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PROCEEDINGS

of the

Biological Society of

Washington

VOLUME 102
1989

Vol. 102(1) published 29 March 1989 Vol. 102(3) published 18 October 1989
Vol. 102(2) published 28 June 1989 Vol. 102(4) published 19 December 1989

WASHINGTON
PRINTED FOR THE SOCIETY

EDITOR

C. BRIAN ROBBINS

ASSOCIATE EDITORS
Classical Languages Invertebrates
GEORGE C. STEYSKAL STEPHEN D. CAIRNS

FRANK D. FERRARI
RAYMOND B. MANNING
Plants Vertebrates
DAvip B. LELLINGER G. DAviID JOHNSON

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 1988-1989

OFFICERS
President
KRISTIAN FAUCHALD

President-Elect
LESLIE W. KNAPP

Secretary
G. DAVID JOHNSON

Treasurer
DON E. WILSON

COUNCIL
Elected Members
GARY R. GRAVES MEREDITH L. JONES
W. RONALD HEYER RAYMOND B. MANNING

W. DUANE HOPE WAYNE N. MATHIS

TABLE OF CONTENTS

Volume 102

Alvarez, Fernando. Smalleyus tricristatus, new genus, new species, and Pseudothelphusa
parabelliana, new species (Brachyura: Pseudothelphusidae) from Los Tuxtlas, Vera-
Feng AMM) a) 4 66 no ea NCS 9 AT Sn Ae Cee Le on SNe eA ies One

Anderson, William D., Jr. and Phillip C. Heemstra. Ellerkeldia, a junior synonym of
Hypoplectrodes, with redescription of the type species of the genera (Pisces: Serranidae:
AN Tut EDR) sa ee eee aa vee eee

Barnard, J. L. Rectification of Halirages regis and H. huxleyanus (Crustacea: Amphip-
oda), from marine Antarctica, with description of a new genus, Austroregid.....................

Barnard, J. L. and James Darwin Thomas. Four species of Synopiidae from the Carib-
bean region (Crustacea: Amphipoda) 1. = ee ee eee,

Barnard, J. L. and James Darwin Thomas. A new species, Ampelisca burkei, (Crustacea,
Amphipoda) from, Plorida..22.000 2.00..) -S eee ee ene

Barr, Douglas J. and Susumu Ohtsuka. Pseudocyclops lepidotus, a new species of demersal
copepod (Calanoida: Pseudocyclopidae) from the northwestern Pacific...

Bieri, Robert. Krohnittellidae and Bathybelidae, new families in the Phylum Chaetog-
natha; the rejection of the family Tokiokaispadellidae and the generea Tokiokaispa-
della, Zahonya, and: Aberrospadella: 2a ee ee ee eee

Boeger, Walter A., Delane C. Kritsky, and Joaber Pereira, Jr. Neotrpical Monogena. 14.
Revision of Callorhynchocotyle Suriano and Incorvaia, 1982 (Hexabothriidae) with
the description of C.amatol....... See eee ee ee ee

Bowman, Thomas E. The high frequency of errors in format in manuscripts submitted
to. Proceedings of the Biological Societyiof Washinelon === ne a ae

Bowman, Thomas E. and Robert Bieri. Paraspadella anops, new species, from Sao anus
Cave, Grand Bahama Island, the second troglobitic chaetognath

Brinkhurst, Ralph O. Varichaetradrilus angustipenis (Brinkhurst and Cook, 1966), new
combination for Limnodrilus angustipenis (Oligochaeta: Tubificidae)...

Brownell, Robert L., Jr. and James G. Mead. Taxonomic status of the delphinid (Mam-
malia: Cetacea) Mursio? panope: Pinilippis SO 5) eee eae ce

Browning, M. Ralph. The type specimens of Hekstra’s OW]S 2... eect

Bullock, Robert C. Rhyssoplax baliensis, a new species of chiton from Indonesia (Mol-
Iusea:: Pobyplacophora: Chitomicac) 2.25 ee ee

Campbell, Jonathan A. A new species of colubrid snake of the genus Coniophanes from
the ‘highlands of Chiapas) Mexico 2: —) 2 a ee ee

Campbell, Jonathan A., William W. Lamar, and David M. Hillis. A new species of
diminutive Eleutherodactylus (Leptodactylidae) from Oaxaca, Mexico...

Carvacho, Alberto. Cancer johngarthi, n. sp. and Cancer porteri (Bell) (Crustacea, De-
capoda): comparisons.and hypothesis-=<2).. ee eee

Chace, Fenner A., Jr. The holotype of Heterocarpus alexandri A. Milne-Edwards (Crus-
tacea: Decapoda: :Pandalidae) (1.3520 5 Pee ee eee

Child, C. Allan. Pycnogonida of the Western Pacific Islands VI. Sericosura cochleifovea,
a new hydrothermal vent species from the Marianas back-arc Dasani eeeeeeeeees

Chirino-Galvez, Luis A. and Raymond B. Manning. A new deep-sea crab of the genus
Chaceon from Chile (Crustacea, Decapoda, Geryonidae)

Colbath, G. Kent. Revision of the family Lysaretidae, and recognition of the family
Ocnonidac Kinberg; 1865 (Eunicida: Polychacta)a == ee

Colbath, G. Kent. A revision of Arabella mutans (Chamberlin, 1919) and related species
(Polychaeta: Arabelln@ac): 20 on ae a TC

Cutler, Edward B. and Norma J. Cutler. A revision of the genus Aspidosiphon (Sipuncula:
Aspidosiphonidae). ee a ee ken nee ee e

Desbruyéres, Daniel and Lucien Laubier. Paralvinella hessleri, new species of Alvinel-
lidae (Polychaeta) from the Mariana back-arc basin hydrothermal vents...

45-49

1001-1017

701-715

362-374

375-384

331-338

973-976

264-271

1067-1068

586-589

311-312

532-534
515-519

808-819

1036-1044

491-499

613-619

84-88

732-737

401-404

116-123

283-299

826-865

761-767

Erséus, Christer. Phallodrilus hessleri, new species (Oligochaeta: Tubificidae), from abys-
Sang OHEHS Mat He sWweSstenm ur aCiNe \OCCAn soe ne Slash pe as
Erséus, Christer. Phallodrilus vescus, new species (Oligochaeta, Tubificidae) from the
STS EN a cat Le See 8 SOS PE Re ee
Erséus, Christer. Four new west Atlantic species of Tubificoides (Oligochaeta,
TEETER ES 0 CS) OS ee ee eens cc eS a de Ee ee ce ae ae
Erséus, Christer and Michael R. Milligan. Three new species of Tubificidae (Oligochaeta)
from an oil seepage area on the continental slope of the northern Gulf of Mexico.......
Etnier, David A. and James D. Williams. Etheostoma (Nothonotus) wapiti (Osteichthyes:
Percidae), a new darter from the southern bend of the Tennessee River system in
BLS TEATS NEE oy ea ne oe ee A es de Ee Re Oe Se ne ee
Fauchald, Kristian. The Second Annual Riser Lecture: Eclecticism and the study of
a UOM ES A i sd ee i a et zs
Fautin, Daphne G. and Robert R. Hessler. Marianactis bythios, a new genus and species
of actinostolid sea anemone (Coelenterata: Actiniaria) from the Mariana vents...
Fitzpack, J. F., Jr. and Craig A. Busack. Hobbseus yalobushensis, a new crawfish from
posers Mississippi (Decapoda: Cambaridae) 2.0 0s
Flint, Oliver S., Jr. and D. G. Denning. Studies of neotropical caddisflies, XL: new
species of Smicridea (Smicridea) from middle America and the West Indies (Tri-
Sanaa eo ol ETAL OPS VC IGAC) ee, nto ae sd a ea
Formas, J. Ramon. A new species of Eupsophus (Amphibia: Anura: Leptodactylidae)
ODSE QE TET Sen CB rl RE Be ee Cee ere aS ene ae eee ae Sa, SO a ee ee
Froeschner, Richard C. A small collection of Heteroptera from the Galapagos Islands,
with the description of the new species Niesthrea ashlocki and a list of Niesthrea
SPETOSY CRNGCR NG A1e) RO es a cae ee ed ee ee
Froeschner, Richard C. Three new species of Colombian lace bugs of the genera Le-
wamenya and Lepropharsa (Heteroptera: Tingidac) 22-25
Gardner, Alfred L. and G. Ken Creighton. A new generic name for Tate’s (1933) Mi-
crotarsus group of South American mouse opossums (Marsupialia: Didelphidae)..
Giere, Olav. The First Annual Riser Lecture: Meiofauna and Microbes—the interactive
relations of annelid hosts with their symbiotic bacteria eect
Gillet, Patrick. Axiothella crozetensis, a new species of maldanid polychaete from Crozet
re RMSE GRE EVE CO) Ce ANT ee see i oe St
Goeke, Gary D. Stenorhynchus yangi, a new western Atlantic species of Arrow Crab
(Crustacea, Brachyura, Majidae) and a redescription Of S. SCtiCOM IIS i.e
Grabe, Stephen A. Some aspects of the biology of Rhopalophthalmus tatersallae Pillai,
1961 (Crustacea, Mysidacea) and extension of range into the Khor Al Sabiya, Kuwait
meats i APAE Cen ITN | seen eee Mice tenn i 8 ee Tere lees 2 OS ee BON ee
Grygier, Mark J. Three new species of Myzostoma (Myzostomida) eee
Hardy, Laurence M. The karyotype of Exilliboa placata Bogert (Tropidopheidae), and
comparisons with the family Boidae (Reptilia: Serpentes)
Hensley, Dannie A. and Kunio Amaoka. A redescription of Pseudorhombus megalops,
with comments on Cephalopsetta ventrocellata (Osteichthyes: Pleuronectiformes:
ee eee IGG tees ee mH itin Pie Mi fete acl Te i ad
Hershler, Robert. Springsnails (Gastropoda: Hydrobiidae) of Owens and Amargosa Riv-
er (exclusive of Ash Meadows) drainages, Death Valley System, California-Nevada..
Heyer, W. Ronald and Alan J. Wolf. Physalaemus crombiei (Amphibia: Leptodactyli-
dae), a new frog species from Espirito Santo, Brazil with comments on the P. signifer

Hobbs, Horton H., Jr. and Daniel J. Peters. New records of entocytherid ostracods
infesting burrowing crayfishes, with the description of a new species, Ascetocythere
BEE LE Se ne ree oad Ree ee OR ee

Hobbs, Horton H., Jr. and Henry W. Robison. On the crayfish genus Fallicambarus
(Decapoda: Cambaridae) in Arkansas, with notes on the fodiens complex and descrip-
HISICHALIV CT HEIVES PECIC Smale. 2 divert ihe pth Th fee

Holsinger, John R. Allocrangonyctidae and Pseudocrangonyctidae, two new families of

131-133

134-136

877-886

887-893

987-1000

742-752

815-825

637-643

418-433

568-576

609-612

968-972

3-7

109-115

866-869

620-636

726-731
793-804

1045-1049

577-585

176-248

500-506

324-330

651-697

holarctic subterranean amphipod crustaceans (Gammaridea), with comments on their
pliylopenetic and zoogeographic-relationships:.<... es ee es
Holt, Perry C. A new species of the cambarincolid genus Sathodrilus from Missouri,
with the proposal of a replacement name for Adenodrilus Holt, 1977 (Clitellata:
Branchiobdellida) 0 2-2) FS NE eB RE ee ee ee a A RON eee
Humes, Arthur G. Acontiophorus excavatus, new species (Copepoda: Siphonostoma-
toida) associated with the soft coral Dendronephthya (Alcyonacea) in the Indo-
PACHIG son OIE hee EST RPE OE eat eee ee ROAR Oe OU nN) Oa ee
Jara, Carlos G. Aegla denticulata lacustris, new subspecies, from Lake Rupanco, Chile
(Crustacea: ‘Decapoda: Anomira Acelidae) es eee 2 ee eee ee
Johnson, G. David and Edward B. Brothers. Acanthemblemaria paula, a new diminutive
chaenopsid (Pisces: Blennioidei) from Belize, with comments on life history...
Kensley, Brian. New genera in the thalassinidean families Calocarididae and Axiidae...
Keppner, Edwin J. Four new species of free-living marine nematodes in the genus
Pareurystomina (Nematoda: Enoplida) with observations on other members of the

Kornicker, Louis S. The adult male of the troglobitic ostracode Spelaeoecia bermudensis
Angel and Iliffe, 1987, from an anchialine cave in Bermuda (Crustacea: Ostracoda:
Halocypridoidea) 2.) Jo3 4) 2, As SE Sa ee ene ee ene ee ee

Kornicker, Louis and Thomas M. Iliffe. Dantya ferox, a new species of mydocopid
ostracode from Niue, central South Pacific (Crustacea: Ostracoda: Sarsiellidae).......

Lowry, James K. and Gary C. B. Poore. First ingolfiellids from the southwest Pacific
(Crustacea: Amphipoda) with a discussion of their Systematics... eeeeeeeecccsneeeeeeeeeeeneeeee

Manning, Raymond B. Sanquerus, a replacement name for Posidon Herklots, 1851
(Crustacea, 'Decapoda-Portumidae) sot cn tle eee ee

Manning, Raymond B. and David K. Camp. Additional records for an Atlantic reef
lobster, Enoplometopus antillensis Liitken, 1865 (Crustacea, Decapoda,
Enoplometopidae) 2.4.22 4 22 See ee eee OE 8 tence O38 es ee ee

Manning, Raymond B. and L. B. Holthuis. Two new genera and nine new species of
geryonid crabs (Crustacea; Decapoda,'Geryoniddae) eee eee

Manning, Raymond B., Marcos Siqueira Taveres, and Elaine Figueiredo Albuquerque.
Chaceon ramosae, a new deep-water crab from Brazil (Crustacea: Decapoda:

CHEF VOL AG) ote Osan mee ePaper Se he Sac leah ene ghee iA aS

Martin, Joel W. and Denton Belk. Eulimnadia ovilunata and E. ovisimilis, new species
of clam shrimps (Crustacea, Branchiopoda, Spinicaudata) from South America...

Mathis, Wayne N. A review of the beach flies of the Caribbean and Gulf of Mexico
(Diptera: Canacidae) 2020 ee ee SF Sioa ee ae Ok, ROO Ee Ae

Mathis, Wayne N. and Jin Zuyin. A review of the shore-fly genus Polytrichophora
Cresson: fromeAsia (Diptera: Ephydridae) = eee ee ee ee

McCranie, James R., Jay M. Savage, and Larry David Wilson. Description of two new
species of the Eleutherodactylus milesi group (Amphibia: Anura: Leptodactylidae)
from: northern: Honduras = 20 Sees SU Se ea eet oy eee ne Se nee eee

McLelland, Jerry A. Eukrohnia calliops, a new species of Chaetognatha from the northern
Gulf of Mexico‘withmiotes on related: specie sees ee eee

Miller, John E. and David L. Pawson. Hansenothuria benti, new genus, new species
(Echinodermata: Holothuroidea) from the troptical western Atlantic: a bathyal, epi-
benthic holothurian’ with swimming) abilitics 2 = eee eee

Miller, Scott E. Paleontological type specimens in the Santa Barbara Museum of Natural

Monniot, Claude and Francoise Monniot. Ascidians collected around the Galapagos
Islands using the Johnson-Sea-Link research submersible... cece
Munroe, Thomas A. and Madhu N. Mahadeva. Symphurus callopterus (Cynoglossidae,
Pleuronectiformes), a new deepwater tonguefish from the eastern Pacific.
Ohwada, Takashi. Redescription of Nephtys squamosa Ehlers (Polychaeta:
Nee polity ihe) es ee koe Th hea eel ad Ns Si ene le
Olson, Storrs L. Two overlooked holotypes of the Hawaiian flycatcher Chasiempis
described by Leonhard Stemneger (Aves: Miyacuticac) ree eet ee

947-959

738-741

916-923

385-393

1018-1030

960-967

249-263

313-323

901-911

933-946

698-700

411-417

50-77

646-650

894-900

590-608

434-446

483-490

33-44

977-986

1-2

14-32

458-467

124-130

555-558

Ortiz, Juan Carlos, Héctor Ibarra- Vidal, and J. Ramon Formas. A new species of Eupso-
phus (Anura: Leptodactylidae) from Contulmo, Nahuelbuta Range, southern Chile...
Ota, Hidetoshi and Ronald I. Crombie. A new lizard of the genus Lepidodactylus (Rep-
tilia: Gekkonidae) from Batan Island, Philippines.
Perkins, Philip D. Adelphydraena, new genus, and two new species from Venezuela, and
remarks on phylogenetic relationships within the subtribe Hydraenina (Coleoptera:
SCE 21e) aa eerie ce aio) ev eae 8 ee Re OL PS 2
Pettibone, Marian H. New species of scale-worms (Polychaeta: Polynoidae) from the
hydrothermal rift-area of the Mariana back-arc basin in the western central Pacific...
Pettibone, Marian H. Polynoidae and Sigalionidae (Polychaeta) from the Guaymas
Basin, with descriptions of two new species, and additional records from hydrothermal
vents of the Galapagos Rift, 21°N, and seep-sites in the Gulf of Mexico (Florida and
eS EEE Sees EEE oes ee eee ee mMRRNT SS | ns Sr ene Oe vee
Pettibone, Marian H. A new species of Benhamipolynoe (Polychaeta: Polynoidae: Lep-
idastheniinae) from Australia, associated with the unattached stylasterid coral Co-
RUPIRIRPERPEIIOL Besos Secon se es 2 atic Pk 2h
Pettibone, Marian H. Two new species of Harmothoinae (Polychaeta: Polynoidae) from
the East Pacific Rise, collected by Alvin dives 2000 and 2003 nee
Rickart, Eric A., Lawrence R. Heaney, and Mark J. Rosenfeld. Chromosomes of ten
species of Philippine fruit bats (Chiroptera: Pteropodidae) oe eeeeeeeceeeeeeeeeeeeeeeee
Rodriguez, Gilberto and Horton H. Hobbs, Jr. Freshwater crabs associated with caves
in southern Mexico and Belize, with descriptions of three new species (Crustacea:
ae, Sennen mee Ee ee es rr ee ee ee
Romero, Raul Castro and Hernan Baeza Kuroki. Neobrachiella anisotremi (Copepoda:
Lernaeopodidae), a new species parasitic on an inshore fish, Anisotremus scapularis,
PEEVE CEE SOR OY a a ee ear Mn ee
Romero, Raul Castro and Hernan Baeza K. Lamelliform structures on the proboscis of
Peniculus and Metapeniculus (Copepoda: Penmellidae) cece eeeeeeeceeeneeeeeeeeeene
Roper, Clyde F. E. and C. C. Lu. Systematic status of Lepidoteuthis, Pholidoteuthis, and
Neranvenoieuins (Cephalopoda: Ocgopsida)...
Rossman, Douglas A., Ernest A. Liner, Carlos H. Trevino, and Alan H. Chaney. Re-
description of the garter snake Thamnophis exsul Rossman, 1969 (Serpentes:
IPE ASS a ee a Fe
Ruff, R. Eugene and Betsy Brown. A new species of Euchone (Polychaeta: Sabellidae)
from the northwest Atlantic with comments on ontogenetic variability.
Russell, David E. A new species of Odontosyllis (Polychaeta: Syllidae) from Twin Cays,

San Martin, Guillermo and Carmen Alos. Paraprocerastea crocantinae, a new genus
and species (Polychaeta: Syllidae: Autolytinae) from the Spanish Mediterranean...
Schotte, Marilyn. Two new species of wood-boring Limnoria (Crustacea: Isopoda) from
Rene ane ARIE eP ICICSE ATG E- RCPILCULIES se
Sheavly, Seba B. and Harold G. Marshall. Plytoplankton composition in a borrow pit
EE CE OSIRIS ere a
Soils-Weiss, Vivianne and Kristian Fauchald. Orbiniidae (Annelida: Polychaeta) from
mangrove root-mats in Belize, with a revision of protoariciin genera
Stauffer, J. R., Jr. and J. M. Boltz. Description of a rock-dwelling cichlid (Teleostei:
Mieaneac)ironirake Malawi Alricn |... te. Noe
Steadman, David W. New species and records of birds (Aves: Megapodiidae, Colum-
bidae) from an archaeological site on Lifuka, Tongo.
Storer, Robert W. Geographic variation in the Yellow-rumped Tanager (Aves: Thrau-

Thomas, James Darwin and J. L. Barnard. Gammaropsis arawakia, a new species of
macine Atmphipoda (Crustacea) from Jamaica
Vari, Richard P. and Ann Williams Vari. Systematics of the Steindachnerina hypostoma
complex (Pieces, Ostariophysi, Curimatidae), with the description of three new
SEE ES te ae eee
Wagele, Johann Wolfgang and Niel L Bruce. Natatolana pastorei (Giambiagi, 1925)

1031-1035

559-567

447-457

137-153

154-168

300-304

305-310

520-531

394-400

106-108

912-915

805-807

507-514

753-760

768-771

870-876

716-725

272-279

772-792

8-13

537-552

553-554

89-94

468-482

(Crustacea, Isopoda, Cirolanidae) from the Straits of Magellan, South America: Re-
description and notes on functional morphology ee
Warén, Anders and Robert Moolenbeek. A new eulimid gastropod, Trochostilifer eu-
cidaricola, parasitic on the pencil urchin Eucidaris tribuloides from the southern
Caribbean 2-45 aden ewer I perett watunensomthcpent fn bs orte wee Bomets Bh EE
Wicksten, Mary K. Synalpheus arostris and Philocheras lapillus, two new species of
caridean shrimp (Crustacea) from the tropical eastern Pacific ee
Wicksten, Mary K. Hippolyte zostericola (Crustacea: Decapoda) in the eastern Pacific.
Williams, Austin B. and P. J. B. Scott. Upogebia corallifora, a new species of coral-
boring shrimp from the West Indies (Decapoda: Upogebiidae)
Wilson, George D. F., Oleg G. Kussakin, and Galina S. Vasina. A revision of the genus
Microprotus Richardson with descriptions of two new species, M. acutispinatus and
M. lobispinatus.(Asellota, Isopoda,. Crustacea) 2s. 2.8 ac eo Bi eS
Zullo, Victor A. and Anna V. Buising. An unusual species of the Balanus amphitrite
Darwin complex (Cirripedia, Balanidae) from the ancestral Colorado River delta in
western Arizona .and ‘southeastern California: 2c. 28 2 aioe ee

95-105

169-175

78-83

644-645

405-410

339-361

924-932

INDEX TO NEW TAXA

Volume 102

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

Si OE SE IS tee tN ga a Ite Pd Bt Fl a Co ee Se
EST IS te se ae a A i IRA RAS Ew REE eS

PLATYHELMINTHES

Trematoda

ea ES TSO SD GON GLC ate ca AS ROPE EAE RE IES SERA ADSI A TH EEN AS RE RA
En DE DOTS G1) TSC a scat Ra a a ee re de Sa a

NEMATODA

ies Oe aM SE CHER DEEN INCE) PIT ICE eee etme Nt ee
PAPEETE EDEL (Se ee re cl PT a orcas Met eee Ree ee
SDT EI TITAS eat el i DAR MR eS NE EEe A eAO ALE SoA Ra A Al Se
LIES SY DE, sac cncenpet ntl det ee et tS NO ee Ret

MOLLUSCA
Polyplacophora

MUIR CREME IIIT COTES 1 Simmer ae Sf a ad ae en eee tee a

2 TEED OTE SES: PUPIL OSS id a ceed eae eh al oR Sn
EET EET EGIL SOO, OG, SS RTS SO NNER Tee GN UDP er ae PMMOM TON TTL LEOE EN We? 2 /
CREE SCALES Scere oe CB De Ree ea IR ch ne Ee eS
ELE YDS I SS os I STS RIT AS Sena til a ele a RE a es ET Os EOE

CETTE LEIS, sata i ct DR NR td Me IE eA CS Atel a LA
FEO GE ST sen ep Bi I a TU at A i le 8 AE ea ce 7A der ant Bes AP Sot Ra dla
DUEL SIC ee oa PT SM RE is ED ER UE AR aN BL SCE OE eT ade eR at

A SEVERE OT Ca AO) a cost a ce a tS I Oe es RE A TA

ee ES ERE T PUTED Bsr oe eh ta Ne ah PS RA Se RE A EE
ETL TT ee eee ee ete en ter ee oe
PONG SOE a nane PSs P Rs i PE Ik e e l A SR e RE

ANNELIDA

Polychaeta

DNTPS SUL EL POTATO se te pr EST Ee ees enn SOC
UCTRE REDS oS OES IS eS eee ee
EMR OE CENE EER EL ACE CNIS Ii err nes ee ees

Beshamipotynoe carpsi 2 * 2 A 2a ee ee ee ee 301

Branchmmolopiutmia DUrKenses ee 144
EE VACENCATRS PRIA SR a a ne ee nc 753
EXURrieee IVETE a a Ree et ee 308
E¥arinrothine WHat ee ee a ey EO ee cee 305
ELEEPTDAST FOEIIN VAP aa a a e 2 300
Lepidonotopockivarm: ryanritetunreg os esa A ee eh te ee me ee 5 142
Pevemsteinielica aise. a a ee ee 139
Macellicephaloides a@ivirer 13 og ee eee ee a Se 162
Qdots Gls LW Pricey CHS IS a a ec 768
COPISTHO UO CHOPOCUS FTTCTIES a re 149
Paralvinella Resslerd 2 eo Ea NE aI ate erro ene oe SD ent cen 761
Paraprocerdstea oe eee ee eee er 872
CFOCGNUIN GG. a ee Ea Tice a ee ne ef ee 874
Bararteia: OCIEZCVISIS ne a ee 789
Petfibonelia on a ee ee 785
VVC TUUTACT ILL oa oh 785
RGEOAUICIA PECTIICIIE AEG: cc Fg 783
Oligochaeta
Tmnodriloides OLCGritis 2 he De no 887
hve raha: eS SE CHU an se es 131
VESCUS 2 aS Ss EEN Dee a 134
Tubificoides
6) gL | | | eam een ue een Merete Ere a ee ne 882
OCPINIUS eee 883
BOLO ss yh 2M re 880
PO ATRLTIICTASIS aa FN a as a 878
PORCH UMTS a Ses ee ee 889
DDCOUCENO AC a on! NR NI oe I a 891
Branchiobdellida
Sea EERO ETN US 1B OKO PULOV EUS a I 738
ARTHROPODA
Pycnogonida
Odontothelphusa wiOnOdOntis xh ee eee ee ee 396
Potamocarcinusleptoriel ies a a ee eee 394
SeriGostina COCHICHOVCG sc. te as a eo 732
Pyphlopseudothelphusa (ya ar 398
Crustacea
AACONTIOPHOFS CXCOVOTLUS isso BO ED BR ei aa ET a Sn 916
A@eia demticurlatanLaCi stra ooh te Se A eS ee eee 385
AL ELOGRANGONY CT LDAE be lana Dato Bl aE Os SF 9 ee 947
Ampelisca burkei. o2 3 I ie oe ee 375
Aseelocy there Stocker hi: 8 BE De 325
PAUSES OV COV 5 ee ot ls nD 703
BSetb ASUS COTO US on See SE op lk Pt 926
GCOISTOCOFIS a Fe EO See hy a oo ee ed 961
Caner JOMMGOTEN xe 2 I ee 613
CCC ON i ie Te ce Lae a Re 51
CAL OPUS as Nhe NS Sr an LN RDN AO EE 53
DS 6) (0) ee oe ee ae ee eee eT ae eee Pee 55
CVAPN ASB Si i Ss IS ac a SEE 401
OCU DSW CU a oh lh te es Se a7
CUM OT A Oo a i tage aH Ta ea eo 61

OORT EET NEE Me ee aaa ae 66

EARS ied Ne ace kB ee SR ee oc ae eee ee 69
POP DN Gee SS DE 2s TN De 646
REGIE DING TG oS NI ae Ee NO ee et 70
we SEE (ERODE Se See TD oi i et 902
CEOS SOPECE EGE DOTTIE lS I De BIE ek ee 894
POOR Sa a ah OE ed a 898
Pat STATA TTI (CO GC ASCUETINES 77) 90 100 a 684
Emaecimnards (balliGamibanuS)) DCHINCONDUS = 661
SOO ART ARTS PAORESSESSECET PR RE CD AUD cecil an ra A i E 89
ODA IE TS SERDS LOVES eS aCe Ss Se IE PT Se eo ee NNN PT ALND ee Te 370
ORES MOE Ps COEME SIT CHEN Sete ae wae ae es ee Pe 637
LL EEE, DOU R ITT Sa Sa SE el li SM PO Se eee a 933
ee EEE 7 em cn nS 2D 938
paid aM ek Sele E ONL GA ETC ECD) nf CGS Ii a ee) FLT
PEE IUD AS tush Bee te AR a ANTE ED SENST ST g IRE OENS AOU ORO PERN PEI PETS 720
NEP IE IEE TEs Pan ae reas Nok ak a ee eo SN ae hk ites 2 962
DMEDIES FAT Dies SS UN) SEI i pe ne POR eh ONT WSN AOS eae Be IE Le or NE REI set UO 963
WAP UPMRCRPMERE ASIN S PRCT YC TIEUZCL LIS ene eh a Sok kd a Be eed thn 354
SUS DEAT OL Sn oa 2 aa Sa MR geet a ee ee eh GS ge ht SE ane eee 349
SET PPS CPS LS COTO OG (y i ct 2 F ea aatot Se RG aOR ct EAE Aa eae A a D 106
SSRmOR TAIT TPITE CS ns Ra toa ls YB De 80
PELE DOTA ce sce Oe ES I ae a 2 A ae eee We Re A Seas eel Seti le 964
ene Ore G Oe TD Ab ees oer eee eee eee 953
PSEC SUES VERT ONO No een ace et eaten a a che ea lee eae oe 0 el Merge nee ear se A oe ane 331
EPEUZIE LEE SVS) GUAT OG TOT sal ee ES eR A Pi cite 46
oo DAE TDG EIS. cent ais ss Ne ae eNO APR Ce a REE SE PSE DETR PET SSRI) AG LM ta SoA 964
ner aneaTie WATT cet ener ete ae ee ere ee ee ee eee eee 698
EE et eR Ss Een SB oa A Ee, eee le ie a. 45
SFLU TILT a re a ot Ot i a lg ee CL eee RDO eRe RA 46
2 EDR ES Tet ETS Te ag a ee Sc Rn 631
LTE SLD ETS GOR ee al oie a OSE Oe ee Oe reek Cee eee, 5 78
a CARTE ED 1) 0) mee ae ee enenne Tr en enn se rte ee ee eee 405
EEN BOVE ac cess a eg ah AN en see a EES AS, a A 2
INLET a Sa a elec Ee Se 73
Insecta
ALP BIL SURE GD) ast So EG yd 5 ns Ne ea a rr 448
Di LG ET Se et a SRE Ws te Se A le ts 450
SORIA ee TS EN Beso UES ae a elt Le ERR oe) 448
RI GI SARI TIAGGICS Eanes ek Beek Oe eee ee Re WA Soho 968
UBD RESS RAPES CTP TICIOIITS e RRN So! Se aig SSCS I EB Re nr re 969
TEES OT! soso SE IE 3 I ES cll NS See I EE 970
“TESS EPSR) EQ DGl Te NA Se ee Sa DS eee Tee oe te a Ne ee Are ELT Pee PU ea 609
Co DEE GSES LOE TTT LE Ss a a a aa nS 599
Ae 8 ey OER RSN yaa sh Sahl ala, AIR 598
TEE SRYESE OS TENCE LEAD CUI 0 a2 Aaa eo 419
CESSES GI Fac Sanat ess dln Micah pees As ll cil te A AO 421
POD Apr wens alan TS cays ba toe les sea estate cel a Me Nimes A a REL a 423
EPP T BOSS Sn I Sc ET Sa I SP 421
SOP TINT 6 a Rs a 429
EAE Tt 2 eee ee ene emnn ane nn ent ne erie fy ee er et 423
APIO ea cee an te Sue ea ea ee A a eS A 426
ELLE TEN ETT a ent gt eg a LL LOE EEA ATT LIE AE 431
ed gn PI PP Baca cs TRC Eh 43]
CRETE sa cau eels Ta Mesh ge SON ae a eel ree . 426
COLES TN a yal SUT Re OS OL) eA ee BS 423

CELE POE IF) oe a8 pe on ne EO a eR ee 419

CHAETOGNATHA

RATELY BE DA es hs Ee ee eT hE eee Seen ee te ee ee ee eee
JESUS oN ga ee ator? 11110) 2) eae Od OO RRR tld MOR gO Ce ROA! cs aan bea Pact adie sac Wire vn tae adode lh,
KRONE ERE TE PAT 5 oe i hd Rs Ie eee eae pie ee RIE ONE ee Set ae LT
Paraspadellainops.2 ee ee Se ee ee

ECHINODERMATA

Holothuroidea

TUNICATA

Ascidiacea
PSC1G NA FAS Cg ha et rm nln dn ere A
OVEN OITA OVA ain se ct as een
Peco yeh ir ts OF IAS ONT i eB
SSL EM G1.) [1] {| a ac ara enone ee mnsee onese stot ene eves ree 0) See eee
Styler SOLfOT IS ccc ce a hh Pn Nee

CHORDATA

Pisces

Acanthemiblemaria paul 2.50253 ey, ey he he I Bll er ln eee
EGE OSUOMNA WPI scsi 8 ha os es es i eo es re ge rose Seek
Pseudotropheus XnstOnta Chis snc. se a ee on verso she pee aa
Stcindachnherima o7acilis. 22-54.) oh oa. bank ile ee pa ee ee tren te on ee

DUGTUIVONUTAS 25s oo sn pe ey SE ates lah SA ee W Ret ae eee

GUGSUIIOU ON 2.58 Sn ee Es Pi set OE Nall tlhe sa ind ae
Syanplrs: CaM Ope eras saa spe 0 Sem Se a a ig eer decane ne

DOU IUUIAU Sea i a Be gd nee eH
EIPSOPHUS COMI OCTISIS 2 an 5 ape eee aC a
CHNIMOPUGIT ia ee gc TE ee
Pi ySala@unvuns: CROWD CL ceo 0 sla me ae One

Cm Oplanies aI V ee 0 ats i Nr ae age oc
Wepidodacty lus: DaQliOD isis tao. A es oo inh settled Se a cla

Megapodius alimentum

Gracilinanus

TRUOA OB OV ON 5 5 5 8 ol pf AD LA a ce
carriebowensis
ramosissima

ERRATA

Hobbs, H. H., & H. W. Robison. 1989. 102(3):651-697. For Fig. 2 (p. 657) and Fig. 7 (p. 673), the same
maps were inadvertently printed. The correct map and caption for Fig. 2 is shown below.

Fig. 2. Distribution of Fallicambarus (F.) dissitus (encircled stars), F. (F.) jeanae (encircled dots), F. (F.)
petilicarpus (triangle) and F. (F.) strawni (dots) in Arkansas. (Some localities listed in text too close to others to
be shown.)

g
4. ww Ye

THE BIOLOGICAL SOCIETY OF WASHINGTON

1988-1989
Officers
President: Kristian Fauchald Secretary: G. David Johnson
President-elect: Leslie W. Knapp : Treasurer: Don E. Wilson
Elected Council

Gary R. Graves Meredith L. Jones

W. Ronald Heyer Raymond B. Manning

W. Duane Hope Wayne N. Mathis

Custodian of Publications: David L. Pawson

PROCEEDINGS

Editor: C. Brian Robbins

Associate Editors

Classical Languages: George C. Steyskal Invertebrates: Stephen D. Cairns
Frank D. Ferrari
Plants: David B. Lellinger Raymond B. Manning

Insects: Wayne N. Mathis
Vertebrates: G. David Johnson

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Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biological
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PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 1-2

PALEONTOLOGICAL TYPE SPECIMENS IN THE
SANTA BARBARA MUSEUM OF NATURAL HISTORY

Scott E. Miller

Abstract.—In accordance with Recommendation 72G of the International
Code of Zoological Nomenclature, which recommends that institutions publish
lists of the type specimens in their collections, this paper lists primary paleon-
tological type specimens in the Santa Barbara Museum of Natural History,
Santa Barbara, California (SBMNH). This list does not include paratypes and
plesiotypes, which include those of Bremner (1933), Chaney and Mason (1933),

and Orr (1968).

Kingdom Plantae
Family Rosaceae

Pyrus hoffmannii Chaney and Mason,
1933:61, pl. 7, figs. 9, 10, 13; syntypes flow-
er, fruit, and leaf, SBMNH Pal. 277-279
(SBMNH Pal. Bot. Types 26-28); Late
Pleistocene, “Carpinteria Formation’’; Car-
pinteria Asphalt Deposit, Santa Barbara
County, California. Synonymized under
Amelanchier pallida Greene by Miller
(1979).

Kingdom Animalia
Class Aves
Family Pseudodontornithidae

Osteodontornis orri Howard, 1957:3, figs.
2-8; holotype nearly complete skeleton in
2 shale slabs, SBMNH Pal. 309; Miocene,
Monterey Formation; Flagstone quarry of
G. Antolini and Sons, west side of Tepus-
quet Creek, Santa Barbara County, Califor-
nia.

Family Phalacrocoracidae

Phalacrocorax rogersi Howard, 1932:118,
fig. 19; holotype coracoid, SBMNH Pal. 131
(SBMNH Pal. Orn. 32.1); Early Pleistocene,
Santa Barbara Formation; Veronica Springs

Stone Quarry (depth 50 feet), Santa Bar-
bara, Santa Barbara County, California.

Class Mammalia
Family Otariidae

Imagotaria downsi Mitchell, 1968:1844,
figs. 1-14; holotype partial skeleton,
SBMNH Pal. 342; Early Pliocene or Late
Miocene, Sisquoc Formation; Quarry of
Great Lakes Carbon Company, near Lom-
poc, Santa Barbara County, California.

Literature Cited

Bremner, C.S.J. 1933. Geology of San Miguel Island,
Santa Barbara County, California.— Santa Bar-
bara Museum of Natural History Occasional Pa-
per 2:1-23.

Chaney, R. W., & H. L. Mason. 1933. A Pleistocene
flora from the asphalt deposits at Carpinteria,
California.— Carnegie Institute of Washington
Publication 415:45-79.

Howard, H. 1932. A new species of cormorant from

Pliocene deposits near Santa Barbara, Califor-

nia.—Condor 34:118-120.

. 1957. A gigantic “toothed” marine bird from

the Miocene of California.—Santa Barbara Mu-

seum of Natural History, Department of Ge-

ology Bulletin 1:1-23.

Miller, S. E. 1979. Reevaluation of Pyrus hoffmannii
(Rosaceae) from the Pleistocene Carpinteria As-
phalt Deposit, California.—Madrono 26:190-
191.

Mitchell, E. D., Jr. 1968. The Mio-Pliocene pinniped
Imagotaria.—Journal of the Fisheries Research
Board of Canada 25:1843-1900.

Orr, P. C. 1968. Prehistory of Santa Rosa Island.—
Santa Barbara Museum of Natural History, San-
ta Barbara, California. xxi + 253 pp.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Santa Barbara Museum of Natural His-
tory, 2559 Puesta Del Sol Road, Santa Bar-
bara, California 93105; Present address:
Bishop Museum, Box 19000-A, Honolulu,
Hawaii 96817.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 3-7

A NEW GENERIC NAME FOR TATE’S (1933)
MICROTARSUS GROUP OF SOUTH AMERICAN
MOUSE OPOSSUMS
(MARSUPIALIA: DIDELPHIDAE)

Alfred L. Gardner and G. Ken Creighton

Abstract. — Until recently, all mouse opossums were considered as belonging
to the genus Marmosa, as revised by Tate (1933). Today, however, we recognize
that Marmosa (s.1.) contains several distinctive species groups whose relation-
ships are best expressed by dividing this complex into five genera as follows:
Marmosa (s.s.), Marmosops, Micoureus, Thylamys, and a complex of species
for which we propose a new name Gracilinanus because no genus-group name

is available.

Systematists have differed in their inter-
pretation of the generic and subgeneric tax-
onomy of the smaller mouse-sized marsu-
pials of the family Didelphidae. Thomas
(1888) grouped the known taxa of mouse
opossums under the subgenus Micoureus in
the genus Didelphys. Matschie (1916) dis-
tributed these taxa among five subgenera of
Didelphis: Marmosa, Grymaeomys, Mar-
mosops, Thylamys, and Caluromys. Ca-
brera (1919) grouped mouse opossums into
two subgenera (Varmosa and Thylamys)
under the genus Marmosa. Tate’s (1933) re-
vision was the most comprehensive. Tate
(1933:22) believed that the genus Marmosa
was a “““good,’ natural genus of didelphids”’
that included all Neotropical mouse opos-
sums. Although Tate acknowledged natural
subunits in his concept of Marmosa, he did
not recognize subgenera and, instead, used
the following names to indicate five infor-
mal groups: cinerea, murina, noctivaga, mi-
crotarsus, and elegans. He also expressed
the opinion (p. 22) that “‘there is always the
chance that subgenera may be later con-
verted into full genera, and thus, in the case
of Marmosa the undoubted unity of the ge-
nus be obscured.”

Over the ensuing half-century, however,
new information has made the division of

Marmosa (sensu Tate 1933) into several
genera a desirable and necessary step to bet-
ter reflect relationships and natural assem-
blages. Gilmore (1941) again divided Mar-
mosa into the subgenera Marmosa and
Thylamys but did not list species. Cabrera
(1958) used Marmosa and Thylamys as sub-
genera as he had earlier (Cabrera 1919) and
followed the species associations of Tate
(1933) except that he divided Tate’s micro-
tarsus group, assigning the microtarsus sec-
tion to Thylamys and the /epida section to
Marmosa. Marshall (1981) listed the names
Marmosa (“murine opossums’’), Micoureus
(‘large murine opossums’’), and Thylamys
(‘small murine opossums’’), as genera
among other didelphids and attributed this
arrangement to a manuscript by Reig et al.
subsequently published in 1985. Reig et al.
(1985) assigned Tate’s (1933) murina and
noctivaga groups to the genus Marmosa
along with the more recently described taxa
M. andersoni, M. cracens (incorrectly spelled
crascens), and M. xerophila. They allocated
Tate’s (1933) cinerea group to Micoureus.
They also assigned the 14 species listed by
Kirsch & Calaby (1977) along with Mar-
mosa lepida, M. emiliae, and M. contrerasi,
to Thylamys. Creighton (1984) also recog-
nized five groups in his revision of the genus

4 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Marmosa (s.1.). He placed four of these (ci-
nerea, murina, noctivaga, and microtarsus)
in the subgenus Marmosa and the fifth (e/e-
gans) in the subgenus Thylamys.

Despite Creighton’s (1984) conservative
treatment of Marmosa (s.l.), each of the
groups he recognized corresponds to a genus
in the sense used by Kirsh & Calaby (1977)
and Reig et al. (1985, 1987). Creighton’s
elegans-group equals Thylamys and corre-
sponds closely to Tate’s elegans group, but
is more restrictive than construed by Ca-
brera (1958), Kirsch & Calaby (1977), and
Reig et al. (1985, 1987). We recognize Thy-
lamys Gray, 1843 as a genus that includes
the species 7. elegans (genotype), 7. ma-
crura (Olfers, 1818:205; Didelphis grisea
Desmarest, 1827, is a Junior synonym), T.
pallidior, T. pusillus, and T. velutinus.
Creighton’s cinerea-group corresponds to
that of Tate (1933) as used by Reig et al.
(1985, 1987) under the generic name M7-
coureus Lesson, 1842 (Micoures Reig et al.
1987, is an incorrect subsequent spelling),
to which we assign the species Micoureus
cinereus (genotype), M. constantiae, M. re-
gina, and M. alstoni. Creighton’s murina-
group is similar to Tate’s (1933) group by
the same name except that Creighton also
included M. lepida. The earliest available
name for Creighton’s murina-group is Mar-
mosa Gray, 1821, which we use here as a
genus in a narrower sense than treated by
Reig et al. (1985, 1987). As we understand
it, Marmosa (s.s.) includes Marmosa an-
dersoni, M. canescens, M. lepida, M. mex-
icana, M. murina (genotype), M. robinsoni,
M. rubra, M. tyleriana, and M. xerophila.
Reig et al. (1985, 1987) combined Tate’s
noctivaga and murina groups under Mar-
mosa. However, Creighton (1984) showed
that Tate’s (1933) noctivaga group, with the
addition of M. parvidens (which Tate had
included in his microtarsus group), and the
more recently described taxa M. cracens and
M. handleyi, is a natural unit of closely re-
lated species for which the name Marmo-
sops Matschie, 1916, is available. Marmo-

sops includes the species M. cracens, M.
dorothea, M. fuscatus, M. handleyi, M. im-
pavidus, M. incanus (genotype), M. invictus,
M. noctivagus, and M. parvidens. This is
similar to Reig et al.’s (1987:7) composition
of Marmosops, which they ranked as a sub-
genus of Marmosa.

Creighton’s (1984) fifth taxon, the micro-
tarsus-group, is nearly identical to Tate’s
microtarsus section with the addition of the
more recently described M. agricolai. How-
ever, Kirsch & Calaby (1977) and Reig et
al. (1985, 1987) followed Cabrera (1958) in
combining Tate’s (1933) elegans group and
microtarsus section under Thylamys, along
with some other taxa that Creighton (1984)
showed belong elsewhere. Creighton includ-
ed nine species in his microtarsus-group,
although here we recognize only six. No ge-
nus-group name is available for this species
complex for which we propose the name:

Gracilinanus, new genus

Type species. —Didelphys microtarsus
Wagner, 1842:359; type locality ““Ypane-
ma’”’ (=Bacaetava), Sao Paulo, Brazil.

Etymology. —From the Latin gracilis
(thin, slender) and Greek nanos (dwarf)
combined to reflect the small size and slen-
der or gracile form of the species included
in this genus.

Distribution. —Gracilinanus contains six
living species whose composite geographic
range is below 3000 m in the tropical and
subtropical zone east of the Andean Cor-
dillera of South America.

Diagnosis and description.—A genus of
small mouse opossums (head and body,
85.0—130.0 mm; tail, 90.0-—150.0 mm). The
ratio of tail length to head-and-body length
is always greater than 1.3 and usually less
than 1.5. Dorsal pelage ranges from bright
reddish brown to dull brownish gray. The
tail has small (more than 40 per cm) round-
ed to square scales, arranged in annular rows.
Triplet interscalar hairs are subequal in
length with the middle hair about three scale
rows long. The diameter of the middle hair

VOLUME 102, NUMBER 1

is about twice that of the lateral hairs, but
not as stout and petiolate as in Marmosops.
The tail is weakly bicolored in some species
(G. agilis and G. marica) and unicolored
fuscous in others (G. aceramarcae, G. dryas,
and G. microtarsus). Claws on the manus
do not extend beyond terminal digital pads.
The central palmar surfaces lack the gran-
ular appearance of those of Thylamys and
Monodelphis. All palmar and plantar tu-
bercles (description by Creighton 1984) are
present and separated by at least a double
row of granules. Sparse granules are present
on the central plantar surface. Granules on
the proximal ventral surfaces of the digits
are fused into transverse bars. The skull lacks
postorbital processes; however, the su-
praorbital margin of the frontals may be
beaded in larger individuals of some species.
The lambdoidal crest is weakly developed
or absent except in the largest individuals.
The hard palate is highly fenestrated, usu-
ally with three pairs of medial fenestrae
(medial, posteromedial, and mesolateral—
described in Creighton 1984). The postero-
lateral fenestrae are moderate in size, usu-
ally about a third to half the breadth of the
last upper molar in length. The nasals are
moderately expanded laterally at the max-
illo-frontal suture. The auditory bullae are
relatively large compared with those of
Marmosa, Micoureus, and Marmosops, but
proportionately smaller than in Thylamys.
A slender anteromedial process of the ali-
sphenoid portion of the auditory bulla is
always present, although frequently dam-
aged during specimen preparation. The sec-
ond upper premolar is always larger than
the third. The incisors increase slightly in
size from 12 through I5. The lower canines
are relatively short compared with those of
Marmosa, Micoureus, and Thylamys, but
are not as short or premolariform as in Mar-
mosops.

Species of Gracilinanus are distinguish-
able from those of Marmosops by the shape
and arrangement of tail scales and bristles,
the non-premolariform lower canine, and

the presence of pectoral mammae. They can
be separated from species of Thylamys by
the absence of seasonal fat deposits in the
tail (thus non-incrassate), absence of dense-
ly granular central palmar and plantar sur-
faces, and relatively longer digits and broad-
er interdigital pads on manus and pes.
Gracilinanus can be distinguished from
Marmosa and Micoureus by the lack of
postorbital processes on the frontals and by
the annular arrangement of minute scales
on the tail.
Included species. —

Gracilinanus aceramarcae (Marmosa ac-
eramarcae Tate, 1931:12; type locality
‘Rio Aceramarca, tributary of Rio Un-
duavi, Yungas,”’ La Paz, Bolivia).

Gracilinanus agilis (Grymaeomys agilis
Burmeister, 1854:139; type locality ““La-
goa Santa,’ Minas Gerais, Brazil).

Synonyms:

Marmosa beatrix Thomas, 1910:502; type
locality ““Ipu,”’ Ceara, Brazil.

Marmosa muscula Shamel, 1930a:83;
type locality ““Kilometro 182” (=Ria-
cho Pilaga, 10 miles northwest of Km
182), Formosa, Argentina.

Marmosa formosa Shamel, 1930b:311;
renaming of M. muscula Shamel,
1930a, preoccupied by Didelphis (Mar-
mosa) musculus Cabanis, 1848.

Marmosa agilis chacoensis Tate, 1931:10;
type locality ““Sapucay,” Paraguay.

Marmosa agilis buenavistae Tate, 1931:
10; type locality ““Buenavista, Depart-
ment of Santa Cruz, Bolivia.”

Marmosa agilis peruana Tate, 1931:11;
type locality ““Tingo Maria, Rio Hua-
llaga,”” Huanuco, Peru.”

Marmosa unduaviensis Tate, 1931:11;
type locality ““Pitiguaya, Rio Unduavi,
Yungas,”’ La Paz, Bolivia.

Marmosa blaseri Miranda-Ribeiro, 1936:
373; type locality “S. Bento,” Goias,
Brazil.

Thylamys rondoni Miranda-Ribeiro,

6 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1936:387; type locality “‘salto do Se-
potuba e S. Joao da Serra do Norte,”
Mato Grosso, Brazil.

Gracilinanus dryas (Marmosa dryas Thom-
as, 1898:456; type locality ““Culata, Me-
rida, Venezuela’’).

Gracilinanus emiliae (Marmosa emiliae
Thomas, 1909:379; type locality “Para,
Brazil’).

Synonym:

Marmosa agricolai Moojen, 1943:2; type
locality ““Crato, Ceara,” Brazil.
Gracilinanus marica (Marmosa marica
Thomas, 1898:455; type locality ““R. Al-
barregas, Merida, Venezuela’’).
Gracilinanus microtarsus (Didelphys micro-
tarsus Wagner, 1842:359; type locality
‘““Ypanema,”’ Sao Paulo, Brazil).

Synonyms:

Marmosa microtarsus guahybae Tate,
1931:10; type locality “‘Island of Gua-
hyba near Porto Alegre, Rio Grande do
Sul,”’ Brazil.

Marmosa herhardti Miranda-Ribeiro,
1936:382; type locality ““Humboldt,”
Santa Catarina, Brazil.

Literature Cited

Burmeister, H. 1854. Systematische Uebersicht der
Thiere Brasiliens, welche wahrend einer Reise
durch die Provinzen von Rio de Janeiro und
Minas Geraés gesammelt oder beobachtat wur-
den von Dr. Hermann Burmeister. I. Sauge-
thiere (Mammalia). G. Reimer, Berlin, x + 342
pp.

Cabanis, J. 1848. Saeugethiere. Pp. 766-786 in R.
Schomburgk, ed., Versuch einer Fauna und Flora
von Britisch-Guiana, in Reisen in Britisch-
Guiana in den jahren 1840-1844. J. J. Weber,
Leipzig, 3:vi1l + 531-1260.

Cabrera, A. 1919. Genera Mammalium. Museo Na-
cional de Ciencias Naturales, Madrid, 177 pp.,

17 pls.

. 1958. Catalogo de los mamiferos de America

del Sur.— Revista del Museo Argentino de Cien-

cias Naturales “Bernardino Rivadavia,”’ Cien-
cias Zoologicas 4(1):iv + 307 pp.

Creighton, G. K. 1984. Systematic studies on opos-
sums (Didelphidae) and rodents (Cricetidae).

Unpublished Ph.D. dissertation, University of
Michigan, xi + 220 pp.

Desmarest, A. G. 1827. Sarigue. Pp. 392-399, Vol.
47 inF. G. Cuvier, ed., Dictionnaire des sciences
naturelles, dans lequel on traite méthodique-
ment des différens étres de la nature considérés
soit en eux-mémes, d’apres |’état actuel de nos
connoissances, soit relativement a l’utilité qu’en
peuvent retirer la médecine, l’agriculture, le
commerce et les arts. F. G. Levrault, Strasbourg;
Le Normant, Paris, 1816-1845, 61 Vols., 12
Vols. pls.

Gilmore, R. M. 1941. Zoology. Pp. 314-319 in J. C.
Bugher, J. Boshell-Manrique, M. Roca-Garcia,
and R. M. Gilmore, eds., The susceptibility to
yellow fever of the vertebrates of eastern Co-
lombia.—American Journal of Tropical Medi-
cine 21:309-333.

Gray, J. E. 1821. On the natural arrangement of ver-

tebrose animals.—London Medical Repository

15:297-311.

. 1843. List of the specimens of Mammalia in

the collection of the British Museum. British

Museum (Natural History), London, xxviii +

216 pp.

Kirsch, J. A. W., & J. H. Calaby. 1977. The species
of living marsupials: An annotated list. Pp. 9-
26 in B. Stonehouse and D. Gilmore, eds., The
biology of marsupials, University Park Press,
Baltimore, viii + 486 pp.

Lesson, R. P. 1842. Nouveau tableau do Régne An-
imal. Mammiféres. Arthus-Bertrand, Paris, 204

pp.

Marshall, L. G. 1981. The families and genera of
Marsupialia. — Fieldiana: Geology, new series 8:
1-65.

Matschie, P. 1916. Bemerkungen iiber die Gattung

Didelphis L.—Sitzungsberichte der Gesellschaft
Naturforschender Freunde, Berlin 1916(1): 259-
272;

Miranda-Ribeiro, A. 1936. Didelphia ou Mamma—
lia—ovovivipara.— Revista do Museu Paulista,
Sao Paulo 20:245-427.

Moojen, J. 1943. Algunos mamiferos colecionados
do nordeste do Brasil com a descriaao de duas
especies novas e notas de campo.— Boletim do
Museu Nacional, Rio de Janeiro, Zoologia 5:1—
14.

Olfers, I. von. 1818. Bemerkungen zu Illiger’s Ue-
berblik der Saugthiere nach ihrer Vertheilung
uber die Welttheile, riicksichtlich Siidamerican-
ischen Arten [chapter 10]. Pp. 192-237 in W.
L. Eschwege, Journal von Brasilien, oder ver-
mischte Nachrichten aus Brasilien, auf
wissenschaftlichen Reisen gesammelt. Vol. 15,
heft 2 in F. T. Bertuch, ed., Neue Bibliotek des
wichtigsten Reisen-beschreibungen zur Erwei-
terung der Erd- und Volkerkunde; in Verbin-

VOLUME 102, NUMBER 1

dung mit einigen anderen Gelehrten gesammelt
und herausgegeben .... Gr. H. S. pr. Landes-
industries-comptoirs, Weimar.

Reig, O. A., J. A. W. Kirsch, & L. G. Marshall. 1985.
New conclusions on the relationships of the
opossum-like marsupials, whith [sic] an anno-
tated classification of the Didelphimorphia.—
Ameghiniana 21(2-4):335-343.

—.. ,& . 1987. Systematic relation-
ships of the living and Neocenozoic American
““opossum-like”” marsupials (suborder Didel-
phimorphia), with comments on the classifica-
tion of these and of the Cretaceous and Paleo-
gene New World and European metatherians.
Pp. 1-90 in Possums and opossums: Studies in
evolution. Surrey Beatty & Sons PTY Limited,
New South Wales, frontispiece, 1xxii + 400 pp..,
4 pls.

Shamel, H. H. 1930a. A new murine opossum from

Argentina.—Journal of the Washington Acad-

emy of Sciences 20:83.

1930b. A new name for Marmosa muscula

Shamel.—Journal of Mammalogy 11:311.

Tate, G. H. H. 1931. Brief diagnoses of twenty-six

apparently new forms of Marmosa (Marsupi-

alia) from South America.— American Museum

Novitates 493:1-14.

. 1933. Asystematic revision of the marsupial

genus Marmosa.—Bulletin of the American

Museum of Natural History 66:1-250, 26 pls.,

1 table (9 sections, pocketed).

Thomas, O. 1888. Catalogue of the Marsupialia and

Monotremata in the collection of the British

Museum (Natural History). British Museum

(Natural History), London, xiv + 401 pp., 33

pls.

. 1898. On seven new mammals from Ecuador

and Venezuela. — Annals and Magazine of Nat-

ural History, Series 7 1:451-457.

. 1909. New species of Oecomys and Marmosa

from Amazonia. — Annals and Magazine of Nat-

ural History, Series 8 3:378-380.

. 1910. On mammals collected in Ceara, N. E.

Brazil by Fraulein Dr. Snethlage.— Annals and

Magazine of Natural History, Series 8 6:500-

503.

Wagner, A. 1842. Diagnosen neuer Arten Brasilischer
Saugethiere.— Archiv fiir Naturgeschichte 8(1):
359-360.

(ALG) Biological Survey Field Station,
National Ecology Research Center, U.S. Fish
and Wildlife Service, National Museum of
Natural History, Washington, D.C. 20560;
(GKC) Division of Mammals, National
Museum of Natural History, Washington,
D.C. 20560.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 8-13

DESCRIPTION OF A ROCK-DWELLING CICHLID
(TELEOSTEI: CICHLIDAE) FROM
LAKE MALAWI, AFRICA

J. R. Stauffer, Jr. and J. M. Boltz

Abstract. —A new species of cichlid fish of the Pseudotropheus zebra complex,
P. xanstomachus, is described from Lake Malawi, Africa. The species is en-
demic to the Maleri Islands. The species is morphologically similar to the
sympatric Pseudotropheus barlow McKaye & Stauffer, but the species are dis-
tinguishable in both coloration and shape.

The rock-dwelling cichlids (“mbuna’’)
which inhabit the rocky shores and rock
outcroppings of Lake Malawi are repre-
sented by many species which are endemic
to particular islands. Ribbink et al. (1983)
published a survey of this group and Lewis
et al. (1986) published a guide to the fishes
of the Lake Malawi National Park, which
discussed the distribution of many of the
mbuna. The purpose of this paper is to de-
scribe a new species closely related to Pseu-
dotropheus zebra (Boulenger) endemic to a
group of islands collectively referred to as
the Maleri Islands, located in Lake Malawi,
Malawi, Africa. Ribbink et al. (1983) rec-
ognized this form and referred to it as P.
zebra “yellow throat.”

Methods and materials. —Standard length
(SL) is used throughout. External counts and
measurements follow Barel et al. (1977).
Scale counts in the lateral line series do not
include scales in the overlapping portion of
the lower lateral line. Except for gill raker
meristics, which were recorded from the
right side, all counts and measurements were
made on the left side of the fish. Vertebral
counts were made from radiographs. All
specimens were collected at Nakantenga Is-
land (34°39’E, 13°55’S), Lake Malawi, Af-
rica (Apr 1984 and Aug 1987).

The new species is compared with P. bar-
lowi by color descriptions of fresh speci-
mens, meristic differences, and shape dif-

ferences. Body shape differences were
compared using sheared principal compo-
nent analysis (Humphries et al. 1981, Book-
stein et al. 1985). This analysis quantifies
shape differences between the two popula-
tions independent of size of the individuals
(Reyment et al. 1984).

Pseudotropheus xanstomachus, new species
Figs. 1, 2; Tables 1, 2

Pseudotropheus zebra, Ribbink et al., 1983:
162 (an part).

Holotype. —National Museum of Natural
History (USNM) 297268, adult male, 63.3
mm, Nakantenga Island, Lake Malawi, 3-
7m, 4 Aug 1987.

Paratypes.—USNM 297269, 11 (47.0-
70.8 mm), data as for holotype; USNM
297270, 4 (63.5-—78.5 mm), Nakantenga Is-
land, Lake Malawi, 12 m, 19 Apr 1984.

Diagnosis. —This species fits the descrip-
tion of the genus Pseudotropheus as given
by Regan (1921) in that: 1) its jaws have
several rows of teeth, with the ones in the
outer row bicuspid and those in the inner
rows tricuspid; 2) it has 16-18 dorsal-fin
spines, 7—8 dorsai-fin rays, 3 anal-fin spines,
and 6-7 anal-fin rays. However, as noted
by Ribbink et al. (1983), Fryer (1957:350)
regarded the genus Pseudotropheus as being
‘“‘rather ill-defined.’ Pseudotropheus xan-

VOLUME 102, NUMBER 1

Fig: 1.

stomachus differs from other members of
the group in coloration. It differs from the
only described sympatric member of the
group in the number of anal-fin rays, shape
of the lower pharyngeal bone, the length of
the dentigerous surface of the lower pha-
ryngeal bone, and in body shape.
Description.— Morphometric ratios and

P. barlowi

Fig. 2. Lower pharyngeal bones of the holotypes of Pseudotropheus barlowi and Pseudotropheus xansto-
machus.

ies fe", ag : 2
eA OS eee
he ES it ES

Holotype (USNM 297268) of Pseudotropheus xanstomachus.

meristics are presented in Table 1. Body
moderately compressed and elongate; jaws
isognathous (Fig. 1). Teeth on lower jaw in
3-4 rows, those on premaxilla in 4 rows;
teeth in outer rows bicuspid, with occasion-
al conical lateral tooth in some individuals;
teeth in inner rows tricuspid; 12 teeth in
outer row of left lower jaw of holotype, 9-

HAA
7)
4

3

ee”

h

lineal
Ate,

P xanstomachus

10 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Morphometric characters and meristics of Pseudotropheus xanstomachus. Range includes holotype

and 15 paratypes.

12 in paratyes. Pectoral fins with 13 rays in
holotype and 10-14 in paratypes; anal fin
with 7 rays in holotype and 6-7 in para-
types; caudal fin emarginate. Vertebrae of
holotype 14 + 14 (abdominal + caudal);
paratypes 12-14 + 14-15. Lower pharyn-
geal bone of holotype triangular in outline,
length of dentigerous surface 59% of lower
pharyngeal bone length; pharyngeal teeth in
left posterior row 23, those in left median
row, 13. Holotype with 5 scale rows on
cheek, paratypes with 5-6; pored scales along

Holotype Mean SD Range
Standard length, mm 63.3 62.9 Tell 47.0-78.5
Head length, mm DOS 20.4 2.4 15.3-26.0
Percent Standard Length
Head length 32.4 32.4 0.7 31.5-33.4
Snout to dorsal By) 33.3 1.2 30.9-35.0
Snout to pelvic 43.4 43.1 3 40.8-45.2
Greatest Body depth 36.7 34.2 1.4 31.8-36.5
Caudal peduncle length 12.8 13.5 i) 11.2-15.5
Least caudal peduncle depth 13.0 12.9 0.3 12.3-13.4
Pectoral-fin length 26.5 25.6 1.8 22.4-28.2
Pelvic-fin length 31.8 ihe 4.5 21.6-36.4
Dorsal-fin base length 60.7 60.4 1.1 58.6-62.2
Percent Head Length
Horizontal eye diameter 30.7 313 1.1 28.5-32.8
Vertical eye diameter 30.2 30.9 123 28.1-—33.3
Snout length 33.7 35.1 1.4 33.2-37.7
Postorbital head length 40.5 40.0 0.8 38.5-41.2
Preorbital depth DAES) 20.3 12 18.2-22.6
Lower jaw length 34.6 33.6 12 32.0-36.5
Interorbital width 28.3 28.3 2.4 25.3-33.6
Cheek depth 30y 30.0 2.8 24.5-34.4
Head depth 98.5 95.1 3.0 91.1-100.1
Counts
Lateral line scales 30 29.9 0.5 29-31
Scale rows on cheek 5 5.4 0.5 5-6
Dorsal-fin spines i 7s 0.5 16-18
Dorsal-fin rays 8 7.9 0.4 7-8
Pectoral-fin rays 13 13.3 1.0 10-14
Anal-fin rays 7 6.9 0.3 6-7
Gillrakers on first ceratobranchial 12 10.8 0.9 9-12
Gillrakers on first epibranchial 3 2.5 0.5 2-3
Teeth in outer row of left lower jaw 12 10.7 ital 9-12
Abdominal vertebrae 14 13.6 0.8 12-15
Caudal vertebrae 14 14.7 0.5 14-15

lateral line of holotype 30, and of paratypes
29-31; pored scales posterior to hypural
plate 2, with exception of two paratypes
which had scales missing from this area.
Gill rakers simple, first gill arch of holotype
with 12 on ceratobranchial (9-12 in para-
types); 3 on epibranchial of holotype and
2-3 in paratypes; and | between epibran-
chial and ceratobranchial.

Body coloration in freshly collected males
light blue with six black vertical bars; head
dark blue with one light blue interorbital

VOLUME 102, NUMBER 1

11

Table 2.—Sheared principal components (shape factors) of the untransformed morphometric data for Pseu-
dotropheus xanstomachus (n = 16) and Pseudotropheus barlowi (n = 10).

Sheared

Size PC 2 PC 3
Standard length O.215 —0.021 —0.082
Head length 0.207 =O —0.108
Horizontal eye diameter 0.183 0.095 —0.154
Vertical eye diameter 0.189 0.048 —0.048
Snout length 0.231 —0.088 eros
Postorbital head length 0.193 —0.107 —0.048
Preorbital depth 0.282 == Ue ly 0.042
Lower jaw length 0.210 0.070 —0.048
Interorbital width 0.275 —0.065 —0.014
Snout to dorsal fin origin 0.212 0.052 —0.149
Snout to pelvic fin origin 0.261 —0.064 etsy
Body depth 0.231 0.136 —0.050
Cheek depth 0.169 —0.594 0.447
Head depth 0.229 0.010 —0.067
Caudal peduncle length 0.219 —0.459 0.206
Least caudal peduncle depth 0.215 0.048 —0.050
Pectoral-fin length 0.258 O55 —0.158
Pelvic-fin length 0.311 0521 0.728
Dorsal-fin base length 0.215 —0.009 —0.069

bar, yellow gular and branchiostegal rays;
dorsal fin medium blue with grey flecks and
light blue marginal band; pectoral fins black;
pelvic fins black anteriorly fading to yellow-
brown posteriorly; anal fin blue; males with
three to five yellow ocelli on anal fin. Body
coloration of females similar, but some in-
dividuals have light brown body coloration
with brownish cast to fins.

The description of the coloration and the
color plate of the males in Ribbink et al.
(1983:162; plate 2a) agree with the above
description, except that they noted between
1-5 ocelli on the anal fin. Ribbink et al.
(1983:162) note that the females are “‘dark
brown sometimes almost black with darker
bars ....’” The differences in color of the
females may be associated with breeding
coloration and may be influenced by the
time of the year when the fish were ob-
served. However, all descriptions of the fe-
male coloration note a yellow chin and gular
region.

Etymology.—The specific epithet is de-
rived from the Greek “‘xanthos” and “‘sto-

machus”’ meaning yellow throat, which char-
acterizes the yellow gular region present in
both sexes.

Discussion. —Pseudotropheus xanstoma-
chus is morphologically similar to the P.
zebra complex in that it has a terminal
mouth, has three or four rows of teeth, of
which the outer rows are bicuspid, with the
occasional conical lateral tooth, and the in-
ner rows are tricuspid (see Ribbink et al.
1983). The only sympatric species of this
complex which is described is P. barlowi,
from which P. xanstomachus differs in col-
oration. P. barlowi males are bright gold,
and the females are uniformly brown
(McKaye & Stauffer 1986). Differences in
color patterns among cichlids, especially the
male’s, generally are recognized to be suf-
ficient to delimit valid species (Greenwood
1981, Hoogerhoud & Witte 1981).

In addition to coloration differences, P.
xanstomachus has six or seven anal rays
while P. barlowi has eight. The shape of the
lower pharyngeal bone between the two
species also differs. The angle of the suture

12 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

SHRD_PC3
-0.025

-0.050
-0.075
-0.100
=O. 125

=O2030

“0.260 ""-0.15>,°-0.10

-0.05 0.00

@

0.05
SHRD_PC2

Fig. 3. Plot of the sheared principal components of Pseudotropheus xanstomachus (solid) and Pseudotropheus

barlowi (open).

of the two posterior halves of the lower pha-
ryngeal bone is more acute and the resultant
depression is greater in P. barlowi (Fig. 2).
Moreover, the length of the dentigerous sur-
face of the lower pharyngeal bone of P. bar-
lowi was 76% (range = 74—78%) of the length
of the bone (n = 3; USNM 274782; USNM
274783), while the length of the dentigerous
surface of the lower pharyngeal bone of P.
xanstomachus was 62% (n = 4; range = 57—
67%).

An attempt was made to determine if there
were differences in body shape between the
two described sympatric species of the P.
zebra complex using a sheared principal
component analysis. Data for P. barlowi
were obtained from the original data sheets
used by McKaye & Stauffer (1986). Sheared
principal component analysis demonstrated
that there was no overlap between species
when the first sheared principal component
was plotted against the second sheared prin-
cipal component (Fig. 3). The first principal
component is interpreted as a size compo-
nent and the sheared components as shape,

independent of size (See Humphries et al.
1981, Bookstein et al. 1985). Thus, these
data were calculated using the untrans-
formed values of the morphometrics rather
than percent standard length or percent head
length. Those morphometrics which have
the highest loadings on the first sheared
principal component are cheek depth, pel-
vic-fin length, and caudal peduncle length,
while those which have the highest loadings
on the second sheared principal component
are pelvic-fin length, cheek depth, and snout
length (Table 2).

All of the specimens reported herein were
captured at a depth between 7—12 m at Na-
kantenga Island. Ribbink et al. (1983) re-
ported this form from all three Maleri Is-
lands: Namkoma, Maleri, and Nakantenga.
They stated that it was most common be-
tween depths of 2-8 m, and was rare in
water deeper than 12 m. Conversely
McKaye & Stauffer (1986) stated that P.
barlowi occurred primarily at depths greater
than 10 m and hypothesized that the oc-
currence of these brightly colored forms in

VOLUME 102, NUMBER 1

deeper water may be an adaptation related
to avoidance of surface predators (i.e., birds
and otters). Therefore, even though P. xan-
stomachus and P. barlowi are indigenous to
the same islands, they are in effect allotopic.

Acknowledgments

We thank the government of Malawi for
providing the facilities to make this research
possible. We thank L. Knapp of the Smith-
sonian Institution who arranged for ship-
ment of specimens from Malawi to the
USNM. We benefited from discussions with
K. R. McKaye. We are especially thankful
to D. Swofford who provided the SAS pro-
gram to conduct the sheared principal com-
ponent analysis. The original art work was
completed by M. Katz.

Literature Cited

Barcel ©. Dp N.. M. J. O. Van Oyen, F. Witte, & E. L.
M. Witte-Mass. 1977. An introduction to the
taxonomy and morphology of the haplochro-
mine Cichlidae from Lake Victoria. Part A.
Text. — Netherlands Journal of Zoology 27:333-
389.

Bookstein, F., B. Chernoff, R. Elder, J. Humphries, G.
Smith, & R. Strauss. 1985. Morphometrics in
evolutionary biology. Special Publication 15. The
Academy of Natural Sciences of Philadelphia.
Philadelphia, Pennsylvania. 277 pp.

Fryer, G. 1957. A new species of Gephyrochromis
(Pisces: Cichlidae) from Lake Nyasa, with notes

13

on its ecology and affinities.—Revue de Zool-
ogie et de Botanique Africaines 55:347-352.

Greenwood, P. H. 1981. The haplochromine fishes
of East African lakes. Cornell University Press,
Ithaca, New York. 839 pp.

Hoogerhoud, R. J. C., & F. Witte. 1981. Revision of
species from the “Haplochromis” empodisma
group. Revision of the haplochromine species
(Teleostei, Cichlidae), from Lake Victoria, Part
II.—Netherlands Journal of Zoology 31:232-
274.

Humphries, J. M., F. L. Bookstein, B. Chernoff, G. R.
Smith, R. L. Elder, & S.G. Poss. 1981. Multi-
variate discrimination by shape in relation to
size.—Systematic Zoology 30:291-308.

Lewis, D., P. Reinthal, & J. Trendall. 1986. A guide
to the fishes of Lake Malawi National Park.
World Wildlife Fund, Gland, Switzerland. 72
pp.

McKaye, K.R., & J. R. Stauffer, Jr. 1986. Description
ofa gold cichlid (Teleostei: Cichlidae) from Lake
Malawi, Africa.—Copeia 1986:870-875.

Regan, C. T. 1921. The cichlid fishes of Lake Nyara.—
Proceedings of the Zoological Society of London
1921:675-727.

Reyment, R. A., R. E. Blackith, & N. A. Campbell.
1984. Multivariate morphometrics. Academic
Press, New York, New York. 233 pp.

Ribbink, A. J., B. A. Marsh, A. C. Marsh, A. C. Rib-
bink, & B. J. Sharp. 1983. A preliminary sur-
vey of the cichlid fishes of the rocky habitats in
Lake Malawi.—South African Journal of Sci-
ence 18:149-310.

