. 2C); ventral seg- 
mental papillae small, rounded, on smaller 
paratype. Pygidium enclosed in parapodia 
of segments 19-21, last 2 smaller, with pair 
of anal cirri (sometimes missing, Fig. 2B). 

Etymology.—The new species is named 
for the collecting site, Santa Catalina Basin. 

Remarks.—Peinaleopolynoe santacatali- 
na differs from P. sillardi in having small 
elytrophores and elytra on segment 19, 


683 


making 10 pairs of elytra, instead of lacking 
elytrophores, elytra and dorsal cirri, and 9 
pairs of elytra. The stout acicular notosetae 
have double rows of spines on one side in 
P. santacatalina and are smooth in P. sil- 
lardi. 


Subfamily Harmothoinae Willey, 1902 
Genus Harmothoe Kinberg, 1856 
Harmothoe craigsmithi, new species 
Fig. 3 


Material.—Santa Catalina Basin, Cali- 
fornia, 33°12'N, 118°30’W, 1240 m, Alvin 
Dives in Whale-fall site, in Nov 1988: AD 
2138, 11 Nov, E2 A 0-1, holotype (USNM 
157590), E6 B 0-10, paratype (CRS); AD 
2133, 6 Nov, WS-6, bone scrapings, para- 
type (USNM 157591). 

Description.— Holotype 22 mm long, 8 
mm wide with setae, 37 segments. Com- 
plete paratype (USNM 157591) 24 mm long, 
8 mm wide, 37 segments. Dorsum darkly 
pigmented, with low ciliated transverse 
bands, 2 per segment, continuing on bases 
of elytrophores and dorsal tubercles. Elytra 
15 pairs, on bulbous elytrophores, on seg- 
ments 2, 4, 5, 7, alternate segments to 23, 
26, 29, 32. First elytra round, with long pa- 
pillae on border and scattered on surface, 
with conical microtubercles throughout but 
more concentrated near borders (Fig. 3B). 
Following elytra subreniform, with long pa- 
pillae on lateral borders, surfaces with con- 
ical microtubercles and long papillae, most- 
ly confined to lateral halves; medial halves 
bare or with scattered small microtubercles 
(Fig. 3C). Prominent bulbous dorsal tuber- 
cles and dorsal cirri on non-elytrigerous seg- 
ments; cirrophores short, bulbous, on pos- 
terior sides of notopodia; styles slender, long, 
extending far beyond setae, with long pa- 
pillae (Fig. 3E). 

Prostomium oval, bilobed, wider than 
long, with small anterior peaks; 2 pairs of 
rather large eyes, anterior pair anterior to 
widest part of prostomium, posterior pair 
near posterior border; median antenna with 


684 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Yea 
X u 
Jeb tiyet 
quia 


hat 


Fig. 3. Harmothoe craigsmithi, holotype (USNM 157590): A, Dorsal view of anterior end, styles of median 


antenna and tentacular cirri missing; B, Right Ist elytron; C, Right middle elytron, with detail of microtubercles 


and papillae; D, Left elytrigerous parapodium, anterior view, acicula dotted; E, Left middle cirrigerous para- 
podium, posterior view; F, Long and short notosetae from same; G, Upper, middle and lower neuroseta from 


same, with detail of parts. Scales = 0.1 mm for A; 0.5 mm for B, C; 0.5 mm for D, E; 0.1 mm for F, G. 


bulbous ceratophore in anterior notch, style 
missing; lateral antennae with ceratophores 
inserted ventrally, styles short, subulate, pa- 
pillate; palps stout, tapering about twice 
length of prostomium; tentaculophores of 
segment I lateral to prostomium, each with 


small acicular lobe and 3 notosetae on inner 

side and dorsal and ventral tentacular cirri 
(missing) (Fig. 3A). Second segment with 
first pair of large elytrophores, biramous 
parapodia and long ventral buccal cirri lat- 
eral to ventral mouth (Fig. 3A). 


VOLUME 106, NUMBER 4 


Notopodium of biramous parapodium 
almost as long as neuropodium, rounded, 
with long acicular process on lower side; 
neuropodium with subconical presetal 
acicular lobe with digitiform supraacicular 
process, postsetal lobe shorter, rounded (Fig. 
3D, E). Notosetae numerous, forming ra- 
diating bundle, stouter than neurosetae, 
shorter, slightly curved to longer, nearly 
straight, with numerous spinous rows and 
short, bare tips; longer ones mostly faint 
split tips (Fig. 3F). Neurosetae numerous, 
forming fan-shaped bundle, with numerous 
spinous rows and bare, slightly hooked tips; 
middle and upper ones with slender sec- 
ondary tooth and more prominent spinous 
rows; lower ones with shorter spinous 
regions and entire tips (Fig. 3G). Ventral 
cirri with cirrophores on middle of neuro- 
podia; styles short, tapered (Fig. 3D, E). 

Etymology. — The species is named for Dr. 
Craig R. Smith, the collector of the poly- 
noids from the whale-fall site, sent for iden- 
tification. 

Remarks.—Harmothoe craigsmithi is 
close to H. tenebricosa Moore (1910:351- 
353), which was described from off Cali- 
fornia in 914-1463 m (also see Pettibone 
1969b:3 1-42) and is widely distributed from 
Japan and the Bering Sea to Lower Califor- 
nia, in 203-1990 m. Harmothoe craigsmithi 
differs from H. tenebricosa in the following: 
the elytra have marginal and surface papil- 
lae and microtubercles, instead of lacking 
them; the eyes are rather large, instead of 
small or absent; the notopodia are almost 
as long as the neuropodia, instead of short- 
er; the neuropodial presetal acicular lobe 
has a long digitiform supraacicular process, 
instead of a small rounded process; the no- 
tosetae have distinct spinous rows, instead 
of nearly smooth or faint spinous rows. 


Genus Subadyte Pettibone, 1969a 
Subadyte mexicana Fauchald, 1972 
Figs. 4, 5 


Subadyte mexicana Fauchald, 1972:27, pl. 
1:figs. a—e. 


685 


Subadyte sp. A. Jones & Thompson, 1987: 
128, fig. 3a (list). 


Material.—Baja California, vicinity of 
Cedros Island, 27°38'N, 115°16’W, 792-844 
m, mud and glauconitic sand, holotype 
(LACM-AHF 1008). 

Santa Catalina Basin, California, 33°12’N, 
118°30'W, 1240 m, Alvin Dives in Whale- 
bone habitat, Nov 1988: 0 m distance from 
whale bones: AD 2133, 6 Nov, E2 A 0-2, 
1 specimen (USNM 157601); AD 2138, 11 
Nov, E6 A 0-10, E6 C 0-10, E6 D 0-10; E7 
D-0-5, 4 specimens (USNM 157602-5). 0.5 
m distance from whale bones: AD 2133, 6 
Nov, Ell B O-1, 1 specimen (USNM 
157600); AD 2135, 8 Nov, E10 C 0-1, 1 
specimen (USNM 157596); AD 2137, 10 
Nov, E6 D 0-1, E9 B 5-10, E9 C 5-10, 3 
specimens (USNM 157597-9); AD 2138, 
11 Nov, El A 0-1, E2 C 0-1, 2 specimens 
(CES). 

Channel Islands, California, R/V Velero 
IV, (as Subadyte sp. A): AHF 22970, Santa 
Rosa Island, 33°51'N, 120°08’W, 368 m, 1 
specimen (LACM): AHF 23000, Santa Rosa 
Island, 33°48'N, 120°04’W, 127 m, 1 spec- 
imen (LACM); AHF 23093, Santa Rosa Is- 
land, 33°39’N, 119°58'W, 113 m, 1 speci- 
men (USNM 157610); AHF 23182, San 
Miguel Island, 33°57’N, 120°22’W, 118 m, 
2 specimens (USNM 157611); AHF 24241, 
San Miguel Island, 33°57'N, 120°23'W, 139 
m, 2 specimens (LACM). 

Southern California, R/V Thomas G. 
Thompson Cruise 113, 1977 (as Subadyte 
sp. A): AHF 82801, off Huntington Beach, 
33°23’N, 117°54'W, 536-543 m, 2 speci- 
mens (LACM); AHF 80201, off Santa Bar- 
bara, 34°22'’N, 119°57'W, 329-340 m, 1 
specimen (LACM); AHF 80546, San Mi- 
guel Island, 33°57’N, 120°26’W, 213-251 
m, 1 specimen (LACM); AHF 81010, Santa 
Cruz Island, 33°46’N, 119°49’W, 444-500 
m, 3 specimens (LACM); AHF 81735, Tan- 
ner Bank, 32°47'N, 119°15'W, 511-530 m, 
1 specimen (LACM). 

Type material. —The holotype consists of 
an anterior fragment 4, 5 mm long, 2 mm 


686 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Subadyte mexicana, A, B (USNM 157601), C (USNM 157603), D-F (USNM 157597): A, Dorsal 
view of anterior end, left tentacular cirri and left dorsal cirrus of segment II missing; B, Ventral view of lateral 
antennae and facial tubercle (not to scale); C, Left Ist elytron from segment II; D, Right middle elytrigerous 
parapodium, anterior view, acicula dotted; E, Short and longer notosetae, from same, with detail of parts; F, 
Neuroseta, from same, with detail of parts. Scales = 0.2 mm for A, C; 0.1 mm for D-F. 


wide without setae, and 12 segments. Most 
of the parapodia are broken off, with elytra 
missing. 

Supplementary description. — All 12 spec- 
imens from Whale-bone site incomplete, 
some with developing posterior ends; fig- 
ured specimen (USNM 157601) 2.5 mm 
long, 3 mm wide with setae, and 9 segments; 
largest specimen with 16 segments, plus 
posterior end of 3 developing segments 
(USNM 157597), 2.5 mm long, 2 mm wide 
with setae. Of 15 specimens from Channel 
Islands, largest specimen incomplete 
(USNM 157610), 8.5 mm long, 4 mm wide 
with setae and 25 segments. 

Body flattened, with long parapodia and 
setae, giving aspect of pelagic form (Fig. 4A, 
D). Elytrophores large, bulbous, on seg- 
ments 2, 4, 5, 7, continuing on alternate 
segments to 23, or more? Elytra mostly 
missing; remaining left elytron on specimen 


from Santa Catalina Basin, oval, with long 
papillae on surface (Fig. 4C), covered with 
foreign material, including filamentous bac- 
teria (Beggiatoa sp.). Elytra remaining on 
specimens from Channel Islands more nu- 
merous: large, delicate, oval, with papillae 
scattered on surfaces and near borders, vari- 
able in size and shape, some short with cla- 
vate tips and some longer, bulbous basally, 
with clavate tips (Fig. SA, B). Cirrigerous 
segments with inconspicuous dorsal tuber- 
cles; cirrophores of dorsal cirri short, cylin- 
drical, on posterior sides of notopodia, with 
styles long, extending beyond setae, papil- 
late, with filamentous tips (Fig. 4A). 
Prostomium (Fig. 4A, B) oval, deeply bi- 
lobed, wider than long, with small anterior 
peaks; 2 pairs of small eyes on posterior half 
of prostomium; median antenna with bul- 
bous ceratophore in anterior notch of pro- 
stomium, style long, papillate, with long fil- 


VOLUME 106, NUMBER 4 


687 


Fig. 5. 


Subadyte mexicana, specimen from San Miguel Island (USNM 157611): A, Left Ist elytron from 


segment 2, with detail of border and surface papillae; B, Left 3rd elytron from segment 6, with detail of border 


and surface papillae. Scale = 0.1 mm. 


amentous tip; lateral antennae with bulbous 
ceratophores, inserted ventrally and nearly 
hidden from view dorsally, styles short, su- 
bulate, papillate. 

Tentaculophores of segment I (Fig. 4A) 
lateral to prostomium, projecting anterior- 
ly, each with small projecting acicular lobe 
and 2 curved notosetae on inner side, long 
dorsal tentacular cirrus, similar to median 
antenna, and shorter ventral tentacular cir- 
rus; small rounded facial tubercle (Fig. 4B) 
between bases of ceratophores of lateral an- 
tennae. Segment II (Fig. 4A) with first pair 
of large elytrophores, biramous parapodia, 
and long ventral buccal cirri, attached ba- 
sally lateral to ventral mouth, similar to 
ventral tentacular cirri. 

Biramous parapodium (Fig. 4D) with no- 
topodium shorter than neuropodium, bul- 
bous basally, with tapering acicular process 
on lower side; neuropodium subconical, 
presetal lobe tapering to pointed acicular 
process, postsetal lobe shorter, rounded. 
Notosetae (Fig. 4D, E) numerous, forming 
radiating bundle, stouter than neurosetae, 
shorter, curved to longer, nearly straight, 
and nearly as long as neurosetae; shorter 
notosetae with up to 15 spinous pockets on 


curved borders and entire tips; longer no- 
tosetae with spinous pockets basally and 
more distally with smaller close-set spines 
and bifid split tips. Neurosetae (Fig. 4D, F) 
numerous, very long, forming fan-shaped 
bundles; neurosetae slender, with basal spi- 
nous pockets or spurs, finely spinous dis- 
tally, tapering to slender bifid split tips. 
Ventral cirri (Fig. 4D) on middle of neu- 
ropodia, short, subulate, smooth. 


Acknowledgments 


My thanks go to Dr. Craig R. Smith and 
Hilary Maybaum of the Department of 
Oceanography, University of Hawaii, at 
Manoa for the polynoid specimens from the 
Whale-fall site in the Santa Catalina Basin 
and for the information packet on the 
Whale-fall research and for being a part of 
this interesting study. Dr. Kenneth L. Smith 
of Scripps Institution of Oceanography fur- 
nished information on the pelagic macelli- 
cephalin polychaete from the Santa Catalina 
Basin. I thank Dr. Jean-Claude Dauvin of 
the Muséum National d’Histoire Naturelle, 
Paris for the loan of the holotype of Pein- 
aleopolynoe sillardi. 1 thank Leslie Harris 


688 


of the Natural History Museum of Los An- 
geles County (AHF-LACM) for the loans of 
the type of Subadyte mexicana and addi- 
tional specimens of Subadyte sp. from the 
Channel Islands. The manuscript benefited 
from the careful reviews of J. A. Blake, C. 
R. Smith, H. Maybaum, and an unnamed 
reviewer. The polychaetes from the Whale- 
fall site were collected under the support of 
NSF grant OCE-90-00162 to Craig R. Smith. 


Literature Cited 


Allison, P. A., C. R. Smith, H. Kukert, J. W. Deming, 
& B. A. Bennett. 1991. Deep-water taphono- 
my of vertebrate carcasses: a whale skeleton in 
the bathyal Santa Catalina Basin.—Paleobiol- 
ogy 17:78-89. 

Desbruyéres, D., & L. Laubier. 1988. Exploitation 
d’une source de matiére organique concentrée 
dans l’océan profond: intervention d’une an- 
nélide polychéte nouvelle.—Comptes Rendus 
de l’Academie des Sciences 307, Serie III (6): 
329-335. 

Fauchald, K. 1972. Benthic polychaetous annelids 
from deep water off Western Mexico and ad- 
jacent areas in the Eastern Pacific Ocean. —Al- 
lan Hancock Monographs in Marine Biology 7:1— 
5) //5¥ 

Hartmann-Schroder, G. 1971. Annelida, Borsten- 
wurmer, Polychaeta.—Die Tierwelt Deutsch- 
lands und der angrenzenden Meeresteile 58:1-— 
549. 

Jones, G. F., & B. E. Thompson. 1987. The distri- 
bution and abundance of Chloeia pinnata Moore, 
1911 (Polychaeta: Amphinomidae) on the 
Southern California borderland.—Pacific Sci- 
ence 41:122-131. 

Kinberg, J.G. H. 1856 (1855). Nye slagten och arter 
af Annelider. —Ofversigt af Kongl. Vetenskaps- 
Akademiens Forhandlingar 12:381-388. 

Kukert, H., & C. R. Smith. 1992. Disturbance, col- 
onization and succession in a deep-sea sediment 
community: artificial-mound experiments. — 
Deep-Sea Research 39(7/8):1349-1371. 

Moore, J. P. 1910. The polychaetous annelids dredged 
by the U.S:S. “Albatross” off the coast of south- 
ern California in 1904: Polynoidae, Aphroditi- 
dae and Segalionidae.— Proceedings of the 
Academy of Natural Sciences of Philadelphia 
62:328-402. 

Nelson, D. C., C. O. Wirsen, & H. W. Jannasch. 1989. 
Characterization of large, autotrophic Beggiatoa 
spp. abundant at hydrothermal vents of the 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Guaymas Basin.— Applied and Environmental 
Microbiology 55:2909-2917. 

Pettibone, M. H. 1969a. Review of some species re- 
ferred to Scalisetosus McIntosh (Polychaeta, 
Polynoidae).— Proceedings of the Biological So- 
ciety of Washington 82:1-30. 

1969b. Remarks on the North Pacific Har- 
mothoe tenebricosa Moore (Polychaeta, Poly- 
noidae) and its association with asteroids (Echi- 
nodermata, Asteroidea).—Proceedings of the 
Biological Society of Washington 82:31—42. 

1976. Revision of the genus Macellicephala 
McIntosh and the subfamily Macellicephalinae 
Hartmann-Schroéder (Polychaeta: Polynoi- 
dae).—Smithsonian Contributions to Zoology 
229:1-71. 

1985a. Additional branchiate scale-worms 
(Polychaeta: Polynoidae) from Galapagos Hy- 
drothermal Vent and Rift-Area off Western 
Mexico at 21°N.— Proceedings of the Biological 
Society of Washington 98:447-469. 

1985b. New genera and species of deep-sea 
Macellicephalinae and Harmothoinae (Poly- 
chaeta: Polynoidae) from the Hydrothermal Rift 
areas off the Galapagos and Western Mexico at 
21°N and from the Santa Catalina Channel. — 
Proceedings of the Biological Society of Wash- 
ington 98:740-757. 

1989. Polynoidae and Sigalionidae (Poly- 
chaeta) from the Guaymas Basin, with descrip- 
tions of two new species, and additional records 
from hydrothermal vents of the Galapagos Rift, 
21°N, and seep-sites in the Gulf of Mexico (Flor- 
ida and Louisiana).— Proceedings of the Bio- 
logical Society of Washington 103:154—-168. 
Smith, C. R. 1985. Food for the deep sea: utilization, 
dispersal, and flux of nekton falls at the Santa 
Catalina Basin floor.—Deep-Sea Research 32: 
417-442. 

1992. Whale Falls chemosynthesis on the 
deep seafloor.— Oceanus 35(3):74—78. 

—, & S.C. Hamilton. 1983. Epibenthic mega- 
fauna of a bathyal basin off Southern California: 
patterns of abundance, biomass, and disper- 
sion. — Deep-Sea Research 30:907-928. 

— ., H. Kukert, R. A. Wheatcroft, P. A. Jumars, & 
J. W. Deming. 1989. Vent fauna on whale 
remains. — Nature 341:27-28. 

Willey, A. 1902. Report on the collections of natural 
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voyage of the Southern Cross. London. XII. 
Polychaeta, pp. 262-283. 


Department of Invertebrate Zoology, Na- 
tional Museum of Natural History, Wash- 
ington, D.C. 20560, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(4), 1993, pp. 689-697 


PROBOPYRUS PACIFICENSIS, A NEW PARASITE SPECIES 
(ISOPODA: BOPYRIDAE) OF 
MACROBRACHIUM TENELLUM (SMITH, 1871) 
(DECAPODA: PALAEMONIDABE) OF THE 
PACIFIC COAST OF MEXICO 


Ramiro Roman-Contreras 


Abstract. —Probopyrus pacificensis, a new species of Bopyridae parasitizing 
the freshwater prawn Macrobrachium tenellum (Smith, 1871) is described from 
the Pacific coast of Mexico. This is the first species of Probopyrus described 
from the Eastern Pacific although P. pandalicola (Packard, 1879) has been 
reported earlier from this slope of the Americas. P. pandalicola, P. palaemo- 
neticola (Packard, 1881), and P. floridensis Richardson, 1904 are differentiated 
from P. pacificensis because they are smaller, by their dorsal pigmentation, 
morphological features and because they parasitize Palaemonetes spp. P. bi- 
thynis Richardson, 1904 exhibits anterolateral projections in its head resem- 
bling horns; this feature makes it different from other species in the genus. P. 
pacificensis is distinguished from P. panamensis Richardson, 1912 by its lam- 
inar and separated edges in the pereomeres 5-7 on the larger side, and a high 
and rounded carina on the seventh leg; P. panamensis has continuous pereo- 
meres and a lower carina. The final hosts of P. bithynis are M. ohione (Smith, 
1874) and M. olfersi(Wiegmann, 1836), while M. acanthurus (Wiegmann, 1836) 
and M. tenellum are parasitized by P. panamensis and P. pacificensis, respec- 


tively. 


The genus Probopyrus Giard & Bonnier 
(1888) comprises a complex of species dis- 
tributed along southeast Asia and the Amer- 
icas. It has been a controversial genus be- 
cause of the difficulty of separating the 
species of the group on the adult morphol- 
ogy. 

Giard & Bonnier (1888) proposed the bo- 
pyrid genus Probopyrus to accommodate 
some western Pacific species. The following 
species have been described from the west- 
ern Atlantic: P. pandalicola, P. palaemo- 
neticola, P. bithynis, P. floridensis, and P. 
panamensis. 

In 1905, Richardson proposed a key to 
identify the American species of the genus 
Probopyrus, but Markham (1985a) stated 
that the key was not suitable because of the 


difficulty distinguishing the adults on the 
basis of morphological features. 

Probopyrus pandalicola, P. floridensis, and 
P. bithynis have been recognized as valid 
species by Dale & Anderson (1982) based 
on observations of behavior, pigmentation 
patterns, and larval morphology; these spe- 
cies have been accepted by Jimenez & Var- 
gas (1990). 

Markham (1985a) pointed out that most 
of the other species of Probopyrus in the 
western Atlantic are synonyms of P. pan- 
dalicola and that it is “‘the only known bran- 
chial parasite of Macrobrachium, Palae- 
mon, and Palaemonetes in the northwestern 
Atlantic, where it infests at least ten differ- 
ent host species.” 

Despite the presence of potential hosts in 


690 


the western slope of the Americas (Holthuis 
1952, Wicksten 1989, Villalobos-Hiriart & 
Nates-Rodriguez 1990, Wicksten & Hen- 
drickx 1992, Markham 1992), no species of 
Probopyrus have been described to date from 
the eastern Pacific, although P. pandalicola 
has been reported earlier from the Pacific 
drainage of the Americas by Markham 
(1974, 1985a, 1992), Campos & Campos 
(1989), Salazar-Vallejo & Leija-Tristan 
(1989), and Jimenez & Vargas (1990). 

From a study on the biology and ecology 
of M. tenellum, one of the most common 
prawns found in fresh and brackish water 
in the coastal area of the Mexican eastern 
Pacific, a large number of specimens were 
infested with a branchial parasite that was 
reported as Probopyrus sp. by Roman (1979, 
1983), and as P. pandalicola by Guzman & 
Roman (1983), and Roman (1991). 

The morphological analysis of approxi- 
mately 2500 specimens of this parasite col- 
lected between 1975 and 1992 in Guerrero 
and Michoacan States, Pacific coast of Mex- 
ico, and of specimens examined in the In- 
stituto de Biologia, Universidad Nacional 
Autonoma de Mexico (UNAM), and com- 
pared with holotypes and paratypes at the 
USNM Smithsonian Institution, lead me to 
the conclusion that the branchial parasite 
found on M. tenellum belonged to an un- 
described species of Probopyrus. 

The terminology and morphological 
characters used in the text are those pro- 
posed by Markham (1985a, 1985b, 1988). 
Figures 2B-I, and 2K—M were made with 
the aid of a photograph taken with scanning 
electron microscopy (SEM); two fully de- 
veloped individuals, male and female, were 
used for descriptions and chosen as type 
specimens. 


Probopyrus pacificensis, new species 
Figs... 2 


Bopyrids, Holthuis, 1954:6, 7 (Rio Zunzal, 
and Rio Conchalio, El Salvador, C.A.); 
infesting M. tenellum. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Probopyrus sp.—Roman, 1979:157; 1983: 
361; infesting M. tenellum. 

Probopyrus pandalicola.—Guzman & Ro- 
man, 1983:345-357 (not P. pandalicola); 
infesting M. tenellum.—Roman, 1991: 
109-119 (not P. pandalicola); infesting M. 
tenellum. 


Holotype female. —USNM 259483, allo- 
type male: USNM 259484. 

Paratypes. —USNM, Smithsonian Insti- 
tution: 184037, 235984, and 241946; San 
Diego Natural History Museum; Instituto 
Nacional da Pesquisas da Amazonia, Brazil 
(INPA); Rikjsmuseum van Natuurlijke His- 
torie, The Netherlands (RMNHL); Mu- 
seum national d’Histoire Naturelle, Paris 
(FMNHN); Instituto de Biologia (IB- 
UNAM), and Instituto de Ciencias del Mar 
y Limnologia ICMyL-UNAM), Mexico. 

Type locality.—Tres Palos Lagoon, Gue- 
rrero, México (16°43’ to 16°49’N, and 99°39’ 
to 99°46'W). 

Host.—The species has been collected 
only from Macrobrachium tenellum. 

Material examined. — Holotype of P. bi- 
thynis (USNM: 29089), holotype of P. flor- 
idensis (USNM: 29090), holotype of P. pan- 
amensis (USNM: 43503), and paratypes of 
P. pandalicola (USNM: 172345, and 
181548); ICMyL-UNAM: 2500 females, 
825 males, 11 cryptoniscus, and 1 epicarid- 
eum, collected between 1975 and 1992; Pa- 
cific coasts of Guerrero and Michoacan 
States, México. 

Range and habitat. —Eastern Pacific from 
México to El Salvador, C.A.; the southern 
part of the range is taken from a report by 
Holthuis (1954). Found in freshwater la- 
goons and associated rivers and ponds, al- 
ways as a branchial parasite of M. tenellum. 

Diagnosis. —Female (Fig. 1). Maximum 
length 13.5 mm, maximum width 11.0 mm. 
Distortion either dextral or sinistral, up to 
30°. Outline subovate. Anterolateral cor- 
ners of head slightly acute. Two pairs of 
subtriangular antennae, first one with three 
articles; second one bigger and with two ar- 
ticles. Maxilliped with acute plectron, palp 


VOLUME 106, NUMBER 4 691 


Fig. 1. Probopyrus pacificensis, new species. A, Holotype female (left) and allotype male (right), dorsal view. 
B, Allotype male (left), holotype female (right), ventral view. 


692 


of maxilliped unsegmented with 8 to 10 se- 
tae. Oostegite 1 relatively large, anterior 
margin concave, falcate and rounded; pos- 
terolateral tip triangular in shape and per- 
pendicular to the former; internal ridge with 
digitate processes. Pereopods with a high 
and rounded carina. Dark brown or black 
pigmentation present dorsally on tergal pro- 
jections of pereomeres 2—4 on short side; all 
oostegites with pigmentation. Small trans- 
verse patches of pigment present on per- 
eomeres 5—7. Pleopods protruding beyond 
border on short side. 

Description of female (Fig. 2A—I).— Head 
longer than wide, broadly cuneate, deeply 
set into first pereomere; anterolateral cor- 
ners usually produced into inconspicuous 
acute or rounded tip, lacking frontal lamina. 
First antennae with 3 articles and tuft of 
terminal setae (Fig. 2B), second antennae 
flat, bigger than first, with 2 articles; distal 
article with rounded tip and terminal setae 
(Fig. 2C). Maxilliped almost rectangular in 
outline, distally segmented (Fig. 2D); short 
nonarticulated setose ovoid palp on margin 
of anteromedial corner with 8 setae (Fig. 
2E); slender plectron with rounded tip ex- 
tending anteriorly (Fig. 2F); exterior border 
of maxilliped with a notch; posterior border 
nearly perpendicular to medial edge, its me- 
dial corner approximately in same line. Bar- 
bula with 2 lateral projections on each side; 
inner small and slender, outer broad and 
blunt; middle region produced into one acute 
and elevated ridge (Fig. 2G). Eyes absent. 

Margin of pereon forming a smooth curve; 
pereomeres 5—7 laminar and rounded, sep- 
arated by deep notches on long side; an- 
terolateral corners of pereomeres 1—4 pro- 
duced into distinct dorsolateral bosses, coxal 
plates narrow. Oostegites completely sur- 
rounding but not enclosing brood pouch; 
oostegite 1 relatively large covering most of 
anterior part of brood pouch; anterior mar- 
gins slightly concave, falcate and rounded 
(Fig. 2H); anterior segment approximately 
4 of total length; posterolateral tip subtrian- 
gular in shape and more or less perpendic- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ular to the former; internal ridge bearing 
many digitate ovoid processes. Small setae 
sometimes present on borders of inner ridge. 
Pigment, if present, in reticular pattern ex- 
cept on margins, or with small patches of 
black or brown pigment. Oostegites 2—4 
suboval, a little smaller than oostegite 1; 
fifth oostegite long and slender, falcate in 
outline, ending in blunt to sharp tip, densely 
setose along posterior margin, extending en- 
tirely across posterior region of brood pouch 
and overlapping opposite number. Pereo- 
pods with all articles distinct, increasing 
slightly in length posteriorly; basis of all per- 
eopods large, with round-shaped prominent 
carina, dactyli deeply set into propodi (Fig. 
21). 

Pleon about 1-1.5 as long as wide; pleo- 
meres separated laterally, margins perfectly 
differentiated; last pleomere bell-shaped, 
slightly notched posteriorly. Five pairs of 
biramous foliate pleopods almost complete- 
ly covering ventral surface of pleon and pro- 
truding on short side (Figs. 1, 2A); in each 
pleopod, endopod somewhat larger and 
overlapping exopod, both rami progres- 
sively smaller from first to fifth pleopods. 
Uropods absent. 

Description of male (Fig. 2J—M).— Length 
2.6 mm, width 1.0 mm. All segments of 
pereon similar in length; all regions of body 
distinct. Head slightly trapezoidal, rounded 
anteriorly, wider than long, extended and 
deeply inserted into first pereomere; an- 
terolateral borders rounded or slightly acute 
(Fig. 2J). Small, conspicuous eyespots; ir- 
regular pigmentation on pereomeres and 
pereopods. First antenna with globose prox- 
imal segment, short second article, and dis- 
tally setose third article (Fig. 2L); second 
antenna with short proximal segment and 
long terminal segment ending in inconspic- 
uous tuft of setae (Fig. 2K). Both antennae 
with scattered scales. Pereomeres 2—7 ap- 
proximately of same length, tips rounded 
and slightly reflexed ventrally; pereopods 
rather small, slightly larger posteriorly, not 
carinate. Pleon unpigmented; 5 pairs of dis- 


VOLUME 106, NUMBER 4 693 


Fig. 2. Probopyrus pacificensis, new species. A-I, holotype female; J—L, allotype male; M, paratype. A, Dorsal 
view; B, Antenna 1; C, Antenna 2; D, Maxilliped; E, palp of same; F, plectron of same. G, Head; H, oostegite 
1, internal view; I, Pereopod 7; J, Dorsal view; K, Antenna 2; L, Antenna 1; M, Pleon, ventral view. 


694 


tinct pleomeres deeply separated laterally, 
overall outline semicircular. The terminal 
pleomere button-shaped; 4 pairs of nearly 
sessile tuberculiform pleopods on pleo- 
meres, often prominent and conspicuous in 
ventral view (Fig. 2M). No uropods. 

Etymology. —Named after the Pacific 
Ocean to emphasize that it represents the 
first species of Probopyrus described from 
the Pacific coast of the Americas. 

Variations. —The young females of P. pa- 
cificensis usually have ocular spots, the 
length-to-width ratio may vary, sometimes 
length and width are equal, and the pleo- 
pods don’t stand out from the pleon edge. 
Adult females have from 8 to 10 setae in 
the maxilliped palpus; the digitate ovoid 
processes of the internal ridge of the first 
oostegite may be absent. The notch in the 
pleotelson may be present or absent in ei- 
ther young or adult females; whenever it is 
present it is shallow. Oostegites 3, 4 or both 
may be unpigmented, and the ridge of the 
barbula can be completely or partially bi- 
furcated. 

Head of males slightly rounded antero- 
lateral corners; pereomeres can be strongly 
pigmented or lack of it. Sometimes the per- 
ecpods have a pigmented spot on the basis, 
the pleon of P. pacificensis is highly variable 
and can be as wide as long; the last pleomere 
sometimes is fused into the fifth pair and 
shows a triangular, button, or slightly bi- 
lobate shape, and the pleopods can vary from 
4 to 5 pairs. 


Discussion 


The species of the genus Probopyrus “‘are 
so plastic and difficult to distinguish that 
their number is subject to considerable dis- 
pute” (Markham 1986), although many au- 
thors have tried to solve this problem. Rich- 
ardson (1905) proposed in her identification 
key the female’s size as the principal char- 
acter to separate species; she distinguished 
P. floridensis, P. bithynis and P. alphei 
(Richardson, 1900) as large-sized species, 
and P. latreuticola as small-sized species. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Chopra (1923) considered the abdominal 
ratio of males and females as a basis for 
differentiating the species of Probopyrus, and 
recognized P. palemoneticola, P. floridensis, 
P. bithynis, and P. panamensis as valid spe- 
cies, but P. pandalicola was not mentioned 
by this author. 

More recently Dale & Anderson (1982) 
recognized the validity of P. bithynis, P. flor- 
idensis and P. pandalicola, P. panamensis 
and P. palaemoneticola, which are recog- 
nized as valid species in this study, were not 
included. 

Although the criteria for separating the 
species still remains controversial, I con- 
sider that they could be identified not only 
through the use of morphological features 
(e.g., size, presence of notch in the last pleo- 
mere of females, etc.), but also through the 
use of additional characters such as the pig- 
mentation patterns of the individual (Bauer 
1981, Knowlton 1986, Knowlton & Mills 
1992), and specificity for the final host 
(Packard 1879; Bonnier 1900; Richardson 
1904, 1905, 1912; Chopra 1923; Markham 
1985a, 1985b), although some species 
seemingly can parasitize more than one host 
species. 

Considering these facts, P. pacificensis can 
be separated from P. palemoneticola be- 
cause in the latter the edges of the pereon 
are continuous, the carina of the seventh leg 
is quadrangular and high, and it parasitizes 
Palaemonetes vulgaris (Gissler, 1882; Rich- 
ardson, 1904), while P. pacificensis has a 
rounded carina and it is a parasite of VM. 
tenellum. 

In P. floridensis the edges of the pereon 
are slightly separated and the seventh leg 
has a rounded mid-sized carina, and it par- 
asitizes Palaemonetes exilipes and P. pal- 
udosus (Richardson, 1904; Dale & Ander- 
son, 1982); P. pacificensis has the edge of 
pereomeres 5—7 separated, and the carina, 
although also rounded, is higher than in the 
former. 

Probopyrus bithynis can be distinguished 
from all the American species of Probopyrus 


VOLUME 106, NUMBER 4 


because it has horn-shaped anterolateral 
processes on its head, a subtriangular and 
extremely high carina on the seventh leg 
(Richardson 1904, 1905), and it is a parasite 
of M. ohione and M. olfersi. 

Probopyrus pacificensis was quoted as P. 
pandalicola by Guzman & Roman (1983), 
and Roman (1983, 1991) from the eastern 
Pacific, but morphological characteristics, 
pigmentation patterns, and its final host al- 
low us to distinguish P. pandalicola in 
agreement with Markham’s review (Mark- 
ham 1985a: figs. 7-10). This species has 3 
articles on each antenna while in P. pacifi- 
censis the second antenna has only 2; in 
addition, the shape of this structure is re- 
markably different between both species; in 
P. pandalicola the barbula has 2 obtuse 
points in middle position, in P. pacificensis 
this structure has a high and acute edge and 
the shape of the plectron is acute. The palp 
of the maxilliped of P. pacificensis is ovoid 
and high (Fig. 2D, E); in P. pandalicola it 
is subtrapezoidal in shape. The first ooste- 
gite of P. pandalicola is curved; in P. pa- 
cificensis the distal part is straight in relation 
to the anterior part (Fig. 2H). The first leg 
of P. pandalicola has no carina, but all the 
legs of P. pacificensis are carinated. The 
pleotelson of some specimens of P. pan- 
dalicola have no notch, but a small tip on 
the posterior edge; in P. pacificensis a slight 
notch is always present. Pigmentation is 
present on the first four dorsal thoracic seg- 
ments of P. pandalicola on both sides; P. 
pacificensis has pigmentation only on seg- 
ments 2-4 on the smaller side (Figs. 1A, 
2A). 

P. pacificensis morphologically is more 
similar to P. panamensis than to the other 
species of the genus, but it is differentiated 
because the edge of the pereon is continuous 
while in P. pacificensis the borders of the 
pereomeres 5-7 are separate. In P. pana- 
mensis the carina of the seventh leg is high 
and middle-sized, in P. pacificensis it is high, 
rounded, and bigger than in the former; P. 
Danamensis parasitizes M. acanthurus. Fe- 


695 


males of P. pacificensis are slightly bigger 
than P. panamensis; both species have the 
anterior margin of head rounded, and the 
anterolateral angles small and acute. In P. 
panamensis, the head is wider than long, 
and the eyes are always absent (Richardson 
1905); in P. pacificensis the head is slightly 
longer than wide, and eyes or ocular spots 
sometimes are present. 

In general females of P. pacificensis show 
a more uniform dorsal pigmentation pat- 
tern than males on the dorsal side; the pleon 
of the males is almost always whitish or 
yellowish, and they have four or five pairs 
of conspicuous pleopods, while males of P. 
panamensis have three pairs of pleopods 
only (Richardson 1912). 