School of Forest Resources, The Penn-
sylvania State University, University Park,
Pennsylvania 16802.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 14-32

ASCIDIANS COLLECTED AROUND THE GALAPAGOS
ISLANDS USING THE JOHNSON-SEA-LINK
RESEARCH SUBMERSIBLE

Claude Monniot and Francoise Monniot

Abstract.—Ascidians were collected for the first time by a manned sub-
mersible on ocean-bottoms between 300 and 800 meters around the Galapagos
Islands. Solitary forms dominate; among the eight species collected, five are
new ones: Polyclinum johnsoni, Ciona pomponiae, Situla rineharti, Ascidia
fusca, and Styela psoliformis. They belong to surprisingly diverse families and

several have deep-sea characteristics.

The fauna of the eastern part of the trop-
ical Pacific Ocean is poorly known, espe-
cially the ascidians. Tokioka (1972) report-
ed on a small collection of ascidians from
the coast of Costa Rica, and recently Millar
(1988) identified some specimens from the
Galapagos and Ecuador. Nothing else is re-
ported about ascidians between Baja Cali-
fornia and Chile.

In November 1986, an expedition to the
Galapagos Islands collected deep-water or-
ganisms for pharmacological studies. Over
600 animals were collected by the R/V Sew-
ard Johnson and Johnson-Sea-Link- I
manned submersible (Harbor Branch
Oceanographic Institution, Fort Pierce, FL)
using a manipulator arm equipped with a
claw, a suction tube, and a grab sampler.
Among this collection there are eight species
of ascidians, five of which are new species.

The material is particularly interesting
because the specimens were collected un-
damaged on rocky bottoms, where a dredge
or any blunt device monitored from a sur-
face boat cannot operate. The animals are
unusually large compared to the size of the
ascidians collected in shallow water by
SCUBA during the same cruise and relative
to the deep-water ascidians on soft bottoms
elsewhere.

Observation from the submersible during
several dives suggests that ascidians are not

evenly distributed on the bottom, but live
in patches in restricted areas. They grow
attached to vertical walls, in small caves, or
on large cobbles; they are directly in contact
with loose sediment.

The solitary forms seem more abundant
in deep water than colonial ones, a situation
opposite to that in shallow water areas out-
side of the Galapagos Islands. This may be
due to the collecting method: solitary ascid-
1ans, especially large ones, are easily seen
from the submersible and can be collected,
but colonial ascidians normally live in crusts
on the undersides of rocks or pebbles, or in
deep crevices, and so escape notice. The col-
onies reported here were attached to cob-
bles.

Of the eight species collected, two are co-
lonial. They both belong to the family Poly-
clinidae (order Aplousobranchiata). One is
a new species of the genus Polyclinum,
named P. johnsoni after the submersible and
ship. The other species is probably Aplidium
californicum (Ritter & Forsyth 1917), one
of the most common Polyclinidae on the
shore nearby, but the samples are too small
and not fully enough developed to verify
their identity. The other ascidians belong to
surprisingly diverse families: one Cionidae,
two Ascidiidae, and one Octacnemidae
among the order Phlebobranchiata; and one
Styelidae and one Pyuridae in the Stolido-

VOLUME 102, NUMBER 1

branchiata. Several of these solitary ascid-
ians have deep-sea characteristics: Ciona
pomponiae, n. sp. has a particularly solid,
noncontractile tunic, and a very small gut
compared to the branchial sac; Situla rine-
harti, n. sp. belongs to a typical bathyal fam-
ily (Octacnemidae); Styela psoliformis, n.
sp. has the same characteristics as other
deep-water species of that genus.

The presence of a deep-water ascidian
fauna around the Galapagos, different from
the shore species collected by SCUBA, is
not surprising. The bottom temperature at
500-700 m was between 7°C and 5.5°C. The
water near the bottom was rich in suspend-
ed particles, but it is not known whether the
particles were plankton except for the nu-
merous crustacean larvae observed.

Type specimens are deposited at the
Smithsonian Institution, National Museum
of Natural History (USNM). Sorted sam-
ples received a Sea Pharm number (SP), in-
dicated here for each specimen.

Polyclinidae
Polyclinum johnsoni, new species
- Fig. 1A, B, C, D

Holotype. —USNM 18249; SP 25-XI-86-
35 iif:

Description. —Two colonies were collect-
ed close to Punta Espinosa, Fernandina Is-
land, at 306 m depth, attached to a sponge
underneath a large piece of rock. The colony
retained as a taxonomic voucher is a cush-
ion 2.5 cm in diameter. Color in life pale
yellow with translucent tunic through which
yellow zooids were clearly visible. Branchial
sac and internal papillae obscured by an or-
ange-yellow pigment. Zooids situated on
either side of cloacal canals that converge
towards the center of the colony where col-
ony’s sole common cloacal aperture opens.

The zooids have a large size range. Tho-
rax large compared to the abdomen and the
post-abdomen (Fig. 1A). Oral siphon star-
shaped with six triangular pointed lobes
above a strong sphincter that closes the si-

15

phon into a narrowed tube. Oral tentacles
vary in number with the zooids and are in-
serted in at least three circles, with three or
four orders of length, the largest being dor-
sal. At least 40 oral tentacles can be counted

‘in the largest zooids. The peripharyngeal

band lined with two high laminae and not
curved at the level of the dorsal tubercle.
Cloacal aperture close to the oral siphon and
provided with a sphincter which may make
the aperture into a tube when contracted.
Dorsal lappet (Fig. 1A) well developed and
inserted on a large base; not wide but its
length, which varies according to the loca-
tion of the zooid in the colony, can reach at
least the length of the thorax; has two lon-
gitudinal muscular bands (Fig. 1A). There
is a rounded button under the cloacal ap-
erture, as is usual in the genus. Thoracic
musculature is weak; longitudinal fibers
from the oral siphon stop very quickly and
do not pass the level of the first row of stig-
mata; there is an average of eight muscles
on each side.

Branchial sac, in both the live and pre-
served animals, visible by transparency; the
transverse vessels underlain by muscular fi-
bers and bear high papillae (Fig. 1B) con-
taining an orange-yellow opaque pigment.
The number of rows of stigmata is variable
according to size of zooids, and divisions of
stigmatal rows commonly seen. In the col-
ony studied there is a maximum of 16-17
rows of stigmata. In this example the fourth
row 1s divided as follows: 25 stigmata on
the right and 23 on the left side with 16
papillae on the right and 15 on the left on
the fourth transverse vessel. Branchial sac
not perforated along the endostyle or under
the rapheal papillae. Dorsal languets short
and wide, only slightly displaced to the left
side. The whole branchial sac has a rect-
angular shape with a right angle at the base
of the endostyle (Fig. 1A).

Abdomen slightly twisted; esophagus
short; the stomach placed anteriorly is sud-
denly cut in its pyloric part (Fig. 1C). There
is a ring-like post-stomach; intestine not

16 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

|)
ane e A
(\ —
ae

j
at

Fig. 1. Polyclinum johnsoni: A, Zooid; B, Detail of the branchial sac; C, Digestive tract; D, Larva. Ciona
pomponiae: E, Animal without tunic, right side; F, Neural region; G, Gonads on the digestive loop; H, Extremities
of gonoducts.

VOLUME 102, NUMBER 1

further differentiated into regions (Fig. 1C).
Anus opens at the level of the 7th or 8th
row of stigmata and has two lips.

Post-abdomen egg-shaped, linked to the
abdomen by a thin peduncle (Fig. 1A); ovary
is anterior followed by a cluster of testes.
The heart is terminal. One or two vascular
appendices extend from the post-abdomen
into the test.

Larvae (Fig. 1D) incubated in the cloacal
cavity. The zooids contain 1-3 larvae. The
trunks measure 550 wm. They have 3 ad-
hesive papillae, 7 pairs of thick epidermal
papillae arranged on a line on each side of
the body except for one pair which is lightly
displaced inside, and there are two areas of
epidermal vesicles on each side, one dorsal
and the other ventral (Fig. 1D).

Remarks.—This species differs from all
other Polyclinum with more than 15 stig-
mata rows in having higher papillae on the
transverse bars, a large number of stigmata
per row, and a peculiar distribution of the
pigments in the colony. The genus Polycii-
num is very homogeneous and the anatom-
ical differences between the species are slight,
sO it is not surprising that deep-water species
do not differ substantially from shallow-
water species. |

There is only one Polyclinum species
known to live deeper than P. johnsoni, n.
sp.; P. neptunium Hartmeyer, 1912, col-
lected at 106 m and 318 m off the Cape of
Good Hope.

The species is named after the submers-
ible from which it was collected.

Cionidae
Ciona pomponiae, new species
Figs. LE, F, G, H,, 2A, B

Holotype. —USNM 18247, SP 21-XI-86-
2-1.

Description. —The unique specimen mea-
sures 15 cm in length; attached by the left
posterior side; oral siphon terminal; cloacal
aperture located '3 the way down the dorsal

17

side (Fig. 2A). Both siphons sessile. Test soft,
gelatinous, colorless in life and free of for-
eign particles; is at least 5 mm thick and
does not appear to consist of several layers
as in Ciona intestinalis. Body contracted in-
side tunic and no longer attached to it, even
at the level of the siphons; size of body re-
duced to 6.9 cm. Tunic much less retractile
than in other Ciona species.

Musculature on each side (Fig. 1E) forms
four muscular ribbons initiated from the oral
siphon and two ribbons from the cloacal
siphon. The two most dorsal muscles of the
oral siphon and the one most ventral muscle
from the cloacal siphon gather to form one
ribbon in the posterior part of the body. The
two ventral muscular bands issuing from
the oral siphon become narrow before
reaching the posterior part of the body and
two muscles reach the bottom of the bran-
chial sac (Fig. 1E). There is a network of
transverse fibers. This small number of
muscular bands is peculiar to this species.

There are few oral tentacles (16 large ones
and several very small) placed without or-
der on acrest. The length of some tentacles
allows them to protrude through the oral
aperture. The peripharyngeal band consists
of two unequal laminae; the anterior thin
and high, the posterior low and thick. There
is a marked dorsal V in which the neural
duct opens as a simple slit (Fig. 1F). Neural
gland close to the nerve ganglion. Dorsal
languets have an increased length poste-
riorly, measuring 6 mm in the anterior 4
and 18 mm near the esophagus.

Branchial sac (Fig. 2B) thin, the trabec-
ulae connecting it to the body wall long.
Transverse vessels of several orders regu-
larly distributed; those of the first order wide
and protrude into the branchial cavity, those
of the third or fourth orders are thin laminae
protruding into the branchial cavity. They
bear large papillae that are flattened longi-
tudinally and form hooks above the longi-
tudinal bars. Longitudinal bars thin; stig-
mata elongated, generally numbering six to

18 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Ciona pomponiae: A, Group of in situ individuals (from a submersible videotape); B, Part of branchial
sac. C: Ascidia clementea part of the branchial sac with dorsal lamina. Situla rineharti: D, Part of the branchial
sac; E, Habitus left side; F, The same without tunic. (Scales: A = 10 cm; B, C, D = 1 mm; E, F = 1 cm.)

VOLUME 102, NUMBER 1

eight in a mesh; most crossed by a high
parastigmatic vessel bearing papillae (Fig.
2B); often divided under this vessel. If the
division is complete, the stigmata on each
side of the vessel are shorter than the others,
and a very thin new parastigmatic vessel
without papillae can occur above them.

Digestive tract clearly situated under the
branchial sac; enlarged stomach has internal
plications visible by transparency; most
posterior fold prolonged and ends in a small
ampulla at its junction with the intestine;
intestinal wall thin and gut contents can be
seen through it. Rectum elongated and ends
in a non-lobated narrow anus; posterior in-
testine encircled by a well developed pyloric
gland.

The pear-shaped ovary is located under
the intestinal loop (Fig. 1E, G). Testis dif-
fuse, consisting of numerous acini which coat
the intestinal wall; ducts join to form a com-
mon sperm duct at the level of the anterior
part of the ovary; i.e., oviduct and common
sperm duct originate at the same point.
Gonoducts contain large numbers of eggs
(400 wm in diameter) and spermatozoa, and
pass beyond the anus; oviduct opens in a
flat horseshoe-shaped papillae (Fig. 1H).
Male papilla has several small apertures;
there is no pigment spot at this level.

Remarks.—This species is very uncom-
mon for a Ciona. Although very few species
occur in shallow waters, the genus is better
differentiated deeper on the continental
slope. In the northeast Atlantic is found
Ciona gelatinosa Bonnevie, 1896 and Ciona
imperfecta Monniot & Monniot, 1977, and
in the northeast Pacific Ciona mollis Ritter,
1907.

Ciona mollis, known from California at
2011 m depth off San Nicolas Island, differs
from the new species in several important
characters: C. mollis’s size is smaller (3 cm
maximum) and its tunic is very soft. There
are 6 muscle bands on each side of the body,
all of them reaching the posterior part where
they fuse. There is circular musculature only
on the siphons. The oral tentacles are very

19

numerous, more than 200, while the present
specimen, much larger, has only 16. This
difference is perhaps not as significant as the
reduction of the number of oral tentacles
even with increased size in Ascidia translu-
cida according to Millar (1960) and in
Cnemidocarpa verrucosa according to Mon-
niot (1978). In C. mollis, furthermore, the
digestive tract does not have the same shape
as in this specimen, C. mollis’s intestine
curves immediately after the stomach, and
its gonoducts do not open near the anus.
Ciona mollis seems closer to the Atlantic
species C. gelatinosa Bonnevie 1896, re-
described by C. Monniot (1970), in the mus-
culature (7 bands), the shape of the gut, and
the location of the genital papillae.

The species is named after Shirley Pom-
poni, co-chief scientist of the cruise.

Octacnemidae
Situla rineharti, new species
Figs. 2D, E,;F, 3A, B, C, D

Holotype. —USNM 18244; paratype
USNM 18245; SP 14-XI-86-1-2 and 18-
XI-86-2-11.

Description. —Among several samples
collected, two specimens have been exam-
ined: one was designated as the holotype,
collected north of Floreana (Santa Maria Is-
land) at 790 m depth, and another (para-
type) collected south of Genovesa (Tower
Island) at 695 m depth. All specimens col-
lected are approximately the same size, al-
most spherical, with a cartilaginous white
tunic that is translucent more than trans-
parent (Fig. 2E). Holotype ovoid, attached
by the small ventral end (7 x 5 xX 5.2 cm).
The part of the tunic attached to the sub-
stratum torn but represents a narrow area
of attachment; in other specimens there is
no particular differentiation of the attach-
ment area. Oral aperture a gaping transverse
slit, made of two large lips located at half
the height of the body (Figs. 2F, 3A). Dorsal
lip protrudes more than ventral one. Cloacal

20 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

aperture not obvious; it is a simple hole at
the upper part of the body slightly displaced
to the dorsal side (Fig. 3A). Tunic on the
two lips of the oral siphon is thin, cartilag-
inous, and glossy, radially wrinkled, and dif-
ferent from the thick test (at least 3 mm)
around the rest of the body. Border between
the two kinds of tunic marked by a groove
at the limit of the lips.

Removed from the tunic, the body ap-
pears composed of three parts: the oral si-
phon formed of two well developed lips, a
branchial cavity widely spread separated
from the oral siphon by a ring of tentacles,
and a “‘nucleus” containing the gut and go-
nads.

The musculature is complex (Fig. 3A, B).
The muscles of the oral siphon, independent
of the body musculature, include radial fi-
bers regularly distributed on both lips; these
end at the circle of oral tentacles and the
peripharyngeal groove. The oral siphon’s
circular fibers are also regularly placed,
spread over both lips and gathered at the
lip corners. At each of those corners is a fan
of fibers belonging to the radial muscula-
ture, connecting the corners to the large
muscle located at the level of the ring of oral
tentacles.

Musculature of the cloacal siphon con-
sists of radial and circular fibers, weakly de-
veloped and anastomosed. It is limited to
the immediate vicinity of the siphon.

A large, strong ribbon of muscle fibers
encircles the body above the ring of oral
tentacles, and has a horseshoe shape opened
on each side of the endostyle at the level
where the body attaches to the substratum
(Fig. 3A). A muscular string binds the rib-
bon’s two free extremities. There are also
two areas of transverse muscles dorsally, and
ventrally two bundles more or less parallel
to the endostyle, connected to the “‘string”’
linking the posterior extremities of the large
horseshoe muscle ribbon.

Oral tentacles (Fig. 3C, D) situated on a
rim located posterior to the horseshoe mus-
cle-band. Dorsally the rim is low; its height

increases ventrally where it is lined by a flat
muscular band. Tentacles have a unique leaf-
shaped structure, narrowed at their inser-
tion on the rim. On the anterior side of each
tentacle is an ampulla which stains very
deeply with hemalum. The tentacles are of
two regularly alternating sizes (Fig. 3C).

The peripharyngeal groove is far from the
ring of tentacles in the dorsal half of the
body but comes close to it in the ventral
part. Dorsally, a series of irregular crests
separate the peripharyngeal groove and the
tentacles and they may be considered as the
anterior rim of the peripharyngeal groove.
Dorsally the peripharyngeal groove is deep-
ly incurved (Fig. 3D) to reach the dorsal
tubercle. The latter is rather far from the
neural ganglion; its aperture is a simple hole.
The neural gland is located on the left pos-
terior side of the neural ganglion.

There is no differentiated dorsal lamina.
The restricted space between the dorsal tu-
bercle and the opening of the esophagus is
marked by transverse elevations continuous
with more or less regular internal transverse
crests on the branchial sac; these elevations
can be considered as transverse vessels.
There is a short endostyle originating near
the oral tentacles and ending far from the
entrance of the esophagus (Fig. 3A, B). At
its posterior extremity, on the left, is a ret-
ropharyngeal groove which initially consists
of a single rim but divides in two wide and
flat ridges.

The entire surface placed between the
peripharyngeal band and the branchial stig-
mata is covered with small papillae. Bran-
chial cavity cup-shaped with the bottom
close to the digestive tract. The perfora-
tions, limited to two small areas in the flat
bottom of the cup on each side of the re-
tropharyngeal band, lie in thick tissue. The
perforations are oval (Fig. 2D) and generally
grouped in pairs; they are not ciliated, and
are larger than in the other known species
of Octacnemidae.

The digestive tract forms a simple loop
(Fig. 3B); esophagus opens by a simple hole

VOLUME 102, NUMBER 1 21

om

Fig. 3. Situla rineharti: A, Right side of the animal without tunic; B, Postero-dorsal side; C, Oral tentacles;
D, Peripharyngeal groove indentation,tentacles and neural ganglion. Ascidia clementea: E, Left side of the body,
tunic removed; F, Dorsal part of the tentacle ring and peripharyngeal groove; G, Intestinal loop with ovary and
oviduct.

a2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

in the branchial sac; stomach is wide and
has a glandular wall which appears some-
what wrinkled but has no internal plications
(Fig. 3A, B); its external oblique ridges may
be due to contraction. The intestine is thin
and transparent, ending in a plain-edged
anus placed beside the esophageal entrance.
The digestive tract and gonads are embed-
ded in a thick mesenchyme. The retro-
pharyngeal groove lies beside the middle of
the gut loop and indicates that the stomach
is on the right side.

The gonads (Fig. 3A, B) consist of a mas-
sive and protrusive ovary located between
the two branches of the digestive loop; testis
is composed of numerous acini; it is flat and
covers the ventral part of the ovary and each
side of the intestine. The oviduct opens on
the medial side of the intestine a short dis-
tance before the anus. The spermduct, full
of spermatozoa, opens next to the oviducal
aperture on a small papilla. The heart lies
alongside the intestine.

Remarks. —Octacnemidae having an oral
aperture with two large lips belong to four
genera: Dicopia Sluiter, 1905; Megalodico-
pia Oka, 1918; Situla Vinogradova, 1969;
and Cibacapsa Monniot & Monniot, 1983.
Cibacapsa is characterized by the total ab-
sence of a branchial sac. The distinctions
between the other genera are less obvious.
The difference between Dicopia and Meg-
alodicopia is the presence in the latter of a
peduncle, which in our opinion is not a dif-
ference of much importance. Dicopia differs
from Situla by the presence in Dicopia of a
true branchial cavity with a conical perfo-
rated area and oral tentacles forming a ring
of a smaller diameter than that of the bran-
chial cavity. In Situ/a the branchial cavity
opens widely and the perforated area is in
a flat disk around the digestive tract. The
Galapagos species is closer to Situla. The
main differences between the species of Di-
copia and Situla are summarized in Table
1. The new species is the only one having a
large imperforate space between the
peripharyngeal groove and the branchial tis-

sue, and its oral tentacles have a very orig-
inal structure.

Ritter (1907) has incompletely described
a species of Octacnemidae collected off San
Diego, California: Benthascidia michael-
seni. His account suggests he did not un-
derstand the anatomy of the animal very
well. The two specimens of that species pre-
served at the Smithsonian Institution are
dissected and their organs isolated, and so
it is impossible to reconstitute their original
disposition in the animal. In any case Rit-
ter’s material cannot be of the same species
as the new Galapagos one.

The species is named after Kenneth Rine-
hart, co-chief scientist of the cruise.

Ascidiidae
Ascidia clementea, Ritter, 1907
Figs. 2C, 3E, F, G

SP 23-XI-86-3-27.

Description. —One specimen was collect-
ed north of Isabella Island at 335 m depth.
The ovoid body is attached to the substra-
tum by the posterior part of the left side.
The maximum length of the animal is 6.8
cm. Oral aperture terminal; cloacal aperture
located at the anterior 14, the distance be-
tween the siphons being 2.9 cm. The tunic
in life is yellowish, covered with small cy-
lindro-conical papillae. Papillae and grooves
of the tunic are pigmented with ochre; both
apertures are 6-lobed. The test is partially
covered with epibionts: foraminifera, poly-
chaete tubes, bryozoans and molluscan egg-
capsules. One of the cheilostome bryozoans
consists of thecal rows, each zooid having
a long ventral extension penetrating deeply
into the ascidian’s tunic; the bryozoan col-
ony is raised above the tunic surface by these
extensions. Thickness of the tunic averages
2 mm; its internal side is soft and translu-
cent.

Body wall thin and transparent; the mus-
culature forms 2 bands of transverse parallel
fibers—a well developed dorsal band, and
a smaller ventral band—and longitudinal

23

VOLUME 102, NUMBER 1

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24 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

muscle fibers extend from the oral siphon.
On the right side all the muscles spread and
intermingle to make a loose network. On
the left side the longitudinal fibers extend
only a short way from the oral siphon, and
most of the left body wall is devoid of mus-
culature.

Seventy oral tentacles arise from an ele-
vated rim that has a muscular ring. The
tentacles are of at least four orders of size,
regularly alternated although the smallest
ones are not developed everywhere. Peri-
pharyngeal groove made of two unequal
laminae, the anterior one being larger, is
close to the tentacles, and the space between
is covered with thin papillae (Fig. 3F). It
does not make a deep ‘V’ dorsally; the dorsal
tubercle is low, J-shaped. Neural ganglion
distant from the dorsal tubercle; neural duct
is twice the length of the neural ganglion.
Rapheal band consists of two laminae which
measure five times the length of the dorsal
tubercle. Posteriorly, its edge is higher and
carries teeth that coincide with the trans-
verse vessels (Fig. 2C). After passing around
the esophageal entrance it becomes irregu-
lar. On the right side close to the esophagus
the transverse vessels end without differ-
entiation.

Branchial sac thin and flat (Fig. 2C). At
the level of the top of the intestinal loop
there are 100 longitudinal vessels. Main pa-
pillae are located at the junctions of the lon-
gitudinal and transverse vessels, and almost
everywhere in the branchial sac there are
smaller intermediate papillae, which are not
linked to the presence of parastigmatic ves-
sels. These vessels exist only where there is
duplication of a stigmata row. There are
four to six stigmata in a mesh, except on
the right side, where the meshes located near
the esophagus have more than 20 stigmata.

The gut forms an open loop with an ac-
centuated secondary curve (Fig. 3E). The
stomach appears plicated externally (Fig. 3E,
G). The intestine is deeply embedded in a
mass of large renal vesicles which spread
over the body wall. The anus is attached to
the body wall and has two slightly scalloped

lips. The specimen was not sexually mature.
The ovary, a network of tubes inside the
intestinal loop, does not spread out onto the
intestine (Fig. 3G). We have not found the
testicular acini that Ritter (1907) shows
against the stomach.

Remarks. —A. clementea has not been re-
ported since its description in 1907 based
on eight specimens: one collected by the A/-
batross near San Clemente Island at 654—
704 fathoms (station 4405), and seven spec-
imens at 21.8 miles to the south of San Nic-
olas Island, 1100 fathoms (station 4425).
There are a few small differences from Rit-
ter’s description: he does not mention the
tunic papillae and counts eight lobes at the
oral siphon instead of six. His figure 33
shows more than 75 small oral tentacles all
of equal size. He describes “‘deep narrow
plications” of the branchial sac, but his fig.
34 does not show this. In his discussion, he
recognizes that these plications do not al-
ways exist. All the other characters of our
specimen are identical to his description.

Ascidia fusca, new species
Fig. 4A, B, C, D

Holotype.—USNM 18248; SP 20-XI-86-
1-25.

Description. — The sole specimen was tak-
en at 375 m depth southeast of San Salvador
(James) Island. Maximum length 5.6 cm; it
was lying on its left side. Apertures distant
from each other (4.6 cm); have an undeter-
mined number of external large soft swell-
ings. Tunic dark brown in life (thus the
species name fusca) and its surface wrin-
kled. Some foraminifera and sedimentary
particles adhere to the settlement surface;
elsewhere on the tunic are small epibionts,
mostly bryozoans. The test is soft and its
internal layer is colorless.

Musculature strong and lies entirely on
the right side; is interrupted before it reach-
es the endostyle and dorsal lamina; com-
prises one ventral and one dorsal field, which
intermix in the middle of the right side. The
fields’ fibers are close together and nearly

VOLUME 102, NUMBER 1

joined; siphonal musculature weak. On the
right side of the oral aperture an array of
about 10 bands blends into both transverse
fields and disappears in the first 4% of the
body.

Oral tentacles arise from posterior part of
a velum linked by a muscular ring; there are
about 40 large ones and as many very small
ones alternating and irregularly developed;
bases crooked; peripharyngeal groove made
of two unequal laminae: anterior high and
thin and the posterior forming a flat ribbon.
There is a large space between the tentacle
ring and the peripharyngeal groove which
is indented into a deep dorsal V (Fig. 4B).
Dorsal tubercle is urn-shaped, opening with
a simple aperture (Fig. 4B). Neural ganglion
far from the dorsal tubercle at a distance of
3 times its length (0.6 cm in the type spec-
imen). Dorsal lamina has 2 rims which ex-
tend to the posterior part of the neural gan-
glion and then form a single lamina that is
thin and high. This carries digitiform pa-
pillae which prolong the transverse sinuses.
The lamina becomes progressively lower and
somewhat irregular as it nears the esopha-
gus, where it forms a low, toothed crest. At
this level on the right side, one of two trans-
verse vessels ends in a large lanceolate pa-
pilla.

Branchial sac flat; at the level of the top
of the intestinal loop there are 70 longitu-
dinal vessels on the right and 55 on the left,
the left side being much more narrow than
the right side. Longitudinal vessels very thin
and carry fingerlike main papillae where they
intersect transverse vessels. There are nei-
ther secondary papillae nor parastigmatic
vessels. The meshes are elongate and con-
tain 2 to 3 stigmata.

Gut (Fig. 4A, C, D) small; stomach, clear-
ly enlarged, has some plications only visible
on its medial side. Intestine inserted inside
the wall of the mantle. The 2 lips of the anus
have many small lobes. In this specimen a
translucent swelling is located close to the
anus. It has no discernable internal structure
and we do not know if it is a specific char-
acter or an artefact.

25

The ovary (Fig. 4C, D) consists of a mass
of tubes projecting internally, with edges
growing partly over the inside of the intes-
tine. The testicular lobules lie on the 2 sides
of the gut. The gonoducts open together next
to the anus. The renal vesicles are small,
almost invisible.

Remarks. —Ascidia fusca differs from
A.vermiformis (Ritter 1913), another deep-
water Ascidia in the East Pacific, by the for-
mer’s external shape, the length ofits cloacal
siphon and the long posterior extension of
its branchial sac. As in several of the rare
Ascidia known from the continental and is-
land slopes, Ascidia fusca has a relatively
small gut compared to the branchial sac, an
open primary gut loop, and a flat branchial
sac.

The species name refers to the dark tunic
of the animal.

Styelidae
Styela psoliformis, new species
Fig. 4E, F, G

Holotype. -USNM 18246, SP 16-XI-
86-1.

Description. —The sole specimen was
found in a cavity of a basaltic cobble near
Espanola island at 567 m depth. The visible
part of the still attached body is an oval disk,
14 x 11 mm, white, covered with low,
round, plate-like papillae which have a vas-
cular dilation full of blood granules. Around
the disk the tunic is thinner and also has
vascular ampullae; it strongly attaches the
animal to the substratum. Both apertures
protrude a distance of 6 mm, but they are
located in depressions, so they do not ex-
tend beyond the body surface after contrac-
tion. Tunic of the central disk opaque and
internally pearly. The rest of the body has
a thinner tunic that is soft and transparent.
Inside the tunic the body is flattened against
the substrate. The dorsal part of the body
wall is opaque and muscular, the ventral
side thin and transparent without muscu-
lature. At the boundary between the dorsal
and ventral side the mantle has a ring of

26 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Ascidia fusca: A, Left side of the body, tunic removed; B, Dorsal tubercle; C, External side of the gut
and gonads; D, Internal side of the gut and gonads. Styela psoliformis: E, Ventral side of the body, tunic removed;
F, Internal side of the body, branchial side removed; G, Gonad.

tissue rich in tan blood lacunae clearly seen velum. There are at least four different sizes
by transparency. which regularly alternate. The oral tentacles

About 30 oral tentacles are attached toa are wide and transversally flattened. The
thin crest located posteriorly to a large oral peripharyngeal groove has 2 equally pro-

VOLUME 102, NUMBER 1

truding rims. It is far from the tentacles on
the ventral part, nearer on the sides, and
dorsally makes a little but distinct V; it is
only curved around branchial folds number
1 and 2. The opening of the dorsal tubercle
has the shape of a C opened on the left. The
neural ganglion is in contact with it. The
dorsal lamina is high and smooth.

Branchial sac has 4 folds on each side in
the anterior part only; fold number 4 dis-
appears in the posterior *4 of the branchial
sac. Longitudinal vessels very numerous and
difficult to count; there are as many as 15
vessels between the folds and more than 20
on folds 1 and 3. Fold 4 has more than 15
longitudinal vessels anteriorly. The folds are
high and the longitudinal vessels close to
each other, with only one or two elongated
stigmata per mesh. Parastigmatic vessels are
generally present.

Gut a narrow loop (Fig. 4E, F). Stomach
and intestine weakly linked to the ventral
side of the body wall but rectum strongly
united to dorsal side. The globular stomach
has 14 obvious plications. A very small cae-
cum is located near the gastric aperture of
the pyloric gland. Intestine long; the short
rectum opens by a wide, lobed anus.

There are two gonads on each side (Fig.
4E, F), weakly linked to the body wall; each
is made of a double series of testes (Fig. 4G).
The central ovary of each gonad spreads
onto the internal part of these lobes, which
makes this structure intermediate between
the genera Styela and Cnemidocarpa. The
gonoducts open together at the margin of
the dorsal disk of the body wall.

Endocarps are evenly distributed on the
ventral side of the body (Fig. 4F). They are
very numerous and protrude along the ring
of tissue rich in tan blood lacunae which
marks the boundary between dorsal and
ventral halves of the body. The dorsal part
is devoid of endocarps. The cloacal aperture
is circled by thin cloacal tentacles arising
from a crest (Fig. 4F).

Remarks. — This species has several char-
acters which are similar to Styela milleri
Ritter, 1907, collected frequently from 900

27

to 4000 m depth along the Pacific coast from
California to Chile and also recorded in the
south Atlantic and the Indian Oceans. The
oral tentacles, dorsal tubercle, dorsal lami-
na, branchial sac, and gut are similar in both
species. In contrast, S. milleri has only one
gonad on each side and they are different in
structure from those of S. psoliformis. The
new species has the same appearance as S.
similis C. Monniot, 1969 from the conti-
nental slope of Europe and West Africa, and
these species have many anatomical char-
acters in common due to their similar body
shape. S. similis also has only one gonad on
each side, but its structure differs.

The gonad of S. psoliformis is morpho-
logically intermediate between typical Sty-
ela gonads and the very peculiar gonad of
Cnemidocarpa pfefferi (Michaelsen, 1904)
shown in Millar 1960; that large (5 cm),
round species lives on the slope of South
Georgia and South Shetland Islands. It has
two gonads on each side, each consisting of
a lobed ovary with each lobe encircled by
testes. The structure of the gonad of S. pso-
liformis resembles that of species of Pyura,
but this is only a convergence: all other char-
acteristics of the present species are typical
of the Styelidae family.

The animal looks like a psolid holothu-
rian after which it is named.

Pyuridae
Halocynthia hispida Herdman, 1881
Pig; 5

Synonymy: see Kott, 1985:342, fig. 169, pl.
7F, Millar (1988).
SP 24-XI-86-3-9 and 27-XI-86-1-9.

Description.—This species has two dif-
ferent types of morphology on the Gala-
pagos Islands: a hard form with a thick tunic
collected in shallow water by SCUBA and
a larger form with a soft hairy tunic collected
from deep water by the submersible. The
two forms also differ in internal structure,
and only a very careful analysis of all char-
acters has convinced us that they belong to
the same species. A reduction of the organs

28 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

linked to increased depth and decreased food
supply has been described in ascidians
(Monniot & Monniot 1978), but the case of
H. hispida in the Galapagos is extreme.

The shallow-water form was collected at
Bainbridge Rocks, east of San Salvador Is-
land, at a depth of 17 m. It was firmly at-
tached by its ventral side; both apertures
are erect, at almost the same level. The spec-
imen measures 10 X 7 X 5 cm, the aper-
tures separated by 4 cm. The side facing the
light is dark red in life. The tunic is wrinkled
and covered with small smooth spines that
become larger near the siphons, the largest
grouped on tubercles. Around the apertures
there are large branched spines 3 to 4 mm
in length. The thickness of the cartilaginous
tunic reaches 5 mm.

The body wall is 2—3 mm thick and fleshy.
The muscles are well developed: the exter-
nal circular ones make a continuous layer,
the internal radial muscles are gathered to-
gether in strong bands. The oral tentacles
are located behind a strong sphincter; they
number about 20 in three orders and are
only weakly pinnate. The peripharyngeal
groove has 2 equal rims and shows a clear
dorsal V. The dorsal tubercle is large, com-
plex and protruding (Fig. 5D). The dorsal
lamina has a double series of languets.

The middle level of the branchial sac has
10 high folds on each side. Anteriorly and
posteriorly, additional ventral folds appear,
so the maximum number of folds in this
specimen is 13 on the right and 12 on the
left side. The distance between 2 longitu-
dinal vessels is the same on the folds as
between them, so it is difficult to distinguish
the base of a fold. In the ventral part of the
branchial sac there are only 2 or 3 longi-
tudinal vessels between folds, the meshes
are larger there and they may have a dozen
stigmata.

The gut (Fig. 5C) is voluminous and oc-
cupies most of the left side. The esophagus
is long and the stomach is covered with a
hepatic gland of two parts: the anterior la-
mellar and the posterior cauliflower-like.

The intestine is attached to the body wall
and has a smooth edged bilobed anus.

The gonads are very numerous (13 on the
right and 10 on the left) and so massed and
interpenetrated (Fig. 5C) that they can only
be counted by their genital papillae. On the
left side they cover the descending limb of
the intestine. Each gonad consists of an ovary
surrounded by lobes of testes.

There are numerous endocarps on the
body wall, inside the gut loop and even pro-
truding from the anterior part of the stom-
ach (Fig. 5C).

The deep-water form is rounded; the larg-
est specimen measures 7 X 5 X 4cm with
a distance of 4 cm between the apertures.
The color in life is tan-brown with some
amounts of red near the siphons. The entire
tunic bears short branched spines with much
larger ones on the siphons; between the
branched spines there are smaller simple
ones, often placed in rings. The test is thin
(1 mm) and flexible. Internal organs are vis-
ible through the thin, transparent body wall.
The external layer of circular muscles is thin.
The internal radial muscles are gathered to-
gether in weak ribbons.

When the tunic is removed, the oral ap-
erture has 4 low lobes with undulated edges.
There is a small plain velum at the base of
the oral siphon. The oral tentacles are not
numerous (36) and occur in at least 4 orders.
Between the largest tentacles on the tentacle
crest there are numerous small digitiform
ones. The largest tentacles measure up to
13 mm but only have about 20 primary
branchings, which are themselves thin and
poorly branched in turn. The peripharyn-
geal groove has 2 equal laminae; it is curved
around each branchial fold and makes a large
dorsal V. The dorsal tubercle is round and
has a C-shaped aperture open on the ante-
rior left side (Fig. 5A). The neural ganglion
is elongated (15 mm), and thin. Nerves from
it are clearly visible: 2 anterior and 2 pos-
terior ones. The visceral nerve follows the
dorsal lamina and is accompanied by a neat
dorsal band; it originates on the right pos-

Fig. 5. Halocynthia hispida: A, Deep-water form, internal side of the body, branchial sac removed; B, Detail
of the right gonad; C, Shallow-water form, internal side of the body; D, Dorsal tubercle of the shallow-water
specimen.

30 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

terior side of the neural ganglion. In its an-
terior part the dorsal lamina consists of a
double row of languets; the left line has true
languets and is more developed than the
right line, which is made of prolongations
of the right transverse branchial vessels.

The branchial sac has 10 folds on each
side. The branchial formula is:

R.E.2(8)2(13)2(15)3(17)2(18)2(20)2(22)2
(21)3(20)3(16)2.DL

L.DL.2(17)2(21)2(22)3(22)2(21)2(19)3(15)
2(13)2(11)1(10)O.R

The 10th (most ventral) fold on the left is
only present in the anterior part of the bran-
chial sac. The folds are very high and largely
cover each other. The longitudinal vessels
are raised, and there are only two or three
between the folds. We counted 24 stigmata
in a median mesh between two folds and 8
to 10 in meshes close to the base of the folds.
It is only at the crest of the fold that the
number of stigmata in a mesh go down to
five or six. We have not observed spiral
stigmata at the crests of folds. The stigmata
are not very elongated; they are crossed by
parastigmatic vessels only when a division
begins in a transverse row. Between the folds
the transverse vessels are inflated into am-
pullae attaining several millimeters, much
like endocarps.

Gut short; a long esophagus leads to a
stomach with a 2-part liver (Fig. 5A), the
anterior one with parallel ridges and the
posterior with papillae on crests. The two
parts of the gland are separated by a nar-
rowing, but they remain united. At the nar-
rowing level there are parallel riges, each
dividing in two in the anterior part (Fig.
5A). At the pyloric end of the stomach the
hepatic gland stops abruptly. The intestine
is short and translucent. Rectum long; at-
tached along the dorsal lamina, and the anus
has two lobed lips.

On each side there is a cluster of four
parallel ovaries united at their proximal

ends. Each cylindrical ovary is surrounded
by ramified testes (Fig. 5B). Each ovary and
its testes are wrapped in a common mem-
brane. The testes give off long thin ducts
which join at the atrial surface of the ovary
to form a common spermduct. The com-
mon spermduct consists of canals coming
from both sides of the ovary. The sperm-
duct opens just behind the oviduct on a
common protruding papilla. The four left
Ovaries originate inside the gut loop and
open onto the intestine (Fig. 5A).

The entire body wall is provided with en-
docarps. The cloacal siphon has a short ve-
lum similar to the oral siphon. We have not
seen cloacal tentacles.

Remarks. — This species is recorded from
the Galapagos by Millar (1988) and the
specimens he describes are intermediate be-
tween our two forms. If we examine point
by point the differences between our shallow
and deep populations, we find that they are
almost all quantitative ones: thickness of
the tunic and body wall, gonad number, size
of the gut, and number of branchial folds.
The difference in the number of oral ten-
tacles has no taxonomic significance. The
most important contrast is the reduction of
the gut in deep-water specimens, and its po-
sition at the posterior part of the body. The
gonads are also posteriorly displaced in deep-
water animals.

The thinning of the tissues in deep-water
animals is probably linked to decreased food
supply. The species of Halocynthia have a
highly evolved branchial sac which is very
efficient in filtering phytoplankton, but the
deep samples certainly have less food avail-
able. In fact, the branchial sac is the only
organ not reduced; on the contrary, the
number of stigmata per mesh increases with
depth. The reduction in size of the gut is
correlated with this nutritional phenome-
non; it is a common characteristic for as-
cidians living on the slope such as in Ascidia
(see below). A similar but less accentuated
situation has been described for Ascidia

VOLUME 102, NUMBER 1

challengeri and Corella eumyota (Monniot
& Monniot 1983) on the slope in the Ant-
arctic Ocean and at Kerguelen.

The shifting of the gut and gonads pos-
teriorly may be a consequence of the loss of
tunic rigidity. In 1973 we reported that deep-
sea ascidians show a tendency to equilibrate
their visceral masses to keep their balance
on the substratum. In other species of Halo-
cynthia the disposition of the gonads on the
right side seems linked to the tunic rigidity:
H. ritteri from Japan has a hard tunic and
gonads located in the middle of the right
side, while H. papillosa, H. aurantium and
H. pyriformis, with thin tunics, have gonads
located posteriorly.

Except for two Arctic species, the tropical
and temperate Halocynthia are found in very
shallow water not deeper than 30 meters. Is
the presence of deep specimens in the Ga-
lapagos exceptional? We cannot answer be-
cause the deep rocky bottoms have not been
sampled enough, but this species represents
the most extreme case of ecological dimor-
phism known in an ascidian.

Conclusions

The bathyal ascidian fauna of the Gala-
pagos does not provide any evidence in sup-
port of an archaic fauna. All species belong
to genera and more advanced families which
are well represented in other oceans and
which show adaptations here to deep-sea
life.

The origin of the fauna is unknown. Rel-
atively few collections have been made along
the South American Pacific coast; we cannot
determine if the new species described here
are endemic or if they have been carried
north by the Humboldt Current. The Ga-
lapagos ascidian fauna differs greatly from
the assemblage of Antarctic ascidians known
from the south coast of Chile. Among the
eight species recorded from the Galapagos,
only three were previously known: Apii-
dium californicum, a tropical shallow-water

31

shallow-water species probably at its depth
limit here, Halocynthia hispida, widely dis-
tributed and known in shallow water from
both the Galapagos and Australia; and As-
cidia clementea, a north Pacific species. Sit-
ula and Ciona are genera which are well
represented in bathyal areas in other parts
of the world. It seems unlikely that the
Humboldt Current contributes to the oc-
currence or distribution of ascidian fauna
around the Galapagos islands.

Observations from the Johnson-Sea-Link
have shown that ascidians are a very small
part of the bathyal biomass which is mainly
comprised of sponges, cnidarians and echi-
noderms. Other filter-feeding animals, such
as bivalves or bryozoans, were also very
seldom seen. Except for Situ/a, which is
phytophagous and carnivorous, the other
ascidians we report have the same diet as
sponges: small suspended particles and dis-
solved substances. Sponges in Galapagos
deep-water habitats are large, abundant, and
diversified, but ascidians remain scarce even
though they are relatively large.

Acknowledgments

The R/V Seward Johnson and Johnson-
Sea-Link-1 cruise was funded by Harbor
Branch Oceanographic Institution, the U.S.
National Cancer Institute, Natural Products
Branch (contract no. NO1-CM-67919), and
SeaPharm Inc. Francoise Monniot is par-
ticularly thankful to the chief scientist of the
cruise, K. L. Rinehart and to Shirley A.
Pomponi who gave her the opportunity to
participate in the submersible dives and in
SCUBA collecting. In gratitude two of the
species reported here are named after them.

Literature Cited

Bonnevie, K. 1896. Ascidiae simplices og Asciidae
compositae fra Nordhavs-Expeditionen. —
Norske Nordhavs-Expeditionen 1876-78 23:1-
16.

Kott, P. 1969. Antarctic Ascidiacea.— Antarctic Re-
search Series 13:1—239.

32 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1985. The Australian Ascidiacea. Part I,
Phlebobranchiata and Stolidobranchiata. —
Memoirs of the Queensland Museum 23:1-440.
Hartmeyer, R. 1912. Die Ascidien der deutschen

Tiefsee-Expedition.— Deutschen Tiefsee-Expe-
dition 7:223-392.

Herdman, W. A. 1881. Preliminary report on the
Tunicata of the Challenger expedition.—Pro-
ceedings of the Royal Society of Edinburgh, part
3 Cynthiadae 11:233-240.

Millar, R. H. 1960. Ascidiacea.— Discovery Reports

30:1-160.

1988. Ascidians collected during the South-
east Pacific Biological Oceanographic Program
(SEPBOP).—Journal of Natural History 22:225-
240.

Monniot, C. 1969. Ascidies récoltées par la ““Tha-

lassa” sur la pente du plateau continental du

golfe de Gascogne: (3-12 Adut 1967).— Bulletin
du Muséum National d’Histoire Naturelle, Paris

(2),41(1):155-186.

1970. Ascidies récoltées par la ““Thalassa”’
sur la pente du plateau continental du golfe de
Gascogne (18-25 Octobre 1968).—Bulletin du
Muséum National d’Histoire Naturelle, Paris
(2),41(5):1131-1145.

1972. Dicopia antirrhinum n. sp. ascidie de
la pente du plateau continental du golfe de Gas-
cogne. Interprétation nouvelle de la famille des
Octacnemidae.— Cahiers de Biologie Marine 13:
9-20.

1978. Ascidies phlébobranches et stolidob-
ranches du sud de Il’Océan Indien.— Annales de
l'Institut Océanographique, Paris 54(2):171-—224.
—., & F. Monniot. 1973. Ascidies abyssales ré-

coltées au cours de la campagne océanogra-
phique Biacores par le “Jean Charcot”.—Bul-
letin du Muséum National d’Histoire Naturelle,
Paris (3), 121 Zoologie 93:289-475.

———,,& . 1977a. Quelques ascidies abyssales
du Sud-Ouest de l’Océan Indien.—Comité Na-
tional Francais de Recherches Antarctiques 42:
305-327.

——., & 1977b. Tuniciers benthiques pro-
fonds du Nord-Est Atlantique. Résultats des
campagnes Biogas.— Bulletin du Muséum Na-
tional d’Histoire Naturelle, Paris (3), 466 Zoo-
logie 323:695-720.

———., & 1978. Recent work on the deep-

sea tunicates. Oceanography and marine biol-

ogy, annual review, 16:181-—228.

——., & . 1983. Ascidies antarctiques et sub-
antarctiques. Morphologie et biogéographie. —
Mémoires du Muséum National d’ Histoire Na-
turelle, Paris (A), Zoologie 125:1-168.

Oka, A. 1913. Zur Kenntnis der zwei aberranten As-

cidiengattungen Dicopia Sluit. und Hexacroby-

lus Sluit.— Zoologischer Anzeiger 43(1):1-10.

1918. Megalodicopia hians n. g., n. sp., eine
sehr merkwurdige Ascidie aus dem japanischen
Meere. — Annotationes Zoologicae Japonense 9:
399-407.

Ritter, W. E. 1907. The ascidians collected by the

United States Fisheries Bureau Steamer Alba-

tross on the coast of California during the sum-

mer of 1904. University of California, Publi-
cations in Zoology, Berkeley 4(1):1—-52.

1913. The simple ascidians from the North-
eastern Pacific in the collection of the United
States National Museum.— Proceedings of the
United States National Museum 45:427-505.
—., & R.A. Forsyth. 1917. Ascidians of the lit-

toral zone of southern California. University of
California Publications in Zoology 16:439-512.

Sluiter, C. P. 1905. Die Tunicaten der Siboga-Ex-
pedition. Supplement zu der I. Abteilung: die
socialen und holosomen Ascidien.—Siboga Ex-
pedition Monographie 56A:129-139.

Tokioka, T. 1953. Ascidians of Sagami Bay collected

by His Majesty the Emperor of Japan. — Tokyo:

Iwanami Shoten 1-315.

. 1972. Onasmall collection of ascidians from

the Pacific coast of Costa Rica. Publications of

the Seto Marine Biological Laboratory 19(6):

383-404.

Vinogradova, N. G. 1969. On the finding of a new

aberrant ascidian in the ultrabyssal of the Ku-

rile-Kamchatka trench.—Bulletin of the Mos-
kovit Society of Natural History, Biological Part

3:27-42.

1975. On the discovery of two new species
of an aberrant deep-water ascidiacean genus Sit-
ula in the South-Sandwich trench.—Transac-
tions of the P. P. Shirshov Institute of Oceano-
logia 103:289-303.

Museum national d’Histoire naturelle,
Laboratoire de Biologie des Invertébrés
Marins et Malacologie, 55 rue Buffon, 75005
Paris, France.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 33-44

EUKROHNIA CALLIOPS, A NEW SPECIES OF
CHAETOGNATHA FROM THE NORTHERN GULF OF
MEXICO WITH NOTES ON RELATED SPECIES

Jerry A. McLelland

Abstract. —Eukrohnia calliops, a new species of Chaetognatha, was discov-
ered in deep-water plankton collections made at four stations in the northern
Gulf of Mexico in August 1987, and in February 1988. The new species appears
to occur in bottom water over the upper continental slope at depths between
200 and 700 m. Its large pigmented eyes easily distinguish it from other Eu-
krohnia species occurring in the Gulf of Mexico. It differs from the closely
related E. kitoui Kuroda, 1981, a species from Japanese waters, chiefly by the
size and shape of the eyes. Thirteen specimens of EF. calliops are studied and
compared with specimens of E. kitoui, E. proboscidea, and E. fowleri, all of
which possess pigmented eyes. The first record of E. proboscidea in the Gulf
of Mexico is documented and affinities within the proposed “‘proboscidea group”
are discussed. Distribution records of four species of Eukrohnia reported from
the Gulf of Mexico prior to this study are presented with a key separating the

Six species now known.

Members of the chaetognath genus Eu-
krohnia typically inhabit deep-water plank-
ton communities throughout the world (Al-
varino 1965, 1969 for zoogeography). Of
the nine currently described species of Eu-
krohnia (Casanova 1986), only three are
known to possess pigmented eyes: E. fowleri
Ritter-Zahony, 1909, E. proboscidea Fur-
nestin & Ducret, 1965, and EF. kitoui Ku-
roda, 1981. Eukrohnia fowleri has a well
known cosmopolitan distribution, while the
latter two species appear restricted to deep
coastal waters of the Atlantic Ocean and the
Pacific coast of Japan, respectively.

One hundred and eighty three specimens
ofan undescribed species of Eukrohnia with
large pigmented eyes were discovered in
plankton samples collected at four study
areas in the northern Gulf of Mexico in Au-
gust 1987 and February 1988 by personnel
of the Gulf Coast Research Laboratory
aboard the R/V Tommy Munro (Fig. 1, Ta-
ble 1). Quantitative samples were obtained
by towing horizontally at depths of 200-500

meters using a Niskin double-trip system
with 1 m diameter, 333 wm mesh nets.
Qualitative samples were obtained near the
bottom with 0.3 m diameter, 183 um mesh
nets attached to crab traps and allowed to
passively filter the bottom current. Only
material from the February 1988 cruise was
used in this work because specimens in the
August 1987 samples were in poor condi-
tion.

Class Sagittoidea Claus and Grobben, 1905
Order Phragmophora Tokioka, 1965
Family Eukrohniidae Tokioka, 1965

Genus Eukrohnia Ritter-Zahony, 1909

Diagnosis. —One pair of lateral fins
emerging at or near ventral ganglion; one
pair of tooth rows; apical gland cell com-
plex, or papillae tuft, at anteroventral point
of head; ventral transverse musculature in
trunk segment extending from neck to at
least posterior edge of ventral ganglion.

34 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

GULF OF MEXICO

YUCATAN

vs 90° 89° 88° 87°

Fig. 1.

86° 35° 84° 83° 82° g1° 80° 79°

Map showing locations of five study areas along the continental slope in the northern Gulf of Mexico

from which deep-water plankton collections were made in August 1987, and February 1988.

Eukrohnia calliops, new species
Figs. 2, 3, Tables 1—4

Material. —Holotype, 21.5 mm (USNM
119293), R/V Tommy Munro, Area 3,
(29°21'N, 86°57'W) northern Gulf of Mex-
ico, 200-500 m, 16 Feb 1988, 1910 CDT,
11 paratypes as follows: 1 specimen, 18.2
mm (USNM 119296), from type locality; 2
specimens 18.4, 15.0 mm (USNM 119294),
from Area 1 (29°00'N, 88°26'W), 677 m, 28
Feb 1988, 1930 CDT; 5 specimens, 14.4,
12.6, 11.7, 10.4, 8.4 mm (USNM 119295),
from Area 2 (29°18'N, 87°32’W), 200-500
m; 13 Feb) 1988; 1035 CDT; ‘bspeciment
16.6 mm (GCRL 1131), from Area 4
(27°50’N, 85°24'W), 400-485 m, 25 Feb

1988, 2345 CDT; 3 specimens, 16.2, 13.0,
11.3 mm (GCRL 1132), from =Arear
(27°50'N, 85°24’W), 677 m, 26 Feb 1988,
1910 CDT.

Description. — Based on type material with
five stages of sexual maturity described in
Table 2. Body firm, translucent, with well-
developed dorsal and ventral longitudinal
muscle bands; widest at trunk-caudal sep-
tum, tapering slightly toward the head;
maximum length recorded 21.5 mm. Tail
length 21.7-29.3% of total body length.
Head wider than long with distinct constric-
tion at neck. Corona ciliata not observed.
Collarette (alveolar tissue) absent.

Eyes large, bulbous, sub-oval to tear-drop
shaped (Figs. 3C, 5B); widely separated on

VOLUME 102, NUMBER 1 35
Table 1.—Station and hydrographic data for Eukrohnia species in the northern Gulf of Mexico.
: Number of
Bottom Sampling Length Bottom Bottom specimens
depth depth oftow salinity temp.
Station Position Date Time (m) (m) (min) (ppt) (°C) Ec. Ef. Ep.
mgea! 29°02’N, 088°27'W 08/02/87 2007 494 200-400* 33 36.0 8.4 mm (Oe OD
29°00'N, 088°26’W 02/28/88 1930 677 Offa ees hr 36.0 - 6.5 = es 8
Area2 29°18'N, 087°34'W 08/04/87 1910 494 200-400* 44 33:0 “8.60 son 0. 0
29°18'N, 087°32'W 02/13/88" 1035 °*677 (200-500 30 34.0 6.3 a Oo 0
29°18'N, 087°32'W 02/14/88 0730 677 Giizze= 23hr (34.0 6.3 2) SU @)
Area3 29°21'N, 086°57'W 08/06/87 0838 677 200-500 54 39.0 6.4 is 0 0
29°21'N, 086°57’'W 02/16/88 1910 677 200-500 94 5602, 6:7 DEW Nera, ©
29°21'N, 086°57'W 02/17/88 1840 677 Site acne. 36.0 ‘6:7 Te On oO
Area4 27°54'N, 085°16'W 08/08/87 1150 494 200-400 31 58:0) 7 1 0
27°50'N, 085°24'W 02/25/88 2345 494 400-485 27 38.0 8.4 040
27°50'N, 085°24'W 02/26/88 1910 677 CUI WeaMinr: S810". 6:5 37 0). 0
Area5 27°00'N, 084°57'W 02/24/88 0720 677 G71 24 hrs 38:0: 6:6 0 4 1

* Net accidentally dragged bottom.

** Passive sampling by 0.3 m diameter nets attached to crab traps.
Ec. = E. calliops, Ef. = E. fowleri, Ep. = E. proboscidea.

dorsal surface of the head. Pigment cups
lunate to hemispherical, dark-brown to black
in color; oriented parallel to longitudinal
body axis with apex of cup on inner median
edge of eyes.

Apical gland cell complex bilobate, pro-
truding anteriorly. Two less conspicuous
gland cell complexes, a smaller anterior one
and a larger posterior one, both with small,
densely packed papillae, situated anterior to
mouth between vestibular ridges (Fig. 3A).

Hooks 12-13 on each side of head; long,
slender, and somewhat straight, with sharp-
ly bent tips (Fig. 3D, E); no denticulate hooks
observed. Anterior teeth absent. Posterior
teeth 9-21 (17-21 in stage HII and IV in-
dividuals); thin, pointed, set close together
(Fig. 3B); located midway between hooks
and apical gland cell complex. Up to 20
vestibular palps in one or two rows on ves-
tibular ridge, beginning ahead of and run-
ning parallel to entire length of tooth row;
those adjacent to teeth elongate, highly
transparent with cup-like terminal ends (Fig.
3B).

Gut extending from mouth to anus, flat-
tened laterally into thin ribbon, appears im-

pregnated with fine darkened granular ma-
terial; usually with distinctly thickened area
directly beneath ventral ganglion (Fig. 2B).
Intestinal diverticulae absent.

Ventral ganglion located at anterior fourth
of body on ventral surface of trunk; small,
rectangular to oval in shape, with fine nerve
fibers emerging at corners. Ventral trans-
verse musculature extending from neck re-
gion past posterior edge of ventral ganglion
by a distance of up to one-fourth the length
of ganglion (Fig. 2B).

Lateral fins extending along body from
posterior third of tail segment to about an-
terior third of ventral ganglion; broadly
rounded at posterior end and tapering an-
teriorly to delicate ridge; posterior portions
entirely rayed to about level of trunk-caudal
septum, rays continuing anteriorly along
distal edge for about half fin length; inner
and anterior portions containing laminar
material and fine fibrils. Tail fin distinc-
tively large, broadly triangular, completely
rayed.

Ovaries short, club shaped, 10-17 ova per
Ovary; Ova arranged in two rows when
viewed laterally (Fig. 2C). Seminal vesicles

36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

=

=

=
=

\ RWI
AW
WOH 1cehhs
Woy
Ni) |
si
\

|| Gos Bne Reid Neko

¥,

age a _—
AL

hy LOU DUVAA GA eT NLEAR DEL LASALLE WH
OOo ATH HA HN

Fig. 2. Eukrohnia calliops from the northern Gulf
of Mexico. A, ventral view of a mature specimen; B,
lateral view of ventral ganglion area with cut-away
showing thickened area of gut tube underlying trans-
verse musculature; C, lateral view of ovary at maturity
stage III; D, detail of mature seminal vesicle.

oval to elongate, about a third their length
overlapped by posterior edges of lateral fins;
separated from tail fin by about a fourth
their length (Fig. 2D).

Etymology.—The specific name is de-
rived from the Greek words “‘kallos”’ (beau-
ty) and “‘ops”’ (pertaining to the eyes), and
is a noun 1n apposition.

Comparative material examined (Table
3).—

Eukrohnia kitoui, 10 specimens, 4.7—25.4
mm; Tokyo Bay, Japan (35°08'N,
139°46’E), 13 Feb 1981, 0-240 m; from
Kazunori Kuroda, Tokai Regional Fish-
eries Research Laboratory, Tokyo, Japan.

Eukrohnia proboscidea, 2 specimens, one
18.0 mm; origin unknown; from Jean-
Paul Casanova, Laboratorie de Biologie
animal (Plancton), Marseille, France.

Eukrohnia proboscidea, 1 specimen, 10.6
mm; Gulf of Mexico (27°00'N, 84°57'W),
14 Feb 1988, 677 m; from Area 5, R/V
Tommy Munro (Table 1).

Eukrohnia fowleri, 7 specimens co-occur-
ring with E. calliops and E. proboscidea
in the northern Gulf of Mexico (Table 1).

Results and Discussion

Affinities. —Morphological characteris-
tics of the four species of Eukrohnia pos-
sessing eye pigmentation are compared in
Table 4 which combines personal obser-
vations with published data.

Eukrohnia calliops is most closely related
to E. kitoui (Figs. 4B; 5C, D). The two species
are very similar in all respects including eye
pigment configuration. In fact, they were at
first considered synonymous until a detailed
comparison was made between non-con-
tracted E. calliops specimens collected in
February 1988 and material of similar con-
dition from Tokyo Bay, Japan. Results of
this comparison appear in Table 3 and Fig.
6. Eukrohnia calliops has greater numbers
of posterior teeth and higher tail segment
percentages in specimens of comparable size.
Obvious differences between the two species
are eye size and the appearance of the pig-

VOLUME 102, NUMBER 1

a0

Fig. 3. Head details of Eukrohnia calliops, holotype: A, ventral view of head; B, vestibular ridge showing
rows of palps and teeth; C, dorsal view of eyes; D, mature hook; E, hook tip variations. (Abbreviations for Fig.
3A, B: gc = gland cell complex, m = mouth, pt = posterior teeth, vp = vestibular palps.)

ment cup within the eye. The pigment cups
of E. calliops are darker in color, larger in
relation to the total area of the eye, and have
a greater three-dimensional aspect causing
them to “‘bulge’’ from the surface of the head.
The larger eye size of E. calliops is dem-
onstrated in Fig. 6 which compares ratios
of eye length to total body length (EL/TL
x 100) between the two species. Eukrohnia
calliops was found to differ from E. kitoui
at the 95% level of significance using a t-test
on the mean ratios with variances pooled.

Eukrohnia proboscidea (Figs. 4C; 5E, F),
originally described from tropical African

waters by Furnestin & Ducret (1965), is ap-
parently a close relative of E. calliops and
E. kitoui as demonstrated by their morpho-
logical similarities (Table 4) and similar
habitats. As such, the three might be loosely
separated from the rest of the genus to form
the “‘proboscidea group.” This grouping
would further subdivide the “‘fowleri group”
proposed by Casanova (1986:827), an ar-
tificial arrangement of five species of Eu-
krohnia based on the presence of similar
light sensitive cells in the eyes, pigmenta-
tion in the digestive tube, and numbers and
coloration of hooks.

38 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Maturity stages for Eukrohnia calliops in February 1988 collections from the Gulf of Mexico.

Stage TL (mm) Male gonads Female gonads
I 8.4-12.6 Testes are small rods originating in anterio- Ovaries short, club-shaped; no ova
lateral corners of tail segment and not ex- formed but oocyte clusters present.
tending over half the length of the seg-
ment; spermatids not enlarged; seminal
vesicles not formed.

II 11.7-14.4 Testes elongated to full length of caudal seg- § Ovaries longer (4% TL); ova small,
ment; spermatids enlarging; seminal vesi- uniform in size, arranged in 2 rows
cles emerging but not enlarged. or clustered.

II = 15.0 Testes completely fill tail segment; sperm Ovaries longer and thicker (S—7% TL);
cells enlarged; seminal vesicles enlarged, some ova enlarged (3-5 per ovary),
fully formed. arranged in 2 rows.

IV >15.0 Tail segment partially empty; seminal vesi- Ovaries at maximum length (10-11%
cles ruptured. TL); all ova enlarged (10-14 per

Ovary), arranged in 2 rows.
Vv > 18.0 Tail segment empty (spent). Ovaries contracted, empty (spent) or

The pigment of E. proboscidea, which is
situated in the posterior portion of the eye,
differs considerably from that of E. calliops
and E. kitoui. It is light-brown in color with
a distinct black edge along the inner border
which lends a ““U”’-shaped appearance when
viewed dorsally.

An interesting affinity within the “‘pro-
boscidea group”’ is the posterior extent from
the neck of the ventral transverse muscu-
lature. Casanova (1986:831) found that for
E. proboscidea the musculature exceeded the
position of the ventral ganglion by half the
length of the ganglion. He considered this
to be a unique feature of that species and
stated that in all other members of the genus
Eukrohnia, the musculature extends only to
the edge of the ganglion. The overreaching
of the musculature by a fourth the length of
the ganglion in E. calliops, while not as great
an extent as that of E. proboscidea, does
demonstrate a similarity. In the largest spec-
imen of the EF. kitoui material (25.4 mm),
the extent of transverse musculature was
similar to that of E. calliops; however, this
characteristic was reported by Kuroda (1981:
181) as “extending from the neck to the
posterior edge of the ventral ganglion.”

with | or 2 large ova; oviducts swol-
len.

The hooks of the “‘proboscidea group”
species are nearly identical. The hook tips
(Fig. 3E), abruptly deflected inward at 45-
90°, are similar to those of E. hamata and
E. bathypelagica. Denticulate hooks, a com-
mon characteristic in juveniles of Eukroh-
nia and Heterokrohnia (Kapp & Hagen
1985:58), were not found on small speci-
mens of the “proboscidea group” in this
study nor were any reported by Kuroda
(1981) for E. kitoui or by Furnestin (1965)
for E. proboscidea. Owre (1973:957), how-
ever, reported that the ventral-most hooks
on 13.0 and 16.5 mm specimens of E. pro-
boscidea from the Caribbean Sea were “‘ser-
rated with 10-18 small hooks.”

The single specimen of E. proboscidea ob-
tained from the bottom sample at Area 5
represents a new record for the Gulf of Mex-
ico. At 10.6 mm, it is the smallest specimen
yet recorded for the species. Meristic counts
were similar to those reported by Owre
(1973) for a 13.0 mm specimen from the
Caribbean Sea but the Gulf specimen had
fewer number of hooks and teeth than the
smallest specimen (13.3 mm) of the type
material from tropical African waters (Du-
cret 1965). The Gulf of Mexico specimen

VOLUME 102, NUMBER 1

Table 3.—Meristic values of Eukrohnia comparison material.

39

TL mm

% Tail

Hooks

Teeth

Eukrohnia calliops, n. sp. from the Gulf of Mexico

Eye Lmm Mat. St. Source
0.26 IV-—V A3, 200-500 m
0.28 III Al, bottom
0.24 Vv A3, 200-500 m
0.22 III A4, 400-440 m
0.24 IV A4, bottom
O27 IV Al, bottom
0.24 II A2, 200-500 m
0.20 II A4, bottom
0.20 I A2, 200-500 m
0.19 II A2, 200-500 m
NA II A4, bottom
0.18 I A2, 200-500 m
0.14 I A2, 200-500 m

2a 24.9 12 21
18.4 27.0 12 17,
18.2 25.6 12 15-17
16.6 26.6 12 17-19
LG? 24.8 12-13 16
15:0 29:3 NA NA
14.4 2335 13 12
13.0 26.4 12 13
12.6 217 12 12-13
ELT 2555 12 7-11
£13 31.0 13 15
10.4 223 11-12 10
8.4 26.7 11 9-10
Eukrohnia kitoui from Tokyo Bay, Japan (35°08'N, 139°46’E), 0-240 m
25.4 24.6 13 16-17
18.4 222 12 14
16.6 23:6 10-11 14
15.0 DEES) i 13
13.4 22.8 11 9-10
10.4 25.4 10 q)
7.9 D522) 10 5
7.0 253 9 4
5.0 26.2 8 5
4.7 27.4 i 1
Eukrohnia proboscidea
18.0 23.0 12 13
NA NA 12 15
10.6 24.2 10 8
Eukrohnia fowleri from the Gulf of Mexico
28.5 23.0 18} 27
28.5 Des 11-13 27
26.6 22.0 11 20
14.4 2333 11 12
14.0 24.0 1hi@)F 12
11.8 20.0 12 (4)* 10
10.7 20.9 12 (4)* 12

* Denticulate hooks (number).

lacked denticulations on the ventral-most
hooks.

Eukrohnia fowleri (Figs. 4D; 5G, H), a
cosmopolitan, mesoplanktonic species, dif-
fers from the “‘proboscidea”’ group in nearly
all morphological features and armature
meristics. The eye pigment is a dark, vari-
ably shaped fleck in the center of the eye.
The hooks are robust, red in color, and have

0.20 IV Kazunori Kuroda
O17 Ill Kazunori Kuroda
0.14 III Kazunori Kuroda
0.14 II Kazunori Kuroda
On II Kazunori Kuroda
0.10 II Kazunori Kuroda
0.08 I Kazunori Kuroda
0.07 I Kazunori Kuroda
0.06 I Kazunori Kuroda
0.07 I Kazunori Kuroda

II J. P. Casanova

II J. P. Casanova

I A5, Gulf of Mex.

II A5, bottom

II AS, bottom

II A3, 200-500 m

I A3, 200-500 m

I A3, 200-500 m

I A5, bottom

I A5, bottom

straight tips. The ventral-most three or four
hooks of small specimens are strongly den-
ticulate (Table 3).

Ecology and distribution.—Based on avail-
able data, the three species of the “‘probos-
cidea group” appear to occupy similar hab-
itats in deep-water areas adjacent to coastal
regions, although E. proboscidea has so far
been found at greater depths, with occur-

40 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

e

Fig. 4. Dorsal views of heads of the four species of Eukrohnia possessing pigmented eyes. A, E. calliops,
holotype; B, E. kitoui Kuroda, 1981; C, E. proboscidea Furnestin & Ducret, 1965; D, E. fowleri Ritter-Zahony,

1909. Scales: A, B, and D = 0.5 mm; C = 0.2 mm.

rences below 1000 m, off the west African
coast (Ducret 1965) and in samples collect-
ed between 750 and 1100 m in the Carib-
bean Sea (Owre 1973, Michel et al. 1976).
Eukrohnia kitoui and E. calliops were col-
lected at similar depths (0-400 and 200-
677 m) in Tokyo Bay and the northern Gulf
of Mexico, respectively, near undersea can-
yon areas where upwelling of water from
deeper regions occurs.

It is worthy to note that at the Area 5
station (Table 1), no specimens of E. cal-
liops were present, but in the February bot-
tom net samples, one immature specimen
of E. proboscidea was found, indicating the

northward extension of a deep-dwelling Ca-
ribbean population of that species. More ex-
tensive sampling near the bottom in these
areas may identify boundaries between these
two species in the Gulf of Mexico.

In the August 1987 cruise, the nets acci-
dentally dragged bottom at the Area 2 sta-
tion near the Desoto Canyon, resulting in a
sample containing a large number (131) of
E. calliops (Table 1). This suggests a pos-
sible hypo-planktonic distribution for the
species in the Gulf of Mexico.

The genus Evkrohnia was not mentioned
in any of the major studies of the chaeto-
gnath fauna of coastal areas of the Gulf of

VOLUME 102, NUMBER 1

41

Table 4.—Characteristics of the four species of Eukrohnia with pigmented eyes from descriptions published
by Ducret (1965), Alvarino (1967), Kuroda (1981), Michel (1984), and from personal observations by the author.

E. fowleri E. proboscidea Furnestin &
Character Ritter-Zahony, 1909 Ducret, 1965 E. kitoui Kuroda, 1981 E. calliops, n. sp.
Total length 40 25 24.5 PAGS
(mm)
% Tail length 22-25 21.5-31 22-24 24-29
Eye shape Elliptical Elongate Circular to sub- Oval to tear-drop
oval shaped

Eye pigment

Apical gland

Hooks

Posterior teeth

Transverse
musculature
extent

Epidermal col-
larette tissue

Lateral fins

Mature ova-
ries

Seminal vesi-
cles

Variably shaped
rectangular fleck
in center of eye,
black

Single mass of cells,
not protruding

11-13, robust,
strongly curved,
reddish, tips
straight

Up to 30

Level with poste-
rior edge of ven-
tral ganglion

Thick band around
body at ventral
ganglion

Extending from an-
terior 4 of ven-
tral ganglion to
middle of tail
segment

Short (13% TL);
ova large, in two
rows

Ovoid; adjacent to
posterior fins and
separated from
tail fin a distance
greater than their
length

Elongate to
U-shaped in pos-
terior part of eye,
light-brown

Bilobate, protrud-
ing anteriorly

10-13, slender,
gently curved,
light-brown,
transparent, tips
curved sharply
inward

Upto 25

Past ventral gan-
gion by up to '2
length of gan-
glion

Absent

Extending from
base of ventral
ganglion to mid-
dle of tail seg-
ment

Short (15% TL);
Ova in 2 rows,
about 14 per
ovary

Ovoid, elongate;
adjacent to inser-
tion of lateral fins
with anterior '4
overlapped by
posterior edge of
fins, separated
from tail fin by
about 4 their
length

Semi-circular in
center of eye,
dark brown to
black

Bilobate, protrud-
ing anteriorly

8-13, slightly
curved, light-
brown, transpar-
ent, tips curved
sharply inward

Up to 17

Usually level with
posterior edge of
ventral ganglion

Absent

Extending from
slightly anterior
to ventral gan-
glion to posterior
third of tail seg-
ment

Shont(13-5% DE):
Ova in 2 rows,
10-15 per ovary

Ovoid; adjacent to
insertion of later-
al fins with ante-
rior 4 over-
lapped by
posterior edge of
fins, separated
from tail fin by
about '/ their
length

Large, lunate, in cen-
ter of eye, dark
brown to black

Bilobate, protruding
anteriorly

11-13, slightly
curved, light-
brown, transpar-
ent, tips curved
sharply inward

Up to 21

Past ventral ganglion
by up to % length
of ganglion

Absent

Extending from ante-
rior third of ven-
tral ganglion to
posterior third of
tail segment

Short (7.5—10.5%
TL); ova in two
rows, 10-14 per
ovary

Ovoid, elongate; ad-
jacent to insertion
of lateral fins with
anterior '/3 over-
lapped by posterior
edge of fins, sepa-
rated from tail fin
by about '2 their
length

42 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Eyes of the four species of Eukrohnia possessing pigmented eyes. A, C, E, and G show relative
distances between eyes and B, D, F, and H show detail of individual eyes. A and B, E. calliops, holotype; C
and D,-E. kitoui Kuroda, 1981; E and F, E. proboscidea Furnestin & Ducret, 1964; G and H, E. fowleri Ritter-
Zahony, 1909. All scales = 0.1 mm.

VOLUME 102, NUMBER 1

30 O E. kitoui Sa

r= 0.989
y = 139.35 x -3.645

BODY LENGTH mm,

® €. calliops

r= 0.865
y = 79.23 x —2.931

EYE LENGTH mm.

05 -10 -15 -20 25 -30

Fig. 6. Comparative relationships of body length
to eye length for Eukrohnia kitoui and E. calliops.

Mexico (Pierce 1951, 1962; Suarez-Caabro
1955; Vega-Rodriguez 1965; McLelland
1984) because sufficient depths were either
not present or not sampled. Deep-water
studies from areas further removed from
the continental slopes that did include four
species of Eukrohnia are presented in Table
5. Based on personal observations and on
characteristics of mature specimens report-
ed in the literature, the following key sep-
arating the species of Eukrohnia occurring
in the Gulf of Mexico is presented.

43

Key to the Species of Eukrohnia of the

la.
1b.
2a.

2D:

3a.

3b.

4a.

4b.

Sa.

Sb.

Gulf of Mexico

Byes with. plement.) is/ye. se. 2
Eyes non-pigmented
Apical gland bilobate and protrud-
ing, causing head to appear point-
ed. Hook tips bent inward at 45 to
90 degree angles. Transverse mus-
culature extending past posterior
edge of ventral ganglion ........ 3
Apical gland not prominent, a sin-
gle lobed mass. Hook tips straight,
transverse musculature even with
posterior edge of ganglion E. fowleri
Eye pigment small, elongate or
‘““U’’-shaped, in posterior region of
BNE cuterh hertinn Se ai. E. proboscidea
Eye pigment large, lunate, encom-
passing most of the median por-
tion of the eye .... E. calliops, n. sp.
Number of hooks greater than 11,
hook tips straight E. bathyantarctica
Number of hooks less than 11,
heok tips bent mward, | «2. .)..-+). . s
Hooks stout, nearly straight; tail
less than 25% of body length
be (iee abies eet tas lout 9 te, £ E. hamata
Hooks long, gently curved; tail
greater than 25% of body length
manent rite l is 2 1 | E. bathypelagica

Table 5.— Distribution records of the genus Evkrohnia in the Gulf of Mexico.

Species

E. hamata

Mobius, 1875

E. fowleri Ritter-
Zahony, 1909

E. bathyantarctica

Area

Central Gulf

Central Gulf
Central Gulf
SW Gulf

Central Gulf

Position
27°15'N, 029°09'W

27°21'N, 088°42'W
24°39'N, 093°09'W
2 ASN; 095725" W
24°39'N, 093°09'W

David, 1958 Central Gulf 27°08'N, 089°55’W
Central Gulf 26°16'N, 087°52’W
Central Gulf 23°41'N, 090°08’W
SW Gulf 21°55’N, 095°25'W

E. bathypelagica
Alvarino, 1962

Central Gulf
Central Gulf

25°59'N, 086°11'W
25°59’'N, 086°11'W

Sampling
depth
(m) Abundance Source
495-749 Common Pierce (1954)
1100 Common Every (1968)
2000 1 specimen Fagetti (1968)
509-1000 1 specimen Fagetti (1968)
2000 2 specimens Fagetti (1968)
1032-2000 1 specimen Fagetti (1968)
2002-2400 2 specimens Fagetti (1968)
1971-2800 1 specimen Fagetti (1968)
509-1900 2 specimens Fagetti (1968)
580-2000 15 specimens Owre (1973)
580 8 specimens Owre (1973)

44 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Acknowledgments

This work was developed with materials
collected under Grant No. NA87WC-H-
06142, Marine Fisheries Initiative (MAR-
FIN), NMFS, NOAA, U.S. Department of
Commerce. I wish to thank Harriet Perry
for providing the MARFIN plankton col-
lections, Dr. Richard Heard and Sara Le-
croy for their review and suggestions on the
manuscript, and Dr. Robin Overstreet, Mary
Ann Pavlov, and Debbie Jackson for their
help in producing the photographs. All of
the above are from the Gulf Coast Research
Laboratory. Dr. Kazunori Kuroda of the
Tokai Regional Fisheries Laboratory (To-
kyo, Japan) and Dr. Jean-Paul Casanova of
the Laboratorie de Biologie animal-planc-
ton (Marseille, France) kindly provided me
with some of the specimens of Eukrohnia
used in this study.

Literature Cited

Alvarino, A. 1965. Chaetognaths. Pp. 115-194 in

Harold Barnes, ed., Oceanography and marine

biology, annual revue, Vol. 3. George Allen &

Unwin Ltd., London.

1967. The Chaetognatha of the NAGA ex-
pedition (1959-1961) in the South China Sea
and the Gulf of Thailand. Part 1—systemat-
ics. -NAGA Reports, University of California,
Scripps Institute of Oceanography 4(2):1-197.

. 1969. Los quetognatos del Atlantico. Distri-

bucion y notas esenciales de sistematica. Tra-

bajos del Instituto Espanol de Oceanografia 37,

290 pp.

Casanova, J. P. 1986. Deux nouvelles espéces d’Eu-
krohnia (Chaetognathes) de l’Atlantique sud-
tropical africain.— Bulletin du Muséum Natio-
nal d’Histoire Naturelle, Paris (4)8(A no. 4):819-
8355

Ducret, F. 1965. Les espéces du genre Eukrohnia
dans les eaux équatoriales et tropicales afri-
caines.—Cahiers O.R.S.T.O.M.—Océanogra-
phie 3(2):63-78.

Every, M. G. 1968. The taxonomy and areal distri-
bution of the Chaetognatha in the oceanic Gulf
of Mexico. M.S. Thesis, Texas A&M Univer-
sity, College Station, 67 pp.

Fagetti, G. E. 1968. New record of Eukrohnia bathy-

antarctica David, 1958, from the Gulf of Mex-
ico and Caribbean Sea. — Bulletin of Marine Sci-
ence 18(2):383-387.

Furnestin, M. L. 1965. Variations morphologiques
des crochets au cours du développement dans
le genre Eukrohnia.—Revue des Travaux. In-
stitut des Péches Maritimes 29(3):275-284.

—., & F. Ducret. 1965. Eukrohnia proboscidea,
nouvelle espéce de Chaetognathe.— Revue des
Travaux. Institut des Péches Maritimes 29(3):
271-273.

Kapp, H., & W. Hagen. 1985. Two new species of
Heterokrohnia (Chaetognatha) from Antarctic
waters. — Polar Biology 4:53-59.

Kuroda, K. 1981. A new chaetognath, Eukrohnia
kitoui n. sp., from the entrance to Tokyo Bay. —
Publications of the Seto Marine Biological Lab-
oratory 26 (1/3):177-185.

McLelland, J. A. 1984. Observations on chaetognath
distributions in the northeastern Gulf of Mexico
during the summer of 1974.—Northeast Gulf
Science 7(1):49-59.

Michel, H. B. 1984. Chaetognatha of the Caribbean
Sea and adjacent areas. NOAA Technical Re-
port NMFS 15, United States Department of
Commerce, National Technical Information
Service, Springfield, Virginia, 33 pp.

—., M. Foyo, & D. A. Haagensen. 1976. Carib-
bean Zooplankton. Office of Naval Research,
Department of the Navy, U.S. Government
Printing Office, Washington, D.C., 712 pp.

Owre, H. B. 1973. A new chaetognath genus and
species, with remarks on the taxonomy and dis-
tribution of others. — Bulletin of Marine Science
23(4):948-963.

Pierce, E. L. 1951. The Chaetognatha of the west

coast of Florida.— Biological Bulletin (Woods

Hole) 100:206-228.

. 1954. Notes on Chaetognatha of the Gulf of

Mexico.—Fishery Bulletin of the U.S. Fish &

Wildlife Service 55:327-329.

. 1962. Chaetognatha from the Texas coast. —

Publications of the Institute of Marine Science,

University of Texas 8:147-152.

Suarez-Caabro, J. A. 1955. Quetognatos de los mares
Cubanos. — Memorias de la Sociedad Cubana de
Historia Natural 22:125-180.

Vega-Rodriguez, F. 1965. Distribucion de Chaetog-
natha en Veracruz, Ver.—Anales del Instituto
de Biologia, Universidad Nacional Autonoma
de México 36(1/2):229-247.

Gulf Coast Research Laboratory, P.O. Box
7000, Ocean Springs, Mississippi 39564.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 45-49

SMALLEYUS TRICRISTATUS, NEW GENUS,
NEW SPECIES, AND PSEUDOTHELPHUSA PARABELLIANA,
NEW SPECIES (BRACHYURA: PSEUDOTHELPHUSIDAE)
FROM LOS TUXTLAS, VERACRUZ, MEXICO.

Fernando Alvarez

Abstract.—One new genus and species, Smalleyus tricristatus, and a new
species of Pseudothelphusa, P. parabelliana, are described from the Los Tuxtlas
region in the State of Veracruz, Mexico. The new species described here belong
to the Tribe Pseudothelphusini. The distribution patterns of both genera are

discussed.

The Los Tuxtlas region of Veracruz lies
adjacent to the coast of the Gulf of Mexico,
and comprises the area between 18°10’N to
18°40'N and 94°45'W to 95°25'W. This re-
gion encompasses a small mountain range
of volcanic origin, where the highest vol-
canic cone is more than 1700 m high. In
this area four genera and six species of pseu-
dothelphusid crabs co-occur: Odontothel-
phusa maxillipes (Rathbun, 1898), Te-
huana veracruzana (Rodriguez & Smalley,
1969), 7. poglayenorum (Pretzmann, 1980),
T. diabolis (Pretzmann, 1980), Smalleyus
tricristatus, and Pseudothelphusa parabel-
liana (described below). Topographic het-
erogeneity, the presence of tropical rainfor-
est, and the fact that this is an isolated
mountain range situated on the coastal plain,
may account for much of the observed di-
versity. Future collections in southern Ve-
racruz and Tabasco will confirm if, in fact,
pseudothelphusid crabs are absent from
these coastal lowlands. This suggests that
Los Tuxtlas may have served as a refuge in
the past and that its brachyuran fauna has
been preserved. The sites where collections
were made are at altitudes that range be-
tween 200 and 1000 m. All the organisms
are deposited in the Carcinologic Collec-
tion, Instituto de Biologia, Universidad Na-
cional Autonoma de México (IBUNAM).

Smalleyus, new genus

Diagnosis.--In cephalic view, proximal
half of gonopod thinner than distal half. In
mesial view, middle portion, along longi-
tudinal axis, narrower than rest. In caudal
view, lateral process also exhibiting con-
striction in middle portion. In lateral view,
proximal half straight with constant thick-
ness; distal half slightly curved caudally.

Apex of gonopod bearing 3 lobes: mesial,
cephalolateral, and lateral. Mesial lobe with
basal *4 of crest disposed at 90° angle to
longitudinal axis of gonopod; distal portion
sloping proximally; small field of spines
along mesial surface of crest; apical margin
of crest serrate, spine size increasing distal-
ly. In cephalic view, mesial process with
cephalic serrate projection, lying oblique to
longitudinal axis of gonopod. Cephalolater-
al lobe with median ridge in apical cavity,
cephalic blade-like projection oriented
cephalolaterally. Apical median ridge
rounded, bearing spines. Cephalolateral
projection with 3 strong teeth. Lateral lobe
forming lateral wall of apical cavity. Field
of spines along lateral crest and lateral sur-
face; spine size increasing laterally and ce-
phalically. Field of terminal pore setae sit-
uated between median ridge and lateral crest.
Sperm channel opening situated caudal to
median ridge.

46 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Type species. —Smalleyus tricristatus.
Etymology.—This genus is named after
Dr. Alfred E. Smalley. Gender is masculine.

Smalleyus tricristatus, new species
Fig. 1

Material examined. —Camino Izquierdo,
Sierra de Santa Marta, Veracruz (18°26’N,
94°57'W), Jul 1985, collector Mardocheo
Palma: male holotype, c.b. 23.4 mm, c.l.
(5-tmm: 2. females. cbs. 272.2232 ine
c.l. 17.0, 14.9 mm, IBUNAM EM-7030. The
large female had 19 juveniles in the abdo-
men.

Gonopod description. —Same as of genus.

Description.—Carapace slightly convex.
Superior frontal border formed by sloping
of carapace; inclined towards median re-
gion. Inferior frontal border well marked,
formed by diffuse tubercles; sinuous in fron-
tal view, arched in dorsal view. Median
groove short, deep between front and post-
frontal lobes. Cervical grooves straight.
Notch on anterolateral margin between or-
bit and cervical groove. Anterolateral mar-
gin with 17-19 small blunt denticles be-
tween cervical groove and epibranchial
region. Pterygostomian region, around third
maxillipeds, covered with setae. Third max-
illiped with ischium/exopod ratio of 0.71.
Five rows of spines on dactylus of walking
legs. Strong heterochely in males, in females
obvious size differences. Cutting margins of
propodus and dactylus of major chelae with
alternating large and small teeth, when
closed large teeth opposing small ones on
opposite finger.

Type.—The holotypic male is deposited
in the Carcinologic Collection, Instituto de
Biologia, Universidad Nacional Autonoma
de México (Catalog No. EM-7029).

Etymology. — From the Latin “‘tri’”’ (three)
and “crista’’ (crest), referring to the three
crests on the apex of the male’s gonopod.

Remarks. —Smalleyus belongs to the
Tribe Pseudothelphusini because it exhibits
a curved sperm channel, a reniform mesial

process, and a discernible lateral lobe. A
new genus is created due to the presence of:
a mesial process whose shape is not com-
pletely reniform as in Pseudothelphusa or
Tehuana and bearing spines along its su-
perior margin; a cephalic serrate projection
of the mesial process; three ridges (crests)
on the apex of the gonopod, the median one
partly filling what would otherwise be the
apex cavity; a cephalolateral lobe, which
seems to be a continuation of the median
apical ridge; a lateral lobe, which corre-
sponds to the lateral lobe in the genus Pseu-
dothelphusa, but in this crab occupying a
more caudal position; and three spine fields,
one on each one of the three apical ridges.
The relationships of this genus with the oth-
er genera of the tribe are not clear. However,
this aberrant form, together with Spirothel-
phusa, Lobithelphusa, Odontothelphusa, and
Disparithelphusa constitute a belt of mono-
typic genera dispersed along the Isthmus of
Tehuantepec. Even though these taxa be-
long to three different tribes, the factors in-
fluencing the origin of these five genera may
be similar. In this case, the isthmus repre-
sents a barrier that these crabs could not
cross (Rodriguez 1987). Epithelphusa with
two species, occurring in the same area, has
more distinct affinities. Rodriguez (1982)
places the genus linking the tribes Hypo-
lobocerini and Pseudothelphusini. The af-
finities of the monotypic genera mentioned
are obscure, with the exception of Odon-
tothelphusa which derives from the genus
Potamocarcinus.

Pseudothelphusa parabelliana, new species
Fig.’ 2

Material examined. —Cerro El Vigia, Los
Tuxtlas Biological Station, Veracruz
(18°35'N, 95°03’W), 7 Jul 1986: male ho-
lotype, c.b. 24.5 mm, c.l. 15.8 mm; 1 male
paratype, c.b. 21.4 mm, c.l._13-0 sami
females, c.b..34.1, 19.2,.c.l. 21.3, Pi Seana
IBUNAM EM-7031. Same locality, 30 Aug
1984, collector Mardocheo Palma: 1 male

VOLUME 102, NUMBER 1 47

Fig. 1. Left gonopod and major chela of Smalleyus tricristatus: a, Total mesial view; b, Cephalic view of
apex; c, Lateral view of apex; d, Caudal view of apex; e, Apical view; f, Right chela.

paratype, c.b. 23.9 mm, c.l. 14.8, IBUNAM in abdomen, c.b. 28.0 mm, c.l. 16.9 mm,
EM-7032; 5 females c.b. 32.3, 29.2, 27.7, IBUNAM EM-7034.

Ue) 20.9 mim, cl. 18.8. 17.6, 16.8, 14.0, Description. —Carapace slightly convex.
13.6 mm, IBUNAM EM-7033. Same lo- Superior frontal border absent, but front
cality, 7 Jul 1986, 1 female with 8 juveniles limited by edge. Front inclined in dorsal

48 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Left gonopod of Pseudothelphusa parabelliana: a, Total mesial view; b, Cephalic view of apex; c,
Lateral view of apex; d, Caudal view of apex.

view. Inferior frontal border smooth, some- frontal lobes. Cervical grooves shallow,
what bilobed in dorsal and frontal views. wide, straight; not reaching anterolateral
Median groove narrow, deep, dividing su- margin. Anterolateral margin with slender
perior edge of front and separating post- notch lateral to orbit. Anterolateral margin

VOLUME 102, NUMBER 1

with 16-19 denticles between cervical
groove and epibranchial region. Ratio is-
chium/exopod of third maxilliped 0.91.
Pterygostomian region covered with setae.
Major chelae right, fingers not gaping.

Gonopod description.—Well developed
reniform mesial process. Marginal process
reduced to acute spine. Mesial crest round-
ed. Lateral lobe completely closing apical
cavity cephalically and ending in 2 tips dis-
tally; inferior sharper than superior. Lateral
lobe with mesial surface triangular. Ter-
minal pore setae 19. Setal field restricted to
lateral portion of cavity. Mesial crest higher
than lateral. Opening of sperm channel cau-
dal.

Type.—The holotypic male is deposited
in the Carcinologic Collection, Instituto de
Biologia, Universidad Nacional Autonoma
de México (Catalog No. EM-7028).

Etymology. —From the Latin “‘para’’ (be-
side) and “‘belliana”’ a freshwater crab spe-
cific name.

Remarks.—Before this species was de-
scribed the genus Pseudothelphusa was
known only from the central part and Pa-
cific slope of Mexico. The presence of P.
parabelliana now widens this distribution
to the 95°W meridian. This species seems
to be closely related to P. belliana Rathbun,
1898, which occurs in the states of Guer-
rero, Michoacan, and México. The gonopod
of P. parabelliana has a lateral lobe with
two acute distal tips, a straight mesial crest,
and the mesial process is much larger in
relation to the marginal process. In P. bel-
liana the lateral lobe bears only one distal
tip, the mesial crest is broadly rounded, and
the mesial process is slightly larger than the

49

marginal process. The occurrence of P.
parabelliana in the Gulf coast of Mexico
agrees well with the morphocline proposed
recently by Rodriguez (1987). He places on
a west-east axis, across central Mexico, a
succession of species (including Tehuana
veracruzana, P. guerreroensis Rathbun,
1933, P. belliana, P. americana de Saussure,
1857, and P. peyotensis Rodriguez & Smal-
ley, 1969) in which a progressive reduction
of the lateral lobe can be seen.

Acknowledgments

I wish to thank Mardocheo Palma for col-
lecting most of the crabs included in this
study, and Dr. Alfred E. Smalley for pro-
viding much helpful advice and reviewing
the manuscript. This study was supported
by a grant from the Mesoamerican Ecology
Institute of Tulane University.

Literature Cited

Pretzmann, G. 1980. Von Dr. Ivo Poglayen-Neuwall
1975 in Mittelamerika gesammelte Krabben. —
Annalen Naturhistorisches Museum Wien 83:
651-666.

Rodriguez, G. 1982. Les crabes d’eau douce d’Amé-

rique. Famille des Pseudothelphusidae. — Faune

Tropicale 22:1-223.

. 1987. Centers of radiation of freshwater crabs

in the Neotropics. Pp. 51-67 in R. H. Gore &

K. L. Heck, eds., Crustacean issues 3: Biogeog-

raphy of the crustacea. A. A. Balkema, Rotter-

dam.

Department of Biology, Tulane Univer-
sity, New Orleans, Louisiana, 70118. Pres-
ent address: Department of Zoology, Uni-
versity of Maryland, College Park, Maryland
20742.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 50-77

TWO NEW GENERA AND NINE NEW SPECIES OF
GERYONID CRABS
(CRUSTACEA, DECAPODA, GERYONIDAE)

Raymond B. Manning and L. B. Holthuis

Abstract. —The family Geryonidae Colosi, 1923 is restricted to three genera:
Geryon Kroyer, 1837, containing two species, G. trispinosus (Herbst, 1803),
and G. longipes A. Milne Edwrds, 1882; Chaceon, new genus, containing 21
species, eight of which are newly described; and Zariquieyon, new genus, con-
taining one new Mediterranean species, Z. inflatus. The new species of Chaceon
and their ranges are: C. atopus, Saint Helena Island; C. bicolor, central Pacific
Ocean; C. crosnieri, Madagascar; C. eldorado, northern South America; C.
inglei, northeastern Atlantic; C. mediterraneus, western Mediterranean Sea; C.
notialis, Uruguay and Argentina; and C. sanctaehelenae, Saint Helena Island.

In addition to results published elsewhere
(see Manning & Holthuis 1984, 1986, 1987,
1988), our studies on deep-sea crabs of the
family Geryonidae have revealed the exis-
tence of nine other undescribed species from
various localities that can be assigned to two
new genera. These taxa are named herein.

Specimens have been deposited in the
British Museum (Natural History), London
(BMNH), the Muséum National d’Histoire
Naturelle, Paris (MNHN), the Rijksmu-
seum van Natuurlijke Historie, Leiden
(RMNH), the National Museum of Natural
History, Smithsonian Institution, Washing-
ton (USNM), and the Zoological Museum,
Copenhagen (ZMC).

We use the following abbreviations: cb,
carapace breadth, measured between the tips
of the posterior pair of anterolateral teeth;
cl, carapace length, measured on the mid-
line; fm, fathoms; leg, collector; m, meter(s);
mm, millimeter(s); P5, fifth leg (fourth
walking leg); sta, station.

Family Geryonidae Colosi, 1923
Geryonidae Colosi, 1923:249.

Diagnosis. —Carapace hexagonal, with
three or five anterolateral teeth on each side.

Suborbital margin with inner tooth only.
Chelipeds unequal, merus with subdistal
dorsal spine, carpus with strong inner spine.
Chelae portunid-like. Walking legs with na-
ked dactyli, lacking segments expanded for
swimming. Abdomens of male and female
with seven free.segments.

Remarks. —The status of the family Ger-
yonidae and of the genera assigned to it have
long posed problems for students of the
group, as we already have pointed out
(Manning & Holthuis 1981:109). That the
family Geryonidae deserves recognition as
a family distinct from the Xanthidae s.]. and
the Goneplacidae s.l. and that its affinities
are with the Portunidae no longer seem to
be in question.

Several genera of uncertain affinities have
been placed in the family Geryonidae (see
Guinot 1971:1077—1078), e.g., Paragalene
Kossmann, 1878, Bathyplax A. Milne Ed-
wards, 1880, Platypilumnus Alcock, 1894,
Progeryon Bouvier, 1922, and Platychelo-
nion Crosnier & Guinot, 1969.

In spite of Guinot’s (1969:692) observa-
tion, “Les deux genres monospécifiques
Progeryon and Platypilumnus nous parais-
sent étroitement apparentés a Geryon, in-
séparables de celui-ci,’’ we do not believe

VOLUME 102, NUMBER 1

that these or any of the genera other than
Geryon and the two new genera established
here belong in the Geryonidae. We believe
that for the moment all other Recent genera
formerly assigned to the Geryonidae should
be placed in the Xanthidae s.l. Several of
them, including Paragalene and Progeryon,
may well belong in a new family.

Members of all of the genera here ex-
cluded from the Geryonidae lack the por-
tunid-like chelipeds and the characteristic
subdistal dorsal spine on the merus of the
chelipeds found in the geryonids proper as
well as in some portunids, and they all can
be distinguished from geryonids as restrict-
ed here by these features alone. Further, all
of the geryonids as restricted here have na-
ked dactyli on the walking legs and lack black
fingers on the chelae, other features that will
distinguish them from members of Para-
galene and Progeryon.

As restricted here, the Geryonidae com-
prise three genera and 24 species: Geryon
Krgyer, with 2 species; Chaceon, new genus,
with 21 species; and Zariquieyon, new ge-
nus, with | species.

Geryon Kroyer, 1837

Geaven, Kroyer, 1837:10, 20, 21 [type
species, by original designation and
monotypy, Geryon tridens Kroyer, 1837,
a subjective junior synonym of Cancer
trispinosus Herbst, 1803].

Chalaepus Gerstaecker, 1856:118 [type
species, by monotypy, Cancer trispinosus
Herbst, 1803].

Definition. —Geryonid crabs with three
anterolateral teeth on each side of the car-
apace. Carapace length half to two-thirds
width. Branchial regions not markedly in-
flated. Frontal teeth poorly developed, small.
Orbits shallow, rounded.

Included species. —Two, both from the
northeastern Atlantic Ocean:

Geryon longipes A. Milne Edwards (1882:

FG:,39);
Geryon trispinosus (Herbst) (1803:43)

51

(=Geryon tridens Kre@yer, 1837, see Man-
ning & Holthuis 1987).

Remarks. —Geryon differs from both
Chaceon and Zariquieyon in having three
rather than five anterolateral teeth on each
side of the carapace, and the two submedian
frontal teeth are smaller than in the repre-
sentatives of either of the other two genera.
Geryon agrees with Chaceon and differs from
Zariquieyon in having the orbits shallow and
rounded rather than rectangular, and Ger-
yon and Chaceon lack the swollen branchial
regions that are characteristic of the only
known species of Zariquieyon.

Chaceon, new genus

Type species.—Geryon fenneri Manning
& Holthuis, 1984.

Name.—The name is given in honor of
our colleague Fenner A. Chace, Jr. The suf-
fix -on is used to make the name similar to
Geryon. This sufhx should not be consid-
ered to be the Greek neuter ending -on. The
gender of the name Chaceon is masculine,
like that of Geryon.

Definition. —Geryonid crabs with five an-
terolateral teeth on each side of the cara-
pace. Carapace length half to two-thirds
width. Branchial regions not markedly in-
flated. Frontal teeth well-developed, large.
Orbits shallow, rounded.

Included species. —Twenty-one, of which
eight are described herein. Nominal species
include the following:

Chaceon affinis (A. Milne Edwards & Bou-
vier, 1894:41), northeastern Atlantic.
Chaceon chuni (Macpherson, 1983:23), Na-

mibia and South Africa.

Chaceon erytheiae (Macpherson, 1984:86),
Valdivia Bank, southeastern Atlantic.
Chaceon fenneri (Manning & Holthuis,

1984:666), northwestern Atlantic.
Chaceon gordonae (Ingle, 1985:90), West
Africa.
Chaceon granulatus (Sakai, 1978:11), Ja-
pan.

52 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Chaceon inghami (Manning & Holthuis,
1986:367), Bermuda.

Chaceon macphersoni (Manning & Hol-
thuis, 1988:83), southwestern Indian
Ocean and South Africa.

Chaceon maritae (Manning & Holthuis,
1981:112), West Africa.

Chaceon atopus, male holotype, cb 124 mm, Saint Helena, dorsal view.

Fig. 1.

Chaceon paulensis (Chun, 1903:531), cen-
tral Indian Ocean.

Chaceon quinquedens (Smith, 1879:35),
northwestern Atlantic.

Two other species originally described in
Geryon, Geryon incertus Miers, 1886, and

VOLUME 102, NUMBER 1

Geryon ischurodous Stebbing, 1923, were
incorrectly assigned to the genus. Manning
& Holthuis (1986:369) showed that G. in-
certus is a synonym of the portunid Bathy-
nectes longispina Stimpson, 1871, and
Manning & Holthuis (1988:78) showed that
G. ischurodous is a species of the goneplacid
genus Carcinoplax.

Remarks. — Accounts of two other species,
one from Chile, and one from Brazil are in
preparation. These 2 species, the 11 known
species, and the 8 new species described be-
low raise the known number of species to
2

Members of Chaceon agree with Zari-
quieyon and differ from Geryon in having
five anterolateral teeth on each side of the
carapace.

Chaceon atopus, new species
Figs. 1-2

Material. —Off Rupert’s Bay, Saint He-
lena Island [15°58’S, 5°43'W], 130 fm (238
m), 5 Aug 1983, leg A. J. Edwards: one male
(holotype, RMNH).

Diagnosis. —A large Chaceon, cl 90 mm,
cb more than 120 mm in only known spec-
imen, with well-developed anterolateral
teeth on the carapace in adults and with
laterally compressed dactyli on the walking
legs. Carapace 1.4 times broader than long,
very inflated, strongly convex from front to
back, appearing smooth. Median pair of
frontal teeth long and sharp, separated by a
narrow, V-shaped emargination, medians
extending further forward than laterals; out-
er margin of lateral frontal tooth with slight
convex projection, indicating position of in-
ner orbital angle. Anterolateral teeth well-
developed, all sharp, fourth smallest but
sharp and distinct. Distance from first to
second tooth about the same as from third
to fourth, distance from first to third tooth
less than distance between third to fifth.
Carapace surface finely granular postero-
laterally. Suborbital tooth strong and sharp-
ly pointed, visible in dorsal view, extending
about to level of lateral frontal tooth; sub-
orbital margin evenly curved, tuberculate.

53

Fig.2. Chaceon atopus, male holotype, cb 124 mm,
Saint Helena: a, Ventral view of orbit; b, P5 dactylus,
posterior view; c, P5 dactylus, dorsal view.

Cheliped merus with sharp spine subdistally
and with distal dorsal spine; carpus rough-
ened dorsally, with distal outer spine, den-
ticulate anterior margin, and strong and
slender distal spine; propodus with distal
dorsal spine broken in left chela, right chela
with distal angled projection. Meri of walk-
ing legs with distinct distal dorsal spine.
Dactyli of walking legs laterally com-
pressed, height at midlength greater than
width. P5: merus 6.5 times longer than high,
length 0.58 cb; carpus with line of erect spi-
nules dorsally; propodus longer than dac-
tylus, length 5.8 times height.

Size. —Unique male holotype, cl 90 mm,
cb 124 mm.

Remarks. —This species resembles C.
gordonae, from the Cape Verde Islands and
Sierra Leone, in many features, including
the large size, the well-developed frontal and
anterolateral teeth on the carapace, and the
compressed dactyli of the walking legs. It
differs from C. gordonae in having much

54 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Chaceon bicolor, male paratype, cb 158 mm, Passe de Saint Vincent: a, Dorsal view; 6; Carapace.

VOLUME 102, NUMBER 1

longer walking legs, with the merus more
than 6 times longer than high, and in its
habitat. Chaceon gordonae, in which the P5
merus is less than 6 times longer than high,
generally occurs in depths in excess of 2000
m, whereas the present new species occurs
in less than 250 m.

This species can be distinguished on sight
from C. sanctaehelenae, described below,
by the long, slender walking legs, the distal
meral spine on the walking legs, and the

well-developed anterolateral spines of the.

carapace.
Name.—The specific name is from the
Greek, atopos, out of place, strange, our re-
action when we learned this. specimen was
specifically distinct from the holotype of C.
sanctaehelenae, described below.

Distribution.—Known only from Saint.

Helena Island, in 238 m. -

Chaceon bicolor, new species
Figs. 3-4

Geryon affinis. —Griffiin & Brown, 1976:256,
figs. 7-9.—Sakai, 1978:9, figs. 18-19, pl.
2, fig. D (color). [Not Geryon affinis A.
Milne Edwards & Bouvier, 1894.]

Geryon quinquedens.—Intés, 1978:7, figs.
5B, 8.— Guinot & Richer de Forges, 1981:
249.—King, 1984:186. [Not Geryon
quinquedens Smith, 1879.]

Geryon.—Intés, 1978: fig. 10.—Guinot &
Richer de Forges, 1981:249.

Previous _records.—Emperor Seamount

Chain: North of Nintoku Seamount,
42°20'N, 170°50’E, 800 m; Jingu Seamount
[38°50’N, 171°15’E], 890-930 m; Kinmei
Seamount, 35°34’N, 171°41’E, 600-640 m,
and 500-700 m; south of Kinmei Sea-
mount, 34°42’N, 171°48’E, 980-1100 m
(Sakai 1978).

Southwestern Pacific: in depths greater
than 600 m (King 1984).

Loyalty Islands [21°00'S, 167°00’E]: (Intés
1978).

New Caledonia [21°30’S, 165°30’E]: 600-

55

Fig.4. Chaceon bicolor, male paratype, cb 148 mm,
Passe de Saint Vincent: a, Ventral view of orbit; b, P5
dactylus, posterior view; c, P5 dactylus, dorsal view.

1000 m (Intés 1978; Guinot & Richter de
Forges 1981).
New South Wales, Australia: Southeast

- of Grafton, 29°49’S, 153°42’E to 29°59’S,

153°38’E, 369 m; east of Broken Bay,
33°40’S,.151°53’E to 33°22’S, 152°09’E, 450
m=33°40'S,. 151°53’E 10 33°35’S, 151°58’E,
450-810 m, and 33°38’S, 151°57’E to
33°34 891152701 E)..774—792) m;: north of

Sydney, 33°43’S, 151°55’E to 33°37’S,

1:52°02'E)}. 675:-m;.. off Sydney, 33°52'S,
152°50’'E to 33°48'S, 152°54’E, 765 m (Grif-
fin & Brown 1976).

Material. —Emperor Seamount Chain:
Kinmei Seamount, 35°17.50'N, 171°25.98’E,
150 fm (275 m), Townsend Cromwell cruise

56 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Cromwell cruise 82-05, sta 57, 11 Nov 1982:
three females (USNM 205974).

New Caledonia: 23°44’S, 166°58’E, 1490-
1620 m, BIOCAL, Jean Charcot sta CP 57,
1 Sep 1985: one juvenile male, one female
(MNHN), one female (USNM 205975).—
23°07'S, 166°51’E, BIOCAL, Jean Charcot
sta CP 32, 825 m, 29 Aug 1985: one male
(MNHN).—Passe de Saint Vincent [22°02’S,
165°57’'E], 800 m, traps, leg B. Richer de
Forges: four males (3 MNHN, 1 USNM
205976).

Australia: Southeast of Newcastle, New
South Wales, 33°11—09’S, 152°24—25’E, 732
m, leg D. E. Brown on FRV Kapala, sta
K77-23-10, 7 Dec 1977: one male (USNM
205978).—East of Broken Bay, New South
Wales, 33°35—33’'S, 152°00-02’E, 823 m, leg
D. E. Brown on FRV Kapala, sta K77-23-
12, 8 Dec 1977: one male, one female
(USNM 205977).

The holotype is the male, cl 144 mm, cb
165 mm, from New Caledonia, Jean Char-
cot sta CP 32 (MNHN). All of the other
specimens are paratypes.

Diagnosis. —A very large Chaceon, cl to
more than 140 mm, cb to 180 mm in adults,
with small, blunt anterolateral teeth on the
carapace in adults and with laterally-com-
pressed dactyli on the walking legs. Cara-
pace 1.1-1.2 times broader than long in
adults, 1.4 times broader than long in ju-
veniles. Median pair of frontal teeth nar-
rower than laterals, separated by U-shaped
emargination, medians extending further
forward than laterals. Second and fourth an-
terolateral teeth reduced, fourth often ob-
solete. Distance from first to second an-
terolateral tooth less than that from third
to fourth, distance from first to third sub-
equal to that from third to fifth. Carapace
with distinct granulation mesial to fifth tooth
and on protogastric, cardiac, and branchial
regions; hepatic region smooth; protogastric
region inflated in large specimens, especially
in females. Suborbital tooth short and blunt
in males, longer and sharper in females and
juveniles, scarcely or not at all visible in

dorsal view, suborbital margin evenly
curved, tuberculate. Cheliped lightly tuber-
culate dorsally; upper margin of merus with
sharp subdistal spine; carpus lacking outer
spine in adult, blunt projection present in
some specimens, with distinct spine in ju-
venile; propodus unarmed distally. Meri of
posterior walking legs with distinct distal
dorsal angled projection, spined in juve-
niles. Dactyli of walking legs laterally com-
pressed, height at midlength greater than
width. P5: merus 4.3—5.1 (mean 4.6) times
longer than high in females, 5.2—5.6 (mean
5.4) times longer than high in males, and
7.0—7.5 times longer than high in juveniles,
usually with blunt distal dorsal projection
in adults, distinct distal spine in juveniles;
merus length 0.49-0.56 cb in females, 0.61-
0.64 cb in males; carpus smoothly tuber-
culate dorsally; propodus distinctly longer
than dactylus, 3.6-4.6 (mean 4.1) times
longer than high in adults, 5.6-6.0 times
longer than high in juveniles.

Size. — Males, cl 116-147 mm, cb 134—
165 mm; females, cl 21-110 mm, cb 30-
122.5 mm; juvenile, cl 7 mm, cb 10 mm.
The largest specimens recorded in the lit-
erature are: male, cl 141 mm, cb 155 mm
(Sakai 1978); a specimen with cb 176 mm
(Griffin & Brown 1976); a specimen with cb
180 mm (Intés 1978).

Color.—Anterior part of carapace pre-
dominantly purple, branchial regions tan,
legs yellowish. Sakai (1978: pl. 2, fig. D)
gave a figure in color.

Remarks. —Chaceon bicolor differs from
all other species of the genus in color pat-
tern, with the anterior part of the body pur-
plish rather than reddish or tan, set off from
the tan branchial regions. Remnants of this
distinctive color pattern are visible in one
of the juveniles from deep water off New
Caledonia. In addition to color pattern, C.
bicolor also differs from the only other
species known to occur in the central Pa-
cific, C. granulatus, in having compressed
rather than depressed dactyli on the walking
legs; also, the hepatic region of the carapace

VOLUME 102, NUMBER 1

in C. granulatus is coarsely granular, where-
as it is smooth in C. bicolor.

The juvenile specimens taken in 1490-
1620 m off New Caledonia differ from the
adults in many features: the teeth of the
carapace are much larger and sharper, there
is a sharp outer spine on the carpus of the
cheliped and a sharp distal spine on the me-
rus of each walking leg, and the legs are
much longer and slenderer. As mentioned
above, one specimen shows traces of the
typical color pattern of the adults, suggest-
ing that these small specimens are juveniles
that occur in much deeper water than do
the adults.

Adult females differ from males in having
much sharper anterolateral teeth on the car-
apace, sharper suborbital spines, and much
shorter legs, with less trace of a distal dorsal
projection on the merus. Also, the carapace
of females is more strongly arched from front
to back and the protogastric regions are no-
ticeably more inflated.

Our specimens were taken in depths of
275, 732, 800, 823, 825, and 1490-1620 m;
the juveniles were collected at the latter
depth. Depth records in the literature in-
clude: 369 to 774-792 m (Grifin & Brown
1976); 500-700 to 980-1 100 m (Sakai 1978);
600-1000 m (Intés 1978, Guinot & Richer
de Forges 1981); and in depths greater than
600 m (King 1984).

Name.—The specific name is from the
Latin, the prefix bi- and color, referring to
the purple and tan color pattern of this
species in life.

Distribution. —Widely distributed in the
central Pacific, from the Emperor Seamount
Chain (42°20'N) to eastern Australia, off
Sydney (33°52’S), in depths between 275 and
about 1600 (1490-1620) m.

Chaceon crosnieri, new species
Figs. 5-6

Material.— Madagascar: 22°21'S, 43°05.5’E,
450-420 m, sta 27, 15 Jan 1986: two males,
three females (MNHN).—22°13.5’S,

57

43°05’E, 530 m, mud, chalutage 62, leg Rudo
von Cosel, 19 Oct 1986: one female
(MNHN).—22°26.1'S, 43°04.6’E, 520 m,
mud, chalutage 65, leg Rudo von Cosel,
1986: one male, four females (MNHN).—
PPV S45 02.5 FE, 560 m, mud, chalu-
tage 75, leg Rudo von Cosel, 23 Oct 1986:
two females (MNHN).—22°24.7’S,
43°04.2’E, 535 m, mud, chalutage 91, leg
Rudo von Cosel, 4 Nov 1986: one male,
two females (USNM 205979).—22°24.6’'S,
43°03.7’E, 590 m, mud, chalutage 93, leg
Rudo von Cosel, 7 Nov 1986: one male,
one female (MNHN).—22°S, 43°E, 790 m,
mud, chalutage 102, leg Rudo von Cosel,
24 Nov 1986: one male, one female
(MNHN).—22°24.7'S, 43°03.7’E, 630 m,
mud, chalutage 115, leg Rudo von Cosel,
28 Nov 1986: one female (MNHN).

The holotype is the male, cl 116 mm, cb
140 mm, from chalutage 93; all other spec-
imens are paratypes.

Diagnosis.—A very large Chaceon, cl to
127 mm, cb to 155 mm in adults, with low,
blunt anterolateral teeth on the carapace in
adults and with laterally compressed dactyli
on the walking legs. Carapace 1.2 times
broader than long, very inflated dorsally,
especially at protogastric regions Median
pair of frontal teeth narrower than laterals,
not set ahead of laterals, both pairs extend-
ing about to same level. All five anterolat-
eral teeth reduced, distance from first to sec-
ond equal to distance from third to fourth,
distance from first to third equal to distance
from third to fifth. Carapace relatively
smooth, with small granules and pits on
branchial, cardiac, and gastric regions, he-
patic regions smooth. Suborbital tooth sharp
in females, very short and blunt in males,
scarcely visible in dorsal view, suborbital
margin concave, tuberculate. Cheliped
lightly granular, not conspicuously rough-
ened, merus with sharp subdistal spine; car-
pus irregular dorsally, with sharp inner
tooth, lacking any trace of outer tooth; prop-
odus with some small tubercles dorsally,
lacking distal spine. Meri of walking legs,

Fig. 5. Chaceon crosnieri, male holotype, cb 140 mm, Madagascar: a, Dorsal view; b, Carapace. Female
paratype, cb i27 mm, Madagascar: c, Carapace, lateral view.

VOLUME 102, NUMBER 1

especially posterior ones, with distal, dorsal
spine. Dactyli of walking legs laterally com-
pressed, height at midlength greater than
width. PS: merus 4.1—5.0 (mean 4.7 in males,
4.5 in females) times longer than high, with
distal dorsal spine, length 0.47-0.55 cb
(mean 0.51) in females, 0.57-0.63 cb (mean
0.59) in males; carpus lacking erect spinules
dorsally; propodus length 3.6-4.4 times
height (mean 3.9 in males, 4.0 in females),
usually longer than dactylus, subequal to
dactylus in large females.

Size. —Males, cl 83-127 mm, cb 101-155
mm; females, cl 82-127 mm, cb 97-148
mm.

Color.—In preservative, some males show
tips of fingers of chela with red band proxi-
mally, apices whitish. Color in life un-
known.

Remarks. —Chaceon crosnieri differs from
C. affinis in that the carapace is much more
inflated, the outer orbital and the suborbital
teeth are stronger, the subdistal tooth on the
merus of the cheliped is stronger and the
cheliped is smoother dorsally, the carpus of
the walking legs lacks dorsal spinules, and
the merus of the walking legs, especially the
posterior ones, has a strong distal dorsal
spine. This new species differs from C. chuni
in being much larger, cb to 155 mm, the
carapace is much more inflated, especially
at the protogastric region, the gap between
the first and second anterolateral tooth of
the carapace is larger (equal to distance from
third to fourth in C. crosnieri, less in C.
chuni), the frontal teeth of the carapace are
stronger, the suborbital spine is smaller, and
the carpus of the cheliped lacks an outer
spine.

Chaceon crosnieri is a much smoother
species than C. bicolor, with much shorter,
stouter legs, the suborbital spine is much
lower and blunter, and the distal projection
on the merus of the walking legs is much
less developed in larger specimens.

A male of C. paulensis in the USNM, cl
so mm), ‘cb 113 mm; ‘from: 38°24.92'S,
77°25.15'E, in 1050-1110 m off Amster-
dam Island in the southern Indian Ocean,

a9

Fig. 6. Chaceon crosnieri, male paratype, cb 155
mm, Madagascar: a, Ventral view of orbit; b, PS dac-
tylus, posterior view; c, P5 dactylus, dorsal view.

is available for comparison. It shares the
compressed dactyli of the walking legs with
C. crosnieri, and little else. It differs in many
features, especially in having strongly de-
veloped, sharp anterolateral teeth on the
carapace and on the meri of the walking legs,
and in having a large, sharp outer spine on
the carpus of the cheliped.

Chaceon crosnieri was taken together with
C. macphersoni at sta 27 and at chalutages
62, 65, 91, and 93. The two species can be
distinguished immediately, as C. macpher-
soni is much rougher dorsally and has the
dactyli of the walking legs depressed, not
laterally compressed.

Our specimens were taken in depths of
420-450 to 790 m, with five samples being
taken in depths between 520 and 590 m.

Name.—We are pleased to dedicate this

60

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig..7:

Chaceon eldorado, male holotype, cb 109 mm, Venezuela: a, Dorsal view; b, Carapace.

VOLUME 102, NUMBER 1

species to our colleague Alain Crosnier, Mu-
séum National d’Histoire Naturelle, Paris,
who made special efforts to save specimens
of geryonids whenever they were encoun-
tered in the field.

Distribution. —Known only from locali-
ties around Madagascar, in depths between
420-450 and 790 m.

Chaceon eldorado, new species
Figs. 7-8

Geryon quinquedens. — Takeda, 1983:15, 18,
31, 164, color fig. on p. 164. [Not Geryon
quinquedens Smith, 1879.]

Previous records. —Off Suriname and
French Guiana, 310-790 m (Takeda 1983).

Material.—Colombia: 12°06'N, 72°55'W,
350-500 fm (641-915 m), Oregon sta 4912,
31 May 1964: one male (USNM 205980).

Venezuela: 12°55'N, 70°16’W, 340 fm
(622 m), Oregon IT sta 11307, 26 Nov 1970:
one female (USNM 205981).—11°53'N,
69°25'W, 350 fm (641 m), Oregon sta 4413,
3 Oct 1963: one male (USNM 205982).—
11°36'N, 62°46'W, 290 fm (531 m), Oregon
sta 2777, 19 Apr 1960: two males (USNM
205983).

French Guiana: 7°37'N, 53°32’'W, 395 fm
(723 m), Oregon IT sta 10616, 13 May 1969:
one male (USNM 205984).

The male, cl 89.5 mm, cb 109 mm, from
Oregon sta 4413, is the holotype (USNM
205982); the other specimens are paratypes.

Diagnosis. —A moderately large Cha-
ceon, cl to 89.5 mm, cb to 109 mm in ma-
terial examined, with low, blunt anterolat-
eral teeth on the carapace in adults and with
dorsoventrally depressed dactyli on the
walking legs. Carapace 1.2—1.3 times broad-
er than long, inflated dorsally, especially at
protogastric regions. Median pair of frontal
teeth about as wide as and extending farther
forward than laterals. Second and fourth an-
terolateral teeth usually reduced, much
smaller and lower than remainder. Distance
from first to second tooth equal to or less

61

Fig. 8. Chaceon eldorado, male holotype, cb 109
mm: a, Ventral view of orbit; 6, PS dactylus, posterior
view; c, P5 dactylus, dorsal view.

than that from third to fourth tooth, dis-
tance from first to third tooth less than that
from third to fifth. Carapace granular pos-
terolaterally and on cardiac and protogastric
regions, hepatic region smooth. Suborbital
tooth short, broad, visible in dorsal view,
lower margin of orbit evenly curved, tuber-
culate. Cheliped lightly granular; merus with
sharp subdistal spine; carpus with sharp in-
ner tooth and distinct angled projection or

62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

low tooth on outer margin; propodus un-
armed distally. Meri of walking legs, espe-
cially posterior ones, with distal dorsal tu-
bercle, lacking distinct distal spine. Dactyli
of walking legs dorsoventrally depressed,
height at midlength less than width. P35:
merus 5.3—5.9 (mean 5.5) times longer than
high, length 0.56 cb in female, 0.61-0.67
(mean 0.63) cb in males; carpus with line
of erect spinules dorsally; propodus length
4.5-5.1 (mean 4.8) times height, longer than
dactylus.

Size. — Males, cl 46-89.5 mm, cb 56-109
mm; only female examined, cl 48 mm, cb
64 mm.

Color. —Takeda’s (1983:164) colored fig-
ure shows a dark red crab.

Remarks. —This species resembles C.
quinquedens in having depressed dactyli on
the walking legs, but differs in that the legs
are shorter (P5 propodus 5-7 times longer
than high) and lack a strong distal spine on
the merus, the propodus of the chela lacks
a distal dorsal spine, and the carapace is
more granular.

The specimens from stations 4912 (male,
cl 47 m), 11,307 (female, cl 48 mm), and
10,616 (male, cl 46 mm) have much stron-
ger anterolateral spines on the carapace than
those from stations 2777 (males, cl 53.5 and
55 mm) and 4413 (female, cl 89.5 mm).

Takeda (1983) reported this species from
depths between 310 and 790 m. Our spec-
imens were taken in depths of 531, 622,
641, 641-915, and 723 m.

Name. —The name refers to the mythical
land of gold, El Dorado, thought by six-
teenth century Spanish explorers to be in
northern South America.

Distribution. —Northern coast of South
America, from Colombia to French Guiana,
in depths between 531 and 641-915 m.

Chaceon inglei, new species
Figs. 9-11

Geryon affinis. —Hansen, 1908:19, pl. 1, fig.
la, b. [Not Geryon affinis A. Milne Ed-
wards & Bouvier, 1894.]

Geryon gordonae Ingle, 1985:90, figs. 3, 4
[part, not material from Sierra Leone and
Cape Verde Islands].

Previous records.—Northeastern Atlan-
tic: 61°33'N, 19°0’W, 1089 fm (1993 m) and
61°30'N, 22°30’W, 975 fm (1784 m) (Han-
sen 1908).—56°49’-56°48'’N, 09°51’—
09°57'W, 2000 m; 51°6.8’-51°6.9'N,
13°16.7'-13°24.4'W, 1817-1930 m; 51°05.3’—
51°06.5'N, 13°04.5’-12°59.5'W, 1925-1960
m; 49°39.5'N, 12°36.9'W, 1857-1910 m;
49°38.6’N, 12°40.9’'W, 1860-1875 m;
49°32.6'-49°33.5'N, 13°7.1'-13°5.9'W, 1630-
1640 m; 49°30.1'’-49°27.7'N, 13°19.9'—
13°17.2'W, 1794-1785 m; and 49°27.3’—
49°30.1’N, 13°21.1'-13°26.8'W, 2045-2110
m (Ingle 1985).

Material.—North Atlantic: 61°33’N,
19°00'W, 1089 fm (1993 m), Ingolf sta 65:
one male (ZMC).—61°30'N, 22°30'W, 975 fm
(1784 m), Ingolf sta 67: one ovigerous female
(ZMC).—56°49’—-56°48'N, 09°51’-09°57'W,
2000 m, Challenger sta 12, leg R. W. Ingle:
one female (BM 1978.102).—55°07.7'N,
15°11.2'W to 55°10.3’N, 15°09.3"W, 2215—
2233 m, COB NORATLANTIQUE, PR
0006, sta CH04, Blake trawl, 10 Aug 1969:
two males (USNM 205986).—49°27.3’—
49°30.1’N, 13°21.1’-13°26.8'W, 2045-2110
m, Challenger sta 50518, 7 Jun 1979: one
male (BMNH).—48°45’N, 11°19.8’W to
48°46.6’N, 11°21.8’W, 1830-1870 m, Thal-
assa sta 2448, 1973: one female (MNHN
B.15835).—48°34'N, 10°51.6’W to 48°32.9’N,
10°49’W, 1975-2070 m, Thalassa sta 2453,
1973: one female, two juveniles (MNHN
B.15834).—47°34.6’N, 08°38.8’W, 2245 m,
BIOGAS IV, Jean Charcot sta CP 01, 25 Feb
1974: one male, one female (MNHN
B.7215).—47°34.5'’N, 08°34.2’W, 2180 m,
CENTOB BIOGAS, sta 011, CV 08: one fe-
male (MNHN _B.17207).—47°34.1'N,
08°40.5'W, 2175 m, CENTOB BIOGAS XI,
sta 1, CP 37: one female (MNHN B.17583).—
47°34'N, 8°41'W, 2100 m, CENTOB, EPITV,
sta 1, CP 40, 3 Sep 1985: one juvenile female
(MNHN B.17585).—47°33.2'N, 08°38.5'W,
2177 m, CENTOB BIOGAS, PR 08, CP:

VOLUME 102, NUMBER 1

63

Fig. 9. Chaceon inglei, male paratype,

cb 61 mm, North Atlantic: a, Dorsal view; 5, Carapace, enlarged.

64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 10. Chaceon inglei, male paratype, cb 34 mm, North Atlantic: a, Carapace; b, Ventral view of orbit; c,
Cheliped, dorsal view; d, Chela, outer view; e, P5; f, P5 dactylus, dorsal view; g, PS dactylus, posterior view.

one female (MNHN B.15821).—47°32.8'N,
08°33.5'W, 2115 m, BIOGAS VI, CP 26:
two males, one female (USNM 205987). —
47°32.7'N, 08°34.2'W, 2034 m, BIOGAS
II, CV 23: one ovigerous female (MNHN
B.17208).—47°31.4'’N, 8°42.7'W, 2100 m,
CENTOB, EPIIV, sta 1, CP 41, 4 Sep 1985:
one male (MNHN B.17586).—47°31'N,
08°16'W, 2200 m, BIOGAS I, CV 06: 2
females (MNHN B.15820).—47°28.1'N,
08°25.1'W to 47°29.6'N, 08°22.6’W, 2149-
2047 m, COB NORATLANTIQUE,
CHOO4, PR 124, BO19, sta 48, Blake Trawl,
31 Oct 1969: one male, one female (MNHN
B.15822).—42°32'N, 08°39.4'W, 2100 m,
CENTOB, EPI I, sta 1, CP 39, 30 Mar 1984:
one female (MNHN B.17584).—39°04.5'N,
32°43.5'W, 2120 m, Blake trawl, BIA-

CORES 1971, sta 131, 24 Oct 1971: one
male, one female (MNHN B.15836).

The female from Challenger sta 12, cl 43
mm, cb 62 mm, is the holotype (BM
1978.102). The other specimens are para-
types.

Diagnosis.—A small Chaceon, cl to 54
mm, cb to 72 mm in adults, with sharp,
well-developed anterolateral spines on the
carapace in adults and with laterally com-
pressed dactyli on the walking legs. Cara-
pace 1.3—1.5 times broader than long. Me-
dian pair of frontal teeth strong, sharp,
separated by U-shaped sinus, medians nar-
rower and extending farther forward than
laterals. Second lateral tooth small, fourth
usually obsolete or even totally absent, fifth
usually largest. Distance from first to second

VOLUME 102, NUMBER 1

65

Rigs 11.
28 mm, dorsal view.

tooth subequal to or less than distance from
third to fourth tooth, distance from first to
third tooth subequal to or less than distance
from third to fifth tooth. Suborbital tooth
large, extending at least to level of outer
frontal tooth, visible in dorsal view, sub-
orbital margin evenly concave. Cheliped
roughened dorsally; merus with sharp sub-
distal and smaller distal spine; carpus with
long inner spine and shorter outer spine;
propodus with distal dorsal spine or angled
projection. Meri of walking legs, especially
fourth and fifth, with distal, dorsal spine.
Dactyli of walking legs laterally com-
pressed, height at midlength greater than
width. P5: merus more than 6 times longer
than high, 6.5—7.2 (mean 6.8) times in males,
6.3-7.5 (mean 6.7) times in females. 8.0—
9.0 times in juveniles, with distal dorsal
spine, length 0.52—0.61 cb (mean 0.56 cb)
in females, 0.53—0.66 cb (mean 0.60 cb) in
males; carpus with line of erect spinules dor-
sally; propodus 5.1—6.7 (mean 5.7 in both
sexes) times longer than high, longer than
dactylus.

Chaceon inglei, North Atlantic: a, Male paratype, cb 18 mm, dorsal view; b, Male paratype, cb ca.

Size.—Males, cl 22-54 mm, cb 34-72
mm; females, cl 15—45 mm, cb 24-62 mm;
ovigerous females, cl 43 and 44 mm, cb 61
mm; juveniles cl 9 mm. Hansen (1908) re-
ported a male with cl 40 mm, an ovigerous
female with cl 42 mm. Ingle (1985) studied
males with cl 11.5—49.5 mm, females with
cl 7-43 mm.

Remarks.—The long anterolateral teeth
of the carapace, the long, slender legs, with
the merus of the fifth leg more than 6 times
longer than high, the sharp outer spine on
the carpus of the claw and the merus of the
walking legs, all will serve to differentiate
this species from C. affinis, the only other
species of Chaceon known to occur in the
northeastern Atlantic off the coast of Eu-
rope.

Chaceon affinis, which, like this species,
has compressed dactyli on the walking legs,
differs from C. inglei of the same size in the
features mentioned above and in having
smaller and blunter frontal teeth on the car-
apace and a much smaller suborbital spine.
A small male, cl 30 mm, cb 42 mm, of C.

66 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 12. Chaceon mediterraneus, male holotype, cb 65 mm, Mediterranean, dorsal view.

affinis available for comparison (USNM) has
a much shorter fifth leg, with the merus less
than 6 times as long as high and the prop-
odus less than 5 times as long as high. The
merus shows no trace of the distal, dorsal
spine that is so prominent in members of
C. ingle.

Chaceon gordonae, a species that occurs
off West Africa, is a much larger species, cb
to at least 161 mm (male from the Cape
Verde Islands, MNHN B.6432) rather than
less than 75 mm, that has much shorter legs,
with the merus less than 6 times longer than
high. In C. gordonae the fourth anterolateral
tooth of the carapace is present, whereas it
usually is obsolete in C. inglei.

This species differs from Chaceon medi-
terraneus, described below, in having com-
pressed rather than depressed dactyli on the
walking legs. That species also inhabits rel-
atively deep water.

Small specimens that lack the second and
fourth anterolateral teeth of the carapace
could be mistaken for Geryon longipes; both
species occur together in the northeastern
Atlantic. The frontal teeth of G. /ongipes are

much smaller than those of C. ing/ei at the
same carapace length.

This species occurs in relatively deep
water, with all records coming from depths
in excess of 1600 m. The specimens re-
ported by Hansen (1908) were taken in 1784
and 1993 m, and Ingle (1985) saw material
from depths ranging from 1630-1640 to
2045-2110 m. Our specimens were taken
in depths between 1784 and 2245 m, with
most lots coming from depths in excess of
2000 m.

Name.—We are pleased to dedicate this
species to our colleague R. W. Ingle, British
Museum (Natural History).

Distribution. —Northeastern Atlantic,
from about 61°N, south of Iceland, to 39°N,
off the Azores, in depths between 1630-1640
m to 2245 m.

Chaceon mediterraneus, new species
Figs. 12-13
Geryon gordonae. —Della Croce, Drago, &
Flocchini, 1988:6, 7, 8 [not Geryon gor-
donae Ingle, 1985].

VOLUME 102, NUMBER 1

ZOD

Fig. 13. Chaceon mediterraneus, male holotype, cb
65 mm, Mediterranean: a, Ventral view of orbit; b, P5
dactylus, posterior view; c, P5 dactylus, dorsal view.

Previous records. —Mediterranean Sea,
north of Sardinia, 1990-2008 m (Della
Croce, Drago, & Flocchini, 1988).

Material.— Mediterranean Sea, 37°56.7'N,
07°31.6’E, 2830 m, POLYMED sta CV 04,
18 May 1970: one male (holotype, MNHN
B.15824). Off Sardinia, 1990-2016 m; N.
Drago: one male (USNM); one male, one
female (BMNH).

Diagnosis. —A moderately large Cha-
ceon, cl to 78 mm, cb to 103 mm, with well-
developed anterolateral teeth on the cara-
pace and with dorsoventrally depressed
dactyli on the walking legs. Carapace 1.3
times broader than long. Median pair of
frontal teeth broad and short, separated by
V-shaped emargination, medians extending
further forward than laterals. Orbit broadly
concave, with low swelling on margin of
inner orbital tooth. First, third, and fifth
anterolateral spines stronger than remain-
der, fifth strongest, slender; gap between first
and second tooth very small. Distance from
first to second tooth subequal to distance
from third to fourth tooth, distance from
first to third tooth less than distance from

67

third to fifth tooth. Raised areas of carapace
pitted and eroded, but not tuberculate. Sub-
orbital tooth strong, sharp, extending about
to level of lateral frontal tooth, suborbital
margin tuberculate, evenly rounded. Che-
liped sparsely tuberculate dorsally; upper
margin of merus with sharp subdistal spine
and distal angled lobe; carpus with strong
inner spine and distinct but smaller outer
spine; propodus with distinct distal spine.
Meri of walking legs broad, upper margin
distinctly convex, with distal, dorsal spine.
Dactyli of walking legs dorsoventrally de-
pressed, width near midlength slightly larg-
er than height. P5: merus 4.5-5.4 times
longer than high, with distal dorsal spine,
length about half cb; carpus with line of
sharp spinules dorsally; propodus 4.1-4.3
times longer than high, slightly longer than
dactylus.

Size.—Males. cl 50-78 mm, cb 65-103
mm; female, cl 59 mm, cb 82 mm.

Remarks. —This species resembles C.
maritae and differs from both C. affinis and
C. inglei in having relatively broad, de-
pressed dactyli on the walking legs. It differs
from C. maritae in having much stronger
anterolateral spines on the carapace (even
when specimens of the same size are com-
pared), in having distal dorsal spines on the
meri of the walking legs, and in having slen-
derer walking legs. The merus of the fifth
leg is more than 5 times longer than high in
C. mediterraneus, about 4 times longer than
high in C. maritae.

Although the dactyli of the walking legs
of C. mediterraneus are not as conspicu-
ously flattened as in C. maritae, they are
distinctly broader than the dactyli in C. gor-
donae or C. inglei.

It is more than a little surprising to find
an undescribed Chaceon from the Mediter-
ranean. Even more surprising is that this
species was collected together with the new
genus and species described below.

The only eastern and central Atlantic
species with strong anterolateral spines on
the carapace in adults are C. atopus, C. gor-
donae, and C. inglei, and all of them have

68

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 14. Chaceon notialis, ovigerous female paratype, cb 110 mm, Argentina: a, Dorsal view; b, Carapace.

VOLUME 102, NUMBER 1

distinctly compressed dactyli on the walk-
ing legs.

Name.—Derived from the type locality
in the Mediterranean Sea.

Distribution. —Known only from the
western Mediterranean Sea, in 1990-2830
m.

Chaceon notialis, new species
Figs. 14-15

Geryon quinquedens.—Juanico, 1973:145,
fig. on p. 147.—Scelzo & Valentini, 1974:
561, figs. 1-2 [part, not specimens from
Brazil].—Boschi, 1976:66, 67; 1979:
140.—Boschi et al., 1981:247.—Barea &
Defeo, 1985:189, fig. 1; 1986:38, fig. 1.
[Not Geryon quinquedens Smith, 1879.]

Previous records.—Uruguay and Argen-
tina: 34°40’—-36°45’S, 250-800 m (Barea &
Defeo 1985).—34°40’—39°04’S, 250-800 m
(Barea & Defeo 1986).—33°56’'—36°25'S,
52°35'-54°51'W (Juanico 1973).—Uru-
guay: 33°38’S, 50°38’W, 790 m; 34°48’S,
52°02'W, 400 m; 35°04'S, 52°06’W, 800 m;
35°04'S, 52°15'W, 600 m (Scelzo & Val-
entini 1974).— Argentina: 36°24’S, 53°58'W,
120 m; 37°45’S, 54°55’'W, 300 m; and
38°55'S, 55°35'W, 170 m (Scelzo & Val-
entini 1974).— Buenos Aires Province, 36°
41°S (Boschi, 1976, 1979).—35°40’S,
52°47'W, 260 m (Boschi et al. 1981).

Material. — Argentina: 37°45'S, 54°55'W,
280-320 m, R/V Cruz del Sur, 17-18 May
1973: one ovigerous female, paratype
(USNM 205702).—38°55’S, 55°35'W, 170
m, R/V Cruz del Sur, 16 Apr 1973: one
male, holotype (USNM 205701).

Diagnosis. — A large Chaceon, cl to about
120 mm, cb to 135 mm in adults, with low,
blunt anterolateral spines on the carapace
and with dorsoventrally depressed dactyli
on the walking legs. Carapace 1.1-1.2 times
broader than long. Median pair of frontal
teeth sharp, projecting forward, separated
by U-shaped sinus, extending farther for-
ward than laterals. Anterolateral teeth low,

69

Fig. 15. Chaceon notialis, male holotype, cb 104
mm, Argentina: a, Ventral view of orbit; b, P5 dactylus,
posterior view; c, PS dactylus, dorsal view.

blunt, fourth tooth very small, almost ob-
solete. Distance from first to second an-
terolateral tooth less than distance from third
to fourth, distance from first to third an-
terolateral tooth less than distance from third
to fifth. Carapace surface strongly granular
posterolaterally. Suborbital tooth low, blunt,
falling short of level of lateral frontal tooth,
suborbital margin evenly curved. Cheliped
with blunt tooth subdistally and with distal
angled lobe on merus; carpus roughened
dorsally, lacking distal outer spine, anterior
margin denticulate; propodus lacking distal
spine or angled projection. Meri of walking
legs with at most a distal dorsal tubercle,
lacking a distinct distal dorsal spine. Dactyli
of walking legs depressed, width at mid-
length much greater than height. PS: merus
less than 5 times longer than high, length
0.60 cb; carpus with line of low but sharp
spinules dorsally; propodus length less than
4 times height, slightly longer than dactylus.

70 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Size. — Male, cl 94 mm, cb 104 mm; ovig-
erous female, cl 99 mm, cb 110 mm. Jua-
nico (1973) reported a male with cl 61 mm,
cb 73 mm; Boschi et al. (1981) studied a
male cl 30 mm. Scelzo & Valentini (1974)
studied 18 specimens, measuring as follows:
males, cl 88-116 mm, cb 97-124 mm; fe-
males, cl 72-99 mm, cb 82-110 mm. Barea
& Defeo (1985) reported catching males with
cb 32-135 mm, females with cb 48-124 mm.

Color.—Scelzo & Valentini (1974) re-
ported that this species is reddish in color.

Remarks.—Chaceon notialis resembles
both C. quinquedens, from the northeastern
Atlantic, and C. maritae, from West Africa,
in having depressed dactyli on the walking
legs, but it differs from both in numerous
features. It is a much shorter-legged species
than C. guinquedens: the merus of the fifth
leg is about three-fifths the width of the car-
apace as opposed to three-fourths the car-
apace width in C. quinquedens, and the
propodus of the fifth leg is less than 4 times
as long as high instead of 5—7 times as long
as high. In C. quinquedens there is a sharp
outer spine on the carpus and a distal dorsal
spine on the propodus of the cheliped, and
each walking leg has a distinct distal dorsal
spine. These spines are not present on the
cheliped of G. notialis, and in this latter
species the meri of the walking legs are armed
with at most a distal dorsal tubercle. The
surface of the carapace also is much coarser
in C. notialis, especially posterolaterally.

Chaceon notialis differs from C. maritae
as follows: the frontal teeth are sharper and
are not as close together; the anterolateral
teeth of the carapace all are sharper, and the
fourth tooth is much more distinct; the sub-
orbital teeth are sharper and longer; and the
walking legs are slightly longer. The merus
of the fifth leg is about 5 rather than 4 times
as long as high, and is about three-fifths
rather than two-thirds of the carapace width.

Barea & Defeo (1985, 1986) provided de-
tails of the occurrence and biology of this
species on joint fishing grounds off Uruguay
and Argentina. Boschi (1976, 1979) char-

acterized it as a cold-temperate species of
the Buenos Aires Province (36—41° South
latitude).

The specimen from off Brazil identified
with Geryon quinquedens by Rathbun (1937:
270) and material from Brazil identified by
Scelzo & Valentini (1974:560) with the same
species is referable to a new species, the de-
scription of which is in preparation. Scelzo
& Valentini reported that their Brazilian
specimens were cream-colored, whereas
their material from Uruguay and Argentina
was reddish. Further, the length/height ratio
of the propodus of the walking legs of their
specimens from Brazil was 4.3—4.5, longer
than that of C. notialis. The range in the
length/height ratio observed by Scelzo &
Valentini for their specimens from Uruguay
and Argentina, with one exception, is as we
found in our material of C. notialis.

Our specimens were taken in depths of
170 and 280-320 m. Boschi et al. (1981)
recorded a specimen from 260 m. Scelzo &
Valentini (1974) reported specimens from
depths of 120, 170, 300, 400, 600, 790, and
800 m, and Barea & Defeo (1985, 1986)
reported this species from fishing grounds
in depths between 250 and 800 m.

Name.—The specific name is from the
Latin, notialis, southern.

Distribution. —Known from the coasts of
Uruguay and Argentina, from about 33°S to
about 41°S, in depths between 120 and
800 m.

Chaceon sanctaehelenae, new species
Figs. 16-17

Material. —Sandy Bay, St. Helena Island
[15°58’S, 5°43’W], trap, 8 Oct 1968, leg F.
N. Martin: one male (holotype, USNM
125510).

Diagnosis. — A large Chaceon, cl 113 mm,
cb 134 mm in adult, with low, blunt an-
terolateral teeth on the carapace in adults
and with laterally compressed dactyli on the
walking legs. Carapace about 1.2 times
broader than long. Median pair of frontal

VOLUME 102, NUMBER 1

be

Fig. 16. Chaceon sanctaehelenae, male holotype, cb 134 mm, Saint Helena: a, Dorsal view; b, Carapace.

teeth broad, obtuse, separated by shallow
U-shaped emargination, medians extending
farther forward than laterals. Second and
fourth anterolateral teeth obsolete, present

as low lobes in adults. Distance from first
to second anterolateral tooth subequal to
distance from third to fourth tooth, distance
from first to third tooth subequal to distance

h2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

yo

jn
¢ b

Fig. 17. Chaceon sanctaehelenae, male holotype,
cb 134 mm, Saint Helena: a, Ventral view of orbit; 5,
P5 dactylus, posterior view; c, P5 dactylus, dorsal view.

from third to fifth. Carapace surface finely
granular, hepatic regions largely smooth.
Suborbital tooth short, blunt, scarcely vis-
ible in dorsal view, falling short of level of
lateral frontal tooth, suborbital margin
evenly rounded. Cheliped with blunt tooth
subdistally on merus; carpus roughened
dorsally, lacking outer spine, anterior mar-
gin smooth; propodus lacking distal angled
projection. Meri of walking legs lacking dis-
tal, dorsal spine. Dactyli of walking legs
compressed, height at midlength greater than
width. P5: merus 4.5 times longer than high,
length 0.6 cb; propodus slightly longer than
dactylus, length 3.3 times height.

Size. —Male, cl 113 mm, cb 134 mm.

Remarks. — This new species differs from
both C. affinis and C. atopus in having much
lower and blunter frontal and anterolateral
teeth on the carapace; in C. affinis the sec-
ond and fourth teeth are well-developed in

large specimens, but in C. sanctaehelenae
the second and fourth teeth are almost com-
pletely obsolete. In addition, in C. sanctae-
helenae the orbit appears to be shallower,
the suborbital tooth is shorter and more ob-
tuse, and the carpus of the walking legs is
much smoother dorsally than in C. affinis.
This new species further differs from C. ato-
pus in having much shorter walking legs (P5
merus 4.5 times rather than 6.5 times longer
than high), and in having the distance from
the first to third anterolateral teeth on the
carapace subequal to the distance from the
third to the fifth tooth; in C. atopus the dis-
tance from the first to third tooth is shorter.

Chaceon sanctaehelenae differs from C.
fenneri in some of these same features. All
marginal teeth of the carapace are lower than
in C. fenneri; in that species the frontal teeth
are much sharper and more prominent, and
the first, third, and fifth anterolateral teeth
are much more prominent. The carapace is
smoother than in C. fenneri, with lower and
fewer granules, and the dorsal ridge on the
carpus of the walking legs is much smooth-
cle

Chaceon chuni, from the southeastern At-
lantic, differs in having a larger suborbital
spine and in having a distinct distal dorsal
spine on the merus of each walking leg.

Name.—The specific name is derived
from the type locality.

Distribution. —Known only from the type
locality.

Zariquieyon, new genus

Type species. — Zariquieyon inflatus, new
species.

Name. —The name is dedicated to the late
Spanish carcinologist, Ricardo Zariquiey
Alvarez, whose studies of Mediterranean
decapods added so much to our knowledge
of the group. As in Chaceon, the ending
-on is added to make the name similar to
Geryon. The gender is masculine.

Definition. —Geryonid crabs with five an-
terolateral spines on each side of the cara-

VOLUME 102, NUMBER 1

pace. Carapace length about seven-tenths
width. Branchial regions markedly inflated.
Frontal teeth well-developed, large. Orbits
deep, rectangular.

Included species. —One, Zariquieyon in-
flatus, n. sp.

Remarks. —Zariquieyon shares with
Chaceon and Geryon the portunid-like che-
lipeds, with the subdistal dorsal spine on
the merus, the strong inner spine on the
carpus, the portunid-like chela, the single
inner suborbital tooth, the slender walking
legs with their naked dactyli, and the seven-
segmented abdomen in both sexes. Zari-
quieyon differs from those genera in having
a comparatively broader carapace, with the
posterolateral margins convex, strongly in-
flated branchial regions, and rectangular
rather than rounded orbits. It also shows far
more pebbling dorsally, visible only under
magnification, than any species of Chaceon
or Geryon.