Males of P. panamensis have the head 
widely rounded while in P. pacificensis it is 
slightly trapezoidal. The pleon of P. paci- 
ficensis is clearly wider than its pereon; in 
P. panamensis it is rather narrow, although 
in both species the shape of the pleon is 
highly variable. 

Probopyrus pacificensis differs from all the 
described species from the Atlantic coast of 
North America, being larger, except for P. 
panamensis, because of the presence of pig- 
mented patches on tergal projections of per- 
eomeres 2-4, well developed pleopods pro- 
truding from the edge, and a high, round- 
shaped carina on all the legs. The males 
have a larger length/width ratio, trapezoidal 
shape of the head, and a bigger number of 
pleopods than P. panamensis. 

Finally, for the reasons stated above, and 
because of the distribution of the species 
compared to P. pacificensis, lagree with Dale 
and Anderson (1982), in recognizing P. pan- 
dalicola, P. floridensis, and P. bithynis; how- 
ever, P. panamensis and P. palaemoneticola 
should also be included as valid species. 


Acknowledgments 


I am indebted to B. Kensley and J. Clark, 
of the National Museum of Natural History 
(USNM) for providing facilities at the mu- 
seum during my short visits; to J. C. Mark- 


696 


ham (Arch Cape Marine Laboratory) for his 
interest in my progress; and to M. Hen- 
drickx (ICMyL, UNAM), and F. Alvarez 
(IB-UNAM), for his invaluable suggestions 
for the manuscript. Three anonymous re- 
viewers, and F. D. Ferrari, Associate Editor 
of PBSW comments improved the final 
manuscript. 


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VOLUME 106, NUMBER 4 


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1983. Impacto de parasitosis causada por 
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Instituto de Ciencias del Mar y Limnolo- 
gia, Universidad Nacional Autonoma de 
México (UNAM). P.O. BOX 70-305, Méx- 
ico, 04510. D.F. MEXICO. 


PROC. BIOL. SOC. WASH. 
106(4), 1993, pp. 698-704 


POTAMALPHEOPS DARWINIENSITS 
(CRUSTACEA: DECAPODA: ALPHEIDAE), THE 
THIRD INDO-WEST PACIFIC SPECIES 


A. J. Bruce 


Abstract.—Two species of the “African”? alpheid shrimp genus Potamal- 
pheops, P. hanleyi Bruce and P. pininsulae Bruce, have been recorded from the 
Indo-West Pacific region. The genus is otherwise known from four West African 
species and one species from a Mexican freshwater cave. A third Indo-West 
Pacific species, P. darwiniensis, from mangroves in Darwin Harbour, Northern 
Territory, is described and illustrated. The new species is closely related to the 
other Indo-West Pacific species, and a key is provided for their identification. 


The first species of the genus Potamal- 
pheops to be discovered was P. haugi, de- 
scribed by Coutiére (1906) as Alpheopsis 
haugi, from Gabon. Subsequently Sollaud 
(1932) described Alpheopsis monodi from 
the Cameroons and Senegal. Hobbs (1973) 
significantly extended the range of the genus 
Alpheopsis when reporting A. stygicola, from 
a freshwater cave at Oaxaca, Mexico. Pow- 
ell (1979) recorded a third African species, 
P. pylorus, from Nigeria and designated the 
genus Potamalpheops to include also the 
earlier described species. Hobbs (1983) con- 
cluded that A. stygicola should also be in- 
cluded in the genus Potamalpheops. Re- 
cently, Bruce (1991) reported the first 
occurrence of the genus in the Indo-West 
Pacific region, with P. hanleyi from Darwin 
Harbour, Australia, and then the presence 
of a second troglobitic species, P. pininsu- 
lae, from the Isle of Pines, New Caledonia. 
The presence of another West African spe- 
cies, in the Calabar River, has been indi- 
cated by Powell (1979), but this species is 
still undescribed. A second Australian spe- 
cies has recently been collected and is here 
described and illustrated. 


Systematic Account 


Alpheidae Rafinesque, 1815 


Potamalpheops Powell, 1979 
Potamalpheops darwiniensis, new species 
Figs. 1-3 


Material examined.—1 6, holotype, sta- 
tion JRH 2-Q3-(M), Hudson’s Creek, Dar- 
win Harbour, Northern Territory, Austra- 
hia, 12°28:75'S; 130°55.67'E; 23ahenaigoe 
intertidal, coll. J. R. Hanley, Northern Ter- 
ritory Museum Cr. 007922. 

Description of holotype.—Small, of sub- 
cylindrical body form, body slightly com- 
pressed, with the distal part of left third 
maxilliped and second pereiopod missing, 
right second pereiopod in early stage of re- 
generation. 

Rostrum (Fig. 2A, B) very short, scarcely 
exceeding anterior margin of cornea, acute, 
broadly triangular in dorsal view, dorsal ca- 
rina obsolete, ventral carina feebly devel- 
oped, unarmed, lateral carinae broad, con- 
fluent with orbital margin. 

Carapace depilate, smooth; orbital mar- 
gin concealing major portion of eye (Fig. 
2B), with very feebly developed, rounded, 
antennal region, pterygostomial angle 
slightly produced, bluntly angular, ventral 
margin with numerous plumose setae; car- 
diac notch distinct. 

Abdomen depilate, glabrous; sixth seg- 
ment (Fig. 2H) about 1.2 times length of 


VOLUME 106, NUMBER 4 


Se —— 


ae 


qo D. 
\- RR 


x 
\s 


Fig. 1. 


fifth, 1.5 times longer than deep, com- 
pressed, posterior lateral angle acute, pos- 
teroventral angle with articulated triangular 
plate; pleura of first four segments broadly 
rounded, fifth subrectangular, posteroven- 
tral angle subacute. Telson (Fig. 2K) about 
1.25 times length of sixth segment, 1.8 times 
longer than anterior width, lateral margins 
straight, posteriorly convergent, with small 
dorsal spines, about 0.07 of telson length, 
anterior pair at 0.3 of telson length, poste- 
rior pair at about 0.6 (spine missing on right); 
posterior margin (Fig. 3G) broad, about 0.6 
of anterior width, with 2 pairs of robust 
subequal subventral spines laterally, about 
3.0 times length of dorsal spines, central 
portion of posterior margin semicircular, 
occupying about half posterior margin 
width, with about 20 long plumose setae, 
with numerous short slender simple spi- 
nules dorsally; anal tubercles feebly devel- 
oped. 


699 


Potamalpheops darwiniensis, new species, holotype male, Hudson’s Creek, Darwin. Scale bar in mm. 


Antennule (Fig. 2E) with peduncle robust, 
about 0.4 of carapace length, distinctly ex- 
ceeding scaphocerite and carpocerite; prox- 
imal segment about 1.4 times longer than 
proximal width, distodorsal margin non- 
dentate, with well developed ventromedial 
carina; statocyst normally developed; with 
broad acute stylocerite reaching distally to 
anterior margin of segment; intermediate 
segment subcylindrical, about 1.1 times 
proximal segment length, 2.0 times longer 
than wide, ventromedial border with long 
plumose setae; distal segment subcylindri- 
cal, about 0.5 of intermediate segment 
length; upper flagellum biramous, proximal 
13 segments fused, shorter ramus with sin- 
gle free segment only, with about 8 groups 
of aesthetascs; lower ramus slender, fili- 
form, subequal to longer upper ramus length 
(tips of rami missing). 

Antenna (Fig. 2F) with stout basicerite, 
with small acute process projecting laterally 


700 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


B 


SSS 


SS 


SS 
>> 


J | 


Fig. 2. Potamalpheops darwiniensis, new species, holotype male. A, Anterior carapace and antennal pedun- 
cles, dorsal. B, Anterior carapace, lateral. C, Eye, lateral. D, Same, dorsal. E, Antennular peduncle. F, Antennal 
peduncle. G, Epipod of first pereiopod. H, Sixth abdominal segment, lateral. I, Endopod of second pleopod. J, 
Same, appendices interna and masculina. K, Telson. L, Uropod. 


VOLUME 106, NUMBER 4 701 


H G . 


Fig. 3. Potamalpheops darwiniensis, new species, holotype male. A, Right first pereiopod. B, Same, chela. 
C, Same, fingers. D, Right third pereiopod. E, Same, propod and dactyl. F, Endopod of first pleopod. G, Posterior 
margin of telson. H, Uropod, diaeresis of exopod, dorsal. 


702 


from anterodorsal margin, with strong acute 
ventrolateral tooth, carpocerite robust, dis- 
tinctly exceeding distal margin of scapho- 
cerite, subcylindrical, slightly compressed, 
about 3.0 times longer than wide, flagellum 
long, slender, 2.2 times carapace length, 
proximal segments not thickened; scapho- 
cerite reaching to about middle of distal seg- 
ment of antennular peduncle, 1.8 times lon- 
ger than wide, suboval, lateral margin 
convex, with small acute distal tooth scarce- 
ly exceeding broadly rounded anterior mar- 
gin of lamella. 

Eyes reduced (Fig. 2C, D), largely con- 
cealed by anterior margin of carapace, an- 
terior surface of cornea only exposed in dor- 
sal view, contiguous in midline, cornea 
small, well pigmented, peduncle short, sub- 
cylindrical, medial surface flattened, with 
small distinct subacute dorsomedial tuber- 
cle, without setae. 

Mouthparts not dissected. Third maxil- 
liped extending to distal end of antennular 
peduncle, terminal segment tapering distal- 
ly, obliquely truncate, with 4 stout spines, 
largest spine strongly cornified; coxa with 
strap-like epipod. 

First pereiopods (Fig. 3A) subequal, sim- 
ilar, chelae appearing to be carried in flexed 
position; chelae (Fig. 3B) robust, slightly 
longer on left, slightly stouter on right, about 
0.4 of carapace length, palm subcylindrical, 
slightly swollen, compressed, smooth, about 
1.65 times longer than deep, fingers (Fig. 
3C) stout, about 0.6 of palm length, feebly 
subspatulate, with very stout, strongly cor- 
nified, blunt, feebly bidentate tips, dactylus 
about 2.8 times longer than deep, curved, 
with entire sharp unarmed cutting edge; car- 
pus stout, about 0.28 of chela length, 0.45 
of palm length, distally excavate, unarmed, 
with about 7 transverse rows of long ser- 
rulate cleaning setae ventromedially; merus 
about 0.5 of chela length, 2.4 times longer 
than wide, uniform, ventral surface feebly 
excavate, unarmed; ischium about 0.38 of 
chela length, 2.0 times longer than distal 
width, tapered proximally, unarmed; basis 
obliquely articulated with ischium, un- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


armed; coxa robust, with 6-setose seto- 
branch dorsally, strap-like epipod (Fig. 2G) 
laterally. 

Second pereiopod (left) with proximal 
carpus and proximal segments showing no 
special features, coxa with setobranch and 
strap-like epipod. 

Ambulatory pereiopods moderately ro- 
bust; third pereiopod (Fig. 3D) slightly ex- 
ceeding antennular peduncle by length of 
dactyl, dactyl (Fig. 3E) slender, simple, 
curved, compressed, about 0.5 length of 
propod, ventral margin sharply carinate, 
without clearly demarcated unguis, with 
single seta distodorsally; propod (Fig. 3E) 
about 0.3 of carapace length, 6.0 times lon- 
ger than wide, uniform, glabrous, with pair 
of short simple distoventral spines, about 
0.12 of dactyl length, ventrolateral row of 
6 small spines, ventromedial row of 4 small- 
er spines; carpus about 0.6 of propod length, 
3.2 times longer than distal width, with small 
distoventral spine; merus 1.35 times longer 
than propod, 5.2 times longer than wide, 
uniform, with large mobile ventrolateral 
spines at 0.38 and 0.68 of length; ischium 
about 0.5 of merus length, 2.4 times longer 
than distal width, strongly tapered proxi- 
mally, with single small mobile spine ven- 
trolaterally; basis normal; coxa robust, with 
6-setose setobranch dorsally, strap-like epi- 
pod laterally. Fourth pereiopod similar to 
third. Fifth pereiopod similar to fourth, 
more slender, dactyl 0.5 of propod length, 
propod 6.0 times longer than wide, 0.55 of 
carapace length; merus with proximal ven- 
trolateral spine; ischium unarmed; coxa with 
setobranch, without epipod. 

Abdominal sternites with narrow trans- 
verse ridges between bases of pleopods, un- 
armed. 

Pleopods normal. Endopod of first pleo- 
pod (Fig. 3F) slender, 3.6 times longer than 
basal width, curved, tapering distally, with 
11 plumose setae medially, 17 similar setae 
distally and laterally, distal setae longer, 
about 0.6 of endopod length. Endopod of 
second pleopod (Fig. 2I) 5.0 times longer 
than wide, with appendices at 0.38 of length; 


VOLUME 106, NUMBER 4 


appendix masculina (Fig. 2J) subcylindri- 
cal, 7.0 times longer than wide, with 4 long 
simple terminal spines, about 0.6 of corpus 
length, 3 similar spines distomedially; ap- 
pendix interna short, reaching to middle of 
appendix masculina corpus, with few distal 
cincinnuli. 

Uropod (Fig. 2L) with protopod bearing 
large acute distodorsal lobe, rami slightly 
exceeding posterior telson margin; exopo- 
dite 2.3 times longer than wide, greatest 
width at 0.6 of length, lateral margin slightly 
convex, with well developed submarginal 
setal fringe, distolateral angle subrectangu- 
lar, with large mobile spine medially, di- 
aeresis (Fig. 3H) well developed with den- 
tate dorsal flange laterally extending across 
about 0.8 of width, with about 24 small 
subuniform acute teeth (tips of many abrad- 
ed), distal lamella large, broadly rounded, 
highly flexible, with short simple spiniform 
marginal setae distolaterally, otherwise with 
long densely plumose marginal setae; en- 
dopodite about 0.9 of exopod length, 2.2 
times longer than wide. 

Measurements. —Carapace length 6.2 
mm, total body length (approx.) 17 mm. 

Etymology. —Specific name derived from 
locality of capture, Darwin Harbour. 

Habitat. —‘‘Mud-mound,”’ amongst 
mangroves. 

Affinities. —Potamalpheops darwiniensis 
is closely related to the two other Indo-West 
Pacific species, both of which belong to the 
species group characterized by the presence 
of two pairs of posterior telson spines in- 
stead of three. This group also includes the 
West African species P. monodi (Sollaud, 
1932), from Cameroon and Senegal. 

Potamalpheops darwiniensis is most 
closely related to P. hanleyi, which is also 
known only from Darwin Harbour. Pota- 
malpheops darwiniensis may be distin- 
guished from P. hanleyi by the following 
features: —(1) the presence of a much short- 
er rostrum, only slightly exceeding feebly 
produced blunt extracorneal teeth; (2) the 
feebly produced antennal and pterygosto- 
mial angles; (3) the presence of numerous 


703 


short plumose setae at the pterygostomial 
angle; (4) the subrectangular, posteroven- 
trally subacute pleuron of the fifth abdom- 
inal segment; (5) dorsal telson spines at 0.3 
and 0.6 of telson length; (6) convex central 
portion of posterior margin of telson more 
than half posterior margin width, with more 
numerous plumose setae; (7) proximal seg- 
ment of antennular peduncle with disto- 
dorsal margin lacking denticulations, ven- 
tromedial carina distally subrectangular; (8) 
basicerite with acute distolateral process 
dorsally; (9) carpocerite distinctly exceeding 
scaphocerite; (10) scaphocerite oval in shape, 
with distolateral tooth scarcely exceeding 
lamella; (11) eye markedly reduced, largely 
concealed by anterior carapace, cornea 
small, eyestalk flattened medially with small 
acute dorsomedial tubercle; (12) first pe- 
reiopods with robust chelae; (13) third am- 
bulatory pereiopod with propod more slen- 
der, 6.0 times longer than distal width, with 
ventromedial and ventrolateral spine rows; 
(14) male first pleopod endopod elongate, 
tapering, curved, with numerous long plu- 
mose setae distomedially; (15) male second 
pleopod endopod with appendices arising 
at less than half medial margin length, ap- 
pendix masculina with 7 longer spines dis- 
tally; (16) exopod of uropod with distolater- 
al angle subrectangular, diaeresis with about 
24 small acute denticles. 

Potamalpheus darwiniensis is readily dis- 
tinguished from the only other Indo-West 
Pacific species, P. pininsulae, by its lack of 
a long, slender, acute, ventrally dentate ros- 
trum. 


Discussion 


Little is known about the precise ecolog- 
ical niche occupied by the various species 
of the genus Potamalpheops. The collection 
of the single specimen of the present species 
from a “mud-mound”’ suggests the possi- 
bility that this species may be associated 
with some burrowing invertebrate. Man- 
grove “mud-mounds” are commonly pro- 
duced by annelids or decapod crustaceans, 
particularly thalassinids. No Indo-West Pa- 


704 


cific caridean shrimps are so far known to 
associate with thalassinideans. 

The discovery of a third species of Po- 
tamalpheops in the Indo-West Pacific re- 
gion clearly establishes this genus a com- 
ponent of its fauna. It seems likely that 
additional specimens or species of these 
shrimps, which are so far known only from 
very shallow coastal or fresh waters, will be 
found in due course, and will link the east- 
ern distribution of the Indo-West Pacific 
species with those with the West African 
species. 

The Mexican species, P. stygicola, may 
be less closely related to the other species 
of the genus, from which it differs by the 
presence of a remarkable longitudinal he- 
patico-branchiostegal groove (Hobbs 1973) 
that cannot be discerned in the other non- 
Mexican species. Potamalpheops stygicola 
could well have had an independent evo- 
lutionary origin from the other species of 
the genus Potamalpheops and may belong 
to a separate genus. 


Key to the Indo-west Pacific Species of 
Potamalpheops Powell 


1. Rostrum well developed, reaching 
to distal margin of proximal seg- 
ment of antennular peduncle, with 
single acute ventral tooth 

P. pininsulae Bruce, 1993 

— Rostrum short, not nearly reaching 
distal margin of proximal segment 
of antennular peduncle, ventrally 
unarmed 

2. Rostrum very short, not reaching 
bases of antennular peduncles; eyes 
reduced, largely covered by anterior 
carapace; extracorneal tooth obso- 
lescent; first pereiopods well devel- 
oped, with robust chelae; diaeresis 
of exopod of uropod with about 24 
smiallacute denticles.... see sas oe 

P. darwiniensis, new species 

— Rostrum exceeding middle of prox- 
imal segment of antennular pedun- 


"2 © © © © ew we ew ew 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


cle; eyes well developed, cornea 
largely exposed by anterior margin 
of carapace; extracorneal tooth 
acute; first pereiopods feebly devel- 
oped, with small chelae; diaeresis of 
exopod of uropod with about 18 
acute teeth .... P. hanleyi Bruce, 1991 


Acknowledgment 


I am most grateful to my colleague, Dr. 
J. R. Hanley, for bringing this specimen to 
my attention. : 


Literature Cited 


Bruce, A. J. 1991. The “African” shrimp genus Po- 
tamalpheops in Australia, with the description 
of P. hanleyi, new species (Decapoda: Alphei- 
dae). — Journal of Crustacean Biology 11(4):629- 

634. 

1992 [1993]. Potamalpheops pininsulae sp. 
Nnov., a new stygiophilic shrimp from New Cal- 
edonia (Crustacea: Decapoda: Alpheidae).— 
Stygologia 87(4):23 1-242. 

Coutiére, H. 1906. Sur un nouvelle espéce d’Alpheop- 
sis, A. haugi, provenant d’un lac d’eau douce du 
bassin de l’Ogoué (Voyage de M. Haug, 1906). — 
Bulletin du Muséum d’Histoire Naturelle, Paris 
12:376-380. 

Hobbs, H. H., Jr. 1973. Two new troglobitic shrimps 

(Decapoda: Alpheidae and Palaemonidae) from 

Oaxaca, Mexico.— Association of Mexican Cave 

Studies, Bulletin 5:73-80. 

1983. The African shrimp genus Potamal- 
pheops in Mexico (Decapoda, Alpheidae).— 
Crustaceana 44(2):22 1-224. 

Powell, C. B. 1979. Three alpheid shrimps of a new 
genus from West African fresh and brackish wa- 
ters: taxonomy and ecological zonation (Crus- 
tacea Decapoda Natantia). — Revue de Zoologie 
Africaine 93:116-—-150. 

Rafinesque, C. S. 1815. Analyse de la Nature ou Ta- 
bleau de l’Univers et de ses Corps organisés, 
Palermo, Sicily, 1-224. 

Sollaud, E. 1932. Sur un alphéidé d’eau douce, Al/- 
pheopsis monodi n. sp., receulli par M. Th. Mo- 
nod au Cameroun.— Bulletin de la Société Zoo- 
logique de France 57:375-386. 


Division of Natural Sciences, Northern 
Territory Museum, P.O. Box 4646, Darwin, 
Australia 0801. 


PROC. BIOL. SOC. WASH. 
106(4), 1993, pp. 705-713 


ON A NEW GENUS AND SPECIES OF XANTHID CRAB 
(CRUSTACEA: DECAPODA: BRACHYURA) FROM 
CHESTERFIELD ISLAND, CORAL SEA 


Peter K. Ne 


Abstract.—A new genus and species of xanthid crab, Cranaothus deforgesi, 
is described from Chesterfield Island. Cranaothus appears to be closely related 
to Paramedaeus, Metaxanthops, Macromedaeus, Medaeops, Neoxanthops, 
Glyptoxanthus and Lipaesthesius, but differs in the form and sculptures on the 
carapace, as well as structures of the sternum, male abdomen and male first 
pleopod. The larger cheliped of Cranaothus also possesses a specialized basal 


cutting tooth on its dactylus. 


A collection of Brachyura from Chester- 
field Island was deposited in the Muséum 
national d’Histoire naturelle (MNHN), Par- 
is, by ORSTOM (Institut Francais de Re- 
cherche Scientifique pour le Developement 
en Coopération). Among these specimens 
was an interesting crab from Chesterfield 
Island with several peculiar features distin- 
guishing it from all other known genera and 
species in the family Xanthidae MacLeay, 
1838 (sensu Guinot 1978). 

The description of this new genus and 
species forms the text of the present paper. 
Abbreviations G1 and G2 are for the male 
first and second pleopods respectively. 
Measurements are reported in millimeters, 
in the sequence carapace width by carapace 
length. 


Systematic Account 


Family Xanthidae MacLeay, 1838 
(sensu Guinot, 1978) 
Subfamily Euxanthinae Alcock, 1898 
(sensu Seréne, 1984) 
Cranaothus, new genus 


Diagnosis. —Carapace quadrate, regions 
not well defined; dorsal surfaces with very 
small squamiform granulations; branchial, 
gastric, cardiac and intestinal regions with 


eroded vermiform granulated ridges; front 
distinctly produced; lobes truncatiform, 
separated by deep fissure extending to epi- 
gastric region; external orbital angle low, in- 
distinct, not clearly demarcating beginning 
of anterolateral margin, joining series of 
smaller granules curving gently downwards 
below supraorbital margin, across subor- 
bital region and towards buccal cavity; an- 
terolateral margin not lobulated or toothed, 
anterior 73 arcuate, posterior 43 subparallel 
to median longitudinal carapace axis; pos- 
terolateral margins gently concave. Ster- 
nites 2-4 broad, sternal suture 1 and 2 com- 
plete, sternal sutures 2 and 3, and 3 and 4 
interrupted medially. Chelipeds distinctly 
asymmetrical, fingers sharp, without pig- 
mentation, larger cheliped with pronounced 
molariform basal cutting tooth on dactylus. 
Lateral margins of fused male abdominal 
segments 3—5 entire, continuous; segment 7 
semicircular, lateral margins strongly con- 
vex, tip rounded. 

Type species. — Cranaothus deforgesi, new 
species, by monotypy. 

Etymology.—The generic name is de- 
rived from the Greek “‘kranaos”’ for “rugged 
and rocky” (alluding to the eroded carapace 
surface), in arbitrary combination with a fi- 
nal syllable of many xanthid genera. Gender 
masculine. 


706 


Remarks.—In its external features Cra- 
naothus, new genus, appears to be closest 
to the genus Paramedaeus Guinot, 1967; 
however, it differs in that the anterolateral 
margin is not cut into distinct teeth (weak 
or otherwise), the front is very produced and 
lamellar, with the median fissure very deep; 
sternites 2—4 distinctly broader, sternal su- 
ture 1 and 2 being distinct (absent in Para- 
medaeus), sternal suture 2 and 3 interrupted 
medially (entire in Paramedaeus); male ab- 
dominal segment 7 semicircular in shape 
(distinctly triangular in Paramedaeus); dis- 
tal part of the G1 is long, slender and strong- 
ly tapering, the tip being relatively sharp 
(distal part stout and tip rounded in Para- 
medaeus) (Guinot 1967, Seréne 1984). 

With regard to the shape of the carapace, 
Cranaothus resembles Indo-West Pacific 
genera such as Metaxanthops Seréne, 1984, 
Macromedaeus Ward, 1942, and Neoxan- 
thops Guinot, 1968, but differs in many key 
aspects. The front margin of Cranaothus is 
somewhat similar to that of Metaxanthops, 
with two broad and truncate lobes separated 
by a deep median fissure. Metaxanthops 
however, differs from Cranaothus in having 
well developed and distinct epibranchial 
teeth and a much smoother carapace sur- 
face. The general shape of the G1 in Meta- 
xanthops, although similar, is distinctly 
stouter and the distal part is not slender and 
tapering (Seréne 1984: fig. 129). The an- 
terolateral margin of Cranaothus resembles 
Macromedaeus nudipes (Milne Edwards, 
1867), but the structure of the front is dif- 
ferent, with the lobes more sinuous and less 
projected forward in Macromedaeus. In 
Macromedaeus, the two frontal lobes are 
also separated only by a cleft, without the 
deep fissure present in Cranaothus. While 
the regions are well defined in Macrome- 
daeus, they are only vaguely so in Crana- 
othus. The G1 of Cranaothus differs sub- 
stantially from those of Macromedaeus, 
being proportionately shorter, stouter and 
different in shape (Seréne 1984: figs. 101- 
104). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


The carapace shape of Cranaothus is per- 
haps closest to Neoxanthops, especially in 
species like N. cavatus (Rathbun, 1907) 
which also has ridges on the dorsal surface. 
In N. cavatus, however, the frontal lobes are 
separated only by a shallow cleft, not a deep 
fissure, and the anterolateral margin is cut 
into distinct lobes. The G1 of Cranaothus 
is also proportionately shorter, with a dif- 
ferent shape, and the distal part is tapered 
and very slender, compared to that of typ- 
ical Neoxanthops (see Seréne 1984: figs. 127, 
128). Neoxanthops cavatus is not a typical 
member of Neoxanthops and should prob- 
ably be transferred to a new genus. It differs 
markedly from the type species of the genus, 
N. lineatus Milne Edwards, 1867, in many 
aspects, viz. the anterolateral margin grad- 
ually becomes more obscure and gently 
curves to end below the orbits, and does not 
meet the external orbital angle or supraor- 
bital margin; the frontal margin is not dis- 
tinctly produced beyond the internal angle 
of the supraorbital margin; the anterolateral 
margin is not distinctly cristiform; the sur- 
face is more distinctly domed, distinctly 
sculptured, appears eroded, and instead of 
gently convex and completely smooth; the 
fingers of the chelipeds are very short in- 
stead of pigmented; and the subdistal part 
of the G1 has only a few short, simple hairs, 
moderately long, setose hairs. 

Cranaothus also differs from Macrome- 
daeus, Neoxanthops and Metaxanthops in 
that the dactyls of the larger cheliped have 
a molariform basal cutting tooth absent in 
the other genera. 

With regard to the sculpturing on the car- 
apace surface, Cranaothus somewhat re- 
sembles the genus Glyptoxanthus Milne Ed- 
wards, 1873-1881, which is represented in 
the Indo-West Pacific region by G. mean- 
drinus (Klunzinger, 1913). The anterolat- 
eral margin of Glyptoxanthus, however, is 
cut into distinct teeth, the front is not pro- 
jecting but is about level with the orbits, the 
frontal lobes are not separated by a deep 
fissure, the dactylus of the cheliped lacks the 


VOLUME 106, NUMBER 4 


special basal cutting tooth, and the last male 
abdominal segment is distinctly triangular, 
with a sharp tip and the lateral margins al- 
most straight (Odhner 1925: pl. 4 fig. 1, 
Guinot 1979: pl. 6 fig. 7). In Cranaothus, 
the last male abdominal segment is semi- 
circular. There is also some resemblance to 
crabs of the genus Medaeops Guinot, 1967, 
although the shapes of the carapaces differ. 
Interestingly, Medaeus granulosus (Has- 
well, 1882) has a weak basal cutting tooth 
on the dactyls of the chelipeds. 

The shape and form of the carapace of 
Cranaothus also bears a marked similarity 
to an eastern Pacific monotypic genus, Li- 
paesthesius Rathbun, 1898, represented by 
L. leeanus Rathbun, 1898. Like Crana- 
othus, the surface of the carapace in Li- 
paesthesius is covered by many small gran- 
ules, forming uneven patterns. Lipaesthesius 
differs from Cranaothus in the following as- 
pects: point of attachment of antennal fla- 
gellum concealed by upper margin of basal 
segment (point of attachment of flagellum 
distinct and not concealed in Cranaothus); 
granule patterns on dorsal surface of cara- 
pace not vermiform; front not produced for- 
ward and without deep median cleft; pos- 
terolateral margin strongly concave (more 
so than in Cranaothus), forming distinct 
““waist’’; junction between antero- and pos- 
terolateral margins not clearly demarcated, 
the anterolateral margin gradually curving 
posteriorly; carpus of cheliped very elon- 
gate; chelae symmetrical; fingers of cheliped 
long and slender and pigmented dark brown 
(Rathbun 1930:272, pl. 112). 

Seréne’s (1984) separation the subfami- 
lies Euxanthinae Alcock, 1898, and Xan- 
thinae MacLeay, 1838, is not satisfactory as 
there appears to be a degree of overlap in 
some of the characters used by him. The 
establishment of Cranaothus further com- 
plicates matters because, while the genus 
seems to belong to what Seréne (1984) de- 
fined as Euxanthinae (cf. Medaeops, Para- 
medaeus, Glyptoxanthus), it also bears a 
striking resemblance to some members of 


707 


the Xanthinae (cf. Neoxanthops, Metaxan- 
thops, Macromedaeus). The genus is placed 
in the Euxanthinae based on the absence of 
a clearly defined anterolateral margin be- 
hind the external orbital angle in Crana- 
othus, with the row of granules curving along 
the suborbital region toward the buccal cav- 
ity. 


Cranaothus deforgesi, new species 
Figs. 1-3 


Paramedaeus noelensis. —? Seréne & Umali, 
1972:68, pl. 7 figs. 7—9 [nec Paramedaeus 
noelensis (Ward, 1934)]. 


Material examined. —Holotype male 
(carapace 8.0 by 5.9 mm), (MNHN), Ches- 
terfield Island, Coral Sea, station 144, 
1927. 73'S, 158°23:28'E,.. ca. 50) mx depth, 
sand and Halimeda algae substrate, dredge, 
leg. B. Richer de Forges, 30 Aug 1988. 

Description of holotype male. —Carapace 
regions not well defined, grooves separating 
gastric and branchial regions shallow, 2F, 
1M, 3M regions low but discernible, 4M 
indistinct, L and R not defined; H-shaped 
groove separating cardiac and gastric regions 
distinct; dorsal surfaces covered with very 
small squamiform granulations; branchial, 
gastric, cardiac and intestinal regions with 
uneven eroded ridges, forming vermiculat- 
ed pattern; pterygostomial, suborbital and 
sub-branchial regions granulose. Density of 
granules and granulations somewhat ob- 
scures sutures between pterygostomial, 
sub-branchial and suborbital regions as well 
as base of chelipeds, ambulatory legs and 
sternum. Front distinctly produced beyond 
imaginary line connecting internal supra- 
orbital angle; distinctly bilobed, lobes sep- 
arated by very deep fissure extending back 
to epigastric region; surfaces smooth, mar- 
gin truncatiform, gently concave. Supraor- 
bital margin with clearly defined rounded 
inner angle, separated from front by distinct 
groove; external orbital angle low, indis- 
tinct, not clearly demarcating beginning of 
anterolateral margin; rest of supraorbital 


708 


margin entire, gently sinuous. Internal in- 
fraorbital angle with distinct tooth visible 
from dorsal view. Anterolateral margin ar- 
cuate along anterior *3, becoming straight 
along posterior '4 (approximately parallel to 
median longitudinal axis of carapace); not 
divided into distinct lobes or teeth, without 
any distinct cristate borders or clefts; 1 tu- 
bercle visible shortly behind external orbital 
angle, followed by 3 larger ones on arcuate 
part of margin; straight part of anterolateral 
margin marked by blunt tubercle on each 
edge; anterolateral margin distinctly sepa- 
rated from gently concave, converging pos- 
terolateral margins. Posterior margin of car- 
apace gently convex. Antennules folding 
transversely, antennular fossae partly cov- 
ered by protruding front from frontal view. 
Antennal flagellum short, attached to stout 
basal segment occupying entire space be- 
tween antennular fossa and internal orbital 
angle. Endostome with weak median lon- 
gitudinal ridge and strong, oblique ridge on 
either side of posterior part, adjacent to 
mouth. 

Entire outer surface of third maxilliped 
finely granulose, that on merus appearing 
more eroded; ischium rectangular, median 
oblique sulcus very shallow, indistinct; me- 
rus quadrate, with median oblique patch of 
eroded granules; outer surface of carpus 
rounded, granuliform; exopod reaching an- 
terior edge of merus, with blunt triangular 
subdistal tooth on inner margin, flagellum 
long. 

Sternum broad, entire surface covered 
with eroded granules, appearing squamate; 
suture between sternites 1 and 2 distinct, 
complete, sutures between sternites 2 and 
3, and 3 and 4 shallow, interrupted medi- 
ally, sutures between sternites 4 and 5, 5 
and 6, and 6 and 7 incomplete; abdomen 
reaching to imaginary line joining posterior 
bases of chelipeds. Gonopore coxal, open- 
ing below abdominal segment 3. 

Chelipeds distinctly asymmetrical; outer 
surfaces of merus, carpus and chelae cov- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ered with very small scale-like granulations 
and vermiform eroded ridges; carpus short, 
rounded, inner distal angle with distinct 
blunt tooth and uneven anterior subdistal 
serrations; fingers not pigmented black or 
brown, appearing white in preservative; 
larger cheliped with pronounced molari- 
form basal cutting tooth on dactylus di- 
rected obliquely outward; minor cheliped 
with elongated fingers. 

Abdomen with segments 3—5 completely 
fused, sutures separating these segments not 
discernible, lateral margins entire, without 
any clefts or discontinuity; segments 1-3 
trapezoidal, segment 6 squarish, lateral 
margins straight, parallel; segment 7 semi- 
circular, lateral margins strongly convex, tip 
rounded; surfaces of all segments slightly 
rugose to squamate. 

G1 relatively short and stout, proximal 
part gradually tapering, distal part straight, 
very slender, distinctly tapering to sharp tip; 
lateral margins of slender distal part lined 
with short spines; subdistal part with nu- 
merous long, stout setiferous hairs. G2 short, 
slender, distal part with petaloid process. 

Etymology. —The species is named after 
Bertrand Richer de Forges, who so kindly 
made the ORSTOM specimen available for 
study. 

Remarks.—The single known specimen 
of Cranaothus deforgesi, new species, is ma- 
ture despite its small size because the gon- 
opods are fully developed. The vermiform 
ridges on the carapace and chelipeds are 
formed by patches of very small granules 
and are easily chipped and scraped off. This 
accounts for the species’ eroded appearance. 

Cranaothus deforgesi bears a striking re- 
semblance to a specimen from the Philip- 
pines identified as Paramedaeus noelensis 
(Ward, 1934) by Seréne & Umali (1972). 
Seréne & Umali’s (1972:68, pl. 7 figs. 7-9) 
specimen, was a male 8.5 by 5.5 mm from 
Maluso Bay, collected from a depth of 25 
m by the Pele Sulu Expedition in 1964. The 
specimen has a very produced and lamellar 


VOLUME 106, NUMBER 4 


709 


Fig. 1. 
Chesterfield Island. A, dorsal view; B, chelae. 


front, and the median fissure appears to be 
very deep. The shape of the carapace, sculp- 
ture of the surface (covered with granulated 
vermiculations) and form of the postero- 
lateral margin, also resemble C. deforgesi. 
Seréne & Umali (1972:69) commented on 
the differences between their Philippine 
specimen and Ward’s (1934) description of 


Cranaothus deforgesi, new genus and species. Holotype male, carapace 8.0 by 5.9 mm, MNHN, 


P. noelensis, noting that in their specimen 
**... the postero-lateral border is concave 
instead of being straight . . . [t]he breadth is 
1.41 its length instead of 1.47, the propor- 
tion in the specimen of Forest & Guinot 
(1961)... . the front in our specimen is more 
salient, more pointed medially with the si- 
nus more open.” It appears that Seréne & 


710 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Cranaothus deforgesi, new genus and species. Holotype male, carapace 8.0 by 5.9 mm, MNHN, 
Chesterfield Island. A, dorsal view of carapace; B, frontal view of carapace; C, buccal cavity showing endostomial 
ridges; D, left third maxilliped (denuded, most of granulation not shown); E, sternites 1-4; F, abdomen; G, coxa, 
basis-ischium and merus of left cheliped; H, right carpus of cheliped (dorsal view); I, right fourth ambulatory 
leg; J, right third ambulatory leg. Small granules and squamate structures on surfaces of carapace, third maxilliped, 
sternum, chelipeds and abdomen omitted. Scales: A, B, E-J, 1.0 mm; C, D, 0.5 mm. 