Zariquieyon inflatus, new species
Figs. 18-19

Material. —Mediterranean Sea, 37°56.7°N,
07°31.6’E, 2830 m, POLYMED sta CV 04,
18 May 1970: one male, two females
(MNHN;; one female USNM 205985).

The holotype is the female, cl 22 mm, cb
35 mm, in the collection of the Muséum
National d’Histoire Naturelle, Paris; the
other specimens are paratypes.

Diagnosis.— A very small geryonid, cl 19—
22 mm, cb 27-35 mm, with strongly de-
veloped anterolateral teeth on the carapace
in adults and with laterally compressed dac-
tyli on the walking legs. Body, except ab-
domen, covered with fine granules, coarser
and larger on posterior branchial regions.
Regions of carapace well-marked, proto-
gastric region, area adjacent to fifth antero-
lateral spine, and branchial regions very in-
flated. Intestinal region with two
prominences. Posterolateral margins
strongly convex. Carapace 1.4—1.6 (mean
1.5) times broader than long. Median pair

73

Fig. 18. Zariquieyon inflatus, female holotype, cb
35 mm, Mediterranean, dorsal view.

of frontal teeth slender, separated by
V-shaped emargination. Second and fourth
anterolateral teeth reduced, fourth smallest;
other teeth progressively increasing in size
posteriorly, fifth much the largest. Distance
from first to second tooth short, about the
same as from third to fourth tooth, distance
from first to third tooth much less than dis-
tance from third to fifth. Orbits rectangular.
Suborbital tooth strong, extending to level
of outer frontal tooth, suborbital margin
denticulate. Cheliped finely granular dor-
sally; merus with sharp subdistal spine, dis-
tal angled projection spinulose; carpus peb-
bled dorsally, with long, sharp inner spine
and distinct outer spine or crest of spinules;
propodus pebbled dorsally, with distal dor-
sal projection. Meri of walking legs with dis-
tinct distal dorsal spine. Dactyli of walking
legs laterally compressed, height near mid-
length greater than width. P5: merus 6.3-
6.6 times longer than high, pebbled dorsal-
ly, with distal dorsal spine, length 0.45 times
cb; carpus with line of spinules dorsally;

74 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 19. Zariquieyon inflatus, female holotype, cb 35 mm, Mediterranean: a, Dorsal view; b, Ventral view
of orbit; c, Third maxilliped; d, Carpus of cheliped; e, P5; f PS dactylus, dorsal view.

propodus length about 5 times height, much
longer than dactylus.

Size.— Male, cl 19 mm, cb 28 mm; fe-
males, cl 20—22 mm, cb 27-35 mm.

Color. —Not recorded.

Remarks.—This species was taken to-
gether with Chaceon mediterraneus at a
depth of 2830 m in the western Mediter-
ranean basin. We know of no deeper record
for a brachyuran crab from the Mediterra-
nean.

Name.—The specific name is from the
Latin, inflatus, swollen.

Distribution. —Known only from the type
locality, in 2830 m.

Remarks

The genus Chaceon has proven to be much
more speciose than even we imagined when
we began studying its members, and much

additional work remains to be done on the
genus. The geographic ranges of most species
remain to be determined; in general, ranges
of each species are restricted latitudinally,
although some species, like C. bicolor, have
relatively wide ranges. Chaceon paulensis
from the southern Indian Ocean should be
redescribed, and the status of the species
reported from off Travancore, India, in 224—
284 fm (410-520 m) by Alcock (1899:85)
needs to be determined. A particularly in-
teresting problem that remains to be solved
is whether the types of Chaceon quinque-
dens, one of which is an ovigerous female
only 22 mm long (Smith, 1879:36), are con-
specific with the large, commercially im-
portant species now identified as C. quin-
quedens.

For most species there is no information
available on growth changes, especially as
it pertains to proportions of the walking legs

VOLUME 102, NUMBER 1

13

Table 1.—Characteristics of known species of Chaceon. Abbreviations are as follows: ? (not known); Color:
B (bicolor, tan and purple), R (red), T (tan), W (whitish); Size (maximum cb in mm); PS dactylus: C (laterally
compressed), D (dorsoventrally depressed); Distal spine on P5 merus: + (large, well developed), — (small,
reduced or absent); Anterolateral teeth of carapace: + (large, well-developed in adults), — (small, reduced or

absent).

Color

Northwestern Atlantic

C. eldorado R

C. fenneri ih

C. inghami R

C. quinquedens R
Southwestern Atlantic

C. notialis R

n. sp. Brazil T
Northeastern Atlantic

C. affinis Gi

C. inglei R

C. mediterraneus 2
Central South Atlantic

C. atopus ?

C. sanctaehelenae 2
Southeastern Atlantic

C. chuni T

C. erytheiae Rp

C. gordonae v

C. maritae Ae
Western Indian Ocean

C. crosnieri ?

C. macphersoni ?

C. paulensis R
Western and Central Pacific

C. bicolor B

C. granulatus ac
Eastern Pacific

n. sp. Chile W

and spination of the carapace and walking
legs. In the case of C. bicolor, for example,
we have identified a small, spiny, long-legged
specimen from deep water with the much
larger adult found in the same area, but the
possibility remains that more than one
species is involved.

Certainly additional species remain to be
recognized, and we believe that a key to the
species now known would be premature and
perhaps even confusing. As the species of

Dactylus Merus Carapace

Size P5 PS spines
109 D = a
190 € = =
110 C <5 =F
178 D 35 =
135 D = =
158 D = a
210 GC = =
2 € + ==
103 D a 55
(124 € =e +
134 € = =
114 ( se =
125 (G =F =
161 C =F =e
172 D = =
155 C == =F
126 D + --
113 Cc ata 2s
180 C = =
146 D == =
A/S) ‘S = 7

Chaceon may be difficult to distinguish in
the absence of comparative material, we
provide here, by broad geographic region, a
summary of the species found in each region
and their major characteristics (Table 1).

Acknowledgments

Many people contributed to this study.
We thank all of those who provided us with
material or access to their collections: E. E.
Boschi, Instituto Nacional de Investigacion

76 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

y Desarollo Pesquero, Mar del Plata, Ar-
gentina; Diane E. Brown, Australian Mu-
seum, Sydney; Alain Crosnier and D. Gui-
not, Muséum National d’Histoire Naturelle,
Paris; Reginald M. Gooding, then of the
NOAA/NMEFS Southwest Fisheries Center,
Honolulu Laboratory; R. W. Ingle, British
Museum (Natural History), London; En-
rique Macpherson, Instituto de Ciencias del
Mar, Barcelona; Torben Wolff, Zoological
Museum, Copenhagen; and Nicolino Dra-
go, Universita di Genova, Italy, who al-
lowed us to add reference to his material
from the Mediterranean in galley.

Our studies on geryonid crabs have been
supported in part by the Food and Agri-
culture Organization of the United Nations
(FAO); that support is gratefully acknowl-
edged. This paper also was supported in part
by the Smithsonian Institution, through its
Research Opportunities Fund.

We thank Roy K. Kropp for taking the
photographs used in Figs. 14 and 16. Fig. 1
was taken by the staff photographer of the
Riksmuseum van Natuurlijke Historie,
Leiden, and Figs. 3 and 5 were taken by the
staff photographer of the Muséum National
d’ Histoire Naturelle, Paris. Lilly King Man-
ning drew the line drawings and prepared
all of the figures for publication.

We thank Fenner A. Chace, Jr., Enrique
Macpherson, and Marilyn Schotte for read-
ing a draft of the manuscript.

Literature Cited

Alcock, A. 1899. An account of the deep-sea Brachy-
ura collected by the Royal Indian Marine Survey
Ship Investigator, pp. 1-85, 1-2, pls. 1-4. Cal-
cutta: Indian Museum.

Barea, L., & O. Defeo. 1985. Primeros ensayos de
captura del crustaceo batial Geryon quinquedens
Smith en el area comun de pesca Argentino-
Uruguaya.—Contribucion, Departamento de
Oceanografia (F.H.C.), Montevideo 2(3):189-
203:

——,, & . 1986. Aspectos de la pesqueria del
cangrejo rojo (Geryon quinquedens) en la zona
comun de pesca Argentino-Uruguaya.— Publi-
caciones de la Comision Técnica Mixta del
Frente Maritimo 1(1):38—46.

Boschi, E. E. 1976. Nuevos aportes al conocimiento

de la distribucion geografica de los crustaceos

decapodos del Mar Argentino.—Physis

(A)35(90):59-68.

1979. Geographic distribution of Argenti-
nian marine decapod crustaceans. — Bulletin of
the Biological Society of Washington 3:134—143.
, M. I. Iorio, & K. Fischbach. 1981. Distri-
bucion e abundancia de los crustaceos decapo-
dos capturados en las campanas de los B/I “Wal-
ther Herwig” y “Shinkai Maru” en el Mar
Argentino, 1978-1979. In Campanas de inves-
tigacion pesquera realizadas en el Mar Argen-
tino por los B/I “Shinkai Maru” y “Walther
Herwig” y el B/P “Marburg,” Anos 1978 y
1979.—Contribuciones del Instituto Nacional
de Investigacion y Desarrollo Pesquero 383:233-
253.

Chun, C. 1903. Aus den Tiefen der Weltmeeres, 2nd
edition. Gustav Fischer, 592 pp.

Colosi,G. 1923. Una specie fossile de Gerionide (De-
capodi brachiuri).— Bolettino della Societa dei
Naturalisti in Napoli, 35 (series 2, volume 15),
37:248-255.

Della Croce, N., N. Drago, & G. Flocchini. 1988.
Ricerche biologiche e geofisiche. Campagna
oceanografica N/R “Minerva” (14-31.8.1987).—
Istituto di Scienze Ambientali Marine, Univ-
ersita di Genova, Rapporto Tecnico 24:1-10.

Gerstaecker, A. 1856. Carcinologische Beitrage.—
Archiv fur Naturgeschichte 22(1):101-—162, pls.
4-6.

Grifin, D. J. G., & D. E. Brown. 1976. Deepwater
decapod Crustacea from eastern Australia:
Brachyuran crabs.—Records of the Australian
Museum 30(11):248-271.

Guinot, D. 1969. Les Goneplacidae (suite et fin). Re-

cherches préliminaires sur les groupements na-

turels chez les Crustacés Décapodes Brachy-
oures, VII.—Bulletin du Muséum National

d’Histoire Naturelle (2)41(3):688—724. pls. 3-5.

. 1971. Synthése et bibliographie. Recherches

préliminaires sur les groupements naturels chez

les Crustacés Décapodes Brachyoures, VIII.—

Bulletin du Muséum National d’Histoire Na-

turelle (2)42(5):1063—-1090.

—, & B. Richer de Forges. 1981. Crabes de pro-
fondeur, nouveaux ou rares, de l’Indo-Pacifique
(Crustacea, Decapoda, Brachyura) (Deuxiéme
partie).— Bulletin du Muséum National d’His-
toire Naturelle, Paris (4)3(A1):227—260, pls. 3-7.

Hansen, H. J. 1908. Crustacea Malacostraca, I.—The
Danish Ingolf-Expedition 3(2):1—120, pls. 1-5.

Herbst, J. F. W. 1799-1804. Versuch einer Natur-
geschichte des Krabben und Krebse, nebst einer
systematischen Beschreibung ihrer verschiede-
nen Arten, 3:215 pp. Berlin and Stralsund.

Ingle, R. W. 1985. Geryon gordonae sp. nov. (Decap-

VOLUME 102, NUMBER 1

oda Brachyura, Geryonidae) from the north-
eastern Atlantic Ocean. —Crustaceana 48(1):88-
98.

Intés, A. 1978. Péche profonde aux casiers en Nou-
velle Calédonie et iles adjacentes. Essais préli-
minaires. ORSTOM, Centre de Nouméa, Rap-
ports Scientifiques et Techniques 2:1-10, figs.
1-10.

Juanico, M. 1973. Hallazgo de Geryon quinquedens
Smith, 1879 (Crustacea Decapoda), en aguas
Uruguayas.— Revista de Biologia del Uruguay
1(2):145-149.

King, M.G. 1984. The species and depth distribution
of deepwater caridean shrimps (Decapoda, Ca-
ridea) near some southwest Pacific islands.—
Crustaceana 47(2):174-191.

Kroyer, H. 1837. Geryon tridens, en ny Krabbe.—
Naturhistorisk Tidsskrift 1:10—21, pl. 1.
Macpherson, E. 1983. Crustaceos decapodos captu-
rados en las costas de Namibia. — Resultados Ex-
pediciones Cientificas (supplement to Investi-

gacion Pesquera, Barcelona) 11:3-80.

1984. Crustaceos decapodos del Banco Val-
divia (Atlantico sudoriental).— Resultados Ex-
pediciones Cientificas (supplement to Investi-

gacion Pesquera, Barcelona) 12:39-10S.

Manning, R. B., & L. B. Holthuis. 1981. West African
brachyuran crabs.—Smithsonian Contributions
to Zoology 306:379 pp.

——,,& . 1984. Geryon fenneri, a new deep-
water crab from Florida (Crustacea: Decapoda:
Geryonidae).—Proceedings of the Biological
Society of Washington 97(3):666-673.

——, & 1986. Notes on Geryon from Ber-

muda, with the description of Geryon inghami,

new species (Crustacea: Decapoda: Geryoni-
dae).— Proceedings of the Biological Society of

Washington 99(2):366-373.

, & 1987. The status of Geryon tri-

spinosus (Herbst) (Geryonidae). — Investigacion

Pesquera, Barcelona 51 (supplement 1):57-62.

,& . 1988. South African species of the

genus Geryon (Crustacea, Decapoda, Geryoni-

ve |

dae).—Annals of the South African Museum
98(3):77-92.

Milne Edwards, A. 1882. Rapport sur les travaux de
la Commission chargée par M. le Ministre de
l’Instruction Publique d’étudier la faune sous-
marine dans les grandes profondeurs de la Médi-
terranée et de l’Océan Atlantique. — Archives des
Missions Scientifiques et Litteraires, Paris (3)9:
1-59.

—., & E.-L. Bouvier. 1894. Brachyoures et ano-
moures. Crustacés décapodes provenant des
campagnes du yacht I’Hirondelle (1886, 1887,
1888). Premiére partie.—Résultats des cam-
pagnes scientifiques accomplies sur son yacht
par Albert Is", Prince Souverain de Monaco 7:
1-112.

Rathbun, M. J. 1937. The oxystomatous and allied
crabs of America. — United States National Mu-
seum Bulletin 166, vi + 278 pp.

Sakai, T. 1978. Decapod Crustacea from the Emperor
Seamount Chain.—Researches on Crustacea 8
(supplement): 1-39, pls. 1-4.

Scelzo, M. A., & A. Valentini. 1974. Presencia de

- Geryon quinquedens Smith en aguas del Oceano
Atlantico sudoccidental (Decapoda, Brachyura,
Geryonidae).—Physis, Buenos Aires (A)33(87):
557-567.

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):27-136.

Takeda, M. 1983. Crustaceans. Pp. 19-185 in Crus-
taceans and mollusks trawled off Suriname and
French Guiana. Japan Marine Fishery Research
Center, Tokyo.

(RBM) Department of Invertebrate Zo-
ology, National Museum of Natural His-
tory, Smithsonian Institution, Washington,
DC 20560; (LBH) Rijksmuseum van Na-
tuurlijke Historie, Postbus 9517, 2300 RA
Leiden, The Netherlands.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 78-83

SYNALPHEUS AROSTRIS AND PHILOCHERAS LAPILLUS,
TWO NEW SPECIES OF CARIDEAN SHRIMP
(CRUSTACEA) FROM THE TROPICAL
EASTERN PACIFIC

Mary K. Wicksten

Abstract.—Two new species of shrimp (Decapoda: Caridea) are described
from the eastern Pacific. Synalpheus arostris is described from specimens from
western Colombia. Philocheras lapillus is the first species of its genus to be

reported in the area.

During examination of specimens from
western Colombia and the Galapagos Is-
lands, I discovered two undescribed species
of shrimp. The new species of Philocheras
is the first report of the genus from the trop-
ical eastern Pacific. I thank Gabriel Ramos,
Universidad del Valle (UV); Raymond
Manning, United States Museum of Natural
History (USNM); and Janet Haig, Allan
Hancock Foundation, University of South-
ern California (AHF) for their interest and
assistance during this study. The figures are
by Gabriel Ramos and Joseph Goy, Texas
A&M University. This project was aided by
a Visiting Fellowship of the Smithsonian
Institution.

Family Alpheidae

Synalpheus arostris, new species
Fig. 1

Type material. —Holotype: female, total
length 10.5 mm. Morro de Los Agujeros,
Bahia de Malaga, Colombia (3°55’N,
77°20'W), rocky intertidal zone, 28 Nov
1981, collector not reported, AHF no. 8111.
Paratype: female, same site and date, UV.

Description. — Rostrum lacking, a slight
raised area present between orbital teeth.
Orbital teeth reaching less than 0.5 x length
of first segment of antennular peduncle,
longer than broad.

Visible part of first antennular segment

1.3 x as long as second antennular article,
third article shorter. Stylocerite reaching just
beyond first antennular segment. Scapho-
cerite with lateral tooth reaching beyond end
of antennular peduncle and exceeding car-
pocerite. Squamous part of scaphocerite
reaching or almost reaching end of anten-
nular peduncle. Inferior tooth of basicerite
reaching as far forward as stylocerite, su-
perior tooth well developed and acute.

Large chela 3 x as long as wide, with fin-
gers occupying about 0.3 entire length.
Superior margin of palm bearing blunt knob
proximal to dactylus. Merus 2x long as
broad, superior margin blunt.

Small chela 2.4 x long as wide, with fin-
gers occupying about 0.4 entire length.
Merus almost 3x long as wide, bearing 3
teeth on superior margin.

Carpal articles of second leg with ratio 10:
3:3:3:4.

Merus of third leg 4 x long as broad, with-
out spines. Carpus without teeth or spines.
Propodus 7 X long as wide, straight, bearing
7-8 spines on inferior margin and a pair of
terminal spines. Dactyl biunguiculate, 0.3 x
propodus, relatively straight; superior and
inferior ungui slender and almost equal in
size and shape. Fourth and fifth legs similar
to third.

Telson only slightly longer than width of
anterior margin, tapering to posterior mar-
gin. Two to three dorsolateral spines per

VOLUME 102, NUMBER 1

79

Pig: 1:
cheliped; E, Large cheliped; F, Third pereopod.

side, pair long spines at distolateral margin,
distolateral margin produced into small
tooth. Posterior margin of telson rounded.

Discussion. —Coutiére (1909) established
six groups in the genus Synalpheus, based
on features of the rostrum, orbital teeth, se-
tae of the smaller chela, ungui of the dactyls,
and other features. Banner & Banner (1975)
reexamined these groups and found that
perhaps only two of them were of taxonom-
ic validity—the others contained such a
range of variation or overlapped other
groups to such an extent that the group cer-
tainly was artificial. The new species illus-
trates the difficulties in finding relationships
among the species of Synalpheus; lacking a

Synalpheus arostris, A, Holotype in lateral view; B, Anterior end in dorsal view; C, Telson; D, Small

rostrum, it cannot easily be compared with
many other species.

The short orbital teeth and equal, slender
ungui of the dactyls of S. arostris are found
in species of Coutiére’s ““Brevicarpus” group,
known only from American waters. Three
species, S. minus (Say), S. obtusifrons Chace,
and S. brevicarpus Coutiére are found in the
Caribbean region. All of these species have
short rostra. Synalpheus minus has a sty-
locerite that greatly exceeds the first article
of the antennular peduncle, but the spine of
the scaphocerite does not reach the end of
the carpocerite. Synalpheus obtusifrons has
a rounded stylocerite and a short scapho-
cerite that does not reach the end of the

80 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Philocheras lapillus, paratype: A, Entire animal in lateral view; B, Carapace in dorsal view; C, Carapace

in lateral view; D, Telson and uropods.

antennular peduncle. The eastern Pacific
species, S. digueti Coutiére, is common in
rocky intertidal areas. Aside from lacking a
rostrum, S. arostris can be distinguished
from S. digueti by having a shorter stylo-
cerite, reaching barely beyond the end of
the first article of the antennular peduncle
rather than to the middle of the second ar-
ticle; and having a longer spine on the
scaphocerite, reaching proportionally far-
ther beyond the blade than in S. digueti.
Only S. arostris has three teeth on the su-
perior margin of the merus of the small che-
liped.

Synalpheus arostris can be distinguished
easily from S. /ockingtoni Coutiére by the
lack of the rostrum (seen in both specimens
of the new species). Should one encounter
a specimen of S. lockingtoni missing a ros-
trum, the species can be distinguished by
the sharp spine at the anterior margin of the

palm of the major chela, above the articu-
lation of the dactyl, in S. arostris. In S. lock-
ingtoni, the anterior margin of the major
chela ends in a knob or tubercle. The pos-
terior margins of the telson of S. arostris are
sharply triangular, those of S. lockingtoni
are weakly pointed. Adult S. /ockingtoni can
be larger than the specimens of S. arostris:
Schmitt (1921) gives a length of 30 mm for
a large female S. lockingtoni.

Family Crangonidae
Philocheras lapillus, new species
Figs. 2-4

Type material. —Holotype male, total
length 13.7 mm. Off Gardner Bay, Hood
Island (Isla Espanola), Galapagos (Archi-
pelago de Colon) 1°22’S, 90°40’W), 46-65
m, rock, 31 Jan 1934, Velero III sta. 201-
34, USNM 234435.

VOLUME 102, NUMBER 1

81

Fig. 3. Philocheras lapillus: A, Antennule; B, Scaphocerite; C, Mandible; D, First maxilla; E, Second maxilla;
F, First maxilliped: G, Second maxilliped; H, Third maxilliped.

Paratypes.—Tagus Cove, Albemarle Is-
land (Isla Isabela) (0°17'S, 91°23'’W), 37 m,
rock and nullipores, 11 Jan 1934, Velero III
sta. 149-34, one specimen. Stephens Bay,
Chatham Island (Isla San Cristobal), (0°48’S,
89°31'W), 59 m, fine sand and coralline al-
gae, Velero III sta. 170-34, 21 Jan 1934,
three specimens. Off Cartago Bay, Albe-
marle Island (0°35’S, 90°54’W), 59 m, mud,
25 Jan 1934, Velero IIT sta. 185-34, five
specimens. Off Cartago Bay, Albemarle Is-
land (0°34’S, 90°53’W), 59 m, sand and nul-
lipores, 25 Jan 1934, Velero IIT sta. 186-34,
two specimens. Off Gardner Bay, Hood Is-
land (same station as holotype), Velero III
sta. 201-34 (four specimens USNM, two
specimens AHF). All specimens USNM ex-
cept as noted.

Description. —Rostrum exceeding or
nearly exceeding end of cornea, distal end
wide. Anterior margin of carapace with
teeth, one suborbital tooth, pair near base
of second antenna and two pterygostomial
teeth. One large tooth on dorsal midline.

Abdominal pleura rounded. Sixth ab-
dominal segment about 1.5 x length of fifth,

slightly shorter than length of telson. Telson
narrow, with two pair dorsolateral, three pair
terminal spines and sharp posteromesial tip.

Basal article of antennular peduncle not
reaching as far as end of cornea. Stylocerite
not as long as basal article of antennular
peduncle, broad and quadrangular. Last two
articles of antennular peduncle short.

Scaphocerite longer than antennular pe-
duncle, outer margin straight, blade longer
than spine. Basicerite short.

Third maxilliped exceeding scaphocerite.
Ultimate segment longer than penultimate,
0.6X antepenultimate. Exopod present.
Other mouthparts as figured.

First pereopod stout, subchelate. Dactyl
about 0.3 X propodus length, subchelar spine
narrow and simple. Carpus 0.25 propo-
dus, with one or no spines. Merus shorter
than chela, with two teeth on distal end near
articulation with carpus and one large tooth
on lower margin. Ischium and basis short,
without teeth. No exopods or epipods on
pereopods.

Second pereopod short, chelate. Dactyl
0.5 X propodus, carpus about equal to chela,

82 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(Tk
Ati

‘)

Fig. 4. Philocheras lapillus: A, First pereopod; B, Second pereopod; C, Third pereopod; D, Fourth pereopod;

E, Fifth pereopod; F, Second pleopod of female.

0.5 merus. Merus longer than ischium.
Third pereopod slender, thread-like. Dactyl
thin, 0.7 X propodus. Propodus 0.6 car-
pus, carpus 1.8x merus, merus 0.6% is-
chium. Fourth and fifth pereopods similar.
Dactyl long and simple, 0.8 propodus.
Carpus 0.8 X propodus, merus 1.2 X carpus,
ischium 0.8 X merus.

Second pleopod with appendix interna,
male with appendix masculina. Uropods
long, outer uropod shorter than inner. Outer
uropod with small posterolateral tooth.

Color in life. —Carapace almost all white,
body Van Dyke brown, speckled. (Field note
for specimen at sta. 149-34, by Waldo L.
Schmitt.)

Discussion. —1 follow the revision of
Chace (1984) in considering Philocheras to
be distinct from Pontophilus. Philocheras
lapillus most closely resembles P. gorei
(Dardeau, 1980) from the Gulf of Mexico

and Georgia. In P. gorei, however, there are
always teeth on the distal margins of the
carpus of the first pereopod. The outer fla-
gellum of the antennular peduncle of P. /a-
pillus is much thicker than that of P. gorei.
The tip of the rostrum of P. /apillus is round-
ed, not spatulate. Pontophilus lapillus also
may be larger than P. gorei: the carapace
length of the holotype is 3.3 mm, that of the
type of P. gorei is 2.5 mm.

The habitat and living color of P. /apillus
suggest that the animal’s mode of life is sim-
ilar to that of species of Crangon and Lis-
socrangon along the coast of California.
These species have a speckled, cryptic color
pattern, which provides camouflage when
the shrimp dig into sand (Kuris & Carlton
1977; Ricketts et al. 1985).

The species epithet means “‘pebble,”’
which the resting animal resembles. The
name is a noun in apposition.

VOLUME 102, NUMBER 1

Literature Cited

Banner, D. M., & A. H. Banner. 1975. The alpheid
shrimp of Australia. Part 2: the genus Synal-
pheus.—Records of the Australian Museum
29(12):267-389.

Chace, F. A., Jr. 1984. The caridean shrimps (Crus-
tacea: Decapoda) of the A/batross Philippine Ex-
pedition, 1907-1910, Part 2: Families Glypho-
crangonidae and Crangonidae.— Smithsonian
Contributions to Zoology No. 397:1-63.

Coutiére, H. 1909. The American species of snapping
shrimps of the genus Synalpheus. — Proceedings
of the United States National Museum 36(1659):
1-93.

Dardeau, M. 1980. A new species of Pontophilus
(Crustacea: Natantia: Crangonidae) from the
Gulf of Mexico and the western Atlantic. — Pro-

83

ceedings of the Biological Society of Washington
93(3):563-572.

Kunis, A. M., & J. T. Carlton. 1977. Description of
a new species, Crangon handi, and new genus,
Lissocrangon, of crangonid shrimps (Crustacea:
Caridea) from the California coast, with notes
on adaptation in body shape and coloration. —
Biological Bulletin 153:540-559.

Ricketts, E. F., J. Calvin, J. W. Hedgpeth, & D. W.

Phillips. 1985. Between Pacific tides, 5th ed.
Stanford University Press, Stanford, Califor-
nia, 652 pp.

Schmitt, W. L. 1921. Marine decapod Crustacea of

California.— University of California Publica-
tions in Zoology 23:1—470.

Department of Biology, Texas A&M Uni-
versity, College Station, Texas 77843.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 84-88

THE HOLOTYPE OF HETEROCARPUS ALEXANDRI
A. MILNE-EDWARDS
(CRUSTACEA: DECAPODA: PANDALIDAE)

Fenner A. Chace, Jr.

Abstract.—The ovigerous female holotype of Heterocarpus alexandri from
off Havana, Cuba, is described and reillustrated. A male of presumably the
same species is recorded from the Bahamas, and probably two related species
are recorded and diagnosed from the Pacific region: H. nesisi Burukovsky from
Baja California and an undescribed species previously recorded as H. alexandri
from Hawaii. The systematic status of H. alexandri is discussed but not con-

summated.

This paper was prompted by the discov-
ery in the collection of the Museum of Com-
parative Zoology, Harvard University, of
the holotype of Heterocarpus alexandri,
which had been rather inadequately de-
scribed and was feared lost. This opportu-
nity was made especially significant by the
revelation that A. Milne-Edwards’ (1883)
original illustration (there was no written
description) was erroneous in several im-
portant characters, thereby precluding as-
surance of the identification of the species
from other material. Although I have of-
fered comparisons of H. alexandri with
specimens of related species from the Pacific
and have discussed the possible systematic
status of those species, my objective in this
contribution has been to record the typical
features of H. alexandri for future reference,
not to define its hierarchical position in the
family.

Family Pandalidae
Genus Heterocarpus A. Milne-Edwards
Heterocarpus alexandri A. Milne-Edwards,
1883
Fig. 1

Heterocarpus Alexandri A. Milne-Edwards,
1883 [unnumbered plate] [type locality:
Blake dredging no. 2; north of Havana;

23°14’00’N, 82°25'00”W; 805 fms (1472
m)].—De Man, 1920:108, 153, 154.

Heterocarpus alexandri. —Faxon, 1896:
161.—Chace, 1985:19, 20, fig. 135. —Bu-
rukovsky, 1986:64, 67, 69.

Description of holotype.—Ovigerous fe-
male with postorbital carapace length of 12.8
mm. Rostrum (Fig. 1a) incomplete, existent
portion nearly three-fourths as long as post-
orbital carapace, overreaching antennal
scale, with sharp lateral carina arising from
orbital margin, armed dorsally with eight
strong teeth, four on carapace posterior to
orbital margin, posteriormost arising slight-
ly anterior to midlength of carapace, bearing
denticle on posterior slope, armed ventrally
with four teeth. Carapace with middorsal
carina extending over anterior two-thirds of
length, posterior one-third smoothly round-
ed; antennal spine flaring sinuously laterad
from origin below orbit, slightly smaller than
anteriorly directed branchiostegal spine,
latter buttressed by submarginal carina ex-
tending to near midlength of carapace; low,
broad elevation, accentuated by rather deep
dorsal and ventral depressions, extending
posteriorly from orbital region and indis-
tinctly joining somewhat more distinct su-
prabranchial ridge reaching nearly to pos-
terior margin of carapace.

VOLUME 102, NUMBER 1

Pig. 1.
aspect; b, Same, dorsal aspect; c, Abdomen, lateral aspect; d, third abdominal somite, dorsal aspect; e, Sixth
abdominal somite, telson, and uropods, dorsal aspect; f Juncture of sixth abdominal somite and telson, lateral
aspect; g, Right second pereopod. (Magnifications: a, c, x3; b, d, e, g, x6; ff x12.)

Abdomen (Fig. lc) with tergum of first
somite rather severely distorted, second so-
mite with distinct, anteriorly convex, pos-
teriorly setose transverse groove crossing
tergum posterior to midlength, third somite
with sharply defined middorsal boss on ter-
gum (Fig. 1d); sixth somite fully one and
three-fourths times as long as fifth somite,
slightly more than twice as long as high,

Heterocarpus alexandri, ovigerous female holotype: a, Carapace and anterior appendages, lateral

dorsal surface shallowly convex and bluntly
distinct from lateral surface (Fig. le), armed
posteroventrally with small but discrete
tooth (Fig. 1f); pleura of four anterior so-
mites with irregularly convex margins, pleu-
ron of fifth somite with sharp posteroventral
tooth. Telson (Fig. le) deeply sulcate mid-
dorsally on anterior half of length, armed
with five pairs of dorsolateral spines, in-

86 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

cluding pair lateral to long, intermediate
posterior spines.

Eye with subspherical cornea slightly
broader than stalk, without ocellus (Fig. 15).
Antennule with stylocerite narrowly taper-
ing to sharp point near level of second third
of second antennular segment. Antennal
scale with lateral margin nearly straight, ter-
minating in strong distolateral tooth not
reaching level of distal oblique margin of
blade. Third maxilliped overreaching an-
tennal scale by length of terminal segment,
exopod reaching to about midlength of an-
tepenultimate segment.

Pereopods with epipods on four anterior
pairs. First pereopods missing or incom-
plete. Right second pereopod (Fig. 1g) over-
reaching antennal scale by length of chela
and about one-third of carpus, carpus com-
posed of 18 articles; left second pereopod
missing. All three pairs of posterior pereo-
pods missing.

Uropod (Fig. le) with branches subequal
in length, slightly overreaching telson, not
including posterior spines.

Eggs measuring about 0.4 x 0.6 mm.

Remarks.—The unique holotype differs
from A. Milne-Edwards’ original illustra-
tion in having the margin of the pleuron of
the fourth abdominal somite rounded in-
stead of posteroventrally acute, the telson
armed with five rather than four pairs of
dorsolateral spines, the blade of the anten-
nal scale oblique rather than transversely
truncate, and the carpus of the right second
pereopod composed of 18 rather than 25
articles.

The specimen assigned to H. alexandri
by Faxon (1896:161) from Blake dredge no.
196; off Martinique; 1030 fms (1884 m) ap-
parently has been temporarily misplaced in
the MCZ collections, but there is a male of
that species in the Smithsonian collections,
taken by the Albatross at station 2629;
mouth of Exuma Sound, Bahamas;
23°48'40"N, 75°10'40’W; 1169 fms (2138
m); 8 Mar 1886. That specimen has a post-
orbital carapace length of 11.0 mm; the an-

terior part of the rostrum is missing beyond
the seventh dorsal and the posterior ventral
teeth. It differs otherwise from the holotype
in having the submarginal carina that but-
tresses the branchiostegal spine less sharp;
the tergum of the first abdominal somite
traversed by a distinct, anteriorly convex
groove similar to the one on the second so-
mite; the margins of the dorsal boss on the
third somite curving laterad more strongly
at the anterior end; the sixth somite nearly
twice as long as the fifth and about two and
one-half times as long as high; the telson
armed with five pairs of dorsolateral spines
on the left side but only four on the right,
including the pair lateral to the intermediate
posterior spines; the stylocerite not over-
reaching the acute lateral projection of the
first antennular segment; the blade of the
antennal scale slightly narrower beyond the
base of the distolateral spine than in the
holotype (but not quite as distinctly so as
indicated in Chace (1985:fig. 135)); the first
pereopod overreaching the antennal scale
by about one-half the length of the chela;
and the appendix interna on the second
pleopod overreaching the appendix mas-
culina by at least one-fourth of its length.
In virtually all other characters, the male
agrees with the ovigerous female holotype.

The Hawaiian specimen recorded as H.
alexandri by Rathbun (1906:918) from A/-
batross station 4181; near Kauai Island; 81 1—
671 fms (1483-1227 m) is an ovigerous fe-
male with a postorbital carapace length of
9.7 mm. It almost certainly represents a dis-
tinct species. The rostrum reaches only about
as far as the distal end of the antennular
peduncle, does not reach the level of the
distal end of the antennal scale, and has a
rostral formula of 4+ 6/4; although the car-
apace is very like that of H. alexandri, the
transverse grooves on the first and second
abdominal somites are much less distinct,
that on the second being~ virtually non-
existent, and there is no suggestion of a boss
on the tergum of the third somite; the sixth
somite is two and one-fourth times as long

VOLUME 102, NUMBER 1

as the fifth and fully two and one-half times
as long as high; the telson is deeply sulcate
anteriorly, but the posterior one-half is
missing.

Another Smithsonian specimen, identi-
fied by Waldo L. Schmitt as H. alexandri,
from “D. 5682,” which almost certainly
refers to an Albatross station by that num-
ber in San Lucas Bay, Baja California; 22°
48'20’N, 109°52'40”W; 491 fms (898 m);
24 Mar 1911, is a large female with a post-
orbital carapace length of 31.5 mm; the
rostrum far overreaches the antennal scale
and has a rostral formula of 4+7/7; the
middorsal postrostral carina reaches poster-
iorly to about the posterior one-sixth of the
postorbital carapace length, and the lateral
postorbital ridge is very like the one in the
holotype of H. alexandri, but the branchio-
stegal spine is situated closer to the antennal
spine and is not supported by a long, cari-
nate buttress; the two anterior abdominal
somites lack transverse grooves; the third
abdominal somite is produced posteriorly
in the midline, but it is without a dorsal
boss on the tergum, except for a vestige sug-
gested by a short, mesially convex furrow
on each side of the midline; the sixth ab-
dominal somite is somewhat transversely
convex dorsally, nearly two and one-fourth
times as long as the fifth, and one and two-
thirds times as long as high; the telson is
rather deeply sulcate dorsally, especially an-
teriorly, armed. with five pairs of dorsolat-
eral spines, including the pair lateral to the
intermediate posterior spines; the eyestalk
widens distally to a nearly hemispherical
cornea; the pereopods have well-developed
epipods on the four anterior pairs; the first
pair slightly overreaching the antennal scale;
the right second pereopod reaches at least
to the distal one-fourth of the antennal scale,
the carpus is composed of five articles; the
left second pereopod overreaches the an-
tennal scale by the length of the chela and
the distal carpal article, and the carpus com-
posed of 15 articles; the third pereopod
overreaching the antennal scale by the

87

lengths of the dactyl and propodus, the dac-
tyl bearing four spines on the flexor margin
proximal to the distal tooth; the uropod with
the lateral branch reaching to the level of
the posterior margin of the telson, not in-
cluding the posterior spines.

It seems probable that the specimen from
the southern tip of Baja California may be
identifiable as H. nesisi Burukovsky, 1986,
previously known only from the male ho-
lotype, with an apparent postorbital cara-
pace length of about 27 mm, found about
1600 kms southwest of the A/batross station
at 13°34'N, 120°33’W, on an apparent sea-
mount in 800 m in an area where total depths
are near 4300 m. Although the Hawaiian
specimen agrees with the one from Baja Cal-
ifornia and disagrees with H. alexandri in
the indistinct or nonexistent grooves on the
first and second abdominal tergites and the
absence of a dorsal boss on the third tergite,
it differs so clearly from H. nesisi in its much
smaller size, shorter rostrum, and much
longer sixth abdominal somite as to suggest
that-it represents an undescribed species.

There is little doubt that Crosnier (1986a:
368) had ample reason to remove H. /aevis
A. Milne-Edwards, 1883, from the genus
Heterocarpus and to transfer it and its Pa-
cific analogue Plesionika chacei Crosnier,
1986 (renamed P. fenneri Crosnier, 1986b),
to the genus Plesionika because of their sim-
ilarity to P. bifurca Alcock & Anderson,
1894, and P. spinidorsalis (Rathbun, 1906).
Similarly, the three species discussed above
probably belong to the same genus as the
four assigned to Plesionika by Crosnier
(1986a), because they lack strong postor-
bital or postantennal carinae and have the
carapace transversely convex rather than
carinate in the dorsal midline of the pos-
terior one-fourth or more of its length. I am
not proposing that change here, however,
because these seven species seem to form a
rather homogeneous group quite different
from Plesionika uniproducta Bate, 1888
(=Acanthephyra ensis A. Milne-Edwards,
1881), the type species of Plesionika. The

88 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

family Pandalidae displays cosiderable
morphological diversity, but some of the
genera are so obscurely differentiated as to
engender the possibility that a group of
species clustered around Heterocarpus lae-
vis and H. alexandri might eventually con-
stitute a valid separate genus. The danger
of prematurely accepting potential generic
characters in the family, however, 1s man-
ifested by the occurrence of a well-defined
boss on the tergum of the third abdominal
somite in those two Atlantic species and the
virtually complete absence of such a boss
in the Pacific species that are obviously
closely related to them: Plesionika fenneri,
Heterocarpus nesisi, and the Hawaiian spec-
imen assigned to H. alexandri by Rathbun
(1906).

Acknowledgments

But for the serendipity provided by Ardis
Johnston of the Museum of Comparative
Zoology in response to my inquiry about
the specimen identified by Faxon (1896:161)
as Heterocarpus alexandri, the holotype of
that species would have remained “‘lost”’ in-
definitely. Iam most grateful to her for that
welcome information and for making the

type specimen available to me. I also thank
Alain Crosnier of the Muséum national
d’Histoire naturelle in Paris and my Smith-
sonian colleague Brian Kensley for reading
the manuscript.

Literature Cited

Alcock, A., & A. R. Anderson. 1894. Natural history
notes from H. M. Indian Marine Survey Steam-
er ““Investigator,’’» Commander C. F. Oldham,
R.N., commanding, series 2(14); An account of
a recent collection of deep sea Crustacea from
the Bay of Bengal and Laccadive Sea.— Journal
of the Asiatic Society of Bengal. new series, 63:
141-185, pl. 9.

Bate, C. S. 1888. Report of the Crustacea Macrura
collected by H.M.S. Challenger during the years
1873-76.— Report on the scientific results of the
voyage of H.M.S. Challenger during the years
1873-76 24:i-xc, 1-942, figs. 1-76, 157 pl.

Burukovsky, R.N. 1986. [Anew shrimp species from
the genus Heterocarpus (Crustacea: Decapoda:
Pandalidae) and a brief review of the genus.]—
Byulleten Moskovskogo Obshchestva Ispytate-
ley Prirody Otdel Bioloogicheskiy 91(5):62-73,
figs. 1-4. [Russian with English summary.]

Chace, F. A., Jr. 1985. The caridean shrimps (Crus-
tacea: Decapoda) of the A/batross Philippine Ex-
pedition, 1907-1910, part 3: Families Thalas-
socarididae and Pandalidae.—Smithsonian
Contributions to Zoology 411:1—143, 62 fig.

Crosnier, A. 1986a. Crevettes de la famille des Pan-

dalidae récoltées durant ces derniéres années en

Polynésie francaise. Description de Plesionika

chacei et P. carsini spp. nov.—Bulletin du Mu-

seum National d’Histoire Naturelle, sér. 4, 8,

sect. A(2):361-377, figs. 1-4.

. 1986b. Plesionika fenneri, nouveau nom pour

Plesionika chacei Crosnier, 1986.— Bulletin du

Muséum National d’Histoire Naturelle, sér. 4,

8, sect. A(3):691.

de Man, J. G. 1920. The Decapoda of the Siboga
Expedition, IV: Families Pasiphaeidae, Stylo-
dactylidae, Hoplophoridae, Nematocarcinidae,
Thalassocaridae, Pandalidae, Psalidopodidae,
Gnathophyllidae, Processidae, Glyphocrango-
nidae and Crangonidae.—Siboga-Expeditie
Monographie 39a:1-318, pl. 1-25.

Faxon, W. 1896. Reports on the results of dredging,
under the supervision of Alexander Agassiz, in
the Gulf of Mexico and the Caribbean Sea, and
on the east coast of the United States 1877 to
1880, to the U.S. Coast Survey Steamer “Blake,”
Lieut.-Commander C. D. Sigsbee, U.S.N., and
Commander J. R. Bartlett, U.S.N., command-
ing. 37. Supplementary notes on the Crusta-
cea.— Bulletin of the Museum of Comparative
Zoology at Harvard College 30(3):153-166.

Milne-Edwards, A. 1881. Description de quelques

Crustacés Macroures provenant des grandes

profondeurs de la mer des Antilles. —Annales

des Science Naturelles, Zoologie, sér. 6, 11(4):

1-16.

1883. Recueil de figures de Crustacés nou-
veaux Ou peu connus, Paris, 3 pp, 44 pls.
Rathbun, M. J. 1906. The Brachyura and Macrura

of the Hawaiian Islands. — Bulletin of the United
States Fish Commission (1903) 23(3):827—930
[reprint with index pages i—vili], figs. 1-79, pl.
1-24.

Department of Invertebrate Zoology, Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 89-94

GAMMAROPSIS ARAWAKIA, A NEW SPECIES OF
MARINE AMPHIPODA (CRUSTACEA)
FROM JAMAICA

James Darwin Thomas and J. L. Barnard

Abstract.—One of the dominant amphipods in algal turf communities at
Discovery Bay, Jamaica, is a new species, Gammaropsis arawakia, character-
ized by long peduncle of uropod 3, small coxa 1, 3-articulate accessory flagellum,
extremely setose gnathopod 2, one pair of dorsal teeth on urosomite 1, pointed
ocular lobe, and oblique upper margin of cephalic sinus for antenna 2.

Gammaropsis arawakia is very abundant
in the algal turf that now (1986) covers the
heavily disturbed reef at Discovery Bay, Ja-
maica. The living coral was damaged by the
hurricane of 1980 and any regenerants al-
most completely exterminated by algal turf
in 1984-1986. This turf developed follow-
ing the death of the herbivorous urchins
Diadema antillarum (information from Di-
rector Dr. Jeremy Woodley). A nearby load-
ing dock for bauxite may also be a factor in
the disturbance, although we are informed
that the coral damage and turf development
are common to most of the north coast of
Jamaica. Discovery Bay is also character-
ized by the presence of large, cold fresh-
water seeps. Other amphipod genera pres-
ent in this turf mat are Lembos, Elasmopus
and Hyale.

Isaeidae
Gammaropsis arawakia, new species
Figs. 1-3

Etymology. —Named for the Arawak,
former Indian tribe of the central Carib-
bean.

Diagnosis. —Ocular lobe projecting mod-
erately, bearing sharp anterior cusp, lower
antennal sinus with oblique dorsal margin;
eyes of medium size, with irregular mass of
purple pigment surrounded sparsely by clear
ommatidia. Accessory flagellum 3-articu-

late, articles 1-2 long, apical article minute.
Epistome sharply produced into long an-
terior spike. Mandibular palp article 3 cla-
vate, with subdistal anterior notch, heavily
setose. Inner plate of maxilla 1 naked, outer
plate with 9 spines. Plates of maxilla 2 broad,
inner with oblique facial row of 6 setae.

Anterior coxae short, coxa | less than half
as wide as coxa 2, rounded-quadrate an-
teroventrally, coxa 3 narrower than coxa 2,
coxa 5 as long as coxa 4. Carpus of gnatho-
pod 1 longer than propodus, dactyl strongly
overreaching oblique palm. Article Z of
gnathopod 2 densely setose mediodistally,
carpus short, propodus greatly enlarged,
palm oblique, convex, defined by tooth
lacking spine, densely setose medially and
anteriorly, these setae projecting dorsally and
medially. Locking spines of pereopods 3—4
absent, of pereopods 5-7 forming unequal
pair. Pereopods 5-7 of similar structure but
increasingly longer, article 2 very narrow
and smooth posteriorly.

Epimera 1-3 similar, each with small pos-
teroventral tooth and setule-notch, poste-
rior margins weakly convex. Urosomite |
with pair of weak dorsal teeth side by side.
Uropod 1 with medium interramal tooth,
lacking on uropod 2; each ramus of uropods
1-2 with elongate apical spine. Peduncle of
uropod 3 elongate, rami lanceolate but al-
most rod-like, subequal to each other but
longer than peduncle, outer ramus with

90

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P|) \ ye

Ny L) OK
PRE

VOLUME 102, NUMBER 1

elongate apical spine-setae about as long as
peduncle of uropod 2. Telson ordinary.

Description of male “‘a’’ 2.09 mm. —Left
mandibular incisor with four teeth, lacinia
mobilis with four teeth, main rakers three.
Apex of palp on maxilla 1 with four spines,
one apicolateral spine-seta, 3 subdistal fa-
cial setae; inner plate short, softly triangu-
lar. Inner plate of maxilliped with 3 medial
setae, 7 apical setae, three stout apical spines,
one ventral facial locking spine and one
ventral seta; outer plate with four thick ser-
rate medial spines, two similar but thinner
apical spines and 2 apical setae, ventral sur-
face with 11 scattered setae; dactyl elongate,
nail of medium length, with three accessory
setules (Figs. 1-3).

Apices of rami on uropods 1-2 as in en-
larged illustration, with medial elongate
spine and short lateral spine, except inner
ramus of uropod 2 with long and short spines
reversed.

Description of female “‘b’” 2.47 mm.—
Oostegites 2 and 5 short, slender, with 2 and
6 setae each, oostegites 3 and 4 twice as long
and wide, with 10 setae each; gills 2-3 and
6-7 sac-like, slender, 3 times as long as
broad, gills 4 and 5 about 4 times as long
as broad, scarcely broader than others.

Holotype. —USNM No. 195175, male “‘a”’
2.09 mm.

Type locality.—Jamaica station 1B, Dis-
covery Bay, backreef, 1 m, 6 Sep 1986, on
Acropora palmata rubble mound near chan-
nel marker, formaldehyde wash of algal turf,
coll. J. D. Thomas and J. Clark.

Material.—Type locality, female “‘b’”’ 2.47
mm, female “‘c’? 2.14 mm, male ‘“‘d”’ 2.03
mm and 15 other specimens. Numerous

—
Fig. 1.

91

specimens from many other similar samples
also deposited in Smithsonian crustacean
collections.

Relationship. —There are 105 species of
Gammaropsis so that it was a laborious task
to distinguish our species from the score of
species that it resembles. To avoid a lengthy
discussion we have eliminated those species
not fitting significant parts in the following
diagnosis:

Ocular lobe pointed, short; eyes small;
antennae of medium length; coxae 1-5 short
and almost evenly extended; carpus of
gnathopod 1 long; carpus of gnathopod 2
short, propodus evenly rectangular, palm
shorter than posterior margin of propodus,
regular, poorly sculptured, defined, medial
surface of hand heavily setose; epimeron 3
with point; urosome with only one pair of
dorsal teeth; peduncle and rami of uropod
3 elongate, armaments of rami very sparse
and simple.

Gammaropsis (=Eurystheus, Kermys-
theus, Podoceropsis, Megamphopus, Se-
gamphopus, Pseudeurystheus, see Barnard
1973) contains a diversity of species not yet
organized cladistically. Note that Paranae-
nia Chilton (1884) is being revived by Bar-
nard & Karaman (1989). Gammaropsis ara-
wakia resembles Segamphopus and
Pseudeurystheus in the elongate carpus of
gnathopod 1 and such species as G. exser-
tipes and G. spinosa in the elongate peduncle
of uropod 3. It also resembles several species
with densely setose gnathopod 2 such as G.
setifera, G. tonichi, and G. tawahi. The nar-
row coxa | suggests certain species of Me-
gamphopus but most members of that “‘ge-
nus” have other anomalies in gnathopods

Gammaropsis arawakia, unattributed figures = holotype male “‘a”’ 2.09 mm; d = male “‘d” 2.03 mm.

Capital letters denote main parts in the following list; lower case letters to right of capital letters or in body of
figure indicate modifications as follows; lower case letters to left of capital letters indicate specimens described
in captions: B, body; D, dactyl of thoracic leg; E, prebuccal lateral; G, gnathopod; H, head; M, mandible; N,
mandibular molar; O, outer ramus; P, pereopod; R, uropod; S, maxilliped; T, telson; V, palp; W, pleon; X,

maxilla; r, right; s, setae removed; t, left.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

92

fj

“a”? 2.09 mm. Letter codes see Fig. 1.

Fig. 2. Gammaropsis arawakia, holotype male

VOLUME 102, NUMBER 1

93

Fig. 3. Gammaropsis arawakia, holotype male ‘“‘a” 2.09 mm. Letter codes see Fig. 1.

or other coxae. We have ignored species of
*“‘Podoceropsis”’ and “‘Kermystheus’”? which
have severely reduced accessory flagella.

Gammaropsis tawahi Barnard (1972),
from New Zealand, has an elongate pedun-
cle on uropod 3 and densely setose gnatho-
pod 2 but the palm bears a tooth near the
hinge, the apical armaments on uropods 1-
3 are all short, mandibular palp article 3
lacks a notch and the antennae are short-
ened and sparsely setose.

Our species is close to G. averus Reid
(1951) from West Africa, but differs in the
broader hand of gnathopod 2 and less sculp-
tured palm.

Our species differs: From G. sutherlandi
Nelson (1981), North Carolina, in the small
even coxae 1-5, strongly subchelate gnatho-
pod 2 and simpler spination of uropod 3;
From G. hirsuta Reid (1951), West Africa,
in the more sculptured palm and long prop-
odus of male gnathopod 2 and the longer
carpus of gnathopod 1; From G. tonichi Bar-
nard (1969), Pacific Mexico, in the presence
of only one pair of teeth on the urosome,

and the lack of radical transformations oc-
curring in male gnathopod 2 and coxa 7;
From G. spinosa Shoemaker (1942), Pacific
Mexico, in the small coxae 1—5 and medium
(versus very short) antennae; From G. chil-
toni and G. haswelli Thomson (1897), New
Zealand, in the sharp ocular lobes, small
coxae 5-6 and the ordinary, not grotesque,
male gnathopod 2; From the pan-tropical
G. atlantica Stebbing (1888) in the normal
and small eyes and longer carpus of gnatho-
pod 1; From G. thomsoni Stebbing (1888),
New Zealand, in the more simplified ar-
maments on the more elongate rami of uro-
pod 3; From G. exsertipes Stebbing (1888),
Kerguelen, in the ordinary condition of cox-
ae 1—5 (versus enlarged), male gnathopod 2
(versus grotesque or elephantine), epimeron
3, regular length of antennae and sharp ocu-
lar lobes; From G. anomala Chevreux
(1926), warm eastern Atlantic, in the nor-
mal gnathopod 2 and pereopod 5 which in
the Lusitanian species are respectively
weakly palmate and elephantine.

The following species also have many

94 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

similarities to ours but differ in two or more
of the characters in the diagnosis presented
just above: G. dilatata and G. pseudoden-
ticulata Ledoyer (1982) from Madagascar;
G. pacifica and G. setifera Schellenberg
(1938) from the middle Pacific Ocean; G.
crenulata and G. ulrici Krapp-Schickel &
Myers (1979) from the Mediterranean Sea;
G. palmata (Stebbing & Robertson, 1891)
from Europe; G. e/lisi Conlan (1983) from
cold-temperate eastern Pacific; and G. lo-
bata Chevreux (1920) from Europe.

Acknowledgments

We thank Janice Clark for her assistance
in the field in Jamaica and Kimberly R.
Cleary for her assistance in our laboratories
at the Smithsonian Institution. Dr. Jeremy
Woodley, Director of the Discovery Bay
Marine Laboratory, and his staff gave us
considerable help in our survey studies.

Literature Cited

Barnard, J. L. 1969. A biological survey of Bahia de
Los Angeles Gulf of California, Mexico, IV.
Benthic Amphipoda (Crustacea).—Transac-
tions of the San Diego Society of Natural His-

tory 15:175-228.

1972. The marine fauna of New Zealand:
Algae-living littoral Gammaridea (Crustacea
Amphipoda).—New Zealand Oceanographic
Institute Memoir 62:1-216.

1973. Revision of Corophiidae and related
families (Amphipoda).—Smithsonian Contri-
butions to Zoology 151:1-27.

, & G. S. Karaman. 1989. The families and
genera of Gammaridean Amphipoda (except
marine Gammaroids).—Smithsonian Contri-
butions to Zoology (In press).

Chevreux, E. 1920. Sur quelques amphipodes nou-

veaux OU peu connus provenant des cdtes de

Bretagne. — Bulletin de la Société Zoologique de

France 45:75-87.

1926. Amphipodes I.—Gammariens (suite).
Voyage de la goélette Melita aux Canaries et au
Senegal 1889-1890.—Bulletin de la Société
Zoologique de France 50:365-398.

Chilton, C. 1884. Additions to the sessile-eyed Crus-

tacea of New Zealand.—Transactions and Pro-
ceedings of the New Zealand Institute 16:249-
265.

Conlan, K. 1983. The amphipod superfamily Coro-
phioidea in the northeastern Pacific region 3.
Family Isaeidae: Systematics and distributional
ecology. — Publications in Natural Sciences, Na-
tional Museum of Natural Sciences, Canada 4:
1-75.

Krapp-Schickel, G., & A. A. Myers. 1979. The Med-
iterranean species of Gammaropsis Liljeborg
(Crustacea, Amphipoda).—Bollettino del Mu-
seo Civico di Storia Naturale, Verona 6:441-
467.

Ledoyer, M. 1982. Crustaces amphipodes gamma-
riens familles des Acanthonotozomatidae a
Gammaridae.—Faune de Madagascar 59(1):1—
598.

Nelson, W. 1981. A new species of the marine am-
phipod genus Gammaropsis from the south-
eastern United States (Photidae).— Proceedings
of the Biological Society of Washington 93:1223-
1229, 3 figs.

Reid, D. M. 1951. Report on the Amphipoda (Gam-
maridea and Caprellidea) of the coast of tropical
West Africa.—Atlantide Report 2:189-291.

Schellenberg, A. 1938. Litorale Amphipoden des tro-
pischen Pazifiks.—Kungliga Svenska Veten-
skapsakademiens Handlingar (3)16(6):105 pp.

Shoemaker, C. R. 1942. Amphipod crustaceans col-
lected on the Presidential Cruise of 1938.—
Smithsonian Miscellaneous Collections 101(1 1):
1-52.

Stebbing, T. R. R. 1888. Report on the Amphipoda
collected by H.M.S. Challenger during the years
1873-—76.—Report on the Scientific Results of
the Voyage of H.M.S. Challenger During the
Years 1873-1876, Zoology 29:xxiv and 1737
pp.

—, & D. Robertson. 1891. On four new British
Amphipoda.—Transactions of the Zoological
Society of London 13:31-42.

Thomson, G. M. 1897. On two new gammarids from
New Zealand.—Annals and Magazine of Nat-
ural History, ser. 6, 20:447-451.

(JDT) The Reef Foundation, P.O. Box
170, Big Pine Key, Florida 33043; (JLB)
Department of Invertebrate Zoology, Na-
tional Museum of Natural History, NHB-
163, Smithsonian Institution, Washington,
DCs 20560.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 95-105

NATATOLANA PASTOREI (GIAMBIAGI, 1925)
(CRUSTACEA, ISOPODA, CIROLANIDAE) FROM THE
STRAITS OF MAGELLAN, SOUTH AMERICA:
REDESCRIPTION AND NOTES ON
FUNCTIONAL MORPHOLOGY

Johann Wolfgang Wagele and Niel L. Bruce

Abstract. —Cirolanids caught with baited traps in the Straits of Magellan and
identified as Natatolana pastorei (Giambiagi) are described. Observations on
living specimens suggest that several features, such as the broadened merus of
the anterior pereopods, the arrangement of spines and setae are adaptations to

digging and respiration in sand.

Giambiagi (1925) described briefly from
“San Sebastian, Tierra del Fuega,” south of
the eastern entrance of the Straits of Ma-
gellan, a cirolanid named Cirolana pastorei.
No other figures than those published by
Giambiagi exist; they are not adequate to
distinguish this species from the many sim-
ilar cirolanids known today. Another ciro-
lanid from the Straits of Magellan was men-
tioned by Pfeffer (1887) in a list of species
collected during a German expedition to
South Georgia (1882-1883). Of this Ciro-
lana magellanica no illustrations or de-
scriptions were ever published. The name
was therefore listed as nomen nudum by
Bruce (1986).

During an expedition to Antarctica (1985—
1986) with R/V Polarstern a large number
of cirolanids were caught in a baited trap in
the Straits of Magellan. Most of these ani-
mals were kept in aquaria for bionomic
studies. Later the same species was found
in collections of the Zoologisches Museum
Hamburg, in vials labelled ““Cirolana ma-
gellanica, cf. types,” which possibly contain
the specimens studied by Pfeffer (1887).
Further material was kindly sent to the first
author by Dr. Gruner from the Zoologisches
Museum Berlin, also with the label “‘Ciro-
lana cf. magellanica.”’

The species clearly belongs to the genus

Natatolana Bruce, 1981 and is related to a
group of large species, the Natatolana valida
group, that have an exclusively southern
hemisphere distribution (Bruce 1986). The
species was compared to drawings of the
types of Natatolana rossi, N. hirtipes and all
subantarctic species and found to be dis-
tinct. The species here described is identical
with the Cirolana pastorei of Giambiagi
(1925); no differences from the original de-
scription or the type specimens could be
found. The species is not identical with the
Cirolana pastorei from South Georgia men-
tioned by Monod (1931).

Natatolana pastorei (Giambiag1)
Figs. 1-7

Cirolana pastorei Giambiagi, 1925:4—7, pl.
1, pl. 3, fig. 3, not Cirolana pastorei sensu
Monod, 1931:22.

Cirolana magellanica Pfeffer, 1887:58, no-
men nudum.—Bruce, 1986:218.

Material examined.—Male, length 33
mm, partly dissected, Magellan Straits,
52°17.49'S, 69°8.96'W, 35 m; 15 Nov 1984;
four adult specimens, 27-36 mm, from the
same locality, Zoological Museum Ham-
burg, K-33324. Further material: Zoologi-
cal Museum Hamburg, K-19219, Magellan

96

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

X)

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Q

ee ae SS Gon on en

",

>

p
1)

WHA,

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SA)

TE vv td Vr VXyt
.

\\\ Wh \|

S

Fig. 1. Natatolana pastorei: A, Male in dorsal view; B, Ventral aspect of cephalothorax; C, Posterior segments,

ventral. Arrow: respiratory current passing through filter of pereopods 5—7. Abbreviations.—A1, A2: antenna 1,2;
La: labrum; Md: mandible; MdP: palp of Md; Mx1, Mx2: maxilla 1,2; Mxp: maxilliped; PJ-7: pereopods 1-

7; Plp1-5: pleopods 1-5; Tel: telson; Urp: uropod.

mens, 13-15 mm, leg. Prof. Carlos Grethe,
Magellan Str., 19 Nov 1908; further mate-
rial collected by Kpt. Paessler: K-23268
(about 52°20’S, 69°W), K-23287 (51°54’S,

Straits, labelled ““Cirolana magellanica, cf.
types,” leg. Kpt. R. Paessler 1886: 44 spec-
imens, 13-29 mm, one male of 30.5 mm,
no mature females; also included: 5 speci-

VOLUME 102, NUMBER 1 97

___0.5mm__,

Fig. 2. Natatolana pastorei, male in lateral view; arrows: respiratory current passing through setal combs of
basipodites, antennule with detail of flagellar setation. For abbreviations see Fig. 1.

98 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

73°42'W), K-23281 (about 47°S, Golfo de
Penas), K-23278 (Hale Cove, Patagonia);
with number K-23242 two samples are de-
posited: leg. Dr. Gassmann, Magellan Str.,
1 Nov 1895, and leg. Umbach, off Punta
Arenas, Chile (53°10’S, 70°54’W), 1 Jan
1903.

Zoological Museum Berlin, Nr. 8118,
Magellan Str., leg. Pohl: three small speci-
mens (10-18 mm); one further specimen
was sent to the Copenhagen Museum, prob-
ably for H. J. Hansen.

Museo Argentino de Ciencias Naturales
‘“‘Bernardino Rivadavia,’’ Buenos Aires, Nr.
12690: nine syntypes, San Sebastian, Tierra
del Fuego.

Description. —Body about 2.5 times as
long as broad. Cephalothorax and eyes as
in Figs. 1 & 2; beneath eye a deep groove
for antennae. Coxae one to three with curved
carina running from frontal area of coxa to
posterolateral point; coxae four to seven with
more pronounced, less curved medial ca-
rina; coxa seven reaching beyond epimeral
plate of pleonite 1 (Fig. 2). Epimeral plate
of pleonite 3 narrower than preceding plate,
posteriorly acute. Outline of telson as in Fig.
7, distal margin with six to seven sensory
spines on each side and between them rows
of four to five setae.

Antennule peduncle of 4 articles (3 in the
sense of Wagele 1983), article 3 longest (Fig.
2); article 4 short, with three plumose setae;
flagellum not much longer than peduncle,
with 17 articles, articles short and broad,
with rows of aesthetascs and each 2nd or
3rd article with a group of short simple se-
tae, last article with simple setae and one
plumose seta (Fig. 2). Antenna peduncle ar-
ticle 2 very short, article 5 longest; fourth
article with lateral row of seven simple se-
tae; flagellum long, extending to pereonite
3, of about 30 articles; last articles more
slender and relatively longer than basal ar-
ticles (Fig. 3).

Mandible palp setation as in Fig. 3. Lat-
eral endite of maxillule gnathal surface with
13 stout spines, medial endite short, with

three stout apical teeth and subapically a
smaller, slender trifurcate spine (Fig. 2).
Maxilla of three lobes; proximal endite with
two rows of setae, first row with 11, second
with 17 setae; medial (second) endite with
15 long setae and a second row with 11
shorter setae; distal lobe bearing 5 setae.
Maxilliped endite with two coupling hooks;
all palp articles bearing large numbers of
simple setae (Fig. 4).

Pereopod 1 merus posterior margin with
14 acute spines; carpus posterodistal margin
with 2 long and 2 short acute spines: pro-
podal palm with 4 large acute and 5 small
acute spines and stout spine opposite base
of dactylus. Pereopods 2 and 3 similar to 1
but spines of merus and carpus larger, pe-
reopod propodal palm with 3 spines, is-
chium posterodistal margin with additional
spines. Pereopod 3 with only 2 small spines
on propodal palm. Pereopods 4—7 essen-
tially similar but basis widening and all ar-
ticles increasing in length towards posterior.
Pereopod 7 basis about 1.8 times as long as
greatest width, posterior margin shallowly
convex, anterior margin feebly sinuate; is-
chium, merus and carpus with clusters of
spines at distal angles and along posterior
margins; propodus with three groups of 2—
4 spines on posterior margin and a group
of 4 spines opposite base of dactylus.

Pleopods conform to the generic condi-
tion; pleopod 2 of male with appendix mas-
culina, apex narrowed, slightly angled,
slightly extending beyond apex of endopod.
Uropod endopod medial margin convex
with six spines and a further apical spine,
lateral margin with two large and two small
spines, apex narrow. Exopod lanceolate, lat-
eral margin with eight spines and a further
small apical spine, medial with four.

Variations of spine armature of tail fan
in adult specimens (25-36 mm): Distal mar-
gin of telson with 6 to 8 spines on each side
(generally 6), lateral margin of uropod ex-
opod with 9 to 11 (usually 10, including
apical spine), medial margin always with 4
(including apical spine of this side), lateral

VOLUME 102, NUMBER 1 99

Fig. 3. Natatolana pastorei, male: Last articles of antenna and apical part of 5th peduncular article (inset,
bottom) of immature adult; Right mandible (r. Md) with detail of setal lobe. For abbreviations see Fig. 1.

100 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Natatolana pastorei, male: Maxilliped with detail of endite; several setae and sensory spines cut off.
For abbreviations see Fig. 1.

VOLUME 102, NUMBER 1 101

XQ \

=S Ulta,

an
Ago rttege »
Set

Fig. 5. Natatolana pastorei, male: Pereopods 2-5, several setae and sensory spines cut off. For abbreviations
see Fig. 1.

102 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Yj
Y

MG

Wh

~——
—
_—-—~_— ——

Soe oe
——

\ é
\S
\ NY
Ys he
SY nh LZ
WY we”
. AY
at\
a \ fina
x
\ wus ‘
a fee >
\v vie OU
\ ne
y ®
VY ae
_ S=
A a
P AIL a ———
hid a =
~U
5 fez
Y
Take
Wa \ =—
ere ON) j Se
A \ g™ Wit iy f A

eee 227111 ar

Fig. 6. Natatolana pastorei, male: Several setae and sensory spines cut off; setae of pleopods shown as simple
setae; dashed line (Plp 2): range of setae. For abbreviations see Fig. 1.

VOLUME 102, NUMBER 1

Tel

, 225 mm _,

103

Urp Wepre at steal

Fig. 7. Natatolana pastorei, male: Setae of pleopods shown as simple setae, dashed line indicating length of
setae; detail of Plp 5 showing apex of endopod; detail of uropod showing a sensory spine. For abbreviations see

Fig. 1.

margin of endopod always with 4, medial
with 4 to 7 spines (including apical spine).

Color.—Purple brown, chromatophore
pattern shown in Fig. 1.

Size. —Immature adults 27-34 mm, males
30.5-—33 mm.

Remarks. —Natatolana pastorei belongs
to the N. valida group of species character-
ized by Bruce (1986). It can be distinguished

from all of those species listed by having a
slightly wider basis to pereopod 7, narrow
apices on the uropod rami, and also differ-
ences in the spination of the uropod rami.
Other large Natatolana species include those
found in the subantarctic region such as N.
meridionalis Hodgson, N. nitida Hale or N.
obtusata VanhOffen (these species were list-
ed by Bruce, 1986). All of these species are

104

characterized by large penial processes,
broad or elongate appendix masculina and
acute upturned posterolateral margins on
pleonite 4.

The present material differs from the ma-
terial discovered in South Georgia and men-
tioned by Monod (1931) as ““Cirolana pas-
torei.”’ An examination of Monod’s material
(collection of the Senckenberg Institute,
Frankfurt) revealed the following differ-
ences: Lateral margin of uropod endopod
with six spines (instead of four in N. pas-
torei); telson narrower; antennule shorter
with only 12 flagellar articles; groove be-
neath the eyes not present; eyes longer and
less wide (number of ommatidia in N. pas-
torei (17.5 mm): longitudinal line with 11-
12, vertical line with 12; in the specimen
from South Georgia: longitudinal line with
14-15, vertical line with 8-9). So the pop-
ulation from South Georgia must belong to
a different species.

Distribution. —Known from Straits of
Magellan and between Hale Cove, Argen-
tina to Punta Arenas, Chile, and from San
Sebastian, Argentina. Occurs in shallow
sublittoral sediments.

Observations on biology and functional
morphology. —About 20 specimens were
observed in aquaria kept at 1—2°C together
with other Antarctic isopods. At this low
temperature the animals were inactive for
many weeks, only sporadically feeding on
pieces of krill and fish. All specimens bur-
rowed in sand; when resting only the tips of
the antennae and sometimes the eyes could
be seen. In coarse sand burrowing is diffi-
cult, as the grains keep rolling back into the
groove, but in fine sand the specimens dis-
appear within 1—2 min. The sand is pushed
with the first pereopods laterally and cau-
dally, the broad merus with its setae being
a very effective broom; the posterior pe-
reopods push the sand caudally, their effec-
tiveness is increased due to the lateral spines
(Fig. 1, bottom). The pleopods help when
the animals dig fast; with their beat a strong
current sweeps away the grains loosened by

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

the pereopods. In a depth of 2—5 cm the
animals stop digging. After feeding the an-
imals can obviously remain inactive for
about two months; in aquaria the animals
were fed only once a week and mostly the
food was not consumed. Though the low
temperature affects the metabolism, in na-
ture this species is probably also able to sur-
vive several weeks without food, living as
an opportunistic carrion feeder.

Some isopods living in sand possess a
protective operculum to cover the respira-
tory pleopods (Chaetiliidae, Bathynatali-
idae, Serolidae, Anthuridae). The cirolanid
genera Conilera and Oncilorpheus also have
such structures, but operculate pleopods are
absent from other cirolanid genera. In Na-
tatolana the setae of the pereopods are ar-
ranged in such a way, that sand grains are
kept away from the thin pleopods. While
on the ventral side gravity will keep many
grains in their place, the sides of the animal
are protected against sand; the marginal se-
tae of the pereopods cover the slits between
the basipodites, where the respiratory cur-
rent passes (Fig. 2). Directly in front of the
pleopods, the setae of the frontal margins
of the basis and ischium of PS5—7 form a fan-
like filter, to intercept grains and detritus.

The pereopods are not used for swim-
ming, animals swim with their dorsum
downwards, propelled only by the beating
pleopods. The broadened merus of P1-3,
flattened basis with the marginal setae of
P5-7, the arrangement of the sensory spines
are adaptations to digging in fine sand.

Acknowledgments

We are grateful to Prof. G. Hartmann
(Hamburg), Dr. M. Turkay (Frankfurt), and
Prof. H. E. Gruner (Berlin, G.D.R.) for the
loan of specimens. Mrs. S. Willig kindly fed
and cultured the living animals. Mr. A.
Pompecki helped to prepare the plates.

This study was supported by the German
Science Foundation (DFG Nr. Wa-530) to
J.-W. Wagele, and a Queen’s Fellowship in
Marine Science to N. L. Bruce.

VOLUME 102, NUMBER 1

Literature Cited

Bruce, N. L. 1981. Cirolanidae (Crustacea: Isopoda)
of Australia: Diagnoses of Cirolana Leach, Me-
tacirolana Nierstrasz, Neocirolana Hale, An-
opsilana Paulian & Deboutteville, and three new
genera—Natatolana, Politolana and Carteto-

lana. — Australian Journal of Marine and Fresh-

water Research 32:945-966.

1986. Cirolanidae (Crustacea: Isopoda) of
Australia.— Records of the Australian Museum
Supplement 6:1—240.

Giambiagi, D. 1925. Resultados de la primera ex-
pedicion a Tierra del Fuego (1921). Crustaceos
IsoOpodos. — Anales de la Sociedad Cientifica Ar-
gentina, Buenos Aires 1925:229-245.

Monod, T. 1931. Tanaidacés et isopodes subantarc-

105

tiques de la collection Kohl-Larsen du Senck-
enberg Museum.—Senckenbergiana 13:10-30.

Pfeffer,G. 1887. Die Krebse von Siid-Georgien nach
der Ausbeute der Deutschen Station 1882/83.
I.—Jahrbuch der Hamburger Wissenschaftli-
chen Anstalten 4:41-150.

Wagele, J. W. 1983. On the homology of antennal
articles in Isopoda.—Crustaceana 45:3 1-37.