VOLUME 106, NUMBER 4 


711 


Fig. 3. Cranaothus deforgesi, new genus and species. Holotype male, carapace 8.0 by 5.9 mm, MNHN, 
Chesterfield Island. A—D, left G1; E, F, left G2. A, C, F, ventral view; B, D, E, dorsal view. Scales: A, B, E, 0.5 


mm; C, D, F, 0.25 mm. 


Umali’s (1972) specimen is actually con- 
specific with Cranaothus deforgesi. Only a 
re-examination of their specimen (suppos- 
edly in the National Museum of the Phil- 
ippines) will resolve this matter with cer- 
tainty. It is clear, however, that Cranaothus 
deforgesi is not conspecific with Parame- 
daeus noelensis (Ward, 1934). 

As regards Paramedaeus noelensis (Ward, 
1934), the species was described from 
Christmas Island, Indian Ocean, by Ward 
(1934:17, pl. 1 fig. 1) as Medaeus, but his 
descriptions are brief and his figures rather 
schematic. Forest & Guinot (1961:56, pl. 1 
fig. 1, text figs. 42, 43, 44a, b) redescribed 
and refigured the species after examining the 
type and additional specimens from Upolu 
and Tahiti. Guinot (1967) subsequently 
proposed transferring the species to a new 
genus, Paramedaeus, and this was followed 
by Sakai (1976) and Seréne (1984). The spe- 
cies is known from Christmas Island, Mad- 
agascar, Upolu, Tahiti, Japan and the Phil- 
ippines. However, there are doubts as to 


whether all the specimens of this species 
reported are conspecific. Certainly, Sakai’s 
(1976:426, fig. 224) descriptions and figures 
of the species differ from those by Forest & 
Guinot (1961) in having a shorter front, 
presence of distinct anterolateral teeth, and 
the smoother carapace. 

Unlike Cranaothus deforgesi, the cara- 
pace surface of Paramedaeus noelensis is 
smoother and has no granulated vermicu- 
lations; the carapace regions are more dis- 
tinct; the front is not lamellar in appearance, 
is less produced and lacks the deep median 
fissure; the posterolateral margin is almost 
straight to slightly convex (distinctly con- 
cave in C. deforgesi); the outer surfaces of 
the carpus of the cheliped are less rugose; 
the sternal structure has a distinctly wider 
space between sternal sutures 2 and 3, and 
3 and 4; the second male abdominal seg- 
ment has no transverse ridges; the lateral 
edges of the fused male abdominal segments 
three and four have a distinct deep cleft on 
each side (entire and continuous in C. de- 


712 


forgesi); the last male abdominal segment 
is distinctly triangular in shape (not round- 
ed); and the distal part of the G1 is not 
produced into a long, slender projection (see 
Forest & Guinot 1961:56, figs. 42, 43, 44a, 
b, pl. 1 fig. 1; Seréne 1984: fig. 51, pl. 12F). 

Paramedaeus noelensis and Cranaothus 
deforgesi seem to be closely related. The car- 
apace of P. noelensis is remarkably similar 
to that of C. deforgesi in shape and general 
armature, the posterior part of the anterior 
lateral margin in both species is almost 
straight and parallel with the median axis 
of the animal. This confers a rather quadrate 
appearance to the carapace of both species. 
The general form of the G1s in both species 
is also similar. Differences in the form of 
the male abdomen and sternum, however, 
suggest that it might be premature to trans- 
fer P. noelensis to Cranaothus, at least for 
the time being. 

The unusual molariform basal cutting 
tooth on the dactylus of the cheliped is rem- 
iniscent of that in crabs of the genus Ca- 
lappa (Calappidae) which is used for “‘peel- 
ing” gastropod shells (Shoup 1956, Ng & 
Tan 1984). In these crabs, the right cheliped 
is almost always the larger one and pos- 
sesses the basal cutting tooth. Ng & Tan 
(1985) suggested that this was because ma- 
rine gastropods have dextral coiling. The 
well developed condition of the cutting tooth 
in Cranaothus strongly suggests that the crab 
is also a “‘peeler”’ like Calappa. Interesting- 
ly, in the specimen recorded by Seréne & 
Umali (1972) (as Paramedaeus noelensis) 
from the Philippines, the right chela is also 
the larger and has a basal cutting tooth. 


Acknowledgments 


The author is very grateful to A. Crosnier 
and B. Richer de Forges (ORSTOM) for for- 
warding the material to him for study, and 
their kind help. A. Crosnier, D. Guinot, P. 
Clark, A. B. Williams and R. B. Manning 
kindly read through the manuscript, and 
their many useful suggestions and criticisms 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


have improved the paper substantially. The 
study has been partially supported by RP 
900360 from the National University of 
Singapore. 


Literature Cited 


Alcock, A. 1898. Materials for a carcinological fauna 
of India. No. 3. The Brachyura Cyclometopa. 
Part I. The Family Xanthidae.— Journal of the 
Asiatic Society of Bengal 67(2)(1):67—233. 

Forest, J., & D. Guinot. 1961. Crustacés Décapodes 
Brachyoures de Tahiti et des Tuamotu. Jn Ex- 
pédition francaise sur les récifs coralliens de la 
Nouvelle-Calédonie. Volume préliminaire. Par- 
is, ditions de la Fondation Singer-Polignac, ix— 
xi + 1-194, figs. 1-178, pls. 1-18, tabls. 1-3, 7 
maps. 

Guinot, D. 1967. Recherches préliminaires sur les 

groupements naturels chez les Crustacés Déca- 

podes Brachyoures. II. Les anciens genres M1- 
cropanope Stimpson et Medaeus Dana.—Bul- 
letin du Muséum national d’Histoire naturelle, 

Paris (2)39(2):345-374. 

1968. Recherches préliminaires sur les 
groupements naturels chez les Crustacés Déca- 
podes Brachyoures. IV. Observations sur quel- 
ques genres de Xanthidae.—Bulletin du Mu- 
séum national d’Histoire naturelle, Paris (2)39(4): 
695-727. 

. 1978. Principes d’une classification évolutive 

des Crustacés Décapodes Brachyoures.— Bulle- 

tin du biologique Francaise et Belgique, new se- 
ries 112(3):211-292. 

. 1979. Données nouvelles sur la morphologie, 

la phylogenése et la Crustacés Décapodes 

Brachyoures.— Mémoires du Muséum national 

d’Histoire naturelle, Paris (A) Zoology 112:1- 

354, pls. 1-27. 

Haswell, W. A. 1882. On some new Australian 
Brachyura.— Proceedings of the Linnean Soci- 
ety of New South Wales 6(3):540-551. 

Klunzinger, C. B. 1913. Die Rindkrabben (Cyclo- 
metopa) des Roten Meeres.—Nova Acta Leo- 
pold Carolia 99(2):97-402, pls. 5-11. 

MacLeay, W.S. 1838. Illustrations of the Annulosa 
of South Africa; being a portion of the objects 
of natural history chiefly collected during an ex- 
pedition into the interior of South Africa, under 
the direction of Dr. Andrew Smith, in the years 
1834, 1835, and 1836; fitted out by the “Cape 
of Good Hope Association for Exploring Cen- 
tral Africa.” In A. Smith, ed., Illustrations of 
the zoology of South Africa investigations, Lon- 
don, Smith, Elder and Co., pp. 1-75, pls. 1-4. 

Milne Edwards, A. 1867. Descriptions de quelques 
espéces nouvelles de Crustacés Brachyures.— 


VOLUME 106, NUMBER 4 


Annales de la Sociéte entomologique Francaise 

(4)7:263-288. 

1873-1881. Etudes sur les Xiphosures et les 
Crustacés de la région Mexicaine. Jn Mission 
scientifique du Mexique et dans l’Amérique cen- 
trale, Recherches Zoologiques pour servir a 
Vhistoire de la faune de l’Amérique centrale et 
du Mexique, 5:1—368, pls. 1-61. 

Ng, P.K.L.,& L.W.H.Tan. 1984. The ‘shell peeling’ 
structure of the box crab Calappa philargius (L.) 
and other crabs in relation to mollusc shell ar- 
chitecture.—Journal of the Singapore National 
Academy of Science 13:195-199. 

——., & 1985. ‘Right Handedness’ in het- 
erochelous calappoid and xanthoid crabs—sug- 
gestion for a functional advantage.—Crusta- 
ceana 49:98-100. 

Odhner, T. 1925. Monographie Gattungen der Krab- 
benfamilie Xanthidae. I.—G6teborgs Kungl Ve- 
tenskaps-och Vitterhets-Samhdalles Handlingar 
(4)29(1):10-92, pls. 1-5. 

Rathbun, M. J. 1898. The Brachyura collected by the 
United States Fish Commission steamer Alba- 
tross on the voyage from Norfolk, Virginia, to 
San Francisco, California, 1887-1888.—Pro- 
ceedings of the United States National Museum 

21:567-616, pls. 41-44. 

. 1907. Report on the Brachyrhyncha, Oxysto- 

mata and Dromiacea. Jn Report on the crabs 

obtained by F.I.S. “Endeavour” on the coasts 
of Queensland, New South Wales, Victoria, 

South Australia and Tasmania, Biological Re- 

sults of the Fishing Experiments carried on by 

the F.I.S. “Endeavour” 1909-14, Sydney 5(3): 

95-156, pls. 16-42. 

. 1930. The cancroid crabs of America. — Unit- 

ed States National Museum Bulletin 152:i-xvi 

+ 1-609, figs. 1-85, pls. 1-230. 


TES 


Sakai, T. 1976. Crabs of Japan and the adjacent seas. 
In three volumes; English Text, pp. xxix + 773 
pp., Japanese Text, pp. 1-461, Plates Volume, 
pp. 1-16, pls. 1-251. Kodansha Ltd., Tokyo. 

Seréne, R. 1984. Crustacés Décapodes Brachyoures 
de l’Océan Indien occidental et de la Mer Rouge. 
Xanthoidea: Xanthidae et Trapeziidae. Adden- 
dum Carpiliidae et Menippidae—A. Cros- 
nier.—Faune Tropicale (ORSTOM) 24:1-400, 
pls. 1-48. 

——.,, & A. F. Umali. 1972. The family Raninidae 
and other new and rare species of Brachyuran 
Decapods from the Philippines and adjacent 
regions. — Philippine Journal of Science 99(1—2): 
21-105, pls. 1-9. 

——, & C. Vadon. 1981. Crustacés Décapodes: 
Brachyoures. List préliminaire, description de 
formes nouvelles et remarques taxonomiques. 
Résultats des Campagnes MUSORSTOM. I. 
Philippines. — Collection Mémoires ORSTOM, 
Paris 91:117-140, pls. I-III. 

Shoup, J.B. 1956. Shell opening by crabs of the genus 
Calappa.—Science 160:887-888. 

Ward, M. 1934. Notes on a collection of crabs from 

Christmas Island, Indian Ocean.—Bulletin of 

the Raffles Museum 9:5—28, pls. 1-3. 

1942. Notes on the Crustacea of the Desjar- 
dins Museum, Mauritius Institute, with descrip- 
tions of new genera and species. — Mauritius In- 
stitute Bulletin 2(2):49-113, pls. 5, 6. 


Department of Zoology, National Uni- 
versity of Singapore, Kent Ridge, Singapore 
0511, Republic of Singapore. 


PROC. BIOL. SOC. WASH. 
106(4), 1993, pp. 714-718 


A NEW DEEP-SEA CRAB OF THE 
GENUS CHACEON FROM INDIA 
(CRUSTACEA: DECAPODA: GERYONIDAE) 


H. C. Ghosh and Raymond B. Manning 


Abstract. —Chaceon alcocki, a species with laterally compressed dactyli on 
the walking legs, is described from a female identified with the European Geryon 
affinis Milne Edwards & Bouvier, 1894 by Alcock in 1899. It is the sixth species 
of the genus to be recognized from the western Indian Ocean. 


Until recently, several species of geryonid 
crabs from different localities had been 
identified with either Geryon quinquedens 
Smith, 1879 or with Geryon affinis Milne 
Edwards & Bouvier, 1894; both of these 
species were transferred by Manning & Hol- 
thuis (1989) to the genus Chaceon. Of the 
24 species of Chaceon now recognized, 14 
at one time in the past had been identified 
with either C. quinquedens or with C. affinis, 
and six species had been identified with both 
C. quinquedens and C. affinis (Manning & 
Holthuis 1988, 1989). In their revision of 
the geryonids, Manning & Holthuis (1989: 
76) pointed out that the status of a species 
identified with Geryon affinis by Alcock 
(1899) needed to be determined. That spe- 
cies, which proved to be new, is described 
below. 

The holotype is in the Zoological Survey 
of India, Calcutta. Measurements are in 
millimeters. Abbreviations used in the ac- 
count below include: cb, carapace breadth; 
cl, carapace length; fm, fathoms; m, meters; 
P5, fifth leg (fourth walking leg). 


Chaceon alcocki, new species 
Figs. 1-3 


Geryon affinis.— Alcock, 1899:85 [not Ge- 
ryon affinis Milne Edwards & Bouvier, 
1894]. 


Holotype. —1°, cl 110.5, cb 126.5; India, 
Travancore coast, 8°37'N, 75°37'30”E; depth 


224-284 fm (410-520 m); sand; Investiga- 
tor station 248; 17 Oct 1898; Zoological 
Survey of India Registry Number C 2907/ 
10. 

Diagnosis. —A large Chaceon, cl to more 
than 110 mm, cb to more than 126 mm, 
with low, obtuse anterolateral teeth on the 
carapace and laterally compressed dactyli 
on the walking legs. Carapace 1.1 times 
broader than long, moderately inflated at 
protogastric regions, convex from front to 
back. Frontal teeth short and blunt, median 
extending further forward than laterals. Dis- 
tance from first to second anterolateral tooth 
subequal to distance from third to fourth, 
distance from first to third tooth equal to 
distance from third to fifth. Carapace largely 
smooth, surface pitted behind front and on 
branchial regions. Orbits shallow, with deep 
incision on posterior margin. Suborbital 
tooth low, obtusely pointed, falling short of 
apices of lateral frontal teeth. Outer angle 
of merus of third maxilliped not projecting 
laterally. Cheliped merus with sharp sub- 
distal spine dorsally, distal spine absent; 
carpus relatively smooth dorsally, lacking 
outer spine or angled projection, inner spine 
well developed; propodus lacking distal 
dorsal spine. Meri of walking legs unarmed 
dorsally. Dactyli of walking legs laterally 
compressed, height at midlength greater than 
width, tips of dactyli blunt, worn off. P5: 
merus 4.5 times longer than high; carpus 
with at most a line of low tubercles dorsally; 


VOLUME 106, NUMBER 4 715 


Fig. 1. Chaceon alcocki, new species. Female holotype, cl 110.5 mm: a, Dorsal view; b, Carapace; c, Buccal 
area. 


716 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Chaceon alcocki, new species. Female holotype, cl 110.5 mm: a, Orbit, ventral view; b, Dactylus of 
P5, dorsal view; c, Dactylus of P5, lateral view. 


VOLUME 106, NUMBER 4 


F417) 


Fig. 3. 
PS, dorsal view; d, dactylus of P5, lateral view. 


propodus 5.0 times longer than high; dac- 
tylus 0.8 times as long as propodus. 

Size. —Unique female holotype, cl 110.5, 
cb 126.5. Measurements of P5: merus length 
60.0, height 13.2; propodus length 50.0, 
height 10.0; dactylus length 40.5, height 4.5, 
width 3.2. 

Color.—Brown. Alcock (1899:85) com- 
mented that “The colours in life are ad- 
mirably shown in the figure given by MM. 
Milne Edwards and Bouvier.” 

Remarks.— Manning (1992) pointed out 
that five species of Chaceon had been re- 
corded from localities in the western Indian 
Ocean. Two of these, C. paulensis (Chun, 
1903) and C. collettei Manning, 1992, differ 
from C. alcockiin having long, sharp frontal 
and anterolateral spines on the carapace and 
longer, slenderer walking legs with a distal 
dorsal spine on the merus of at least the 
posterior three legs. Chaceon alcocki resem- 


Chaceon alcocki, new species. Female holotype, cl 110.5 mm: a, P5; b, Carpus of PS; c, Dactylus of 


bles C. chuni (Macpherson, 1983) and C. 
crosnieri Manning & Holthuis, 1989, and 
differs from C. macphersoni (Manning & 
Holthuis, 1988) in having laterally com- 
pressed dactyli on the walking legs, but C. 
alcocki differs from both C. chuni and C. 
crosnieri in lacking distal dorsal spines on 
the meri of the walking legs. 

Etymology. —Named for Alfred William 
Alcock, 23 June 1859-24 March 1935, En- 
glish surgeon and naturalist, Superintendent 
of the Indian Museum from 1893 to 1907, 
one of the pioneers in carcinological re- 
search in India. 

Distribution. —Known only from the type 
locality. 


Acknowledgments 


We thank Prof. M. S. Jairajpuri, Director 
of the Zoological Survey of India, for his 


718 


support of this project. Lilly King Manning 
prepared the figures for publication. 


Literature Cited 


Alcock, A. 1899. An account of the deep-sea Brachy- 
ura collected by the Royal Indian Marine Survey 
Ship Investigator. Indian Museum, Calcutta, pp. 
1-85, 1-2, pls. 1-4. 

Chun, C. 1903. Aus den Tiefen der Weltmeeres, 2nd 
edition. Gustav Fischer, 592 pp. 

Macpherson, E. 1983. Crustaceos decapodos captu- 
rados en las costa de Namibia. — Resultados Ex- 
pediciones Cientificas (supplement to Investi- 
gacion Pesquera, Barcelona) 11:3-80. 

Manning, R. B. 1992. A new geryonid crab from 
Walters Shoals, southwestern Indian Ocean 
(Crustacea: Decapoda: Brachyura).—Proceed- 
ings of the Biological Society of Washington 105: 
86-89. 

—,&L.B.Holthuis. 1988. South African species 
of the genus Geryon (Crustacea, Decapoda, Ger- 
yonidae).— Annals of the South African Muse- 
um 98(3):77-92. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


——. & 1989. Two new genera and nine 
new species of geryonid crabs (Crustacea, De- 
capoda, Geryonidae).— Proceedings of the Bi- 
ological Society of Washington 102:50-—77. 

Milne Edwards, A., & E.-L. Bouvier. 1894. Brachy- 
oures et anomoures. Crustacés décapodes prov- 
enant des campagnes du yacht I’ Hirondelle (1886, 
1887, 1888). Premiére partie. — Résultats des ca- 
pagnes scientifiques accomplies sur son yacht 
par Albert I", Prince Souverain de Monaco 7:1- 
PU2: 

Smith, S. I. 1879. The stalk-eyed crustaceans of the 
Atlantic coast of North America north of Cape 
Cod.—Transactions of the Connecticut Acade- 
my of Arts and Sciences 5(1):27—36. 


(HCG) Zoological Survey of India, 27 Ja- 
waharlal Nehru Road, Calcutta 700 016, 
India; (RBM) Department of Invertebrate 
Zoology, National Museum of Natural His- 
tory, Smithsonian Institution, Washington, 
D.C..20560;Ws- A: 


PROC. BIOL. SOC. WASH. 
106(4), 1993, pp. 719-727 


CAMBARUS (JUGICAMBARUS) SUBTERRANEUS, A 
NEW CAVE CRAYFISH (DECAPODA: CAMBARIDAE) 
FROM NORTHEASTERN OKLAHOMA, WITH A KEY 

TO THE TROGLOBITIC MEMBERS OF THE 
SUBGENUS JUGICAMBARUS 


H. H. Hobbs III 


Abstract.—Cambarus (Jugicambarus) subterraneus, a new albinistic, trog- 
lobitic, crayfish, is described from three cave stream habitats in Delaware 
County, northeastern Oklahoma. Its closest affinities are with four other trog- 
lobitic species occurring in the Ozarks, particularly with C. (J.) aculabrum and 
C. (J.) setosus, all of which are allopatric. 


As early as 1951, a population of trog- 
lobitic crayfishes of the genus Cambarus had 
been known from a “Cave [=Twin Cave] 
between Spavinaw and Jay, Delaware Co.,”’ 
Oklahoma (Hobbs & Barr 1960:27). This 
obligate cavernicole was tentatively iden- 
tified by Hobbs & Barr (1960) as Cambarus 
(Jugicambarus) setosus Faxon (1889:237). 
They realized that it was not a typical pop- 
ulation of individuals and that there were 
“reasons to question this [species] desig- 
nation” (see also Hobbs et al. 1977). In May 
1972 I collected a troglobitic Form I male 
from Twin Cave and also recognized that it 
exhibited atypical setosus features. Field in- 
vestigations were conducted from Septem- 
ber 1989 to spring 1992 by the Oklahoma 
Biological Survey (Mehlhop-Cifelli 1990, 
Vaughn & Certain 1992) which revealed two 
additional populations of this species in Jail 
and Star caves. Based on data from DNA 
analyses by Koppleman (1990) and on those 
of morphological investigations conducted 
by me on a limited number of available 
specimens, these populations are consid- 
ered an undescribed species designated 
herein as Cambarus (Jugicambarus) subter- 
raneus. This is the thirty-first troglobitic 
crayfish to be named from North America 
(north of Mexico); the second obligate cave 
crayfish to be discovered from Oklahoma; 


and the eleventh and sixth, respectively, of 
the genus Cambarus and of the subgenus 
Jugicambarus, known to be restricted to hy- 
pogean waters of the United States (Hobbs 
III 1993). 


Cambarus (Jugicambarus) subterraneus, 
new species 
Figs..1.. 2a, b, ¢; k: o,-p, Table 


Diagnosis. —Albinistic; eyes reduced, 
carapace subcylindrical. Body and pereio- 
pods bearing conspicuous stiff setae. Ros- 
trum broadest at base, with small marginal 
spines symmetrically to slightly asymmet- 
rically situated, lacking median carina; ros- 
trum reaching base of ultimate podomere 
of antennular peduncle (reaching proximal 
to ultimate podomere in small individuals). 
Carapace lacking cervical spines; postorbit- 
al ridges depressed and terminating ce- 
phalically in weak tubercles. Areola 12.2- 
18.7 times as long as broad and constituting 
40.9-42.7% of total carapace length (47.6- 
49.8% of postorbital carapace length) and 
with 1 or 2 punctations in narrowest part. 
Antennal scale about 1.9 times longer than 
wide, broadest distal to midlength. Chela of 
first pereiopod with moderately inflated 
palm bearing 17-24 tubercles scattered in 
an irregular mesial arrangement of two 


720 


somewhat distinct rows; dorso-longitudinal 
ridges of fingers moderately well developed. 
Hooks on ischia of third pereiopods of males 
compressed, not reaching basioischial artic- 
ulation, and not opposed by tubercles on 
basis. Caudomesial boss on coxae of fourth 
pereiopods. First pleopods of male sym- 
metrical, not continguous at base, termi- 
nating in two parts (central projection and 
mesial process), both recurved at angles 
greater than 90°. Central projection mod- 
erately long and slender, corneous, and with 
well defined subapical notch; mesial process 
with broad base not greatly inflated and with 
distal third tapering and projecting 108—123° 
to shaft of pleopod; proximolateral lobe of 
gonopod not set off from shaft by groove. 
Annulus ventralis subsymmetrical in out- 
line, caudal part slightly movable; cephalic 
half traversed by deep submedian longitu- 
dinal trough; sinus originating on right cau- 
dolateral side of trough, crossing median 
line, continuing to fossa, crossing median 
line, and ending on caudal wall of annulus. 

Holotypic male, form I.—Cephalothorax 
subovate in cross section, markedly de- 
pressed (Fig. 1a, j). Abdomen narrower than 
thorax (9.7 and 12.5 mm in widest parts, 
respectively). Greatest width of carapace at 
slightly less than 0.2 length of areola from 
cephalic margin where width is greater than 
height (12.5 and 7.9 mm, respectively). Are- 
ola very narrow, 18.7 times as long as wide; 
length of areola 42.7% of entire length of 
carapace (49.8% of postorbital carapace 
length). Cephalic section of carapace 1.3 
times as long as areola length. Rostrum with 
convergent margins, slightly thickened; 
small corneous asymmetrical marginal 
spines delimiting base of corneous up-turned 
acumen extending cephalic to basal part of 
ultimate podomere of antennular peduncle; 
dorsal surface of rostrum excavate and bear- 
ing few, small setiferous punctations. Sub- 
rostral ridges weakly developed and evident 
in dorsal aspect for only short distance at 
base of rostrum. Postorbital ridges short, 
lacking spines, terminating cephalically in 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


very weak tubercles. Suborbital angle ab- 
sent; branchiostegal spine small but acute. 
Cervical spines absent. Surface of carapace 
weakly punctate dorsally and granulate lat- 
erally. 

Abdomen only slightly longer than car- 
apace. Epimeron of first abdominal segment 
barely overlaped by that of second; pleura 
of fourth through fifth abdominal segments 
rounded anteroventrally, subangular pos- 
teroventrally. Cephalic section of telson with 
single fixed spine in each caudolateral cor- 
ner, smaller spine immediately mesial to 
each and movable; mesial ramus of uropod 
with distolateral spine falling short of distal 
margin. 

Anteromedian lobe of epistome (Fig. 1k) 
broader than long with cephalomedian de- 
pression, elevated (ventrally) margins with 
several distinct projections. Antennule of 
usual form with prominent submedian spine 
on ventral surface of basal segment distal 
to midlength. Antennae extending caudad 
well beyond caudal margin of telson. An- 
tennal scale (Fig. 11) 1.9 times as long as 
broad, greatest width distal to midlength 
with lamellate portion suddenly broadened 
in distal half; heavy lateral portion termi- 
nating cephalically in moderately long spine 
slightly overreaching tip of acumen. 

Right chela (Fig. 11) almost 4 times as long 
as wide, subovate in cross section and palm 
somewhat inflated. All surfaces with seti- 
ferous punctations bearing conspicuously- 
long setae. Inner margin of palm with about 
24 tubercles arranged roughly in two stag- 
gered rows; ventral surface of palm with 
longitudinal row of widely spaced subme- 
dian tubercles increasing in size distally, two 
distalmost ones corneous; lateral margin 
with row of low tubercles along basal half, 
otherwise punctate. Fingers provided with 
moderately well defined ridges dorsally and 
ventrally. Opposable margin of fixed finger 
with row of 15 tubercles situated more 
densely in proximal fourth, those in distal- 
most segment corneous tipped; seventh tu- 
bercle from base largest; single longitudinal 


VOLUME 106, NUMBER 4 


721 


= 2 
ee = 


Fig. 1. Cambarus (J.) subterraneus (a, b, f-1 from holotype; c, e from morphotype, and d from allotype): a, 
Lateral view of cephalothorax; b, c, Mesial view of first pleopod; d, Annulus ventralis; e, f, Lateral view of first 
pleopod; g, Ventral view of basal podomeres of third and fourth pereiopods; h, Caudal view of first pleopods; 
i, Right antennal scale; j, Dorsal view of carapace; k, Epistome; 1, Dorsal view of distal podomeres of right 


cheliped. 


row of minute denticles extending entire 
length of finger. Opposable margin of dactyl 
with single row of 17 tubercles along prox- 
imal four-fifths of finger, interspersed with 
single row of minute denticles; third tuber- 
cle from base largest. 


Carpus (Fig. 11) of right cheliped longer 
than broad with shallow longitudinal fur- 
row on dorsal surface; with scattered setifer- 
ous punctations. Mesial surface with one 
large tubercle and smaller one ventrome- 
sially; smaller one distal to and another 


722 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Measurements (mm) of Cambarus (Jugicambarus) subterraneus, new species. 


Holotype Morphotype Paratype Paratype 
al all al al 
Twin Cave Twin Cave Twin Cave Star Cave 
Carapace 
Entire length 26.2 18.2 20.2 14.9 
Postorbital length 22:5 Ses L735 12.8 
Width 12.5 8.4 9:3 6.7 
Height 79 5.9 6.4 4.5 
Areola 
Width 0.6 0.9 0.5 0.5 
Length 11.2 7.6 8.5 6.1 
Rostrum 
Width 33 2.4 pF | 1.8 
Length 3.6 2.9 2.6 ee 
Right chela 
Length, palm (regen.) 
mesial margin 10.4 5.6 6.0 — 
Palm width 7.4 52 4.0 — 
Length, lateral margin 29.4 PSsh 16.6 — 
Dactyl length 18.2 8.9 9.7 — 
Abdomen 
Width 9.8 qe 13 5.0 
Length 28.1 14.1 2168 15.4 


proximal to large tubercle; lower mesiodis- 
tal margin with prominent spike-like tu- 
bercle; lower laterodistal margin with two 
pronounced flat projections, mesialmost 
opposing articular knob on palm; several 
small tubercles arranged in arc proximal to 
laterodistal tubercle. 

Merus of right cheliped with many small 
tubercles forming row on proximodorsal 
surface; lateral and mesial surfaces irregular 
with few small punctations and tubercles; 
lower mesial surface with row of 11 spike- 
like tubercles and lateral one of 7, distal two 
of which extending mesiodistally across dis- 
tal part of podomere. Ischium with ventral 
row of two small tubercles; otherwise weak- 
ly punctate. 

Hooks (Fig. 1g) on ischiopodites of third 
pereiopods only; hooks strong, simple, not 
reaching basioischial articulation and not 
opposed by tubercles on basis. Coxae of 
fourth pereiopods with moderately promi- 
nent, rounded caudomesial boss; coxae of 
fifth pereiopods with prominences. 


First pleopods (Fig. 1b, f) symmetrical, 
not contiguous at base, barely reaching cau- 
dal portion of coxae of third pereiopods 
when abdomen flexed, and terminating in 
central projection and mesial process. See 
Diagnosis for description. 

Allotypic female. —The only female spec- 
imen available (from Jail Cave) is badly dis- 
torted in preservation following a recent 
molt that occurred prior to capture and thus 
no description is attempted. The annulus 
ventralis (Fig. 1d) as described in “Diag- 
nosis.” 

Morphotypic male, form II.—Differing 
from holotype in following respects: small 
corneous marginal spines of rostrum sym- 
metrical; elevated margin of cephalic lobe 
of epistome with small median prominence, 
lateral ones very weak; inner margin of palm 
of left chela (both chelae regenerated but 
right particularly distorted) with 17 tuber- 
cles, proximal and distalmost ones spini- 
form; opposable margin of fixed finger with 
row of 12 corneous tubercles situated in 


VOLUME 106, NUMBER 4 


proximal two-thirds, absent from distal 
third, fifth tubercle from base largest; op- 
posable margin of dactyl with sixth tubercle 
from base largest; lower mesial surface of 
merus with row of 14 spike-like tubercles 
and lateral row of 6. Hook on ischium of 
third pereiopod smaller but otherwise sim- 
ilar to that of holotype. First pleopod (Fig. 
lc, e) with more robust terminal elements 
but both disposed similarly as in holotype. 

Type locality. —Twin Cave, Delaware 
County, Oklahoma (Choleta Quadrangle, T. 
23N, R. 22E), protected by The Nature Con- 
servancy. This is a large solution cave in 
Mississippian limestone (Boone Forma- 
tion—cherty limestone) with a recharge area 
of 6 km? (Aley & Aley 1990). Cambarus (J.) 
subterraneus occurs in the main pool, the 
visible portion is approximately 20 m in 
diameter with a maximum depth of 3-4 m. 
The substrate consists of clay covered with 
silt. This cave supports a diverse fauna 
(Black 1971) including a millipede, collem- 
bolan (Folsomia candida Willem), cricket 
(Ceuthophilus utahensis Thomas), dipteran, 
beetles Gncluding Scaphinotus elevatus), a 
bullhead catfish, blind Ozark Cavefish (Am- 
blyopsis rosae (Eigenmann)), Cave Sala- 
mander (Eurycea lucifuga Rafinesque), 
Grotto Salamander (Typhlotriton spelaeus 
Stejenger), Pickerel Frog (Rana palustris 
LeConte), two endangered bat species, My- 
otis grisescens Howell and M. sodalis Miller 
& Allen, and the Eastern Pipistrel Bat, Pip- 
istrellus subflavus Cuvier. 

Disposition of types.—The holotype, al- 
lotype, and morphotype are deposited in the 
National Museum of Natural History, 
Smithsonian Institution, Washington, D.C., 
(USNM) USNM 260249, 260250, and 
260251, respectively. Paratypes, consisting 
of 1 6 I from Twin Cave, 1 4 I from Star 
Cave, and 1 ¢ II from Jail Cave (see 
*“Range’’), are also deposited in the Smith- 
sonian Institution. 

Range. —This troglobitic crayfish is 
known from three caves in Delaware Coun- 
ty, Oklahoma. In addition to the type lo- 
cality, C. (/.) subterraneus is found in Jail 


723 


and Star caves. In Jail Cave (Choleta Quad- 
rangle, T. 23N, R. 22E) specimens were cap- 
tured in a single, 5 m diameter, mud-bot- 
tomed pool of unknown depth. Recharge 
area for this cave is 3.9 km? (Aley & Aley 
1990) and a stream is present but little of it 
can be accessed for sampling. Historically 
this privately owned cave was a maternity 
site for the Gray Bat (Myotis grisescens), and 
a diplopod, dipterans, Amblyopsis rosae, a 
salamander, and Rana palustris are known 
from this cave (see Black 1971). Star Cave 
(Jay Quadrangle, T.23N, R. 23E) isa stream 
cave with six small pools in which crayfish 
have been observed. Cave passage width is 
generally 3-4 m and stream depth is rarely 
greater than 1 m, usually less than 25 cm; 
substrate is gravel and mud. The cave should 
be entered with extreme caution since loose 
rock slabs occur in the crawlways. Black 
(1971:28) listed Typhlotriton spelaeus, from 
this locality, and Ceuthophilus utahensis, a 
beetle (Platynus sp.), and Amblyopsis rosae 
have been observed. 

Aley & Aley (1990) identified six poten- 
tially hazardous sites that would introduce 
contaminants into the groundwater system 
in the delineated recharge area for Twin 
Cave and five within the drainage basin for 
Jail Cave. They concluded that disposal of 
untreated animal wastes is probably the 
greatest single potential threat to aquatic life 
in these caves and recommended that man- 
agement attention should be focused on ef- 
forts to minimize groundwater quality im- 
pacts, mainly from large hog farms and/or 
poultry houses. 

Size. —The largest specimen is the holo- 
type from Twin Cave, having a carapace 
length of 26.2 mm (postorbital carapace 
length 22.5 mm). The smallest first form 
male has a carapace length of 11.5 mm. 
Mehlhop-Cefelli (1990: table 2) presented 
measurements for individuals of this spe- 
cies that were captured and released from 
these three known localities. These data are 
summarized in Table 2. 

Seasonal data.—First form males have 
been observed and/or collected from Twin 


724 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 2.— Measurements (mm) of Cambarus (Jugicambarus) subterraneus, new species, captured and released 
from the three Oklahoma caves from which it is known (data from Mehlhop-Cefelli 1990). 


Carapace Areola Chela 

Cave Sex Length Width Length Width Length Width 
Twin ) | 16.3 9.4 7.8 2a | 7.4 3.8 
él L72 8.1 9.4 ZS 6.0 5.3 

él 18.1 6.9 9.1 [2 4.7 2.6 

Q 14.0 6.0 7.4 1.4 _ _ 

Q 15.4 6.4 8.0 rs 6.0 2S 

Q 15.6 19 10.4 3.6 5.7 2.8 

Q 16.3 ial 8.6 12 6.0 pes) 

g 18.3 6.9 9.5 ZO 5-5 | es: 

Q | I >) 9.4 10.1 2.8 8.1 4.2 

g 20.5 8.0 10.2 22) 6.0 4.6 

Q 954 | 10.5 Piri SP 11.3 55 

Jail fo) | ley Da ye 1.0 4.5 2.4 
éll 116-5 6.9 8.5 1.6 6.0 3.4 

Q Es Dye) 7.0 je: 3.9 L.5 

Star | 18.6 Bs 8.0 1.4 4.9 p29) 
éll beli3 4.2 5.1 0.9 4.1 1.4 

éll 14.1 6.4 6.7 1.1 3.4 2 

éll 17.0 7.0 8.3 3 6.2 3.0 

2 13.6 4.9 Day) 0.9 5:3 2.6 

Q 16.5 kgs PA 12 5.7 2.8 

Q ey! 7.4 8.3 1.3 6.2 bP 

Q Dibred | 8.2 Lat — — 

g 21.8 9.0 8.5 1.6 8.1 4.6 

Q 23.4 PL.3 11.6 1.4 11.0 2 

Q 24.6 10.6 EES 3.0 10.1 4.8 


Cave on: 12 May 1972, 17 Aug 1975, 30 
Sept 1989; and from Star Cave on 28 Oct 
1989. Females with ova or young have not 
been observed. 

Variations. —Individuals from the three 
caves examined are relatively similar in most 
morphological features. Perhaps the antero- 
median lobe of the epistome (Figs. 1k, 20, 
p) is the most variable structure, being un- 
evenly and asymmetrically crenate but gen- 
erally truncate, broader than long, and nev- 
er in the form of an isosceles triangle as in 
C. VJ.) aculabrum Hobbs & Brown (1987). 
The number of tubercles on several podo- 
meres of the cheliped is variable as is the 
projected angle of the mesial process rela- 
tive to the shaft of the appendage. Angle of 
projection of mesial process relative to shaft 
of first pleopod is variable. Without addi- 


tional specimens from the three known lo- 
calities, no attempt is made to assess mor- 
phological variations among populations. 

Relationships. —The five troglobitic cray- 
fishes known from the Ozark Region are 
probably more closely allied to one another 
than any one is to other hypogean or epigean 
species (see Hobbs & Brown 1987 for a dis- 
cussion of the troglobitic members of the 
subgenus Jugicambarus and their possible 
origins). 

Cambarus (J.) subterraneus differs from 
other troglobitic members of the subgenus 
from the Ozarks only in a combination of 
features. Clearly it demonstrates close affin- 
ities with C. (J.) setosus and C. (J.) acula- 
brum and is more distantly related to C. (J/.) 
tartarus Hobbs & Cooper (1972) and C. (J/.) 
zophonastes Hobbs & Bedinger (1964), yet 


VOLUME 106, NUMBER 4 


cN | 
a b 


ahh? 


725 


Fig. 2. Epistome and secondary sexual features of troglobitic species of subgenus Jugicambarus (a—h, Mesial 
view of first pleopod of male, Form I: i-n, Caudal view of first pleopod of male, Form I; 0, p, Epistome): a, b, 
Paratypes of C. (J/.) subterraneus from Star and Twin caves, respectively; c, i, Holotype of C. (J.) zophonastes;: 
d, j, Holotype of C. (J.) aculabrum; e. k, Holotype of C. (J.) subterraneus: f, 1, C. (J.) setosus from Smallins 
Cave; g,m, Holotype of C. (J.) tartarus; h, n, Holotype of C. (J.) cryptodytes; 0, Morphotype of C. (J.) subterraneus; 
p. Paratype of C. (J.) subterraneus from Twin Cave (c, d. fj, I-n after Hobbs & Brown 1987). 


it shares morphological similarities with all 
species. As in C. zophonastes and C. acu- 
labrum, the proximolateral lobe of the first 
pleopod is not set off from the main shaft 
by a transverse groove; however, the new 
species can be distinguished from the for- 
mer by its shorter central projection and 
rostrum and from the latter by its slenderer 
central projection and more truncate epli- 
stome. As in C. tartarus, the central projec- 
tion of the first pleopod bears a distinct sub- 
apical notch but lacks a transverse groove 
and the epistome has a less ornate antero- 
median lobe. The epistome of C. setosus is 
similar to that of C. subterraneus yet the 
projections are more pronounced in C. se- 
tosus,; also, C. subterraneus has a pro- 
nounced subapical notch on the distal end 


of the central projection but lacks a trans- 
verse groove on the first pleopod. 

Results of electrophoretic analyses of 
troglobitic crayfishes from Arkansas, Mis- 
souri, and Oklahoma (Koppleman 1990, 
pers. comm.) demonstrate that the speci- 
mens from Jail, Star, and Twin caves in 
Delaware County, Oklahoma are genetical- 
ly similar to C. aculabrum in Benton Coun- 
ty, Arkansas. They are also genetically anal- 
ogous to C. setosus from southwestern 
Missouri, yet these Delaware County pop- 
ulations are dissimilar enough to be consid- 
ered a distinct species. Genetic similarities 
and differences are based on the use of Rog- 
ers (1972) distance measure. 

The following key should be useful in 
identifying first form males of the six trog- 


726 


lobitic species of the subgenus Jugicamba- 
rus. 


Key to the Troglobitic Members of the 

Subgenus Jugicambarus (Based on first 

pleopods of first form males— modified 
from Hobbs & Brown 1987) 


la. Central projection directed at 
right angle to shaft of pleopod 
(Fig é2hy a) ae ee ee 
Nesites eee C. (/.) cryptodytes Hobbs 
1b. Central projection directed 


caudally more than 90° to shaft 
of pleopod (Fig. 2a—g) 
Proximolateral lobe of shaft of 
pleopod set off from shaft by 
transverse groove (Fig. 21, m) 3 
2b. Proximolateral lobe of shaft of 
pleopod not set off from shaft 
by transverse groove (Fig. 2i— 
Ka) ose Ge ee ee 4 
Central projection short, 
heavy, not tapering, truncate 
apically (Fig. 2g, m) 
C. VJ.) tartarus Hobbs & Cooper 
3b. Central projection moderately 
long and tapering somewhat to 
rounded apex (very shallow 
subapical notch rarely devel- 
oped) (Fig. 2f, 1) 
af SUE C. (J.) setosus Faxon 
Central projection short and 
lacking subapical notch (Fig. 
De HR FPL IG A AT aes 
C. (J.) zophonastes Hobbs & Be- 
dinger 
Ab. Central projection moderately 
long bearing distinct subapical 
noteh (Fig. Za byds.c) 
Central projection relatively 
heavy (Fig. 2d, j); anterome- 
dian lobe of epistome in form 
of isosceles triangle and pro- 
duced anteriorly in acute or 
subacute apex 
a. a4 C. (J.) aculabrum Hobbs & 
Brown 


2a(1b). 


3a(2a). 


oe © © © © © we we 8 


4a(2b). 


5a(4b). 


©) 0; <9) jes ba! ne Je Ge, 6 wie) le 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Sb. Central projection relatively 
slender; anteromedian lobe of 
epistome generally truncate 
with several distinct projec- 
tions, not triangulate, and 
rarely produced anteriorly in 
acute or subacute apex (Fig. 2a, 


b, k, 0, p) ... C. VU.) subterraneus 


Etymology.—Cambarus (Jugicambarus) 
subterraneus takes its name from the Latin 
subter (below, beneath), alluding to its spe- 
lean existence. 


Acknowledgments 


I am most grateful to Jeffrey B. Kopple- 
man of the Missouri Department of Con- 
servation and to Caryn Vaughn and David 
L. Certain of the Oklahoma Biological Sur- 
vey for their continued assistance and en- 
couragement during this study. Apprecia- 
tion is extended to Horton H. Hobbs, Jr. of 
the Smithsonian Institution for encouraging 
me to conduct this study and for his valu- 
able comments and criticisms of the manu- 
script. 


Literature Cited 


Aley, T., & C. Aley. 1990. Hydrogeology of Ozark 
cavefish caves in Oklahoma. Final Report to 
The Nature Conservancy. Ozark Underground 
Laboratory, Protem, 70 pp. 

Black, J. H. 1971. The cave life of Oklahoma. A 
preliminary study (excluding Chiroptera).— 
Oklahoma Underground, Central Oklahoma 
Grotto 4(1, 2):2—53, 25 text fig., 8 figs. 

Faxon, W. 1889. Cambarus setosus Faxon. In Samuel 
Garman, Cave animals from southwestern Mis- 
souri. — Bulletin of the Museum of Comparative 
Zoology at Harvard College 17(6):237, pl. 1:figs. 
1—3- pl 2: fie. I 

Hobbs, H. H., Jr., & T. C. Barr, Jr. 1960. The origins 
and affinities of the troglobitic crayfishes of North 
America (Decapoda, Astacidae), I: The genus 
Cambarus.— American Midland Naturalist 64: 
12-33. 

—, & M.S. Bedinger. 1964. A new troglobitic 
crayfish of the genus Cambarus (Decapoda, As- 
tacidae) from Arkansas with a note on the range 
of Cambarus cryptodytes Hobbs.— Proceedings 


VOLUME 106, NUMBER 4 


of the Biological Society of Washington 77(3): 
9-15. 

—.,&A.V.Brown. 1987. A new troglobitic cray- 

fish from northwestern Arkansas (Decapoda: 

Cambaridae).— Proceedings of the Biological 

Society of Washington 100:1040-1048. 

, & M. R. Cooper. 1972. A new troglobitic 

crayfish from Oklahoma (Decapoda: Astaci- 

dae).— Proceedings of the Biological Society of 

Washington 85(3):49—56. 

—, H. H. Hobbs III, & M. A. Daniel. 1977. A 
review of the troglobitic crustaceans of the 
Americas.—Smithsonian Contributions to Zo- 
ology 244:v + 183 pp. 

Hobbs, III, H. H. 1993. Biogeography of subterra- 
nean decapods in North and Central America 
and the Caribbean region (Caridea, Astacidea, 
Brachyura). — Hydrobiologia (in press). 

Koppleman, J. B. 1990. A biochemical genetic anal- 
ysis of troglobitic crayfish (Cambarus spp.) in 
Missouri, Oklahoma, and Arkansas.— Report 
for the Missouri Department of Conservation, 


1p 


Oklahoma Natural Heritage Inventory, and Ar- 
kansas Game and Fish Commission. Missouri 
Department of Conservation, Columbia, 13 pp. 

Mehlhop-Cifelli, P. 1990. A survey and species de- 
terminations of cave crayfish (Cambarus spp.) 
in Oklahoma.—Report for the Oklahoma De- 
partment of Wildlife Conservation, Norman, 68 
pp. 

Rogers, J.S. 1972. Measures of genetic similarity and 
genetic distance. — Studies in Genetics, Univer- 
sity of Texas Publication 7213:145-153. 

Vaughn, C. C., & D. L. Certain. 1992. Survey and 
species determination of cave crayfish (Cam- 
barus spp.) in Oklahoma.—Final report for the 
Oklahoma Department of Wildlife Conserva- 
tion, Norman, 8 pp. 


Department of Biology, Wittenberg Uni- 
versity, P.O. Box 720, Springfield, Ohio 
45501-0720, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(4), 1993, pp. 728-739 


LITTORAL COMPOUND ASCIDIANS (TUNICATA) 
FROM SAO SEBASTIAO, ESTADO DE 
SAO PAULO, BRAZIL 


Sergio de Almeida Rodrigues and Rosana Moreira da Rocha 


Abstract. 


—Compound ascidians were collected during a two-year survey of 


intertidal and shallow subtidal habitats near Sao Paulo, Brazil. Among the 
fourteen species recorded, Distaplia bermudensis and Botryllus giganteum are 
new to Sao Sebastiao, and Didemnum psammathodes, Trididemnum orbicu- 
latum, and Symplegma rubra are new to South America. 


This paper describes species encountered 
in the Sao Sebastiao Channel (Fig. 1), near 
Sao Paulo, Brazil, during a study of recruit- 
ment, growth and reproduction of com- 
pound ascidians there (Rocha 1988, 1991). 
The specimens were collected in the inter- 
tidal and subtidal zones, by diving or during 
low tides, monthly from January 1985 to 
March 1987. Some specimens were taken 
from ceramic settling-plates, others from the 
natural rocky substrate and from under 
boulders. Not every species occurred at each 
locality. This may reflect differences in ecol- 
ogy or just collecting success. 

Ascidians from this region have been de- 
scribed by Van Name (1945), Bjornberg 
(1956), Millar (1958), Rodrigues (1962, 
1966, 1977) and Rocha (1988). Ascidians 
from other Brazilian waters have been de- 
scribed by Van Name (1945), Moure et al. 
(1954), Millar (1961, 1977), C. Monniot 
(1969/1970), Costa (1964, 1969a, 1969b, 
1969c, 1969d, 1969e, 1969f), and Simoes 
(1981). References to some older papers that 
mention Brazilian ascidians will be found 
in Van Name’s (1945) monograph. 

The synonymy of some species is exten- 
sive. We provide references to recent West 
Atlantic records of the species we recorded. 
For a more comprehensive but older list of 
synonymies see Van Name (1945). 

Of the fourteen species collected, Didem- 
num psammathodes (Sluiter, 1895), Tridi- 


demnum orbiculatum (Van Name, 1902), 
and Symplegma rubra C. Monniot, 1972 are 
new records for South America. Distaplia 
bermudensis Van Name, 1902 and Botryllus 
giganteum Aron & Sole-Cava, 1991 are re- 
ported for the first time from Sao Sebastiao. 
Didemnum speciosum (Herdman, 1886), 
Diplosoma listerianum (Milne-Edwards, 
1841), Symplegma brakenhielmi (Michael- 
sen, 1904), Botryllus tuberatus Ritter & For- 
syth, 1917, and Botryllus niger (Herdman, 
1886) were already recorded from Sao Se- 
bastiao under the names Didemnum can- 
didum Savigny, 1816, Diplosoma macdon- 
aldi Herdman, 1886, Symplegma viride 
Herdman, 1886, Botryllus primigenus Oka, 
1928, Botrylloides nigrum Herdman, 1886, 
respectively. Of the compound ascidians 
registered on the previous surveys of Sao 
Sebastiao, only Polyandrocarpa anguinea 
(Sluiter, 1898), recorded by Van Name 
(1945) and Rodrigues (1962) under the name 
Polyandrocarpa maxima (Sluiter, 1904), was 
not found in the present survey. 

Vouchers of the material examined have 
been deposited in the Museu de Zoologia, 
Universidade de Sao Paulo, Sao Paulo, Bra- 
zil (MZUSP), and in the National Museum 
of Natural History, Smithsonian Institu- 
tion, Washington, D.C., U.S.A. (USNM). 


Family Polyclinidae 


Polyclinum constellatum Savigny, 1816 


VOLUME 106, NUMBER 4 


TROPIC | OF _CAPRICORN 


SAO PAULO 


Oo 10 20 30km 
a 


ATLANTIC OCEAN 
46°w 


CEBIMar 33. 
PRAIA DO SEGREDOy= 


Fig. 1. 
Sao Paulo (CEBIMar). 


Polyclinum constellatum Savigny, 1816:189, 
pl. 4, fig. 2, pl. 18, fig. 1.—Van Name, 
1945:68, fig. 28, pl. 13, figs. 2-3. — Bjorn- 
berg, 1956:164.— Millar, 1958:498, 1962: 
62.—Rodrigues, 1962:194.—Costa, 
1969a:192, fig. 1.—F. Monniot, 1972: 
958, fig. 4, 1983b:417, pl. 1.—Simoes, 
1981:19, figs. 4-6.— Goodbody, 1984:29. 


Material examined. — Eight colonies from 
Praia do Araca, one from Praia Grande, and 
two from Praia do Cabelo Gordo de Dentro. 
One specimen in USNM (20037), two in 
MZUSP (11190, 11191). Colonies collected 
in October 1985 and 1986 and July 1986 
had well-formed gonads. October colonies 
had larvae. 

Distribution and habitat. —Distributed 
widely throughout warm waters. In the At- 
lantic: Florida, Mexico (Yucatan), Colom- 
bia (Sabanilla), Caribbean Islands, Bermu- 
da. In Brazil: Rio de Janeiro (Urca, Niter6oi) 
and Sao Paulo (Ubatuba, Sao Sebastiao, 
Santos, Cananeia). Colonies found in shal- 
low water, often exposed during very low 
tides. 


CANAL DE SAO SEBASTIAO 


729 


PRAIA DO CABELO aa 
GORDO DE DENTRO |-~ 


Map showing localities where specimens were collected and the marine station of the University of 


Family Didemnidae 
Didemnum psammathodes (Sluiter, 1895) 


Leptoclinum psammathodes Sluiter, 1895: 
Hk. 

Didemnum psammathodes: F. Monniot, 
1983a:31, fig. 13, pl. 2, figs. B and E.— 
Goodbody, 1984:30. 

Didemnum candidum: Van Name, 1945 (in 
part):83, fig. 35. 


Material examined. —Five colonies from 
Praia do Segredo, three from Praia do Ca- 
belo Gordo de Dentro, two from Praia do 
Araca, one from Praia Grande. One speci- 
men in USNM (20036), three in MSUSP 
(11193, 11194, 11195). 

External appearance. —Macroscopic 
structure of colonies closely resembles that 
of D. speciosum, differing only in color, 
which is brown or grayish-brown due to large 
accumulations of fecal pellets in common 
cloacal cavities and even in the substance 
of the test; these pellets absent around com- 
mon cloacal apertures, where tunic is trans- 
parent. Spicules scarce, more abundant 


730 


around branchial openings, small (5—30 um), 
nearly spherical with numerous rays having 
irregular tips. 

Internal structure.—Zooids 0.4 to 0.75 
mm long. Branchial aperture six-lobed, atri- 
al aperture exposing part of branchial sac. 
Testis undivided, spermiduct with six to 
eight spiral turns. Larvae bearing three pa- 
pillae and four pairs of ampullae, trunk 
length 0.4 to 0.5 mm, present in colonies in 
January. 

Remarks. —F. Monniot (1983a) recorded 
this species for the first time in the Carib- 
bean. Earlier, Van Name (1945), under D. 
candidum, described “‘colonies containing 
large accumulations of dark-colored fecal 
pellets,’ but he considered this “merely the 
result of some abnormal or pathological 
condition” attributable to the inability of 
the water current to carry off waste material. 
In Sao Sebastiao the species is very com- 
mon. It grows side by side with D. specio- 
sum and attains large sizes, so that a “‘patho- 
logical condition”’ seems highly unlikely. 

Distribution and habitat. —Recorded 
widely from tropical waters under many dif- 
ferent names (see Eldredge 1967). In the 
Atlantic: Guadeloupe, Curagao. In Brazil: 
Sao Paulo (Sao Sebastiao). Colonies com- 
mon intertidally, under boulders or on ver- 
tical surfaces, and growing over other as- 
cidians, sponges and bryozoans. 


Didemnum speciosum (Herdman, 1886) 


Leptoclinum speciosum Herdman, 1886: 
274, pl. 36, figs. 1-8. 

Didemnum speciosum: Millar, 1977:197, fig. 
20.—Simoes, 1981:29, figs. 14-17. 

Didemnum candidum: Van Name, 1945 Gn 
part):83, fig. 35, pl. 13, fig. 4.—Moure et 
al., 1954:235.—Bjornberg, 1956:164.— 
Rodrigues, 1962:194.—Costa, 1969b:202, 
fig. 4. 


Material examined. —Eleven colonies 
from Praia do Segredo, one from Praia do 
Cabelo Gordo de Dentro, two from Praia 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


do Araca, one from Praia Grande. One spec- 
imen in USNM (20033). 

External appearance. —Colonies very 
thin, usually about 1 mm thick; rarely more 
than 5 cm across. Color snow white or milky 
white. Spicules very abundant, chiefly in 
upper layer of test. Spicules mostly stellate 
with blunt tips, 5-40 um across, usually very 
regular but sometimes less regular and with 
only four to six rays. 

Internal structure. —Zooids 0.75 to 1 mm 
long. Branchial aperture six-lobed, atrial ap- 
erture a wide cleft, exposing much of bran- 
chial sac. Stigmata difficult to see clearly. 
Testis undivided; spermiduct with six or 
rarely seven spiral turns. Larvae with three 
papillae and four pairs of ampullae; larval 
trunk 0.35 mm long. Larvae present in col- 
onies from May to September. 

Remarks.—Van Name (1945) identified 
southern Brazilian didemnids like this one 
as Didemnum candidum. D. candidum’s 
presence in the Caribbean was contested by 
Millar (1962), who called the material from 
Curacao D. conchyliatum (Sluiter, 1898), as 
did F. Monniot (1983a) in identifying spec- 
imens from Guadeloupe. Millar (1977) res- 
urrected Herdman’s (1886) D. speciosum, 
an abundant didemnid from the northern 
and northeastern Brazilian shelf. He point- 
ed out that the only difference between D. 
speciosum and D. conchyliatum is the size 
of the larval trunk: 0.25—0.40 mm for the 
former species and 0.43—0.50 mm for the 
latter. Examination of specimens of D. con- 
chyliatum from Guadeloupe deposited in 
the National Museum of Natural History, 
Paris, showed that both zooids and larvae 
are slightly larger than the present material. 
The characteristics of our specimens are very 
similar to what Millar (1977) notes. Colors 
of the colonies, according to F. Monniot 
(1983a), also can be different. We believe 
that D. conchyliatum and D. speciosum are 
both valid species. 

Distribution and habitat.—The overall 


' distribution of Didemnum speciosum must 


remain uncertain until its extensive syn- 


VOLUME 106, NUMBER 4 


onymy is resolved. In Brazil: at least off the 
northern coast from Para to Sergipe, Bahia 
(type locality), Rio de Janeiro (Baia de Gua- 
nabara, Cabo Frio, Ilha Grande), Sao Paulo 
(Ubatuba, Sao Sebastiao), Parana (Baia de 
Paranagua), Santa Catarina (Florianopolis, 
Garopaba). Common intertidally and in 
shallow water, growing directly on rocky 
substrates or as an epibiont on barnacles, 
bivalves, bryozoans, polychaete tubes and 
solitary ascidians. 


Polysyncraton amethysteum 
(Van Name, 1902) 


Polysyncraton amethysteum Van Name, 
1902:366, figs. 62, 64-67. pl. LVIII, fig. 
102.—Rodrigues, 1962:195.— Millar, 
1977:200, fig. 21. 

Didemnum (Polysyncraton) amethysteum: 
Van Name, 1945:95, fig. 41, pl. 18, fig. 
3.—Moure et al., 1954:236.— Millar, 
1958:499.—Costa, 1969b:203, fig. 5. 

Didemnum amethysteum: Plough, 1978:67. 


Material examined. —Eight colonies from 
Praia do Segredo, two from Praia Grande. 
One specimen in USNM (20035), three in 
MZUSP (11204, 11205, 11206). Gonads 
present during most of the year. Larvae 
present in fall and spring specimens. 

Remarks. —Our specimens agree well with 
Van Name’s (1902, 1945) descriptions. The 
colonies, however, are larger, with more 
common cloacal openings, and the zooids 
with fewer testicular lobes. This last char- 
acter seems to be rather variable: five (Van 
Name 1902), five to six (Van Name 1945), 
four to six (Moure et al. 1954), four to five 
(Millar 1977). 

Distribution and habitat. — Bermuda (type 
locality), Puerto Rico, Colombia, Florida. 
In Brazil: Ceara, Pernambuco (Recife), Ba- 
hia, Rio de Janeiro (Rio de Janeiro, Cabo 
Frio), SAo Paulo (Sao Sebatiao, Cananéia), 
Parana (Baia de Paranagua), Santa Catarina 
(Garopaba). Colonies common intertidally, 
in crevices and abandoned sea-urchin holes, 


731 


under boulders, and among encrusting 
sponges, bryozoans and polychaete tubes. 


Trididemnum orbiculatum 
(Van Name, 1902) 
Figs. 2—5 


Didemnum orbiculatum Van Name, 1902: 
300, pl: 31, fies: 32,38) pl G1. fies: 127a, 
128. 

Trididemnum orbiculatum: Van Name, 
1945:103, fig. 47.—F. Monniot, 1983a: 
i2ehe. 3. pl. A. fie, A; 


Material examined.—Twenty-three col- 
onies from Praia do Cabelo Gordo de Den- 
tro, seven from Praia do Segredo, one from 
Praia do Araca, one from Praia Grande. One 
specimen in USNM (20039), two in MZUSP 
81202514203). 

External appearance. —Colonies encrust- 
ing of variable thickness (1-4 mm), up to 
40 mm across. Test firm, transparent, gen- 
eral color light or dark gray, yellowish, 
greenish or dun-colored. In some colonies 
a black spot is easily visible on the anterior 
end of the endostyle, and sometimes other 
darkened spots mark each of the six lobes 
of the zooids’ branchial apertures. Spicules 
usually of regular shape, 7-58 um across, 
abundant, lying in superficial layer of test. 
Spicules sometimes absent or scarce, of ir- 
regular shape, with striated rays and broken 
or divided tips (Fig. 2). 

Internal structure.—Zooids 0.6 to 0.85 
mm long (Fig. 3). Branchial siphon with six 
lobes; atrial siphon forms a short, smooth- 
margined tube, projecting from middle of 
thorax, usually directed backward. Round- 
ed lateral organs at level of second row of 
stigmata, distinguishable by aggregation of 
spicules around them. Testis undivided, 
surrounded by five to seven (usually six) 
coils of spermiduct (Fig. 4). Larval trunk 
0.4-0.6 mm long.with three papillae and 
four pairs of ampullae (Fig. 5). 

Remarks. — Millar (1962) identified spec- 
imens from Curacao as 7. orbiculatum. 
However, the irregularly shaped spicules and 


732 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 2-5. 
larva. 


the two-lobed testis of his specimens put his 
determination in doubt. Millar (1977) de- 
scribed northern Brazilian material as 77i- 
didemnum sp. The spicules (SO—-88 um) in 
that material are larger than in our speci- 
mens; the lack of mature gonads and larvae 
in Millar’s specimens precludes a definitive 
conclusion about their identity. F. Monniot 
(1983a), in her discussion of T. orbiculatum, 
points out that this species is very similar 
to T. tethidis Van Name, 1945, of which 
only immature Florida specimens are 
known, and that larvae of this last species 
are needed to confirm its identity. In our 
material we found some mature colonies 
that agree well with the description of T. 
tethidis. In these colonies, spicules are scarce 
or absent and zooids have a black spot on 
the anterior end of the endostyle. The larvae 
from these colonies are identical to larvae 
from the other T. orbiculatum colonies, sug- 
gesting that 7. tethidis and T. orbiculatum 
are indeed synonymous. The specimens 
from Guadeloupe (F. Monniot 1983a) seem 
to differ from the present material only in 
their more narrow range of spicule size and 


Trididemnum orbiculatum: 2, spicules; 3, immature zooid in lateral view; 4, zooid with testis; 5, 


in having a few more coils (eight) in their 
spermiducts. 

Distribution and habitat.— Bermuda (type 
locality), Curacao, Guadeloupe. In Brazil: 
Sao Paulo (Sao Sebastiao). Colonies under 
low intertidal boulders or on subtidal ce- 
ramic plates. 


Diplosoma listerianum 
(Milne-Edwards, 1841) 


Leptoclinum listerianum Milne-Edwards, 
1841:295. 

Diplosoma macdonaldi: Herdman, 1886: 
315, pl. 42, figs. 1-4.—Van Name, 1945: 
109, fig. 51, pl. 12, fig. 5.—Costa, 1969b: 
201, fig. 1.—Plough, 1978:67 fig. 29, pl. 
VIII.—Simoes, 1981:33, figs. 18-20.— 
Rocha, 1988:30, fig. 5. 

Diplosoma listerianum: Rowe, 1966:457, 
figs. 1-5; Millar, 1978:104.—F. Monniot, 
1983a:41, fig. 18.—Goodbody, 1984:31. 


Material examined.—Thirty-nine colo- 
nies from Praia do Segredo, two from Praia 
Grande, one from Praia do Araca. One spec- 
imen in USNM (20040), four in MZUSP 


VOLUME 106, NUMBER 4 


(11198, 11199, 11200, 11201). Gonads and 
larvae present in specimens from the whole 
year round. 

Remarks. —The type locality of D. mac- 
donaldi Herdman, 1886 is in Bahia. Van 
Name (1945) used this name for specimens 
from Sao Sebastiao. However, according to 
Rowe (1966), who examined the type spec- 
imen, and F. Monniot (1974, 1983a), who 
compared the specimens from the Antilles, 
Bermuda, Azores, and France, all reports of 
D. macdonaldi are in fact of D. listerianum: 
the species are synonymous. 

Distribution and habitat. —Bermuda, 
South Carolina, Florida, West Indies, Gui- 
ana Shelf. In Brazil: Bahia, Rio de Janeiro 
(Rio de Janeiro, Niteroi), Sao Paulo (Sao 
Sebastiao). Colonies in shallow water under 
boulders, sometimes attached directly to 
rocky substrate but more often growing on 
other organisms such as ascidians and bry- 
ozoans, and on artificial substrate (plastic, 
ceramic, or wood). 


Family Polycitoridae 


Distaplia bermudensis Van Name, 1902 


Distaplia bermudensis Van Name, 1902:349, 
pl. 49, figs. 15, 18, 19, pl. 59, figs. 108, 
111, pl. 62, fig. 130b. 

Distaplia bermudensis: Van Name, 1945: 
146, fig. 70, pl. 16, fig. 2.—Millar, 1958: 
500, fig. 3, 1962:68.—Costa, 1969c:279, 
fig. 3.—F. Monniot, 1972:960, fig. 4.— 
Millar, 1977:188, fig. 14.—Plough, 1978: 
60, fig. 27, pl. II, 1V.—F. Monniot, 1983c: 
1000, fig. 3. 


Material examined.—Eighteen colonies 
from Praia do Segredo, four from Praia 
Grande. One specimen in USNM (20032), 
fourin MZUSP (11209, 11210, 11211, 
1212); 

Distribution and habitat. — Bermuda (type 
locality), North Carolina, Florida, Antilles, 
Guadeloupe. In Brazil: along Para and Ma- 
ranhao coasts, Rio de Janeiro, Sao Paulo 
(Ubatuba, Sao Sebastiao). Colonies in shal- 
low water, on the undersurfaces of boulders. 


133 


Clavelina oblonga Herdman, 1880 


Clavelina oblonga Herdman, 1880:724. 

Clavelina oblonga: Van Name, 1945:136, 
figs. 63, 64, pl. 16, fig. 6.—Bjornberg, 
1956:165.—Millar, 1958:500.— Millar, 
1962:68.—Rodrigues, 1962:196.—Cos- 
ta, 1969c:277, fig. 1.—Plough, 1978:58, 
fig. 25, pl. II.—Simoes, 1981:36, figs. 21- 
23.—F. Monniot, 1983c:1000.—Rocha, 
1988:32, fig. 6. 


Material examined. —Twelve colonies 
from Praia do Cabelo Gordo de Dentro, two 
from Praia do Araca. One specimen in 
USNM (20034), two in MZUSP (11207, 
11208). Gonads present in colonies collect- 
ed from November to July. Larvae present 
from November to May. 

Distribution and habitat. — Bermuda (type 
locality), North Carolina to Florida, West 
Indies, West Africa. In Brazil: Rio de Ja- 
neiro (Niteroi, Rio de Janeiro), Sao Paulo 
(Ubatuba, Sao Sebastiao, Santos), Santa Ca- 
tarina (Florianopolis). Colonies in shallow 
water and intertidal, attached to vertical 
walls and to artificial substrates such as 
buoys, iron pilings, and ceramic plates. 


Family Styelidae 


Symplegma brakenhielmi 
(Michaelsen, 1904) 


Diandrocarpa brakenhielmi Michaelsen, 
1904:50. 

Symplegma brakenhielmi: C. Monniot, 
1983:429, fig. 3A—E. 

Symplegma viride. Van Name, 1945:232, 
figs. 139, 140c, 140d, pl. 18, fig. 2.—Cos- 
ta, 1969f:321 (part).—Rocha, 1988:35, 
He. 7. 


Material examined. —Twenty-three col- 
onies from Praia Cabelo Gordo de Dentro, 
one from Praia do Araca. One specimen in 
USNM (20029), three in MZUSP (11185, 
11186, 11187). Ripe gonads and larvae were 
found in summer and fall specimens. 

Remarks. —C. Monniot (1983) recogniz- 
es three species in the tropical West Atlan- 


734 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 6-7. Symplegma rubra: 6, group of living zooids in dorsal view (the coarser dots represent the red 


rings); 7, ventral side of a zooid. 


tic: Symplegma viride Herdman, 1886, S. 
brakenhielmi (Michaelsen, 1904), and S. ru- 
bra C. Monniot, 1972. There are several 
records from the Brazilian coast under the 
name S. viride. The accounts of Bjornberg 
(1956) and Millar (1958, 1977) do not pro- 
vide sufficient information for correct iden- 
tification of their material. Rodrigues (1962) 
mentions three populations, but without de- 
scription; Costa (1969f) gives a table with 
some characteristics of these populations, 
one of them having transparent colonies. 
Re-examination of Rodrigues’ (1962) ma- 
terial from this population showed less dense 
packing of zooids in the colony, pyriform 
or only slightly lobed testes, and no ovary. 
These differences may reflect immaturity of 
these colonies or be representative of an- 
other species. Additional material is re- 
quired to clarify how many Symplegma spe- 
cies are present at Sao Sebastiao. The 
material from Niter6i, Rio de Janeiro (Sim- 
oes 1981), is described as light or dark green 
with a coral-colored patch between the si- 
phons. This pattern and the outline of the 
testis (Simoes 1981, fig. 40) agree with C. 
Monniot’s (1983) interpretation of S. viride. 

Distribution and habitat. —Mexico (Vera 
Cruz: type locality), Florida, Puerto Rico, 
Bermuda, Guadaloupe. In Brazil: Rio de 
Janeiro (Ilha Grande), Sao Paulo (Sao Se- 


bastiao). Colonies in shallow water or in- 
tertidal, under boulders, on mollusc shells, 
and on artificial substrates such as buoys, 
ropes, iron pilings, and ceramic plates. 


Symplegma rubra C. Monniot, 1972 
Figs. 6—7 


Symplegma rubra C. Monniot, 1972:622, 
fig. 2, E-I, fig. 3; 1983:429, fig. 3F. 

Symplegma viride: Rodrigues, 1962:202 
(part).—Costa, 1969f:321 (part). 


Material examined. — Three colonies from 
Praia do Cabelo Gordo de Dentro, one from 
Praia do Segredo. One specimen in MZUSP 
(11188). 

External appearance. —Encrusting colo- 
nies about 2 mm thick and 10 cm across. 
Tunic rose with a bright pink ring linking 
both siphons. Zooids’ arrangement (Fig. 6) 
as in S. brakenhielmi. 

Internal structure. —Zooids up to 2.5 mm 
long. Branchial tentacles 16—18, of two sizes. 
Branchial sac with 12-13 rows of stigmata. 
Stomach with 16 external folds and a small, 
curved pyloric caecum. Gonads with very 
prominent and multilobed testis (Fig. 7); 
Ovaries absent from our specimens. 

Remarks. —The present material resem- 
bles S. rubra C. Monniot, 1972 in general 
color and in the shape of the testis, but the 


VOLUME 106, NUMBER 4 


conspicuous pink ring around the siphons 
has not been noted in the material from 
Bermuda and Guadeloupe (C. Monniot 
1972, 1983). Our zooids are smaller, and 
they have more branchial tentacles and more 
external stomach folds. The absence of ova- 
ries may be evidence of sequential her- 
maphroditism, which C. Monniot (1972) 
suggests may be typical of S. rubra. Re-ex- 
amination of Rodrigues’ (1962) material re- 
vealed only immature specimens. 

Distribution and habitat. —In Brazil: Rio 
de Janeiro (Ilha Grande), Sao Paulo (Sao 
Sebastiao). Colonies in shallow water, on 
rocks or artificial (ceramic) surfaces. 


Botryllus giganteum 
Aron & Sole-Cava, 1991 


Botryllus giganteum Aron and Sole-Cava, 
991-271. 

Metrocarpa nigrum var. Giganteum: Pérés, 
1949:205, figs. 26, 27. 

Botrylloides nigrum var. giganteum: C. 
Monniot, 1969:628, fig. 3.—Simoes, 
1981:54, figs. 35-37. 


Material examined. —Twelve colonies 
from Praia do Cabelo Gordo de Dentro, one 
colony from Praia Grande. One specimen 
in USNM (20038), five in MZUSP (11175, 
PPi76. 11177, 11178, 11179). 

External appearance.—Fleshy colonies, 
4-15 mm thick, irregularly rounded, 3-15 
cm in diameter, bright red or reddish orange 
overall. Tunic tough, somewhat rubbery, 
pale, translucent. Systems elongated, 
rounded or irregular. The greater the num- 
ber of zooids, the more irregular are the 
systems. 

Internal structure. —Zooids 2.5—3.5 mm 
long. Branchial aperture smooth and round. 
Atrial aperture a narrow horizontal slit in 
the young zooids, wide open; its anterior 
border produced into a broad and long lan- 
guet in older zooids. Branchial tentacles 24, 
of three sizes in fully developed zooids. 
Branchial sac with 11-17 (usually 14-16) 
rows of stigmata; each row with four or five 


735 


stigmata near endostyle and dorsal lamina, 
and two or three between lateral internal 
longitudinal vessels. Stomach with nine folds 
and a very small caecum. Anus bilobed and 
sometimes enlarged at the tip. Testis com- 
prises 5—15 rounded lobes, dorsal to ova- 
ries; ovaries hold only one egg each. Testis 
present throughout year but ovaries found 
only in July, October and November, in col- 
onies that had settled on ceramic plates. 

Remarks. —Aron & Sole-Cava (1991) 
raised the variety giganteum, described by 
Pérés (1949), to species status based upon 
specimens collected at Rio de Janeiro (Urca) 
and Espirito Santo (Vitoria). Our specimens 
agree with the description given by Simoes 
(1981) and Aron & Sole-Cava (1991, table 
4) except that our colonies are thinner and 
smaller, and our zooids have fewer rows of 
stigmata and gastric folds. The species seems 
to be rare. It was not found by Rodrigues 
(1962), and it was found only once on nat- 
ural substrate in the present survey. All our 
other colonies settled instead on experi- 
mental ceramic plates (Rocha 1988). 

Distribution and habitat.—Senegal (Da- 
kar: type locality). In Brazil: Espirito Santo 
(Vitoria), Rio de Janeiro (Niterdi, Rio de 
Janeiro). The only colony found on natural 
substrata was growing on the side of a boul- 
der, exposed during low tide. 


Botryllus niger (Herdman, 1886) 


Botrylloides nigrum Herdman, 1886:50, pl. 
1, fig. 8, pl. 3, figs. 19-21.—Van Name, 
1945:227, figs. 136c, 137.— Millar, 1962: 
71.—Rodrigues, 1962:201, pl. 2, figs. 5— 
7.—Costa, 1969e:300, fig. 1.—C. Mon- 
niot, 1972:618.— Plough, 1978:89, fig. 43: 
Simoes, 1981:51, figs. 32—34.—Good- 
body, 1984:36.—Rocha, 1988:38, fig. 8. 

Botryllus niger. Aron & Sole-Cava, 1991: 
274, t. 4. 