(JWW) Arbeitsgruppe Zoomorphologie,
Fachbereich 7, Universitat Oldenburg,
P.O.B. 2503, 2900 Oldenburg, FRG; (NLB)
Australian Museum, Division of Inverte-
brate Zoology, P.O.B. A285, Sydney South,
N.S.W. 2000, Australia.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 106-108

NEOBRACHIELLA ANISOTREMI
(COPEPODA: LERNAEOPODIDAE), A NEW SPECIES
PARASITIC ON AN INSHORE FISH,
ANISOTREMUS SCAPULARIS, OFF THE
CHILEAN COAST

Raul Castro Romero and Hernan Baeza Kuroki

Abstract. — Neobrachiella anisotremi, parasitic on Anisotremus scapularis, an
inshore fish of the northern coast of Chile, is described and illustrated. The
new species can be distinguished from all the species in the genus by a com-
bination of characters including second maxilla, trunk-cephalothorax length
ratio, and specially the posterior processes.

The existing literature contains very little
information on the genus Neobrachiella Ka-
bata, 1979, in Chilean waters. Only five
species have been recorded. Krgyer (1863)
gave a sketchy decription of Anchorella
(=Neobrachiella) appendiculata, without
recording its host. Ho (1983) described Neo-
brachiella amphipacifica, a species occur-
ring in both the North and South Pacific. In
the latter it was found on Psichrolutes sio
Nelson, a deep water fish, in northern Chile.
Castro & Baeza (1986) added to the list Neo-
brachiella exilis (Shiino, 1956) parasitic on
Mugil cephalus L., 1758; N. chevreuxii (van
Beneden, 1891), from Paralonchurus peru-
anus (Steindachner, 1785) and N. paralich-
hyos, Castro & Baeza, 1986. Recently we
decribed four other species collected from
sciaenid hosts from the Chilean coast (Cas-
tro & Baeza 1987).

In the course of our studies of the parasitic
copepods of Chilean fishes, we discovered
another species of Neobrachiella, parasitic
on an inshore fish, Anisotremus scapularis
(Tschudi). This species is described and il-
lustrated below. Terminology used is that
proposed by Kabata (1979).

Neobrachiella anisotremi, new species
Figs. 1-7

Material examined. —Four ovigerous fe-
males (8 Jan 1982); one female (12 Mar

1982) and three females (14 Jul 1982). Types
deposited in National Museum of Natural
History (USNM), Smithsonian Institution.
Holotype USNM 210517; Paratype USNM
210158.

Habitat.—Branchial arches of A. scapu-
laris, collected by beach seining.

Locality. —Isla Santa Maria, Antofagasta
(2327-5, FO225 Why:

Description. —Female (Fig. 1). Cephalo-
thorax subcylindrical, arching with slightly
expanded cephalic region, mean length 2636
um (range 2242-2908 um), width 455 wm
(range 364-545 um); dorsal shield well de-
limited, subrectangular. Trunk much longer
than wide, mean length 2909 um (range
2121-3636 um), width 1136 wm (range 909-
1333 um); its length varying from 75 to 123%
of that of cephalothorax; posterior extrem-
ity with two pairs of processes (Fig. 7A, B);
first pair ventral to egg sacs, mean length
736 um (range 606-805 um); its length 20—
38% that of trunk; second pair dorsal to egg
sacs and shorter, mean length 570 um (range
483-606); its length 73-83% that of first;
genital process of varying length, mean 186
um (range 113-145 um); poorly developed
in some specimens. Egg sacs subcylindrical,
multiseriate, mean length 2651 um (range
1818-3515 um); their length 71—112% that
of trunk.

First antenna (Fig. 2A, B) apparently 4—

VOLUME 102, NUMBER 1

107

Figs. 1-4. Neobrachiella anisotremi: 1, Female, lat-
eral view; 2A, First antenna; 2B, First antenna, detail
of apical armament; 3A, Second antenna; 3B, Second
antenna, detail of apical armament; 4, Mandible.

segmented; first and second segments in-
distinctly separated, second armed with
spine on distoventral margin; third short,
unarmed; fourth segment with apical ar-
mament as follows: tubercles 1 and 3 well
developed, 4a centrally situated, digitiform
seta; 6a single slender seta, and 5a complex,
consisting of a bifid seta and an additional
simple process; gibber absent. Second an-
tenna (Fig. 3A, B) strongly sclerotized, its
long axis running through sympod and ex-
opod; sympod bearing short robust spine on
its distoventral border; exopod bulbous,
apex spinulose on dorsal side; endopod
2-segmented, distal segment with apical spi-
niform process (reduced hook one, see Ka-
bata 1979) and two setae (spinulation on
ventral surface of distal segment can be seen
on distended cuticle in Fig. 3B). Mandible
(Fig. 4) with dental formula P1,P1,S1,P1,
S1,B3. First maxilla (Fig. 5) with papilli-

Figs. 5-7. Neobrachiella anisotremi: 5, First max-
illa, ventral; 6A, Maxilliped; 6B, Maxilliped, detail of
myxal area and claw; 7A, Posterior end of trunk, fe-
male; 7B, Lateral view of same, showing processes and
egg sacs.

form exopod situated ventrally, bearing
single seta; endopod armed with two sub-
cylindrical, setiferous papillae, ventral
papilla stouter than dorsal; no spinulation
observed. Second maxilla (Fig. 1) fused only
at tip, slender, less than half length of ceph-
alothorax; mean length 1051 wm (range 909-
1273 um); without conspicuous collars (bul-
la with short manubrium and anchor of nar-
row diameter).

Maxilliped (Fig. 6A, B) corpus strong,
slightly tapering, with spiniform process and
denticulate pad in myxal area; patch of spi-
nules near maxilla with subchela; shaft of
subchela with barb and spinulose pad on
distomedial part, claw gently curved, with
two accessory teeth.

Male. —Unknown.

Etymology. —The specific name anisotre-
mi, is derived from the generic name of the
host.

108

Discussion. —To establish the identity of
Neobrachiella anisotremi it must be com-
pared with all the species of the genus pos-
sessing two pairs of posterior processes. A
comparison within this group must be based
on a combination of gross morphological
characters (cephalothorax length; trunk
length; relationships between the length of
posterior process and other parts of the
body), since descriptions of the appendages
are not available for all species. Using these
criteria, the present species can be distin-
guished from all of them by a combination
of characters. It resembles most closely N.
gulosa (Wilson, 1915) but can be differen-
tiated from this species by the posterior pro-
cesses.

Wilson’s species has ventral processes as
long as the trunk, whereas in N. anisotremi
they are only 20-38% of the trunk length.
Slight differences exist also in the structure
of the second maxilla, that of N. gulosa, in
contrast to N. anisotremi, having a promi-
nent swelling accommodating the excretory
glands.

Another difference can be found in the
cephalothorax-trunk length ratio. In N. gu-
losa the cephalothorax is considerably long-
er than the trunk, whereas in anisotremi the
trunk is longer than the cephalothorax.

A unique feature of N. anisotremi is the
presence of a short spine on the distolateral
border of the sympod of the second antenna.

The four recently described species, para-
sitic on host fishes of the genus Sciaena (N.
oralis, N. auriculata, N. fasciata, and N. dis-
par), which are included in the same group
as the present one, can be differentiated by

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

a combination of characters that clearly dis-
tinguish between each one and N. anisotre-
mi. Their differences are given in the key to
the species of Neobrachiella (Castro & Baeza
1987).

Acknowledgments

The authors wish to thank Dr. Z. Kabata
for his review of the manuscript and spe-
cially for inviting the first author to spend
some time at the Pacific Biological station,
Nanaimo, B.C., Canada where this paper
was completed. They are also grateful to the
Universidad de Antofagasta for making the
journey to Canada possible.

Literature Cited

Castro, R., & Baeza, H. 1986. Some species of Neo-
brachiella Kabata, 1979 (Copepoda, Lernaeo-
podidae), parasitic on Chilean fishes, with de-
scription of Neobrachiella paralichthyos sp. nov.
from Paralichthys adspersus (Steindachner).—
Crustaceana 51(3):245-253.

——., & 1987. Four new species of Neo-
brachiella (Copepoda: Lernaeopodidae), para-
sitic on Sciaena genus (Teleostei: Sciaenidae) in
the South Pacific. — Estudios Oceanologicos 6(1):
1-24.

Ho, J. S. 1982. Copepod parasites of Psychrolutes
(Pisces: Scorpaeniformes) from deep waters in
the eastern Pacific.— Parasitology 85:451-458.

Kabata, Z. 1979. Parasitic copepoda of British fish-
es.— The Ray Society, London, No. 152:1-469.

Kroyer, H. 1863. Bidrag til Kundskab om snyltek-
rebsene. — Naturhistorisk Tiddsskrift, ser. 3(2):
75-426, pls. 1-18.

Universidad de Antofagasta, Instituto de
Investigaciones Oceanologicas, Casilla 170,
Antofagasta, Chile.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 109-115

THE FIRST ANNUAL RISER LECTURE:
MEIOFAUNA AND MICROBES— THE INTERACTIVE
RELATIONS OF ANNELID HOSTS WITH THEIR
SYMBIOTIC BACTERIA

Olav Giere

Abstract.—The gutless marine tubificid Phallodrilus leukodermatus (Oligo-
chaeta) harbors two kinds of chemoautotrophic sulfur bacteria beneath its
cuticle. The annelid hosts live predominantly around the redox potential dis-
continuity layer in calcareous sediments, providing their bacterial endosym-
bionts with access to both oxygen and reduced sulfur compounds. The bacteria
produce energy by oxidation of these sulfur compounds, and use this energy
for the fixation of CO, and production of carbohydrates. The worms utilize
the exudates of the bacteria and also ingest a portion of the bacterial population
in the epidermal cytoplasm. Further interactions in this apparently obligate
symbiosis are discussed. The transmittance of the bacteria through the host
generations by “infection” of freshly deposited eggs is correlated with a modified
reproductive biology. The studies on this symbiosis reveal a well-coordinated

balance between the annelid hosts and their extracellular bacterial partners.

Serendipity let us find a new species of
small (2-3 cm long) white annelid worms
in the calcareous sands of Bermudian reefs.
These tubificid oligochaetes, described as
Phallodrilus leukodermatus (Giere, 1979),
were unique among known annelids. They
lacked mouth, gut and anus and had no ex-
cretory organs. But the most striking feature
was a thick, subcuticular ‘mantle’ of extra-
cellular bacteria living between extensions
of the epidermal cells (Fig. 1).

This bacterial association, present in each
worm, even in the youngest, apparently

The Riser Lecture Series.—In 1985 the annual Riser
Lecture was initiated by members, alumni and friends
of the Marine Science Center, Northeastern University
at Nahant, Massachusetts. The occasion was the official
retirement of Professor Nathan W. Riser. As teacher,
biologist and founder of the facility, ‘““Pete’’ Riser en-
dowed the laboratory with a legacy—the importance
of considering the whole organism regardless of one’s
special focus. We dedicate these annual lectures to that
principle.

consists of two different gram-negative pro-
karyotes (Fig. 2), a smaller, rod-shaped form
(maximally 1.9 x 0.4 wm) with a thick cell
envelope, and a larger oval one (about 3 x
2 wm) characterized by a thinner body wall
and, more conspicuously, by the rich supply
of vesicles in its cytoplasm (Giere et al.
1984).

This peculiar bacterial layer raises ques-
tions about the nutrition of these worms and
about the structural implications of com-
plete reduction of the gut in worms from
sands that otherwise are richly populated by
other meiofauna, including oligochaetes with
normal guts. Also the microdistribution of
these exceptional worms is quite unusual
(Giere et al. 1982). We found rich popula-
tions in an area where, despite strong cur-
rents, anoxic horizons occurred at 5 cm sed-
iment depth. Only a few worms were present
in the upper, oxygenated strata, while the
bulk of the population always occurred in
the deeper layers where they were exposed
to considerable amounts of toxic hydrogen

110 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Doi am
7

Avance me

VOLUME 102, NUMBER 1

sulfide (around 100 uM S~-) with oxygen
present only in minimal concentrations.

What promotes this exceptional distri-
bution? How are the bacteria involved in
this scenario? The answers require integra-
tion of morphological studies with ecolog-
ical, physiological and microbiological
analyses, i.e., consideration of the whole
animal’s biology. These studies were per-
formed by a multidisciplinary team during
several research periods at the Bermuda Bi-
ological Station for Research and in mem-
bers’ respective home laboratories.

Structural and ecological similarities sug-
gest that the symbiotic prokaryotes are sul-
fur bacteria similar to those found in the
giant vestimentiferan Riftia (Cavanaugh et
al. 1981) from hydrothermal vents. The
bacteria obtain energy by oxidation of re-
duced-sulfur compounds, enabling them to
fix ambient CO, with subsequent produc-
tion of carbohydrates.

For Phallodrilus leukodermatus, confir-
mation of this pathway results from phys-
10logical analyses: Ribulosebiphosphate-
carboxylase in high activities in the bacteria
enables a notable incorporation of CO, with
a most rapid transfer into the malate pool
(Felbeck et al. 1983). The high activity of
enzymes like ATP-sulfurylase and sulfite
oxidase, involved in the sulfur metabolism,
indicates more specifically that sulfur com-
pounds become metabolized by the sym-
biotic bacteria. Moreover, it can be shown
that the products of this metabolism get
stored in the numerous bacterial vesicles:

—_—

Lit

Energy-dispersive X-ray microanalysis
demonstrates high sulfur peaks in mem-
brane-bound bacterial vesicles, suggesting
deposition of polysulfanes or polythionates
as easily available reserve products (Steudel
1985, Steudel et al. 1987).

Sulfur also is found in the internal tissues
of the worms where it accumulates in the
““chloragosomes,” annelid structures known
to store xenobiotic substances. Thus, sulfur
is transported from the ambient pore water
(perhaps via the symbiotic bacteria?) to the
internal worm tissues. Sulfate, the end prod-
uct of oxidative sulfur metabolism, is ex-
creted by the worms into the ambient sea
water (Giere et al. 1988).

Bacteria oxidizing sulfur compounds are
micro-aerophilic, gram-negative microor-
ganisms thriving in those horizons where
they have access to both reduced sulfur and
oxygen, 1.e., around the redox potential dis-
continuity layer. Microdistribution of the
worm population presents ecological con-
firmation of these sulfur-based chemoau-
totrophic processes in the annelid sym-
bionts. The bulk of the worm population is
always concentrated around this transition-
al zone between the oxygenated surface lay-
er and the anoxic depth with its poisonous
hydrogen sulfide. In this zone the prokary-
otic symbionts can synthesize their reserve
substances. In addition to reduced-sulfur
compounds, the bacterial cytoplasm
abounds with unwalled globules (Fig. 2) filled
with polyhydroxybutyric acid. Thus, in in-
cident light worms from this zone have well-

Figs. 1-3. Phallodrilus leuakodermatus: 1, Cross section through the gutless worm with “mantle” of subcu-
ticular bacteria, scale bar = 50 um; chl = chloragocytes, coe = coelomic cavity, cu = cuticle, ep = epidermis,
mu = musculature, ne = ventral nerve cord, scu = subcuticular space filled with bacteria; 2, The two kinds of
bacteria endosymbionts; note difference in thickness of cell envelope, scale bar = 0.5 um; Iba = large bacterium,
phb = vesicle filled with polyhydroxybutyric acid, sba = small bacterium, sve = sulfur vesicle; 3, Typical zonation
pattern of subcuticular bacteria with free, active bacteria in peripheral position and enclosed, lytic bacteria in
innermost position, scale bar = 5 wm; blys = bacteria in lysis, zba = zone of (free) bacteria, zint = intermediate
zone bacteria become enclosed by epidermal strands, zlys = zone of bacterial lysis: bacteria are enclosed in
vacuoles of epidermal cells and become digested (Fig. 1: light microscopy; Figs. 2+3: transmission electron
microscopy).

112

fed bacteria and appear shiny white due to
the rich content of light-refractile sub-
stances in the bacterial cytoplasm, which
can be almost 6% of the worm’s dry weight
(Giere et al. 1988). When kept in pure, ox-
ygenated seawater, the worms (through their
bacteria) continue to take up CO, for a while
from the environment, using up energy
stored in the reserve substances. However,
this uptake decreases soon, the worms be-
come (after about 24 h) transparent pale-
grayish, indicating rapid loss of their re-
serves, which in these “pale worms” is only
1.7% of their dry weight (Giere et al. 1988).

These results emphasize the biological in-
teractions between the oligochaete host and
its prokaryotic symbionts and lead to ques-
tions for the biological function of the bac-
teria/host system. The presence of bacteria
in all members of the population and the
ubiquity of bacterial incorporation in gut-
less oligochaete species—several species
mainly from the Caribbean, but also from
Australian reefs have been examined—sug-
gests an obligate symbiosis with mutual
benefits.

What are the benefits to the microorgan-
isms of this incorporation into the worms’
bodies?

-In a changing sedimentary system with
steep, overlapping gradients of oxygen, sul-
fide, thiosulfate and CO,, the highly motile
worms are able to quickly adjust position
relative to environmental fluctuations,
maintaining an optimal combination of
gradients required for growth of sulfur-ox-
idizing bacteria.

—It is conceivable that the prokaryotes take
up organic substances excreted by their host
which, in turn, has lost its excretory or-
gans. The highly vesiculate epidermal cy-
toplasm of the worms and the multitude
of membrane-bound secretory granules
found extracellularly in their subcuticular
lumen suggest this physiological pathway.

—The favorable position of the bacteria un-
derneath the cuticle affords both shelter
and easy access to nutrients from the am-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

bient pore water through the highly perme-
able cuticle.

On the other hand, what benefits might
the oligochaetes derive from close associa-
tion with bacteria?

—Since the bacterial oxidation of reduced
sulfur combines production of energy with
detoxification of poisonous hydrogen sul-
fide, the microorganisms enhance the
chances of the worms to invade a biotope
of high ecological potential, rich in dis-
solved nutrients, but with little competi-
tion or predation. Our physiological ex-
periments demonstrate that the worms can
actively and selectively take up dissolved
hexoses (particularly glucose and galac-
tose) and acidic free amino acids (partic-
ularly glutamic acid and aspartate). These
purely transcuticular absorption processes
seem so intensive that, compared to cor-
responding figures from pogonophorans of
the same size, they “contribute signifi-
cantly” to the nutrition of P. leukoder-
matus (Liebezeit et al. 1983, Liebezeit, pers.
comm.).

—Exuded bacterial products could serve as
a valuable and ever-present source of fur-
ther dissolved organic matter.

—Digestion of a certain portion of the inter-
nal bacterial population would provide an
additional trophic pathway. Comparison
of bacteria in the deeper layer of the epi-
dermis with those in the subcuticular areas
reveals a zonation pattern (Fig. 3): Periph-
eral bacteria are extracellular in a subcu-
ticular lumen, where they accumulate re-
serve substances and multiply intensively
by division. In the middle zone they tend
to be surrounded by strands of the epider-
mal cells. the innermost bacteria are al-
ways found enclosed in intracellular vac-
uoles, apparently “‘engulfed’’ by the
epidermal cells, and clearly undergoing lysis
(Giere and Langheld 1987).

This coordinated system, combining bac-
terial growth and multiplication with lysis
and digestion, is similar to those described

VOLUME 102, NUMBER 1

Lh3

Figs. 4-5. Phallodrilus leukodermatus: 4, Cross section through genital pad in clitellar region, scale bar =
50 um; cl = clitellar glands in epidermis, gepa = genital pad filled with bacteria, ne = ventral nerve cord; 5,
Cross section through young embryo with some free subcuticular bacteria (both kinds) and many lytic bacteria
enclosed in host cells, scale bar = 1 um; balys = bacteria enclosed in embryonic cells, mostly in lysis, foa = free
bacteria in peripheral layer (Fig. 4: light microscopy; Fig. 5: transmission electron microscopy).

for some thyasirid bivalves (Southward
1986), for the vestimentiferans Riftia
pachyptila (Bosch & Grassé 1984a, b) and
Tevnia sp. (Bassaglia 1986) and is compa-
rable to the processes found in the root nod-
ules of Leguminosae (Sprent 1979).

Thus, it seems that by their symbiosis
with sulfur-oxidizing bacteria the worms
have acquired a tripartite nutritional basis
via transepidermal uptake of dissolved or-
ganic matter, utilization of bacterial meta-
bolic products, and digestion of bacterial
cells. This flexible trophic pattern seems to
allow for the complete loss of digestive and
excretory organs. Moreover, the worms
gained access to a new habitat, the sulfidic
sediments. Examination of four additional
species within the approximately 40 species
of gutless marine tubificids showed that in
each case associations with subcuticular
bacteria are established, suggesting that this
symbiosis is geographically as ubiquitous as
it is evolutionarily favorable.

An intricate, mutualistic and well-estab-
lished symbiosis as seen in P. leukoder-
matus might be predicted to have evolved
a fairly secure method of transmitting the
bacterial partners through the generations
of their hosts rather than depending on more
or less randomly repeated de-novo-infec-
tions of the worms. A regular transmission
would require specific adaptative trends in
the reproductive biology of these oligo-
chaetes (Giere & Langheld 1987).

In fact, investigation of fully mature
worms reveals formation of a ventral “‘gen-
ital pad’’ (Fig. 4), a blister-like structure
packed with masses of fully active bacteria
ensheathed only by a thin cuticular enve-
lope. In a few cases deposition of single eggs
was observed. They are covered by a sticky
mucus, but not enclosed in a cocoon as
would be typical for oligochaetes. These eggs
were already infected by the characteristic
bacteria. These first bacteria are mostly in-
tracellular, within the cleavage cells, and

114

clearly in a lytic stage (Fig. 5). Only a few
peripheral bacteria maintain their subcutic-
ular, extracellular position and have a
healthy appearance. Probably the latter stock
is the source of the rich supply of extracel-
lular, active prokaryotes in adult worms. In
fortuitous sections of freshly laid eggs, still
adhering to the body, the bacteria appear to
be just about to enter the egg membrane
through deep invaginations. Since we found
free bacteria only during egg deposition and
the earliest developmental stages, we con-
clude that prokaryotic symbionts are re-
leased by rupture or, more probably, by lysis
of the genital pad adjacent to the oviducal
pore. Subsequently, they are transmitted by
invasion of the freshly deposited egg.

Although some microbiological and zoo-
logical details are not yet fully understood,
we envisage the following scenario: While
in early stages a defense mechanism initially
causes intracellular lysis of the bacterial-in-
truders, a balance between production of
extracellular bacteria and their intracellular
digestion becomes established in the grow-
ing worms, resulting in a coordinated sym-
biotic relationship between these aberrant
marine tubificids and chemoautotrophic
sulfur bacteria.

Acknowledgments

As pointed out, this lecture is based on a
series of studies performed by a group of
colleagues from various institutes and dis-
ciplines. I owe sincere thanks for intensive
collaboration and stimulating discussions
to R. Dawson, now Chesapeake Biological
Laboratory, University of Maryland, Sol-
omons, Maryland, USA; H. Felbeck, Scripps
Institution of Oceanography, La Jolla, Cal-
ifornia, USA; H. Jannasch and C. Wirsen,
both Woods Hole Oceanographic Institu-
tion, Woods Hole, Massachusetts, USA; C.
Langheld, G. Liebezeit and C. Schmidt, all
University of Hamburg, Hamburg, FRG.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Literature Cited

Bassaglia, Y. 1986. Le trophosome d’un Vestimen-
tifére. Structure et physiologie d’une symbiose.
— Université Paul & Marie Curie, D.E.A. d’O-
cénologie Biologique Paris, 42 pp. + 18 pp. of
appendices and bibliography + 8 pls.

Bosch, C., & P.-P. Grassé. 1984a. Cycle partiel des
bactéries chimiautotrophes symbiotiques et leurs
rapports avec les bactériocytes chez Riftia
pachyptila Jones (Pogonophore Vestimentifére).

I. Le trophosome et les bactériocytes. Comptes

Rendus de l’Académie des Sciences, Paris 299

(sér. 3):371-376.

1984b. Cycle partiel des bactéries chimiau-
totrophes symbiotiques et leurs rapports avec
les bactériocytes chez Riftia pachyptila (Pogono-
phore Vestimentifére). II. L’évolution des bac-
téries symbiotiques et des bactériocytes.—

Comptes Rendus de l’Académie des Sciences,

Paris 299 (sér. 3):413-419.

Cavanaugh, C. M., S. L. Gardiner, M. L. Jones, H. W.
Jannasch, & J. B. Waterbury. 1981. Prokary-
otic cells in the hydrothermal vent tube worm
Riftia pachyptila Jones: Possible chemoauto-
trophic symbionts.—Science 213 (4505):340-
34)

_ Felbeck,.H., G. Liebezeit, R. Dawson, & O. Giere.

1983. CO, fixation in tissues of marine oligo-
chaetes (Phallodrilus leukodermatus and P.
planus) containing symbiotic, chemoautotro-
phic bacteria.— Marine Biology 75:187-191.

Giere, O. 1979. Studies on marine Oligochaeta from
Bermuda, with emphasis on new Phallodrilus-
species (Tubificidae).— Cahiers de Biologie Ma-
rine 20:301-314.

— ; H. Felbeck, R. Dawson, & G. Liebezeit. 1984.
The gutless oligochaete Phallodrilus leukoder-
matus Giere, a tubificid of structural, ecological
and physiological relevance.— Hydrobiologia
115:83-89.

—,&C. Langheld. 1987. Structural organisation,
transfer and biological fate of endosymbiotic
bacteria in gutless oligochaetes.— Marine Biol-
ogy 93:641-650.

— ., G. Liebezeit, & R. Dawson. 1982. Habitat
conditions and distribution pattern of the gut-
less oligochaete Phallodrilus leukodermatus. —
Marine Ecology Progress Series 8:291-299.

—., C. O. Wirsen, C. Schmidt, & H. W. Jannasch.
1988. Contrasting effects of sulfide and thio-
sulfate on symbiotic CO,-assimilation of Phal-
lodrilus leukodermatus (Annelida). — Marine Bi-
ology 97:413-419.

Liebezeit, G., H. Felbeck, R. Dawson, & O. Giere.
1983. Transepidermal uptake of dissolved car-

VOLUME 102, NUMBER 1

bohydrates by the gutless marine oligochaete
Phallodrilus leukodermatus (Annelida).—
Océanis 9:205-211.

Southward, E. C. 1986. Gill symbionts in thyasirid
and other bivalve molluscs. — Journal of the Ma-
rine Biological Association of the United King-
dom 66:889-914.

Sprent, J. I. 1979. The biology of nitrogen-fixing or-
ganisms. McGraw-Hill Book Company, Lon-
don, 196 pp.

Steudel, R. 1985. Neue Entwicklungen in der Chemie
des Schwefels und des Selens.— Nova Acta Leo-
poldina, N.F. 59, No. 264:231-246.

115

—,G. Holdt, T. Gobel, & W. Hazeu. 1987. Chro-
matographische Trennung hoherer Polythionate
S,0O,2> (n = 3... 22) und deren Nachweis in
Kulturen von Thiobacillus ferrooxidans; molek-
ulare Zusammensetzung bakterieller Schwefe-
lausscheidungen. — Angewandte Chemie 99:143-
146.

Zoologisches Institut und Zoologisches
Museum, Universitat Hamburg, Martin-
Luther-King-Platz 3, D-2000 Hamburg 13,
Federal Republic of Germany.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 116-123

REVISION OF THE FAMILY LYSARETIDAE, AND
RECOGNITION OF THE FAMILY
OENONIDAE KINBERG, 1865
(EUNICIDA: POLYCHAETA)

G. Kent Colbath

Abstract.— As recognized by previous authors, the family Lysaretidae Kin-
berg, 1865 includes genera with dissimilar jaw morphology and composition.
Lysaretidae is restricted here to include only Lysarete Kinberg, characterized
by jaws mineralized with calcite and relatively short carriers with transverse
musculature. Oenone Savigny, Halla Costa and Tainokia Knox & Green are
removed into the family Oenonidae Kinberg, 1865. These genera are charac-
terized by heavily sclerotized jaws which are not mineralized, and by long
carriers with oblique-longitudinal musculature.

Hartman (1944) published a family-level
revision of the eunicoid polychaetes which
was widely accepted. Drawing on the work
of Kinberg (1865) and Ehlers (1868), she
placed heavy emphasis on the configuration
of the jaws as a taxonomic character.

Hartman recognized three basic maxil-
lary jaw patterns within the superfamily Eu-
nicea: the labidognath pattern ascribed to
the Eunicidae, Onuphidae, and Lumbriner-
idae; the prionognath pattern in the
Arabellidae and Lysaretidae; and a third
pattern (later named ctenognath by Kielan-
Jaworowska 1966) found only in the Dor-
villeidae. Labidognath jaws are character-
ized by having two relatively short, broad
carriers, in contrast to the two long, slender
carriers with a ventral median ligament (or
“third carrier’’) which characterize priono-
gnath jaw apparatuses. In the ctenognath
condition the maxillae consist of two to four
rows of numerous separate denticles.

During a study of the composition and
fossilization potential of polychaete jaws, I
divided eunicoid jaws into three groups
based on their composition (Colbath 1986).
Labidognath jaws are mineralized with cal-
cium carbonate in the form of aragonite in
the Onuphidae and Eunicidae, and calcite
in the Lumbrineridae. Labidognath jaws are

also weakly sclerotized, and the outer pro-
tein wall is readily leached in 2% KOH
(Hartmann-Schroder 1967, Colbath 1986).
In contrast, prionognath and ctenognath
jaws are amorphous to x-rays, do not react
with HCl, and are relatively resistant to
leaching in KOH (Colbath 1986).

The jaws of species of Lysarete Kinberg,
1865 are mineralized with calcite (Colbath
1986), and exhibit other labidognath mor-
phological characters. Other genera previ-
ously included within the Lysaretidae have
jaws of the prionognath type, prompting a
revision of the family, and recognition of
the family Oenonidae Kinberg, 1865.

History of Study

Early classifications of the eunicoid poly-
chaetes were based exclusively on features
of the external anatomy, and produced ar-
bitrary groupings of genera which are dis-
similar internally (Hartman 1944:1). First
to emphasize the importance of jaw parts
in classification was Kinberg (1865). Kin-
berg recognized ten families which he di-
vided into four major groups (labeled in-
formally as A-D). He defined group C by
the presence of mandibles with separated
right and left halves (an ontogenetically
variable feature), and included only La-

VOLUME 102, NUMBER 1

randa Kinberg, 1865, which was considered
unrecognizable by Hartman (1944:10, 1948:
98). Kinberg’s other groups correspond to
the labidognath (A), prionognath (B), and
ctenognath (D) jaw types presently recog-
nized. Kinberg considered the Lysaretae (sic)
and Oenonidea (sic) distinct, including the
former in group A and the latter in group B.

Ehlers (1868:280—282) also emphasized
the importance of jaw configuration in clas-
sifying these polychaetes, but did not for-
malize the distinctions. Instead, he recog-
nized a single family Eunicea (sic), which
he informally divided into the Eunicea lab-
idognatha and Eunicea prionognatha. Eh-
lers (1868) did not stress the morphology of
maxillary carriers as strongly as did Kin-
berg, but rather subdivided the eunicoids
according to whether the anterior maxillae
are arranged in a semicircle (labidognatha)
or in parallel rows (prionognatha).

In contrast to Kinberg, Ehlers (1868)
placed the genus Lysarete close to Aglau-
rides Ehlers (~Oenone Savigny) in the
prionognath subdivision. He later (Ehlers
1887:107-108) acknowledged the close
similarity between the jaws of Lysarete and
those of Lumbriconereis Grube (=Lumbri-
neris Blainville), but did not emend his ear-
lier taxonomic treatment.

Hartman (1944:2) essentially formalized
the 1868 scheme of Ehlers using the family-
level nomenclature of Kinberg. She retained
six of the ten families recognized by Kin-
berg, placing the Oenonidae into synonymy
with the Lysaretidae.

Fauchald (1970:118) removed Iphitime
Marenzeller from the Lysaretidae, and
erected the family Iphitimidae. He later
(Fauchald 1977) erected the order Eunicida
to include the seven families within the su-
perfamily Eunicea, plus two additional fam-
ilies (Histriobdellidae, Ichthyotomidae)
which were not assigned to a superfamily.

Materials and Methods

All lysaretids in the collection of the U.S.
Museum of Natural History (USNM) were

| GF

examined for the present study, and selected
specimens were dissected for examination
of the jaws. Additional specimens of Ly-
sarete were obtained on loan from the Flor-
ida Department of Natural Resources
(FSBC). The holotype of L. brasiliensis Kin-
berg, 1865 was obtained from the Swedish
National Museum, Stockholm.

One specimen of Lysarete brasiliensis with
jaws in excellent condition (USNM 55884)
was used for the chemical and mineralogical
analyses, and for scanning electron micros-
copy. The maxillary apparatus was re-
moved, dehydrated in an alcohol series, and
prepared for SEM by critical point drying.
The specimen was initially coated with car-
bon only, and chemically analyzed using
x-ray dispersion. The specimen was then
coated with gold palladium for routine scan-
ning electron microscopy. The mandibles
were removed, photographed with a Wild
400 wide-field optical microscope, and ana-
lyzed with the Gandalfi x-ray spectrometer
in the Department of Mineralogy, Smith-
sonian Institution.

Taxonomic Characters

Jaw morphology and composition. — X-ray
dispersion analysis of the maxillae of Ly-
sarete brasiliensis indicates the presence of
abundant calcium carbonate. The mandi-
bles are mineralized with calcite, as are jaws
in the lumbrinerids (Colbath 1986). Calci-
fication is also evident in L. australiensis
Benham, 1915, in that the blades of the
mandibles are white, and the maxillae are
dark brown rather than black. The presence
of calcite is inferred by comparison with L.
brasiliensis.

In contrast to Lysarete, jaws in species of
Oenone Savigny, Halla Costa and Tainokia
Knox & Green (commonly included in the
Lysaretidae) are not mineralized, but rather
are composed almost exclusively of sclero-
protein. As a consequence, both maxillae
and mandibles are jet black in color. The
distinction between heavily sclerotized and
calcified jaws is apparent upon inspection,

118 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

and was described by Benham (1915:233,
236-237). Benham made no comment about
the utility of this distinction above the
species level, however.

Other aspects of jaw morphology are cor-
related with the difference in mineraliza-
tion. In Lysarete the maxillary carriers are
relatively short (approximately equal to the
length of the maxillae) and broad (Fig. 1b),
with no ventral ligament (=“‘third carrier’).
A thick band of transverse muscles joins the
carriers dorsally (Fig. 2f), allowing them to
be pulled closed in the “trap”? motion de-
scribed by Wolf (1980) for other labido-
gnath jaws. The MI (maxilla I) elements in
the maxillary apparatus are the same length
(Fig. 2c), which allows the anterior tips to
come together in a pincer-like fashion as
part of the trap motion. The posterior dorsal
surfaces of the MIs exhibit an interlocking
ridge and furrow system (Fig. 2e), which
may help to lock the apparatus in a closed
position (Wolf 1980).

In species of Oenone, Halla and Tainokia
the carriers are narrow and much longer than
the combined length of the maxillae, and
are not equiped with a transverse muscle
band. Instead, a series of long, oblique-lon-
gitudinal muscles are attached along the
length of the carriers. A ventral ligament is
present. The right and left MI elements may
be of different lengths (Crossland 1924;
Hartman 1944; Kielan-Jaworowska 1966),
and no basal ridge and furrow system is
present to lock the jaws in a closed position.
Such jaws are presumably incapable of per-
forming a trap motion, as Wolf (1980) also
concluded for the arabellids.

Soft-part morphology.—The genera tra-
ditionally included within the Lysaretidae
share an elongated post-setal lobe and
prominent dorsal cirrus on posterior para-
podia. The morphology of the head, how-
ever, is distinctly different in each of the
four genera under discussion. Although the
difference between Lysarete and the other
genera may not merit consideration as a
family-level character, the anterior mor-

phology is none-the-less a useful key char-
acter.

In Lysarete, three antennae emerge from
the posterior margin of the prostomium, and
may lie flat in a V-shaped notch (Fig. 1a).
In adult specimens the notch extends pos-
teriorly across the peristomial rings, and in
some cases across the first one or two seti-
gers. Halla has three antennae which may
lie back in a semicircular notch in the two
peristomial rings (Fig. le). Tainokia has a
single antennae, and a slight invagination
which extends posteriorly only partially
across the first of two peristomial rings
(Knox & Green 1972, fig. 1). In Oenone,
three antennae are tucked into a nuchal fold
(Fig. 1d), no notch is present, and dorsally
the peristomium has only one distinct ring.

Setae and acicula.—All of the worms
considered here have simple geniculate se-
tae. In addition, bifid hooks have been re-
ported in Oenone and Halla. In Lysarete,
Oenone, and Tainokia the geniculate setae
are smooth (Knox & Green 1972, figs. 6, 7;
pers. observ.). In Halla, however, the dorsal
setae are denticulate, and the ventral setae
are marked by distinctive oblique striae (Fig.
2a, b; Fauvel 1923:427, fig. 169 g, h).

Both described species of Lysarete have
black acicula. In Oenone fulgida (Savigny,
1818) and Halla parthenopeia (Chiaje, 1828)
the acicula are amber, while Knox & Green
(1972:433) described the acicula in Taino-
kia iridescens as colorless. Black acicula are
thus useful for distinguishing Lysarete from
the other three genera, supporting the fam-
ily-level distinction based on jaw morphol-

ogy.

Systematics

Order Eunicida Fauchald, 1977
Family Lysaretidae Kinberg, 1865

Emended diagnosis. —Eunicoid poly-
chaetes with jaws mineralized with calcite.
Mandibles flat with semicircular growth
rings anteriorly (Fig. 2d). Maxillary carriers
no longer than combined length of maxillae,

VOLUME 102, NUMBER 1

119

Fig. 1.

a-c, Lysarete brasiliensis, holotype, Brazil; a, Dorsal view, anterior end, illustrating V-shaped notch

in peristomium; b, Dorsal view, MI elements and carriers, muscle attachment stippled; c, Labial view, MIIL-
MVL. d, Oenone fulgida, USNM 17729, Dry Tortugas, dorsal view, anterior end, note tips of antennae protruding
from nuchal fold. e, Halla parthenopeia, USNM 5147, Bay of Naples, dorsal view, anterior end, note semicircular

notch in peristomium. All scale bars = 1 mm.

articulated by prominent transverse mus-
cular bundle, no ventral ligament. Maxillary
jaws symmetrical. MI elements of identical
length, distally falcate with prominent sec-
ondary fang, additional small denticles may
be present at base. Three short antennae
arise from base of prostomium. Elongate
post-setal lobe and flattened dorsal cirrus
(or notopodium) developed in posterior
parapodia. Simple geniculate setae present,
bifid hooks absent.

Comparison.—The Lysaretidae are dis-
tinguished from the Oenonidae based on

jaw morphology and mineralogy. Lysarete,
the only genus presently recognized within
the family, is further distinguished from the
Oenonidae in having black acicula, and a
V-shaped notch in the dorsal surface of the
peristomium.

The Lysaretidae are similar to the Lum-
brineridae in having: 1) jaws mineralized
with calcite; 2) flat mandibles with a semi-
circular blade; 3) symmetrical maxillary
jaws; 4) transverse musculature on the car-
riers. The Lysaretidae are distinguished from
the Lumbrineridae by: 1) well-developed

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. a, b, Halla parthenopeia, USNM 5147, Bay of Naples; a, SEM dentate setae from dorsal bundle,
posterior parapodium, scale bar = 10 um; b, SEM striate setae from ventral bundle, scale bar = 50 um. c-g,
Lysarete brasiliensis, USNM 55884, Gulf of Mexico; c, SEM maxillary apparatus, dorsal view, scale bar = 1
mm; d, Light photomicrograph mandibles, ventral view, scale bar = 1 mm; e, SEM MI elements, oblique-lateral
view, note ridges and grooves on bases of MIs, scale bar = 0.5 mm; f, SEM carriers, oblique-lateral view, carriers
canted at an angle to MIs, note transverse muscle band, scale bar = 0.2 mm; g, SEM anterior maxillary jaws,

oblique-lateral view, scale bar = 0.2 mm.

dorsal cirri; 2) a prominent secondary fang
on the MI elements of the maxillae; 3) the
absence of hooded hooks. Kuwaita Moham-
mad, 1973 is intermediate between the Ly-
saretidae and Lumbrineridae in having an-

tennae on the prostomium, but clearly
belongs in the Lumbrineridae based on the
3 criteria above (Mohammad 1973:34, 36,
figs. 6, 7).

Distribution.—Lysarete includes one

VOLUME 102, NUMBER 1

species recorded from the east coast of North
and South America in low latitudes, and a
second species from a single locality east of
Australia. Larymna Kinberg, 1865 may also
belong in the Lysaretidae based on jaw mor-
phology (Hartman 1948:99). The type and
only specimen (from Mozambique) is so
poorly preserved that, according to Hart-
man, attribution cannot be made below the
family level, and Larymna should be re-
garded as unrecognizable.

Genus Lysarete Kinberg, 1865

Type species.—L. brasiliensis Kinberg,
1865:570.

Lysarete brasiliensis Kinberg, 1865
Figs. la—c, 2c-g

Lysarete brasiliensis Kinberg 1865:570;
1910:49, pl. 17, fig. 30.—Ehlers 1887:107-
108, pl. 33, figs. 1-8.—Rioja 1944:130-
131, figs. 47-50.—Hartman 1951:65, pl.
14, figs. 7, 8.—Orensanz 1975:106-108,
fig. 8.— Gardiner 1976:213, figs. 29b-e. —
Gilbert 1984:43-6—43-7, figs. 43-3, 43-4.

?Oenone brevimaxillata. —Treadwell 1931:
1-3, figs. 4-8.

Remarks.—In her re-appraisal of Kin-
berg’s type material, Hartman (1948:8) did
not consider the holotype of Lysarete bra-
siliensis, but instead referred to an earlier
revision by Ehlers (1887:107-108, pl. 33).
Ehlers did not figure the Brazilian type of
L. brasiliensis, but rather figured a specimen
collected from Florida. The posthumous
publication of Kinberg’s plates (Kinberg
1910) did not include illustrations of the
jaws of the holotype. The maxillae are there-
fore illustrated here (Fig. 1b, c).

On the holotype, maxillary jaws III and
IV each have a prominent anterior denticle,
with two progressively smaller denticles
posteriorly. The jaws in a specimen from
Argentina (Rioja 1944, fig. 50) are similar,
except that the MIII elements have four
denticles instead of three.

Most specimens examined from the east

121

coast of the United States have five to six
denticles on the MIII and four to seven den-
ticles on the MIV elements (Fig. 2g). The
anterior three denticles on the MIII element
are sub-equal in length, unlike those of the
type.

One specimen from Florida (FSBC I
31008) has MIII jaws like the type, how-
ever, and a specimen from Argentina illus-
trated by Orensanz (1975, fig. 8-3) has MIIIs
and IVs indistinguishable from most of the
North American specimens. There is thus
some overlap in jaw morphology between
the North and South American populations.
The number and shape of denticles on an-
terior jaws is variable in eunicoid poly-
chaetes, and the two populations are ten-
tatively considered conspecific in spite of
their disjunct geographic ranges.

The type and only specimen of Oenone
brevimaxillata Treadwell, 1931 was never
deposited (H. S. Feinberg, AMNH, pers.
comm. 1985), and is presumed lost. Tread-
well’s (1931, fig. 8) illustration of the max-
illae 1s schematic, and insufficient for de-
tailed comparison.

Family Oenonidae Kinberg, 1865

Emended diagnosis. —Jaws jet black,
composed primarily of scleroprotein, no
crystalline mineral phases present. Mandi-
bles H-shaped. Maxillary carriers much
longer than anterior jaws, attached to bun-
dles of oblique-longitudinal muscles, ven-
tral ligament present. Maxillae commonly
asymmetrical, MIs differ slightly to mark-
edly in length. One or three antennae arise
from base of prostomium. Elongate post-
setal lobe and prominent dorsal cirri de-
veloped on posterior parapodia. Geniculate
setae present, bifid hooks present or absent.

Key to the genera of Oenonidae

la. One or 3 antennae recline in notch
in peristomium, 2 peristomial rings
dorsally, bifid hooks present or ab-
sent
lb. Three antennae tucked into nuchal

122

fold, 1 peristomial ring dorsally,
bifid hooks present. 424.222 -: Oenone
2a. One antenna, peristomial notch
weakly developed, bifid hooks ab-
sent, setae smooth Tainokia
2b. Three antennae, distinct semicir-
cular peristomial notch, bifid hooks
present (small individuals only),
dorsal setae dentate, ventral setae
SUGIALCE toot aes 2s Re eR he Halla

o) = @ e je @ = ©

Genus Halla A. Costa, 1844

Type species.—Halla_ parthenopeia
(Chiaje) Costa, 1844:63.—Lysidice parthe-
nopeia Chiaje, 1828:175.

Remarks. —Fauvel (1923:427, fig. 169f)
described Halla parthenopeia as having bi-
fid hooks, in addition to the setae figured
here (Fig. 2a, b). Hooks are absent in para-
podia from the large specimen of H. par-
thenopeia examined here (USNM 5147), nor
were hooks recorded from the large speci-
men figured by Ehlers (1868:28, figs. 27,
28). The type of Halla okudai Imajima
(1967:437) is also a large worm, and lacks
hooks (Imajima & Hartman 1964:267-—268).
Individuals of Halla which attain large size
may lose their bifid hooks. A similar con-
dition prevails in several large species with-
in the Eunicidae (K. Fauchald, pers. comm.
1985).

Distribution. — All western hemisphere
specimens deposited in the USNM collec-
tion as Halla proved to be misidentified
members of Oenone or Lysarete. Halla may
be restricted to the Old World, with docu-
mented reports from the Mediterranean
(Fauvel 1923) and Japan (Imajima 1967).

Acknowledgments

I thank Kristian Fauchald for extensive
discussions about the taxonomy of eunicoid
polychaetes, and for critically reading the
manuscript. Thomas H. Perkins arranged
the loan of material from the Florida De-
partment of Natural Resources. H. S. Fein-
berg, American Museum of Natural His-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

tory, provided information concerning the
type of Oenone brevimaxillata Treadwell.
This research was supported by a post-doc-
toral fellowship, Smithsonian Institution.

Literature Cited

Benham, W. B. 1915. Report on the Polychaeta ob-
tained by the F. I. S. “Endeavour” on the coasts
of New South Wales, Victoria, Tasmania and
South Australia, part I.—Biological Results of
the Fishing Experiments carried on by the F. I.
S. “Endeavour,” 1909-14:173-273.

Chiaje, S. delle. 1828. Memorie sulla storia a notomia
degli animali senza vertebre del regno di Napoli,
3. Napoli, 232 pp.

Colbath, G. K. 1986. Jaw mineralogy in eunicean
polychaetes (Annelida).— Micropaleontology 32:
186-189.

Costa, A. 1844. Cerni intorno alle osservazioni zool-
ogiche fatte durante intre mesi vernali del
1844.—Annales Accademia aspirantes Natu-
rales Napoli 2:40-68.

Crossland, C. 1924. Polychaeta of tropical East Af-
rica, the Red Sea and Cape Verde Islands. The
Lumbriconereidae and Staurocephalidae. —
Proceedings of the Zoological Society of London
1924:1-106.

Ehlers, E. 1864-1868. Die Bornstenwiirmer (Anne-

lida, Chaetopoda). Leipzig, 748 pp.

1887. Reports on the results of dredging in
the U.S. Coast steamer “Blake.” XX XI. Report
on the annelids.—Memoirs of the Museum of
Comparative Zoology, Harvard 15:1-335.
Fauchald, K. 1970. Polychaetous annelids of the fam-
ilies Eunicidae, Lumbrineridae, Iphitimidae,
Arabellidae, Lysaretidae and Dorvilleidae from
western Mexico.— Allan Hancock Monographs
in Marine Biology 5:1-335.

. 1977. The polychaete worms.— Natural His-

tory Museum of Los Angeles, Science Series 28:

1-188.

Fauvel, P. 1923. Faune de France 5, polychetes er-
rantes.—Office Central de Faunistique, Paris,
488 pp.

Gardiner, S. L. 1976. Errant polychaete annelids from
North Carolina. — Journal of the Elisha Mitchell
Scientific Society 91:77—220.

Gilbert, K. M. 1984. Family Lysaretidae Kinberg,
1865. Pp. 43-1-43-7 in J. M. Uebelacker & P.
G. Johnson, eds., Taxonomic guide to the poly-
chaetes of the Northern Gulf of Mexico, Volume
6. Barry A. Vittor and Associates, Mobile, Al-
abama.

Hartman, O. 1944. Polychaetous annelids. Part V.
Eunicea.— Allan Hancock Pacific Expedition 10:
1-339.

VOLUME 102, NUMBER 1

1948. The marine annelids erected by Kin-
berg, with notes on some other types in the
Swedish State Museum.—Arkiv For Zoologi
42A:1-137.

. 1951. The littoral marine annelids of the Gulf
of Mexico.—Publications of the Institute for
Marine Science, University of Texas 2:7-124.

Hartmann-Schroder, G. 1967. Feinbau und Funktion
des Kieferapparates der Euniciden am Beispiel
von Eunice (Palola) siciliensis Grube (Poly-
chaeta).— Mitteilungen aus dem Hamburg Zool-
ogischen Museum und Institut 64:5—27.

Imajima,M. 1967. Errant polychaetousannelids from
Tsukumo Bay and vicinity of Noto Peninsula,
Japan.— Bulletin of the National Science Mu-

seum, Tokyo 10:403-441.

, & O. Hartman. 1964. The polychaetous an-

nelids of Japan.—Allan Hancock Foundation

Occasional Papers 26:1-452.

Kielan-Jaworowska, Z. 1966. Polychaete jaw appa-
ratuses from the Ordovician and Silurian of Po-
land and a comparison with modern forms.—
Paleontologica Polonica 16:1—152.

Kinberg, J. G. H. 1865. Annulata nova.—Ofversigt

af Svenska Kongliga Vetenskaps-Akademiens

Forhandlingar 21:559-574.

1910. Kongliga Svenska Fregatten Eugenies

123

Resa omkrig jorden. Vetenskapliga Iakttagelser.
II. Zoologi. 3. Annulater, Uppsala, 78 pp.

Knox, G. A., & K. M. Green. 1972. The polychaetes
of New Zealand part 3: Lysaretidae.— Journal
of the Royal Society of New Zealand 2:43 1-434.

Mohammad, M.-B. 1973. New species and records
of polychaete annelids from Kuwait, Arabian
Gulf. — Zoological Journal of the Linnean So-
ciety 52:23-44.

Orensanz, J. M. 1975. Los anelidos poliquetos de la
provincia biogeographica Argentina. VII. Eu-
nicidae y Lysaretidae.— Physis 34:85-111.

Rioja, E. 1944. Estudios anelidologicos. X. Estudio
de algunos poliquetos del Museo Argentino de
Ciencias Naturales.—Anales del Instituto de
Biologia, México 15:115-138.

Treadwell, A. L. 1931. New species of polychaetous
annelids from California, Mexico, Puerto Rico,
and Jamaica.—American Museum Novitates
(N.Y.) 1023:1-7.

Wolf, G. 1980. Morphologische Untersuchungen an
den Kieferapparaten einiger rezenter und fos-
siler Eunicoidea (Polychaeta). —Senckenber-
giana Maritima 12:1—182.

2195 Deborah Way, Upland, California
91786, USA.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 124-130

REDESCRIPTION OF NEPHTYS SQUAMOSA EHLERS
(POLYCHAETA: NEPHTYIDAE)

Takashi Ohwada

Abstract. —Confusion about the identity of Nephtys squamosa Ehlers, 1887
requires redescription of the species based on the syntypes. A lectotype is
designated. Comparisons are made with similar species.

Nephtys squamosa was originally de-
scribed by Ehlers (1887) as having scale-like
structures on both dorsum and ventrum, a
character unique to the family Nephtyidae.
Many subsequent authors have described
N. squamosa on the basis of specimens from
temperate and tropical waters of the Atlan-
tic and the East Pacific (Augener 1918;
Monro 1933; Fauvel 1936; Hartman 1940,
1950; Pettibone 1963; Nonato & Luna 1970;
Fauchald 1972, 1976; Gardiner 1976; Tay-
lor 1984; Lana 1986). Although all agree on
the presence of the scale-like lamellae, dis-
crepancies exist among descriptions in the
parapodial morphology. None of these de-
scriptions after that of Augener (1918) men-
tioned the presence of infrapreacicular la-
mellae on neuropodia, which is clearly stated
in the original description (Ehlers 1887).

Types of N. squamosa have been exam-
ined in the present study to clarify its iden-
tity. The presence of infrapreacicular la-
mellae has been confirmed on neuropodia,
and suprapreacicular lamellae have been also
found on notopodia. The present exami-
nation has further shown the presence of
tiny, slender dorsal cirri on the first noto-
podia, overlooked by previous workers.

Nephtys squamosa Ehlers, 1887
Figs. 1-3

Nephthys squamosa Ehlers, 1887:128-131,
pl. 37, figs. 7-10.

Material examined. — Original specimens
collected from Florida during the U.S. Coast

Survey 1868-1869, L. F. Pourtales, assist.:
off French Reef, Cast No. 6, 103 fms, 3 Apr
1869, paralectotype (MCZ 735).—off French
Reef, Cast No. 4, 50 fms, 3 Apr 1869, para-
lectotype (MCZ 740).—off Alligator Reef,
Cast No. 9, 113 fms, 8 May 1869, paralec-
totype (MCZ 836).—off American Shoal, 55
fms, lectotype (MCZ 1202).

Measurements. —Lectotype complete,
48.4 mm long, 3.4 mm wide including para-
podia with 120 setigers. Paralectotypes
(MCZ 735, 740, 836) incomplete posterior-
ly, respectively, 21.7 mm long, 3.5 mm wide
with 47 setigers, 54.4 mm long, 2.9 mm
wide with 105 setigers, 30.7 mm long, 2.4
mm wide with 96 setigers.

Description. —Body creamy white to
brown, lacking pigmentation in alcohol. Eyes
absent.

Prostomium pentagonal with anterior
margin convex, thin, spatulate (Fig. la, b);
lateral margins rounded, broadest between
second antennae (Fig. 2a), posterior margin
with V-shaped middorsal projection ex-
tending to posterior border of setiger 1. An-
terior part between first antennae translu-
cent, marked by an intricate pattern of
opaque tissue (Figs. 1b, 2a). First antennae
small, flattened, attached obliquely, ap-
pearing as raised lobes originating some dis-
tance from and fused with frontal prosto-
mial margin (Figs. 1b, 2a). Second antennae
large, basally flattened and distally conical,
slightly constricted at bases, much larger
than lst antennae, directed laterally at mid-
dle of ventral surface of prostomium, well

VOLUME 102, NUMBER 1 125

a
@'

=

i Soe

O
QR

* =

PANNE RE PAT Hy cana

lL :
|

Fig. 1. Nephtys squamosa (setae omitted): a, c, d, lectotype (MCZ 1202), b, paralectotype (MCZ 735): a,
Anterior end, dorsal view, ventrally dissected and somewhat flattened, dorsal cirrus missing and preacicular
lobe twisted outside on Ist right notopodium (acL~acicular lobe; prL—preacicular lobe); b, Same, ventral view,
3rd right ventral cirrus missing (notopodia omitted); c, 72nd to 76th setiger, dorsal view (neuropodia omitted);
d, 74th to 78th setiger, ventral view (notopodia omitted).

behind Ist antennae and slightly anterior to podia reduced, with triangularly pointed
Ist neuropodia (Figs. la, b; 2a, b). Mouth acicular lobes, short preacicular and post-
covered with two thick lateral lips and thick acicular lobes (preacicular lobes concealed
rugose midventral lip at level of setiger 1 behind acicular lobes and usually not rec-
(Fig. 1b); rugose area extending to setiger 5. ognizable from above); dorsal cirri (barely

Setiger 1 modified (Figs. la, 2b); noto- visible unless stained) tiny, slender, point-

126

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Nephtys squamosa: a, c, paralectotype (MCZ 735), b, lectotype (MCZ 1202): a, Anterior half of
prostomium, dorsal view (setae omitted); b, Ist left parapodium, dorsal view, preacicular lobe obscured by
acicular lobe (acL—acicular lobe; dc—-dorsal cirrus; tips of setae omitted); c, 12th right neuropodium, dorsal view

(setae omitted).

ed, arising from outer side of acicular lobes
near their base between preacicular and
postacicular setal fascicles (Fig. 2b). Neu-
ropodia of setiger 1 on anterior margin, lat-
eral to prostomium, with slender rudimen-
tary acicular lobes projecting anteriorly
(Figs. la, 2b); acicular lobes supported by
thin acicula whose tip curves inward. Ven-
tral cirri large, similar to or smaller than
2nd antennae, digitate but tips may be con-
ical, lateral to neuropodia and continuous
with anterior setigerous margins, directed
laterally or bent backward (Figs. la, 2b).
Notosetae of setiger 1 including preacicular
barred (laddered) capillaries and very finely
serrated slender postacicular capillaries;
neurosetae long, thin, smooth capillaries,
surrounding acicular lobes (Fig. 2b).
Parapodia similar throughout from se-
tiger 2; both rami with preacicular and post-
acicular setal fascicles; branchiae (interra-
mal cirri) may be lacking on setiger 2. Both
notopodial and neuropodial acicular lobes
conical (Fig. 3a—c), becoming acutely coni-

cal and pointed posteriorly (Fig. 3e). No-
topodial presetal lamellae thin, rather broad
and slightly bilobed, distally rounded, ex-
tending from lower margins of notopodia,
as long as acicular lobes on anterior para-
podia (Fig. 3a—c), posteriorly reduced and
much shorter than acicular lobes (Fig. 3e).
Small lamellae dorsal to notopodial presetal
lobes, thinner than presetal lobes except on
middle parapodia, rounded on anterior no-
topodia (Fig. 3a, b), posteriorly reduced and
becoming much thinner and smaller (Fig.
3c, e), barely recognizable on posterior se-
tigers (Fig. 3e). Preacicular setae appearing
mostly beyond dorsal margins of presetal
lamellae and usually partially covering small
suprapreacicular lamellae (Fig. 3a—c, e).
Neuropodial presetal lamellae thin, larger
than notopodial presetal lamellae, extend-
ing from upper margins of neuropodia rath-
er triangularly on middle parapodia (Fig.
3b, c), distally rounded on all but especially
on anterior and posterior parapodia (Fig.
3a, e), as long as acicular lobes on anterior

VOLUME 102, NUMBER 1

127

Fig. 3. Nephtys squamosa (tips of setae omitted): a, b, paralectotype (MCZ 735), c-e, lectotype (MCZ 1202):
a, Left parapodium, setiger 6, anterior view; b, Same, setiger 32, anterior view; c, Same, setiger 40, anterior
view; d, Same, posterior view (preacicular setae omitted); e, Right parapodium, setiger 79, anterior view.

neuropodia (Fig. 3a, b), posteriorly reduced
and much shorter than acicular lobes (Fig.
3e). Lamellae ventral to neuropodial pre-
setal lobes, broad, thin, shorter than presetal
lobes, slightly bilobed on anterior parapodia

(Fig. 3a, b), posteriorly reduced and becom-
ing narrower and much thinner, thinner than
presetal lobes, hardly recognizable on pos-
terior setigers (Fig. 3e). Preacicular setae ap-
pearing mainly beyond ventral margins of

128

presetal lamellae and usually partially cov-
ering infrapreacicular lamellae (Fig. 3a-c,
e). Notopodial postsetal lamellae thin, lig-
ulate, extending dorsolaterally from seg-
mental wall above and longer than acicular
lobes, distally rather rounded on anterior
setigers (Fig. 3a), posteriorly becoming more
or less pointed (Fig. 3b—e). Neuropodial
postsetal lamellae large, thin, extending from
behind and longer than acicular lobes (Fig.
3a-e), posteriorly extending much further
(Fig. 3c, e), but degree of extension variable
between specimens (Fig. 3b, c), with rather
pointed tips except on anterior parapodia
where distally rounded (Fig. 3a). Small, short
lobe on superior edge of neuropodia be-
tween presetal and postsetal lamellae (Figs.
2c, 3a, d, e) except on setiger 1, distinct on
anterior neuropodia but posteriorly reduced
and barely recognizable. Dorsal cirri flat-
tened, thin, triangularly elongate with rather
pointed tips, smaller than notopodial post-
setal lamellae, extending laterally from bas-
al part of branchiae behind acicular lobes
(Fig. 3a—e). Ventral cirri digitiform on an-
terior several parapodia (Figs. 1b, 3a),
thereafter foliaceous, constricted at bases
(Figs. 1b, 3b-d).

Branchiae first present on setigers 2-3
usually as processes slightly curved out-
ward, completely recurved within a few se-
tigers (Fig. 3a), thereafter gradually increas-
ing in size and fully developed by setigers
25-27 (Fig. 3c-e), then (in lectotype) de-
creasing gradually in size after setigers 89-
92, and not recurved after setigers 106-107,
still present as very small rudimentary ridge
on setiger 116, completely absent on last 4
setigers.

Scale-like dorsal lamellae broad, folia-
ceous (Fig. Ic), originating from dorsolat-
eral wall of body (Fig. 3b-e), extending
mainly posterolaterally over notopodia,
partially overlapping successive segments in
shingle-like arrangement (Fig. lc), begin-
ning on setigers 5-9, becoming larger more
posteriorly but not fully developed until

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

middle setigers (Fig. 3c, d), gradually de-
creasing in size (in lectotype) over posterior
several setigers but still recognizable on se-
tiger 117, absent on last 3 setigers. Similar
but much smaller lamellae present on ven-
trolateral surface of body (Figs. 1d, 3c, d),
foliaceous, extending somewhat postero-
laterally, only slightly covering neuropodia
and successive segments (Fig. 1d), begin-
ning on setiger 2, increasing in size more
posteriorly but not fully developed until
middle setigers (Fig. 3c, d), gradually re-
duced (in lectotype) over posterior several
setigers but still recognizable on setiger 117,
absent on last 3 setigers.

Both preacicular and postacicular setae
occurring in spreading fascicles, with post-
acicular fascicles wider than preacicular ones
(Fig. 3a—c, e); preacicular setae much shorter
and fewer than postacicular setae. Most
preacicular setae consisting of barred cap-
illaries; a few, similarly long, very finely ser-
rated capillaries at upper and lower end of
preacicular fascicles. Postacicular setae con-
sisting of finely serrated long capillaries. No
lyrate seta found. Acicula single in rami,
completely embedded in conical acicular
lobes (Fig. 3b-e).

Proboscis not everted in specimens ex-
amined. Proboscis in lectotype previously
dissected, damaged, incomplete, with ter-
minal circlet of 20 bifid papillae and simple
middorsal papilla, with at least 20 rows of
5-6 cirriform papillae subterminally, and
more distal cirriform middorsal papilla;
middorsal papilla not noticeably larger nor
longer than others. Pair of horny jaws pres-
ent inside proboscis. Pharyngeal trepans ab-
sent.

Remarks.—In the original description,
Ehlers (1887:130) stated that dorsal cirri
were about half as long as notopodial post-
setal lamellae. This is the case in the middle
parapodia of the lectotype (Fig. 3c, d). In
two paralectotypes (MCZ 735, 836), how-
ever, notopodial postsetal lamellae do not
extend beyond dorsal cirri (Fig. 3a, b). The

VOLUME 102, NUMBER 1

relative length of notopodial postsetal la-
mellae to dorsal cirri is thought to vary
greatly between specimens.

There are obvious problems with descrip-
tions of Nephtys squamosa. Although the
presence of suprapreacicular lamellae on
notopodia has been clearly recognized in the
lectotype and paralectotypes in the present
study, neither the original description (Eh-
lers 1887) nor any subsequent description
mentioned this structure, whereas Ehlers
described infrapreacicular lamellae on neu-
ropodia (p. 130, Taf. 37, fig. 8). Examina-
tion of the lectotype and paralectotypes also
revealed some other structures such as tiny,
slender dorsal cirri on Ist notopodia and
small, short lobes on the superior edge of
neuropodia between presetal and postsetal
lamellae, which had been overlooked by
previous workers.

Descriptions subsequent to Augener
(1918) did not mention the presence of in-
frapreacicular lamellae on neuropodia.
Hartman’s figure (1940: pl. 41, fig. 98) of
the 25th parapodium of N. sguamosa from
the Gulf of California showed only preacic-
ular lamellae both on the notopodium and
the neuropodium, and she stated (p. 237)
that the prostomium of her specimens dif-
fered from that originally described for the
species. Neither is an additional preacicular
lamella shown on either the notopodium or
the neuropodium in Taylor’s figure (1984:
fig. 35-8b) of the 20th parapodium of N.
squamosa from the Gulf of Mexico. Taylor
also stated (p. 11) that dorsal cirri were lack-
ing on the Ist notopodia of his specimens,
whereas dorsal cirri, although tiny and slen-
der, are present on Ist notopodia of the lec-
totype and paralectotypes. Additional pre-
acicular lamellae are neither described nor
figured also in Lana’s description (1986:139,
figs. 4, 5) of N. squamosa from Brazil. Since
additional preacicular lamellae, especially
suprapreacicular lamellae on notopodia, are
difficult to observe in some specimens, the
lack of description may not necessarily im-

129

ply absence in specimens concerned. Never-
theless, it seems to be necessary to reex-
amine these specimens for certain
identification.

Nephtys squamosa is closely related to N.
acrochaeta Hartman, 1950, N. imbricata
Grube, 1857 and N. serratifolia Ehlers, 1897
in the possession of scale-like dorsal la-
mellae. Nephtys squamosa, however, differs
from N. acrochaeta in that branchiae are
present from setigers 2—3 in the former and
9-10 (Hartman 1950, Fauchald 1976) in the
latter. Neuropodial postsetal lamellae are
lacinated in N. acrochaeta (Hartman 1950)
and N. serratifolia (Ehlers 1897), whereas
they are entire in N. squamosa. Capillary
setae with a large spur are present in post-
acicular fascicles both in N. acrochaeta
(Hartman 1950) and N. serratifolia (Hart-
man-Schroder 1965), but absent in N. squa-
mosa. Nephtys imbricata is ‘too incomplete-
ly known for certain identification’
(Hartman 1950:111), but according to the
original description (Ehlers 1887), N. squa-
mosa differs from N. imbricata in parapo-
dial morphology. I have examined types of
both species, however further study will be
needed to fully characterize N. imbricata.

Acknowledgments

I am grateful to Dr. Ardis B. Johnston,
Museum of Comparative Zoology, Harvard
University (MCZ) for the loan of the spec-
imens. The manuscript benefited from the
helpful suggestions by Dr. Mary E. Petersen,
Zoological Museum, University of Copen-
hagen, Denmark and the critical review by
Dr. Thomas H. Perkins, Florida Depart-
ment of Natural Resources.

Literature Cited

Augener, H. 1918. Polychaeta.—Beitrage zur Kennt-
nis des Meeresfauna West-Afrikas 2:67-625, 6
pls.

Ehlers, E. 1887. Report on the annelids of the dredg-
ing expedition of the U.S. coast survey steamer
Blake.—Memoirs of the Museum of Compar-

130

ative Zoology of Harvard College 15:vi + 335

pp., 60 pls.

1897. Polychaeten.—Hamburger Magalla-
nische Sammelreise. Hamburg, Friedrichsen &
Co. 148 pp., 9 pls.

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.

1976. Some Nephtyidae (Polychaeta) from
Ubatuba, Brasil.— Bulletin of the Southern Cal-
ifornia Academy of Sciences 75:16-19.
Fauvel, P. 1936. Contribution a la fauna des Anné-

lides polychétes du Moroc. — Mémoires de la So-
ciété des Sciences Naturelles du Moroc 43:1-
143.

Gardiner, S. L. 1976. Errant polychaete annelids from
North Carolina. — Journal of the Elisha Mitchell
Scientific Society 91:77—220.

Grube, E. 1857. Annulata Orstediana Enumeratio
Annulatorum, quae in itinere per Indiam occi-
dentalem et Americam centralem annis 1845-
1848 suscepto legit cl. A. S. Orsted, adjectis spe-
ciebus nonnullis a cl. H. Kréyero in itinere ad
Americam meridionalem collectis.— Videnska-
belige Meddelelser fra Dansk naturhistorisk
Forening i Kjgbenhavn 1857:1-29.

Hartman, O. 1940. Polychaetous annelids. Part 2.

Chrysopetalidae to Goniadidae.—Allan Han-

cock Pacific Expeditions 7:173-—287.

1950. Polychaetous annelids. Goniadidae,
Glyceridae, Nephtyidae.—Allan Hancock Pa-
cific Expeditions 15:1-181.
Hartmann-Schroder, G. 1965. Zur Kenntnis des Sub-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

litorals der chilenischen Kiiste unter besonderer
Beriicksichtigung der Polychaeten und Ostra-
coden. Teil II. Die Polychaeten des Sublito-
rals.— Mitteilungen aus dem Hamburgischen
Zoologischen Museum und Institut 62 (Supple-
ment):59-305.

Lana, P.C. 1986. Nephtyidae (Annelida; Polychaeta)
do litoral do estado do Parana (Brazil). — Neri-
tica 1:135-154.

Monro, C. C. A. 1933. The Polychaeta Errantia col-
lected by Dr. C. Crossland at Colon, in the Pan-
ama region, and the Galapagos Islands during
the expedition of the S. Y. ‘St. George’.— Pro-
ceeding of the Zoological Society of London
1933:1-96.

Nonato, E. F., & J. A. C. Luna. 1970. Anelideos
poliquetas do nordeste do Brasil. 1. Poliquetas
bent6nicos da costa de Alagoas e Sergipe.—
Boletim do Instituto Oceanografico, Sao Paulo
19:57-130.

Pettibone, M. H. 1963. Marine polychaete worms of
the New England Region. 1. Aphroditidae
through Trochochaetidae.— United States Na-
tional Museum Bulletin No. 227:1-356.

Taylor, J. L. 1984. Family Nephtyidae Grube, 1850.
Pp. 1-20 in J. M. Uebelacker & P. G. Johnson,
eds., Taxonomic guide to the polychaetes of the
Northern Gulf of Mexico, Vol. 5, Chapt. 35.
Barry A. Vittor & Associates, Inc., Mobile, Al-
abama.

Ocean Research Institute, University of
Tokyo, 15-1, 1-Chome, Minamidai, Na-
kano-ku, Tokyo 164 Japan.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 131-133

PHALLODRILUS HESSLERI, NEW SPECIES
(OLIGOCHAETA: TUBIFICIDAE), FROM ABYSSAL
DEPTHS IN THE WESTERN PACIFIC OCEAN

Christer Erséus

Abstract.—Phallodrilus hessleri is described from abyssal clay at 5916 m
depth east of the Philippines. It is characterized by (1) slender, sharply pointed
somatic setae with very thin (or absent?) upper teeth, (2) somewhat sigmoid
penial setae in segment XI, 2 per bundle, and (3) very small, spindle-shaped
atria. The species represents the second deepest record of marine oligochaetes

in the world.

In a collection of benthic invertebrates
from a deep-sea area near the Philippine
Trench in the western Pacific Ocean, de-
posited in the National Museum of Natural
History (USNM), Washington, D.C., the
author found one well preserved individual
of a new species of Phallodrilus Pierantoni,
1902 from a station almost 6000 m deep.
Oligochaetes from such great depths are in-
deed rare. It was therefore considered ap-
propriate to describe the species on the basis
of this specimen.

The worm was stained in paracarmine and
mounted whole in Canada balsam.

Phallodrilus hessleri, new species
Fig. 1

Holotype. —USNM_ 104128, whole-
mounted specimen from E of the Philippine
Trench, West Pacific Ocean, 9°06.0'N,
127°43.6’E, 5916 m, abyssal red clay (13
Mar 1975).

Etymology. —Named for Dr. Robert R.
Hessler (Scripps Institution of Oceanogra-
phy, La Jolla, California), who collected the
sample.

Description. —Length more than 2.5 mm,
more than 16 segments (worm not com-
plete). Width at XI (compressed) 0.22 mm.
Prostomium large, rounded. Body wall
bearing scattered, very small particles. Cli-
tellum extending over '2X—XII. Somatic se-

tae (Fig. 1A, B) bifid, with teeth almost par-
allel, upper tooth shorter and much thinner
than lower, sometimes not visible (or ab-
sent?; cf. right bundle in Fig. 1B). Bifids 50—
85 um long, 1.5—2.5 wm thick at node, three
to four per bundle anteriorly, two (or three)
per bundle in postclitellar segments. Penial
setae (Fig. 1C, D, ps) two per bundle, some-
what sigmoid, sharply single-pointed, 65—
75 um long, about 3 wm thick at node, di-
rected in an almost antero-posterior direc-
tion. Male pores inconspicuous, paired ven-
trally and posteriorly in XI. Spermathecal
pores paired, in line with ventral setae(?) in
anteriormost X.

Pharyngeal glands in IV—VII. Male gen-
italia (Fig. 1D) paired. Vas deferens 8-10
um wide, coiled, much longer than atrium,
entering apical end of latter. Atrium pear-
shaped, about 60 um long, 28-33 um wide,
with very thin outer lining and ciliated inner
epithelium; latter densely granulated in ec-
tal, wide part of atrium. Prostate glands
compact, round, one at each end of atrium.
Atrium opening to exterior through simple
pore; penis or pseudopenis absent. Sper-
mathecae (Fig. 1D, s) not seen in their full
length (hidden behind gut), but consisting
of short, narrow duct, and slender ampulla.

Remarks. —The diagnostic characters of
this species are (1) the slender, sharply
pointed somatic setae with very thin upper

132
B C
A
ae ec)
Fig. 1.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Phallodrilus hessleri: A, Free-hand drawing of tip of somatic seta; B, Two bundles of somatic setae

(from postclitellar segments); C, Penial setae; D, Horizontal view of spermathecae (only parts visible) and male
genitalia in segments X—XI. Abbreviations: a, atrium; pr 1, anterior prostate gland; pr 2, posterior prostate
gland; ps, penial seta; s, spermatheca; sf, sperm funnel; vd, vas deferens.

teeth (or sometimes absent?), (2) the bisetal
penial bundles, and (3) the small atria. Oth-
er species within the genus with similar fea-
tures are all deep-sea forms, viz. P. profun-
dus Cook, 1970 (N Atlantic), P. remus
Erséus, 1979 (NW Atlantic), P. altus Erséus,
1980 (Beaufort Sea, Arctic), and P. segon-
zaci Erséus, 1986 (S Indian Ocean), and they
are probably the closest relatives of P. hess-
leri. However, in all these other species, the
penial setae are clearly longer than the so-
matic setae (setae about equally long in hess-
leri). The new species is further distin-
guished from all of them except P. remus
by its very short atria (atria not very large,
but elongate in the others), from P. remus
by the shape of its penial setae (those of
remus oar-shaped, with wide inner ends),
and from P. profundus and P. segonzaci by
its lack of modified genital setae in segments
IX and X.

Distribution and habitat.—Known only
from the type-locality E of the Philippines,
W Pacific Ocean. Abyssal clay, 5916 m.

Discussion

More than 30 species of Tubificidae have
been described from the deep sea, all within
the last 20 years (Cook 1970a, b; Erséus

1979a, b, 1980, 1982a, b, 1983a—c, 1984,
1986, 1988; see also review by Erséus
1985). Most of them are bathyal forms, but
four are known from depths greater than
4000 m: the North Atlantic Bathydrilus at-
lanticus Erséus, 1979 (depth range 1600-
4632 m; Erséus 1983), the two Indian Ocean
species Phallodrilus segonzaci and P. stilus
Erséus, 1986, both from a single station at
4910 m (Erséus 1986), and the North Pacific
B. hadalis Erséus, 1979, which was collected
at 7298 m in the Aleutian Trench (Erséus
1979a). Phallodrilus hessleri (from 5916 m)
thus represents the second deepest record
of marine oligochaetes in the world.

Acknowledgments

I am indebted to Dr. K. Fauchald for
bringing my attention to the Philippine
Trench collection in the USNM, and for the
loan of the specimen studied; to Dr. R. R.
Hessler for providing station data; and to
Ms. Barbro Lofnertz and Mrs. Aino Falck-
Wahlstrom for technical assistance.

Literature Cited

Cook, D.C. 1970a. Peloscolex dukei n. sp. and P.
aculeatus n. sp. (Oligochaeta, Tubificidae) from
the N.W. Atlantic, the latter being from abyssal

VOLUME 102, NUMBER 1

depths. — Transactions of the American Micro-
scopical Society 88(1969):492-497.

. 1970b. Bathyal and abyssal Tubificidae (An-
nelida, Oligochaeta) from the Gay Head-Ber-
muda transect, with descriptions of new genera
and species.— Deep-Sea Research 17:973-981.
Erséus, C. 1979a. Taxonomic revision of the marine
genera Bathydrilus Cook and Macroseta Erséus
(Oligochaeta, Tubificidae), with descriptions of
six new species and subspecies.— Zoologica
Scripta 8:139-151.

1979b. Taxonomic revision of the marine
genus Phallodrilus Pierantoni (Oligochaeta,
Tubificidae), with descriptions of thirteen new
species. — Zoologica Scripta 8:187—208.

1980. New species of Phallodrilus (Oligo-
chaeta, Tubificidae) from the Arctic deep sea
and Norwegian fjords.—Sarsia 65:57—60.
. 1982a. Atlantidrilus, a new genus of deep-sea
Tubificidae (Oligochaeta).—Sarsia 67:43-46.

1982b. Taxonomic revision of the marine
genus Limnodriloides (Oligochaeta: Tubifici-
dae).— Verhandlungen des Naturwissenschaft-
lichen Vereins in Hamburg (Neue Folge) 25:
207-277.

1983a. New records of Adelodrilus (Oligo-
chaeta, Tubificidae), with descriptions of two
new species from the Northwest Atlantic. — Hy-
drobiologia 106:73-83.

. 1983b. Deep-sea Phallodrilus and Bathydri-

133

lus (Oligochaeta, Tubificidae) from the Atlantic
Ocean, with descriptions of ten new species. —
Cahiers de Biologie Marine 24:125-146.
. 1983c. A new bathyal species of Coralliodri-
lus (Oligochaeta, Tubificidae) from the S.E. At-
lantic.— Proceedings of the Biological Society of
Washington 96:273-275.
. 1984. New and little-known species of deep-
sea Tubificidae (Oligochaeta) from the North-
west Atlantic.— Zoologica Scripta 13:101-106.
. 1985. Distribution and biogeography of Oli-
gochaeta. Pp. 365-367 in L. Laubier and C.
Monniot, eds., Peuplements profonds du Golfe
de Gascogne. Institut Francais de Recherche pour
V’Exploitation de la Mer (IFREMER), Brest.
. 1986. Two new abyssal species of Phallodri-
lus (Oligochaeta, Tubificidae) from the South
Indian Ocean.— Bulletin du Muséum National
d’Histoire Naturelle (4° sér.) 8 (section A, n° 3):
567-572.
—. 1988. Deep-sea Tubificidae (Oligochaeta)
from the Gulf of Mexico.— Proceedings of the
Biological Society of Washington 101:67-71.

Zoo-Tax, Swedish Museum of Natural
History, Stockholm, and (postal address):
Department of Zoology, University of G6-
teborg, Box 25059, S-400 31 Goteborg,
Sweden.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 134-136

PHALLODRILUS VESCUS, NEW SPECIES
(OLIGOCHAETA, TUBIFICIDAE) FROM THE
GULF OF MEXICO

Christer Erséus

Abstract.—Phallodrilus vescus (subfamily Phallodrilinae) is described from
the continental slope of the northern part of the Gulf of Mexico. The species
is distinguished from the closely related P. davisi Erséus, 1984 by (1) its fewer
setae, (2) its very small anterior prostate glands, (3) its very short penial papillae,
and (4) the markedly narrow inner part of its spermathecal ampullae.

Species of Phallodrilinae from the north-
ern Gulf of Mexico continental slope, orig-
inating from a project at LGL Ecological
Research Associates, Inc. (Bryan, Texas),
were recently described by Erséus (1988).
Subsequently, further material was sent to
this author for identification. Three worms
proved to represent an additional new
species of Phallodrilus Pierantoni, 1902,
which is described herein.

The specimens, weakly stained in Rose
Bengal, were mounted whole in Canada bal-
sam by the author. They are deposited in
the United States National Museum of Nat-
ural History (USNM), Smithsonian Insti-
tution, Washington, D.C.

Phallodrilus vescus, new species
Fig. 1

Holotype.—USNM 118716, from south
of Louisiana, 27°49'36’”N, 90°07'06’W, 841
m (12 Apr 1984).

Paratypes.—USNM 118717, 118718, one
from type locality, and one from
27°49'24”’N, 90°07'00’"W, 840 m (12 Apr
1984).

Etymology. —The epithet vescus is Latin
for “thin, slender,” a feature of the new
species.

Description. —Length (two complete, fixed
specimens) 3.4—-5.6 mm, 31-38 segments.
Width at XI (compressed, whole-mounted

specimens) 0.08-0.13 mm. Prostomium
rounded triangular, pygidium blunt-ended,
clitellum extending over 2X—XII. Setae (Fig.
1A) bifid, with upper tooth slightly thinner
and shorter than lower. Setae 35-50 um long,
1—1.5 um thick, three per bundle anteriorly,
two per bundle in postclitellar segments.
Ventral setae absent from XI. Male pores
paired in line with ventral setae, posterior
to middle of segment X.

Pharyngeal glands poorly developed. Male
genitalia (Fig. 1B) paired. Vas deferens about
5 wm wide, long and coiled, entering apical
end of atrium. Atrium comma-shaped,
about 30 um long, about 15 um wide, with
thin outer (muscular) lining and thick, gran-
ulated inner epithelium; ciliation not seen.
Anterior prostate gland small, inconspic-
uous, located near junction between vas def-
erens and atrium. Posterior prostate gland
large, conspicuous, attached to ectal part of
atrium. Atrium terminating in small penial
papilla, about as long as wide (10-15 um),
enclosed in simple penial sac. Spermathecae
(Fig. 1B, s) slender, 65-90 um long, con-
sisting of short, 11-13 wm wide ducts, and
11-18 wm wide ampullae; latter with inner
end very narrow in (at least) two of the spec-
imens studied. Lumen of ampullae incon-
spicuously and irregularly incised in a few
places. A few spermatozoa, arranged in a
bundle, present in each ampulla.

Remarks. —This species is distinguished

VOLUME 102, NUMBER 1

Fig. 1.

135

| 50 pm

Phallodrilus vescus: A, Free-hand drawing of somatic seta; B, Lateral view of spermatheca and male

genitalia in segments X—XI. Abbreviations: a, atrium; p, penial papilla; pr 1, anterior prostate gland; pr 2,
posterior prostate gland; s, spermatheca; vd, vas deferens.

from the closely related P. davisi, described
by Erséus (1984) from sublittoral depths (78-
80 m) off Massachusetts, by: (1) having only
two setae per bundle in postclitellar seg-
ments (up to four setae in davisi), (2) its very
small anterior prostate glands (in davisi,
these glands are conspicuous and in fact
larger than the posterior prostates), (3) its
very short penial papillae (penes slender and
conspicuously hollow in davisi), and (4) the
narrow inner end of its spermathecal am-
pullae (these ends rounded, not markedly
narrow in davisi).

The genitalia of P. vescus bear resem-
blance to those of P. constrictus Erséus, 1988,
which also occurs on the upper continental
slope in the northern Gulf of Mexico (Erséus
1988). The latter species, however, pos-
sesses a very thick muscular lining on the
atria, copulatory sacs and spermathecae;
penial setae in segment XI; and a very dis-
tinct constriction on the male ducts between
the atria and the copulatory sacs; these fea-
tures are all absent in P. vescus.

The spermathecal ampullae are markedly

incised or compartmented in P. davisi (Er-
seus 1984:fig. 6B) and P. constrictus (Erséus
1988:fig. 1C). A similar, but less developed,
compartmentalization is present in the
spermathecae of P. vescus (Fig. 1B).

Distribution and habitat. —N Gulf of
Mexico. Upper continental slope sediment,
840-841 m.

Acknowledgments

The LGL specimens reported on in this
publication were collected through funding
by the U.S. Department of Interior, Min-
erals Management Service, Gulf of Mexico
Regional OCS Office under Contract Num-
ber 14-12-0001-30046 and 14-12-0001-
30212. I am indebted to Dr. Linda H. Pe-
quegnat (LGL) for placing the material at
my disposal, and to Ms. Barbro Lofnertz
and Mrs. Aino Falck-Wahlstrom for tech-
nical assistance.

Literature Cited

Erséus, C. 1984. Taxonomy of some species of Phal-
lodrilus (Oligochaeta: Tubificidae) from the

136 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Northwest Atlantic, with description of four new
species. — Proceedings of the Biological Society
of Washington 97:812-826.

—. 1988. Deep-sea Tubificidae (Oligochaeta)
from the Gulf of Mexico.— Proceedings of the
Biological Society of Washington 101:67-71.

Zoo-Tax, Swedish Museum of Natural
History, Stockholm, and (postal address):
Department of Zoology, University of G6-
teborg, Box 25059, S-400 31 Goteborg,
Sweden.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 137-153

NEW SPECIES OF SCALE-WORMS
(POLYCHAETA: POLYNOIDAE) FROM THE
HYDROTHERMAL RIFT-AREA OF THE
MARIANA BACK-ARC BASIN IN THE
WESTERN CENTRAL PACIFIC

Marian H. Pettibone

Abstract. —Polynoid polychaetes collected by the Mariana Back-Arc Basin
Expedition in the western Central Pacific in 1987 are described. They include
four new species belonging to three subfamilies and four genera of Polynoidae:
Macellicephalinae: Levensteiniella raisae,; Lepidonotopodiinae: Lepidonoto-
podium minutum,; and Branchinotogluminae: Branchinotogluma burkensis and
Opisthotrochopodus marianus. Included also is a brief summary of the subfam-
ilies, genera, and species associated with deep-sea hydrothermal vents, deep
seep-sites, trenches, abyssal basins, experimental deep wood panels, and caves.

In her revision of the genera and species
that had been grouped in Macellicephalinae
Hartmann-Schroder, 1971, containing
mostly abyssal species, Pettibone (1976)
separated Macellicephalinae and five ad-
ditional subfamilies: Macellicephaloidinae
(four species), Macelloidinae, Bathyedithi-
nae, Polaruschakovinae (two species), and
Bathymacellinae. In the remaining Macel-
licephalinae, in addition to Macellicephala
(seven species), six new genera were added:
Bathykermadeca, Bathyeliasona (three
species), Bathyvitiazia, Bathyfauvelia,
Bathycatalina, and Bathykurila.

Subsequently, additional new subfami-
lies, genera, and species have been added.
Pettibone (1979) emended and added to
Macellicephalinae Bruunilla natalensis
(Hartman, 1971). Levenstein (1975) added
two new species of Macellicephala from the
deep-sea trenches in the Antarctic: M. tri-
cornis and M. grandelytris. The latter species
was later transferred to Bathyfauvelia by
Levenstein (1978a:77). Levenstein (1975)
also added the new species Macellicepha-
loides sandvichensis. Levenstein (1978b)
added the new species Macellicephala alia
from the Palau trench and the new genus

Bathymariana for B. zebra from the Ryuku
trench. The latter genus was placed in
Bathyedithinae by Levenstein (1984).

From the deep Canadian basin in the Arc-
tic Ocean, Levenstein (1981) added the new
genus Bathypolaria for B. carinata in Ma-
cellicephalinae; the new species Bathyedi-
thia tuberculata in Bathyedithinae; and the
new genera Bathymiranda for B. microceph-
ala and Bathycanadia for B. diaphana in
Polaruschakovinae. In the same subfamily,
Loshamn (1981) added the new genus Di-
placonotum for D. paucidentatum (Eliason,
1962) from the Skagerrak.

From the Japanese trench, Levenstein
(1982a) added a new species of Macelli-
cephaloides, M. villosa. From the Tasman
trench, Levenstein (1982b) included a key
to the genera of Macellicephalinae and added
two new genera: Bathytasmania for B. in-
solita, and Bathynotalia for B. perplexa. The
latter genus and species were referred to
Harmothoinae by Pettibone (1985a:130),
based on the figure of the prostomium
showing long lateral antennae (not men-
tioned in the text): the prostomium is har-
mothoid, with cephalic peaks; prostomial
lateral antennae are lacking in Macelli-

138

cephalinae. From the Kurile-Kamchatka
trench, Levenstein (1983) added a new
species of Macellicephaloides, M. improvi-
sa, and included a key to the seven species
of the genus.

Levenstein (1984) summarized the dis-
tributional patterns for the 19 genera and
38 species of polynoid polychaetes belong-
ing to the six deep-sea subfamilies covered
in Pettibone (1976), along with the addi-
tional genera and species indicated above.
They were widely distributed on the abyssal
plains and in the deep-sea trenches. The
Macellicephalinae had the greatest number
of genera (10) and species (21). Three
subfamilies were represented by fewer gen-
era and species: Macellicephaloidinae (one
genus, seven species), Bathyedithinae (two
genera, three species), and Polaruschako-
vinae (four genera, five species). Two
subfamilies were monotypic: Macelloidinae
and Bathymacellinae. These groups were
defined as ancient or primary abyssal be-
cause all the representatives lived only in
abyssal depths except for a few that ascend-
ed secondarily. Members of the Macelli-
cephalinae showed the greatest vertical range
and panoceanic distributions, mostly in the
Northern Hemisphere. This group appeared
to be the most ancient of the abyssal poly-
noids.

Since the summary of the group by Lev-
enstein (1984), additional new subfamilies,
genera, and species have been added, chiefly
in connections with studies on the hydro-
thermal vents in the northeastern Pacific,
deep seep-sites in the Gulf of Mexico, and
deep wood panel and cave studies in the
northwestern Atlantic.

From the hydrothermal rift-area off west-
ern Mexico at 21°N, Pettibone (1983) added
the new subfamily Lepidonotopodiinae for
Lepidonotopodium fimbriatum. From the
Galapagos vent, the new subfamily Bran-
chipolynoinae for Branchipolynoe symmy-
tilida, commensal with the deep-sea mussel,
was added by Pettibone (1984a). Also from
the vents, Pettibone (1984b) added two new

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

species of Lepidonotopodium: L. riftense and
L. williamsae and included a key to the
species of the genus.

From the North Atlantic, Pettibone
(1985a) added to Macellicephalinae two new
genera: Pelagomacellicephala for P. iliffei,
from a cave in the Bahamas, and Bathy-
bahamas for B. charleneae from off the Ba-
hamas, associated with experimental wood
panels, and a new species of Bathykerma-
deca, B. turnerae, from off the Virgin Is-
lands, also associated with wood panels.

Pettibone (1985b) added the new subfam-
ily Branchiplicatinae for Branchiplicatus
cupreus from the vents off western Mexico
at 21°N. From the vents off the Galapagos
and western Mexico, Pettibone (1985c)
added the third new subfamily having ar-
borescent branchiae, Branchinotogluminae
for two new genera: Branchinotogluma, with
three new species: B. hessleri, B. sandersi,
and B. grasslei; and Opisthotrochopodus for
O. alvinus. A key to the three branchiate
subfamilies was included.

Pettibone (1985d) added to Macelli-
cephalinae the new genus Levensteiniella for
L. kincaidi, and the new species Macelli-
cephala galapagensis from the Galapagos,
and from the Santa Catalina Channel, the
new genus Natopolynoe for N. kensmithi.
Included also was a summary of the poly-
noid polychaetes collected by DSRV Alvin
in the Galapagos Rift in 1979 and East Pa-
cific Rise at 21°N in 1982, along with a key
to the subfamilies, genera, and species.

From the deep seep-sites at the Florida
escarpment in the Gulf of Mexico, Petti-
bone (1986) added a new species of Bran-
chipolynoe, B. seepensis, commensal in the
mantle cavities of the Gulf of Mexico mus-
sels, along with a key to the two species of
the genus.

From the hydrothermal vents in the
Northeast Pacific Explorer and Juan de Fuca
Ridges, Pettibone (1988) added two new
species: Lepidonotopodium piscesae in the
Lepidonotopodiinae, and Opisthotrochop-
odus tunnicliffeae in Branchinotogluminae.

VOLUME 102, NUMBER 1

A new species of Macellicephala from the
Antarctic, M. australis, was added by Wu
and Wang (1987).

Thus to the summary by Levenstein
(1984), four new subfamilies have been
added (total 10), nine new genera (total 28),
and 20 new species (total 58). Four of the
new genera were added to Macellicephali-
nae (total 14) and seven new species (total
28). Two of the new species were referred
to Macellicephala, one to Bathykermadeca.

Additional polynoids were collected by
the Mariana Back-Arc Basin Expedition
during April and May 1987, where active
hydrothermal vents were found in the west-
ern Pacific off the Mariana Islands about
18°N and 144°E. The polynoid polychaetes
were collected during dives of DSRV Alvin
and sent to me for study by Robert Hessler,
along with information on the collecting
sites. Four species of polynoids were col-
lected on two A/vin dives: dive 1836, re-
ferred to as Snail Pits in Burke Field (1
species), and dive 1843, designated as Alice
Springs (3 species). The polynoids agree with
some of the subfamilies and genera that were
previously described from the hydrother-
mal vents in the eastern Pacific.

Burke Field (Alvin dive 1836, 27 Apr,
18°10.9'N, 144°43.2’E, 3660 m) was an ex-
tensive area of low temperature vents at the
southern end of an active spreading zone
and dominated by pillow basalts. The Snail
Pits vent had a water temperature of 15°C,
with small, dense aggregations of “hairy”
snails that clogged the vent openings. The
hot vent water was crystal clear. The single
polynoid collected is described below under
Branchinotogluminae: Branchinotogluma
burkensis, new species.

Alice Springs (A/vin dive 1843, 4 May,
18°12.6'N, 144°42.4’E, 3640 m) was an ex-
tensive area dominated by pillow basalts
with sharp cliffs up to 40 m high. Some low
temperature vent sites (10—25°C) were char-
acterized by cloudy, particle-filled water and
dense concentrations of white anemones.
Some high temperature vents (up to 250°C)

139

had dense aggregations of “hairy” snails,
crabs, and shrimps, with emerging crystal
clear water. The three species of polynoids
collected are described below under three
subfamilies: Macellicephalinae: Levenstei-
niella raisae, new species (1 specimen): Lep-
idonotopodinae: Lepidonotopodium minu-
tum, new species (9 specimens); and
Branchinotogluminae: Opisthotrochopodus
marianus, new species (12 specimens).

The types are deposited in the National
Museum of Natural History, Smithsonian
Institution (USNM).

Subfamily Macellicephalinae
Hartmann-Schroder, 1971
emended Pettibone, 1976

Levensteiniella Pettibone, 1985

Levensteiniella was erected for the single
species L. kincaidi from off western Mexico
at 21°N and the Galapagos Rift (Pettibone
1985d:741). An additional species is added
from the Mariana Back-Arc Basin.

Levensteiniella raisae, new species
Figs. 1, 2

Material examined. —Western Central
Pacific in Mariana Back-Arc Basin, Alvin
dive 1843, 4 May 1987, Alice Springs,
18°12.6'N, 144°42.4’E, 3640 m, R. Hessler,
collector, holotype (USNM 118362).

Description. —Length of holotype 21 mm,
width 11 mm with setae, segments 27, last
one very small (Fig. 1D). Body flattened,
tapering anteriorly and more so posteriorly,
with parapodia about as long as body width.
Elytra 11 pairs, on segments 2, 4, 5, 7, con-
tinuing on alternate segments to 21, large,
covering dorsum, oval to subreniform; most
of surface covered with unusual type of
macro- and micro-tubercle-papillae, with
chitinous bases and bulbous papillar tips,
larger ones on posterior border; bases cov-
ered with brownish foreign material and
bacterial ‘‘hairs” (Fig. 1G). Elytrophores
large and prominent (Figs. 1A, C, 2A, D).
Dorsal cirri on segments lacking elytra, with
cylindrical cirrophores on posterodorsal

140 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig: 1.

—— ee

Levensteiniella raisae, holotype, USNM 118362: A, Dorsal view of anterior end; B, Ventral view of

anterior end, bases of palps only shown; C, Dorsal view of left half of segments 8 (cirrigerous) and 9 (elytragerous);
D, Dorsal view of posterior end, including parapodia of segments 24-27, last one very small; E, Ventral view
of left side of segments 1 1-13 showing ventral papillae on segments 11 and 12, tip of long papilla on segment
12 hidden from view; F, Dorsal row of border papillae and jaws of pharynx (cut open), inner view; G, Left
elytron, with detail of macro- and micro-tubercle-papillae (not to scale). Scales = 1.0 mm for A-E; 0.5 mm for

F; 2.0 mm for G.

faces of notopodia; long styles extending be-
yond neurosetae, with filamentous tips; dor-
sal tubercles prominent, nodular to truncate
(Figs. 1C, D, 2B, C). Dorsal transverse cil-
lated ridges, up to five per segment, con-
tinuing onto bases of elytrophores and dor-
sal tubercles (Fig. 1A, C, D).

Prostomium oval, deeply bilobed: lobes
projecting anteriorly, with frontal filaments;
ceratophore of median antenna short, oval,
inserted in anterior notch, with style short,
tapered, shorter than palps; palps stout, long,
tapered; lateral antennae and eyes lacking
(pair of tannish spots resembling “eyes’’)

VOLUME 102, NUMBER 1

141

E

Fig. 2. Levensteiniella raisae, holotype, USNM 118362: A, Right elytragerous parapodium of segment 2,
anterior view, acicula dotted; B, Right cirrigerous parapodium of segment 3, posterior view; C, Same, from
segment 8; D, Right elytragerous parapodium of segment 9, anterior view, acicula dotted; E, Notosetae; F,
Supraacicular neuroseta, with detail of parts; G, Subacicular neurosetae. Scales = 1.0 mm for A—D; 0.1 mm for

E-G.

(Fig. 1A). Tentacular segment not distinct
dorsally; tentaculophores lateral to prosto-
mium, lacking setae; each with pair of ten-
tacular cirri, dorsal one about as long as
palps, ventral one shorter, both with long
slender tips; large bilobed facial tubercle
medial to palps (Fig. 1A, B).

Second or buccal segment bearing first pair
of elytrophores, biramous parapodia, with
notopodium shorter than neuropodium;
ventral buccal cirri similar to tentacular cir-
ri, longer than following ventral cirri (Figs.
1A, B, 2A). Pharynx not extended (cut open);
seven pairs of border papillae, none elon-
gated; two pairs of light amber-colored, en-
tire jaws (not denticled) (Fig. 1F).

Both rami of biramous parapodia well de-

veloped, notopodium on dorsoposterior side
of larger neuropodium, rounded basally with
projecting acicular process on lower side;
neuropodium with conical presetal lobe with
projecting acicular process, postsetal lobe
shorter, rounded, deeply cut on dorsal side
(Fig. 2B-D). Notosetae very numerous,
forming radiating bundle, short to longer,
much stouter than neurosetae, straight to
slightly curved, tapering to bare blunt tips,
with widely spaced spines along one side,
variable in number (1-18) (Fig. 2E); tips
often covered with reddish-brown foreign
material and bacterial “‘hairs.’? Neurosetae
slender, numerous, forming fan-shaped
bundle; supraacicular neurosetae very slen-
der, with longer spinous regions, consisting

142

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Lepidonotopodium minutum, A-F, holotype, USNM 118363; G, paratype, USNM 118364: A, Dorsal
view of anterior end, right dorsal and left ventral tentacular cirri missing; B, Right 1st elytron from segment 2,
with detail of micropapillae; C, Right 2nd elytron from segment 4; D, Right 6th elytron from segment 11, with
detail of micropapillae; E, Right 10th elytron from segment 19; F, Right 11th elytron from segment 21; G,
Ventral view of extended pharynx. Scales = 0.5 mm for A; 1.0 mm for B—F; 0.2 mm for G.

of two rows of long, widely spaced spines
along borders, and with shorter spines near
tapered tips (Fig. 2F); subacicular neuro-
setae with shorter spinous regions, stouter,
with minute, close-set spines along one bor-
der, and with slightly hooked bare tips (Fig.
2G). Ventral cirri short, tapered, attached
on middle of neuropodia (Fig. 2C, D).

Anus dorsal, medial to parapodia of last
two segments (26, 27); pygidium small, rect-
angular, with pair of long anal cirri (Fig. 1E).
Two pairs of long ventral papillae on seg-
ments 11 and 12, their tips sometimes ex-
tending dorsally between parapodia (Fig.
LE).

Etymology.—The species, as well as the
genus, is named for Raisa J. Levenstein in
recognition of her numerous contributions
to the study of the deep-sea polychaetes.

Remarks. —Levensteiniella raisae differs
from L. kincaidi in the form of the elytra,
where the surfaces are nearly covered with
unusual macro- and micro-tubercle-pa-
pillae; in L. kincaidi the elytra are more
delicate with only scattered micropapillae.
The notosetae differ in having more distinct
and fewer spines along the border, rather
than more numerous spinous rows in L.
kincaidi.

Subfamily Lepidonotopodiinae
Pettibone, 1983
Lepidonotopodium Pettibone, 1983

Lepidonotopodium was erected for L. fim-
briatum Pettibone, 1983 from the East Pa-
cific Rise at 21°N. Two additional species
were added by Pettibone (1984b) from the
Galapagos Rift, as well as 21°N: L. william-
sae and L. riftense, and a fourth species from
the North East Pacific Explorer and Juan de
Fuca Ridges by Pettibone (1988), L. pisce-
sae. An additional species is added from the
Mariana Back-Arc Basin.

Lepidonotopodium minutum, new species
Figs. 3, 4

Material.—Western Central Pacific in
Mariana Back-Arc Basin, A/vin dive 1843,
4 May 1987, Alice Springs, 18°12.6'N,
144°42.4’E, 3640 m, R. Hessler, collector,
holotype (USNM 118363) and 8 paratypes
(USNM 118364, 118365).

Description. —Holotype 7.5 mm long, 3.5
mm wide with setae, with 23 segments;
slightly smaller paratypes 6—6.5 mm long,
3-4 mm wide, with 22—23 segments; small-
est paratype 5.2-5.5 mm long, 3-3.2 mm
wide, with 21-22 segments. Body short,

VOLUME 102, NUMBER 1

143

See |

H

Fig. 4. Lepidonotopodium minutum, holotype, USNM 118363: A, Right elytragerous parapodium from
segment 2, anterior view; B, Right cirrigerous parapodium from segment 3, posterior view; C, Right elytragerous
parapodium from segment 9, anterior view; D, Right cirrigerous parapodium from segment 10, posterior view;
E, Right elytragerous parapodium from segment 11, posterior view, notopodium and neuropodium separated,
acicula dotted; F, Upper, middle and lower notosetae; G, Short anterior notosetae; H, Upper and lower neurosetae.

Scales = 0.2 mm for A—-E; 0.1 mm for F—H.

suboval in outline, flattened dorsoventrally
and slightly tapered anteriorly and poste-
riorly. Elytra 11 pairs, on segments 2, 4, 5,
7, continuing on alternate segments to 21,
attached eccentrically on prominent elyt-
rophores (Figs. 3A, 4A, C, E). Elytra oval
to subreniform, imbricated, covering dor-
sum, variable in size, last pair very small,
Opaque, smooth, with branching “‘veins’’
emanating from place of attachment to elyt-
rophores. Surface of first pair of elytra and
posterior half on rest of elytra covered with
scattered micropapillae, globular with short
to longer tapered or cylindrical bases (Fig.
3B-F). Elytral surface and borders covered
with foreign material and bacterial “‘hairs.”’
Dorsal cirri witih cylindrical cirrophores at-
tached on dorsoposterior sides of notopo-
dia; styles tapered, extending beyond tips of
neurosetae: dorsal tubercles elongated, in-
flated (Fig. 4B, D).

Prostomium deeply bilobed, anterior
lobes subtriangular with small frontal fila-
ments; ceratophore of median antenna in
anterior notch, short, bulbous, with short,
subulate style; palps cylindrical, with ta-
pered tips, nearly two times longer than pro-
stomium; lateral antennae and eyes lacking
(Fig. 3A). Tentacular segment not visible
dorsally; tentaculophores lateral to prosto-
mium, achaetous, each with pair of tenta-
cular cirri, dorsal ones nearly as long as palps,
ventral ones slightly shorter, both with long
tapering tips (Fig. 3A). Without distinct fa-
cial tubercle.

Second or buccal segment with first pair
of elytrophores, biramous parapodia, and
ventral or buccal cirri attached basally on
prominent cirrophores lateral to ventral
mouth, with styles similar to tentacular cir-
ri, longer than following ventral cirri (Figs.
3A, 4A). Opening of extended pharnyx en-

144

Table 1.—Comparison of Lepidonotopodium rif-
tense and L. minutum.

L. riftense L. minutum

Borders of with short pa- lack papillae
notopodi- pillae
al bracts
Papillae of 7-9 pairs, none 7 pairs, unequal,
pharynx elongated some elongated
Jaws of with numerous smooth, without
pharynx teeth teeth
Long ven- 2 pairs on seg- not present
tral pa- ments 11 &
pillae 12 (on some
specimens)

circled with seven pairs of unequal-sized pa-
pillae; dorsal with three medial ones long,
tapered, and two short lateral pairs; ventral
with three middle ones and lateral pair short,
next to lateral pair long, tapered; two pairs
of hooked jaws, without teeth or serrations
(Fig. 3G).

Biramous parapodia with shorter noto-
podia on anterodorsal sides of longer neu-
ropodia (Fig. 4A—D). Notopodia subconi-
cal, with projecting acicular lobe, hidden
anteriorly by very numerous notosetae; well-
developed flaring bracts enclosing bases of
notosetae; ventro-anterior and dorso-pos-
terior bracts attached basally to acicular lobe
(Fig. 4A, C, E). Neuropodia with conical
presetal acicular lobe and shorter, rounded
postsetal lobe (Fig. 4B—E); distal borders with
bacterial “‘hairs.”’ Notosetae very numer-
ous, forming thick radiating bundle, stouter
than neurosetae, with widely spaced spines
along one side; notosetae arranged in four
groups: upper (about 8), middle (11), lower
(8) (Fig. 4F), and short anterior group (Fig.
4G). Notosetae covered with foreign ma-
terial and bacterial “‘hairs.’’ Neurosetae nu-
merous, forming fan-shaped bundle; neu-
rosetae with two rows of numerous spines
along one side and slightly hooked bare tips;
upper neurosetae longer, with longer spi-
nous regions (Fig. 4H). Ventral cirri at-
tached on middle of posterior face of neu-
ropoda, short, tapered (Fig. 4B—E).

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Segmental or nephridial papillae not ob-
vious. Without elongated ventral papillae
on segments 11 and 12. Pygidium small,
rectangular, between posterior parapodia,
with pair of long anal cirri.

Etymology.—The specific name minu-
tum refers to the relatively small size of the
species.

Remarks.— Lepidonotopodium minutum
is closest to L. riftense. Both are of small
size with relatively few segments (up to 7
mm in length and 23 segments). The elytra
lack macrotubercles and micro-tubercles but
have clavate micropapillae, differing from
the other species of Lepidonotopodium. The
notosetae have widely spaced spines along
one side (also found in L. fimbriatum). The
two species may be separated by the char-
acters indicated in Table 1.

Subfamily Branchinotogluminae
Pettibone, 1985
Branchinotogluma Pettibone, 1985

Branchinotogluma was established by
Pettibone (1985c) for three species from the
hydrothermal vents off western Mexico at
21°N and the Galapagos: B. hessleri, B. san-
dersi, and B. grasslei, with B. hessleri as the
type species. B. grassleiand B. sandersi were
also reported from the North Pacific Ex-
plorer and Juan de Fuca Ridges by Petti-
bone (1988). An additional species is added
from the Mariana Back-Arc Basin.

Branchinotogluma burkensis, new species
Figs. 5, 6

Material. —Western Central Pacific in
Mariana Back-Arc Basin, Alvin dive 1836,
27 Apr 1987, Burke Field, Snail Pits,
18°10.9’N, 144°43.2’E, 3660 m, R. Hessler,
collector, holotype (USNM 118366).

Description. — Length of holotype 30 mm,
width with setae 16 mm, segments 21, with
additional small parapodium on right side
(Fig. 5B). Body flattened, tapering anteriorly
and posteriorly, with parapodia longer than
body width. No color except for golden-col-
ored setae. Dorsum with transverse ciliated

VOLUME 102, NUMBER 1

145

~

——

G

Fig. 5. Branchinotogluma burkensis, holotype. USNM 118366: A. Dorsal view of anterior end: B, Dorsal
view of posterior end including segments 18-21: C, Ventral view of left side of segments 1 1—13, showing small
segmental ventral papillae: D. Right 1st elytron from segment 2: E, Right 7th elytron from segment 13; F, Dorsal
view of left side of cirrigerous segment 14, only base of style of dorsal cirrus shown: G. One of jaws removed
from body: H, Right elytragerous parapodium from segment 2, anterior view, acicula dotted. Scales = 2.0 mm

for A—E- 0.5 mm for F. G: 1.0 mm for H.

bands, 2-4 per segment, extending onto
elytrophores. dorsal tubercles. and bran-
chial bases (Fig. 5A, B, F).

Elytra 10 pairs, attached on large bulbous
elytrophores on segments 2. 4. 5. 7. con-
tinuing on alternate segments to 19 (Figs.
5A, B, H, 6B). Elytra large. overlapping.
covering dorsum, round to oval. delicate.
with branched “‘veins.”” without tubercles or
papillae (Fig. 5D. E). Dorsal cirm on non-
elytragerous segments, with cylindrical cir-
rophores on posterodorsal sides of noto-

podia. and long smooth styles with slender
tips extending beyond tips of neurosetae:
dorsal tubercles nodular. projecting (Figs.
5B, F, 6A, C). Branchiae arborescent, in two
groups: on dorsal bases of notopodia and
on lateral sides of elytrophores and dorsal
tubercles (Figs. 5F, 6B). Branchiae begin-
ning on segment 3 as two small groups (Fig.
6A). becoming larger (Figs. 5F, 6B. C), con-
tinuing to posterior end and becoming very
small (Fig. 5B).

Bilobed prostomium with prominent cy-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

146

(4
D 46
y
i
sense Z | LF
i 2 Z
E | an
B / / g
. i | i
\ \ { f
\ } i :
iy, * ema oe F |; Y
~ i \ ; ;
\ I
= Sy j f | ¥ ; Y,
SS I}
: } | E
2S j | £
SSG j | j
So | ¢
/
gs o~ Z ! | é
— ———S- bs
—— | [

Cc —— = Sg
= E Ee

Fig. 6. Branchinotogluma burkensis, holotype. USNM 118366: A, Right cirrigerous parapodium from seg-
ment 3, posterior view; B, Right elytragerous parapodium from segment 9, anterior view, acicula dotted; C,
Right cirrigerous parapodium from segment 10, posterior view; D, notosetae; E, Tip of long supraacicular
neuroseta, with detail of parts; F. Subacicular neuroseta, with detail of parts. Scales = 1.0 mm for A-C; 0.1 mm

for D-F.

smooth, about three times length of prosto-
mium; without eyes, but non-ocular paired
oval shaded areas present (Fig. 5A). Ten-
tacular segment not distinct dorsally; ten-

lindrical anterior lobes with terminal fila-
ments; median antenna with bulbous cer-
atophore in anterior notch and short style
tapered to slender tip: palps long, tapered,

VOLUME 102, NUMBER 1

taculophores lateral to prostomium, achae-
tous, each with pair of dorsal and ventral
tentacular cirri about as long as palps, dorsal
tentacular cirri slightly longer than ventral
ones (Fig. 5A). Second or buccal segment
with first pair of large elytrophores, bira-
mous parapodia, and long, slender buccal
cirri similar to tentacular cirri, longer than
following ventral cirri; notopodium with
prominent bract fused with acicular lobe (not
distinct, as usual in genus) and small bundle
of stout notosetae; neuropodium similar to
following (Fig. 5A, H). Pharynx not everted
(cut out), with five border papillae around
opening: three dorsal and two ventral; two
pairs of prominent dark reddish amber-col-
ored jaws, minutely denticled on inner bor-
der (about 50) (Fig. 5G).

Biramous parapodia with notopodia
shorter than neuropodia, both rami with
golden-colored setae (Figs. 5H, 6A—C). No-
topodia conical, with prominent projecting
acicular processes, without notopodial bracts
except on segment 2 with bract fused to acic-
ular process. Notosetae numerous, forming
radiating bundles, not as long as but much
stouter than neurosetae, straight, tapered,
acicular, shorter ones smooth, longer ones
with some spines near distal tips (Fig. 6D).
Larger and longer neuropodium with sub-
conical presetal acicular lobe with project-
ing acicular process and shorter rounded
postsetal lobe, lobes deeply separated dor-
sally and ventrally. Neurosetae very nu-
merous, forming fan-shaped bundles, slen-
der, with slightly hooked tips. Supraacicular
neurosetae with two rows of widely spaced
prominent spines and finely spinous hood
on slightly hooked tip (Fig. 6E).Subacicular
neurosetae finely spinous along border, with
slightly hooked tip (Fig. 6F). Ventral cirri
short, tapering, attached on middle of neu-
ropodia (Fig. 6A-C).

Posterior end not modified or com-
pressed. Pygidium small lobe between para-
podia of last segment (21), with dorsal anus
and pair of anal cirri (Fig. 5B). Segmental
ventral papillae seven pairs, small squarish,

147

on ventral bases of parapodia of segments
11-17, none elongated (Fig. 5C).

Etymology. —The species is named bur-
kensis for the collecting area, Burke Field
on the Mariana Back-Arc Basin.

Remarks.—B. burkensis is closest to B.
grasslei Pettibone (1985c). Both species have
a pair of anal cirri and posterior segments
that are not compressed or modified; bran-
chiae are present on all segments from 3 on;
and both lack elongated ventral papillae. B.
burkensis has seven pairs of small segmental
ventral papillae on segments 11-17, com-
pared to five pairs on segments 11-15 in B.
grasslei. In B. burkensis, the notopodial bract
on segment 2 is fused with the acicular lobe
and not distinct, as in B. grasslei, where the
notopodial bract encloses the acicular lobe
and notosetae (Pettibone 1985c, fig. 6A).
The long notosetae in B. burkensis have dis-
tal spines, whereas they are smooth in B.
grasslei.

Opisthotrochopodus Pettibone, 1985,
emended

Opisthotrochopodus was established by
Pettibone (1985c) for O. alvinus from the
hydrothermal vents off western Mexico at
21°N and the Galapagos. O. tunnicliffeae,
from the hydrothermal vents of the Ex-
plorer and Juan de Fuca Ridges, was added
by Pettibone (1988). A third species is added
from the Mariana Back-Arc Basin and the
genus 1s emended.

Diagnosis.—Body short, with 21 seg-
ments, first achaetous. Elytra 10 pairs, on
segments 2, 4, 5, continuing on alternate
segments to 19. Dorsal cirri with short cir-
rophores and long style, and dorsal tuber-
cles, in line with elytrophores, on segments
3, 6, continuing on alternate segments to 20
and 21. Arborescent branchiae two pairs per
segment, attached to lateral sides of elyt-
rophores, dorsal tubercles, and dorsal side
of notopodia, beginning on segment 3 and
continuing to segment 15-18. Prostomium
bilobed, with triangular anterior lobes bear-

148

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 7.

Opisthotrochopodus marianus, A-I, paratype, USNM 118369; J, holotype, USNM 118367: A, Dorsal

view of anterior end; B, Ventral view of segments 12-18; C, Right Ist elytron from segment 2; D, Right 2nd
elytron from segment 4; E, Right 4th elytron from segment 7; F, Right 6th elytron from segment 11; G, Right
9th elytron from segment 17; H, Right 10th elytron from segment 19; I, Ventral view of extended pharynx; J,
Dorsal jaws, inner view. Scales = 0.5 mm for A, B, I; 0.5 mm for C—H; 0.1 mm for J.

ing frontal filaments, with ceratophore of
median antenna in anterior notch, with
paired ventral palps, without lateral anten-
nae or eyes. First or tentacular segment not
visible dorsally; tentaculophores lateral to
prostomium, achaetous, each with dorsal
and ventral tentacular cirri. Second or buc-
cal segment with first pair of elytra, bira-
mous parapodia, and ventral or buccal cirri
attached to basal parts of parapodia lateral
to mouth; styles longer than following ven-
tral cirri. Parapodia biramous, with noto-
podia shorter than neuropodia. Notopodia
of elytragerous segments 2-17 with promi-
nent rounded bracts enclosing conical acic-
ular lobes and notosetae, only on segment
2 or lacking. Neuropodia with longer con-
ical presetal and shorter rounded postsetal
lobes. Notosetae stouter than neurosetae,
straight, acicular, smooth or with spines.
Neurosetae long, slender, finely spinous,
with slightly hooked tips. Ventral cirri short,
tapered. Posterior 3 to 6 segments modified,
including wheel organs on segment 20 or
lateral lamellae on segment 21. Pharynx with
5 to 8 border papillae; 2 pairs of dorsal and
ventral hooked jaws, with or without den-

ticles. Elongated ventral papillae on seg-
ment 12 and rounded ventral lamellae on
some following segments. Pygidium with or
without pair of anal cirri.

Opisthotrochopodus marianus, new species
Figs. 7-9

Material.—West Central Pacific, Mar-
iana Back-Arc Basin, A/vin dive 1843, 4 May
1987, Alice Springs, 18°12.6'N, 144°22.4’E,
3640 m, holotype (USNM 118367), para-
type (USNM 118369), paratype (USNM
118368), 9 paratypes (3 small, USNM
118370).

Description. —Holotype 6 mm long, 3 mm
wide, with 21 segments. Two adult para-
types 5—5.5 mm long, 3 mm wide, with 21
segments. Two young paratypes 2.5—3 mm
long, 2-3 mm wide, with 13-14 segments
plus small growing zone. Body flattened, ta-
pering slightly anteriorly, anterior parapo-
dia (segments 2-13) longer, tapering grad-
ually (segments 14-18), narrowing and
projecting posteriorly with extra long setae
(segments 19-21) (Figs. 7B, 9H, I). Dorsum
with transverse ciliated bands, 1—2 per seg-

VOLUME 102, NUMBER 1

149

Fig. 8. Opisthotrochopodus marianus, paratype, USNM 118369: A, Right elytragerous parapodium from
segment 2, anterior view, acicula dotted; B, Right cirrigerous parapodium from segment 3, posterior view, style
of dorsal cirrus broken off; C, Left elytragerous parapodium from segment 9, anterior view, acicula dotted; D,
Left cirrigerous parapodium from segment 10, posterior view; E, Short and long notosetae; F, Supraacicular
neuroseta; G, Subacicular neurosetae; H, Right elytragerous parapodium from segment 15, anterior view; I,
Right cirrigerous parapodium from segment 16, posterior view; J, Right elytragerous parapodium from segment
17, posterior view; K, same, anterior view. Scales = 0.3 mm for A-D, H-J; 0.1 mm for E—G; 0.2 mm for K.

ment, extending onto elytrophores and dor-
sal tubercles (Figs. 7A, 9H).

Elytra and prominent bulbous elytro-
phores 10 pairs (Figs. 7A, 8A, C, H, J, K,
9H). Elytra large, overlapping, covering
dorsum, except for posterior modified par-
apodia, round to oval, larger on middle ely-
tra, very small on segments 17 and 19, del-
icate, showing branched “‘veins”’ emanating
from scars of attachment to elytrophores,
without tubercles or papillae (Figs. 7C—H,
9H). Dorsal cirri with cylindrical cirro-
phores on posterodorsal sides of notopodia,
long smooth styles with slender tips ex-
tending beyond setae; dorsal tubercles elon-
gate, tapered, with long cilia (Fig. 8B, D, I).
Branchiae delicate, arborescent, on lateral
bases of elytrophores and dorsal tubercles
and dorsal bases of notopodia, beginning on

segment 3 as two small groups (Fig. 8B),
becoming larger (Fig. 8C, D), and continu-
ing to segment 15 as single small group (Fig.
8H).

Prostomium bilobed, anterior lobes tri-
angular with slender frontal filaments; me-
dian antenna with bulbous ceratophore in
anterior notch, with slender short style; palps
stout, tapered, about two times longer than
prostomium; without eyes (Fig. 7A). Ten-
taculophores lateral to prostomium, achae-
tous, with 2 pairs of tentacular cirri, dorsal
pair longer than palps, ventral pair shorter
(Fig. 7A). Second or buccal segment with
first pair of prominent elytrophores, bira-
mous parapodia, similar to but smaller than
following: ventral buccal cirri similar to ten-
tacular cirri, longer than following ventral
cirri; notopodium without notopodial bract

150 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 9. Opisthotrochopodus marianus, A—G, paratype, USNM 118369; H-I, paratype, USNM 118368: A,
Right cirrigerous parapodium from segment 18, anterior view, acicula dotted; B, Right elytragerous parapodium
from segment 19, anterior view, acicula dotted, elytrophore not shown; C, Right cirrigerous parapodium from
segment 20, anterior view, acicula dotted; D, Tips of long notosetae from same; E, Neuroseta from same; F,
Left cirrigerous parapodium from segment 21, anterior view, acicula dotted; G, Same, cirrigerous parapodium
from segment 21, anterior view, acicula dotted; G, Same, enlarged: H, Dorsal view of posterior end including
segments 19-21, acicula dotted: I, same, ventral view. Scales = 0.3 mm for A-C, F; 0.1 mm for D, E, G; 0.2

mm for H, I.

(Figs. 7A, 8A). Notosetae stout, similar to
those of following segments; lower neuro-
setae with capillary tips, rest similar to up-
per neurosetae of following segments. Ex-
tended pharynx encircled with four pairs of
papillae, dorsal with two medial ones larger
and lateral ones smaller: ventral with four
small papillae; two pairs of curved reddish
jaws occupying most of opening, without
denticles on inner border (Fig. 7I, J).
Parapodia of segments 2-16 biramous,
with notopodia shorter than neuropodia,
both rami with light yellow amber-colored

setae (Fig. 8A—D, H, I). Notopodia rather
long, conical, without notopodial bracts en-
closing acicular lobe and notosetae (as in O.
alvinus). Notosetae numerous, stouter than
neurosetae, short, smooth and longer,
smooth or with faint spinous rows, tapered
to blunt tips (Fig. 8E). Larger neuropodia
with presetal lobe long, subconical, with
projecting acicular process, postsetal lobe
Shorter, rounded. Neurosetae numerous,
long, slender, wider basally, with slightly
hooked tips. Supraacicular neurosetae more
slender, with longer spinous regions (Fig.

VOLUME 102, NUMBER 1

Table 2:—Comparison of three species of Opisthotrochopodus.

151

O. alvinus Pettibone,
1985c (Figs. 7-9)

O. tunnicliffeae Pettibone,
1988 (Figs. 6-9)

O. marinus n. sp. (Figs. 7-9)

Notopodial bracts on
elytragerous parapo-
dia

Branchiae on segments

Pharynx:

Papillae

Jaws

Segment 20:
Notopodia

Neuropodia

segments 2-17 (Figs.
7A, G)

3-17

5 small: 3 dorsal, 2
ventral

with minute denticles
(Fig. 8H, I)

achaetous (Fig.
9B, F—L)

with “wheel” organ,
incl. stout, acicular

segment 2 only (Fig.
6A, C)

short notosetae (Figs.
8B, C, 9A-F)

with “balloon” organ
including stout har-

without

3-15

8: 4 dorsal, 2 medial
larger; 4 ventral small

without denticles (Fig.
7I, J)

long notosetae
(Fig. 9C-E)

long, slender, acicular
lobe & short capillary

& slender spinous

neurosetae _.
Ventral cirrus without
Segment 21:
Lamellae small notopodial
& neuropodial
(Fig. 9M, N).
Neurosetae few, short
Ventral cirrus without

8F). Subacicular neurosetae decreasing in
length ventrally, finely spinous more basal-
ly, with longer spines distally; few shorter
lower ones with capillary tips (Fig. 8G).
Ventral cirri short, tapered, attached on
middle of neuropodia (Fig. 8B—D, H, I).

Elongated ventral papillae on segment 12,
extending posteriorly to segment 15 (Fig.
7B); 6 pairs of short, flat ventral lamellae
on segments 13-18, ciliated on upper side
(Figs. 7B, 8H, J, K).

Parapodia from segment 15 on more or
less modified. Parapodium 15 with only
small branchia on lateral side of elytro-
phore; notopodium almost as long as neu-
ropodium; supraacicular neurosetae with
capillary tips (Fig. 8H). Parapodium 16, and
following parapodia, without branchiae; all
neurosetae with capillary tips (Fig. 81). Par-
apodium 17 with small neuropodium short-
er than ventral cirrus, with bundle of long

poon & slender spi- neurosetae
nous neurosetae
present without

without (Fig. 9C) large lateral neuro-

podia (Fig. 9G-—I)

same
present

long capillary
without

slender capillary neurosetae; notopodium
small, with three shorter and seven longer
stout notosetae extending to tip of neuro-
podium (Fig. 8J, K).

Parapodium 18 with small notopodial
acicular lobe fused to cirrophore of dorsal
cirrus, ciliated on lower side, with few (4)
short stout notosetae on upper side; conical
neuropodium shorter than ventral cirrus,
ciliated on upper side, with small bundle of
long capillary neurosetae (Fig. 9A).

Parapodium 19 with elytrophore and
small elytron, inflated oval notopodium with
short acicular lobe and few (3-4) short stout
notosetae near tip, ciliated on underside;
neuropodium similar to preceding parapo-
dium, with small bundle of long capillary
neurosetae (Fig. 9B, H, I).

Parapodium 20 directed posteriorly, with
small notopodial acicular lobe fused to cir-
rophore of dorsal cirrus; neuropodium with

£52

long slender acicular lobe nearly as long as
dorsal cirrus, with short ventral cirrus (Fig.
9C, H, I). Long notosetae wider subdistally,
spinous, tapering to slender tips (Fig. 9D).
Shorter neurosetae slender, wider subdis-
tally, spinous, tapering to capillary tips (Fig.
9E).

Parapodium 21 directed posteriorly, with
notopodial acicular lobe fused to cirrophore
of dorsal cirrus, without notosetae; neuro-
podial conical acicular lobe with slender
capillary neurosetae and large, oval, delicate
lateral lamella; without ventral cirrus (Fig.
9F-I). Pygidium small rounded lobe with
dorsal anus medial to parapodia of segment
21, and pair of long anal cirri (Fig. 9H, I).

Etymology.—The specific name mari-
anus is based on the collecting site of the
Mariana Back-Arc Basin.

Remarks. —O. marianus can be separated
from the other two species of Opisthotro-
chopodus according to Table 2.

Acknowledgments

My thanks go to the members of the Mar-
iana Back-Arc Basin Expedition, the crew
of the Alvin, and especially Robert R. Hes-
sler and Michel A. Boudrias, who sent me
the polynoid specimens and furnished sta-
tion data and information on the collecting
area. The manuscript benefited from the re-
views by James A. Blake and Jerry D. Ku-
denov.

Literature Cited

Levenstein, R. J. 1975. [The polychaetous annelids
of the deep-sea trenches of the Atlantic sector
of the Antarctic Ocean.]—Trudy Institut
Okeanologii P.P. Shirshov Academii Nauk SSSR
103:119-142 [in Russian, English summary].

. 1978a. [Annelida (Polychaeta) from the deep
waters of the Pacific region of Antarctica.]—
Trudy Institut Okeanologii P.P. Shirshov Aca-
demii Nauk SSSR 113:73-87 [in Russian, En-
glish summary].

. 1978b. [Polychaetes of the family Polynoidae
(Polychaeta) from the deep-water trenches of the
western part of the Pacific.|—Trudy Institut

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Okeanologii P.P. Shirshov Academii Nauk SSSR
112:162-174 [in Russian, English summary].

1981. [Some pecularities in the distribution
of polychaetes of the family Polynoidae in the
Canadian basin of the Arctic Ocean.]—Trudy
Institut Okeanologii P.P. Shirshov Academii
Nauk SSSR 115:26-36 [in Russian, English
summary].
. 1982a. [On the polychaete fauna of the family
Polynoidae from the trench of Japan.]— Trudy
Institut Okeanologii P.P. Shirshov Academii
Nauk SSSR 117:59-62 [in Russian, English
summary].

1982b. [New genera of the subfamily Ma-
cellicephalinae (Polychaeta, Polynoidae) from
the Tasman Trench]—Zoologichesky Zhurnal
61(9):1291-1296 [in Russian, English summa-
ry]

1983. [Macellicephaloides improvisa sp. n.

(Polychaeta, Polynoidae) from the Kurile-Kam-

chatka Trench]—Zoologichesky Zhurnal 62(9):

1419-1421 [in Russian, English summary].

. 1984. On the ways of formation of the deep-

sea polychaete fauna of the family Polynoidae.

Pp. 72-85, in P.A. Hutchings, ed., Proceedings

of the First International Polychaete Confer-

ence, Sydney, Australia, 1983. The Linnean So-
ciety of New South Wales.

Loshamn, A.-A. 1981. Descriptions of five polynoid
species (Polychaeta) from the coasts of Norway
and Sweden, including three new species, one
new genus and one new generic name.—Zoo-
logica Scripta 10(1):5—13.

Pettibone, M. H. 1976. Revision of the genus Ma-

cellicephala McIntosh and the subfamily Ma-

cellicephalinae Hartmann-Schroder (Polychae-
ta: Polynoidae). —Smithsonian Contributions to

Zoology 229:1-71.

1979. Redescription of Bruunilla natalensis
Hartman (Polychaeta: Polynoidae), originally
referred to Fauveliopsidae. — Proceedings of the
Biological Society of Washington 92(2):384-388.

1983. A new scale worm (Polychaeta: Poly-
noidae) from the hydrothermal rift area off
Western Mexico at 21°N.—Proceedings of the
Biological Society of Washington 96(3):392-399.

1984a. 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(1):226-239.

1984b. Two new species of Lepidonotopo-
dium (Polychaeta: Polynoidae: Lepidonotopo-
dinae) from hydrothermal vents off the Gala-
pagos and East Pacific Rise at 21°N.—
Proceedings of the Biological Society of Wash-
ington 97(4):847-863.

VOLUME 102, NUMBER 1

. 1985a. Polychaete worms from a cave in the
Bahamas and from experimental wood panels
in deep water of the North Atlantic (Polynoidae:
Macellicephalinae, Harmothoinae).—Proceed-
ings of the Biological Society of Washington
98(1):127-149.

1985b. An additional new scale worm (Po-
lychaeta: Polynoidae) from the hydrothermal rift-
area off Western Mexico at 21°N.— Proceedings
of the Biological Society of Washington 98(1):
157-164.

1985c. Additional branchiate scale-worms
(Polychaeta: Polynoidae) from Galapagos hy-
drothermal vent and rift-area off Western Mex-
ico at 21°N.— Proceedings of the Biological So-
ciety of Washington 98(2):447-469.

1985d. New genera and species of deep-sea
Macellicephalinae and Harmothoinae (Poly-
chaeta: Polynoidae) from the hydrothermal rift
areas of the Galapagos and western Mexico at
21°N and from the Santa Catalina Channel. —
Proceedings of the Biological Society of Wash-
ington 98(3):740-757.

153

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: Branchipolynoinae).—Pro-
ceedings of the Biological Society of Washington
99(3):444-451.

. 1988. New species and new records of scaled

polychaetes (Polychaeta: Polynoidae) from hy-

drothermal vents of the Northeast Pacific Ex-
plorer and Juan de Fuca Ridges. — Proceedings

of the Biological Society of Washington 101(1):

192-208.

Wu, B. & Wang, Y.-H. 1987. Two new species of
Polychaeta from South Ocean.—Acta Zootax-
onomica Sinica 12(1):23—29 [in Chinese and En-
glish].

Department of Invertebrate Zoology, Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 154-168

POLY NOIDAE AND SIGALIONIDAE (POLYCHAETA) FROM

THE GUAYMAS BASIN, WITH DESCRIPTIONS OF
TWO NEW SPECIES, AND ADDITIONAL RECORDS FROM
HYDROTHERMAL VENTS OF THE GALAPAGOS
RIFT, 21°N, AND SEEP-SITES IN THE GULF OF
MEXICO (FLORIDA AND LOUISIANA)

Marian H. Pettibone

Abstract. —Eight species of scaled polychaetes are reported from hydrother-
mal vents of the Guaymas Basin in the Gulf of California: Neoleanira racemosa
in the Sigalionidae and seven species of Polynoidae, including two new species,
Bathykurila guaymasensis, in Macellicephalinae, and Macellicephaloides alvini,
in Macellicephaloidinae. The latter genus and subfamily are reviewed, and a
Key to the seven species in the genus is provided. Included are additional
records of some species of Polynoidae from hydrothermal vents from the Ga-
lapagos and 21°N in the eastern Pacific, as well as seep-sites in the Gulf of

Mexico Florida Escarpment and off Louisiana.

A small collection of scaled polychaetes
from the Guaymas Basin in the Gulf of Cal-
ifornia or the Sea of Cortez was sent to me
for study by J. F. Grassle. Hot vents and
hydrocarbon seeps were discovered in this
area in 1980 and sampled in 1982 during
dives of the submersible DSRV Alvin (Lons-
dale 1984, Grassle 1986). The area, a con-
tinuation of the East Pacific Rise into the
Gulf of California, is covered by a thick
layer of pelagic muddy sulfide sediment. The
study site consists of mounds of hydrother-
mal precipitates in the spreading center
troughs at about 2000 meters. The fauna
included large numbers of the giant vesti-
mentiferans, Riftia, giant clams, such as Ca-
lyptogena pacifica, nuculanid bivalves, al-
vinellid polychaetes, as Paralvinella grasslei,
and fluffy mats of the large bacterium Beg-
giatoa.

Eight species of scaled polychaetes are re-
ported herein from the Guaymas Basin, sev-
en of them belonging to the Polynoidae and
one to the Sigalionidae: Neoleanira race-
mosa (Fauchald), previously reported from
the Guaymas Basin by Fauchald (1972, as

Sthenolepis). Of the polynoid species, five
were previously described by me from other
vent areas: one, Branchiplicatus cupreus,
from the tropical eastern Pacific at 21°N;
one, Lepidonotopodium riftense, the same
area and the Galapagos Rift; and three,
Branchinotogluma grasslei, B. sandersi, and
Levensteiniella kincaidi, from the above two
areas and also the Explorer and Juan de
Fuca Ridges. The other two polynoid species
are described below as new species: Bathy-
kurila guaymasensis, in Macellicephalinae,
and Macellicephaloides alvini, in Macelli-
cephaloidinae. The specimens were collect-
ed during A/vin dives in 1982 and 1985, in
washings from rock with worms, Bead Ex-
periment washings, washings from Rock
Chimney, washings from Riftia, Calypto-
gena, and alvinellid polychaetes, from Box
cores taken on the slopes of the mounds,
some with numerous nuculanid bivalves.
Some additional records of previously de-
scribed vent fauna are included in this re-
port, such as Branchipolynoe symmytilida,
removed from some Galapagos mussels and
sent to me in July 1985 by J. F. Grassle.

VOLUME 102, NUMBER 1

Also, specimens of Branchipolynoe seepen-
sis were removed by Ruth Turner from Gulf
of Mexico mussels collected in the seep-sites
at the Florida Escarpment and sent to me
in October 1986, by Barbara Hecker. An
additional specimen of this species was col-
lected in May 1987, by the US Navy sub-
marine NR-1 and R/V Gyre and sent to me
by T. H. Perkins. Additional vent polynoids
from the Galapagos and the eastern Pacific
Rise at 21°N, collected during A/vin dives
in 1979 and 1982, chiefly among fauna in
washings from mussels and crab traps, were
sent to the Smithsonian Sorting Center for
further sorting and transferred to me in De-
cember 1987. They supplement my pre-
vious studies on this group.

Types and additional specimens are de-
posited in the collections of the Department
of Invertebrate Zoology, National Museum
of Natural History, Smithsonian Institution
(USNM).

Family Polynoidae
Subfamily Branchipolynoinae Pettibone,
1984
Genus Branchipolynoe Pettibone, 1984,
emended Pettibone, 1986
Branchipolynoe symmytilida Pettibone

Branchipolynoe symmytilida Pettibone,
1984a:227, figs. 1-8.—Kenk & Wilson,
1985:264.—Desbruyéres et al. 1985:104,
113, 114.—Grassle, 1986:325, 327, 336.

Material. —East Central Pacific, from
dives of the A/vin on three vent areas along
the Galapagos Rift in 1979, associated with
deep-sea mussels, Branchimodiolus ther-
mophilus Kenk & Wilson: Mussel Bed,
00°47.89'N, 86°09.21'W: Dive 880, 21 Jan,
2493 m, from 9 mussels, 9 specimens
(USNM 98577). Dive 888, 13 Feb, 2493 m,
from mussel, 1 specimen (USNM 98578).
Dive 986, 3 Dec, 2494 m, mussel bucket
washings, 1 specimen (USNM 118168).
Dive 989, 6 Dec, 2482 m, from small mus-
sels, 5 specimens (USNM 118171). Dive
991, 8 Dec, 2490 m, washings from 68 mus-

155

sels in mesh-lined milk crates, 5 specimens,
(USNM 118170). Garden of Eden,
00°47.69'N, 86°07.74'W: Dive 883, 25 Jan,
2482 m, from mussel, 1 specimen (USNM
98579). Dive 884, 25 Jan, 2482 m, from 6
mussels, 7 specimens (USNM 98580) and
3 young specimens (USNM 118172). Dive
993, 10 Dec, 2518 m, slurp sample from
base of vestimentiferans, 1 specimen
(USNM 118166). Rose Garden, 00°48.25’N,
86°13.48'W: Dive 892, 17 Feb, 2454 m,
from 3 mussels, 3 specimens (USNM
98581). Dive 894, 19 Feb, 2457 m, from
12 mussels, 12 specimens (USNM 98582).
Dive 983, 30 Nov, 2457 m, mussel bucket
residue, 2 young specimens (USNM
118167). Dive 984, 1 Dec, 2451 m, mussel
washings left in clam bucket overnight, 13
specimens (USNM 118169); residue from
retrieved box containing vestimentiferans,
1 small specimen (USNM 118173).

Remarks.—Branchipolynoe symmytilida
has been found living commensally in the
mantle cavities of the giant deep-sea vent
mussels, Branchimodiolus thermophilus
Kenk & Wilson (Pettibone 1984a, Kenk &
Wilson 1985). Of the 37 polynoids taken
from 36 mussels (USNM 98577-98582), 1
had a length of 40 mm; 14, lengths of 29-
33 mm; 9, lengths of 21—25 mm; 10, lengths
of 15-20 mm; 2, lengths of 11 mm; and 1,
length of 6 mm. The two polynoids taken
from a single mussel had lengths of 25 and
15 mm. The additional 31 specimens from
8 Alvin dives, taken mostly from mussel
washings (USNM 118166-73), consisted of
smaller worms: adults up to 1 1 mm in length,
3 mm in width, with 20 segments and 10
pairs of elytra; juveniles 3—4 mm in length,
1.5—2.2 mm in width, with 14-17 segments;
and very young specimens 1.2—2 mm in
length, 1 mm in width, with 11-12 seg-
ments.

According to observations by Desbru-
yéres et al. (1985:104) on B. symmytilida,
gut contents included pieces of mussel gills,
coccoid and filamentous bacteria-like cells,
and a mixture of crustacean molts, uniden-

156

tifiable worm setae, diatom frustules, and
amorphous organic matter consistent with
a mixed diet including living tissue and
pseudofeces of mussels. Special features are
their red blood, segmental arborescent
branchiae, 10 pairs of elytra, and 20-21 seg-
ments.

Distribution. —East Central Pacific along
Galapagos Rift, associated with Galapagos
deep-sea mussels, in 2451-2518 m. Re-
ported for 13°N by Desbruyéres et al. (1985:
114).

Branchipolynoe seepensis Pettibone

Branchipolynoe seepensis Pettibone, 1986:
445, figs. 1-4.
Branchipolynoe sp. Hecker, 1985:470.

Material. —Gulf of Mexico, from dives of
the A/vin in Oct 1986, Florida Escarpment,
26°01'46’"N, 84°54'36”"W, 3270 m: Dive
1753, commensal with mussels, 15 speci-
mens (USNM 10151). Dive 1755, with
mussels, 19 specimens (USNM 101503).
Dive 1756, in washings and from broken
mussels, 25 specimens (USNM 101502).
Dive 1758, with mussels, 2 juveniles
(USNM 101504).

Gulf of Mexico off Louisiana, 27°47'N,
91°30'’W, 1097 m, Fla. Dept. Nat. Res., 5
Mar, EJ 87-127, 1987, Bush Hill Box 1,
among mussels, collected by US Navy sub-
marine NR-1, recovered on surface by
R/V Gyre, cruise 87-G-2, one specimen
(USNM 102844).

Remarks. — The additional 61 specimens
from 4 Alvin dives (USNM 10151-4), as-
sociated with the unnamed Gulf of Mexico
giant mussels (to be described and named
by Ruth Turner and Barry Wilson) include
adults and juveniles; some are larger than
previously reported. Larger adults are 31-
40 mm long, 14-17 mm wide, with 21 seg-
ments; smaller adults, 9-17 mm long, 4-8
wide, with 21 segments; larger juveniles, 5-
9 mm long, 3-4.5 mm wide, with 17-19
segments; smaller juveniles, 3-4.5 mm long,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

2-2.5 mm wide, with 15-16 segments; and
the smallest juvenile, 2 mm long, 1.5 mm
wide, with 14 segments. The single speci-
men from Bush Hill off Louisiana (USNM
102844) is 20 mm long, 8 mm wide, with
21 segments; it differs by having long ven-
tral papillae on segment 12 only, not on
both segments 11 and 12. The smaller ju-
veniles lack both pairs of ventral papillae.

Distribution. —Gulf of Mexico, near hy-
persaline seeps at base of Florida Escarp-
ment, associated with Gulf of Mexico mus-
sels, in 3266-3270 m and off Louisiana, in
1097 m.

Subfamily Lepidonotopodiinae Pettibone,
1983

Genus Lepidonotopodium Pettibone, 1983 —

Lepidonotopodium riftense Pettibone

Lepidonotopodium riftense Pettibone,
1984b:852, figs. 2-4.

Material. —East Central Pacific, from
dives of the A/vin on 3 vent areas along the
Galapagos Rift in 1979: Mussel Bed,
00°47.89'N, 86°09.21'W: Dive 986, 3 Dec,
2494 m, mussel bucket washings, one spec-
imen (USNM 118180). Garden of Eden,
00°47.09’N, 86°07.74'W: Dive 993, 10 Dec,
2518 m, slurp sample from base of vesti-
mentiferans, three specimens (USNM
118181). Rose Garden, 00°48.25'N,
86°13.48'W: Dive 896-4, 21 Feb, 2460 m,
six specimens (USNM 118184). Dive 983,
30 Nov, 2457 m, mussel bucket washings
and crab trap residue, 13 specimens (USNM
118183). Dive 990, 7 Dec, 2451 m, wash-
ings from crab trap, 1 specimen (USNM
118185).

East Pacific Rise off western Mexico,
20°50’N, 109°06’W, Oasis Alvin dives in
1982: Dive 1220, 26 Apr, 2617 m, Riftia
and Calyptogena washings, nine specimens
(USNM 118182). Dive 1229, 14 May, 2615
m, Riftia, Calyptogena and Alvinella wash-
ings, eight specimens (USNM 118186).

Gulf of California, Guaymas Basin, dive

VOLUME 102, NUMBER 1

of the A/vin in Jan 1982, 27°01'N, 111°25'W,
2020 m. Dive 1170, rock sample, two small
specimens (USNM 118708). Dives of the
Alvin in 1985, 27°00'N, 111°24’'W, 2000-
2009 m: Dive 1603, 25 Jul, Angel Rock,
bacterial mat, Riftia washings, three spec-
imens (USNM 118707). Dive 1607, 29 Jul,
Lutz 2, bacterial mat, washings from rocks
worms, one specimen (USNM 118706).
Dive 1609, 1 Aug, Lutz 2, sediment, bottom
of bucket of Bead Exp. no. 1, one minute
specimen (USNM 118705). Dive 1615, 7
Aug, smoker in hydrothermal area, Riftia
washings, 16 specimens (USNM 118709).

Remarks.—Some specimens from the
Guaymas Basin are larger than previously
reported: up to 19 mm long, 9 mm wide,
with 26 segments, compared to 13 mm long,
7 mm wide, with 25 segments.

Distribution. —Gulf of California, East
Pacific Rise at 21°N, and East Central
Pacific in the Galapagos Rift, in 2020-
2633 m.

Lepidonotopodium williamsae Pettibone

Lepidonotopodium williamsae Pettibone,
1984b:857, figs. 5-7.

Material. —East Central Pacific, from
dives of the A/vin on two vent areas along
the Galapagos Rift in 1979: Mussel Bed,
00°47.89'N, 86°21'W: Dive 991, 8 Dec, 2490
m, washings from mussels in mesh-lined
milk crates and slurp samples from mussel
wall, eight specimens (USNM 118118-—90).
Rose Garden, 00°48.25'N, 86°13.48'W:
Dive 990, 7 Dec, 2451 m, residue from re-
trieval box containing vestimentiferans, two
specimens (USNM 118187).

East Pacific Rise off western Mexico,
20°50’N, 109°06’W, Oasis Alvin dive in
1982: Dive 1220, 26 Apr, 2617 m, Riftia
and Calyptogena washings, three specimens
(USNM 118188).

Distribution. —Hydrothermal vents of
Tropical East Pacific, Galapagos Rift and
East Pacific Rise at 21°N, in 2451-2633 m.

£SZ

Subfamily Branchinotogluminae Pettibone,
1985
Genus Branchinotogluma Pettibone, 1985
Branchinotogluma grasslei Pettibone

Branchinotogluma grasslei Pettibone,
1985b:457, figs. 5, 6; 1988: 215.

Material. —East Central Pacific, Galapa-
gos Rift, Alvin dives in 1979: Mussel Bed,
00°47.89'N, 86°09.21'W: Dive 991, 8 Dec,
2490 m, two slurp samples from mussel wall,
three specimens (USNM 118151). Rose
Garden, 00°48.25’N, 86°13.48’W: Dive 983,
30 Nov, 2457 m, mussel bucket washings,
three specimens (USNM 118152).

East Pacific Rise off western Mexico, Oa-
sis Alvin dive in 1982, 20°50'N, 109°06’W:
Dive 1220, 26 Apr, 2617 m, Riftia and Ca-
lyptogena washings, one specimen (USNM
118150).

Gulf of California, A/vin dive in Guaymas
Basin in Jan 1982, 27°01'N, 111°25'W: Dive
1172, 2010 m, Box core no. 1, on slope of
mound in area of sulfides, sediment covered
with white material, one specimen (USNM
118699).

Distribution. —Hydrothermal vents of
Tropical East Pacific, Galapagos Rift and
East Pacific Rise at 21°N; Gulf of California;
Northeast Pacific, Explorer and Juan de Fuca
Ridges, in 1495-2633 m.

Branchinotogluma sandersi Pettibone

Branchinotogluma sandersi Pettibone,
1985b:453, figs. 3, 4; 1988: 217.