Material examined.—Thirteen colonies 
from Praia do Cabelo Gordo de Dentro, 
four from Praia do Segredo, one from Praia 
Preta, one from Praia do Araca. One spec- 


736 


fie 
, i) 
Wide ots 
Li { 
> 
i] 


OF ee ee 
~ 
rd 
? 
4s 
4 oie 
/ an 
Mw 
TD) 
7 min 
‘, 


Ps ae 
7 
/ 
/ 
] 
/) 
r re! 
a 
pb 
“/ "y 
yet fy 
1 i 
-/” ‘Ot 
No ee 


ae 

1 

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A 

it ~—— \ 
il 

5S 

{ 

if 


0.5mm 


Fig. 8. Botryllus tabori: mature zooid in lateral view. 


imen in USNM (20031), three in MZUSP 
(11180, 11181, 11182). Gonads found in all 
months, except September and November; 
larvae present in April and August. 

Remarks. —This species is widely known 
as Botrylloides nigrum. However, C. & F. 
Monniot (1987) argue convincingly that the 
characters used to distinguish the genus Bo- 
trylloides from Botryllus are too unreliable 
to maintain this separation and that the 
name of the older genus, Botryllus, should 
prevail in all cases. 

Distribution and habitat. — Distributed 
widely throughout warm regions. In the At- 
lantic: Bermuda, Florida, the Caribbean Sea. 
In Brazil: Espirito Santo (Vitoria), Rio de 
Janeiro (Niter6oi, Rio de Janeiro), Sao Paulo 
(Sao Sebastiao, Santos), Santa Catarina 
(Florianopolis). Very common in shallow 
water, sometimes growing directly on rocky 
surfaces but more often on other organisms 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


such as algae, sea grasses, sponges, poly- 
chaete tubes, and ascidians. 


Botryllus tabori Rodrigues, 1962 
Fig. 8 


Botryllus tabori Rodrigues, 1962:199, pl. 3, 
figs. 8-11; Costa, 1969e:301, fig. 5. 


Material examined.—Six colonies from 
Praia do Segredo, five from Praia do Arag¢a, 
one from Praia Grande. One specimen in 
USNM (20030), one in MZUSP (11183). 

Remarks. —The zooids (Fig. 8) differ from 
the description of Rodrigues (1962) in hav- 
ing more branchial tentacles (twelve instead 
of eight) and fewer testis lobes (from five to 
nine instead of twelve). The atrial aperture 
most commonly was a wide horizontal 
opening; rarely, it opened at the end of a 
siphon. 

Distribution and habitat.—In Brazil: Rio 
de Janeiro (Urca), Sao Paulo (Sao Sebastiao: 
type locality). Intertidal colonies on the un- 
dersurfaces of boulders, and on algae (main- 
ly Ulva sp.), sponges, and solitary ascidians. 


Botryllus tuberatus 
Ritter & Forsyth, 1917 


Botryllus tuberatus Ritter & Forsyth, 1917: 
461, pl. 39 figs. 10, 12, pl. 40, fig. 22.— 
Van Name, 1945:225, fig. 135.—C. Mon- 
niot, 1983:426, fig. 2. 

Botryllus primigenus: Van Name, 1945:223, 
fig. 134.—Millar, 1958:505.—Costa, 
1969e:301, fig. 4. 

Botryllus schlosseri: Bjornberg, 1956:164. 


Material examined. — Three colonies from 
Praia do Segredo, one from Praia Grande. 
One specimen in MZUSP (11184). 

Remarks.—C. Monniot (1983) decided 
that all western tropical Atlantic Botryllus 
with four rows of stigmata are B. tuberatus. 
He noted that the zooids of this species form 
systems around common cloacas, in con- 
trast to those of B. primigenus, which send 
their atrial apertures separately to the col- 
ony surface. In the present material only a 


VOLUME 106, NUMBER 4 


few zooids within a colony have atrial ap- 
ertures that open independently in this way; 
by far most zooids form systems like those 
in colonies from Guadeloupe (C. Monniot 
1983). C. Monniot (1983) does not remark 
on the enlargement of the gut near the stom- 
ach, regularly present in our specimens, but 
it is apparent in his figure. 

The same material that Bjornberg (1956) 
attributed to B. schlosseri was later de- 
scribed by Millar (1958) as B. primigenus. 
By implication, if Monniot’s (1983) syn- 
onymy holds, those specimens are B. tube- 
ratus. 

Distribution and habitat. —California 
(Santa Barbara: type locality), Florida, Tor- 
tuga Islands, Guadeloupe. In Brazil: Rio de 
Janeiro (Urca), Sao Paulo (Santos, Cana- 
néia, Sao Sebastiao). Colonies were found 
intertidally on the undersurfaces of boul- 
ders, growing directly on the rock or on 
sponges and algae. They also settle on ce- 
ramic plates submerged continuously in 
shallow water. 


Discussion 


All the compound ascidians already re- 
corded from intertidal and shallow subtidal 
habitats at Sao Sebastiao occur as well in 
the Caribbean region and/or other tropical 
seas (Rodrigues 1962). Only Botryllus tabori 
seems to be endemic to the Brazilian coast. 
New records are reported in this paper for 
Didemnum psammathodes, Trididemnum 
orbiculatum, Distaplia bermudensis, and 
Symplegma rubra, all known previously 
from the Caribbean, and some of them dis- 
tributed elsewhere, as well, in warm waters. 
Botryllus giganteus is known previously 
from West Africa. 

Species distributed widely in warm wa- 
ters may be newcomers. Worldwide ship- 
ping has introduced exotic species into many 
ports (C. Monniot et al. 1991): Sao Sebas- 
tiao, since the sixties, has become Brazil’s 
biggest maritime oil terminal. 

In general, the littoral marine fauna of 
Sao Paulo can be considered an impover- 


TAT 


ished Caribbean fauna. However the sub- 
littoral ascidian fauna presents antiboreal 
cold water species (Rodrigues 1966). Evi- 
dence of an upwelling of subtropical water 
in the region of the continental shelf be- 
tween 20° and 26° S has been known since 
Emilsson (1961). 


Acknowledgments 


We are indebted to the following insti- 
tutions for providing logistical support: 
Centro de Biologia Marinha da Universi- 
dade de Sao Paulo e Departamento de Ecol- 
ogia Geral do Instituto de Biociéncias, 
Universidade de Sao Paulo. One of us 
(RMR) also thanks her advisor Dr. A. Ce- 
cilia Z. Amaral, and Drs. Claude and Fran- 
coise Monniot, who kindly received her at 
the Muséum National d’Histoire Naturelle, 
Paris, for a four-month training in ascidian 
systematics, and for helpful suggestions. 
Todd Newberry made a careful and com- 
prehensive revision to the manuscript and 
provided copies of missing literature, for 
which we are indebted. The research was 
supported by grants from CAPES and FA- 
PESP (86/1476-0) to R. M. Rocha and 
formed part of her studies for the MSc de- 


gree. 


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VOLUME 106, NUMBER 4 


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739 


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(SAR) Departamento de Ecologia Geral, 
Instituto de Biociéncias, and Centro de 
Biologia Marinha, Universidade de Sao 
Paulo, C. Postal 11461, 05422-970 Sao 
Paulo, SP, Brasil; (RMR) Departamento de 
Zoologia, Instituto de Biologia, Universi- 
dade Estadual de Campinas, C. Postal 6109, 
13081-970 Campinas, SP, Brasil. 


PROC. BIOL. SOC. WASH. 
106(4), 1993, pp. 740-748 


ACTIVITY AND REPRODUCTIVE PATTERNS OF 
AMPHIBIANS AND REPTILES FROM THE ENGARE 
ONDARE RIVER REGION OF CENTRAL KENYA, 
DURING THE DRY SEASON 


Laurence M. Hardy 


Abstract. — This is the first report of the reproductive patterns and activities 
of a sample of frogs and lizards from the remote region of Isiolo District in 
central Kenya during the dry season. Seventeen species were active and feeding; 
one species was dormant and not feeding. Five species were reproductive and 
thirteen species were clearly not reproductive. Adaptive partitioning of the 
reproductive cycle of some species seems to be occurring. 


There are no reports on the herpetofauna 
of the remote region of Isiolo District in 
central Kenya nor is there any information 
concerning the activity, reproduction, and 
relative abundance of reptiles and amphib- 
ians during the summer dry season in this 
arid habitat. Western (1974) studied lizards 
for 17 days near Lokori in South Turkana, 
Kenya. That area is 235 km NW of our site 
and separated from it by the Suguta Valley 
(300-400 m) and the Lerochi Plateau (2000-— 
2600 m). Even though taxonomic literature 
for amphibians and reptiles in Kenya is 
abundant, few papers give reproductive and 
feeding data, especially during the dry sea- 
son. 

Hebrard (1980) reported on the habitats 
of Chamaeleo during the dry season in lo- 
calities south of Nairobi. Greer (1967) stud- 
ied comparative ecology of two species of 
Lygodactylus in northwestern Kenya during 
the dry season. The report by Loveridge 
(1929) does not present reproductive nor 
activity data and includes only a few records 
in central Kenya. Western (1974) reported 
biomass data for lizards but did not include 
reproductive or feeding data. Other reports 
concerned only the rainy season (Bogert 
1942) or only the rain forest (Loveridge 


1935, 1936; Drewes 1976). This paper con- 
cerns some species of frogs and lizards from 
Isiolo District, Eastern Province, of central 
Kenya during the middle of the dry season 
(July 1987). 

It is of interest to know which species are 
reproductive and feeding during the dry sea- 
son, which species are active but not repro- 
ductive (are they feeding?), and which are 
dormant. Until we know how each species 
responds to seasonal change, we cannot dif- 
ferentiate populational changes that might 
be due to natural biological interactions from 
those resulting from the ever-increasing im- 
pacts from over-grazing by domestic ani- 
mals. The objectives are to identify 1) those 
species of amphibians and reptiles that are 
actively feeding versus those that are dor- 
mant and 2) those that are reproductive ver- 
sus those that are not reproductive. 

Hardy and John L. Darling arrived on 11 
July 1987 and departed on 3 August 1987. 
The camp was 20.6 km west and 1.5 km 
south of Isiolo, Isiolo District, at an ele- 
vation of 1120-1140 m, on the eastern bank 
of the Engare Ondare River, which forms 
the boundary between Isiolo and Laikipia 
Districts (Fig. 1). During that period no rain 
fell and the camp workers reported that the 


VOLUME 106, NUMBER 4 


741 


<= 
°o 
°o 
oe 
J 

7 
c 
o 
=x 


37° 30) 


Fig. 1. Map showing vicinity of Isiolo, Isiolo District, Eastern Province, Kenya. Uppercase letters identify 
collection locations (Table 1). Heavy stipling represents area above 1400 m; fine stipling represents areas between 


1200 and 1400 m elevation. 


last rain was in March or April and none 
was expected until October. The annual 
rainfall in this general region of Kenya is 
50-100 cm. 

The Engare Ondare is a permanent stream 
near our camp; however, as the dry season 
progressed the stream dried from the north- 
ern lower end toward the mountains to the 
south. During our stay, the lower end of the 
stream advanced (by drying) to within 2-3 
km north of camp. At our location the 
stream was 1-3 m wide, 10-20 cm deep, 
and flowing over a rocky or compacted mud 
substrate. The flood channel of the stream 
was 10-50 m wide with sandy, eroded banks 
up to 10 m high; the floor and sides were 
composed of sand with mixed gravel and 
rocks. 

Acacia trees, up to 10 m high, were abun- 
dant and formed a riparian buffer between 
the stream and the surrounding desert. 
Among the Acacia were several shrubs, in- 
cluding Euphorbia, sparse grass, and a va- 
riety of perennials and annuals. The Acacia 


forest was 50-100 m wide on each side of 
the river. Beyond the Acacia forest, the des- 
ert was a short thorn forest consisting of 
several species of sparse Acacia (2-3 m high), 
Euphorbia, and small-leafed perennial 
shrubs. Abundant grass (not Cynodon) was 
found only at distances of 1-2 km from the 
river and away from areas grazed by goats; 
Cynodon was in small clumps on the river 
bank and in a wet meadow. 


Methods 


All preserved specimens are in the Mu- 
seum of Life Sciences of Louisiana State 
University in Shreveport (LSUS). Collec- 
tion localities (A—E) are defined in Table 1. 

One testis and the largest ovum were 
measured to 0.01 mm with dial calipers un- 
der a dissecting microscope. In frogs, ovi- 
ducal eggs or amplexus was interpreted as 
an accurate indication of relatively imme- 
diate reproduction and oviposition. En- 
larged ovarian eggs could occur over a long 


742 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Collection localities. Letters A-E are used in the text to identify the following localities. 


A) Isiolo Dist.: 20.6 km W, 1.5 km S Isiolo. 
B) Isiolo Dist.: 20.0 km W, 0.4 km S Isiolo. 
C) Isiolo Dist.: 16.6 km W, 1.2 km N Isiolo. 
D) Isiolo Dist.: Jct. Kenya hwy. A2 & Ewaso Ngiro.? 
E) Laikipia Dist.: 1-2 km E 01 Doinyo Lossos peak.? 


2 Locality not shown in Fig. 1. 


period of time and would not always indi- 
cate immediate reproduction. Calling of 
anurans and enlarged seminiferous tubules 
and vas deferentia indicate a non-dormant 
condition, probably feeding, but not nec- 
essarily immediately reproductive (the fe- 
males may not yet be receptive). The pres- 
ence of recognizable food items in the 
stomach was used as an indication of rela- 
tively recent (probably within 24 hours) 
feeding activity. All measurements are in 
mm. Sexes are combined for statistics of 
small samples. 


Results 


Bufo garmani Meek.—(LSUS 4776, lo- 
cality A; 4777, B; 4778-4809, A; 4865, A; 
total n = 35). SVL was 53-74 (X = 66.0, n 
= 22) in males and 62-89 (X¥ = 72.4, n = 
12) in females. Testes were 0.8-2.1 (¥ = 
1.6) x 9.1-16.1 (X¥ = 11.9). Four females 
with SVL < 66 had non-pigmented ova (0.4 
diameter or less) and eight with SVL > 68 
had pigmented ova (0.7 dia. or greater). The 
two specimens (4781, 4808) with the largest 
ova (1.4 dia.) had ovulated (oviducal eggs) 
and were 80 and 83 SVL. One metamorph 
(4865), 11.3 SVL with a regressing tail 3.1 
long, was found under a rock in wet sand 
on 29 July at 1605-1650 h. The sex ratio 
of the 34 adults (SVL > 52) was 1 female 
to 1.8 males. All specimens lacked bright 
scarlet on the posterior surface of the thigh 
and dark spots on the dorsum of the snout, 
thus eliminating B. regu/aris and had reni- 
form paratoids, not like B. kerinyagae (Keith 
1968). All specimens except two (6%; one 
male and one female) had food in the stom- 


achs. Two specimens (LSUS 4776 on 12 
July, 4792 at 0950 h on 19 July) were flushed 
from holes in sandy soil by flooding with a 
bucket of water. Another (LSUS 4777) was 
found under a log in a grassy meadow (1100 
h) on 14 July. A pair in amplexus (LSUS 
4780-8 1) was found along the river at 2200-— 
2400 h, 14 July. One male was calling from 
within a burrow at the base of the arroyo 
bank well away from the water; it was dug 
out at 1920 h (14 July). All other specimens 
were found along the edge of the water in 
sandy areas; none was found in the thick 
Bermuda grass (Cynodon) that grows in 
small patches along the river. 

Tomopterna cryptotis (Boulenger).— 
(LSUS 4810-11, A). Two males were found 
on wet sand along the edge of the river (LSUS 
4810 on 14 July; LSUS 4811 at 1930-2100 
h, 25 July); neither was calling nor moving. 
Testes were 0.5 X 2.4 and 1.0 X 4.5 re- 
spectively. One stomach contained insects; 
the other was empty. 

Hemisus marmoratum (Peters).—(LSUS 
4876; A). A male (SVL = 32; testis spher- 
ical, 1.7 dia.) with an empty stomach was 
dug from moist sandy loam, at a depth of 
15-16 cm, in a sand bar next to the river 
(0900 h on 26 July). Because of its empty 
stomach, small gonads, depth at which it 
was buried, and clean appearance (as if it 
were in a chamber, rather than embedded 
in sand) we assumed that it was dormant. 
No eggs were found. 

Ptychadena anchietae (Bocage).—(LSUS 
4813-26, A; 4827-30, B; 4831-60, A; 4861- 
64, D; total nm = 52). Males had SVLs of 27— 
AO (X = 35.5; n = 21); females 18-55 (X = 
37.5; n = 30). All males 35 SVL or larger 


VOLUME 106, NUMBER 4 


had testes greater than 1.0 < 2.7 (max. size 
= 1.4 x 4.1); those with smaller testes were 
less than 34 SVL. Some females 35 SVL or 
greater had enlarging oocytes (greater than 
0.1 dia.), other specimens from 18 to 45 
SVL did not contain enlarged oocytes. All 
females larger than 48 SVL had pigmented 
oocytes up to 1.3 diameter (probably ready 
for ovulation), but none had ovulated. Even 
though some males were calling, this species 
was probably not reproducing at that time; 
none of the females had ovulated. Food was 
present in the stomachs of 48 specimens; 
two females and two males had empty 
stomachs. The male with the largest testis 
had an empty stomach. In life, most spec- 
imens were yellow postero-ventrally with 
yellowish-green stripes on the posterior sur- 
face of the thigh; some had a green mid- 
dorsal stripe. All were found near the water; 
some in the water, along the stream margin 
on sand, others in the thick bermuda grass 
(Cynodon) on the bank. 

Ptychadena mascareniensis (Dumeril and 
Bibron).—(LSUS 4867-73, A). SVL X = 
43.9, range = 28-55, n = 7. The smallest 
individual was a non-reproductive female 
(SVL = 28); all others (SVL > 41) were 
reproductive. Three mature females con- 
tained enlarged ovarian ova and one con- 
tained oviductal eggs. Two males had testes 
measuring 4.7 < 1.3 and 3.9 x 1.3 with 
black tunica albuginea. One mature female 
had lost the right rear foot. All contained 
food (unrecognizable insects) in the stom- 
achs. All specimens were found along the 
river; one (LSUS 4867) was under a rock in 
the river at 1820 h on 14 July. The mid- 
dorsal stripes were tan (LSUS 4871-72) or 
yellow (LSUS 4868). 

Rana angolensis Bocage.—(LSUS 4812; 
B). A female (SVL = 77) containing large, 
pigmented, ovarian eggs and an enlarged 
oviduct, was found on 15 July. The stomach 
contained insects. 

Phrynobatrachus mababiensis_ Fitz- 
Simons.—(LSUS 4866, 4874-75; A). This 
is the first report of this taxon as a distinct 


743 


species in Kenya. Duff-MacKay (1980) re- 
ported mababiensis as a subspecies of P. 
ukingensis. Two females (4866, 4875; SVL 
= 17.3, 17.6, respectively) and one male 
(SVL = 14.8) were found. Ovarian eggs were 
enlarged (1.0, 0.8, respectively) and pig- 
mented. The testis of the male was 0.9 x 
1.7. All contained food in the stomachs. 
These specimens were caught on 29 July 
from beneath rocks in wet sand along the 
edge of the water between 1605-1650 h. 

Chamaeleo gracilis Hallowell.—(LSUS 
4951; C). A male (SVL = 100, tail = 80, 
tail/total length = 0.44; black testis, 6.3 x 
4.4) was found on a small bush near a house, 
25 July. The stomach was packed with in- 
sects. No others were found after thorough 
searches in the area. Local residents re- 
ported that this species was common at this 
time of the year. 

Hemidactylus brookii Gray.—(LSUS 
4893-94; A). A female (4893; SVL = 49), 
found on 12 July, contained ova 1.0 or 
smaller in diameter; a male (SVL = 45) cap- 
tured on 20 July had an unusually fat tail 
and the right testis was 1.8 x 4.2. Both 
contained food in the stomachs. 

Hemidactylus mabouia (Moreau de 
Jonnés).—(LSUS 4891-92; A). A female 
(4891; SVL = 75; ova to 2.0) was found on 
19 July at 2320-2350 h and a male (4892; 
SVL = 48; testis 1.6 x 3.2) on 20 July (air 
temperature 20.2°C at 2153 and at 2253 h). 
Both were found on Acacia trees at night 
with food in the stomachs. Both had incom- 
plete tails. 

Lygodactylus picturatus Peters.—(LSUS 
4895-99, A; n = 5). In one female (LSUS 
4895; SVL = 34) the tip of the tail was spat- 
ulate in shape with five pairs of ventral la- 
mellae, which had the same appearance as 
the subdigital lamellae. She had two ovi- 
ductal eggs (5 x 7). The other specimens 
had normal tails. Two specimens (4897-8) 
contained ova up to 3.1 and 1.1 in diameter, 
respectively. Two males (4896, 4899) had 
testes 1.1 X 2.6 and 1.6 x 2.4, respectively. 
The vas deferens of one (4896) was greatly 


744 


enlarged. Three specimens contained in- 
sects in the stomachs. The gravid female 
contained unidentifiable debris in the large 
intestine and one male contained insect re- 
mains only in the small intestine. Three 
specimens (4895, 4896, 4898; 13, 14, and 
20 July, respectively) had yellow midven- 
tral stripes and the yellow extended onto 
the venter of each leg in 4895. On LSUS 
4896 the yellow was restricted to a median 
ventral stripe, the throat and chin were black, 
and the sides of the throat and chest had 
black streaks; this gecko was found on a tree 
trunk at 1400 h. Three females (SVL Y = 
34.0, range = 33-50) had black streaks on 
the chins and two males (SVL = 34, 35) had 
solid black chins. One female had a com- 
plete tail (tail/total length = 0.47). The males 
had 7 and 9 preanal pores. 

Agama agama (Linnaeus).—(LSUS 4930- 
48, A; 4949, D; 4950, E). Three females 
under 100 SVL were not reproductive; two 
larger females contained shelled oviductal 
eges (SVL = 103) and ova to 2.4 (SVL = 
110). The testes in 11 males under 100 SVL 
were less than 2.0 X 2.0; four males over 
100 SVL (X = 119) had testes 3.3-6.0 x 
4.3-6.1. SVL for males was X = 71.9 (range 
= 35-142, n = 15) and for females 73.2 
(range = 33-110, m = 6). The yolk-sac scar 
was evident on one female (SVL = 33) and 
one male (SVL = 35). All contained food. 
Two juveniles (4931-32; 12 July) had four 
red dorsal stripes; two adult males (4933- 
34; 13 July) had four orange dorsal stripes, 
green spots on the head, and a pale yellow 
throat. A female (4935) and two males 
(4936-37) caught at night (2200-2300 h; 13 
July) in a crack of a large dead tree had red 
heads, blue chests, yellowish-brown necks, 
and bluish-green tails. Another male (4944; 
21 July) had a red throat with pink longi- 
tudinal stripes, blue venter, and golden- 
brown neck. Four specimens (4939-42) were 
found under one rock near the river at 2200 
h (20 July). This diurnal species was found 
at night under bark of dead trees, in cracks 
in trees and rocks, and under large rocks; 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


none was seen in small mammal burrows. 
All were extremely wary and took refuge at 
the slightest disturbance. 

Agama rueppelli Vaillant.—(LSUS 4924- 
28, A; 4929, E). All six specimens were found 
during 20-28 July (SVL X = 70.5, range = 
38-88; tail ¥ = 163.0, range = 157-169, n 
= 3; tail/total length Y = 0.67, range = 0.66— 
0.67). Ova of three females were 0.6 (SVL 
= 52), 1.6 (83), and 4.7 (88); testes of three 
males were 1.1 x 1.7 (38), 5.7 X 7.7 (81), 
and 6.1 x 9.5 (81). All had food in the 
stomachs. One female (4925) had a pair of 
dark, brownish-black chest spots, yellow 
chin, grayish-tan middorsal stipe, yellow 
nuchal sides, and a pale pink loreal region. 
A male (4926) was similar, except paler, and 
the back of the head was pink. Both speci- 
mens were found in the shade of shrubs in 
the desert away from the river (1200-1230 
h). In contrast to A. agama, individuals of 
this species were docile and easily caught. 

Eremias spekii Guinther.—(LSUS 4907— 
23; A). SVL was 31-48 (X¥ = 39.3, n= 11) 
in males and 33-53 (X = 46.7, n = 6) in 
females. Four males less than 38 SVL had 
small testes (1.3 < 1.9), whereas five others 
with SVLs greater than 40 (X = 46) had 
testes 2.1 x 3.0 or larger; the two largest 
specimens (48 SVL) had greatly enlarged 
vas deferentia. Four females had ova 2.2 or 
smaller; one (4921) had an SVL of 53 and 
ova as large as 4.6. Males probably reach 
sexual maturity at approximately 45 SVL 
and females at approximately 53. All 17 
specimens were found in the desert away 
from the river and contained food in the 
stomachs, except one female (SVL = 51) 
that had food only in the large intestine. One 
female (4907; 12 July), active under a bush 
in the morning, had a yellowish-orange sub- 
caudal surface changing to yellowish-green 
at the base of the tail. Two (4908-9) were 
active between 0830-0900 h. Most were 
found among low grayish-green shrubs 
(Acacia sp.) in low foothills above the river 
valley before noon on 20 July. This area 
was less grazed by goats than the area bor- 


VOLUME 106, NUMBER 4 


dering the river. Lizards escaped into rodent 
burrows, but apparently stayed just inside 
the entrance; many were easily dug out by 
suddenly scooping all of the sand from just 
behind the burrow entrance into an open 
area. 

Latastia longicaudata (Reuss).—(LSUS 
4900-06, A). SVL X¥ = 68.0, range = 44- 
79; tail ¥ = 148.3, range = 117-191 (3 fe- 
males); tail/total length ¥ = 0.73, range = 
0.72-0.73 (3 females). The male had a long 
regenerated tail that was 71 percent of its 
total length. Six females contained ova 0.8- 
1.8 diameter and the single male (4906; SVL 
= 79) contained a testis 1.7 x 3.1. All had 
food in their stomachs. One female (4900) 
was found under a small bush about 1600 
h, 14 July. Six other specimens were found 
on 20 July at the same time and in the same 
habitat as that described for Eremias spekii. 

Lygosoma sundevallii (Smith).—(LSUS 
4889-90; A). An adult male (4890; SVL = 
79, tail = 58, tail/total length = 0.42) was 
in a hollow log brought to camp for firewood 
on 26 July. This male had a testis 2.6 x 7.0 
and enlarged vas deferens and contained a 
20 mm insect larva. A small female (4889; 
SVL = 47, tail = 27, tail/total length = 0.36) 
with ova to 0.6 and several food items in 
the duodenum was dug out of the sand in 
the morning of 20 July by digging in loose 
sand at the bases of small bushes; food items 
in the intestine suggest that it may not have 
been foraging that day. 

Mabuya quinquetaeniata (Lichten- 
stein).—(LSUS 4877, A; 4878-84, C; 4885, 
E). SVL X = 68.7, range = 44-89, n = 9; 
tail ¥ = 100.4, range = 70-132, n = 5; tail/ 
total length ¥ = 0.60, range = 0.58-0.61, 
= 5. Maximum ova size for three females 
was 1.7; maximum testis size for six males 
was 1.6 x 2.9. All contained food in the 
stomachs. One specimen (4877; 18 July) was 
found on the NW slope of a rocky hill W 
of camp; several others were extremely wary. 
On 19 July, seven specimens were collected 
from a dark lava flow along a dry creek bed. 
All were captured by stunning with large 


745 


rubber bands. Their skins were fragile and 
all were damaged during capture. 

Mabuya planifrons (Peters).—(LSUS 
4886, C; 4887-88, A; 4952, B). SVL was 
93, 104 for the two mature males (4886, 
4888; testes 7.8 x 5.0 and 11.6 x 5.0, re- 
spectively) and 116 for the single mature 
female; tail was 130, 119, 221, 156+, re- 
spectively. The female contained oviductal 
ova up to 11.0 diameter. All specimens had 
arthropod remains in the stomachs. One 
large adult female (4952) was in thick ber- 
muda grass of a large wet meadow (approx- 
imately 0.8 x 0.3 km) on 14 July, at 1000 
h. The Loborua River flows through the 
meadow but was reduced to a meter or so 
in width by 25 July. This female was pale 
yellow around the ear openings. A male 
(4886; 19 July) was caught with M. quin- 
quetaeniata on the lava flow. Two others 
(4887-88) were caught on 21 July: one of 
these, a juvenile male (4887) with testis 1.5 
x 3.0, had a yellowish wash on the lips and 
soles of the feet. 


Discussion 


This sample represents a small propor- 
tion of the total herpetofauna (more than 
340 species) in Kenya. However, it probably 
represents many of the species of frogs and 
lizards that normally are active during the 
dry season in the Isiolo area. Since signifi- 
cant migration is not a documented re- 
sponse for terrestrial amphibians and rep- 
tiles, means of individual survival during 
seasonal drought include dormancy, change 
in diet, change in microhabitat use, or no 
change. One of the frogs, Hemisus mar- 
moratum, appeared to be in a state of dor- 
mancy, but the other species were active and 
foraging regularly (Table 2). Without com- 
parable study during the rainy season, it 1s 
unknown if any of these species undergo 
seasonal shifts in diet, microhabitat use, or 
activity. Interspecific competition is as- 
sumed to occur among some species of this 
herpetofauna and is presumably different in 
the wet and dry seasons. Hebrard & Madsen 


746 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 2.—Seasonal activity and reproductive condition of 7 species of amphibians (above line) and 11 species 
of reptiles from the vicinity of the Engare Ondare River, Isiolo District, Kenya. Feeding activity is measured 
as the percent of stomachs containing recognizable food. 


Percent 
stomachs 


with Repro- 

Species n food Active ductive Most reliable reproductive condition 
Bufo garmani 34 94 Yes Yes Oviductal eggs; amplexus observed 
Ptychadena mascareniensis 4 100 Yes Yes Oviductal eggs 
Tomopterna cryptotis Z 50 Yes No Testis to 1.0 x 4.5 
Ptychadena anchietae 52. 92 Yes No Calling; pigmented ovarian eggs 
Rana angolensis 1 100 Yes No Pigmented ovarian eggs 
Phrynobatrachus mababiensis 3 100 Yes No Pigmented ovarian eggs 
Hemisus marmoratum 1 0 No No Testis 1.7 


Lygodactylus picturatus 5 60 Yes Yes Oviductal eggs 

Agama agama 21 100 Yes Yes Oviductal eggs 

Mabuya planifrons 4 100 Yes Yes Oviductal eggs 

Chamaeleo gracilis 1 100 Yes No Testis 4.4 x 6.3 
Hemidactylus brookii 2 100 Yes No Testis 1.8 x 4.2; ova to 1.0 
Hemidactylus mabouia 2 100 Yes No Testis 1.6 x 3.2; ova to 2.0 
Agama rueppelli 6 100 Yes No Testis to 6.1 xX 9.5; ova to 4.7 
Eremias spekii £7 94 Yes No Testis to 2.8 x 4.1; ova to 4.6 
Latastia longicaudata 7 100 Yes No Testis 1.7 x 3.1; ova to 1.8 
Lygosoma sundevallii 2 50 Yes No Testis 2.6 x 7.0; ova to 0.6 
Mabuya quinquetaeniata 9 100 Yes No Testis to 1.6 xX 2.9; ova to 1.7 


(1984) related differential microhabitat dis- 
tribution in Chamaeleo dilepis in Kenya to 
environmental stresses imposed on the pop- 
ulation during the dry season. During the 
wet season microhabitat diversity increases 
due to increased water availability and in- 
creased plant diversity (activity); thus, rel- 
ative interspecific competition probably in- 
creases during the dry season owing to a 
concentration of species in available micro- 
habitat and using reduced food resources 
(see also Hebrard & Madsen 1984). Garcia 
& Drummond (1988) found Thamnophis 
eques to be more euryphagous when allo- 
patric with congeners and more stenopha- 
gous when sympatric with congeners. Toft 
& Duellman (1979) suggested that the uti- 
lization of reproductive resources is affected 
by seasonality. Hebrard & Madsen (1984) 
found that niche expansions by Chamaeleo 
is permitted by the absence of competing 
species. During the dry season, microhab- 
itat diversity is at a minimum due to water 
shortage (restricted to the flowing river) and 


decreased botanical structural complexity 
(loss of leaves, fewer flowers and fruits, and 
restricted vegetative growth); however, rel- 
ative interspecific competition might con- 
tinue to be high because the fewer active 
species compete for highly reduced re- 
sources (most plants are not growing and 
flowering to produce suitable microhabitat 
and food supply). Jenssen (1973) recorded 
niche shifts in tropical lizards (Anolis) due 
to competition. 

Tropical snakes (Henderson et al. 1978) 
and arboreal frogs (Toft & Duellman 1979) 
are probably most adversely affected by sea- 
sonal drought and, therefore, rely on dor- 
mancy to survive the dry period. More ter- 
restrial frogs (less exposed to lowered relative 
humidity; 1.e., Ptychadena, Rana, Tomop- 
terna, and Phrynobatrachus) probably 
change diet or microhabitat utilization. 

Adaptations to the dry season include 
dormancy by some species and normal ac- 
tivity by others. Active species have greater 
relative energy demands and are feeding 


VOLUME 106, NUMBER 4 


whereas those unable to sustain activity (due 
to inadequate food or water) are dormant 
during the dry season. Thus, three activity 
categories for amphibians and reptiles can 
be recognized during the dry season: 1) ac- 
tive and reproductive, 2) active, but not re- 
productive, and 3) dormant. 

Of the active amphibians encountered in 
this study, only Bufo garmani and Ptycha- 
dena mascareniensis were reproductive 
(Table 2); however, all of the other active 
species had pigmented ovarian eggs or en- 
larged testes. All of the active frogs that were 
not reproductive were probably approach- 
ing reproduction and would have repro- 
duced when the summer rains began. Hem- 
isus 1s in category three. 

Both species of Hemidactylus were not 
reproductive, but Lygodactylus was ready 
for oviposition (Table 2). Agama agama was 
reproductive, but A. rueppelli was not. Ma- 
buya quinquetaeniata was not reproductive 
(gonads tiny, apparently regressed), but M. 
planifrons was ready for oviposition (Table 
Z). For some pairs of species in a genus (Pty- 
chadena, Agama, Mabuya) one species is 
reproductive and the other is not. The non- 
reproductive condition of several species of 
amphibians and reptiles and the reproduc- 
tive condition of others suggest an adaptive 
partitioning of the reproductive season by 
congeners, which would result in reduction 
of demands on the associated resources, and 
potentially would reduce competition. 


Acknowledgments 


I thank Mrs. Mwongo (Office of the Pres- 
ident, Kenya), Dr. Olinde (Director, De- 
partment of Wildlife, Kenya), and Dr. M. 
Buvi (Assistant Director, Department of 
Wildlife, Kenya) for the research permit and 
exportation permits for Kenya. I also thank 
Dr. Richard Leakey and Mr. Alex Duff- 
MacKay (National Museum of Kenya) for 
assistance during the process of obtaining 
the research permit. For help with collecting 
I thank James, Shadrack, Raoul, Konoso, 
and Charlie. Vaughn Langman (LSUS) 


747 


waived a portion of the commercial camp 
fee during the stay in his camp. I am most 
indebted to John L. Darling for his untiring 
assistance and perseverance during this ex- 
pedition. For permission to examine spec- 
imens in their care I thank Dr. Barry Clarke, 
British Museum (Natural History) (BM), and 
Dr. W. Ronald Heyer, U.S. National Mu- 
seum of Natural History (USNM). I thank 
LSU in Shreveport for a faculty research 
grant (no. 757-05-6002) for partial support 
of this project. 


Literature Cited 


Bogert, C. M. 1942. Snakes secured by the Snyder 
East African expedition in Kenya colony and 
Tanganyika territory.—American Museum 
Novitates 1178:1-5. 

Drewes, R.C. 1976. Report on an expedition to Ka- 
kamega forest.— East Africa Natural History So- 
ciety Bulletin pp. 122-126. 

Duff-MacKay, A. 1980. Amphibia.—National Mu- 
seum of Kenya Conservation Status Report (1): 
1-44. 

Garcia, C. M.,& H. Drummond. 1988. Seasonal and 
ontogenetic variation in the diet of the Mexican 
garter snake, Thamnophis eques, in Lake Te- 
cocomulco, Hidalgo.—Journal of Herpetology 
22(2):129-134. 

Greer, A. E. 1967. The ecology and behavior of two 
sympatric Lygodactylus geckos. — Breviora (268): 
1-19. 

Hebrard, J. J. 1980. Habitats and sleeping perches 
of three species of chameleons in Kenya.— 
American Zoologist 20(4):842. 

—., & T. Madsen. 1984. Dry season intersexual 
habitat partitioning by flap-necked chameleons 
(Chamaeleo dilepis) in Kenya.— Biotropica 16(1): 
69-72. 

Henderson, R. W., J. R. Dixon, & P. Soini. 1978. On 
the seasonal incidence of tropical snakes. — Mil- 
waukee Public Museum, Contributions to Bi- 
ology and Geology (17):1-15. 

Jenssen, T. A. 1973. Shift in the structural habitat 
of Anolis opalinus due to congeneric competi- 
tion.— Ecology 54(4):863-869. 

Keith, R. 1968. A new species of Bufo from Africa, 
with comments on the toads of the Bufo regu- 
laris complex.—American Museum Novitates 
(2345):1-22. 

Loveridge, A. 1929. East African reptiles and am- 
phibians in the United States National Muse- 
um.—Bulletin of the United States National 
Museum 151:1-135. 


748 


1935. Scientific results of an expedition to 
rain forest regions in eastern Africa. I. New rep- 
tiles and amphibians from east Africa.—Bulle- 
tin of the Museum of Comparative Zoology 
79(1):3-19. 