Material. —East Central Pacific, Galapa-
gos Rift, Alvin dives in 1979: Mussel Bed,
00°47.89'N, 86°09.21'W: Dive 991, 8 Dec,
2490 m, two slurp samples from mussel wall,
two specimens (USNM 118157). Rose Gar-
den, 00°48.25'N, 86°13.48’W: Dive 983, 30
Nov, 2457 m, mussel bucket washings, one
specimen (USNM 118156). Dive 988, 5 Dec,
2450 m, slurp sample from base of vesti-
mentiferans, one specimen (USNM
118158).

158

East Pacific Rise off western Mexico, Oa-
sis Alvin dive in 1982, 20°50'N, 109°06'W:
Dive 1220, 26 Apr, 2617 m, Riftia and Ca-
lyptogena washings, three specimens
(USNM 118159).

Gulf of California, A/vin dive in Guaymas
Basin in 1985, 27°00'N, 111°24’W: Dive
1599, 13 Jul, 2009 m, Lutz 2, bacterial mat,
washing from Rock Chimney, one specimen
(USNM 118700).

Distribution. —Hydrothermal vents of
Tropical East Pacific, Galapagos Rift and
East Pacific Rise at 21°N; Gulf of California;
Northeast Pacific, Explorer and Juan de Fuca
Ridges, in 1592-2633 m.

Branchinotogluma hessleri Pettibone

Branchinotogluma _ hessleri Pettibone,

1985b:450, figs. 1, 2.

Material. —East Central Pacific, Galapa-
gos Rift, dives of A/vin in 1979: Mussel Bed,
00°47.89'N, 86°09.21'W: Dive 991, 8 Dec,
2490 m, two slurp samples from mussel wall,
three specimens (USNM 118164). Garden
of Eden, 00°47.69°N, 86°07.74'W: Dive 993,
10 Dec, 2518 m, slurp sample from base of
vestimentiferans, four specimens (USNM
118162). Rose Garden, 00°48.25’N,
86°13.48'W: Dive 983, 30 Nov, 2457 m,
mussel bucket washings, three specimens
(USNM 118161). Dive 988, 5 Dec, 2450
m, slurp sample from base of vestimenti-
ferans, one specimen (USNM 118163).

East Pacific Rise off western Mexico, Oa-
sis A/vin dives in 1982, 20°50'N, 109°06'W:
Dive 1220, 26 Apr, 2617 m, Riftia and Ca-
lyptogena washings, 13 specimens (USNM
118160). Dive 1229, 14 May, 2615 m, Rif-
tia, Calyptogena and Alvinella washings, 2
specimens (USNM 118165).

Distribution. —Hydrothermal vents of
Tropical East Pacific, Galapagos Rift and
East Pacific Rise at 21°N, in 2450-2633 m.

Opisthotrochopodus Pettibone, 1985
Opisthotrochopodus alvinus Pettibone

Opisthotrochopodus alvinus Pettibone,
1985b:459, figs. 7-9.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Material. —East Central Pacific, Galapa-
gos Rift, dives of A/vin in 1979: Mussel Bed,
00°47.89'N, 86°09.21'’W: Dive 991, 8 Dec,
2490 m, two slurp samples from mussel wall,
one specimen (USNM 118179). Garden of
Eden, 00°47.69'N, 86°07.74'W: Dive 993,
10 Dec, 2518 m, slurp sample from base of
vestimentiferans, 2 specimens (USNM
118177). Rose Garden, O0°48225 Ne
86°13.48’W: Dive 983, 30 Nov, 2457 m,
mussel bucket washings, 11 specimens
(USNM 118174), crab trap residue, 2 spec-
imens (USNM 118176).

East Pacific Rise off western Mexico, Oa-
sis Alvin dive in 1982, 20°50'N, 109°06’W:
Dive 1220, 26 Apr, 2617 m, Riftia and Ca-
lyptogena washings two specimens (USNM
118178).

Distribution. —Hydrothermal vents of
Tropical East Pacific, Galapagos Rift and
East Pacific Rise at 21°N, in 2451-2633 m.

Branchinotogluma sp. A
[Young of B. hessleri or O. alvinus]

Branchinotogluma sp. A Pettibone, 1985b:
466.

Material. —East Central Pacific, from
dives of A/vin along Galapagos Rift in 1979:
Mussel Bed: Dive 991, 8 Dec, 2490 m, slurp
sample from mussel wall, nine young spec-
imens (USNM 118154). Garden of Eden:
Dive 993, 10 Dec, 2518 m, slurp sample
from base of vestimentiferans, two young
specimens (USNM 118155). Rose Garden:
Dive 983, 30 Nov, 2457 m, mussel bucket
washings, 10 young specimens (USNM
118153).

Remarks. —Adults of the above two
species were collected on the same dives.

Branchinotogluma sp. B
[Young of B. grasslei or B. sandersi|

Branchinotogluma sp. B Pettibone, 1985b:
466.

Material. —East Central Pacific, from
dives of Alvin along Galapagos Rift in 1979:
Rose Garden: Dive 983, 30 Nov, 2457 m,

VOLUME 102, NUMBER 1

mussel bucket washings, 11 young speci-
mens (USNM 118149).

Gulf of California, A/vin dive in Guaymas
Basin in 1985, 27°00’N, 111°24’W: Dive
1603, 25 Jul, 2004 m, Angel Rock, bacterial
mat, Riftia washings, one small specimen
(USNM 118701).

Remarks. —Adults of the above two
species were collected on the same dives.

Subfamily Branchiplicatinae Pettibone,
1985
Genus Branchiplicatus Pettibone, 1985
Branchiplicatus cupreus Pettibone

Branchiplicatus cupreus Pettibone, 1985a:
151, figs. 1-4.

Material. —Gulf of California, Alvin dive
in Guaymas Basin in 1982, 27°01'N,
111°25’W: Dive 1169-001, 11-20 Jan, 2020
m, rock sample, one specimen (USNM
118702). Alvin dive in 1985, 27°00’N,
111°24’W: Dive 1615, 7 Aug, 2000 m,
smoker 1, hydrothermal area, Riftia wash-
ings, one specimen (USNM 118703).

Distribution. —Hydrothermal vents in
Gulf of California and East Pacific Rise at
21°N, in 2000-2633 m.

Subfamily Macellicephalinae Hartmann-
Schroder, 1971,
emended Pettibone, 1976
Genus Levensteiniella Pettibone, 1985
Levensteiniella kincaidi Pettibone

Levensteiniella kincaidi Pettibone, 1985c:
471, figs. 1-3; 1988:215.

Material. —East Central Pacific, Galapa-
gos Rift, dives of A/vin in 1979: Mussel Bed,
00°47.89'N, 86°09.21'W: Dive 986, 3 Dec,
2494 m, mussel bucket washings, three
specimens (USNM 118196). Dive 991, 8
Dec, 2490 m, slurp samples from mussel
wall, four young specimens (USNM
118194). Garden of Eden, 00°47.69'W,
86°07.74'W: Dive 993, 10 Dec, 2518 m,
slurp sample from base of vestimentiferans,
29 young specimens (USNM 118191). Rose
Garden, 00°48.25'W, 86°13.48'W: Dive 983,

159

30 Nov, mussel bucket washings, 20 young
specimens (USNM 118197). Dive 988, 5
Dec, 2450 m, slurp sample from base of
vestimentiferans, one specimen (USNM
118192).

East Pacific Rise off western Mexico,
20°50’N, 109°06’W, Oasis Alvin dives in
1982: Dive 1220, 26 Apr, 2617 m, Riftia
and Calyptogena washings, 10 specimens (4
young, USNM 118195). Dive 1229, 14 May,
2616 m, Riftia, Calyptogena and Alvinella
washings, small specimen (USNM 118193).

Distribution. —Hydrothermal vents of
Tropical East Pacific: Galapagos Rift and
East Pacific Rise at 21°N; Northeast Pacific,
Explorer and Juan de Fuca Ridges, in 1818-
2633 m.

Genus Bathykurila Pettibone, 1976
Bathykurila guaymasensis, new species
Bigs. h.2

Material. —Gulf of California, Alvin dive
in Guaymas Basin in Jan 1982, 27°01'N,
111°25’°W: Dive 1176, 2020 m, Box core 2,
taken over vent, with numerous nuculanid
bivalves, 2d, holotype (USNM 118694) and
paratype (USNM 118695); sec. B, paratype
(USNM 118696). Alvin dive in 8 Aug 1985,
27°00’N, 111°24’W: Dive 1614, 2004 m,
Angel Rock, bacterial mat, Box core 5, sub-
core B, 0-5 cm, paratype (USNSM 118697),
subcore, C, 0-3 cm, paratype (USNM
118698).

Description. —Length of holotype 8 mm,
width with setae 5 mm, segments 15; length
of two larger paratypes 7 mm, width 5 mm,
segments 14; length of two smaller para-
types 3-4 mm, width 3-3.5 mm, segments
15. Body fusiform, flattened, with parapo-
dia and setae very long, projecting, longer
than body width (Fig. 1A, B).

Elytra (mostly missing) and large, bul-
bous elytrophores (Figs. 1A, B, 2A) seven
pairs, on segments 2, 4, 5, 7, 9, 11, and 13.
Elytra large, oval, covering dorsum, deli-
cate, with larger conical to rounded tuber-
cles variable in size on posterior and lateral
borders, with smaller ones scattered on sur-

160

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig-T.

Bathykurila guaymasensis, A, B, holotype, USNM 118694; C, paratype, USNM 118698; D, paratype,

USNM 118695: A, Dorsal view of anterior end, acicula in tentaculophores of lst segment dotted; B, Dorsal
view of posterior end (segments 12-15); C, Ventral view of posterior end (segments 10-14), anal cirri and most
of ventral cirri broken off; D, Border papillae on dorsal side of pharynx (dissected). Scales = 0.5 mm for A, B;

1.0 mm for C; 0.1 mm for D.

face (Fig. 2F): larger tubercles not usual type,
not chitinous, not sharply set off from sur-
face. Dorsal tubercles on cirrigerous seg-
ments large, bulbous to nodular (Figs. 1A,
B, 2B).

Prostomium oval, deeply bilobed; trian-
gular anterior continuations of prostomium
with small frontal filaments; median anten-
na with small ceratophore in anterior notch,
with short subulate style; without lateral an-
tennae and eyes; palps stout, tapered, about
two times longer than prostomium (Fig. 1A).
First or tentacular segment fused to prosto-
mium, not visible dorsally; tentaculophores
lateral to prostomium, each with prominent
acicular lobe on inner side, without setae,
and pair of subulate tentacular cirri, shorter
than palps; upper lip large, bilobed (Fig. 1A).
Second or buccal segment with first pair of
elytrophores, biramous parapodia and ven-
tral or buccal cirri lateral to ventral mouth,
longer than following ventral cirri (Fig. 1A).

Parapodia biramous, with notopodia
shorter and smaller than neuroodia (Fig. 2A,
B.) Notopodium rounded basally, with pro-
jecting acicular lobe on lower side; neuro-
podium with conical projecting presetal
acicular lobe and shorter, rounded postsetal
lobe. Notosetae numerous, short to longer,
forming radiating bundles, stouter than
neurosetae, with row of widely spaced teeth
(4-11) along one side and blunt tips (Fig.
2C). Neurosetae very numerous, forming
fan-shaped bundles; supraacicular neuro-
setae with longer spinous regions and two
rows of long spines, tapering to bluntly
pointed bare tips (Fig. 2D); subacicular neu-
rosetae with shorter spinous regions, close-
set short spines, and tapered bare tips (Fig.
2E). Dorsal cirri with cylindrical cirro-
phores on posterior sides of notopodia,
shorter than notopodia, with styles wider
basally and long slender tips extending to
tips of setae or beyond (Figs. 1A, B, 2B).

VOLUME 102, NUMBER 1

161

¢
c
c
¢
is
€ ©
¢ ie
ee ee ee
SSS)

Mm

C D

Fig. 2. Bathykurila guaymasensis, holotype, USNM 118694: A, Right elytragerous parapodium, anterior
view, acicula dotted; B, Right cirrigerous parapodium, posterior view; C, Notosetae; D, Supraacicular neuroseta;
E, Subacicular neuroseta; F, Right elytron. Scales = 0.5 mm for A, B; 0.1 mm for C-E; 0.5 mm for F.

Ventral cirri attached on middle of neuro-
podia, short, tapered (Fig. 2A, B).

Pair of long ventral papillae usually pres-
ent on segment 11 (present on four of five
specimens), sometimes extending far be-
yond posterior end, with yellowish secretion
inside (Fig. 1C). Posterior 2 segments (14,
15) smaller, enclosed in elytragerous seg-
ment 13 (Fig. 1B); segment 14 with dorsal
cirri; segment 15 smaller, without dorsal cirri
(Fig. 1B), or segment absent (Fig. 1C); py-
gidium long, slender, wedged between par-
apodia of segment 15, with pair of slender,
short anal cirri (Fig. 1B).

Pharynx (dissected) with opening encir-
cled by eight pairs of border papillae (Fig.
1D), and two pairs of heavy, hooked jaws.

Etymology.—The species is named for the
collecting site, the Guaymas Basin.

Remarks.—Bathykurila guaymasensis is
close to B. zenkevitchi (Uschakov 1955, fig.
la-—c; Pettibone 1976, fig. 24a—d), the type
species of Bathykurila, from the Kurile-
Kamchatka Trench in the northwest Pacific,
in 8100 m. They differ as indicated in Ta-
ble 1.

Subfamily Macellicephaloidinae
Pettibone, 1976
Genus Macellicephaloides Uschakov, 1955

Type species.—Macellicephaloides gran-
dicirrus Uschakov, 1955 (as grandicirra).
Gender: masculine (according to the Code,
Art. 30.a.i1, names ending in -oides should
be masculine, not feminine, as indicated by
Pettibone 1976:42).

Table 1.— Differences between Bathykurila zenkevit-
chi (Uschakov) and Bathykurila guaymasensis, n. sp.

Character B. zenkevitchi B. guaymasensis
Tentacular about as long as _ shorter than palps
cirri palps
Notopodia as long as or shorter than neu-
longer than ropodia
neuropodia
Cirrophore longer than no- shorter than no-
of dorsal topodia topodia
cirri
Ventral pa- 4 pairs on seg- | pair on segment
pillae ments 10-13 11
Length 15-21 mm 3-8 mm
Width 12 mm 3-5 mm
Segments 1S 14-15

162

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Macellicephaloides alvini, holotype, USNM 118704: A, Dorsal view of anterior end, styles of median
antenna, tentacular cirri, and dorsal cirri of segment 3 missing, right parapodia of segments 2 and 3 damaged
or not developed; B, Ventral view of anterior end, ventral cirri missing; C, Dorsal view of posterior end (segments
13-17), elytra and styles of dorsal cirri missing. Scale = 1.0 mm for A-C.

Macellicephaloides alvini, new species
Figs. 3, 4

Material. —Gulf of California, Alvin dive
in Guaymas Basin, F. Grassle, collector,
Dive 1614, 6 Aug 1985, 27°00’N, 111°24'W:
2004 m, Angel Rock, bacterial mat, Box
core 5, subcore D, holotype (USNM
118704).

Description. — Length of holotype 7.5 mm,
width with setae 4 mm, segments 17. Body
flattened dorsoventrally, tapering slightly
anteriorly and more so posteriorly, with
parapodia of 5 posterior segments directed
posteriorly (Fig. 3C). Integument smooth,
not papillate. Elytra (all missing) and large,
bulbous elytrophores eight pairs, on seg-
ments 2; 4,5, 7,9, 115 13sand 15 (Figs.3A,
C, 4A). Dorsal tubercles on cirrigerous seg-
ments indistinct.

Prostomium (poor condition) withdrawn
and fused to tentacular segment, oval,
slightly bilobed; ceratophore of median an-
tenna in middle of prostomium, style miss-
ing; palps on short palpophores, rather short,
tapering; without lateral antennae and eyes
(Fig. 3A, B). Tentacular segment distinct
dorsally, ventrally forming upper, lateral,
and posterior lips of ventral mouth; tenta-

culophores of dorsal and ventral cirri short,
lateral to prostomium, styles missing (Fig.
3A, B). Segment 2 with first pair of elytro-
phores and elongate parapodia; ventral cirri
missing; deep depression in middle of ven-
tral surface, covered by large, oval flap at-
tached to segment 3 (Fig. 3A, B).

Parapodia subbiramous; notopodium
bulbous basally, with projecting, stout, am-
ber-colored notoaciculum, without noto-
setae (Fig. 4A—C); neuropodium elongated,
with projecting acicular lobe; neurosetae
numerous, delicate, slender, slightly wider
basally, tapering to slender tips, with two
rows of delicate spines (Fig. 4A, B, D). Pos-
terior five segments with notoacicula stou-
ter, projecting beyond neuropodia (Fig. 3C).
Dorsal cirri with rather long, thick, cylin-
drical cirrophores on posterior dorsal sides
of notopodia; styles missing (Fig. 4B). Ven-
tral cirri missing.

Pygidium rounded, enclosed in parapo-
dia of last segment (Fig. 3C). Pharynx (dis-
sected) large, muscular; anterior end encir-
cled with papillae, number and arrangement
not evident but with longer lateral pair (not
extra long as in M. vitiazi); two pairs of long
V-shaped hooked jaws; dorsal pair fused
medially, with additional median piece (Fig.

VOLUME 102, NUMBER 1

163

Fig. 4. Macellicephaloides alvini, holotype, USNM 118704: A, Right elytragerous parapodium from segment
7, anterior view, acicula dotted, elytron and ventral cirri missing; B, Right cirrigerous parapodium of segment
8, posterior view, styles of dorsal and ventral cirri missing; C, Notoaciculum from same; D, Neuroseta from
same, with detail of parts; E, Dorsal jaws, outer view; F, Same, inner view; G, Ventral jaws, outer view; H,
Same, inner view. Scales = 0.5 mm for A, B; 0.1 mm for C—H.

4E, F); ventral pair denticulate on inner side
(Fig. 4G, H).

Etymology.—The species is named for the
collecting submersible, DSRV Alvin.

Remarks on the genus Macellicephaloides
Uschakov, 1955, and the subfamily Macel-
licephaloidinae Pettibone, 1976: Of partic-
ular interest is the find of a single specimen
from the Guaymas Basin that seems to fit
into the genus Macellicephaloides, estab-
lished by Uschakov (1955) for three
ultra-abyssal species, M. grandicirrus, M.
verrucosus, and M. vitiazi, from the Kurile-
Kamchatka Trench in 7210-9950 m. Us-
chakov placed the genus under Aphroditi-

dae: Polynoinae. An additional species, M.
uschakovi, was added to the genus from the
same area in 8120 m by Levenstein (1971).

In her catalogue, Hartman (1959:93)
placed Macellicephaloides under Polynoi-
dae: Lepidonotinae, and selected M. gran-
dicirrus as the type species. Hartmann-
Schroder (1974), in her revision of the
subfamily Macellicephalinae Hartmann-
Schroder, 1971, included Macellicepha-
loides in the subfamily. As a part of my
revision of the genus Macellicephala and the
subfamily Macellicephalinae Hartmann-
Schroder, I established the new subfamily
Macellicephaloidinae for Macellicepha-

164

loides and the above four species, with sup-
plementary descriptions, based on speci-
mens sent in exchange by P. Uschakov and
R. Levenstein.

Levenstein (1975) added an additional
species, M. sandvichensis, from the South
Sandwich Trench in 7200-7934 m. Another
species, M. villosus, was added by Leven-
stein (1982) from the Japan Trench in 7370-
7380 m. Levenstein (1983) added a seventh
species, M. improvisus, from the Kurile-
Kamchatka Trench in 8035-8120 m and
included a Key to the seven species of Ma-
cellicephaloides. It probably should be not-
ed that Uschakov (1982) did not recognize
the subfamily Macellicephaloidinae and
other subfamilies proposed by Pettibone but
included the genus under Macellicephali-
nae.

In her review of the genera and species of
the deep-sea polynoid subfamilies in the
world oceans, Levenstein (1984:83, fig. 4)
indicated that the distributional pattern of
the Macellicephaloidinae showed a western
Pacific origin, since six of the seven species
of the genus Macellicephaloides were known
to inhabit trenches of the western Pacific
and only one species was known to inhabit
the Atlantic sector of the Antarctic Ocean,
the South Sandwich Trench. She suggested
that representatives of this genus (or its
ancestor) were probably widely distributed
in the abyssal region of the Pacific Ocean
and gave rise to species adapted to life in
the oceanic trenches, producing the ultra-
abyssal fauna of this subfamily. The pres-
ence of M. sandvichensis in the South Sand-
wich Trench could be explained on the basis
of a close connection between the deep-sea
faunas of the Pacific and Atlantic sectors of
the Antarctic. The new species, M. alvini
from 4000-5000 m shallower waters in the
Gulf of California, might be considered to
be close to the ancestor of the genus. It is
considerably smaller than representatives
from the other areas.

All eight species have a relatively small
number of segments (16-21, 1st achaetous)
and eight pairs of elytrophores (elytra all

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

missing) on segments 2, 4, 5, 7, 9, 11, 13,
and 15. The prostomium is relatively small,
bilobed, with the ceratophore of the median
antenna in the middle of the prostomium
and with palps but without lateral antennae
or eyes. The first or tentacular segment en-
closes the prostomium and is distinct dor-
sally, with two pairs of tentacular cirri on
tentaculophores, but without acicular lobes
or setae; ventrally the tentacular segment
forms the upper, lateral, and posterior lips
of the ventral mouth. The second segment
has the first pair of elytrophores, large par-
apodia, and ventral cirri, with a deep mid-
ventral depression covered by a large oval
flap attached to segment 3 (as in M. alvini,
Fig. 3B) or a rectangular flap (as in M. gran-
dicirrus, Pettibone, 1976, fig. 25b). Para-
podia are elongate, subbiramous, the no-
topodium represented by a conical lobe with
a more or less projecting, stout notoacicu-
lum, without notosetae; the neuropodium
is conical, with numerous, slender neuro-
setae. Dorsal cirri on the non-elytragerous
segments have moderately to very long, cy-
lindrical cirrophores and short styles; dorsal
tubercles are indistinct or rarely nodular, as
in M. verrucosus. Ventral cirri are short. The
pygidium is enclosed in the long parapodia
of a variable number of posterior segments,
and without anal cirri. The integument is
smooth, rarely papillate dorsally, as in M.
villosus, or with middorsal nodular tuber-
cles, asin M. verrucosus. The large muscular
pharynx has two pairs of hooked jaws and
distal papillae, the lateral pair of papillae
longer than the others. Lateral papillae
sometimes very long, as in M. vitiazi (Pet-
tibone, 1976, fig. 26c) and M. sandvichensis
(Levenstein, 1975, fig. 3a), and similar to
those of the pelagic polychaete family AI-
ciopidae.

Key to the Species of

Macellicephaloides
b. Segments '6tor 7.22 eae 2
=) ‘Segments tS: 071 20-21), 2a eee 6
2. Segments. 16) b. &. HO) See 3
=, Segments l7is 24. Bi ee ee 4

VOLUME 102, NUMBER 1 165

SS
Cle oS
“et WSss =
=

D

Fig. 5. Neoleanira racemosa, USNM 118714: A, Dorsal view of anterior end, ventral tentacular cirri not
visible; left dorsal cirrus on segment 3 broken off; B, Right tentacular parapodium, outer view, aciculum dotted;
C, Right elytragerous parapodium from segment 2, anterior view, acicula dotted; D, Right cirrigerous parapodium
from segment 3, posterior view; E, Right elytragerous parapodium from segment 4, anterior view, acicula dotted.
Scales = 1.0 mm for A; 0.5 mm for B—E. au, auricle; br, branchia; ct, ctenidium; dc, dorsal cirrus; dtc, dorsal
tentacular cirrus; itl, inner tentacular lobe; lan, lateral antenna; man, median antenna; pa, palp; st, stylode; vtc,
ventral tentacular cirrus.

3. With paired, nodular dorsal tuber- ments. Pharynx with lateral papillae
cles on cirrigerous segments 6-14 MISE ERA IOs ee cen age NE ee 3
and middorsal tubercles on seg- 5. Cirrophores of dorsal cirri extend-
ments 3-15 (Uschakov 1955, fig. 3; ing far beyond tips of neuropodia
1982, pl. 12; Pettibone 1976, fig. 27 (Uschakov 1955, fig. 2; 1982, pl. 11:

oe eee M. verrucosus Uschakov Petibone, 197.6. fie.25)=...: -h4 enn

—wymnout Gorsal tubercles On. Gifi- .... | 6 nk beet eus- M. grandicirrus Uschakov
gerous segments and middorsal tu- — Cirrophores of dorsal cirri not as
bercles (Uschakov 1955S, fig. 4; 1982, long, extending to tips of neuropo-
pl 13; Pettibone 1976, fig. 26)... dia or shorter (Figs. 3, 4) ........

a ee M. vitiazi Uschakov ieee tegen lla VINE HCW SPeGiEs

4. Notoacicula not extra stout or long. 6. Segments 18. Cirrophores of dorsal
Cirrophores of dorsal cirri extend- cirri very short. Notopodia with
ing to about tips of neuropodia. flexible, thin acicula (Levenstein
Pharynx with pair of very long lat- POSS ie Ae) ee Sy ae Bee
peal papillae (Pevensicein lOVS AGS... 6) on de Suan woe ce M. improvisus Levenstein
SW Rha ve: 2. M. sandvichensis Levenstein — Segments 20-21. Notopodia other-

— Notoacicula stout, extending far be- WISE darian Paces ss ari Sa, Se 7

yond neuropodia in posterior seg- 7. Dorsal side of body with conical pa-

166

LL

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Zz
SS
—— ———-

7 <a = |||

Fig. 6. Neoleanira racemosa, USNM 118714: A, Right parapodium from posterior region (about segment
35), anterior view, acicula dotted; B, Same, posterior view; C, Upper, middle, and lower compound neurosetae,
with detail of distal part of stems. Scales = 0.5 mm for A, B; 0.1 for C. br, branchia; ct, ctenidium; st, stylode.

pillae (Levenstein 1982, fig. 2)
pit iiehiisicel Late M. villosus Levenstein
— Integument smooth (Levenstein
1971, fig. 6) ..M. uschakovi Levenstein

Family Sigalionidae
Genus Neoleanira Pettibone, 1970
Neoleanira racemosa (Fauchald)
Figs. 5, 6

Sthenolepis racemosa Fauchald, 1972:34,
pl. 2, figs. a-d.

Neoleanira racemosa.—Fauchald & Han-
cock, 1981:23.

Remarks. — The species was originally de-
scribed by Fauchald (1972) under Stheno-
lepis, based on specimens collected in the
Guaymas Basin in 1774—2259 m and later
referred to Neoleanira by Fauchald and
Hancock (1981), with additional specimens
reported from off central Oregon in 2000-
2900 m. The additions to the description of
the species are based on additional speci-
mens from the Guaymas Basin. All speci-
mens are incomplete posteriorly and elytra
are missing in both the previous and the
present collections.

Material. —Gulf of California, Alvin dives

in Guaymas Basin, Jan 1972, 27°01'N,
111°25’W, 2020 m: Dive 1170, 2 m from
vestimentiferans, Box core 1, small speci-
men (USNM 118713). Dive 1173, on slope
of mound, Box core 1, one specimen (USNM
118714), Box core 2b, subcore D, two spec-
imens (USNM 118715). Dive 1174, about
15 m from mound, Box core la, one spec-
imen (USNM 118712), Box core lc, one
small specimen (USNM 118710). Dive
1177, about 3 m from mound with tube
worms, Box core 2, one specimen (USNM
118711).

Supplementary description. —Incomplete
specimen of 44 segments, 16+ mm long, 3
mm wide with setae (USNM 118714). Ely-
tra all missing; elytrophores on segments 2,
4, 5, 7, continuing on alternate segments to
23, then on all segments. Prostomium
rounded, ceratophore of median antenna on
anterior margin, with pair of lateral auricles
and long tapering style; parapodia of first or
tentacular segment partially fused to lateral
and anterior sides of prostomium, with long
lateral antennae attached on dorsal, inner
side; long dorsal and short ventral tenta-
cular cirri with short inner tentacular lobe;
long tapering palps inserted ventral to ten-

VOLUME 102, NUMBER 1

tacular parapodia; with aciculum and 2 ra-
diating bundles of capillary setae (Fig. 5A,
B).

Parapodia of segments 2 and 3 directed
anteriorly (Fig. 5A). Segment 2 with first
pair of elytrophores, biramous parapodia,
and ventral buccal cirri longer than follow-
ing ventral cirri (Fig. 5A, C). Parapodium
of segment 3 with long dorsal cirrus on short
cirrophore and small oval ventral ctenid-
ium medial to ventral cirrus (Fig. 5A, D).
Segment 4 with 3 oval ctenidia between dor-
sal base of notopodium and elytrophore,
with very small lateral branchia (Fig. 5E).
Anterior neuropodia with numerous fim-
briae or stylodes on tips of acicular lobes
and bilobed posterior bracts (Fig. 5C—E).

More posterior parapodia with well-de-
veloped branchiae attached to lateral sides
of elytrophores or dorsal tubercles and 3
ciliated ctenidia in curved part; notopo-
dium with stylodes and long capillary no-
tosetae, finely to coarsely spinous; neuro-
podium conical, with stylodes, fan-shaped
bundle of slender neurosetae, ventral cirrus,
and small bulbous ventral ctenidium (Fig.
6A, B). Neurosetae compound spinigers,
upper and lower ones with shorter blades,
middle ones with longer blades, none can-
aliculate; long slender shafts distally dentate
(Fig. 6C).

Distribution. —Gulf of California and off
Central Oregon. In 1774-2900 m.

Key to the Species of Neoleanira

. Blades of compound spinigerous
neurosetae not canaliculate. Ventral
ctenidia small, bulbous. Branchiae
without prominent spurlike pro-
cesses. Elytra missing (Figs. 5, 6;
Fauchald 1972, figs. a-d) ........

| Le N. racemosa (Fauchald)

— Blades of compound spinigerous

neurosetae canaliculate

2. Elytra with borders entire, without

lateral fringes of papillae. [Ventral

ctenidia small, clavate. Branchiae
without prominent basal spurlike

processes] (Pettibone 1970, fig. 7)

a

167

SURE AL N. magellanica (McIntosh)
— Elytra with delicate lateral fringes of
Pal ae ee ey ee et ev eens! £h 3
3. Ventral ctenidia small, clavate.
Branchiae without prominent basal
spurlike processes (Pettibone 1970,
figs. 1-4) ...... N. tetragona (Oersted)
— Ventral ctenidia large, boot-shaped.
Branchiae with prominent basal
spurlike processes (Pettibone 1970,
MESES. 6)! 2 SF N. areolata (McIntosh)

Acknowledgments

My thanks go to J. Fred Grassle, Susan
Brown-Leger, and Rosemarie Petrecca of the
Woods Hole Oceanographic Institution for
the scaled polychaetes from the Guaymas
Basin and for the additional polynoids com-
mensal with the Galapagos mussels, sepa-
rated by Vida Kenk. I also thank Barbara
Hecker for the additional polynoids com-
mensal with the Gulf of Mexico mussels,
separated by Ruth Turner. Additional col-
lections from the Galapagos and the East
Pacific Rise at 21°N, received from WHOI
through the Smithsonian Sorting Center, are
appreciated. The manuscript benefited from
the careful reviews of Thomas H. Perkins
and Nancy J. Maciolek.

Literature Cited

Desbruyéres, D., F. Gaill, L. Laubier, & Y. Fouquet.
1985. Polychaetous annelids from hydrother-
mal vent ecosystems: An ecological overview. —
Bulletin of the Biological Society of Washington
6:103-116.

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:
1-575.

—, & D. R. Hancock. 1981. Deep-water poly-
chaetes from a transect off Central Oregon. —
Allan Hancock Foundation Monograph 11:
1-73.

Grassle, J. F. 1986. The ecology of deep-sea hydro-
thermal vent communities.— Advances in Ma-
rine Biology 23:301-362.

Hartman, O. 1959. Catalogue of the polychaetous
annelids of the world, Part I.—Allan Hancock
Foundation Publications Occasional Paper 23:
1-353.

168

Hartmann-Schréder, G. 1974. Die Unterfamilie Ma-
cellicephalinae Hartmann-Schroéder, 1971 (Po-
lynoidae, Polychaeta). Mit Beschreibung einem
neuen Art, Macellicephala jameensis n. sp., aus
einem Hohlengewasser von Lanzarote (Kanar-
ische Inseln).— Mitteilungen aus dem Hambur-
gischen Zoologischen Museum und Institut 71:
75-85.

Hecker, B. 1985. Fauna from a cold sulfur-seep in
the Gulf of Mexico: Comparison with hydro-
thermal vent communities and evolutionary
implications. — Bulletin of the Biological Society
of Washington 6:465-473.

Kenk, V. C., & B. R. Wilson. 1985. A new mussel
(Bivalvia, Mytilidae) from hydrothermal vents
in the Galapagos rift zone.— Malacologia 26(1-
2):253-271.

Levenstein, R. J. 1971. [Polychaete worms of the

genera Macellicephala and Macellicephaloides

(Family Aphroditidae) from the Pacific Ocean.

In Fauna of the Kurile-Kamchatka Trench.]—

Trudy Institut Okeanologii P.P. Shirshov Aca-

demiia Nauk SSSR 92:18-—35 [in Russian, En-

glish summary].

. 1975. [Polychaetous annelids of the deep-sea

trenches of the Atlantic Sector of the Antarctic

Ocean.]— Trudy Institut Okeanologii P.P. Shir-

shov Academiia Nauk SSSR 103:119-142 [in

Russian, English summary].

. 1982. [On the polychaete fauna of the family

Polynoidae from the trench of Japan.]— Trudy

Institut Okeanologii P.P. Shirshov Academia

Nauk SSSR 117:59-62 [in Russian, English

summary].

1983. [Macellicephaloides improvisa sp. n.
(Polychaeta, Polynoidae) from Kurile-Kam-
chatka Trench.]— Zoologichesky Zhurnal 62(9):
1419-1421 [in Russian, English summary].

. 1984. On the ways of formation of the deep-

sea polychaete fauna of the family Polynoidae.

Pp. 72-85 in P. A. Hutchings, ed., Proceedings

of the First International Polychaete Confer-

ence, Sydney, Australia, 1983. The Linnean So-
ciety of New South Wales.

Lonsdale, P. 1984. Hot vents and hydrocarbon seeps
in the Sea of Cortez.— Oceanus 27(3):21-24.

Pettibone, M. H. 1970. Two new genera of Sigalio-

nidae (Polychaeta). — Proceedings of the Biolog-

ical Society of Washington 83(34):365-386.

1976. Revision of the genus Macellicephala
McIntosh and the subfamily Macellicephalinae
Hartmann-Schroéder (Polychaeta: Polynoi-
dae).—Smithsonian Contributions to Zoology
229:1-71.

1983. A new scale worm (Polychaeta: Poly-
noidae) from the hydrothermal rift-area off
Western Mexico at 21°N.—Proceedings of the
Biological Society of Washington 96(3):392-399.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1984a. 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(1):226-239.

1984b. Two new species of Lepidonotopo-
dium (Polychaeta: Polynoidae: Lepidonotopo-
dinae) from hydrothermal vents of the Gala-
pagos and the East Pacific Rise at 21°N.—
Proceedings of the Biological Society of Wash-
ington 97(4):847-863.

1985a. An additional new scale worm (Po-
lychaeta: Polynoidae) from the hydrothermal rift-
area off Western Mexico at 21°N.— Proceedings
of the Biological Society of Washington 98(1):
150-157.

1985b. Additional branchiate scale-worms
(Polychaeta: Polynoidae) from Galapagos hy-
drothermal vent and rift-area off Western Mex-
ico at 21°N.—Proceedings of the Biological So-
ciety of Washington 98(2):447—-469.

1985c. New genera and species of deep-sea
Macellicephalinae and Harmothoinae (Poly-
chaeta: Polynoidae) from the hydrothermal rift
areas of the Galapagos and Western Mexico at
21°N and from the Santa Catalina Channel.—
Proceedings of the Biological Society of Wash-
ington 98(3):740-757.

1986. A new scale-worm commensal with
deep-sea mussels in the seep-sites at the Florida
Escarpment in the eastern Gulf of Mexico (Po-
lychaeta: Polynoidae: Branchipolynoinae). —
Proceedings of the Biological Society of Wash-
ington 99(3):444-451.

. 1988. New species and new records of scaled
polychaetes (Polychaeta: Polynoidae) from hy-
drothermal vents of the Northeast Pacific Ex-
plorer and Juan de Fuca ridges.— Proceedings
of the Biological Society of Washington 101(1):
192-208.

Uschakov, P. V. 1955. [Polychaete worms of the fam-
ily Aphroditidae from the Kurile-Kamchatka
Trench.]—Trudy Institut Okeanologii Akade-
miia Nauk SSSR 12:311-321 [in Russian:
translated in 1969 by the Bureau of Commercial
Fisheries, 10 pages].

1982. [Polychaetes of the Suborder Aphro-
ditiformia of the Arctic Ocean and the north-
western part of the Pacific, families Aphroditi-
dae and Polynoidae.] Jn Fauna of the USSR:
Polychaetes, Vol. 2, Part I, Academy of Sciences
of the USSR. Zoological Institute, 272 pp. [in
Russian].

Department of Invertebrate Zoology, Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 169-175

A NEW EULIMID GASTROPOD,
TROCHOSTILIFER EUCIDARICOLA, PARASITIC
ON THE PENCIL URCHIN EUCIDARIS TRIBULOIDES
FROM THE SOUTHERN CARIBBEAN

Anders Warén and Robert Moolenbeek

Abstract.—Trochostilifer eucidaricola (Prosobranchia, Eulimidae) is de-
scribed from Aruba, an island north of Venezuela. The parasite lives on the
ventral side of the host, close to the peristome, permanently attached by its
snout. This is the first Atlantic species of Trochostilifer, a genus previously

known only from the Indo-Pacific area.

The family Eulimidae contains a large
number of species, all of which as far as
known, parasitize echinoderms (Warén
1984). Many species have been described
from the western Atlantic including the Ca-
ribbean, but at present the host is known
for only a small number of them. The eu-
limids represent all different levels of par-
asitism, from species with normal gastro-
pod anatomy that suck body fluids from the
host for a comparatively short time and then
leave the host to digest their meal, to species
that are permanent endoparasites and can
be recognized as gastropods only from the
veliger larvae.

The genus Trochostilifer Warén, 1980
contains five species, all parasitic on cida-
roid sea urchins. Four of these species live
in galls in the spines of their hosts. The fifth
species, Trochostilifer striatus (Hedley, 1901)
lives attached to the test of the host close
to the peristome in a way similar to the
present new species.

Trochostilifer eucidaricola, new species
Figs. 1, 3-8, 14

Holotype. —Zoological Museum, Am-
sterdam (ZMA Moll. 3.88.004). Female
from host number 9 (Fig. 4). Allotype.—
Zoological Museum, Amsterdam (ZMA
Moll. 3.88.005). Male attached with holo-
type (Fig. 6).

Paratypes. —Remaining specimens in
Zoological Museum, Amsterdam, Swedish
Museum of Natural History and U.S. Na-
tional Museum of Natural History, see Ta-
ble 1.

Type locality. —Parasitic around peri-
stome of Eucidaris tribuloides (Lamarck).
Caribbean, Aruba, Malmok, Sandy Thalas-
sia flat with Porites, 2-7 m depth, under
corals and diorite boulders (12°36'N,
70°3.5 W). Leg. I. Peters, Dec 1986.

Material examined.—Known only from
the type locality and the material examined,
listed in Table 1.

Description. —Female: Shell of medium
size, conical, transparent, rather solid, with
high, distinctly shouldered whorls and
sharply set off larval shell. Larval shell (Figs.
1, 3) of typical eulimid shape, consisting of
about four slightly convex, perfectly smooth
whorls, of which 650 um are visible above
the teleoconch (Fig. 3). Holotype with about
five teleoconch whorls, of which the apical
2.5 whorls are distinctly shouldered and the
lower 1.5 almost perfectly flat. Between these
there is one transitional whorl. Body whorl
distinctly angulated at the periphery with a
demarcated basal surface. Shell surface
marked by very shallow, indistinct spiral
furrows and more obvious, sharp incre-
mental lines plus irregularly appearing,
strong incremental scars. Aperture strongly

170 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Specimens of Eucidaris tribuloides para-
sitized by culimids. Explanation: F—female; M—male;
O-—sex characteristics not developed. Numbers in pa-
renthesis are register numbers in ZMA— Department
of Malacology, Zoological Museum of Amsterdam;
SMNH-— Department of Invertebrate Zoology, Swed-
ish Museum of Natural History; USNM-— Division of
Mollusks, U.S. National Museum of Natural History.
Other specimens kept in ZMA and SMNH.

Specimen mid
number T. eucidaricola S. troglodytes Nn. sp
1 (Aug F3.6mm, — 3

1986) M 2.4 mm

(SMNH
3847, left on
host)
2 (Dec F 3.6 mm, — i
1986) M 2.0 mm,
eggs (SMNH
3848)
3 (May O 1.6 mm 2 apical, emp- i
1986) (ZMA Moll. ty galls; 1
3.88.006 basal gall
(Fig. 13),
F 4.6 mm,
M 2.2 mm,
M 2.3 mm,
M 2.2 mm,
O 0.9 mm
(Figs. 9-11)
(galls never
closed)
4 (29? — (1 open apical —
1986) gall, F 2.8
mm, M 2.2
mm, eggs)
>(Dec 1 pair lost; — _
1986) F 4.0 mm,
M 2.6 mm,
eggs (ZMA
Moll.
3.88.007—
008, left on
host)
6 (Dec O 2.0 mm, — —
1986) O 2.0 mm
(ZMA Moll.
3.88.009—
010)
vaCDee O27 nim. — —
1986) O 2.2 mm
(ZMA Moll.
3.88.01 1-
012)

—_—_—_—_—_—_—_—_—_—_—_—_—_—_—_—_——_—_

Table 1.—Continued.

Specimen mid
number T. eucidaricola S. troglodytes n. sp
8 (Dec O 1.6 mm, — —
1986) O 0.6 mm
(ZMA Moll.
3.88.013-
014)
9 (Dec F 3.6 mm, empty gall 2
1986) M 2.0 mm
(USNM
859313);
F 4.8 mm
(holotype),
M 2.6 mm
(allotype)
(ZMA Moll.
3.88.004
and 005)
10 (Dec — O 0.6 mm, on —
1986) spine
11 (Dec — 1 gall with —
1986) M 2.2 mm,
M 2.2 mm,
M 1.8 mm,
M 1.8 mm
(Fig. 15).

prosocline. Outer lip thickened, with the
most protruding part situated one-seventh
of total height from suture. Inner lip thick-
ened, reflected over columella. Height of
holotype 4.68 mm (larval shell broken).
Soft parts (based on rehydrated speci-
mens): Head-foot can be almost completely
retracted into pallial cavity. Animal pig-
mented with bright red, especially along right
side of visceral mass. Tentacles short, flat-
tened, with subcutaneous eyes situated
somewhat laterally to center of bases, meet-
ing above snout to form a “V.”’ Snout large,
cylindrical, with distal attachment disc with
anterior notch and central proboscis open-
ing. Foot smaller than snout, shapeless but
with distinct opercular lobes and large oper-
culum. One large pedal fold on each side of
foot. Body of holotype consisting of 2.5
whorls, of which pallial cavity occupies 0.5.
Holotype female with large closed pallial
Oviduct, no penis. Gill with about 25 low

VOLUME 102, NUMBER 1

Figs. 1-6. 1, Trochostilifer eucidaricola new species, recently settled larva, from host 8, 0.67 mm; 2, Sabinella
troglodytes, recently settled larva from host 10, 0.67 mm; 3-6, Trochostilifer eucidaricola new species; 3, apex
of allotype, 0.65 mm of larval shell visible; 4, holotype, 4.68 mm; 5, female paratype from host 2, 3.6 mm; 6,
male paratype from host 2, 2.6 mm.

We

Figs. 7-8. Trochostilifer eucidaricola, operculum,
female and male, maximum diameter 1.4 and 0.79 mm
respectively.

leaflets, occupying almost all space between
osphradium and oviduct. Hypobranchial
gland anteriorly occupying small, short, tri-
angular area between gill and oviduct, pos-

Figs. 9-11.
respectively.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

teriorly forming a very narrow band. Os-
phradium a simple low ridge in shallow
furrow to the left of and along whole length
of gill. Operculum (Figs. 7—8) yellowish, thin,
paucispiral with thickened part inside the
nucleus serving as a muscular attachment.

Male: In fully grown pairs it is less than
0.6 the size of the female. Shell consisting
of up to 2.5 teleoconch whorls of shape
identical to corresponding whorls of female.
Soft parts similar to those of female, but
with a simple tentacle-like penis curving
forwards, to the right and then backwards
along right side of pallial cavity. No trace
of female reproductive organs seen. Height
of allotype 2.72 mm.

Remarks. —Specimens of T. eucidaricola
and Sabinella troglodytes can easily be dis-
tinguished by the apical teleoconch whorls
which are strongly shouldered in 7. euci-
daricola and evenly convex in S. troglo-

Sabinella troglodytes, female (Fig. 9) and two males from host 3, height 4.6, 2.2, and 2.3 mm

VOLUME 102, NUMBER 1 173

Figs. 12-15. 12, Epidermal fold formed by the host, normally covering distal part of snout of Sabinella
troglodytes, detail of lower left scar in Fig. 13; 13, Spine of Eucidaris tribuloides with deformed (cf. Fig. 15) gall
caused by S. troglodytes, apical view. The spine has formed 3 “‘secondary”’ spines at a right angle to the axis of
the original spine. Four attachment scars of epidermal tissue visible. Maximum diameter of spine 5 mm; 14,
Eucidaris tribuloides, host 1, with male and female of Trochostilifer eucidaricola. Primary spine just to the right
and dorsal to the pair has been removed to show the pair better. Diameter of test 20 mm; 15, normally formed
gall caused by Sabinella troglodytes, host 11, length 9.7 mm.

dytes. Monogamus minibulla (Olsson & much smaller species, ca. 1.5 mm; it has a
McGinty, 1958) bears some resemblance to larval shell of about 1.0 whorl and is para-
T. eucidaricola in having the first part ofthe _ sitic on the sea urchin Echinometra lucunter
teloconch distinctly shouldered, but is a (L.) (Warén, unpub. data).

174

Among the described species of Trochos-
tilifer only T. striatus bears any resemblance
to T. eucidaricola. The other species of the
genus have a shell of typical ““7rochus shape”
and live in galls in the spines of their hosts.
Trochostilifer striatus can, however, easily
be distinguished by having a much more
distinct spiral sculpture, a larval shell of 1.5
whorls and by having evenly convex apical
whorls.

The newly settled larva (Fig. 1) is virtually
indistinguishable from that of Sabinella
troglodytes (Fig. 2), but it 1s possible that
they can be distinguished by the preferred
site on the host, because the specimen in
Fig. 2 had already started to deform the
spine where it was found. This is also the
basis for our determination of these two
specimens. These determinations are not
contradicted by the shape and size of the
larval shells of specimens with enough post-
larval growth to allow identification by the
shape of the teleoconch.

Several dozen specimens of the host ur-
chin were examined and of these 8 were
parasitized by this new species. These spec-
imens are enumerated in Table 1. During
the examination two more species of eu-
limids were found on Eucidaris, viz. Sabi-
nella troglodytes (Thiele, 1925) (Figs. 9-13,
15) and specimens of an undescribed, small,
slender, straight-sided species. Because of
problems with generic assignment of such
species, and because they seemed juvenile,
the latter are left undescribed.

A few years ago the senior author ex-
amined all material of Eucidaris tribuloides
in USNM and found several specimens par-
asitized by Sabinella troglodytes from the
Caribbean area. No specimens of Trochos-
tilifer eucidaricola were then noticed and we
assume it to be a more rare and/or locally
distributed species.

Sabinella troglodytes lives in galls of the
spines of the host and has been reported in
the literature several times (Warén 1984).
We figure the latter species for comparison,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

together with two galls. These specimens are
also listed in Table 1.

Trochostilifer eucidaricola lives attached
to the test of the host by the disc-shaped
distal part of the snout, and it seems very
unlikely that it is able to move. In the center
of the scar on the test of the host there is
a very narrow hole, probably caused by the
proboscis, but in these dried specimens it
proved impossible to trace the proboscis to
find the precise way of parasitism. The spec-
imens were almost always attached in cou-
ples, between a row of primary spines and
the peristome. In two cases also the egg cap-
sules, small globular sacs, were present, at-
tached to the test by a short stalk. Each
capsule contained 100-300 eggs, embryos,
or veliger larvae ready to hatch. The di-
ameter of the shell of the veligers, consisting
of about 0.5 whorls is about 90 wm. This
clearly shows that the species has plankto-
trophic development.

In the five cases when the sex of the spec-
imens in a pair could be determined, the
couples consisted of a small male and a larg-
er female. In one case the sex could not be
determined, because the specimens had not
begun to develop a penis or pallial oviduct.
A single specimen was found only on one
host, which clearly shows that the distri-
bution is not random, but that larvae prefer
to settle on a host already parasitized by a
single specimen.

Acknowledgments

We thank Mrs. Ineke Peeters, Aruba, who
collected the sea urchins and most kindly
allowed us to describe their molluscan fau-
na, and Frere Fredericus, who forwarded
the material to ZMA.

Literature Cited

Hedley, C. 1900. Studies on Australian Mollusca.
2.—Proceedings of the Linnean Society of New
South Wales 26:16-25.

Olsson, A. A., & T. L. McGinty. 1958. Recent marine

VOLUME 102, NUMBER 1

mollusks from the Caribbean coast of Panama
with the description of some new genera and
species.— Bulletins of American Paleontology
39(177):1-58.

Thiele, J. 1925. Gastropoda der deutschen Tiefsee-
Expedition. II. Teil. — Wissenschaftliche Ergeb-
nisse der deutschen Tiefsee-Expedition auf dem
Dampfer “Valdivia” 1898-1899 7(2):36—-382.

Warén, A. 1980. Descriptions of new taxa of Eulim-
idae (Mollusca, Prosobranchia) with notes on
some previously described genera.— Zoologica
Scripta 9:283-306.

Wes)

. 1984. A generic revision of the family Eulim-
idae (Gastropoda, Prosobranchia).—The Jour-
nal of Molluscan Studies, Supplement 13. 96
pp.

(AW) Swedish Museum of Natural His-
tory, Box 50007, S-10405 Stockholm, Swe-
den; (RM) Zoological Museum, Postbus
20125, 1000 HC Amsterdam, The Neth-
erlands.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 176-248

SPRINGSNAILS (GASTROPODA: HYDROBIIDAE)
OF OWENS AND AMARGOSA RIVER
(EXCLUSIVE OF ASH MEADOWS) DRAINAGES,
DEATH VALLEY SYSTEM, CALIFORNIA-NEVADA

Robert Hershler

Abstract. — Thirteen springsnail species (9 new) belonging to Pyrgulopsis Call
& Pilsbry, 1886, and Tryonia Stimpson, 1865 are recorded from the region
encompassing pluvial Owens and Amargosa River (exclusive of Ash Meadows)
drainages in southeastern California and southwestern Nevada. Discriminant
analyses utilizing shell morphometric data confirmed distinctiveness of the
nine new species described herein, as: Pyrgulopsis aardahli, P. amargosae, P.
owenensis, P. pertubata, P. wongi, Tyronia margae, T. robusta, T. rowlandsi,
and 7. salina. Of the 22 springsnails known from Death Valley System, 17
have very localized distributions, with endemic fauna concentrated in Owens
Valley, Death Valley, and Ash Meadows. A preliminary analysis showed only
partial correlation between modern springsnail zoogeography and configuration
of inter-connected Pleistocene lakes comprising the Death Valley System.

This constitutes the second part of a sys-
tematic treatment of springsnails from the
Death Valley System, a large desert region
in southeastern California and southwest-
ern Nevada integrated by a series of lakes
during Pleistocene times. An earlier paper
(Hershler & Sada 1987) dealt with the Ash
Meadows faunule, while this document pro-
vides descriptions of fauna collected during
1985-1987 survey of much of remaining
portions of the System, including waters in
Mono, Adobe, Long, Owens, Indian Wells,
Panamint, and Death Valleys; Amargosa
River drainage; and some areas adjacent to
the above (Fig. 1). A brief discussion of
springsnail zoogeography also is provided,
although a more extensive treatment will be
given following survey of remaining por-
tions of Death Valley System (notably Mo-
jave River drainage) and additional periph-
eral areas.

List of Recognized Taxa

Pyrgulopsis aardahli, new species.
P. amargosae, new species.

P. micrococcus (Pilsbry, 1893).

P. owensensis, new species.

P. perturbata, new species.

P. cf. stearnsiana (Pilsbry, 1899).
P. wongi, new species.

Tryonia margae, new species.

T. protea (Gould, 1855).

T. robusta, new species.

T. rowlandsi, new species.

T. salina, new species.

T. variegata Hershler & Sada, 1987.

Materials and Methods

Localities visited, consisting of low- to
mid-elevation (<2500 m) springs and pe-
rennial streams, are shown in Figs. 2—7 and
listed in the Appendix. Snails were found

VOLUME 102, NUMBER 1

Deep Springs
Valley
oO

(e)
=
rs)
=
o

50
Kilometers

Fig. 1.
area. Stippled areas indicate mountain ranges.

in springs of varying sizes and appearance
as well as low- to moderate-energy (spring-
fed) streams. Photographs of representative
sites are in Figs. 8 and 9.

Snails were relaxed in the field with men-
thol crystals, fixed in 4% buffered formalin
and preserved in 70% ETOH. Material was
collected by author unless otherwise indi-

177

Map showing desert basins of southeastern California and southwestern Nevada comprising study

cated. Water temperature and conductivity
were measured with a temperature-com-
pensated, HACH 16300 conductivity me-
ter.

Methods of anatomical study and pho-
tography of shells and other morphologic
features are routine (Hershler & Sada 1987).
Generalized radular formulae are based on

178 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Selected shell parameters for Pyrgulopsis species. Shell height (SH) and width (SW) are given in
mm. Data for each population include mean values (upper line) and standard deviations (lower). Number of
populations and specimens for which data were obtained are given in parentheses. NW = number of whorls, T
= translation rate, D = distance of generating curve from coiling axis, AS = aperture shape, W = whorl expansion

rate.

Species NW SH

P. aardahli 3.98 2.90
Ci To) G5 O21
P. amargosae 3.86 L977,
(3, 45) 0.20 0.28
P. crystalis S51 2.02
€.3) 0.29 0.24
P. erythropoma 3.46 2.00
(5, 68) O27 0.31
P. fairbanksensis 3:29 2.67
(1, 14) 0.34 O21
P. isolata 4.00 245
(1; 13) 0.14 0.13
P. micrococcus 4.16 2.16
(27, 362) 0.33 0.31
P. nanus 5555) 1.74
(3, 42) 0.24 1.74
P. owensensis 3.61 2.16
(6, 86) 0.26 0.28
P. perturbata 4.37 3:29
(3,39) 0.19 0.39
P. pisteri 3.97 2-3
(2; 31) O:27 0.18
P. cf. stearnsiana 4.14 22
(25) 0.15 0.18
P. wongi 3.86 ZA
(19, 286) 0.27 0.39

Parameter

SW a D AS W
2.20 4.24 0.56 es 2A
0.20 0.54 0.04 0.04 0.19
1.45 4.43 0.57 1.14 2.03
0.25 0.67 0.05 0.06 0.21
£95 3.44 0.50 1.18 2.56
0.18 0.32 0.05 ~ 0.06 0.76
1.82 S332 0.56 Ici 2.13
0.24 0.64 0.05 0.06 0.87
2.28 4.23 0.61 1.09 2.80
0.15 1.03 0.09 0.13 0.07
2.18 3.81 0.58 Le 2.09
0.11 0.36 0.03 0.04 0.20
1.42 S17 0.62 1.19 1.89
0.17 1.05 0.06 0.08 0.28
53 S252 0.55 1.09 2.44
1253 3.52 0.55 1.09 0.34
1.62 4.56 0.59 1.16 2.08
0.18 0.98 0.05 0.05 0.24
apap 5.26 0.62 1.18 2.01
0.31 0.85 0.62 1.18 0.19
1.99 3.46 0.54 1.10 Dest,
0.13 0.45 0.04 0.04 0.31
1.68 I 0.61 1.24 |S
0.12 1s | 0.05 0.06 0.21
ey, 4.10 0:59 | be 2.28
0.24 0.70 0.05 0.05 0.29

examination of SEM micrographs from
varying numbers of populations, and are
presented in following order of tooth types:
centrals, laterals, inner marginals, outer
marginals.

Whorl counts, standard measurements,
and Raupian parameters were obtained from
selected series of live-collected, adult spec-
imens, recognizable by their complete and
thickened inner shell lips. After whorls were
counted (NW), shells were imbedded in clay
with aperture facing up and coiling axis per-
pendicular to the observer’s line of sight.
Shell outlines were traced using camera lu-
cida (25 x or 50) and points on these dig-

itized for calculation of the following pa-
rameters (largely following Kohn & Riggs
1975): shell height (SH), shell width (SW),
length of body whorl (LBW), width of body
whorl (WBW), aperture length (AL), aper-
ture width (AW), translation rate (T), whorl
expansion rate (W), distance of generating
curve from coiling axis (D), and aperture
shape (SA). In order to facilitate compari-
sons, these parameters were also obtained
from the entire set of shells of Ash Meadows
springsnails used in an earlier morphomet-
ric study (which involved generation of a
subset of the above parameters using a
somewhat different methodology; Hershler

VOLUME 102, NUMBER 1 iZ9

Table 2.—Selected shell parameters for 7ryonia species. Shell height (SH) and width (SW) are given in mm.
Data for each population include mean values (upper line) and standard deviations (lower). Number of popu-
lations and specimens for which data were obtained are given in parentheses. NW = number of whorls, T =
translation rate, D = distance of generating curve from coiling axis, AS = aperture shape, W = whorl expansion
rate.

Parameter

Species NW SH SW T D AS WwW
T. angulata 5:10 3.24 1.56 1.99 0.58 1.50 1.66
39) 0.52 O37 0.19 29 0.07 0.08 O22
T. elata noo 1.96 0.82 8.96 0.59 1.34 1.42
(2, 30) 0.53 0.29 0.07 ts 0.05 0.08 0.11
T. ericae 4.71 1.40 0.76 6.42 0.57 125 1.65
(2, 19) 0.66 0.19 0.06 1.06 0.06 0.06 0.11
T. margae 6.21 218 0.98 10.80 0.59 1.48 | Bes
(1;°6) 0.10 Qeh2 0.02 0.93 0.59 0.03 0.19
T. protea 5233 3.53 eg 9.45 0.66 1.38 L351
(1, 6) 0.52 0.45 0.13 0.86 0.04 0.05 0.19
T. robusta 4.22 1.63 1.06 5.03 0.60 1225 1.81
C28) 0.34 0.31 0.18 0.83 0.04 0.04 OT
T. rowlandsi 4.34 1295 1.19 Al 0.62 IH | 1.69
Cl, 16) 0.18 0.16 0.07 1205 0.06 0.08 0.15
T. salina 4.83 2.89 1.70 5212 0.53 1225 1.64
(1,15) 0.20 0.14 0.08 0.60 0.04 0.03 0.09
T. variegata 6.81 4.27 1.69 8.93 0.56 1.50 1.44
Gi 79226) 0.91 1.18 0.32 2.02 0.07 Oat O15

& Sada 1987). Digitizing was done using
CONCH (methodology described in Chap-
man et al., in prep) and a GI'CO Micro-
Digi Pad 12 x 12 linked toa KAYPRO 2000
microcomputer.

Descriptive statistics for all morpholog-
ical variables were obtained for each pop-
ulation and species, and a selection of these
are summarized (by species) in Tables 1 and
2. Discriminant analysis (DA) incorporated
all of the shell morphometric parameters
and were constructed using all specimens
measured for each species (considered as
separate groups). All computations were
performed using SYSTAT (Wilkinson 1986)
on an IBM-AT.

Systematics

Generic diagnoses are in Hershler & Sada
(1987). Species accounts include common
name, synonomy, material examined (lo-
calities numbered [in parentheses] as in Ap-

pendix), diagnosis, description, remarks,
type locality, distribution and habitat,
etymology, and remarks. Asterisked (*) lots
were used for shell measurements. Descrip-
tions emphasize distinctive features from
Shell, radula, body pigmentation, and re-
productive systems of both sexes.

Family Hydrobiidae Troschel, 1857
Genus Pyrgulopsis Call & Pilsbry, 1886
Pyrgulopsis aardahli, new species
Benton Valley springsnail
Figs. 10-14

Material examined.—California: Mono
Co.; Spring at Bramlette Ranch (87), USNM
860406 (holotype), *857951 (paratypes), 21
Apr 1987.

Diagnosis.—A large-sized species with
broadly conical shell. Penis with elongate
filament, very small lobe, and ventral swell-
ing. Penial glandular ridges, 1-3.

Description. —Shell (Fig. 10) 2.6-3.4 mm

180

Fig. 2. Sampling localities in southern Owens Val-
ley and western Indian Wells Valley. Solid lines indi-
cate selected elevation contours (light) and modern
drainage (dark; stipple indicates lakes); dashed lines
indicate historic drainage, including dry lake beds.

high, height/width, 140-—170%. Whorls,
3.75—4.25, moderately convex, moderately
shouldered. Body whorl inflated, height 76-
82% of that of shell. Aperture ovate, aper-
tural plane near-parallel to coiling axis (Fig.
10c). Inner lip thickened, slightly reflected,
adnate to small portion of or slightly sep-
arated from body whorl. Outer lip straight,
thin. Umbilicus moderately open.

Dark, grey-black epithelial (melanic) pig-
ment on most of snout (to just posterior of
cephalic tentacles), along anterior edge of
foot, on operculigerous lobe (Fig. 11). Cen-
tral portions of sides of head/foot lightly

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Sampling localities in northern Owens Val-
ley.

dusted or unpigmented. Tentacles unpig-
mented except for dark ring along bases.
Brown-black subepithelial pigment gran-
ules sometimes forming dark band along
posterior edge of “‘neck.”

Radular (Fig. 12) formula: 5—1—-5/1-1,
2(3)-1-3, 22-24, 28-32 (from paratypes).
Central tooth broadly trapezoidal; basal
process moderately excavated. Penis (Fig.
13b-e) large (extending beyond mantle col-
lar), thin, considerably longer than wide.
Filament slender, moderate in length. Lobe
reduced or absent, with blunt distal edge.
Large, elongate glandular ridge borne on
elongate swelling of ventral penial surface.
Single, smaller ridges sometimes found on

VOLUME 102, NUMBER 1

Oke
WM Ag
ON 4

Mono Valley

Fig. 4. Sampling localities in Mono, Long, and
Adobe Valleys.

tip of lobe, and on dorsal penial surface
proximal to base of filament. Filament
darkly pigmented with subepithelial gran-
ules (Fig. 13b). Capsule gland sub-equal to
albumen gland (Fig. 13a). Seminal recep-
tacle minute, positioned entirely anterior to
bursa copulatrix. Bursa pear-shaped, mod-
erate in size relative to pallial oviduct, po-
sitioned partly posterior to albumen gland.

Type locality. —Spring at Bramlette Ranch
in Benton Valley, Mono County, California.

Distribution and habitat.—Endemic to
type locality (Fig. 14). Common in dense
watercress in uppermost portion of outflow
of small, highly degraded spring (20°C, 190
micromhos/cm).

Etymology. —Named after Jeffrey B. Aar-
dahl, Bureau of Land Management (Cali-
fornia Desert District), who provided di-
verse forms of assistance throughout the
tenure of this project.

Remarks. — Distinguished from similar P.
perturbata by shell appearance and highly
reduced penial lobe.

181

Fig. 5. Sampling localities in Saline, Eureka, and
Deep Springs Valleys.

Pyrgulopsis amargosae, new species
Amargosa springsnail
Figs. 15.46. b7a. bs ks. 19

Material Examined. —California: San
Bernardino Co.; Saratoga Spring (204),
USNM 860401 (holotype), *853515 (para-
types), 7 Feb 1985; 857972, 10 Jul 1986.—
Inyo Co.; Spring crossing path on south side
of Amargosa Gorge (208), *853516, 13 Mar
1986; 857973, 16 Mar 1987.—Spring feed-
ing Amargosa River in Amargosa Gorge
(208), 853517, 13 Mar 1985.—Spring in
marsh east of Grimshaw Lake (209),
*853518, 13 Mar 1985.

Diagnosis.—A small-sized species with
globose to low conical shell. Penis large rel-
ative to head/foot; penial lobe reduced; fil-
ament long, wide. Penial glandular ridges,
0-2.

Description. —Shell (Fig. 15) 1.5-2.7 mm
high, height/width, 1 10-160%. Whorls, 3.5-

182

Fig. 6. Sampling localities in Panamint and north-
ern Indian Wells Valleys.

4.25, convex, moderately shouldered. Body
whorl 74-85% of shell height. Apex, parts
of teleoconch whorls often appearing white
due to erosion. Aperture broadly ovate,
apertural plane near-parallel to coiling axis
(Fig. 15b). Inner lip thickened, reflected
above, adnate to small portion of or slightly
separated from body whorl. Outer lip
straight, thin. Umbilicus chink-like to
broadly open. Spiral striations sometimes
prominent on teleoconch.

Dark, brown-black epithelial pigment on
most of snout (except distal tip), along an-
terior and posterior edges of “‘neck”’ (central
area lighter), on part or all of operculigerous
lobe. Tentacles lightly pigmented.

Radular (Fig. 16) formula: 5(6)—1—5(6)/
1-1, 3-1-4, 27-28, 24—26 (from paratypes).
Central tooth broadly trapezoidal; basal
process moderately excavated. Penis (Figs.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

17a, b, 18b—-e) thickened, longer than wide.
Filament much longer than lobe, about equal
to remaining penis length. Lobe narrow,
variable in shape (Fig. 18b—e). Filament
having highly muscular appearance; surface
lined with weak striations. One or two small
glandular ridges sometimes present at tip of
lobe. Dark, grey-black subepithelial pig-
ment filling filament length almost to distal
tip as well as portions of remaining distal
penis (Fig. 18b). Albumen and capsule
glands about equal in length (Fig. 18a). Sem-
inal receptacle minute, positioned well an-
terior to bursa copulatrix. Bursa copulatrix
club-shaped, small, positioned partly pos-
terior to albumen gland.

Type locality. —Saratoga Spring, southern
Death Valley, San Bernardino Co., Califor-
nia.

Distribution and habitat. — Found in a few
near-brackish springs along lower course of
Amargosa River at Saratoga Spring, Amar-
gosa Gorge, and Grimshaw Lake area (Fig.
19). Type locality an extensive marsh area
(with several large, open pools) fed by sev-
eral small springs. Snails were common in
westernmost spring, both on rocks and sub-
mergent macrophytes in small springpool
(28°C, 5000 micromhos/cm), and on emer-
gent sedges and firm bottom in shallow
stream outflow. At other localities, consist-
ing of small rheocrenes (16-—27°C, 2000-10,
150 micromhos/cm), snails were common
on all substrates.

Etymology. —Referring to distribution
along Amargosa River.

Remarks. — Distinguished from P. micro-
coccus by globose shell; large penis; large
penial filament; small, narrow penial lobe;
and highly reduced glandular ridge. Ranges
of these very similar snails (previously con-
fused by Hershler and Sada 1987:788) are
contiguous.

Pyrgulopsis micrococcus (Pilsbry, 1893)
Oasis Valley springsnail
Figs. 17c, d, 20-25

Amnicola micrococcus Pilsbry in Stearns,
1893:277.

VOLUME 102, NUMBER 1

183

Fig. 7. Sampling localities in Death Valley and Amargosa Basin.

Material examined. —Nevada: Nye Co.;
Spring in Oasis Valley (204), USNM
*850297, 18 Nov 1985; 857961, 16 Mar
1987.—Cane Spring (239), *857936, 10 July
1986:— California: Inyo Co.; Northernmost
of Tecopa Hot Springs (210), *853502, 12-
13 Mar 1985; 857962, 16 Mar 1987.—
Shoshone Spring (211), *853501, 12 Mar
1985; 857962, 16 Mar 1987.—Grapevine
Springs (241): northwest-most spring in
complex, 853503, 1 Mar 1985; Spring
emerging below limestone bench northwest
of ranch, 853504, 26 Feb 1985; Spring on
bench above ranch, 853505, 25 Feb 1985;
Spring at ranch, *853506, 25 Feb 1985;
857964, 14 Mar 1985.—Spring 1.6-km east
of Scotty’s Castle (242), *853507, 26 Feb
1985; 857965, 14 Mar 1987.—Hanaupah
Sprime (222), *8§53508, 16 Feb 1985;

857966, 21 Mar 1987.—Lower spring in
Johnson Canyon (220), *853509, 18 Feb
1985; 857967, 21 Mar 1987.—Stream in
Jail Canyon (196), *853513, 14 Feb 1985;
857970, 19 Mar 1987.—Stream in Surprise
Canyon (194), *857937, 19 Mar 1987;
Stream in Pleasant Canyon (190), *857940,
19 Mar 1987.—China Garden Spring (184),
853511, 9 Feb 1985.—Stream below Dar-
win Falls (184), 857969, 23 Mar 1987.—
Spring above Darwin Falls (184), *853512,
10 Feb 1985.—Stream in Snow Canyon
(182), *857938, 16 Sep 1987, J. Aardahl! —
Stream in Knight Canyon (180), *857939,
14 Apr 1987.—Tennessee Spring (178),
*S 53014, 04Mart, 1985: 5/971, 13, May
1987.—Saline Marsh (152), *853510, 10 Feb
1985; 857968, 28 Mar 1987.
Diagnosis. —A small-sized species, with

184

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 8. Photographs of selected springsnail localities: a, stream in Cottonball Marsh, Death Valley, type
locality of T. salina (3-10-87: photo by P. Rowlands); b, Grapevine Springs, Death Valley, type locality of T.
margae and T. rowlandsi is on the large travertine mound (3-14-87); c, Saline Marsh, Saline Valley (3-29-87);
d, Sand Canyon stream, Indian Wells Valley (3-26-87); e, Fish Slough, Owens Valley, spring sources (including
type locality of P. perturbata) visible in lower portion of photograph (ca. 1975; aerial photo by P. Pister); f,
““NE” Spring at Fish Slough (2-7-85).

globose to ovate-conic shell. Penis with
moderate-sized lobe; distal edge of lobe usu-
ally ornamented with glanduiar ridge.
Description. —Shell (Figs. 20, 21) 1.1-3.1
mm high, height/width, 99-190%. Whorls,
3.25-5.0, well-rounded, with slight shoul-
ders. Spire often irregularly convex due to

bulging of whorls. Body whorl 65-89% of
shell height. Aperture ovate, apertural plane
near-parallel to coiling axis (Fig. 21b). Inner
lip moderately thickened and reflected, usu-
ally slightly separated from body whorl.
Outer lip straight, thin. Umbilicus chink-
like to moderately open.

VOLUME 102, NUMBER 1 185

Fig. 9. Photographs of springsnail localities: a, Warm Springs, Owens Valley (4-19-87); b, Small spring S of
Warren Lake, Owens Valley (4-18-87); c, Spring in Owens Gorge, Owens Valley (5-7-87); d, Hot Creek, Long
Valley (4-23-87).

186 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 10. SEM micrographs of P. aardahli from spring at Bramlette Ranch: a, Holotype, USNM 860406 (shell
height, 3.26 mm); b—d, Paratypes, USNM 857951 (scale of “‘b” and “‘c”’ as in “‘a”’; bar = 150 um).

VOLUME 102, NUMBER 1

Head/foot epithelial pigment ranging from
absent to near-uniformly black (lighter on
central portions of sides of “‘neck’’).

Radular (Figs. 22, 23) formula: (4—7)-1-
(4-7)/1-1, (2-4)-1-(3-6), 17-26, 19-32
(from numerous populations). Central tooth
moderately to broadly trapezoidal; basal
process moderately excavated.

Penis (Figs. 17c, d, 24b—1) barely extend-
ing beyond mantle collar, moderately thick-
ened, longer than wide. Filament slender,
usually extending well beyond edge of lobe.
Lobe highly variable in size and shape, ta-
pering slightly distally. Glandular ridge
(rarely absent) small, positioned on or near
distal edge of lobe. Capsule gland smaller
than albumen gland (Fig. 24a). Seminal re-
ceptacle small, positioned largely anterior
to bursa copulatrix. Bursa pear-shaped,
small to fairly large relative to pallial ovi-
duct, most of length positioned on albumen
gland.

Type locality. —Small spring in Oasis Val-
ley, Nye County, Nevada (precise location
unknown).

Distribution and habitat.—A widespread
form (contrary to Taylor 1985:317) found
in small springs and stream outflows (see
Figs. 8b, c) of Amargosa River drainage as
well as interior drainage in Death, Pana-
mint, and Saline Valleys (5—32°C, 300—4000
micromhos/cm) (Fig. 25). Typically com-
mon on stone, bits of travertine, watercress,
and plant debris.