1936. Scientific results of an expedition to 

rain forest regions in eastern Africa. VII. Am- 
phibians.— Bulletin of the Museum of Compar- 
ative Zoology 79(7):369-430. 
Toft, Catherine A., & W. E. Duellman. 1979. An- 
urans of the lower Rio Liullapichis, Amazonian 
Peru: a preliminary analysis of community 
structure. — Herpetologica 35(1):71-77. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Western, David. 1974. The distribution, density and 
biomass density of lizards in a semi-arid envi- 
ronment of northern Kenya.—East African 
Wildlife Journal 12:49-62. 


Museum of Life Sciences, One University 
Place, Louisiana State University in 
Shreveport, Shreveport, Louisiana 71115- 
2399 Sane 


PROC. BIOL. SOC. WASH. 
106(4), 1993, pp. 749-751 


A PETREL-LIKE BIRD FROM THE LATE EOCENE OF 
LOUISIANA: EARLIEST RECORD FOR THE 
ORDER PROCELLARITFORMES 


Alan Feduccia and A. Bradley McPherson 


Abstract. —The earliest known certain record of the avian order Procellariifor- 
mes is a Pterodroma-like fossil represented by the distal end of a tibiotarsus, 
reported herein, from the late Eocene Yazoo Formation of Louisiana. Sedi- 
ments bearing this bird appear to have been deposited on the continental shelf 
far from the Eocene coast line of the Gulf of Mexico, and associated fossils 
include a sea snake, a primitive whale, numerous chondrichthian and marine 
teleost fishes, and a large marine invertebrate fauna. 


Given the relative abundance of the mod- 
ern “‘tube-nosed swimmers,” it is somewhat 
surprising that the Procellariiformes do not 
have a more comprehensive fossil record 
(Brodkorb 1963, Olson 1985a). This may be 
due in part to their pelagic habits, and also 
to the fact that most major finds of fossil 
birds have been from the Northern Hemi- 
sphere, whereas the Procellariiformes are 
predominantly distributed in the Southern 
Hemisphere. 

The order is thought to be ancient, but 
the fossil record has yet to provide much 
support for this assumption. The avifauna 
of the late Cretaceous greensands of New 
Jersey, recently revised by Olson & Parris 
(1987), includes 8 or 9 genera and 9 or 10 
species, of which two genera and species 
show some similarities with the Procellar- 
uiformes. However, given the fragmentary 
nature of the material and the mosaic nature 
of many early Paleogene birds, positive 
identification must await the discovery of 
more complete material. Paleogene procel- 
lariiform fossils are rare. Albatrosses (Di- 
omedeidae) range only back to the Upper 
Oligocene (Olson 1985a). Excluding fossils 
of neospecies, the storm-petrels Oceanod- 
roma hubbsi (Oceanitidae) from the late 
Miocene of California, and a species of 
Oceanites from the early Pliocene of South 


Africa, as well as a diving-petrel Peleca- 
noides (Pelecanoididae) from the early Plio- 
cene of South Africa (Olson 1985b), all oth- 
er fossil species of the order are contained 
within the Procellariidae. With the excep- 
tion of the present find, the fossil record of 
the Procellariidae extended back only to the 
early Oligocene of Belgium, by a fossil hu- 
merus described as Puffinus raemdonckii 
(Brodkorb, 1962). Thus, the find of a fossil 
close in morphology to Pterodroma from a 
marine late Eocene locality in Louisiana is 
of particular interest, and extends the range 
of the order Procellariiformes back one geo- 
logic epoch. 

The fossil reported here is the distal end 
(28.5 mm) ofa left tibiotarsus (Fig. 1), CC VC 
(Centenary College Vertebrate Collection) 
001004, collected by BMP during February 
of 1984, from a pit in an oil well field west 
of the village of Tullos, Louisiana (La Salle 
Parish); west of the Union Pacific Railroad, 
but within the “‘city” limits. The locality is 
from the late Eocene Jackson stage (early 
Priabonian age) of the Tulos Member of the 
Yazoo Formation (Murray 1961). 

Other amniote vertebrates from Tullos 
include a zeuglodontid whale (either Zygo- 
rhiza kochii or Basilosaurus cetoides) (Har- 
ris & Veatch 1899), and the giant sea snake 
Pterosphenus schucherti (McPherson & 


750 


Fig. 1. 
petrel-like (Pterodroma-like) bird (CCVC #001004) 
herein reported. Actual length of fossil, 28.5 mm. Left, 
anterior view; right, posterior view. 


Distal end of left tibiotarsus of late Eocene 


Manning, 1986), which is thought to have 
reached a length of some seven to eight me- 
ters. Additional vertebrates include eight 
species of Selachii, three species of Batoi- 
dea, and four teleost fishes (McPherson & 
Manning 1986). In addition, there is a large 
fauna of benthic foraminifera, some plank- 
tonic foraminifera (Godfrey 1983), two 
mollusks and four echinoderms (fauna list- 
ed in McPherson & Manning 1986). The 
Tullos fauna is similar to that of two other 
Yazoo Formation localities, Montgomery 
Landing and Copenhagen (Beard 1978). 
With the exception of the procellariiform 
bird described here and a marine crocodil- 
ian from the Copenhagen locality (Beard 
1978), the sharks, rays, teleost fishes, and 
whale all appear in various localities (Mc- 
Pherson & Manning 1986). Thus, the Tullos 
site represents a marine setting that depos- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ited as its primary facies a prodeltaic clay, 
probably on the continental shelf far from 
the Eocene coast line of the Gulf of Mexico. 
The relative absence of planktonic forami- 
nifera in the fauna is thought to be account- 
ed for by turbidity and relatively shallow 
water depths (Godfrey 1983). 

The distal tibiotarsus from the Tullos lo- 
cality agrees most favorably in size and 
morphology with Pterodroma externa, the 
White-necked Petrel of the South Pacific 
(casual off Pacific coast of Middle America, 
A.O.U., 1983), differing only in minor mor- 
phological detail. Measurements, in mm, of 
the fossil, compared with means from ten 
specimens (five male and five female) of P. 
externa are as follows: 


Greatest width across __ fossil 5.9; P. ex- 


condyles: terna 5.87 

Width external condyle: fossil 6.0; P. ex- 
terna 6.09 

Width internal condyle: fossil 5.2; P. ex- 
terna 5.41 

Least width shaft: fossil 3.2; P. ex- 
terna 2.89 


Morphologically, the fossil differs from P. 
externa, as follows: tendinal groove broader 
and extends further proximally up shaft; 
slighter greater excavation of tendinal groove 
at base of tendinal bridge; external condyle 
more greatly excavated; and less notching 
and excavation on mid-margin of internal 
condyle. Because the tibiotarsi of many spe- 
cies of Pterodroma are very similar mor- 
phologically, comparison specifically with 
P. externa is not thought necessarily to in- 
dicate a specific close relationship with that 
species, rather to indicate the general size 
and proportions of the fossil. 

Feduccia examined all of the genera in 
the collections of the Smithsonian’s Na- 
tional Museum of Natural History, and con- 
cluded that generic allocation within the 
Procellariiformes based on the distal tibio- 
tarsus is possible, but identification of this 
fossil must remain tentative until additional 


VOLUME 106, NUMBER 4 


material is discovered. The fossil agrees in 
general morphology with the species of Pte- 
rodroma examined, and differs from other 
procellariform genera. The tibiotarsus of 
Calonectris has the anterior intercondylar 
area much more excavated with a deeper 
tendinal groove; in Pachyptila, the external 
condyle in external view is elevated around 
the rim and the condylar region is not as 
tapered as in Pterodroma. In Bulweria, there 
is relatively less intercondylar distance and 
the tendinal groove is less deeply excavated. 
Procellaria is a larger genus, and the distal 
condylar region ends with slight internal in- 
flexion, and the internal condyle is rela- 
tively wider. The species of Puffinus are also 
relatively large, and the shaft of the tibio- 
tarsus is relatively broader as it meets the 
condylar region, and is more tapered than 
in Pterodroma. 

With the exception of this fossil and two 
species of Fulmarus, all other Tertiary Pro- 
cellariidae from the Northern Hemisphere 
have been referred to the genus Puffinus (Ol- 
son 1985a), and to our knowledge this is the 
only Tertiary procellariiform reported any- 
where between Bone Valley, Florida, and 
Argentina. Olson (1985a) examined a hu- 
merus of a Bulweria-like bird from the up- 
per Oligocene of South Carolina, and con- 
cluded that, ‘“... some diversity of 
shearwater and petrel-like birds existed in 
the Atlantic as early as the Oligocene.”’ With 
the discovery of the late Eocene Tullos bird 
described herein, it is apparent that this di- 
versity extends much further back in time, 
as expected, and the probability of finding 
many more earlier procellariiform fossils ap- 
pears to be very good. 


Acknowledgments 


We thank Storrs L. Olson, Division of 
Birds, Department of Vertebrate Zoology, 
National Museum of Natural History, 
Smithsonian Institution, for providing ac- 
cess to the bird skeletal collections under 
his care. Storrs L. Olson and Kenneth I. 


fey | 


Warheit provided helpful comments on the 
manuscript. The photographs were taken by 
Susan Whitfield. 


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North American birds. 6th ed. American Or- 
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Beard, S.Q. 1978. Macrofaunal ecology, climate, and 
biogeography of the Jackson Group in Louisiana 
and Mississippi. Unpublished MS thesis, 
Northeast Louisiana University, Monroe, x + 
159 pp., 4 pls. 

Brodkorb, P. 1962. The systematic position of two 

Oligocene birds from Belgium.— Auk 79:706- 

107% 

‘963. Catalogue of fossil birds. Part I.—Bul- 
letin of the Florida State Museum 7(4):179-293. 
Godfrey, W.M. 1983. Foraminifera of Jackson Stage 

sediments of Montgomery Landing, Louisiana. 
Unpublished MS thesis, Louisiana State Uni- 
versity, Baton Rouge, xi + 160 pp., 14 pls. 

Harris, G. D., & A. C. Veatch. 1899. A preliminary 
report on the geology of Louisiana. Section 2, 
General Geology, pp. 45-138. Report of the 
Geological Society of Louisiana. Part 5. State 
Experimental Station, Baton Rouge, 354 pp. 

McPherson, A. B., & E. M. Manning. 1986. New 
records of Eocene sea snakes (Pterosphenus) 
from Louisiana. Pp. 197—207 in J. A. Schiebout 
& W. van den Bold, eds., Proceedings, 1986 
Symposium, Gulf Coast Association of Geolog- 
ical Societies, Baton Rouge, Louisiana. 

Murray, G.E. 1961. Geology ofthe Atlantic and Gulf 
Coastal Province of North America. Harper & 
Brothers, Publishers, New York, 692 pp. 

Olson, S. L. 1985a. The fossil record of birds. Pp. 

79-238 in D. S. Farner, J. R. King, & K. C. 

Parkes, eds., Avian biology, volume 8. Academ- 

ic Press, New York and London, 238 pp. 

1985b. Early Pliocene Procellariiformes 

(Aves) from Langebaanweg, south-western Cape 

Province, South Africa.—Annals of the South 

African Museum, 95(3):123-145. 

—., &D.C. Parris. 1987. The Cretaceous birds 
of New Jersey.—Smithsonian Contributions to 
Paleobiology, 63:1-—22. 


(AF) Department of Biology, Coker Hall 
CB#3280, University of North Carolina, 
Chapel Hill, North Carolina 27599-3280, 
U.S.A.; (ABM) Department of Biology, 
Centenary College of Louisiana, P.O. Box 
41188, Shreveport, Louisiana 71134-1188, 
U.S.A. 


PROC. BIOL. SOC. WASH. 
106(4), 1993, pp. 752-761 


A NEW GENUS AND SPECIES OF RAT FROM 
BORNEO (RODENTIA: MURIDAE) 


Louise H. Emmons 


Abstract. — A new genus and species of arboreal rat from Sabah, Pithecheirops 
otion, 1s described. It is most closely allied to the Sunda Shelf endemic genus 
Pithecheir, but it differs from this genus in lacking both diagnostic features of 
the arrangement of the inner ear bones, and the inflation of the auditory bulla. 


The fragmented lands of the Sunda Shelf 
(landmasses and islands from 10°N to 10°S 
and about 96°E to 120°E that are united by 
seas shallower than 200 m) formed a single, 
dry continental landmass or were connected 
to varying degrees during epochs of low sea 
levels during the Pleistocene. At least 14 
genera and 40 species of native murid ro- 
dents are known from this region. Five gen- 
era are endemics narrowly restricted to Sun- 
daland (Musser & Newcomb 1983). Over 
the past two decades the systematics of these 
and other Asian genera has been clarified in 
an extensive series of publications by Guy 
Musser. As a result, it has become relatively 
easy to recognize new taxa and key mor- 
phological features. The Sundanese genus 
Pithecheir Cuvier, 1838 includes two spe- 
cies, P. melanurus Cuvier, 1838 known only 
from Java, and P. parvus Kloss, 1916 known 
only from West (Peninsular) Malaysia. 
Characters of the genus and species were 
outlined by Kloss (1916), Muul & Lim 
(1971), and Musser & Newcomb (1983). In 
1991, I collected a rat closely allied to Pi- 
thecheir on the island of Borneo. This spec- 
imen has several features that warrant dis- 
tinguishing it within both a new species and 
genus. 


Materials and Methods 


Specimens examined are in the National 
Museum of Natural History (USNM), 
Smithsonian Institution, Washington D.C. 
Many Asian genera of Muridae were ex- 


amined for comparison but only the indi- 
vidual specimens of the genus Pithecheir are 
listed: Pithecheir melanurus USNM 239661 
(1). Java, Born at Museum Buitenzorg, Bo- 
gor, locality unknown; Pithecheir parvus 
USNM 488796 to 488819 (24). Malaysia, 
Selangor, various localities. 

Terminology of tooth cusps and cranial 
features follows Musser & Newcomb (1983) 
and for the inner ear, Voss (1988) and Carle- 
ton (1980). Abbreviations for lengths in text 
are as follows: HB, head and body; T, tail; 
HF, hindfoot; E, ear. 


Pithecheirops, new genus 


Type species.—Pithecheirops otion, new 
species. 

Included species. —The type species only. 

Etymology.— A combination of the Greek 
sufhx -ops, having the appearance of, with 
the generic name Pithecheir, of a genus of 
Sundaland rats. 

Diagnosis. —Morphologically a strongly 
arboreally-adapted rat with: 1) fur long, 
dense and soft, body fur extending well onto 
base of tail; 2) tail robust, slickly naked, and 
at least partially prehensile; 3) hindfeet 
broad, with large pads, hallux separated by 
a gap from other digits, with a claw and 
toepad expanded medially, forefeet also 
broad, with large palmar pads, pollux with 
a broad shiny nail; 4) cranium with no post- 
glenoid vacuity; 5) supraorbital ridges 
strongly flared; 6) squamoso-mastoid fora- 
men absent; 7) auditory bullae small, with 


VOLUME 106, NUMBER 4 


no inflation, such that medial base of bulla 
has a large exposed contribution from pe- 
trosal; 8) inner ear with orbicular apophysis 
present and manubrium of malleus tipped 
anteriorly (parallel configuration); 9) pars 
flaccida present; 10) upper molar M! with 
labial cusps t6 and t9 reduced and joined 
by an enamel ridge; 11) upper M? with la- 
bial cusps t3 and t9 absent; 12) first two 
lower molars (the third lower molar un- 
known) with chevron-shaped pairs of sep- 
arated cusps. 


Pithecheirops otion, new species 


Holotype. —USNM 574500, a juvenile 
male, collected 21 Sep 1991 by Louise H. 
Emmons, Field no. LHE 673. A skin with 
right feet only; a skull; and the entire body, 
with left feet and organs, in fluid. The top 
of the head was damaged in the trap, making 
a large hole in the skin, which is sewn up. 
The animal was kept alive for a day and 
during this time the top layer of bone was 
resorbed from the skull under the wound. 
The lower right M1 is missing, apparently 
congenitally. M3 is just beginning to erupt 
and lies below the level of the gumline. The 
holotype is the only known specimen. 

Type locality. —Malaysia: State of Sabah; 
Danum Valley Field Centre, 4°58'N, 
117°48’E (approximately 75 km W Lahad 
Datu), elevation about 150 m. About 1 km 
NE of the field centre by road, and about 
500 m NW of the road on a study plot trail. 

Habitat.—The holotype was trapped at 
1.5 m on top of a large stump in dense viny 
roadside secondary brush on an abandoned 
logging road. The forest section had been 
selectively logged from primary forest in 
1989. Primary lowland dipterocarp forest 
still occurs within 600 m and is the domi- 
nant vegetation type of the entire surround- 
ing region. 

Etymology. — From the Greek ofion, little 
ear, referring to the small auditory bullae. 

Diagnosis.—The same as for the genus. 

Description. —Because the holotype is a 
juvenile, pelage and body and cranial pro- 


753 


portions can be expected to differ from those 
of adults. Body measurements: HB = 113, 
£ = 11/7, HF = 25. — 15; WT = 36‘e. 
Cranial measurements: total length = 32.2; 
condylo-basal length = 29.4; least interor- 
bital constriction = 6.0; diastema = 8.4; zy- 
gomatic breadth = 16.9; nasals length = 
10.8; auditory bulla length = 5.3; alveolar 
length toothrow 7.3; M! length = 3.4; M! 
width = 2.0. Testes abdominal. Head and 
body covered with long, soft, dense and 
slightly wavy hair. Hair dull and not glossy, 
about 1.0 cm at midback, guard hairs to 1.5 
cm. Body hair extends prominently onto the 
tail base for 2.5 cm; legs thickly haired to 
the wrists and heels (Fig. 1). Ears thickly 
haired at the base and thinly haired on both 
surfaces at the tips. Dorsal fur rusty-red- 
dish, slightly paler on sides than midback. 
Guard hairs with broad brown subterminal 
bands that impart a dusky cast to red up- 
perparts. Base of dorsal fur dark slate gray 
sharply demarcated from the paler tips. Un- 
derparts whitish, with hair white to the roots. 
Throat and abdomen faintly tinged pale 
rusty. Inner forelegs whitish, inner thighs 
rusty with pale hair base. Forefeet clothed 
above with pure rusty hairs, hindfeet with 
darker, more brownish-red hairs except 
whitish hairs at toetips. Soles of feet and 
toetips unpigmented. Toes pigmented above 
and below giving feet a dusky cast. All digits 
except pollux with curved, sharp claws. Pol- 
lux with a broad, shiny nail. Vibrissae dark 
brown, stiff and robust, the longest reaching 
the shoulder. Genal and superciliary groups 
sparse, including but one longer whisker in 
each. Tail robust, unicolored brown, smooth 
and slightly shiny, thick at the tip, with scales 
in narrow, even rings. In life it was at least 
semi-prehensile, the rat twisted it around 
branches (Fig. 1) but was not seen to coil it. 
On the dorsal surface of the tailtip, there 
seems to be a slightly specialized hairless 
region of wider scales that could form a grip- 
ping surface. The feature is unclear due to 
wrinkling. 

Cranium, viewed dorsally, with wide in- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Pig: 


terorbital region (Fig. 2); supraorbital ridges 
strongly flared into shelves; squamosal with 
a salient point at parietal-frontal suture. Na- 
sals broad anteriorly, tapering only slightly 
to a straight posterior border. Viewed lat- 
erally (Fig. 3), zygomatic arches dip deeply 
ventrally in a smooth curve, slightly oc- 
cluding view of the posterior molar. Maxilla 
deep at its lowest point below jugal. No 
postglenoid vacuity, auditory bulla solidly 
fused to squamosal (Fig. 4). No squamoso- 
mastoid foramen, but a small indentation 
in the suture at the place where this foramen 
would occur (Fig. 4). Ventrally (Fig. 2), au- 


Pithecheirops otion holotype. Note use of prehensile tail. 


ditory bulla tight to the basioccipital, with 
no medial vacuities except under the eusta- 
chian tube. Foramen ovale small (possibly 
due to youth of the specimen), with no ex- 
ternal strut of the alisphenoid (Fig. 4). Pos- 
terior opening of alisphenoid canal well an- 
terior to auditory bulla, separated from it 
by a shelf of bone. Pterygoid fossa shallow, 
but likely to deepen with age. Incisive fo- 
ramen reaching to about level of anterior 
edge of first molar; premaxillary portion of 
septum within long, about *% of length of 
foramen. Posterior palatine foramen slight- 
ly posterior to first root of second molar. 


VOLUME 106, NUMBER 4 


735 


Fig. 2. A) Pithecheir parvus adult female USNM 488803; B) P. parvus juvenile/subadult USNM 488817; C) 
P. melanurus juvenile USNM 239661; D) Pithecheirops otion holotype, juvenile USNM 574500. The juvenile 
P. parvus is older than the P. otion, as its third molar is fully erupted. The top of the braincase of D) is damaged 
from a wound. All photos on each figure are to same scale. 


Ectotympanic portion of the auditory bulla 
not hypertrophied, petrosal exposed. Inner 
ear with orbicular apophysis present and 
manubrium of malleus tipped anteriorly 
(parallel configuration) and pars flaccida 
present. 

Incisors pale orange, almost whitish, ap- 
proximately orthodont. Lower molars with 
cusps forming a series of chevron-shaped 


rows of pairs, labial and lingual cusps sep- 
arated, of approximately equal size (Fig. 5). 
Upper molars likewise composed of dis- 
cretely separated cusps (Fig. 5), a central 
row of large cusps flanked on either side by 
rows of smaller cusps. Lingual cusps tl, t4, 
and t7 of M! and M? large and of equal size. 
Large medial cusps t2, t5, and t8 of M', and 
t2 and t5 of M’, also of equal size and spac- 


756 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. A) Pithecheir parvus, adult; B) P. parvus juvenile; C) P. melanurus juvenile; D) Pithecheirops otion 


juvenile holotype. The same specimens as Fig. 2. 


ing. On M!, t3 widely separated from t6; t6 
and t9 reduced, close together, and joined 
by an enamel crest. T1 bis present as a tiny 
shelf. On M7’, t3 and t9 absent and t6 large. 


Comparison with Other Species 


Musser & Newcomb (1983) compared and 
discussed many of the characters of Pithe- 
cheir in relation to other Sundanese genera, 
and included a hypothesis for the polarity 
of the characters they define. The following 
section draws frequently on their work. 

Pithecheirops otion shares with Pithecheir 
spp. characters 1-5 and 12 of the diagnosis, 
and differs from Pithecheir in characters 6— 
Ihe 

Externally, Pithecheirops otion is not dis- 
tinguishable from Pithecheir spp. on the ba- 
sis of the juvenile specimen. The following 
differences await verification from a series. 
The hind and forefeet of Pithecheirops are 
dusky above, those of Pithecheir white or 
whitish. The tail of Pithecheirops otion seems 
much shorter: a similar-sized Pithecheir 


parvus (USNM 488796) has head and body 
length = 122 mm, tail = 157 mm (T = 129% 
of HB), and a younger one (USNM 488800) 
85 mm and 95 mm respectively (112%); 
versus 113 mm and 117 mm for the Pi- 
thecheirops (104%). The mean tail length of 
adult Pithecheir melanurus (126% of HB) is 
relatively much longer than that of P. parvus 
(113%, adult measurements from Musser & 
Newcomb 1983). A prehensile tail is found 
in no other Sundanese genus but Pithecheir. 
Two New Guinea rats, Pogonomys and Chi- 
ruromys have this derived character. The 
tail of Pogonomys is hairier distally, and a 
section of its dorsal tip is hairless and spe- 
cialized into a gripping surface of broad 
scales. There is a lesser tendency towards 
this condition in both Pithecheirops and Pi- 
thecheir. 

Pithecheirops, like Lenothrix and Pi- 
thecheir, has a claw on the hallux, rather 
than a nail as in the specialized arboreal 
genera Chiropodomys, Hapalomys, Kadar- 
sonomys and Abditomys (Musser 1982, 
Musser & Newcomb, 1983). The claw on 


VOLUME 106, NUMBER 4 


Kp W | 


Fig. 4. Region of the auditory bulla A) Pithecheir parvus juvenile/subadult; B) P. melanurus juvenile; C) 
Pithecheirops otion holotype. The same specimens as in Fig. 2. Note the squamoso-mastoid foramen (arrow) in 
B), its absence in C); the perpendicular malleus partially visible in A), the ossified area of the pars flaccida in 
A) and B), and lack of same in C); and the similarity between all crania in the foramen ovale, and postglenoid 


regions. 


the hallux of P. otion seems narrower, larger 
and less modified than that of Pithecheir, 
which is short, with a broad base that shows 
a tendency toward nailishness (Pithecheir 
was listed as having a nail on the hallux in 
Musser 1982). 

The cranium is similar in size and shape 
to that of Pithecheir parvus of similar dental 
stage but shorter than one of P. melanurus 
(Figs. 2, 3). The most dramatic difference 
between the crania of Pithecheirops and Pi- 
thecheir is the lack of inflation of its ecto- 
tympanic auditory bulla (Fig. 2). Greatly 
enlarged bullae are rare among Asian Muri- 
dae. Of 14 Sunda Shelf genera, they occur 
in two others, both arboreal, Kadarsanomys 
and Hapalomys (Musser & Newcomb 1983). 
The exposed wedge of petrosal at the medial 
base of the auditory bulla in Pithecheirops 
is absent in Pithecheir, where it has been 
suppressed by hypertrophy of the ectotym- 


panic. Consequently, the carotid canal 
emerges partway down the medial wall of 
the bulla in the former, but above the base 
of the bulla in the latter. Likewise, the pos- 
terior opening of the alisphenoid canal is 
well forward of the small bulla of P. otion 
but crowded right under the anterior edge 
of the ballooning ectotympanic of Pithe- 
cheir spp. Both genera, however, share a 
general absence or reduction of all vacuities 
that may surround the base of the auditory 
bulla, such as in the carotid, alisphenoid, 
and squamosal regions (Figs. 2, 4). Larger 
vacuities are present in all other Sundaland 
genera. Lack of a postglenoid vacuity was 
deemed the plesiomorphic state by Musser 
& Newcomb (1983) and Pithecheirops shares 
this with about half of the Sundanese murid 
genera. 

The perpendicular manubrium and as- 
sociated lack of orbicular apophysis of the 


758 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 5. Upper and lower left toothrows of like-aged individuals of A) Pithecheir parvus juvenile USNM 
488800; B) P. melanurus; and C) Pithecheirops otion. B) and C) the same specimens as in Fig. 2. 


VOLUME 106, NUMBER 4 


malleus and the lack of a pars flaccida of 
the tympanic membrane (see Carleton 1980 
and Voss 1988 for discussions of these char- 
acters) in Pithecheir are features found only 
in Hapalomys among 23 genera of Asian 
murids examined (see also Carleton & Mus- 
ser 1984, for other incidences). Pithechei- 
rops therefore has the more common con- 
dition for these features (pars flaccida 
present, manubrium of malleus in “‘paral- 
lel’’ configuration). 

A squamoso-mastoid foramen (Fig. 4) is 
a derived feature present only in Mus and 
Pithecheir of Sunda Shelf mice (Musser & 
Newcomb 1983). It is absent in the holotype 
of P. otion, but there is a suggestion of a 
notch in the suture where it would occur. 
In Pithecheir the posterior palatine foramen 
seems more anteriorly positioned than in 
Pithecheirops and the premaxillary portion 
of the septum in the incisive foramen is 
shorter. 

Pale incisors (should this character prove 
constant in P. otion), is a derived state pre- 
viously noted only in Bery/mys among Sun- 
daland murids (Musser & Newcomb 1983). 

The upper molars are in general form like 
those of Pithecheir, but differ in important 
details. In both species of Pithecheir, cusps 
t6 and t9 of M! are usually well separated 
by a notch, but as pointed out by Musser & 
Newcomb (1983) an occasional individual 
of P. parvus shows a tendency for these cusps 
to join on one or both sides; in P. otion they 
are connected by an enamel crest. In M? of 
P. parvus t3 and t9 are prominent nubbins 
or columns, in P. melanurus t3 seems great- 
ly reduced (only one specimen seen) while 
t9 is prominent. Both are absent in P. otion. 
Musser & Newcomb (1983) consider the 
presence of t3 and t9 to be primitive states, 
absence of these derived, while presence of 
t7 is derived. t7 is smaller in P. otion than 
in Pithecheir spp. The lower molars are sim- 
ilar to those of Pithecheir except the pos- 
terior labial cusplet of M, is slightly larger, 
and the anterior labial cusplet slightly small- 
er. 


759 
Discussion of Systematic Relationships 


The characters of the genus Pithecheir, 
apart from a short comment listing three 
characters supporting specific status of P. 
melanurus (Muul & Lim 1971), have been 
discussed recently only by Musser & New- 
comb (1983). The teeth were discussed by 
Misonne (1969). To give a perspective on 
the relationships of the new genus, it is use- 
ful to redefine the genus Pithecheir. The fol- 
lowing list includes two characters men- 
tioned by Muul & Lim (1971), those 
discussed by Musser & Newcomb (1983), 
Kloss (1916) and some reported here. Be- 
cause only one subadult cranium of P. me- 
lanurus was available for the present study 
(skin on body in fluid in poor condition), I 
rely on published measurements and illus- 
trations of that species (Musser & Newcomb 
1983). 

The genus Pithecheir is characterized by: 
1) A naked, prehensile tail, scales of the 
terminal, dorsal 0.5 cm expanded to form 
a smooth surface; 2) arboreally adapted feet; 
hallux widely separated from other toes, 
toepad large, expanded medially, with a 
claw; 3) ectotympanic of auditory bulla in- 
flated; 4) tympanic membrane with no pars 
flaccida, this region ossified; 5) malleus with 
no orbicular apophysis, manubrium per- 
pendicular; 6) squamoso-mastoid foramen 
present; 7) postglenoid vacuity absent; 8) 
rows of major cusps on cheek teeth with 
cusps separate, not fused into laminae in 
unworn teeth; 9) upper molars with an even 
row of large, approximately equal-sized lin- 
gual cusps tl, t4, t7 on each tooth; 10) labial 
cusps of upper molars reduced and irregular 
in size, t6 large, t3 large in M', reduced in 
M?, t9 reduced but present in M! and M?, 
t6 and t9 of M! usually separate in unworn 
teeth. 

Pithecheirops otion is clearly quite closely 
related to Pithecheir by many traits, includ- 
ing both overall appearance, which results 
from many similarities in features such as 
texture, color and distribution of the pelage, 


760 


shape of ears, feet, and body, and appear- 
ance of the head and vibrissae. It shares with 
Pithecheir the derived features of a prehen- 
sile tail, flared supraorbital ridges, long in- 
cisive foramina, and arrangement of the lin- 
gual cusps of the upper molars. It also shares 
a number of plesiomorphic traits that are 
nonetheless unusual among Sundanese gen- 
era, including absence of a postglenoid va- 
cuity, virtually identical design of the lower 
molars and central cusps of the upper mo- 
lars. Pithecheirops does not seem to closely 
resemble any other genus but Pithecheir. In 
tooth characters and in lesser inflation of 
the ectotympanics, Pithecheir melanurus is 
closer to Pithecheirops otion than is Pithe- 
cheir parvus. It is also closer in geographic 
range. 

Of the major characters by which it differs 
from Pithecheir, and which warrant its 
placement in a separate genus, the character 
states in Pithecheirops, including lack of in- 
flated ectotympanic, lack of squamoso- 
mastoid foramen, and lack of t3 on M2, are 
all considered by Musser & Newcomb (1983) 
to be the less derived conditions. In con- 
trast, the lack of cusps t3 and t9 on the 
second molar are apomorphic states (Mus- 
ser & Newcomb 1983). A perpendicular 
malleus and lack of a pars flaccida are con- 
sidered by Carleton (1980) to be plesiomor- 
phic, therefore Pithecheirops has the apo- 
morphic condition. However, Voss (1988) 
argues that the evidence is inconclusive for 
these two characters, and I here consider 
the polarity as unknown. Overall, Pithechei- 
rops seems a more primitive relative of Pi- 
thecheir, with a few derived traits, but the 
opposite could be argued. The polarity of 
the inner-ear characters is the pivotal ques- 
tion. 

Of other Asian arboreal rats, only the In- 
dochinese rat Hapalomys shares the inner- 
ear traits of Pithecheir. It also shares a num- 
ber of derived characters associated with 
arboreality (and therefore perhaps homo- 
plastic) with both Pithecheir and Pithechei- 
rops, and some tooth-features (Musser & 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Newcomb 1983). Among Sunda Shelf gen- 
era, Pithecheirops seems closer to Lenothrix 
than is Pithecheir, because it shares the 
characters of the auditory apparatus. The 
polarity of the inner-ear structures is central 
to determining these relationships. The New 
Guinea genus Pogonomys also shares a mix- 
ture of features with Pithecheirops and Le- 
nothrix. More complete analysis of the re- 
lationships of Pithecheirops awaits the 
capture of a good series of specimens. 

From the perspective of the characters of 
all Sunda Shelf rodent genera, Musser & 
Newcomb (1983:543) concluded that: “On 
the Sunda Shelf, Lenothrix may be closer to 
Pithecheir than to any other Sundanese gen- 
era. Lenothrix, and even Pithecheir, have 
the aspects of old endemics, relicts left over 
from an earlier time in the history of the 
Sunda region and the early evolution of rats 
there.’’ One can speculate that specialized 
arboreal habits have allowed the persistence 
of a number of relict genera in the face of 
competition from more recent radiations of 
highly successful advanced rats. 


Acknowledgments 


I thank Yayasan Sabah and the Sabah 
Wildlife Department for hosting my re- 
search in Sabah and at Danum Valley Field 
Centre. The new rat species was collected 
while trapping for treeshrews during an eco- 
logical study. Fieldwork was funded by The 
National Geographic Society and the Dou- 
roucouli Foundation. Michael D. Carleton 
made many helpful comments and pointed 
out the characters of the inner ear. David 
Schmidt photographed the specimens. 


Literature Cited 


Carleton, M. D. 1980. Phylogenetic relationships in 
Neotomine-Peromyscine rodents (Muroidea) 
and a reappraisal of the dichotomies within New 
World Cricetinae. — Miscellaneous Publications 
of the Museum of Zoology, University of Mich- 
igan, No. 157. 

—., & G.G. Musser. 1984. Muroid rodents. Pp. 
289-379 inS. Anderson & J. K. Jones, Jr., eds., 


VOLUME 106, NUMBER 4 


Orders and families of recent mammals of the 
world. John Wiley & Sons. 

Kloss, C. B. 1916. On two rodents new to the fauna 
of the Malay Peninsula, with the description of 
a new subspecies-species, Pithecheirus melanu- 
rus parvus. — Journal of the Federation of Malay 
States Museums 6:249-252. 

Misonne, X. 1969. African and Indo-Australian 
Muridae. Evolutionary trends.—Museé Royale 
de l’Afrique Centrale Tervueren, Zoologie, No. 
72-219 pp. 

Musser, G. G. 1982. Results of the Archbold Expe- 
ditions. No. 107. A new genus of arboreal rat 
from Luzon Island in the Philippines.— Amer- 
ican Museum Novitates No. 2730, 23 pp. 

—, & C. Newcomb. 1983. Malaysian Murids 


761 


and the Giant rat of Sumatra.— Bulletin of the 
American Museum of Natural History 174:327- 
598. 

Muul, I., & Lim Boo Liat. 1971. New locality records 
for some mammals of West Malaysia. — Journal 
of Mammalogy 52:430-437. 

Voss, R.S. 1988. Systematics and ecology of ichthyo- 
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Division of Mammals, Smithsonian In- 
stitution MRC 108, Washington, D.C. 
20560, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(4), 1993, pp. 762-774 


A NEW THOMASOMYS (MAMMALIA: RODENTIA) 
FROM THE PERUVIAN ANDES 


Alfred L. Gardner and Monica Romo R. 


Abstract. — We describe a new species, Thomasomys macrotis, from the east- 
ern slope of the Andes of northcentral Peru (department of San Martin, Rio 
Abiseo National Park). This rodent is a large-bodied, big-eared, but relatively 
short-tailed thomasomyine inhabiting the wet montane forest near the upper 
limit of the continuous forest zone. It is the second large, previously-unde- 
scribed Thomasomys to be found in the park. 


Resumen. —Describimos una nueva especie, Thomasomys macrotis, de la 
vertiente oriental de los andes al norte del Peru (departamento de San Martin, 
Parque Nacional Rio Abiseo). Este roedor thomasomino de cuerpo y orejas 
grandes, pero de cola relativamente corta, habita el bosque humedo montano 
cerca al limite superior de la zona de bosque continuo. Este es el segundo 
Thomasomys grande, anteriormente desconocido, encontrado en el Parque. 


The Peruvian Association for the Con- 
servation of Nature (APECO) recently com- 
pleted a 4-year (1987-1990) biological in- 
ventory of the Rio Abiseo National Park, 
San Martin, Pert. A new giant thomaso- 
myine, 7Thomasomys apeco, was found dur- 
ing the first year and recently described by 
Leo & Gardner (1993). A second species 
represented by one specimen, also collected 
during the first year and sympatric with T. 
apeco, 1s a large, big-eared, and relatively 
short-tailed Thomasomys unlike any thus 
far known. Subsequently, we found three 
additional specimens in the mammal col- 
lection of Louisiana State University Mu- 
seum of Zoology, Baton Rouge (LSUMZ), 
collected in 1981 from a nearby locality in 
the park. The Rio Abiseo National Park, 
centered at approximately 07°45’S, 77°15'W, 
covers 2745 square kilometers on the east- 
ern (Amazonian) slope of the Andes and 
encompasses most of the Rio Abiseo wa- 
tershed, which drains into the Rio Huallaga 
(Leo & Gardner 1993). The western bound- 
ary of the park lies along the border between 
the departments of La Libertad and San 
Martin. The elevational range is from near 


- 1000 to more than 4000 m and includes at 


least five habitat zones according to the 
Holdridge Life Zone classification (Tosi 
1960). Young & Leon (1988) suggest that 
two additional life zones (Tropical Montane 
Wet Forest and Tropical Lower Montane 
Wet Forest) be recognized in the park. Sur- 
veys during the first year of study were at 
higher elevations (from about 3000 to 3600 
m) in Tropical Subalpine Pluvial Paramo 
and Tropical Montane Rain Forest (Paramo 
Pluvial Subalpino and Bosque Humedo 
Montano, respectively; Tosi 1960). The new 
thomasomyine we describe here was caught 
in tropical montane elfin forest habitat; it 
may be known as: 


Thomasomys macrotis, new species 


Holotype. —Adult female, LSUMZ 27286, 
from Puerta del Monte, ca. 30 km NE [fof] 
Los Alisos, ca. 3250 m [Parque Nacional 
Rio Abiseo], San Martin, Peru. Collected 
by Linda J. Barkley on 13 August 1981. The 
holotype is a well-made skin in excellent 
condition with cranium, dentaries, and par- 
tial post-cranial skeleton that includes a 


VOLUME 106, NUMBER 4 


complete left forefoot and complete right 
hind foot (Fig. 1; right forefoot and left hind 
foot remain in the skin). 

Paratypes. — Young adult female, LSUMZ 
27285; adult male, LSUMZ 27287, both 
from the type locality and collected by L. J. 
Barkley on 8 August and 15 August (re- 
spectively), 1981. Young adult female, 
USNM 567243, from Pampa del Cuy, 24 
km NE of Pataz, 3380 m, Parque Nacional 
Rio Abiseo, San Martin, Peru, collected by 
A. L. Gardner on 11 August 1987. 

Distribution. —Known only from elfin 
forest habitat in the Pampa del Cuy Valley. 

Etymology.—The name refers to the un- 
usually large ears, the most obvious feature 
of the species when encountered in the field. 

Diagnosis.—Thomasomys macrotis is a 
large-bodied, big-eared thomasomyine with 
a long hind foot and comparatively short 
(1.34 times head and body; n = 4), basally 
dark brown and distally white tail. It is in- 
termediate in size between the smaller-bod- 
ied, but longer-tailed 7. aureus (Tomes, 
1860) and the much larger 7. apeco, both 
of which are sympatric (Table 1). Diagnostic 
features include broad, spatulate nasals that 
extend only barely beyond the anterior plane 
of incisors and terminate behind premax- 
illary-frontal sutures; postorbital constric- 
tion with rounded upper borders and lack- 
ing ridges (Fig. 2); jugal strongly developed; 
tooth-bearing portion of maxilla unusually 
deep above M1 and tapering posteriorly to 
root zone of M3 where maxilla is dorso- 
ventrally thin (Fig. 3); auditory bullae me- 
dium-sized to large with posterior and up- 
per anterior border of auditory meatus 
unusually inflated (Fig. 4); zygomatic plate 
terminating on a line behind procingulum 
of first upper molar (M1); procingulum of 
M1 with smooth anterior face; anterome- 
dial flexid of first lower molar (m1) shallow 
and narrow (dental terminology follows Reig 
1977; also see Carleton & Musser 1989): 
cusps on all teeth robust and separated by 
relatively narrow flexi and flexids; oblique 
orientation of paralophs, metalophs, and 


763 


Fig. 1. Left pes of the holotype of Thomasomys 
macrotis, new species, LSUMZ 27286. Vertical line 
equals 10 mm. 


metalophids in first two molars, and ento- 
lophid of m1; lateral orientation (at right 
angle to median murid) of entolophid of 
m2; M3 wider than anterior-posterior 
length; posteroloph in m1 straight, not cres- 
cent shaped; m3 relatively broad posteriorly 
with hypoflexid extending across approxi- 
mately three quarters width of tooth; mar- 
gins of lophs, lophids, mures, and murids, 
“‘wrinkled”’ or crenulated (especially evi- 
dent in relatively unworn teeth; see Fig. Sa, 
b). 


764 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Measurements of Thomasomys aureus complex, Thomasomys macrotis new species, and Tho- 
masomys apeco. Linear measurements are in millimeters and mass in grams; values are the mean followed by 
range (in parentheses) and sample size. See “Additional specimens examined” for sources of 7. apeco and T. 
aureus. 


Measurements T. aureus T. macrotis T. apeco 
Length 
Total 374.3 362.2 514.3 
(350-421) 14 (345-387) 4 (469-560) 9 
Tail 220.1 207.0 307.0 
(208-248) 14 (193-219) 4 (279-329) 9 
Hind foot (cu) 36.5 46.0 54.0 
(33-41) 18 (44-48) 4 (50-59) 9 
Ear (from notch) 2321 31.2 29.4 
(21-24) 18 (28-33) 4 (27-31) 9 
Skull (greatest) 38.2 40.9 48.3 
(34.6-41.8) 20 (38.0-42.8) 4 (44.3-51.0) 9 
Condylobasal 35.8 38.7 45.4 
(31.9-40.1) 21 (35.1-41.3) 4 (41.9-47.8) 9 
Condyloincisive 34.1 38.2 44.9 
(30.7—38.6) 21 (34.3-41.0) 4 (41.5—47.6) 5 
Palatilar 15.9 17.8 PAKS 
(13.9-18.4) 21 (16.4-19.2) 4 (20.1—22.6) 5 
Postpalatal 13.9 15.0 7S 
(12.5-16.5) 21 (13.3-16.2) 4 (15.5-18.7) 5 
Incisive foramina 8.1 8.5 10.6 
(7.1-9.1) 21 (7.6-9.1) 4 (9.4—11.7) 9 
Nasal 14.4 14.4 17.9 
(12.4—15.8) 20 (13.1-15.4) 4 (16.3-19.1) 9 
Rostrum: 14.2 15.5 18.3 
(12.4—16.0) 20 (13.8-16.3) 4 (16.1-19.8) 5 
Maxillary toothrow Ta 8.1 9.7 
(6.8-7.8) 21 (8.0-8.3) 4 (9.5—10.0) 5 
Mandible 21.6 22.8 28.5 
(19.5—23.5) 21 (22.3-26.3) 4 (27.0—29.8) 5 
Breadth 
Zygomatic 20.0 22M D558 
(18.6—21.4) 21 (20.4—23.2) 4 (23.6—27.4) 9 
Mastoidal 14.8 16.7 17.9 
(13.9-15.9) 18 (15.9-17.3) 4 (17.2-18.6) 8 
Postorbital 4.8 5.9 52 
(4.3-5.3) 21 (5.5-6.4) 4 (4.9-5.4) 9 
Across molars Ved op 10.0 
(M2-M2) (7.3-8.5) 21 (8.6-9.7) 4 (9.2—10.6) 5 
Palate (post-dental 6.0 G2 7.4 
constriction) (5.4-6.8) 21 (6.2-6.3) 4 (6.9-7.9) 5 
Rostral 6.8 LT 8.9 
(6.1-7.4) 21 (7.2-8.2) 4 (7.8-9.9) 9 
Zygomatic plate 3}. 3.9 4.1 
(2.5-4.0) 21 (3.5-4.2) 4 (3.2-4.4) 5 
Depth 
Braincase 11.1 22 13.4 
(10.2-11.9) 21 (11.5-12.8) 4 (13.0-14.0) 5 
Mass 91.8 bkS 232.0 
(58-136) 9 (64-166) 4 (164-335) 9 


VOLUME 106, NUMBER 4 765 


Fig. 2. Dorsal, ventral, and lateral views of the cranium, and lateral view of the mandible of the holotype 
of Thomasomys macrotis, new species, LSUMZ 27286. Vertical line equals 10 mm. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Lateral view of tooth-bearing portion of left maxillae of Thomasomys macrotis, new species, LSUMZ 
27286 (a), and T. aureus, MLL 390 (5). Horizontal line equals 5 mm. 


Measurements of holotype. —Measure- 
ments are in millimeters and weight (mass) 
in grams; external measurements are those 
made by the preparator recorded on the la- 
bel: Total length, 372; head and body, 153; 
tail, 219; hind foot (with claws), 47; ear (from 
notch), 33; greatest length of skull, 42.8; 
condyloincisive length, 41.0; palatilar length, 
19.2; post palatal length, 16.2; length inci- 
sive foramina, 9.1; zygomatic breadth, 23.0; 
braincase breadth (taken below parietal root 
of zygomatic arches), 17.7; interorbital con- 
striction, 6.0; breadth across molars (M2- 


M2), 9.6; palatal breadth (across post-dental 
constriction), 6.3; rostral breadth, 8.0; ros- 
tral length, 16.2; breadth of zygomatic plate, 
4.2; breadth of mesopterygoid fossa, 2.6; 
depth of braincase (from lower surface of 
basisphenoid and basioccipital to top of cra- 
nium), 12.3; length of dentary, 26.1; length 
of maxillary toothrow, 8.3; alveolar length 
of maxillary toothrow, 8.7; alveolar length 
of mandibular toothrow, 8.6; mass, 140 g. | 

Additional measurements. —See Table 1 
for additional measurements. 

Description. —Fur long (as long as 15.0 


VOLUME 106, NUMBER 4 


767 


Fig. 4. Lateral view of left auditory bullar region of Thomasomys macrotis, new species, LSUMZ 27286 (a), 
and T. aureus, USNM 507266 (b). Horizontal line equals 5 mm. 


mm middorsally) and lax; individual hairs 
with extensive dark-gray base and paler tips. 
Guard hair is also gray based, but broadly 
darker tipped. Dorsum Bone Brown to Se- 
pia (capitalized color terms from Ridgway 
1912) finely streaked with pale brown im- 
parting an agouti pattern. Color darkest 
middorsally becoming gradually paler lat- 
erally where it grades into Sudan Brown or 
Argus Brown over sides and flanks, and 
blends into Pinkish Cinnamon on venter 


and inside of legs. Fur on throat dark gray 
at base, tipped with white, and conspicu- 
ously paler than remainder of venter. Hair 
on margin of lips pale buff; lips otherwise 
blackish and conspicuously darker than col- 
or of throat and base of vibrissae. Area sur- 
rounding eye and anterior to and below eye 
at base of vibrissae also conspicuously darker 
than remainder of head, which otherwise is 
pigmented like the dorsum. Ears large, 
clothed inside and out with dusky brown 


768 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ee 


Fig. 5. Left maxillary (upper row) and mandibular toothrows (lower row) of Thomasomys macrotis, new 
species, USNM 567243, a, b; LSUMZ 27286, c, d; T. apeco, MUSM 7197, e, f; T. aureus, USNM 507266, g, 
h. Vertical line equals 5 mm. 


VOLUME 106, NUMBER 4 


hair, and lack evidence of a pale rim. Head 
also with a weakly-defined Pinkish Cinna- 
mon spot below and just behind ears. Dor- 
sal surface of hind feet dark, colored like 
dorsum of body, but with paler claws and 
digital bristles; metacarpals of forefeet dark 
with paler toes, claws, and digital bristles. 
Tail longer than head and body (126 to 
143%), and uniformly dark brown except 
for terminal one-third to two-fifths where 
both scales and hairs are white. Mystacial 
vibrissae long (longest 65 to 70 mm), a few 
uniformly dark brown throughout, but most 
with broadly pale tips. Superciliary and sub- 
ocular vibrissae few in number and shorter 
than mystacial vibrissae. Hind feet long with 
plantar surface naked to ankle. Hallux 
shortest toe, base of claw extending just be- 
yond metatarsal-phalanx joint of digit II and 
tip of claw not reaching joint between sec- 
ond and third phalanges. Digit II slightly 
shorter than subequal digits III and IV. Claw 
tip of digit V extends to approximately mid- 
length of second phalanx of digit IV (Fig. 
By. 

Skull, although large and robust, typically 
thomasomyine (sensu Steadman & Ray 
1982:15) in dental topography, length of 
hard palate, lack of overhanging postorbital 
ridges (Fig. 2), and presence of strut from 
mastoid extending anteriorly above and 
across ectotympanic to overlap squamosal 
in front of hamular process of squamosal 
(Fig. 4). Rostrum comparatively broad and 
deep, and with tapering, anteriorly-truncat- 
ed and spatulate nasals whose tips protrude 
only slightly beyond anterior plane of upper 
incisors. Dorsal branches of premaxillae 
short, not reaching distal ends of nasals and 
terminating at or anterior to level of zygo- 
matic notch (Figs. 2, 6). Zygomatic plate 
broad, its posterior margin terminating at 
approximate level of protoflexus of M1, but 
its anterior border not occluding openings 
to nasolacrimal capsule when viewed lat- 
erally. Jugal large, its greatest depth ap- 
proximately equal to or greater than dis- 
tance between maxillary and squamosal 


769 


portions of zygomatic arch (Fig. 7). Supra- 
orbital region elevated laterally with round- 
ed, non-overhanging borders and a shallow 
median depression extending from near an- 
terior border of interparietal to anterior third 
of nasals. Dorsal exposure of lacrimals com- 
paratively broad, each with a posterior me- 
dian projection. Alisphenoid strut present, 
but weak (incomplete on left in holotype). 
Auditory bullae medium-sized to large for 
genus. Upper border of incomplete ecto- 
tympanic ring forming auditory meatus (be- 
low second [posterior] dorsal postglenoid 
fossa) notably swollen and its terminus blunt 
(Fig. 4). Ventral surface of periotic broadly 
exposed. Mastoid fenestra small. Squamo- 
sal-alisphenoid groove (for supraorbital 
branch of stapedial artery) and sphenofron- 
tal foramen absent; stapedial foramen small. 
Mesopterygoid fossa of uniform width, not 
expanded anteriorly; its width equal to or 
less than width of either parapterygoid fossa 
(Fig. 2). 

Molar-bearing portion of maxilla unusu- 
ally deep at level of M1 and tapering pos- 
teriorly to level of M3 where the bone is 
dorsoventrally comparatively thin (Fig. 3a). 
Roots of M3 and posterior roots of M2 ex- 
tend through maxilla into orbital fossa. 

Upper incisors slightly procumbent, 
without grooves; upper and lower molars 
large and robust; all except M3 longer than 
wide and forming a graded series (Fig. 5a, 
b). M3 is wider than long. Procingulum of 
M1 lacks any trace of anterior cuspules, 
styles, or cingular ridge. Anterolabial and 
anteromedial conules separated anteriorly 
by a deep, narrow anteromedial flexus whose 
internal limit lies close to corresponding end 
of relatively narrow protoflexus. Antero- 
flexus short and narrow, and separated from 
anteromedial flexus by a small enamel is- 
land. Anteroloph well developed, but com- 
paratively thin and may or may not be ex- 
panded antero-posteriorly at the labial 
margin of M1. Paraflexus long and termi- 
nating before level of medial penetration of 
hypoflexus in M1 and M2, but extending 


770 


(right). Vertical line equals 5 mm. 


posteriorly beyond level of hypoflexus in 
M3. Mesoflexus moderate in length, not 
reaching median mure, and extending across 
tooth approximately one-half the width of 
paracone in M1 and M2. Mesoflexus rela- 
tively longer in M3 and may meet median 
mure. Metaflexus long, reaching postero- 
loph in M1 and M2, but poorly developed 
in M3 as a shallow flexus and one or more 
small enamel islands. Protoflexus and hy- 
poflexus of M1 narrow; the latter longer and 
extending approximately half way across 
tooth in all upper molars. Paraloph joins 
median mure at juncture with mesoloph. 


os 


Fig. 6. Nasals of Thomasomys macrotis, new species, LSUMZ 27286 (left), and 7. aureus, USNM 507266 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ai 
i 4H 
S3 


By me J 
‘ dS Wie 


= 


M3 wider than long (antero-posterior axis), 
with small hypocone and weakly-developed 
to absent metacone. General occlusal pat- 
tern created by protoloph, hypoloph, pos- 
teroloph, and anterior and median mure, 
takes the form of a sigma (2) in upper left 
M1 and M2. The pattern in the center of 
M1 and M2 is an “X”’ with mesoloph and 
anterior median mure forming one leg and 
protocone, protoloph, hypoloph, hypocone, 
and connecting median mure forming the 
other. 

Lower molars also form a graded series 
with m3 the smallest and having a complex 


VOLUME 106, NUMBER 4 


He 


Fig. 7. Lateral view of left zygomatic arches of Thomasomys macrotis, new species, LSUMZ 27286 (a), and 
T. aureus, USNM 507266 (6). Horizontal line equals 5 mm. 


sigmoid cuspid, murid, and lophid pattern 
because of an under-developed entoconid 
(Fig. 5b, d). First lower molar characterized 
by a shallow and narrow anteromedial flex- 
id, and anterolingual and anterolabial con- 
ulids coalesce early in wear. Protolophid 
short, narrow, and not reaching margin of 
tooth. Outer anterior margin of procingu- 
lum in ml dominated by ridge extending 
posteriorly from anterolabial conulid to 
form anterior margin of protoflexid. This 
ridge appears to be characteristic of Thom- 
asomys (s.l.) and has essentially the same 
position as the anterolabial cingulum of m2 
and m3. Anteroflexid obsolete. Metaflexid 
defined by a shallow indentation on lingual 
surface of m1 and a transversely elongated 


internal enamel island. Mesoflexid termi- 
nating before reaching level of inner margin 
of protoflexid. Mesolophid may or may not 
reach lingual margin of m1 and m2; where 
it does, its stylid not fused to either meta- 
conid or entoconid. Entoflexid variable in 
length and tends to be shorter in m1 than 
in m2, where it is transversely in line in- 
ternally with medial penetration of hypo- 
flexid. Metalophids ofall three lower molars 
and the entolophid of m1 oriented slightly 
anteriorly toward the midline. Entolophid 
of m2 oriented at right angle to axis of tooth 
and joins the median mure in line with pos- 
terior enamel margin of hypoflexid. Hy- 
poflexid narrow in all molars. Posterolo- 
phid variable, usually straight in ml, with 


772 


a terminal hook in m2, and crescent shaped 
in m3 with its terminus fused with rudi- 
mentary entoconid and mesolophid. 
Comparisons. — Except for the tail, whose 
terminal third is white, and relatively larger 
ears and hind feet, 7. macrotis resembles 
several species of much smaller Thomaso- 
mys such as T. ischyurus Osgood, 1914, and 
T. incanus (Thomas, 1894). Although av- 
eraging larger (see Table 1), 7. macrotis is 
most similar in size to T. aureus from which 
it can be distinguished externally by darker 
brown and softer fur dorsally and ventrally, 
significantly longer (no overlap in measure- 
ments) ears and hind feet, and bicolored 
relatively-shorter tail (averages 1.34 times 
length of head and body versus 1.43 times 
[1 = 14] head and body in T. aureus). Thom- 
asomys aureus has reddish fur, a distinctly 
ochraceous venter, and a longer and mono- 
colored tail. Cranially, 7. macrotis can be 
readily distinguished from 7. aureus by 
wider nasals that terminate distinctly be- 
hind premaxillae (compare in Fig. 6), 
broader postorbital constriction with more 
rounded borders, larger and deeper jugal 
(Fig. 7), conspicuously deeper maxilla above 
M1 than above M3 (Fig. 3), smooth anterior 
face of procingulum of M1, M3 wider than 
long, shallow anteromedial flexid and 
straight posterolophid on m1, and entolo- 
phid of m2 joining median murid at right 
angle to axis of tooth (Fig. 5). In contrast, 
T. aureus has narrower nasals that extend 
forward of anterior face of incisors and ter- 
minate posteriorly at the same level as pre- 
maxillae (Fig. 6), narrower postorbital con- 
striction with sharper and more elevated 
borders, narrower jugal (Fig. 7b), maxilla 
only little deeper above M1 than above M3 
(Fig. 35), conspicuous anterior cingular ridge 
with accessory cuspules and styles on pro- 
cingulum of M1, M3 longer than wide (Fig. 
5g), deep anteromedial flexid and crescent- 
shaped posterolophid on m1, and entolo- 
phid of m2 joining mesolophid more or less 
in line with axis of tooth (Fig. 5/). In mea- 
surements, 7. macrotis averages larger than 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


T. aureus in most dimensions. In the sam- 
ples at hand, there is no overlap in length 
of ear, hind foot, and maxillary toothrow, 
or in breadth across molars, and mastoidal 
and postorbital breadth, even when young 
adults of the former are included (see Table 
1). 

There is little external, cranial, or dental 
similarity between T. macrotis and T. ape- 
co, except that both have a thick jugal and 
terminally white tail. Thomasomys apeco is 
significantly larger with no overlap in most 
dimensions (compare measurements in Ta- 
ble 1) and has coarser, reddish pelage with 
conspicuous guard hair. Cranially and den- 
tally, 7. apeco shares more features with the 
smaller 7. aureus than it does with T. mac- 
rotis and appears to be an outsized member 
of the JT. aureus complex. Other than size, 
T. macrotis differs from T. apeco in many 
of the same ways that it differs from T. au- 
reus. Despite its overall smaller size, T. 
macrotis 1s larger than T. apeco in length of 
ear and breadth of postorbital constriction 
(Table 1). Thomasomys macrotis shares 
some dental features with the much larger, 
extinct Megaoryzomys curioi (Niethammer, 
1964) known only from the Islas Galapagos 
of Ecuador (Steadman & Ray 1982). These 
features include smooth anterior face of 
procingulum of M1, weakly developed an- 
teromedial flexid in m1 (retained only as an 
internal enamel island in M. curioi), and 
more medial orientation of paralophs, 
metalophs, metalophids and entolophids. 
Nevertheless, on the basis of dental and cra- 
nial features, 7. macrotis is more closely 
related to T. ischyurus Osgood, 1914, and 
related Thomasomys, than with M. curioi 
or members of the 7. aureus complex. 

Remarks. —We refer to specimens iden- 
tified as 7. aureus as the T. aureus complex 
in the text and Table 1 because we recognize 
that the name is used currently for three or 
more morphologically similar species. A 
thorough review and examination of the 
group is required before the number of spe- 
cies and their relationships can be estab- 


VOLUME 106, NUMBER 4 


lished. Thomasomys macrotis is the second 
to be described of at least seven new mam- 
mals discovered during the Rio Abiseo fau- 
nal inventory. It is remarkable that the fau- 
na of this region contains such a high number 
of new taxa, and unfortunate that most of 
them are represented by single specimens. 
Label information for LSUMZ 27285, a 
female collected 8 Aug 1981, includes the 
notation that she was nonparus with a vagi- 
nal plug. Corresponding information for two 
males is as follows: LSUMZ 27286, 13 Aug 
1981, testes = 9.0 mm, abdominal; LSUMZ 
27287, 15 Aug 1981, testes = 17.0 mm, 
scrotal. USNM 567243, a female, showed 
no overt evidence of reproductive activity 
when collected on 11 Aug 1987. 
Additional specimens examined. —Other 
specimens we examined are deposited ei- 
ther in the collections of the Museo de His- 
toria Natural, Universidad Nacional Mayor 
de San Marcos, Lima, Pera (MUSM); the 
National Museum of Natural History, 
Washington, D.C., U.S.A. (USNM [Divi- 
sion of Mammals], USNM-P [Paleobiolo- 
gy]); or are indicated by field numbers (MLL 
[Mariella Leo L.] and MRR [Monica Romo 
R.]). The majority of the latter will be de- 
posited in the MUSM, which previously was 
commonly referred to in the literature as 
the Museo de Historia Natural “Javier Pra- 
do.” Thomasomys aureus complex. CO- 
LOMBIA: Cundinamarca, Bogota, USNM 
251957, topotype of Thomasomys princeps 
(Thomas, 1900); Cundinamarca, Choachi, 
USNM 251956; Cundinamarca, Laguna 
Vergon [=Laguna del Verjon], USNM 
251976. ECUADOR: Pichincha, Rio Con- 
dor Huachana, 3.45 km NE of Lloa, USNM 
513588, 513589. PERU: San Martin, Rio 
Montecristo, ca. 28 km NE of Pataz, Parque 
Nacional Rio Abiseo, MLL 312; San Mar- 
tin, Puerta del Monte, ca. 26 km (at 60°) 
from Pataz, Parque Nacional Rio Abiseo, 
MLL 249; San Martin, Las Palmas, ca. 32 
km NE of Pataz, Parque Nacional Rio Abi- 
seo, MRR 579 and 594, MLL 340 and 343; 
San Martin, Las Papayas, ca. 32 km NE of 


(73 


Pataz, Parque Nacional Rio Abiseo, MLL 
390; Junin, Rio Palca, 15 km W of San 
Ramon, USNM 507265 and 507266; Cuz- 
co, Torontoy, 3260 m, USNM 194818, 
194820, and 194821; Cuzco, Tocopoque 
[=Tocopogueyu], Occobamba Valley, 
USNM 194822-194828. Thomasomys ape- 
co. PERU: San Martin, Valle de Los Cho- 
chos, ca. 25 km NE of Pataz, 3280 m, Par- 
que Nacional Rio Abiseo, MUSM 7197, 
7199; San Martin, Pampa del Cuy, ca. 24 
km NE of Pataz, 3260-3380 m, Parque Na- 
cional Rio Abiseo, MUSM 7196, 7198, 
7201, 7202, 7203, 7204; San Martin, Puerta 
del Monte, ca. 26 km (at 60°) from Pataz, 
3250 m, Parque Nacional Rio Abiseo, 
MUSM 7200. Megaoryzomys curioi. EC- 
UADOER: Islas Galapagos, Isla Santa Cruz, 
USNM-P 284204 (15 molars), USNM-P 
284213 (mandible with m2 and m3), 
USNM-P 284276 (right M1), USNM-P 
284283 (right mandible); USNM-P 284287 
(right ml), USNM-P 284343 (mght man- 
dible with all molars), USNM-P 284346 
(right mandible with m1). 


Acknowledgments 


The inventory of Parque Nacional Rio 
Abiseo was conducted by the Asociacion 
Peruana para la Conservacion de la Natura- 
leza (APECO), supported by The Abiseo 
River National Park Research Project from 
the University of Colorado, the David and 
Lucile Packard Foundation, and the Pew 
Charitable Trust. We gratefully acknowl- 
edge the assistance of the many persons who 
were part of the inventory team. We also 
thank the personnel from the Parque Na- 
cional Rio Abiseo for their help at several 
stages of the project, as well as the Direccion 
General Forestal y de Fauna for authorizing 
and facilitating research in the park. We are 
indebted to Dr. Mark S. Hafner, Museum 
of Natural Sciences, Louisiana State Uni- 
versity (LSUMZ), Baton Rouge, and to Dr. 
Clayton E. Ray and Mr. Robert W. Purdy 
of the Department of Paleobiology, Nation- 
al Museum of Natural History, who made 


774 


critical material available. Guy Musser, 
Bruce Patterson, and Robert Fisher re- 
viewed drafts of this report and made nu- 
merous helpful suggestions. 


Literature Cited 


Carleton, M. D., & G. G. Musser. 1989. Systematic 
studies of oryzomyine rodents (Muridae, Sig- 
modontinae): a synopsis of Microryzomys.— 
Bulletin of the American Museum of Natural 
History 191:1-83. 

Leo L., M., & A. L. Gardner. 1993. A new species 
of a giant Thomasomys (Mammalia, Muridae, 
Sigmodontinae) from the Andes of northcentral 
Peru.— Proceedings of the Biological Society of 
Washington 106:417-428. 

Reig, O. A. 1977. A proposed unified nomenclature 
for the enamelled components of the molar teeth 
of the Cricetidae (Rodentia).—Journal of Zo- 
ology, London 181:227-241. 

Ridgway, R. 1912. Color standards and color no- 
menclature. Washington, D.C., iv + 43 pp., 53 
pls. 

Steadman, D., & C. Ray. 1982. The relationships of 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Megaoryzomys curioi, an extinct cricetine ro- 
dent (Muroidea: Muridae) from the Galapagos 
Islands, Ecuador.—Smithsonian Contributions 
to Paleobiology 51:1-—23. 

Tosi, J. A., Jr. 1960. Zonas de vida natural en el 
Pera.—Instituto Interamericano de Ciencias 
Agricolas de la OEA Zona Andina, Boletin Téc- 
nico No. 5, vi + 271 pp. 

Young, K., & B. Leon. 1988. Vegetacion de la Zona 
Alta del Parque Nacional Rio Abiseo, San Mar- 
tin.— Revista Forestal del Pert 15(1):3-20. 


(ALG) Biological Survey Field Station, 
National Ecology Research Center, U.S. Fish 
and Wildlife Service, National Museum of 
Natural History, Washington, D.C. 20560, 
U.S.A.; (MRR) Asociacion Peruana para la 
Conservacion de la Naturaleza, Parque José 
de Acosta 187, Magdalena, Lima 17, Peru. 
Present address: Department of Biology, 
University of Missouri-St. Louis, 8001 
Natural Bridge Road, St. Louis, Missouri 
63121-4499, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(4), 1993, pp. 775-785 


THREE NEW GENERA OF VERNONIEAE FROM 
SOUTH AMERICA, DASYANDANTHA, DASYANTHINA, 
AND QUECHUALIA (ASTERACEAE) 


Harold Robinson 


Abstract. —Three new genera of Vernonieae are described from South Amer- 
ica, each having Type A pollen and hairs inside the corolla. They are placed 
in two different subtribes, the arborescent genus Dasyandantha of Venezuela 
in the Piptocarphinae, the herbaceous Dasyanthina of eastern Brazil and shrub- 
by or scandent and Quechualia of the central Andes, in the Vernoniinae. Das- 
yandantha has a thyrsoid inflorescence with glomerules of sessile heads. Das- 
yanthina and Quechualia have thyrsoid inflorescences with corymbiform 
branches and have glanduliferous anther thecae, but differ in the shape and 
distribution of the hairs on their corollas. Dasyandantha cuatrecasiana is trans- 
ferred from Vernonia and Piptocarpha, Dasyanthina serrata, D. palustris, Que- 
chualia cardenasii, Q. fulta, and Q. trixioides are transferred from Vernonia, 
and Quechualia smithii is described as new. 


Progress toward systematic resolution of 
the Neotropical genera of the Vernonieae 
has nearly eliminated the excessively para- 
phyletic aspects of Vernonia Schreb. in the 
Western Hemisphere (Robinson 1980, 
1987a, 1987b, 1987c, 1988a, 1988b, 1989a, 
1989b, 1990, 1992, 1993; Robinson & Funk 
1987). The remaining elements needing re- 
moval from Vernonia represent compara- 
tively isolated small groups that sometimes 
contain notable species such as V. fulta Gri- 
seb. of western South America or V. serrata 
of Brazil. The latter two species, their closest 
relatives, and the rare V. cuatrecasasiana of 
Venezuela show distinctive pubescence on 
the inner surface of the corolla throat. The 
species with hairs inside their corollas do 
not form a single related group, but form 
three distinct groups named here as the gen- 
era Dasyandantha, Dasyanthina and Que- 
chualia. Hairs are consistently present in- 
side the corollas in Dasyandantha and 
Dasyanthina, but in Quechualia the pubes- 
cence is lacking in the corollas of the two 
rarest species. In spite of the lack of im- 
mediate relationship of all the genera and 


the partial unreliability of the corolla pu- 
bescence as a character in Quechualia, it is 
convenient to name the three genera to- 
gether at this time. With the establishment 
of the present three genera, it is believed 
that fewer than ten small Neotropical ele- 
ments remain in need of removal from Ver- 
nonia. 

The three genera described here have the 
characters of the broad, excessively para- 
phyletic, traditional concept of Vernonia, 
including alternate leaves, non-stellate hairs, 
discrete heads, epaleaceous receptacles, reg- 
ular corollas, and a pappus of many capil- 
lary bristles. One of the three genera, Das- 
yandantha, is based on a species that was 
placed in Piptocarpha R.Br. by Badillo 
(1974) on the basis of the basally tailed an- 
ther thecae, but such tails occur in all three 
genera. The tails are not the sharp, sclerified 
type seen in true Piptocarpha, but are the 
unsclerified type with denticulate bases seen 
in many species of Critoniopsis Sch.-Bip. of 
the Piptocarphinae and in Vernonanthura 
H.Rob. of the Vernoniinae that were in- 
cluded in the broad concept of Vernonia. 


776 


All three of the new genera are among 
those elements within the traditional con- 
cept of Vernonia that have Type A pollen 
grains. These grains, like those found in Ver- 
nonia sens. str. and Vernonanthura, are tri- 
colporate and spinulose with a perforated 
tectum continuous over the non-colpar ar- 
eas. The genera all have elongate raphides 
in the achene walls, which differ from the 
subquadrate raphides in Vernonia and Ver- 
nonanthura. The final character, the one by 
which the present three genera are selected 
here for description, is the presence of hairs 
inside the corolla throat, a feature not found 
in other Neotropical species falling within 
the traditional concept of Vernonia. Such 
hairs are accompanied by hairs or spicules 
on the outer surfaces of the corolla lobes 
like those of many relatives of Lepidaploa 
(Cass.) Cass. 

The three genera described here with hairs 
inside their corollas can be best placed in 
the Vernonieae as follows. Dasyandantha 
has subcoriaceous leaves with short, felty 
pubescence and densely thyrsoid inflores- 
cences with glomerules of sessile heads. The 
genus is generally similar to Critoniopsis 
Sch.-Bip. in appearance. The arborescent 
habit, deciduous inner involucral bracts, and 
complete lack of glands on the anthers are 
characteristic of the subtribe Piptocarphi- 
nae (Robinson et al. 1980), where I place 
both genera. The hairs on the stems are un- 
branched. The elongate shape of the raph- 
ides in the achene walls of Dasyandantha is 
the only character not in complete com- 
formance with the placement of the genus. 
The raphides of the Piptocarphinae are oth- 
erwise rather subquadrate or shortly rhom- 
boidal. Dasyanthina and Quechualia are 
herbs or weak to scandent shrubs with per- 
sistent inner involucral bracts and glands 
on the connectives or appendages of the an- 
thers, as in the subtribe Vernoniinae (Rob- 
inson 1992), where I place the genera. The 
two genera have hairs of the stem and some 
other organs T-shaped, one of the types 
found in the Vernoniinae but not in the Pip- 
tocarphinae. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


The stated characteristics of deciduous 
inner involucral bracts and hairs inside the 
corolla immediately distinguish Dasyan- 
dantha among the genera of the Vernonieae. 
It is the only member of the Piptocarphinae 
with hairs inside the corolla. It is also dis- 
tinct in the very long, rather lanate pubes- 
cence of the stems, the corolla lobes not 
longer than the throat, and the elongate 
raphides of the achenes. The phyletic po- 
sition is considered close to Critoniopsis, 
which is phyletically a core-genus for the 
subtribe. 

Dasyanthina and Quechualia are distin- 
guished within the subtribe Vernoniinae 
from all other genera by the hairs inside the 
corollas, but their relationship within the 
subtribe is not certain. Most genera of the 
subtribe can be placed in one of two groups. 
The group containing Vernonia and Ver- 
nonanthura has corollas glabrous, raphides 
in the achene walls strictly subquadrate, and 
pollen strictly Type A. The second group 
has been referred to as the Lepidaploa Com- 
plex (Robinson 1987a, 1987b, 1987c, 1988a, 
1988b, 1988c, 1990) and is characterized 
by often strongly cymose branches of the 
inflorescence, corolla lobes pubescent out- 
side, raphides in the achene wall often elon- 
gate, and the pollen mostly lophate. The 
new genera Dasyanthina and Quechualia re- 
semble Vernonanthura in the tails on the 
bases of the anther thecae, but they differ 
from both Vernonia and Vernonanthura in 
the hairs on either the inner or outer sur- 
faces of the corolla. They also differ in the 
elongate raphides in the achene walls. The 
two new genera differ from most members 
of the Lepidaploa Complex by their Type 
A pollen and by their corymbose rather than 
seriate-cymose inflorescences. 

The phyletic position of Dasyanthina and 
Quechualia could be considered closer to 
the Lepidaploa group in the Vernoniinae 
because of the hairs on the outside of the 
corolla lobes, the elongate raphides in the 
achene walls, and the sometimes thickened 
vein apices in the corolla lobes, but none of 
these characters is convincing. Hairs on the 


VOLUME 106, NUMBER 4 


corolla lobes occur widely in the tribe out- 
side of the subtribe Vernoniinae. Thickened 
veins at the tips of the corolla lobes seem 
to be of limited systematic value; they are 
common in most members of the Lepida- 
ploa relationship and are lacking in Ver- 
nonia and Vernonanthura. However, a lim- 
ited survey has revealed that distally 
thickened veins occur in a number of other 
genera of the tribe in both hemispheres. The 
thickened veins occur in Quechualia but not 
in Dasyanthina. Elongate raphides in the 
achene wall occur in most of the genera of 
the Lepidaploa generic alliance, but Das- 
yanthina and Quechualia do not closely re- 
semble any of those genera. The elongate 
raphides in Dasyanthina and Quechualia 
may be of separate origin in the Vernoni- 
inae, as they evidently are in Dasyandantha 
in the Piptocarphinae. At present, Dasyan- 
thina and Quechualia are regarded in the 
Vernoniinae as a separate group from either 
the Vernonia/Vernonanthura group or the 
Lepidaploa group. 

The closest relationships of Dasyanthina 
and Quechualia seem to be with each other. 
Both genera have T-shaped hairs or a more 
complex, derived form of zigzag hairs on 
vegetative parts. Both genera have more 
corymbiform inflorescences than seen in e1- 
ther the Vernonia/Vernonanthura group or 
the Lepidaploa group. Both genera have 
heads with obvious peduncles, broadly 
campanulate involucres, imbricate involu- 
cral bracts in 5—6 series, and usually 25-60 
florets. The achenes of both genera have 
numerous setulae, as well as elongate raph- 
ides. 

In spite of the similarities, differences be- 
tween the two new Vernoniinae genera lend 
a considerable degree of uncertainty as to 
their relationship. Geography is an obvious 
difference. Dasyanthina is restricted to the 
coast of southeastern Brazil, whereas Que- 
chualia ranges from northern Peru south 
through Bolivia to northern Argentina. Das- 
yanthina seems to be strictly herbaceous, 
whereas Quechualia is more shrubby and 
often cited as a vine. The inflorescence of 


777 


Dasyanthina is a short, rounded, thyrsoid 
panicle, whereas that of Quechualia is an 
elongate, pyramidal, thyrsoid panicle. The 
raphides in the achene walls are much more 
numerous in Quechualia, and the achenes 
are larger with more ascending setulae. Ac- 
tually, the hairs of the corolla, the seeming 
unifying character of the two genera, pro- 
vide important differences in their details. 
Dasyanthina has the inner surface of the 
corolla throat always densely covered with 
stiff, unicellular hairs, but in Quechualia the 
inner surface of the throat is sparsely pu- 
bescent with long, multicellular hairs in two 
of the species and is glabrous in the other 
two species. 

The detailed structure of the hairs in each 
of the three new genera is of special interest. 
The hairs mentioned are all technically uni- 
seriate. The T-shaped hairs on the stems 
and inflorescence branches of Dasyanthina 
and Quechualia (Fig. 2F) contrast strongly 
with the sublanate pubescence in Dasyan- 
dantha. The corollas of Dasyandantha have 
bi- or tri-cellular hairs on both the outside 
of the throat and lobes and the inside of the 
throat. These hairs have the short, thinner- 
walled basal cells. The corollas of Dasyan- 
thina have unicellular hairs outside of the 
lobes and inside of the throat (Fig. 1E). Que- 
chualia has unicellular hairs outside of the 
lobes (Fig. 2H), but the hairs inside the co- 
rolla throat in two of the species are mul- 
ticellular (Fig. 2G). The long multicellular 
internal hairs of Quechualia are particularly 
distinctive in their weak, thin-walled basal 
cells and the contortions in the upper parts 
near the widely spaced septations. In Q. ful- 
ta the hairs can sometimes be numerous. 
The long hairs often seem to detach, and a 
few loose hairs have been seen in prepara- 
tions of the species in which no attached 
hairs have been seen. The unicellular hairs 
on the corolla lobes and throat of Dasyan- 
thina (Fig. 1E, I) and on the corolla lobes 
of Quechualia (Fig. 2H) are structurally the 
same, and their similarity helps further sug- 
gest that relationship between the genera. 
Their bases are broadened, and they char- 


778 


acteristically have a peg that inserts into the 
thin-walled tissue of the corolla. There are 
no separate basal cells in the latter hairs. 

The three new genera are as follows. The 
pollen grains were measured in Hoyer’s So- 
lution. 


Dasyandantha H. Robinson, gen. nov. 
Type: Vernonia cuatrecasasiana Aristeg. 


Plantae arborescentes; caules sublanati; 
folia simplices alterna breviter petiolata. In- 
florescentiae dense pyramidaliter thyrso- 
ideo-paniculatae; bracteae involucri interi- 
ores deciduae; capitula homogama; corollae 
discoideae in lobis distaliter extus et in fau- 
cibus intus et extus dense pilosulae, pilis 
uniseriatis et bi- vel tri-cellularibus; thecae 
antherarum base tenuiter caudatae; pili sty- 
lorum omnino obtusi; achenia setulifera, ra- 
phidis elongatis. Grana pollinis in 
subtypo A. 

Trees to 6 m tall; stems sublanate with 
sordid hairs, with solid pith. Leaves simple, 
alternate, petioles ca. 1 cm long; blades sub- 
coriaceous, obovate, to 30 cm long, 10 cm 
wide, base narrowly cuneate, margins entire 
or with a few minute mucronate-denticu- 
lations distally, apex shortly acute, upper 
surface densely pilosulous, lower surface 
finely velutinous on veins and veinlets; ve- 
nation pinnate with ca. 10 secondary veins 
on each side, veinlets prominulous in dense 
reticulum. Inflorescence without foliose 
leaves or bracts, densely pyramidally thyr- 
soid-paniculate, bearing glomerules of ses- 
sile heads. Heads homogamous, discoid, 
broadly campanulate; involucre 0.3—0.4 cm 
wide, with bracts ca. 30, imbricate in ca. 4 
series, inner bracts deciduous; receptacles 
epaleaceous. Florets ca. 12; corollas white, 
narrowly funnelform, basal tube slender, 
throat distinct, broad, as long as lobes, lobes 
narrowly triangular, with scattered small 
glands outside, veins not thickened distally; 
outer surface of throat, lobe tips, inner sur- 
face of throat, and inner surface of lobe bas- 
es pilosulous with stiff, uniseriate hairs, hairs 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


bi- or tri-cellular with short, thinner-walled 
basal cells, with long, stiff, thicker-walled 
distal cell; anthers without glands, bases of 
thecae spurred and caudate, tails denticu- 
late, with thin-walled cells; apical anther ap- 
pendages with thin-walled cells; style with 
sclerified basal ring; shaft of styles with hairs 
distally; style branches with sweeping hairs 
all blunt-tipped. Achenes 8-ribbed, covered 
with biseriate setulae, with glandular dots 
mostly near base, raphides in walls elongate; 
carpopodium doliiform, not extending 
above constriction, glabrous, with short and 
thick-walled cells; pappus with a short, nar- 
rowly squamulose outer series, inner cap- 
illary bristles easily deciduous, broadened 
distally. Pollen grains tricolporate, spinu- 
lose, type A, ca. 40 wm in diam. 

The generic name is derived from dasys 
meaning shaggy and antha meaning flower, 
with an intervening and refering to the An- 
des. The genus contains the following single 
known species. 


Dasyandantha cuatrecasasiana (Aristeg.) 
H. Robinson, comb. nov. 


Vernonia cuatrecasasiana Aristeg., Acta 
Biol. Venez. 3:363. 1963. Piptocarpha 
cuatrecasasiana (Aristeg.) Badillo, Rev. 
Fac. Agron. (Maracay) 7(3):14. 1974. 
VENEZUELA: Edo. Lara, Falcon, Yara- 
cuy and Cojedes (Badillo, 1989). 


The species is well illustrated by Ariste- 
guieta (1963). 


Dasyanthina H. Robinson, gen. nov. 


Type: Vernonia serrata Less. 
(Fig. 1 A—H) 


Plantae herbaceae perennes; caules in pi- 
lis appresse T-formibus sparse obsiti. Folia 
simplicia alterna longe petiolata. Inflores- 
centiae rotundate thyrsoideae, ramis cor- 
ymbiformis. Capitula homogama tenuiter 
pedunculata; bracteae involucri interiores 
plerumque persistentes; flores ca. 25; corol- 
lae discoideae in faucibus intus et lobis ex- 


VOLUME 106, NUMBER 4 779 


es 
Tae 


wees 


ae 
_— 


rn 
/ 


Kes 
= 


—Ly 


Fig. 1. Dasyanthina. A-H. D. palustre (Gardner) H. Robinson. A. Habit. B. Head. C. Corolla showing anthers 
and style. D. Section of corolla showing tailed anthers, glands on connective, and short hairs on inner surface 
of corolla. E. Short hairs for inner surface of corolla throat. F. Style showing enlarged basal ring. G. Achene. H. 
Raphides from cells of achene wall. I. Hairs from inner surface of corolla throat in D. serrata (Less.) H. Robinson. 


780 


tus dense pilosulae, pilis unicellularis; the- 
cae antherarum base tenuiter caudatae in 
connectivis glanduliferae; achenia setuli- 
fera, raphidis elongatis. Grana pollinis in 
typo A. 

Perennial herbs 2—4 m tall; stems with 
very sparse pubescence of appressed, 
T-shaped hairs, with solid pith. Leaves sim- 
ple, alternate, petioles elongate, sometimes 
winged to base; blades thinly herbaceous, 
ovate, to 50 cm long, mostly 15—25 cm long 
and 4.5—16.0 cm wide, base and apex acu- 
minate, margins closely and sharply serru- 
late, surfaces sparsely puberulous with small 
often T-shaped hairs, venation pinnate with 
ca. 10 secondary veins per side. Inflores- 
cence with foliose bracts only at lower pri- 
mary nodes, rounded thyrsoid with cor- 
ymbiform cymose branches and slender 
peduncles. Heads discoid, broadly campan- 
ulate; involucre 0.5—1.0 cm wide, with bracts 
ca. 60, imbricate in 5 or 6 series, inner bracts 
mostly persistent; receptacles epaleaceous. 
Florets ca. 25; corollas white, narrowly fun- 
nelform, throat cylindrical, somewhat 
shorter than linear lobes, veins not thick- 
ened at tips of lobes, inner surface of throat 
and distal outer surface of lobes densely 
covered with stiff, unicellular hairs (Fig. 1D, 
E, I), rarely with a few multicellular hairs 
inside of throat; anther thecae with thin- 
walled tails at base, with glands on connec- 
tive (Fig. 1C, D), apical appendage without 
glands, with thin-walled cells; style with 
sclerified basal ring, upper part of shaft and 
branches with mostly pointed hairs. Achenes 
ca. 8-ribbed, ca. 2 mm long, densely covered 
with short, spreading, biseriate setulae, 
raphides elongate (Fig. 1H); carpopodium 
doliiform, with short, thick-walled cells, with 
glandular dots crowded around upper edge; 
pappus rather fragile, with outer series of 
short squamellae, inner pappus series of 
white capillary bristles, only slightly broad- 
ened distally. Pollen grains tricolporate, spi- 
nulose, type A, 37—40 um in diam. 

The name is derived from dasys meaning 
shaggy and anthina meaning little flower. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


The genus contains the following two spe- 
cies. 


Key to the Species of Dasyanthina 


1. Petioles without wings; corolla 
throat with long unicellular hairs in- 
side, lobes with shorter unicellular 
hairs restricted to outer surface ... 

eae ep ie GT D. serrata 

1. Petioles winged to base; corolla 
throat with hairs inside not longer 
than those distally on lobes, lobes 
distally with hairs on both inner and 
Outer'suriaces 0. (5st ee D. palustris 

Dasyanthina palustris (Gardner) 
H. Robinson, comb. nov. 


Vernonia palustris Gardner, London J. Bot. 
5:215. 1846. BRAZIL: Bahia, Espirito 
Santo, Minas Gerais. 


Dasyanthina serrata (Less.) 
H. Robinson, comb. nov. 


Vernonia serrata Less., Linnaea 4:275. 1829. 
BRAZIL: Rio de Janeiro, Sao Paulo. 


Quechualia H. Robinson, gen. nov. 


Type: Vernonia fulta Griseb. 
(Figs. 2A—-K, 3) 


Plantae perennes frutecentes vel scan- 
dentes; caules in pilis appresse T-formibus 
sparse obsiti. Folia simplicia alterna, pe- 
tiolis angustis. Inflorescentiae elongate pyr- 
amidaliter paniculatae, ramis corymbosis. 
Capitula homogama pedunculata; bracteae 
involucri interiores persistentes; flores 30— 
55; corollae discoideae in faucibus intus in- 
terdum sparse pilosae, pilis elongatis mul- 
ticellularibus, pilis in lobis unicellularibus; 
thecae antherarum base tenuiter caudatae 
in connectivis glanduliferae; achenia setu- 
lifera, raphidis elongatis. Grana pollinis in 
typo A. 

Perennial shrubs or vines to 5 m tall; stems 
sparsely to densely pilose with simple or 


VOLUME 106, NUMBER 4 


T-shaped hairs, with solid pith. Leaves al- 
ternate, petioles narrow, usually short; 
blades thinly herbaceous, ovate to elliptical 
or obovate, mostly 10-17 cm long, 3.5—9.0 
cm wide, margins with remote, antrorse ser- 
rulations, upper surface sparsely pilosulous, 
lower surface sparsely to densely pilose with 
simple or T-shaped hairs, with or without 
many obvious glandular dots, venation pin- 
nate, with 4—6 secondary veins on each side. 
Inflorescence usually a narrowly pyramidal, 
thyrsoid panicle with corymbosely cymose 
branches (Fig. 2A), peduncles obvious, usu- 
ally slender. Heads discoid, broadly cam- 
panulate; involucre 1—2 cm wide, with bracts 
60-90, subimbricate to imbricate in 5 or 6 
series, outer bracts appressed or spreading, 
inner bracts persistent, outer surfaces of 
bracts sometimes bearing seriately T-shaped 
hairs (Figs. 2C, 3); receptacle epaleaceous. 
Florets 30—55; corollas lavender, narrowly 
funnelform, throat shorter than linear lobes, 
veins thickened near tips of lobes, throat 
with or without sparse pilosity of long, mul- 
ticellular hairs inside (Fig. 2D, E, G), lobes 
with short, stiff, unicellular hairs outside 
(Fig. 2H); anther thecae spurred and with 
thin-walled, denticulate tails, connective 
with glands (Fig. 2E); apical appendage with 
mostly thin cell walls; style with sclerified 
ring at base, upper style shaft and branches 
with mostly pointed hairs. Achenes 8—10- 
ribbed, 3—4 mm long, surface with many 
ascending, biseriate setulae, with glandular 
dots in grooves and near base, raphides 
elongate (Fig. 2K); carpopodium long-do- 
liiform, with short, thick-walled cells; outer 
pappus with short crowded squamellae, in- 
ner pappus series of white, capillary bristles, 
only slightly broadened distally. Pollen 
grains tricolporate, spinulose, type A, 40- 
50 um in diam. 

Chromosome number of n = 17 is re- 
ported from Q. fulta (Jones 1979). 

The two most common species of the ge- 
nus have long hairs inside the corolla, but 
such hairs have not been seen in the two 
rarer species, Q. cardenasii and Q. smithii. 


781 


The latter two species, both known from 
only three collections, have all the other 
characters of the genus, including the shrub- 
by to scandent habit, the narrowly thyrsoid 
inflorescence, the long peduncles, the heads 
with involucres 1—2 cm wide, 30-55 florets 
in the head, the hairs on the outside of the 
corolla lobes, anther thecae with basal tails, 
connectives with glandular dots, and elon- 
gate raphides in the achene walls. The two 
rarer species differ in appearance from the 
common Q. fulta only in the less appressed 
and more pubescent involucral bracts and, 
in Q. smithii, in the more foliose branches 
of the inflorescence. Only Dasyanthina 
shares most of the characters of Quechualia, 
but that Brasilian genus has a more herba- 
ceous habit, a more rounded inflorescence, 
and smaller unicellular hairs consistently 
present inside the throat of the corolla. 

The generic name is derived from the 
Quechua-speaking indians that dominated 
the area from Peru to northern Argentina 
where the genus occurs. The genus contains 
the following four species. 


Key to the species of 
Quechualia 


1. Outer involucral bracts all ap- 
pressed or with stiff, spreading tips; 
peduncles less than 1 mm wide; 
throats of corollas usually with 
sparse pubescence of long hairs on 
inner surface; pollen grains ca. 40 
um in diam. 

2. Leaf blades with distinct glan- 
dular dots below; outer involu- 
cral bracts obtuse to shortly acute 

Brits ieee 1 Bay eds, 2 sch QO. fulta 

2. Leaf blades without evident 
glandular dots below; outer in- 
volucral bracts sharply acute to 
shorty acuminate .... Q. trixioides 

1. Outer involucral bracts with weakly 
to strongly recurved tips; peduncles 
ca. 1 mm wide; throats of corollas 
with few or no hairs on inner sur- 


782 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Anjali 


Ry 
Upp, 


Se - 


WSS 
NUMA py 
ANY, 7 
Wy 


yy 


We 
WW) i We, 


SA 

AW 

\ ANS 
oS \ 


aA 


— 


Fig. 2. Quechualia fulta (Griseb.) H. Robinson. A. Habit. B. Head. C. Hair from outer surface of involucral 
bract. D. Corolla showing anther tips and style. E. Section of floret showing tailed anthers, glands on connective, 
and long hairs on inner surface of corolla. F. T-shaped hair from outer surface of corolla. G. Long hairs from 
inner surface of corolla. H. Hairs from outer surface of corolla lobe. I. Style. J. Achene. K. Raphides from cells 
of achene wall. 


VOLUME 106, NUMBER 4 


783 


Fig. 3. 
two of the hairs lacking base or tip. 


face; pollen grains 45-5O um in 
diam. 
3. Headsin clusters on axillary, leaf- 
less, lateral branches of inflores- 
cence; tips of involucral bracts 
often tightly recurved, densely 
tomentellous; plants slender 
shrubs; pollen grains ca. 50 wm 
memetanMd LIC 10). Q. cardenaslii 
3. Heads few at tips of leafy branch- 
es; tips of involucral bracts only 
partially recurved, densely pi- 
lose; plants scandent; pollen 
grains ca. 45 wm in diam. ..... 
ee Aer rorpe tes Ott: Q. smithii 


Quechualia cardenasii (H. Robinson) 
H. Robinson, comb. nov. 


Vernonia cardenasii H. Robinson, Phyto- 
logia 49:262. 1981. 


The species is still known only from the 
type collection from the Cordillera Chi- 
moré, Cochabamba, Bolivia. The stems and 
involucral bracts of the species have a dis- 
tinct tomentum, but microscopically the 
hairs are a modified T-form (Fig. 3). Above 
the basal stalk, the cap cells are in a series 


Quechualia cardenasii (H. Robinson) H. Robinson. Hairs from outer surface of involucral bracts, 


mounted at or near their ends. The indi- 
vidual hairs have a zigzag shape. 


Quechualia fulta (Griseb.) 
H. Robinson, comb. nov. 


Vernonia fulta Griseb., Symb. Fl. Argent, 
Goett. Abhandl. 24:164. 1879. Vernonia 
senecionaefolia Rusby, Bull. Torrey Bot. 
Club 18:331. 1891. 


The species occurs throughout the range 
of the genus. Specimens range from Depts. 
of San Martin and Amazonas in northern 
Peru to the Prov. of Tucuman, Argentina. 
The bases of the leaf blades are usually ob- 
tuse to shortly acute, but in plants from the 
Provs. of Salta and Jujuy in northern Ar- 
gentina the bases of the blades are narrowly 
acuminate. 


Quechualia trixioides (Rusby) 
H. Robinson, comb. nov. 


Vernonia trixioides Rusby, Mem. Torrey 
Bot. Club 6:54. 1896. 


The species is resurrected from the syn- 
onymy of Q. fulta for material from near 
Mapiri in the Prov. La Paz, Bolivia and near 
Machu Picchu in the Prov. Cuzco, Peru that 


784 


lack glands on the leaf undersurfaces and 
have more pointed outer involucral bracts. 


Quechualia smithii H. Robinson, sp. nov. 


Type: BOLIVIA: Santa Cruz, Prov. Ma- 
nuel Maria Caballero, 50 km al norte de 
Mataral (en la carretera Santa Cruz—Coma- 
rapa) pasando por San Juan del Potrero y 
bajando a la cuenca del alto Rio Ichilo, 
2000-2100 m, 26 May 1989, D. N. Smith, 
G. Quintana & V. Garcia 13451 (US; iso- 
types LPB, MO). 

Plantae volubiles; folia alterna, laminis 
base anguste acutis subtus glandulo-punc- 
tatis. Inflorescentiae in ramis lateralibus fo- 
luferis terminales paucicapitatae, pedun- 
culis ca. 1 mm latis; capitula ca. 1.5 cm alta; 
bracteae involucri exteriores apice leniter 
recurvatae dense pilosae; corollae in fauci- 
bus intus non pilosae. Grana pollinis in 
diam. ca. 45 um. 

Vines; stems subglabrous, with small 
T-shaped hairs mostly near nodes. Leaves 
alternate, petioles 1.0-1.5 cm long; blades 
elliptical to somewhat obovate, 8-14 cm 
long, 2-5 cm wide, base narrowly acute, 
margins remotely sharply serrate distally, 
apex narrowly shortly acuminate, upper 
surface sparsely pilosulous, lower surface 
sparsely pilose on veins, with sparse and 
sometimes obscure glandular dots; second- 
ary veins ca. 4 on each side, strongly as- 
cending. Inflorescence terminal on leafy lat- 
eral branches, corymbiform cymes with few 
to 20 heads; peduncles 1.2—4.5 cm long, ca. 
1 mm wide. Heads ca. 1.5 cm high; invo- 
lucre green, bracts ca. 80, in ca. 6 series, 
narrowly oblong to linear, 3-12 mm long, 
1.0—2.5 mm wide, tips obtuse to short-acute, 
basal bracts densely brownish pilose, with 
tips slightly recurved, larger bracts more 
sparsely brownish pilose outside, hairs 
T-formed. Florets ca. 25 in a head; corollas 
dark magenta, ca. 13 mm long, tube ca. 5 
mm long, throat ca. 4 mm long, sparsely 
glandular-dotted outside, without evident 
hairs inside, lobes linear, ca. 5 mm long, ca. 
0.7 mm wide, distally scabrid outside with 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


many short, unicellular hairs; anther thecae 
ca. 3 mm long, with short, basal, denticulate 
tails of thin-walled cells, connective bearing 
many glands, apical appendage ca. 1 mm 
long, glabrous. Achenes submature, ca. 2.5 
mm long, densely setuliferous; pappus with 
outer series capillary, ca. 1 mm long, inner 
series fragile, capillary, 6-7 mm long, not 
or scarcely wider distally. Pollen grains ca. 
45 um in diam. 

Paratype: BOLIVIA: Cochabamba: Prov. 
Carrasco. Chua Khocha, ca. 17°48’S, 
64°43’'W, 2300 m, trepadora de flores color 
lila, 5 Sep 1990, Ramiro Renejel 12 (NY, 
US). 

The type specimen is cited from “‘Bosque 
primario, bosque pluvial montana; laderas 
con abundante Prumnopitys y pastizales 
anthropogenicos.’’ The species seems most 
distinct in the inflorescence of few heads 
born on leafy lateral branches. The scandent 
habit might also be distinctive if the differ- 
ence proves consistent. The larger heads on 
stouter peduncles and the apparent lack of 
hairs inside the corolla throat place the spe- 
cies closest to Q. cardenasii, but the latter 
is a Slender subshrub or shrub with numer- 
ous heads on lateral branches lacking leafy 
bracts. The stems, leaf undersurfaces, and 
involucral bracts of the latter are more 
densely tomentose, and the involucral bracts 
have strongly recurved tips. 

The species is named after the first listed 
collector of the type specimen, the late Da- 
vid N. Smith. 


Acknowledgments 


The illustrations were prepared by Alice 
Tangerini of the Department of Botany, Na- 
tional Museum of Natural History. John 
Pruski of the Department of Botany is 
thanked for his numerous suggestions. 


Literature Cited 


Aristeguieta, L. 1963. Tres especies de Compositae 
de Venezuela nuevas para la ciencia.—Acta 
Biologica Venezuelica 3:363-369. 

Badillo, V. M. 1974. Blumea viscosa y Piptocarpha 


VOLUME 106, NUMBER 4 


cuatrecasasiana dos nuevas combinaciones en 
Compositae.— Revista de Facultad Agronomia, 
Universidad Central (Maracay) 7(3):9-16. 
. 1989. Enumeracion de las Vernonieae (Com- 
positae) de Venezuela.—Ernstia 53:1-54. 
Jones, S. B. 1979. Chromosome numbers of Ver- 
nonieae (Compositae).— Bulletin of the Torrey 
Botanical Club 106:79-84. 

Robinson, H. 1980. Re-establishment of the genus 
Critoniopsis (Vernonieae: Asteraceae).— Phy- 
tologia 46:437-442. 

1987a. Studies of the Lepidaploa Complex 
(Vernonieae: Asteraceae). I. The genus Steno- 
cephalum Sch. Bip.— Proceedings of the Biolog- 
ical Society of Washington 100:578-583. 

1987b. Studies of the Lepidaploa Complex 
(Vernonieae: Asteraceae). I]. A new genus, 
Echinocoryne.—Proceedings of the Biological 
Society of Washington 100:584-589. 

1987c. Studies of the Lepidaploa Complex 
(Vernonieae: Asteraceae). III. Two new genera, 
Cyrtocymura and Eirmocephala.— Proceedings 
of the Biological Society of Washington 100: 
844-855. 

1988a. Studies of the Lepidaploa Complex 
(Vernonieae: Asteraceae). IV. The new genus, 
Lessingianthus.— Proceedings of the Biological 
Society of Washington 101:929-951. 

1988b. Studies of the Lepidaploa Complex 
(Vernonieae: Asteraceae). V. The new genus, 
Chrysolaena.—Proceedings of the Biological 
Society of Washington 101:952-958. 


785 


1988c. Studies of the Lepidaploa Complex 
(Vernonieae: Asteraceae). VI. A new genus, Ay- 
nia. —Proceedings of the Biological Society of 
Washington 101:959-965. 

1989a. Two new genera of Vernonieae (As- 
teraceae) from the northern Andes with dis- 
sected corolla limbs Cuatrecasasanthus and Jo- 
seanthus. — Revista de la Academia Colombiana 
de Ciencias Exactas, Fisicas y Naturales 17:207- 
213: 

1989b. Acilepidopsis, a new genus of Ver- 
nonieae from South America (Asteraceae).— 
Phytologia 67:289-292. 

1990. Studies of the Lepidaploa Complex 
(Vernonieae: Asteraceae). VII. The genus Lep- 
idaploa. — Proceedings of the Biological Society 
of Washington 103:464498. 

. 1992. Anew genus Vernonanthura (Vernoni- 
eae: Asteraceae).— Phytologia 73:65-76. 

1993. A review of the genus Critoniopsis in 
Central and South America (Vernonieae: As- 
teraceae).— Proceedings of the Biological Soci- 
ety of Washington 106:606-627. 

, & V.A. Funk. 1987. A phylogenetic analysis 
of Leiboldia, Lepidonia, and a new genus Stra- 
mentopappus (Vernonieae: Asteraceae).—Bo- 
tanische Jahrbiicher fur Sytematik, Pflanenge- 
schichte und Pflanzengeographie 108:213-228. 


Department of Botany, National Muse- 
um of Natural History, Smithsonian Insti- 
tution, Washington, D.C. 20560, U.S.A. 


PROC. BIOL. SOC. WASH. 
106(4), 1993, pp. 786-788 


INTERNATIONAL COMMISSION ON ZOOLOGICAL 
NOMENCLATURE 


% The Natural History Museum 
Cromwell Road 
London, SW7 5BD, U.K. 
Tel. 071-938 9387 
30 June 1993 


Call for nominations for new members of the International 
Commission on Zoological Nomenclature 


The following members of the Commission reach the end of their terms of service 
at the close of the XX V General Assembly of the International Union of Biological 
Sciences to be held in Paris in September 1994: Dr. F. M. Bayer (U.S.A., Corallia); 
Prof. J. O. Corliss (U.S.A., Protista); Prof. Dr. G. Hahn (Germany, Trilobita); Prof. 
Dr. O. Halvorsen (Norway, Parasitology); Dr. Ya. I. Starobogatov (Russia, Mol- 
lusca); Dr. V. A. Trjapitzin (Russia, Hymenoptera). 

The addresses and specialist fields of the present members of the Commission 
may be found in the Bulletin of Zoological Nomenclature, 50(1) (March 1993). Under 
Article 3b of the Commission’s Constitution a member whose term of service has 
ended is not eligible for immediate re-election unless the Council of the Commission 
has decided to the contrary. 

The Commission invites nominations, by any person or institution, of candidates 
for membership. Article 2b of the Constitution prescribes that: 


‘The members of the Commission shall be eminent scientists, irrespective of 
nationality, with a distinguished record in any branch of zoology who are known 
to have an interest in zoological nomenclature.’ 


(It should be noted that ‘zoology’ here includes the applied biological sciences (med- 
icine, agriculture, etc.) which use zoological names). 

Nominations made since June 1990 will be reconsidered automatically and need 
not be repeated. Additional nominations, giving the date of birth, nationality and 
qualifications (by the criteria mentioned above) of each candidate should be sent by 
1 June 1994 to: The Executive Secretary, International Commission on Zoological 
Nomenclature, % The Natural History Museum, Cromwell Road, London SW7 
SBD, U.K. 


Applications published in the Bulletin of Zoological Nomenclature 


The following Applications were published on 30 June 1993 in Vol. 50, Part 2 of 
the Bulletin of Zoological Nomenclature. Comment or advice on these Applications 
is invited for publication in the Bulletin and should be sent to the Executive Secretary, 
I.C.Z.N., % The Natural History Museum, Cromwell Road, London SW7 5BD, 
U.K. 


Case No. 

2823 Acineta Ehrenberg, [1834] and Tokophrya Butschli, 1889 (Ciliophora, Suc- 
toria): proposed conservation; Acineta tuberosa Ehrenberg, [1834] 
and Podophrya quadripartita Claparéde & Lachmann, 1859 (cur- 


VOLUME 106, NUMBER 4 787 


rently Tokophrya quadripartita): proposed conservation of the spe- 
cific names. 

Turbo politus Linnaeus, 1758 (currently Melanella polita; Mollusca, Gas- 
tropoda): proposed conservation of usage of the specific name, so 
conserving the specific name of Buccinum acicula Miller, 1774 
(currently Cecilioides acicula). 

Termes lacteus Froggatt, 1898 (currently Coptotermes lacteus; Insecta, Isop- 
tera): proposed conservation of the specific name. 

Aradus caucasicus Kolenati, 1857 (Insecta, Heteroptera): proposed replace- 
ment of syntype by a neotype, so conserving usage of the specific 
name and that of A. hieroglyphicus Sahlberg, 1878. 

Notonecta obliqua Gallén in Thunberg, 1787 (Insecta, Heteroptera): proposed 
conservation of the specific name. 

Corisa nigrolineata Fieber, 1848 (currently Sigara (Pseudovermicorixa) ni- 
grolineata; Insecta, Heteroptera): proposed conservation of the spe- 
cific name. 

2831 Corisa sexlineata Reuter, 1882 (currently Sigara (Tropocorixa) sexlineata; 
Insecta, Heteroptera): proposed conservation of the specific name. 

2777 Dytiscus biguttatus Olivier, 1795 (currently Agabus biguttatus; Insecta, Co- 
leoptera): proposed conservation of the specific name. 

2869 HEMIDACTYLIINI Hallowell, 1856 (Amphibia, Caudata): proposed con- 
servation. 

2882 Lagomeryx Roger, 1904 (Mammalia, Artiodactyla): proposed designation of 
L. ruetimeyeri Thenius, 1948 as the type species. 

2883 Procervulus Gaudry, 1877 (Mammalia, Artiodactyla): proposed designation 
of Antilope dichotoma Gervais, 1849 as the type species. 


Opinions published in the Bulletin of Zoological Nomenclature 


The following Opinions were published on 30 June 1993 in Vol. 50, Part 2 of the 
Bulletin of Zoological Nomenclature. Copies of these Opinions can be obtained free 
of charge by writing to the Executive Secretary, I.C.Z.N., % The Natural History 
Museum, Cromwell Road, London SW7 SBD, U.K. 


Opinion No. 

1717 Bucephalus Baer, 1827 and B. polymorphus Baer, 1827 (Trematoda): con- 
served in their accepted usage. 

1718 Balea Gray, 1824 (Mollusca, Gastropoda): conserved. 

1719 Xeromunda Monterosato, 1892 (Mollusca, Gastropoda): Helix candiota Pfeif- 
fer, 1849 designated as the type species. 

1720 Cycloceras M’Coy, 1844 (Mollusca, Nautiloidea): C. Jaevigatum M’Coy, 1844 
designated as the type species, and a neotype designated for C. 
laevigatum. 


788 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


BELEMNITIDAE Owen, 1838 (Mollusca, Coleoidea): ruled to be unavail- 
able; Belemnites Lamarck, 1799 and B. paxillosa Lamarck, 1801 
suppressed. 

Acrolocha Thomson, 1858 (Insecta, Coleoptera): conserved, and Coprophilus 
Latreille, 1829: Staphylinus striatulus Fabricius, 1792 designated as 
the type species. 

Carabus mollis Marsham, 1802 (currently Calathus mollis; Insecta, Cole- 
optera): specific name conserved. 

Helophorus Fabricius, 1775 (Insecta, Coleoptera): conserved as correct orig- 
inal spelling. 

Meladema Laporte, 1835 (Insecta, Coleoptera): conserved. 

Myceloporus Mannerheim, 1831 (Insecta, Coleoptera): Tachinus punctus 
Gravenhorst, 1806 designated as the type species; Jschnosoma Ste- 
phens, 1829 conserved; and Mycetoporus given precedence over 
Ischnosoma. 

Schizopus Le Conte, 1858 (Insecta, Coleoptera): placed on the Official List 
of Generic Names. 

Planoplatyscelis Kaszab, 1940 (Insecta, Coleoptera): Platyscelis margelanica 
Kraatz, 1882 designated as the type species. 

Platyscelis Latreille, 1818 (Insecta, Coleoptera): Tenebrio hypolithus Pallas, 
1781 designated as the type species, so conserving Oodescelis Mot- 
schulsky, 1845. 


Graptolithus clintonensis (currently Monograptus clintonensis; Graptoli- 
thina): specific name attributed to Hall, 1852 and a lectotype des- 
ignated. 

Monograptus crenulatus (currently Monoclimacis crenulata; Graptolithina): 
specific name attributed to Elles & Wood, 1911, and a lectotype 
designated. 

Scylliorhinus atlanticus Koefoed, 1927 (currently Apristurus atlanticus; 


Chondrichthyes, Carcharhiniformes): specific name conserved. 
Amphiuma tridactylum Cuvier, 1827 (Amphibia, Caudata): specific name 
conserved. 
Ichthyosaurus trigonus Owen, 1840 (currently Macropterygius trigonus; Rep- 
tilia, Ichthyopterygia): neotype replaced by rediscovered holotype. 
Anniella pulchra Gray, 1852 (Reptilia, Squamata): neotype designated. 
Hylobates entelloides 1. Geoffroy Saint-Hilaire, 1842 (Mammalia, Primates): 
specific name conserved. 
Palaeopropithecus ingens G. Grandidier, 1899 (Mammalia, Primates): ge- 
neric and specific names conserved. 


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per column. One ms. page = approximately 0.4 printed page. 


CONTENTS 


A new sponge species, Ceratopsion crustosum (Demospongiae: Raspailiidae), from deep waters 

of the Gulf of Mexico Belinda Alvarez and R. W. M. Van Soest 

A new species of freshwater planarian from Chile (Platyhelminthes: Tricladida), with a no- 
menclatural note on Girardia festae (Borelhi, 1898) 

Alejandro C. Curino and Néstor J. Cazzaniga 

Octopus ornatus Gould, 1852 (Cephalopoda: Octopodidae) in Australian waters: morphology, 

distribution, and life history Mark D. Norman 


Grassleia hydrothermalis, a new genus and species of Ampharetidae (Annelida: Polychaeta) 


from the hydrothermal vents off the Oregon coast (U.S.A.), at Gorda Ridge 
Vivianne Solis-Weiss 
A new species of scale-worm, Harmothoe commensalis (Polychaeta: Polynoidae), from mantle 
cavities of two Chilean clams N. Rozbaczylo and J. I. Canete 
Scalispinigera oculata Hartman, 1967 (Scalibregmatidae: Polychaeta): senior synonym of La- 
cydonia antarctica (Lacydoniidae) Hartmann-Schréder & Rosenfeldt, 1988 
Fredrik Pleijel and Kristian Fauchald 
Polynoid polychaetes associated with a whale skeleton in the bathyal Santa Catalina Basin 
Marian H. Pettibone 
Probopyrus pacificensis, a new parasite species (Isopoda: Bopyridae) of Macrobrachium tenellum 
(Smith, 1871) (Decapoda: Palaemonidae) of the Pacific coast of Mexico 
Ramiro Roman-Contreras 
Potamalpheops darwiniensis (Crustacea: Decapoda: Alpheidae), the third Indo-West Pacific 


species A. J. Bruce 
On anew genus and species of xanthid crab (Crustacea: Decapoda: Brachyura) from Chesterfield 
Island, Coral Sea Peter K. L. Ng 


A new deep-sea crab of the genus Chaceon from India (Crustacea: Decapoda: Geryonidae) 
H. C. Ghosh and Raymond B. Manning 
Cambarus (Jugicambarus) subterraneus, a new cave crayfish (Decapoda: Cambaridae) from 
northeastern Oklahoma, with a key to the troglobitic members of the subgenus Jugicambarus 
H. H. Hobbs III 
Littoral compound ascidians (Tunicata) from Sao Sebastiao, estado de Sao Paulo, Brazil 
Sergio de Almeida Rodrigues and Rosana Moreira da Rocha 
Activity and reproductive patterns of amphibians and reptiles from the Engare Ondare River 
region of central Kenya, during the dry season Laurence M. Hardy 
A petrel-like bird from the late Eocene of Louisiana: earliest record for the order Procellari- 
iformes Alan Feduccia and A. Bradley McPherson 
A new genus and species of rat from Borneo (Rodentia: Muridae) Louise H. Emmons 
A new Thomasomys (Mammalia: Rodentia) from the Peruvian Andes 
Alfred L. Gardner and Monica Romo R. 
Three new genera of Vernonieae from South America, Dasyandantha, Dasyanthina, and Que- 
chualia (Asteraceae) Harold Robinson 
International Commission on Zoological Nomenclature 
Table of Contents, Volume 106 
Index to New Taxa, Volume 106 


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17 LIBRARIES SMITHSONIAN INSTITUTION NOILALILSNI NVINOSHLINS S3IYVYSIT LIB 


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