Pyrgulopsis owensensis, new species
Owens Valley springsnail
Figs. 26a—-d, 27-32

Material examined. —California: Inyo
Co.; Spring ca. 1.6 km south of Mule Spring
(92), USNM 853541, 5 Dec 1987, D. Giu-
liani.— Mule Spring (93), *853540, 19 Nov
1985, D. Wong; 857983, 20 Apr 1987.—
Spring at Graham Ranch (94), *853543, 8
Feb 1985; 857985, 19 Apr 1987.—Wilk-
erson Springs (96), *853542, 8 Feb 1985;
857984, 18 Apr 1987.—Spring at Toll House

187

Fig. 11. Right lateral view of head-foot of P. aar-
dahli, USNM 857951, spring at Bramlette Ranch. Stip-
ples indicate pigmented areas.

(97), *857950, 26 Apr 1987.— Warm Springs
(98), Main spring, *853544, 7 Feb 1985,
857988, 19 Apr 1987; Small spring north
of above, 853545, 7 Feb 1985; 857989, 19
Apr 1987. Mono Co.: Stream in canyon S
of Piute Creek (104), 860404 (holotype),
*857955 (paratypes), 8 May 1987.—Springs
on bench south of Piute Creek (103), North
spring, 857986, 21 Apr 1987; South spring,
857987, 22 Apr 1987.

Diagnosis.—A small- to moderate-sized
species with globose to ovate-conic shell.
Penis large relative to head/foot, filament
short, lobe enlarged, ventral swelling pres-
ent. Penial glandular ridges, 2-6.

Description. —Shell (Figs. 26a—d, 27, 28)
1.5-—2.8 mm high, height/width, 110-160%.
Whorls, 3.0—4.25, well-rounded, sutures
slightly impressed. Protoconch sometimes
slightly tilted. Body whorl inflated, height
74-90% of that of the shell. Apical whorls
sometimes eroded, shell often covered by
deposits. Aperture ovate, angled above;
moderate to large in size. Aperture plane
near-parallel to somewhat tilted relative to
coiling axis (Figs. 27b, 28b). Inner lip slight-
ly to considerably thickened and reflected,
adnate to or (more commonly) slightly sep-
arated from body whorl. Outer lip thin,
straight or slightly sinuate. Umbilicus chink-
like to moderately open.

Dark brown or grey-black epithelial pig-
ment on snout, peripheral portions of

188 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

‘ ee
is : By
4 ‘ =
ad 1% 4 ri
is %, oe)
=
; a)
is ~ 4
2 =
: ¥ an .
at
J a

Fig. 12. Radula of P. aardahli, USNM 857951, spring at Bramlette Ranch: a, Centrals (bar = 12 wm); b,
Laterals (bar = 8.6 um); c, Inner marginal (bar = 6.7 um); d, Outer marginals (bar = 6.0 um).

“‘neck,”’ and dorsal surfaces of tentacles. Radular (Figs. 29, 30) formula: 5(6)—1-
Dark subepithelial granules prominent in 5(6)/1-1, 3-1-3(4), 18-23, 25-37 (from
central areas of sides and sometimes along three populations). Central tooth broadly
posterior edges of “‘neck.”’ trapezoidal; basal process well-excavated.

VOLUME 102, NUMBER 1

Penis (Fig. 31b—1) extending well beyond
mantle collar, elongate. Narrow filament
shorter than lobe. Lobe massive, tapering
only slightly, distal edge blunt. Edge of lobe
ornamented with single elongate or two
shorter glandular ridges. Dorsal penis some-
times having small glandular ridges on lobe
and/or near base of filament. Ventral swell-
ing small, located filament base. Ridge or-
namenting swelling variably-sized. Fila-
ment darkly pigmented with sub-epithelial
granules along most of length (Fig. 31b).
Capsule gland equal to (Fig. 31a) or slightly
smaller than albumen gland. Seminal re-
ceptacle minute, positioned largely or en-
tirely anterior to bursa. Bursa pear-shaped,
small- to moderate-sized, with about half
of length posterior to albumen gland.

Type locality.—Stream in canyon S of
Piute Creek, Mono Co., California.

Distribution and habitat.—Found along
escarpments of White and Inyo Mountains
east side of Owens Valley (Fig. 32). Habitat
consisting of small springbrooks (10—29°C,
300-780 micromhos/cm) (Fig. 9a) where
snails typically common in watercress and/
or on bits of travertine and stone. Syntopic
with P. wongi in spring at Toll House.

Etymology—Referring to occurrence in
Owens Valley.

Remarks. — Distinguished from similar P.
perturbata by shell appearance. more en-
larged penial lobe, and (usually) more nu-
merous penial glandular ndges.

Pyrgulopsis perturbata, new species
Fish Slough springsnail
Figs. 26e-g, 33-36

Material examined.—California: Mono
Co.; Fish Slough: “Northwest Springs.”
South spring (76), USNM 860407 (holo-
type), *853546 (paratypes). 7 Feb 1985:
857990, 24 Apr 1987: “Northwest Springs.”
North spring (76), 853547, 7 Feb 1985:
857991, 20 Apr 1987; ““NE Spring” (77),
*853548, 7 Feb 1985: 857992. 20 Apr 1987:
“BLM Spring” (75), *853549, 7 Feb 1985.

Fig. 13. Reproductive anatomy of P. aardahli,
USNM 857951, spring at Bramlette Ranch: a, Left
lateral view of pallial oviduct complex: b-e, Penes (b.
c, dorsal: d, e. ventral). Stippled areas on penes indicate
glandular ridges (enclosed stipples) and pigmented re-
gions (open). Ag = albumen gland: Bu = bursa copu-
latrix; Cg = capsule gland: Cga = capsule gland open-
ing: Emc = posterior end of pallial cavity; Ov = oviduct:
Pf =penial filament: Plo = penial lobe: Sr = seminal
receptacle: Vc = ventral channel of capsule gland.

Diagnosis.—A large-sized species with
thickened, low-conical shell. Penis large,
lobe and filament small, ventral swelling
present. Penial glandular ridges. 3 or 4.

Description. — Shell (Figs. 26e-g. 33) 2.7-
4.00 mm high, height/width. 130—-180%.
Whorls, 4.25—5.0, only slightly convex, with
pronounced sub-sutural angulations. Body
whorl inflated. 71—85% of shell height. Ap-
erture broadly ovate. Apertural plane tilted
relative to coiling axis (Fig. 33b). Inner lip
usually well-thickened and reflected, broad-
ly adnate to body whorl. Outer lip straight,
thin or moderately thickened. Umbilicus
chink-like.

Dark brown-black epithelial pigment
prominent on snout, tentacles, and periph-
eral portions of sides of head/foot. Pigment

190

Kilometers

Fig. 14. Map showing distribution of P. aardahii.

either light or absent on central portions of
sides of head/foot.

Radular (Fig. 34) formula: 5—1—5/1-1, 3-
1-4(5), 18-19, 29-31 (from paratypes).
Central tooth broadly trapezoidal, basal
process well-excavated. Penis (Fig. 35b—e)
extending well beyond edge of mantle collar,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

thin, considerably longer than wide. Fila-
ment both narrow and short, barely extend-
ing beyond tip of lobe. Lobe tapering only
slightly distally, bearing elongate glandular
ridge along distal edge. Dorsal surface of
penis with small ridge proximal to base of
filament. Ventral swelling arising slightly

VOLUME 102, NUMBER 1 191

Fig. 15. SEM micrographs of P. amargosae from Saratoga Spring: a, Holotype, USNM 860401 (2.28 mm);
b, c (bar = 136 um), Paratypes, USNM 853515.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

¥ ab
\

Fig. 16. Radula of P. amargosae, USNM 853515, Saratoga Spring: a, Centrals (bar = 12 wm); b, Laterals
(bar = 8.6 um); c, Inner marginals (bar = 8.6 wm); d, Outer marginal (bar = 3.8 wm).

proximal to base of filament; ridge (rarely
two) on swelling elongate. Filament dark-
ened along much of length by sub-epithelial
pigment granules. Capsule gland sub-equal

to albumen gland (Fig. 35a). Bursa small,
with ca. 50% of length posterior to albumen
gland. Seminal receptacle minute, posi-
tioned entirely anterior to bursa.

VOLUME 102, NUMBER 1

Fig. 17. SEM micrographs of head-foot and dorsal penes of P. amargosae and P. micrococcus: a (bar = 0.43
mm), b (bar = 0.38 mm), P. amargosae, USNM 853515, Saratoga Spring; c (bar = 0.3 mm), d (bar = 200 um),
P. micrococcus, USNM 857961, spring in Oasis Valley.

194 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Type locality. —‘‘Northwest Springs,”’
south springpool, Fish Slough, Mono Co.,
California.

Distribution and habitat. — Found in three
of the four main springs in Fish Slough (ex-
tinct in ““BLM Spring’) (Figs. 8e, f, 36).
Springs issuing relatively cool (22—26°C),
soft (200-360 micromhos/cm) water. Snails
found only in small vestiges of rheocrene
habitat at various small orifices in north-
west springs (rare at both sites on vegetation
and stones), and at beginning of outflow from
“NE Spring” (common in Chara).

Etymology. — Referring to various distur-
bances to endemic locale, including severe
reduction of original rheocrene habitat as
unanticipated side effect of efforts to con-
serve Owens pupfish (Miller and Pister
1971).

Pyrgulopsis cf. stearnsiana
(Pilsbry, 1899)
Figs. 37-40

Paludestrina stearnsiana Pilsbry, 1899:124.

Fontelicella stearnsiana. —Gregg & Taylor
1965:108.

Pyrgulopsis stearnsiana.—Hershler &
Thompson 1987:30.

Material examined. —California: Kern
Co.; Stream in Sage Canyon (1), USNM
*853520, 26 Mar 1987; 857975, 10 Nov
1987.—Stream in Sand Canyon (6),
*853519, 26 Mar 1987; 857974, 22 Dec
1987, J. Aardahl.

Diagnosis. —A small-sized species with
low conical shell. Penis small relative to
head/foot, penial lobe reduced, filament
elongate, ventral swelling present. Penial
glandular ridges, 1-3.

Description. — Pertaining only to popula-
tions in study area. Shell (Fig. 37) 2.0-2.9
mm high, height/width, 130—170%. Whorls,
4.0-4.5, convex. Body whorl ca. 75% of shell
height. Shell surface encrusted with thick,
black deposits. Aperture broadly ovate,
somewhat angled above, about half as tall

Fig. 18. Reproductive anatomy of P. amargosae:
a, Left lateral view of pallial oviduct complex, USNM
853515, Saratoga Spring; b, c, Dorsal penes, USNM
853515, Saratoga Spring; d, e, Dorsal penes, USNM
857973, spring in Amargosa Gorge. “‘b” through “e”’
drawn to same scale.

as body whorl. Inner lip slightly thickened
and reflected, adnate to small portion of or
slightly separated from body whorl. Outer
lip straight, thin; apertural plane slightly
tilted relative to coiling axis (Fig. 37b).
Dark, grey-black epithelial pigment on
most of snout (except distal tip), proximal
portion of cephalic tentacles, along anterior
and posterior edges of “neck,” part or all of
operculigerous lobe. Pigment on central
portions of sides of neck absent to dark (sub-
epithelial pigment cluster dense in area).
Radular (Fig. 38) formula: 5(6)—1—5(6)/
1-1, 2(3,4)-1-3(4,5), 23-24, 23-29 (from
single population). Central tooth broadly
trapezoidal; basal process moderately ex-
cavated. Penis (Fig. 39b—e) rarely protrud-
ing beyond edge of mantle collar, longer than
wide. Filament slender, sub-equal to re-
maining penis length. Reduced lobe scarcely

VOLUME 102, NUMBER 1

Kilometers

118°

Fig. 19. Map showing distribution of P. amargosae.

projecting distal to base of filament. Tip of
lobe often ornamented with small glandular
ridge; similar, single ridges on dorsal penial
surface about halfway from base of penis to
base of filament, on ventral surface near
(sometimes on) inner edge. The latter ridge
enlarged, borne on low swelling (Fig. 39e).

195

Wu

Filament with dark sub-epithelial pigment
streak (Fig. 39b). Albumen gland sub-equal
to capsule gland (Fig. 39a). Seminal recep-
tacle small, positioned anterior to bursa
copulatrix. Bursa copulatrix small relative
to capsule gland, positioned partly posterior
to gland.

196 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 20. Camera lucida drawings of shells of P. micrococcus: a, USNM 857936, Cane Spring (2.24 mm); b,
USNM 853501, Shoshone Spring (1.96 mm); c, USNM 853502, Tecopa Hot Springs (1.90 mm); d, USNM
853506, Grapevine Springs (2.16 mm); e, USNM 853508, Hanaupah Spring (2.26 mm); f, USNM 853510,
Saline Marsh (1.96 mm); g, USNM 853512, spring above Darwin Falls (1.94 mm); h, USNM 857937, Surprise

Canyon stream (1.88 mm).

Type locality. —Near Oakland, Alameda
County, California.

Distribution and habitat. —Distribution
given by Taylor (1981:152) as ““Central Cal-
ifornia, from Sonoma County to Monterey
County along the coast, and inland in the
foothills of the Sierra Nevada.” Found in
the study area in two moderate-sized, cool
streams (16—20°C, 550 micromhos/cm) sep-
arated by 25 km along eastern slope of Sier-
ra Nevada in Indian Wells Valley (Fig. 40).
Snails common in watercress and Chara.

Remarks.—Shells of the Indian Wells
Valley snails closely resemble P. stearnsiana
from the San Francisco Bay area and their
penes conform to a general description for
the species provided by Gregg and Taylor
(1965:107). Additional study, including ex-
amination of anatomy of topotypes or near-
topotypes, will be necessary to confirm

identity of the snails from Indian Wells Val-
ley.

Pyrgulopsis wongi, new species
Wong’s springsnail
Figs. 41-47

Material examined. —California: Inyo.
Co.; Spring at Little Lake, east of HW 395
(10), USNM *853521, 9 Feb 1985.—Hog-
back Creek (Monache Mtn. quad.) (20),
*853522, 28 Jun 1985, D. Wong.—Summit
Creek (21), 853523, 12 Apr 1987.—Spring
ca. 3.0 km north of Summit Creek (21),
*853524, 12 Apr 1987.—Spring at Cabin
Bar Ranch (25), 853525, 12 Oct 1987, J.
Goldberg and M. Shumway.—Lubkin Creek
(34), 853526, 16 Apr 1987.—Spring feeding
Lubkin Creek (34), 857942, 25 Apr 1987.—
Spring along east side of Tuttle Creek (35),

VOLUME 102, NUMBER 1 197

Fig. 21. SEM micrographs of P. micrococcus: a, USNM 853502, Tecopa Hot Springs (1.98 mm); b, USNM
853512, spring above Darwin Falls: c, USNM 853512. Tecopa Hot Springs (bar = 120 um).

198 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

f™ i —_ — i i

Fig. 22. Central radular teeth of P. micrococcus: a, USNM 857936, Cane Spring (bar = 7.5 um); b, USNM
857963, Tecopa Hot Springs (bar = 7.5 um); c, USNM 857965, spring E of Scotty’s Castle (bar = 7.5 um); d,
USNM 857968, Saline Marsh (bar = 8.6 um).

857943, 30 Apr 1987.—Hogback Creek Giuliani.—Spring on hill south of Warren
(Lone Pine quad.) (36), *853527, 8 Feb 1985; Lake (57), *853530, 18 Apr 1987.—Spring
857976, 16 Apr 1987.—Boron Springs (40), in canyon south of Shannon Canyon (59),
*853528, 17 Apr 1987.—Stream in Charlie 857944, 20 Apr 1987.—Spring along north
Canyon (43), *853529, 12 Jun 1987, D. side of upper Pine Creek (69), *853531, 24

VOLUME 102, NUMBER 1

Fig. 23. Lateral and marginal radular teeth of P. micrococcus: a, Laterals, USNM 857936, Cane Spring (bar
= 6.0 um); b, Laterals, USNM 857966, Hanaupah Spring (bar = 7.5 um); c, Inner marginals, USNM 857937,
Surprise Canyon stream (bar = 5.0 um); d, Outer marginals, USNM 857963, Tecopa Hot Springs (bar = 4.3

pm).

Apr 1987; 857977, 15 Sep 1987,D. Wong.— 857945, 3 May 1987.—Spring in Owens
West spring in Birchim Canyon (70), 860403 Gorge (74), *853532, 14 May 1985, D.
(holotype), *857941 (paratypes), 3 May Wong; 857978, 8 May 1987.—French Spring
1987.—East spring in Birchim Canyon (70), (89), *857947, 4 May 1987.—Barrel Springs

200

Fig. 24. Reproductive anatomy of P. micrococcus:
a, Left lateral view of pallial oviduct complex, USNM
857961, spring in Oasis Valley; b—-i, Penes (all dorsal
except “1’’) (b, 1, USNM 857936, Cane Spring; c, USNM
853505, Grapevine Springs; d, USNM 857962, Shosh-
one Spring; e, USNM 857963, Tecopa Hot Springs; f,
USNM 857967, Johnson Canyon spring; g, USNM
857971, Tennessee Spring; h, USNM 853512, spring
above Darwin Falls).

(91), *857948, 18 Aug 1987.—Spring at Toll
House (97), *857949, 26 Apr 1987.—Corral
Springs (160), *853535, 8 Feb 1985; 857979,
29 Mar 1987.—Antelope Spring (162),
*853536, 8 Feb 1985. Mono Co.; Spring
along Marble Creek (82), *853533, 22 Apr
1987.—Springs at north end of Blind Spring
Valley (86); North Spring, *857946, 21 Apr
1987; South spring, 853534, 21 Apr 1987.—
Layton Springs (122), *853537, 6 Feb 1985;
857980, 23 Apr 1987.—River Springs (132),
*853538, 6 Feb 1985; 857981, 21 Apr 1987.

Diagnosis. —A small- to moderate-sized
species with globose to low conical shell.
Penis massive, with large filament and mod-
erate-sized lobe. Ventral penis with two
prominent swellings bearing glandular
ridges. Glandular ridges, 7-12.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Description. —Shell (Figs. 41, 42) 1.2—3.0
mm high, height/width, 100—160%. Whorls,
3.25—4.5, well rounded, with slightly an-
gulated shoulders. Body whorl 76-87% of
shell height. Aperture broadly ovate, angled
above. Apertural plane slightly tilted rela-
tive to coiling axis (Fig. 42c). Inner lip
slightly thickened, usually only minimally
reflected, either adnate to portion of or
slightly separated from body whorl. Outer
lip straight, thin. Umbilicus chink-like to
moderately open.

Dark grey-black epithelial pigment on
snout, proximal tentacles, and peripheral
portions of sides of head/foot. Entirety of
head/foot often very dark. Central portions
of sides of head/foot usually with densely
clustered dark, subepithelial pigment gran-
ules.

Radular (Figs. 43, 44) formula: (4—7)—1-
(4-7)/1-1, (2-—5)—1-(3-6), 17-26, 20-34
(from numerous populations). Central tooth
relatively square-shaped; basal process well-
excavated. Penis (Figs. 45b-—l, 46) flattened,
longer than wide. Filament fairly broad,
short relative to length of remaining penis.
Lobe slightly shorter than filament, broad,
often widening distally. Glandular ridge ar-
rangement typically as follows: 1) elongate
ridge lining edge of lobe; 2) similar, stalked
ridge on ventral surface just proximal to
lobe; 3) similar ridge (rarely very reduced)
on fleshy projection of ventral surface prox-
imal to above; 4) elongate ridge covering
proximal-most two-thirds of dorsal fila-
ment; 5) very elongate ridge on dorsal sur-
face curving from just proximal to filament
to lobe; 6) shorter ridges on dorsal surface
along right side of (5) and in area between
base of filament and edge of lobe. Modifi-
cations of above include addition of a few
small ridges on central dorsal surface, ad-
dition of a single small ridge (sometimes
stalked) on ventral surface, apparent frag-
mentation of ridges (particularly “1,” ““5’’);
apparent merging of adjacent ridges on dor-
sal surface. Filament darkened with sub-ep-
ithelial pigment along most of length. Cap-
sule gland sub-equal to albumen gland (Fig.

VOLUME 102, NUMBER 1

(@)
Kilometers

Fig. 25.

45a). Seminal receptacle small, positioned
lateral to anterior bursa copulatrix. Bursa
pear-shaped, massive, with about one-third
of length posterior to albumen gland.

Type locality.—Spring in Birchim Can-
yon north of Bishop, Inyo Co., California.

Distribution and habitat. — Widespread in
Owens Valley along eastern escarpment of
Sierra Nevada from Pine Creek south to

201

Map showing distribution of P. micrococcus. Snail occurs at 15 localities in Ash Meadows.

Little Lake, and along western side of val-
ley from French Spring to Marble Creek.
Also found in a few sites in Long, Adobe,
and Deep Springs Valleys (Fig. 47). Habitat
includes seeps and spring-fed streams of
small-moderate size (9.5-—22°C, 85-450
micromhos/cm) (Fig. 9b, c). Snails typically
common in watercress and/or on small bits
of travertine and stone.

202

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 26. Camera lucida drawings of shells of P. owensensis and P. perturbata: a—d, P. owensensis (a, USNM
853540, Mule Spring [2.1 mm]; b, USNM 857950, spring at Toll House [2.3 mm]; c, USNM 857955, stream
in canyon S of Piute Creek [2.08 mm]; d, USNM 853543, spring at Graham Ranch [2.16 mm]); e-g, P. perturbata
(e, USNM 853546, Fish Slough, ““Northwest Springs,” [3.93 mm]; f, USNM 853549, Fish Slough, ““BLM Spring,”
[3.22 mm]; g, USNM 853548, Fish Slough, ““NE Spring” [3.97 mm]).

Etymology. —Named after Darrell Wong,
California Fish and Game, for his assistance
during the project and interest in conserving
Owens Valley springsnails.

Remarks. —Similar in penial morphology
to P. californiensis (Gregg & Taylor) from
western California (Taylor 1981:154). Dis-
tinguished from above by globose shell, and
more numerous stalked glandular ridges on
ventral penial surface.

Genus Tryonia Stimpson, 1865
Tryonia margae, new species
Grapevine Springs elongate tryonia
Figs. 48-50, 5la, 52

Material examined. —California: Inyo
Co.; Grapevine Springs (241); (upper) Warm

spring on limestone bench, USNM 860408
(holotype), *857952 (paratypes), 14 Mar
1987; Spring on limestone bench above
ranch 853555, 25 Feb 1987; 857996, 14
Mar 1987.

Diagnosis.—A small-sized species with
narrow, turriform shell. Penis small, with 3
papillae on inner curvature, and a single
papilla (sometimes absent) on outer cur-
vature. Distal tip of penis with elongate
swelling on inner side.

Description. — Shell (Fig. 48) 1.5-—2.9 mm
high, height/width, 250—300%. Whorls, 4.5-
6.25, rounded, with angulated shoulders.
Sutures often deeply impressed. Apex blunt,
protoconch slightly depressed (Fig. 48g).

VOLUME 102, NUMBER 1

203

Fig. 27. SEM micrographs of P. owensensis from stream in canyon S of Piute Creek: a, Holotype, USNM
860404 (2.0 mm); b-d, Paratypes, USNM 857955 (bars = 120 um, 176 um).

Body whorl 45-55% of shell height. Marl
frequently covering apical whorls, with near-
entirety of shell sometimes blanketed. Ap-
erture narrowly ovate. Inner lip well-reflect-
ed, slightly thickened, adnate to or slightly
separated from body whorl. Outer lip sin-
uate, thin (Fig. 48d, f). Umbilicus chink-
like. Protoconch near-smooth (Fig. 48g).
Collabral growth lines well developed on

teleoconch, often elevated at irregular in-
tervals. Spiral sculpture consisting of weak
striations.

Dark brown-black epithelial pigment
covering most of head/foot, although some-
times absent on tentacles, and lighter on
snout and central portion of sides of ‘“‘neck.”

Radular (Figs. 49, 50) formula: 4(5)}-1-
4(5)/1(2)-1(2), 2(3}-1-3(4), 20-21, 22-28.

204

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 28. SEM micrographs of P. owensensis from Warm Springs: a (1.92 mm), b, d, USNM 853544, main
spring; c, USNM 853545, spring N of above (bar = 136 um).

Central tooth broadly trapezoidal, basal
cusps large, basal process well excavated.
Central cusps of lateral tooth only slightly
enlarged (Fig. 49c, d). Penis (Fig. 51a) ex-
tending only slightly beyond edge of mantle
collar. Enlarged papilla located on outer edge
of penis at base. Two smaller papillae po-
sitioned along distal half on inner edge; sim-
ilar small papilla usually found near base

on inner edge. Brown-black epithelial pig-
ment usually prominent near distal tip, at
bases of two distal papillae. Lighter pigment
dusting much of dorsal surfaces of penis and
2 proximal papillae.

Type locality. —Warm spring on lime-
stone bench in Grapevine Springs complex,
northern Death Valley, Inyo Co., Califor-
nia.

VOLUME 102, NUMBER 1

205

Fig. 29. Radula of P. owensensis, USNM 857955, stream in canyon S of Piute Creek: a, Centrals (bar = 10
um); b, Laterals (bar = 6.7 um); c, Inner marginal (bar = 6.0 wm); d, Outer marginals (bar = 6.0 um).

Distribution and habitat.— Found in two
sites (Separated by about 1 km) among
Grapevine Springs in northern Death Val-
ley (Fig. 52). Habitat consists of upper por-
tions of shallow spring brooks flowing over
travertine (Fig. 8b), on which snails were
moderately common. Springs were highly

mineralized (1050-1075 micromhos/cm),
but differed considerably in temperature
(west spring, 26—27°C; east, 36—37°C). Syn-
topic with 7. rowlandsi in the thermal spring.

Etymology. —From Latin marga, mean-
ing marl, and referring to occurrence of snail
in travertine springs.

206 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 30. Radula of P. owensensis, USNM 857988, Warm Springs (main spring): a, Centrals (bar = 6.7 um);
b, Laterals (bar = 6.0 um); c, Inner marginal (bar = 6.0 wm); d, Outer marginals (bar = 3.8 um).

VOLUME 102, NUMBER 1

Tryonia protea (Gould, 1855)
Desert tryonia
Figs. 52-54

Amnicola protea Gould 1855:129.
Melania exigua Conrad 1855:269.
Pyrgulopsis blakeana Taylor 1950:30.
Pyrgulopsis cahuillarum Taylor 1950:31.

Material examined. —California: Mono
Co.; Hot Creek, USNM *857954, 23 Apr
1987.

Diagnosis.—A moderate to large-sized
species (shell height typically 3-7 mm) with
elongate-conic to turriform shell. Shell
smooth or with spiral and/or collabral
sculpture varying from faint striations to
elevated ridges.

Description of Hot Creek population. —
Shell (Fig. 53) 3.0-4.5 mm high, height/
width, 210—242%. Whorls, 4.75-6.25,
rounded. Sutures moderately impressed.
Body whorl 55-61% of shell height. Pro-
toconch slightly depressed (Fig. 53b), apex
of adults invariably highly eroded. Aperture
ovate, apertural plane tilted relative to coil-
ing axis. Inner lip only slightly thickened
and reflected, broadly adnate to body whorl.
Outer lip thin, slightly sinuate. Umbilicus
chink-like, near-absent. Smooth surface of
protoconch lined with well-spaced rows of
numerous, shallow perforations (Fig. 53b,
c). Teleoconch with strong growth lines; 10-
20 spiral lines prominent on middle whorls,
sometimes extending to aperture.

Brown epithelial pigment dense on most
of snout. Black patches of subepithelial pig-
ment along edges of sides of head/foot. Pal-
lial roof with paired, black pigment patches
extending along length of ctenidium.

Radular (Fig. 54) formula: 5(6)—1-5(6)/
2(3)—2(3), (3-5)—1-(6-8), 35-36, 28-31
(from single population). Central tooth
broadly trapezoidal; basal cusps large, basal
process moderately excavated. No males
found among the more than 100 specimens
collected.

Type locality.—Colorado Desert (proba-

207

1.0 mm

Fig. 31. Reproductive anatomy of P. owensensis:
a, Left lateral view of pallial oviduct complex, USNM
857955, stream in canyon S of Piute Creek; b—i, Penes
(b [dorsal], i [ventral], USNM 857955, stream in can-
yon S of Piute Creek; c [dorsal], f [ventral], USNM
857983, Mule Spring; d [dorsal], g [ventral], h [ventral],
USNM 857988, Warm Spring (main spring); e [ven-
tral], USNM 857989, Warm Springs [spring N of
above]). ““b’’ through “i” drawn to same scale.

bly in Riverside County, California [based
on Blake 1857]).

Distribution and habitat. —‘‘Western Utah
to southeastern California, adjacent Baja
California, and southeastern Arizona’”’
(Taylor 1981:155), with only two living
populations known from California: that of
Hot Creek (Fig. 52) and Dos Palmas Spring
in Riverside County (Taylor 1981:154). At
Hot Creek (Fig. 9d), snails were rare on var-
ious submergent macrophytes lining sides

208

Kilometers

Fig. 32. Map showing distribution of P. owensensis.

of the swiftly flowing thermal stream (32°C,
480 micromhos/cm).

Remarks. —The taxonomic concept of this
form is vague (Taylor 1966:53—54) and sta-
tus of the few living (and mostly widely sep-
arated) populations (Taylor 1985:fig. 35) re-
mains unclear. The Hot Creek population
is referable to 7. protea on basis of shell
form, size, and sculpture, but confirmation

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

by anatomical comparison with snails from
Dos Palmas Spring (that represent near to-
potypes) would be desirable.

Tryonia robusta, new species
Robust tryonia
Figs. 51d, 52, 55-58

Material examined. —California: Inyo
Co.; Nevares Springs (230); Spring on trav-

VOLUME 102, NUMBER 1 209

Fig. 33. SEM micrographs of P. perturbata from Fish Slough, ““Northwest Springs,” south spring: a, b,
Holotype, USNM 860407 (3.57 mm); c, Paratype, USNM 853456 (bar = 200 um).

ertine mound, USNM 860411 (holotype), 1985—Travertine Springs; South spring,
*853557 (paratypes), 2 Feb 1985; 857999, *853559, 2 Feb 1985; 857959, 12 Mar 1987;
16 Nov 1985; 857958, 12 Mar 1987;Spring North spring, 857960, 12 Mar 1987.

issuing from base of mound, 853558, 2 Feb Diagnosis.—A small-sized species with

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 34. Radula of P. perturbata, USNM 857990, Fish Slough, “‘“Northwest Springs,” south spring: a, Centrals
(bar = 13.6 wm); b, Laterals (bar = 10 wm); c, Laterals and inner marginals (bar = 12 wm); d, Outer marginals
(bar = 10 wm).

unusually broadly conical shell for genus. high, height/width, 140-170%. Whorls,
Penis moderate-sized, with single papillaon 3.75-4.75, rounded, with pronounced
outer curvature. shoulders. Sutures impressed. Body whorl

Description. —Shell (Fig. 55) 1.1-2.1mm 68-80% of shell height. Protoconch (Figs.

VOLUME 102, NUMBER 1

56a-c) simple, sometimes eroded. Aperture
near-ovate, sometimes sharply angled
above; apertural plane slightly tilted relative
to coiling axis. Inner lip thickened, mod-
erately reflected, usually slightly separated
from body whorl. Outer lip thin, near-
straight. Umbilicus chink-like. Protoconch
surface wrinkled (Figs. 56a—c). Teleoconch
often with weak spiral striations, particu-
larly on upper whorls.

Operculum (Fig. 56d) thin, paucispiral,
with 2.5—3.0 rapidly expanding whorls.

Brown-black melanic pigment dark on
snout and posterior edge of “‘neck,”’ variable

on central portions of sides of head/foot; -

sub-epithelial pigment granules prominent
in latter. Mantle roof epithelial pigment typ-
ically uniformly dark brown-black.

Radular (Fig. 57) formula: (5—7)-1-(5-7)/
1-1, 3(4)-1-5(6), 24, 22—27 (from two pop-
ulations). Central tooth relatively square-
shaped; central cusps elongate, basal cusps
small, basal process highly excavated. Ce-
phalic tentacles covered with sparse, irreg-
ular patches of hypertrophied cilia, rarely
forming short tracts. Penis (Figs. 51d, 58)
considerably longer than wide, moderately
thickened. Attachment area broad, with rest
of narrow length coiled in tight, clockwise
fashion. Papilla small, conical, positioned
along outer curvature just proximal to mid-
length. Distal tip blunt, with inner side
slightly swollen; surface of tip densely cov-
ered with elongate cilia. Distal tip of papilla
simple, lightly ciliated (Fig. 58b).

Type locality.—Nevares Springs, Death
Valley, Inyo County, California.

Distribution and habitat.— Found at Ne-
vares and Travertine Springs along base of
Funeral Mountains in mid-eastern Death
Valley (Fig. 52). Snails found commonly on
plant debris and travertine in shallow out-
flows of four thermal springs (30—37°C, 900-—
975 micromhos/cm). Extinct at Texas
Springs (located between above complexes).

Etymology. —Referring to unusually
broad shell for genus.

Remarks. —Shell form and penial shape

211

Fig. 35:

Reproductive anatomy of P. perturbata
from Fish Slough, “‘ Northwest Springs”: a, Left lateral
view of pallial oviduct complex, USNM 857990, south
spring; b-e, Penes (b [dorsal], c [dorsal], d [ventral],
USNM 857990, south spring; e [ventral], USNM
857991, north spring).

and lobation pattern are highly distinctive,
and relationship between this local endemic
and other congeners in region is unclear.
Species closely conforms to typical Tryonia
in other features (including details of female
reproductive morphology).

Tryonia rowlandsi, new species
Grapevine Springs squat tryonia
Figs. 51b, 52, 59, 60

Material examined. —California: Inyo
Co.; Grapevine Springs (241); (upper) Warm
spring on limestone bench, USNM 860409
(holotype), *857953 (paratypes), 14 Mar
1987; (lower) Warm spring on limestone
bench, 857997, 14 Mar 1987.

Diagnosis.—A small-sized species with
elongate-conic shell. Penis large, with single
papilla on outside curvature and 2 papillae
on inside curvature.

Description. —Shell (Fig. 59) 1.7—2.3 mm

212

50
Kilometers

Fig. 36. Map showing distribution of P. perturbata.

high, height/width, 140—190%. Whorls, 4.0—
4.75, rounded, somewhat shouldered. Su-
tures impressed. Body whorl 62-74% of shell
height. Protoconch tilted relative to teleo-
conch. Aperture ovate, apertural plane
slightly tilted relative to coiling axis. Inner
lip thin, only very slightly reflected, adnate
to or slightly separate from body whorl.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Outer lip thin, near-straight. Umbilicus nar-
row, chink-like. Protoconch mottled with
series of short, predominantly spiral stria-
tions (Fig. 59e). Teleoconch with weakly de-
veloped spiral lines, particularly on upper
whorls.

Head/foot and visceral coil usually uni-
formly coated with dark brown epithelial

VOLUME 102, NUMBER 1

213

Fig. 37. SEM micrographs of P. cf. stearnsiana, USNM 853519, Sand Canyon (a, 3.04 mm; bar = 136 um).

pigment. Central portions of sides of head/
foot consisting of either a thin band or broad
patch, often lighter or unpigmented.
Radular formula (Fig. 60): 4(5)—1-4/1(2)}+
1, 2(3)}-1-2(3), 16, 20 (from paratypes).

Central tooth broadly trapezoidal; basal
cusps small, basal process only moderately
excavated. Lateral tooth having unusual
““claw-like’’ appearance, with cusps few and
widely separated. Penis (Fig. 51b) elongate,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 38. Radula of P. cf. stearnsiana, USNM 857974, Sand Canyon: a, Centrals (bar = 10 wm); b, Laterals
(bar = 6.0 um); c, Inner marginal (bar = 8.6 wm); d, Outer marginals (bar = 6.0 wm).

distal tip with slightly swollen inner edge.
Attachment area broad, with remaining pe-
nis near-totally uncoiled. Single, small pa-
pillae located on outer curvature near mid-
point, and on inner curvature near distal

tip. Inner curvature also bearing enlarged,
basal papilla. Penial pigmentation consist-
ing of small, dark patch near distal tip and/
or light dusting on broad area near base.
Type locality.—Grapevine Springs,

VOLUME 102, NUMBER 1

Grapevine Mountains, Death Valley, Inyo
Co., California.

Distribution and habitat. — Found in two
thermal, highly mineralized springs on
limestone bench in northeast portion of
Grapevine Spring complex (32-—36°C, 1 100-
1050 micromhos/cm) (Fig. 52).

Etymology.—Named after Peter Row-
lands, Resources Management Division,
National Park Service, Death Valley, for his
efforts in both assisting this study and con-
serving local aquatic gastropods in the Mon-
ument.

Tryonia salina, new species
Cottonball Marsh tryonia
Figs. 5le, 52, 61, 62

Tryonia sp.— Taylor in LaBounty and Dea-
con 1972:775.

Material examined.—California: Inyo
Co.; Spring in Cottonball Marsh (231),
USNM 860410 (holotype), .*853556 (para-
types), 22 Feb 1985; *857998, 11 Mar 1987.

Diagnosis. —A moderate-sized-species.
with elongate-conic shell bearing well-de-

veloped spiral striae. Penis with three pa-
pillae on inner curvature and single papilla
on outer curvature.

Description. —Shell (Fig. 61) 2.4—3.1 mm
high, height/width, 150—190%. Whorls, 4.5-
5.25, well-rounded, shouldered above. Su-
_.tures impressed. Shell loosely coiled, with
slight separation of upper whorls common.
Body whorl 61-68% of shell. height. Pro-
toconch tilted relative to teleoconch. Ap-
erture ovate, apertural plane slightly tilted
relative to coiling axis. Inner lip thin, only
slightly reflected, narrowly adnate to or
slightly separated from body whorl. Outer
lip thin, near-straight. Umbilicus chink-like
to moderately. open. Protoconch smooth
(Fig. 61c). Teleoconch with strong, regular-
ly-spaced spiral striations, usually continu-
ing onto body whorl, interacting with oc-
casionaHy highly pronounced growth: lines
to produce a weakly cancellate appearance.

Brown epithelial pigment dark on snout,

2AS

0.5 mm

Fig. 39. Reproductive anatomy of P. cf. stearn-
siana, USNM 857974, Sand Canyon: a, Left lateral
view of pallial oviduct complex; b-e, Penes (b, c, dorsal
aspects; d, e, ventral aspects).

dorsal tentacles, and all but narrow central
strip of head/foot.

Radular (Fig. 62) formula: 5(6)—1-5(6)/
2(3)—2(3), 3(4)-1-4(5), 19-26, 26-28 (from
paratypes). Central tooth broadly trapezoi-
dal, basal cusps small, basal process weakly
excavated. Penis (Fig. 5le) longer than wide,

- moderately thickened. Distal tip with

prominent swelling of inner side. Inner cur-
vature with two small papillae positioned
on inner curvature near tip, and larger basal
papilla. Outer curvature with enlarged basal
papilla. Dorsal penis variably pigmented by
brown epithelial melanin, with especially
dark cover on basal penis and papillae.

Type locality.—Spring brook in Cotton-
ball Marsh, near base of Panamint Moun-
tains at west side of salt pan in Death Valley,
Inyo Co., California.

Distribution and habitat. — Found in a few
small, cool, brackish (17°C, >5000 mi-
cromhos/cm) spring brooks emerging in

216 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

50
Kilometers

Fig. 40. Map showing distribution of P. cf. stearnsiana in the study area.

Cottonball Marsh, ca. 1.6 km out onto salt oped shell sculpture, penial lobation pat-
pan east of Salt Springs. Snails common on __ tern, and morphology of lateral radular teeth.
large, actively growing tufa blocks. Highly
remote area in near-pristine condition.
Etymology. —Referring to highly miner-
alized spring habitat. Amargosa Se
Remarks.—Very similar to T. rowlandsi Figs. 51c, 52, 63, 64
from northern Death Valley, but distin- Zryonia variegata Hershler & Sada 1987:
guished by larger size, more highly devel- 817.

Tryonia variegata
Hershler and Sada, 1987

VOLUME 102, NUMBER 1

217

Fig. 41.

Camera lucida drawings of shells of P. wongi: a, USNM 853531, spring alongside upper Pine Creek

(2.18 mm); b, USNM 853524, spring N of Summit Creek (2.24 mm); c, USNM, 853527, Hogback Creek (Lone
Pine) (1.74 mm); d, USNM 853530, spring S of Warren Lake (2.08 mm); e, USNM 853521, spring at Little
Lake (1.44 mm); f, USNM 857947, French Spring (2.32 mm); g, USNM 857949, spring at Toll House (1.66
mm); h, USNM 857946, spring (north) at N end of Blind Spring Valley (2.34 mm); 1, USNM 853536, Antelope
Spring (1.90 mm); j, USNM 853535, Corral Springs (1.80 mm); k, USNM 853538, River Springs (2.66 mm);

1, USNM 853537, Layton Spring (1.78 mm).

Material examined. —California: San
Bernardino Co.; Saratoga Spring (204),
USNM *853554, 27 Feb 1985; 857995, 10
Jul 1986. Inyo Co.; Spring crossing path on
south side of Amargosa Gorge (208), 853553
(empty shell).—Spring in marsh east of
Grimshaw Lake (209), *853551, 13 Mar

1985; 857994, 22 Mar 1987.—Shoshone
Sprme (11); *853550, 12 Mar 1985;
857993, 16 Mar 1987.—Resting Spring
(212). .853552,.13 Mar 1985.
Diagnosis.—A variably-sized species with
turriform-aciculate shell. Penis large, with
3 or 4 papillae on inner curvature and single

218 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

>

Fig. 42. SEM micrographs of P. wongi from west spring in Birchim Canyon: a, Holotype, USNM 860403
(2.14 mm); b, c, Paratypes, USNM 857941 (bar = 136 um).

papilla (sometimes absent) on outer cur- 4.75-9.75, slightly to moderately rounded,
vature. sometimes slightly shouldered. Sutures im-

Description. —Shell (Fig. 63) 1.2-7.5 mm __ pressed. Body whorl 37-63% of shell height.
high, height/width, 170-—350%. Whorls, Aperture narrowly ovate, apertural plane

VOLUME 102, NUMBER 1 219

Phe ay
rhe.
Fig. 43. Central radular teeth of P. wongi: a, USNM 857941, west spring in Birchim Canyon (bar = 10 um);

b, USNM 853521, spring at Little Lake (bar = 7.5 um); c, USNM 853536, Antelope Spring (bar = 7.5 wm); d,
USNM 857981, River Springs (bar = 12 um).

near-parallel with coiling axis. Inner lip Umbilicus chink-like to moderately open.
slightly thickened and reflected, adnate to Protoconch smooth to irregularly wrinkled
or slightly separated from body whorl. Out- (Fig. 63d). Growth lines often highly pro-
er lip thin, slightly to moderately sinuate. nounced.

220 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

"
‘

35 .

“a
7

Fig.44. Lateral and marginal teeth of P. wongi: a (laterals), c (laterals and inner marginals), d (outer marginals),
USNM 857941, west spring in Birchim Canyon (bars = 6.0 um); b, Laterals, USNM 853536, Antelope Spring
(bar = 6.0 um).

VOLUME 102, NUMBER 1

Fig. 45.

221

Reproductive anatomy of P. wongi: a, Left lateral view of pallial oviduct complex, USNM 857941,

west spring in Birchim Canyon; b-—l, Dorsal penes (b, USNM 857941, west spring in Birchim Canyon; c, USNM
853530, spring S of Warren Lake; d, USNM 857981, River Springs; e, USNM 857979, Corral Springs; f, USNM
857980, Layton Spring; g, USNM 857947, French Spring; h, USNM 857949, spring at Toll House; i, USNM
853534, spring (north) at northern end of Blind Spring Valley; 3, USNM 857978, spring in Owens Gorge; k,
USNM 857942, spring alongside Lubkin Creek; 1, USNM 853523, Summit Creek). “‘c’—“‘l’? drawn to same

scale.

‘

Radular (Fig. 64) formula: (4—7)—1-(4-7)/
2(3)—2(3), 3(4)-1-4(5), 17-30, 22-33 (from
numerous populations). Central tooth typ-
ically broadly trapezoidal; basal cusps mod-
erate to large-sized, basal process moder-
ately excavated.

Epithelial pigment usually dark brown-

black on snout and sides of head/foot, with
lighter cover on central portions of latter.
Type locality. —Five Springs, Ash Mead-
ows, Nye County, Nevada.
Distribution and habitat. —Amargosa
River drainage, from Ash Meadows south
to Saratoga Spring. Snails typically common

222

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 46. Penes (ventral aspects) of P. wongi: a, USNM 857941, west spring in Birchim Canyon; b, USNM
857947, French Spring; c, USNM 853536, Antelope Spring; d, USNM 857942, spring alongside Lublin Creek;
e, USNM 853534, spring (north) at northern end of Blind Spring Valley; f, USNM 857981, River Springs; g,
USNM 857949, spring at Toll House; h, USNM 857979, Corral Spring; 1, USNM 853523, Summit Creek. “a’—

““e” and “‘g’—“‘1” drawn to same scale.

on various substrates in small springpools
and upper portions of outflows (7—-32°C,
430-10,150 micromhos/cm).

Remarks. —Populations from lower
Amargosa River drainage are clearly as-
signable to 7. variegata on basis of shell,
radular, and penial features.

Morphometrics

Discriminant analyses were performed on
each of three groups of congeners from giv-
en portions of the study area: a) P. amar-
gosae and P. micrococcus, very similar forms
with contiguous ranges; b) Pyrgulopsis fau-
na of Owens Valley (P. aardahli, P. owen-
sensis, P. perturbata, P. cf. stearnsiana, P.

wongl); and c) small-sized Tryonia spp., in-
cluding those of northern Death Valley (T.
salina, T. rowlandsi, T. margae) and Ash
Meadows (T. ericae, T. elata).

Results (Summarized in Tables 3-5) sup-
port taxonomy presented herein, insofar as
high correct classifications (81-87% [over-
all]) indicate that these snails are well-dif-
ferentiated in shell features. In the analysis
involving P. amargosae and P. micrococcus,
correct classification was 89% (40/45) and
80% (291/362) for each species, respective-
ly. Correct classification for two populations
of P. amargosae (Amargosa Gorge, Grim-
shaw Lake) from Tecopa area (where the
ranges abut) was high (83%), as was that
(92%) for two nearby populations of P. mi-

VOLUME 102, NUMBER 1

Kilometers

Fig. 47. Map showing distribution of P. wongi.

crococcus (Tecopa Hot Springs, Shoshone
Spring). Misclassified specimens of P. mi-
crococcus were concentrated in a few pop-
ulations in northern Death, Panamint, and
Saline Valleys (well separated from the range
of P. amargosae), and indicate local differ-
entiation of this relatively widespread form.

223

In this analysis, discrimination was based
largely on height of shell and aperture.
For the Owens Valley Pyrgulopsis, correct
classification was 84% overall (382/451),
and ranged from 79-87% by species. The

sole species having <80% correct classifi-

cation, P. owensensis, was most frequently

224 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 48. SEM micrographs of T. margae: a—d, Grapevine Springs, upper warm spring (a, holotype, USNM
860408, [3.52 mm]; b-d, Paratypes, USNM 857952); e-g, USNM 853555, Grapevine Springs, spring above
shack (bar = 86 um).

VOLUME 102, NUMBER 1 225

Fig. 49. Central and lateral radular teeth of T. margae: a,c, USNM 857953, Grapevine Springs, upper warm

spring (a, centrals, bar = 6.7 um; c, laterals and inner marginals, bar = 6.7 um); b, d, USNM 857996, Grapevine
Springs, spring above shack (b, centrals, bar = 6.7 um; d, laterals, bar = 6.7 um).

(10/14) misclassified as P. wongi, which nant function were related to size and width
cannot be confused with the former when of shell and aperture.

penial morphology is considered. Parame- Classification of each of the three small-
ters weighing heavily on the first discrimi- sized Tryonia spp. from northern Death

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 50. Marginal radular teeth of T. margae: a, c, USNM 857953, Grapevine Springs, upper warm spring
(a, laterals and inner marginals, bar = 10.0 um; c, outer marginal, bar = 3.8 um); b, d, USNM 857996, Grapevine
Springs, spring above shack (b, laterals and inner marginals, bar = 7.5 um; d, outer marginal, bar = 3.8 um).

Valley was 100%, while that of 7. elata and
T. ericae from Ash Meadows was 93% and
95%, respectively. Despite their general
similarity in form, 7. margae and T. elata

were well-discriminated, with only a single
specimen (of the latter) misclassified (as the
former). Parameters weighing heavily on the
first discriminant function were shell height,

VOLUME 102, NUMBER 1

0.5 mm
1.0 mm

Fig. 51. Penes (dorsal aspects) of Tryonia spp.: a,
T. margae, USNM 857952, Grapevine Springs, upper
warm spring; b, 7. rowlandsi, USNM 857953, Grape-
vine Springs, upper warm spring; c, 7. variegata, USNM
857994, spring at Grimshaw Lake; d, T. robusta, USNM
857999, Nevares Springs, upper spring; e, 7. salina,
USNM 857998, Cottonball Marsh. “a,” “b,”’ “‘d,”’ “‘e”’
drawn to same scale.

width of body whorl, and aperture size and
shape.

Discussion

Modern drainage in the Death Valley area
includes Owens, Amargosa, and Mojave
Rivers, the latter two of which are dry along
most of their courses. Although now iso-
lated from one another, these river systems
were integrated during wetter, pluvial times
during the Pleistocene as a series of lakes
overflowed along their courses to terminate
in Death Valley. This Death Valley System
(named by Miller 1943:69) was initially
proposed by Gale (1914) and Blackwelder
(1933), and has been discussed in detail by
many others (Hubbs & Miller 1948; Miller
1946, 1948; Smith 1978; Soltz & Naiman

227

Table 3.—Results of discriminant function analysis
on Pyrgulopsis amargosae and P. micrococcus. Also
given are the canonical correlation, number of speci-
mens used (by species, ordered as above), and percent
correct classification (overall).

Standardized
Variables coefficients Correlations

WH 0.365 0.554
SH 1.194 0.354
SW 0.398 —0.094
LBW =—§: 717 0.176
WBW 0.749 0.146
AL 1-928 0.055
AW —0.067 —0.108
WwW 0.176 ==§:292
D 0.798 0.536
a: —0.390 0.425
SA 0.871 0.351
C. correlation 0.475
N 45,362
% correct

classification 81

1978; Smith & Street-Perrott 1983; Minck-
ley et al. 1986; Taylor 1986). Configuration
of the System is portrayed in Fig. 65 as pre-
lude to discussion of springsnail zoogeog-
raphy.

Springsnail distributions reflect pro-
nounced local endemism of the modern fau-
na. Twenty of 22 species known from Death
Valley System area are restricted to its con-
fines. Of these, 10 are endemic to single
springs or spring complexes, and another
seven occur in relatively small portions of
single modern drainage basins. Locally en-
demic Pyrgulopsis are concentrated in Ow-
ens Valley and Ash Meadows, while endem-
ic 7ryonia occur in the latter and northern
Death Valley. Three forms are relatively
widespread in the system: P. wongi, of plu-
vial Owens River drainage (Long, Adobe,
Owens Valleys); P. micrococcus, widely dis-
tributed from Amargosa Valley west to Pan-
amint Valley; and 7. variegata, of pluvial
Amargosa River drainage (Ash Meadows,
Tecopa Basin, Saratoga Spring [southern
Death Valley]).

Endemic Pyrgulopsis of Owens Valley and

228

Kilometers

Fig. 52. Map showing distribution of 7Jryonia spp.

in Ash Meadows.

Amargosa Basin are sharply differentiated
on basis of penial morphology and represent
separate local radiations. Owens. Valley
forms, characterized in part by possession
of a ventral penial swelling bearing glan-
dular ridge (absent in forms endemic to

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

T. margae o
T. protea a
T. robusta =
-T. rowlandsi o
T. salina a
i _T. variegatae

= —

in study area. Tyronia variegata occurs at 18 localities

Amargosa drainage), include a group com-
prising P. cf. stearnsiana and similar, en-
demic forms (P. aardahli, P. owensensis, P.
perturbata); and P. wongi, which has no close
relatives in the study area. Ash Meadows
(Amargosa Basin) endemics represent at

VOLUME 102, NUMBER 1

Fig. 53. SEM micrographs of T. protea, USNM 857954, Hot Creek. Shell “a”

229

7 sj ] - -
7s
s 3
. . j
~ 7 j}°
* * ‘ >
.
7 .
- - .
_ A pe
+ afr A > -
7 4 ;
s
Ps
. 2s
-
—
* >
J 7
“

is 3.64 mm tall. “b,” “‘c”

show sculpture on protoconch of embryonic shell (bars = 60, 17.6 um).

least two lineages (both including Flumin-
icola-like representatives): snails with re-
duced or absent penial lobes and a glandular
ridge on mid-ventral penial surface (P.
erythropoma, P. crystalis, P. pisteri); and
those having a ridge on terminal portion of
moderate-large sized penial lobes (P. fair-

banksensis, P. nanus, P. isolatus). The latter
may be related to the probable sister species
pair of widespread P. micrococcus and P.
amargosae from southern Amargosa River
drainage.

Relationships among regional 7ryonia are
more speculative. Endemic 7Jryonia in

Table 4.—Results of discriminant function analysis on Owens Valley Pyrgulopsis (P. aardahli, P. owensensis,
P. perturbata, P. cf. stearnsiana, P. wongi). Also given are canonical correlations, number of specimens used
(by species, ordered as above), and percent correct classification (overall).

Standardized coefficients

Variable Function |
WH 0.580
SH Sie T/he
SW 1.037
LBW O51S
WBW 1.367
AL 2.937
AW een
WwW 0.075
D 0.285
T —0.143
SA —0.810
C. correlation 0.771
N 15, 86, 39, 25, 286
% correct

classification 84

Correlations

Function 2 Function 1 Function 2

—0.841 Se | ee (7 =—O714
0.489 —0.708 —0.534
2.483 —0.549 —0.466

2 501 —0.641 == | Ta PA

=29 —0.558 0.333

=0:550 i553 —0.424
1.822 —0.617 —0.387

=@.3f7 0.255 —0.068
0.453 OOS —0.120
0.904 =0-397 —0.042
0.600 0.085 —0.164
0.687

230 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 54. Radula of T. protea, USNM 857954, Hot Creek: a, Centrals (bar = 12 um); b, c, Laterals and inner
marginals (bars = 8.6 um); d, Outer marginals (bar = 7.5 wm).

VOLUME 102, NUMBER 1

231

Fig. 55.
(1.92 mm); b-d, Paratypes, USNM 853557.

northern Death Valley and Ash Meadows,
apart from enigmatic T. robusta, apparently
comprise a single local radiation. T. protea
from west of Death Valley (Long and [fossil;
Taylor 1985:317] Panamint Valleys) is quite
similar (in shell) to at least one represen-

SEM micrographs of 7. robusta from Nevares Springs, upper spring: a, Holotype, USNM 860411

tative from the east, 7. variegata (contrary
to Taylor 1985:317) and may not represent
a separate lineage (as implied by Taylor
1985:317).

Pluvial waters of the Death Valley System
provided aquatic continuity (and thus the

Table 5.—Results of discriminant function analysis on small-sized Tryonia (T. elata, T. ericae, T. margae,
T. rowlandsi, T. salina). Also given are canonical correlations, number of specimens used (by species, ordered

as above), and percent correct classification (overall).

Standardized coefficients

Variable Function 1
WH —0.044
SH P92
SW 0.393
LBW 0.423
WBW = |\-)5)//
AL ieols
AW 0.003
W —0.031
D 0.286
Tp —0.071
SA 0.736
C. correlation 0.994
N 30: 19: 6, 16, 15
% correct

classification 97

Correlations

Function 2 Function 1 Function 2
0.672 0.109 0.487
0.600 —OaWs 0.699
O735 —0.580 0.432

02227 —0.402 O:535

—0.908 —0:521 0.493

—0.341 —O:510 0.580
0.634 —0:622 0.405
0.081 =U. a a=) eel
0.191 0.032 —0.030
0.429 0.131 0.427
0.828 0.073 0.353
0.975

232 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 56. SEM micrographs of T. robusta, Nevares Springs, upper spring, USNM 853557: a-c, Views of apex
(bars = 75, 43, 100 um); d, Dorsal operculum (bar = 200 um).

VOLUME 102, NUMBER 1 233

Fig. 57. Radula of 7. robusta, Nevares Springs, upper spring, USNM 857999: a, Centrals (bar = 6.0 um);
b, c, Laterals and inner marginals (bars = 3.8, 5.0 um); d, Outer marginals (bar = 3.8 wm).

234 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 58. SEM micrographs of penes of T. robusta, USNM 857999, Nevares Springs, upper spring: a, Dorso-
right lateral aspect, with attachment to head on lower left (bar = 136 um); b, Distal tip of papilla (bar = 23.1
um); c, d, Distal tips of penis, showing ciliation (bars = 30, 20 um).

VOLUME 102, NUMBER 1 235

Fig. 59. SEM micrographs of 7. rowlandsi from Grapevine Springs, upper warm spring: a, Holotype, USNM
860409 (2.18 mm); b-e, Paratypes, USNM 857953 (bar = 100 um).

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 60. Radula of 7. rowlandsi, USNM 857953, Grapevine Springs, upper warm spring: a, Centrals (bar =
15 um); b, Laterals (bar = 13.6 wm); c, Inner marginal (bar = 6.0 um); d, Outer marginals (bar = 5.0 um).

means for potential exchange ofaquatic fau- ern springsnail zoogeography indicates that
na) across current drainage dividesimposed the above did not facilitate much mixing
by the region’s predominant north-south between pluvial Owens and Amargosa Riv-
trending basin and range topography. Mod- er faunas. These results support earlier con-

VOLUME 102, NUMBER 1

Fig. 61.

clusions of Taylor (in Miller 1981:58, 1985:
317-318), although his observation that
“Spillover from Lake Panamint into Death
Valley ... had no recognizable effect on
mollusc distribution” (Taylor 1985:318) is
contradicted by distribution of P. micro-
coccus (and possible close affinities between
T. protea and congeners to the east).
Taylor (1985:317) suggested that faunal
exchange between Amargosa and Owens
River drainages might have been precluded
by a saline dispersal barrier consisting of an
arm of Bouse embayment (from former Gulf
of California) extending into Death Valley.
Although this hypothesis appears plausible,
it may not need to be invoked to explain
patterns described above. Apart from ad-
ditional influences on snail distribution im-
posed by far older lacustrine episodes than
considered above (Smith 1984) and other
aspects of complex regional geologic/hydro-
logic history (Minckley et al. 1986), fun-
damental questions relating to types of hab-
itat occupied by Pleistocene springsnails and
plausibility of dispersal of such (frequently
stenotopic) organisms along chains of plu-

23ST

SEM micrographs of T. salina from Cottonball Marsh: a, Holotype, USNM 860410 (3.27 mm); b,
c, Paratypes, USNM 853556 (bar = 120 um).

vial lakes have not been thoroughly inves-
tigated (see Hershler and Minckley 1986).
Details such as size, number, and salinity
of inter-connected lakes; and length and gra-
dient of lake outlets obviously would affect
probability of snail dispersal. Further, other
modes of dispersal undoubtedly occurred
and could have produced distributions
comparable to or conflicting with those pre-
dicted by pluvial drainage models. The
presence of fauna in isolated basins recently
created or devastated by volcanic activity
(Long, Adobe Valleys, respectively), or those
that either lacked pluvial connections (Sa-
line Valley, Frenchman Flat [Cane Spring];
Hubbs and Miller 1948) or connected with
snailless areas (Deep Springs Valley, whose
pluvial lake spilled into Eureka Valley; Mil-
ler 1928) attests to importance of factors
other than late Pleistocene pluvial drainage
systems in effecting modern snail distribu-
tions in the region. A detailed zoogeograph-
ic analysis encompassing the above consid-
erations will be provided following
completion of springsnail survey of remain-
ing portions of the Death Valley System.

238 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 62. Radula of T. salina, USNM 857998, Cottonball Marsh: a, Centrals (bar = 10 um); b, Laterals and
inner marginal (bar = 7.5 wm); c, Inner and outer marginals (bar = 8.6 wm); d, Outer marginal (bar = 5.0 um).

VOLUME 102, NUMBER 1 239

Fig. 63. SEM micrographs of 7. variegata: a (2.8 mm), b, USNM 853554, Saratoga Spring; c (2.86 mm), d,
USNM 853550, Shoshone Spring (bar = 86 um).

|
|
|

passin

Fig. 64. Radula of 7. variegata: a,b, USNM 857995, Saratoga Spring (a, centrals, bar = 10.0 um; b, laterals,
bar = 7.5 wm); c, d, USNM 857993, Shoshone Spring (c, laterals and inner marginals, bar = 6.0 wm; d, outer

marginal, bar = 4.3 wm).

—

Fig. 65. Map of Pleistocene Death Valley System showing drainage relations (not necessarily contempora-
neous). Stippled areas encircled by dashed lines may not have contained lakes for significant portions of the
pluvial period. In the northeast, Lake Russell (Mono Valley) spilled into Lake Adobe (Adobe Valley), which in
turn overflowed (through a gap in Benton Range) to enter the (now mostly dry) north fork of ancestral Owens
River (Putnam 1949; Hubbs and Miller 1948; Gilbert et al. 1968). Lake Long Valley filled shortly after Long
Valley caldera was created by explosive eruption of Bishop tuff 700,000 BP (Bailey et al. 1976), and spilled into
south fork of Owens River. Outflow from Owens Lake, possibly initiated as a result of increased flow due to

240

C5

5 fe) 25 50 75
L. Long Valley

| | | | Kilometers

L. Searles

L. Mojave

to
Colorado R.

Base map from
Snyder et al. 1964
118°

capture of upper San Joaquin River drainage in Long Valley (Huber 1981, Smith et al. 1983), passed between
Sierra Nevada and Coso Range to enter and fill China and Searles Lakes. Searles Lake overflowed into Panamint
Valley, which held a large lake in its southern portion that spilled into Death Valley during several periods
between 120,000 and 20,000 BP (Gale 1914, Hooke 1972, Smith 1976, Hale 1984). To the west, the lush
wetland of Ash Meadows (that did not have a large pluvial lake; Mifflin and Wheat 1979, Hay et al. 1986)
drained at least intermittently to Amargosa River during Pliocene-Pleistocene. Further downflow the river was
ponded in mid- to late-Pleistocene by alluvial fans above Amargosa Gorge to form Lake Teopa (Sheppard and
Gude 1968, Starkey and Blackmon 1979), which spilled into southern Death Valley. Mojave River filled a series
of three pluvial lakes, with Lake Mojave overflowing toward southern Death Valley during several intervals of
late Pleistocene from > 14,500—9000 BP (Ore and Warren 1971). Overflow from Death Valley System to Colorado
River during an extreme pluvial period is conjectural, and has often been hypothesized based on distribution
of aquatic biota (Miller 1981, Taylor 1986). Hale (1984) described an old (pre-Pleistocene) fluvial channel at
Ash Hill near Ludlow, CA, that could be a product of such outflow. The channel is at 594 m elevation, which
would require a vast pluvial lake (> 300 km long and >650 m maximum depth) extending to northern Death
Valley: additional geological evidence is needed to confirm this remarkable possibility.

241

242

Acknowledgments

Fieldwork in the Death Valley area was
supported by Smithsonian Institution Re-
search Opportunity Fund (ROF), Wildlife
Conservation International of the New York
Zoological Society, and contracts awarded
by U.S. Fish and Wildlife Service, Great
Basin Complex (Order No. 14320-0182);
Bureau of Land Management, California
Desert District (CA-060-CT5); and Califor-
nia Fish and Game (C-1922). Collecting
permits were provided by National Park
Service (Death Valley National Monument)
and State of California, and access to sites
on Nevada Test Site, China Lakes Naval
Weapons Center, and Fort Irwin is also
gratefully acknowledged. Bureau of Land
Management (District offices in Ridgecrest
and Bishop), California Fish and Game
(Bishop office), and National Park Service
(Death Valley National Monument) loaned
field vehicles and provided other forms of
logistic support. Field assistance was pro-
vided by numerous individuals, especially
J. Aardahl, R. Brown, W. Cassidy, T. Ford,
D. Giuliani, D. Herbst, B. Kohfield, P.
Rowlands, and D. Wong. Heidi Wolf sorted
material and produced excellent SEM mi-
crographs. Paul Greenhall digitized shell
material. Molly Ryan drew shells and draft-
ed maps (with assistance from C. Flamer).

Literature Cited

Bailey, R. A., G. B. Dalrymple, & M. A. Lanphere.
1976. Volcanism, structure, and geochronology
of Long Valley Caldera, Mono County, Califor-
nia.— Journal of Geophysical Research 81:725-
744.

Blackwelder, E. 1933. Lake Manley: An extinct lake
of Death Valley.— Geographical Review 24:464—
471.

Blake, W. P. 1857. Geological Report.— United States
War Department Pacific Railroad Surveys 5:1-
310.

Chapman, R. E., M. G. Harasewych, & R. Hershler.
[In Preparation.] CONCH: An interactive com-
puter program for the analysis of shell coiling
parameters.

Conrad, T. A. 1855. Description of a new species of

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Melania. — Proceedings of the Academy of Nat-
ural Sciences of Philadelphia 7:269.

Gale, H. S. 1914. Salines in the Owens, Searles, and
Panamin Basins, southeastern California. —
United States Geological Survey Contributions
to Economic Geology (1913):251-323.

Gilbert, C. M., M. N. Christensen, Y. Al-Rawi, & K.
R. Lajoie. 1968. Structural and volcanic his-
tory of Mono Basin, California-Nevada. —Geo-
logical Society of America Memoir 116:275-
329.

Gould, A. A. 1855. New species of land and fresh-
water shells from western America.—Proceed-
ings of the Boston Society of Natural History 5:
127-130.

Gregg, W. O., & D. W. Taylor. 1965. Fontelicella
(Prosobranchia: Hydrobiidae), a new genus of
West American freshwater snails. — Malacologia
3:103-110.

Hale, G. R. 1984. Reassessment of the Death Valley-
Colorado River overflow hypothesis in light of
new evidence. Ph.D. dissertation (unpublished),
University of California, Berkeley, 291 pp.

Hay, R. L., R. E. Pexton, T. T. Teague, & T. K. Kyser.
1986. Spring-related carbonate rocks, Mg clays,
and associated minerals in Pliocene deposits of
the Amargosa Desert, Nevada and California. —
Geological Society of America 97:1488-1503.

Hershler, R., & W. L. Minckley. 1986. Microgeo-
graphic variation in the banded spring snail (Hy-
drobiidae: Mexipyrgus) from the Cuatro Cie-
negas Basin, Coahuila, Mexico.—Malacologia
27:357-374.

——., & D. W. Sada. 1987. Springsnails (Gastro-
poda: Hydrobiidae) of Ash Meadows, Amar-
gosa Basin, California-Nevada.— Proceedings of
the Biological Society of Washington 100:776—
843.

—, & F.G. Thompson. 1987. North American
Hydrobiidae (Gastropoda: Rissoacea): Rede-
scription and systematic relationships of Tryon-
ia Stimpson, 1865 and Pyrgulopsis Call and Pils-
bry, 1886.—Nautilus 10:25-32.

Hewett, D. F. 1954. Geology of the Natural Prov-
inces. Pp. 5—20 in R. H. Jahns, ed., Geology of
Southern California, California Division of
Mines Bulletin 170, Chapter II.

Hooke, R. L. 1972. Geomorphic evidence for Late-
Wisconsin and Holocene tectonic deformation,
Death Valley, California.— Geological Society
of America Bulletin 83:2073-2098.

Hubbs, C. L., & R. R. Miller. 1948. Correlation be-
tween fish distribution and hydrographic history
in the desert basins of western United States.
Pp. 17-166 in The Great Basin, with emphasis
on Glacial and Postglacial times. Bulletin of the
University of Utah 38, Biological Series 10.

VOLUME 102, NUMBER 1

Huber, N. K. 1981. Amount and timing of Late Ce-
nozoic uplift and tilt of the Central Sierra Ne-
vada, California—evidence from the Upper San
Joaquin River Basin. — United States Geological
Survey Professional Paper 1197:1-28.

Kohn, A. J., & A. C. Riggs. 1975. Morphometry of
the Conus shell.—Systematic Zoology 24:346-
B59:

LaBounty, J. F., & J. E. Deacon. 1972. Cyprinodon
milleri, a new species of pupfish (Family Cy-
prinodontidae) from Death Valley, Califor-
nia.—Copeia 1972:769-780.

Mifflin, M. D., & M. M. Wheat. 1979. Pluvial lakes
and estimated pluvial climates of Nevada. — Ne-
vada Bureau of Mines and Geology 94:1-57.

Miller, R.R. 1943. Cyprinodon salinus, a new species

of fish from Death Valley, California.—Copeia

1943:69-78.

1946. Correlation between fish distribution
and Pleistocene hydrography in eastern Cali-
fornia and southwestern Nevada, with a map of
the Pleistocene waters. — Journal of Geology 54:
43-53.

1948. The cyprinodont fishes of the Death
Valley System of eastern California and south-
western Nevada.— Miscellaneous Publications
of the Museum of Zoology, University of Mich-
igan 68:1-155.

1981. Coevolution of desert and pupfishes
(genus Cyprinodon) in the American Southwest.
Pp. 39-94 inR. J. Naiman and D. L. Soltz, eds.,
Fishes in North American deserts. John Wiley
and Sons, New York.

—, & E. P. Pister. 1971. Management of the
Owens Pupfish, Cyprinodon radiosus, in Mono
County, California. — Transactions of the Amer-
ican Fisheries Society 100:502-509.

Miller, W. J. 1928. Geology of Deep Spring Valley,
California.— Journal of Geology 36:510-525.

Minckley, W. L., D. A. Hendrickson, and C. E. Bond.
1986. Geography of western North American
freshwater fishes: Description and relationships
to intracontinental tectonism. Pp. 519-613 in
C. H. Hocutt and E. O. Wiley, eds., Zoogeog-
raphy of North American freshwater fishes. John
Wiley and Sons, New York.

Ore, H. T., & C. N. Warren. 1971. Late Pleistocene-
Early Holocene geomorphic history of Lake Mo-
jave, California.— Geological Society of Amer-
ica Bulletin 82:2553-2562.

Pilsbry, H. A. 1899. Catalogue of the Amnicolidae
of the Western United States. — Nautilus 12:121-
127.

Putnam, W.C. 1949. Quaternary geology of the June
Lake District, California.— Bulletin of the Geo-
logical Society of America 60:1281-1302.

Sheppard, R. A., & A. J. Gude. 1968. Distribution

243

and genesis of authigenetic silicate minerals in
tuffs of Pleistocene Lake Tecopa, Inyo County,
California.— United States Geological Survey
Professional Paper 597:1-38.

Smith, G. I. 1984. Paleohydrologic regimes in the

southwestern Great Basin, 0-3.2 my Ago, com-

pared with other long records of “global’’ cli-

mate.— Quaternary Research 22:1-17.

, V. J. Barczak, G. F. Moulton, & J. C. Liddicoat.

1983. Core KM-3, a surface-to-bedrock record

of Late Cenozoic sedimentation in Searles Val-

ley, California.— United States Geological Sur-

vey Professional Paper 1256:1-24.

—., & F. A. Street-Perrott. 1983. Pluvial lakes of
the United States. Pp. 190-212 in H. E. Wright,
ed., Late-quaternary environments of the United
States, Volume 1, The Late Pleistocene (S. C.
Porter, ed.). University of Minnesota Press,
Minneapolis.

Smith, G. R. 1978. Biogeography of intermountain
fishes. Pp. 17-42 in K. T. Harper and J. L. Re-
veal, eds., Intermountain biogeography, a sym-
posium. Great Basin Naturalist Memoir 2.

Smith, R. S. 1976. Late-Quaternary pluvial and tec-
tonic history of Panamint Valley, Inyo and San
Bernardino Counties, California. Ph.D. disser-
tation (unpublished), California Institute of
Technology, 295 p.

Snyder, C. T., G. Hardman, & F. F. Sdenek. 1964.
Pleistocene lakes in the Great Basin.— United
States Geological Survey Miscellaneous Geo-
logical Investigations Map I-416.

Soltz, D. L., & R. J. Naiman. 1978. The natural
history of native fishes in the Death Valley Sys-
tem.— Natural History Museum of Los Angeles
County, Science Series 30:1-76.

Starkey, H. C. & P. D. Blackmon. 1979. Clay min-
eralogy of Pleistocene Lake Tecopa, Inyo Coun-
ty, California.— United States Geological Sur-
vey Professional Paper 1061:1-34.

Stearns, R.E.C. 1893. Report on land and freshwater
shells collected in California and Nevada by the
Death Valley Expedition, including a few ad-
ditional species obtained by Dr. C. Hart Mer-
riam and assistants in parts of the southwestern
United States. —North American Fauna 7:269-
283.

Taylor, D. W. 1950. Three new Pyrgulopsis from the

Colorado Desert, California.— Leaflets in Mal-

acology 1:27-33.

. 1966. Summary of North American Blancan

nonmarine mollusks. — Malacologia 4:1-172.

1981. Freshwater mollusks of California: A
distributional checklist.—California Fish and

Game 67:140-163.

1985. Evolution of freshwater drainages and

molluscs in western North America. Pp. 265-

244

321 inC. J. Hocutt and A. B. Leviton, eds., Late
Cenozoic History of the Pacific Northwest.
American Association for the Advancement of
Science, San Francisco, California.

Wilkinson, L. 1986. SYSTAT: The system for sta-
tistics. SYSTAT, Inc., Evanston, IL.

Department of Invertebrate Zoology,
NHB STOP 118, National Museum of Nat-
ural History, Smithsonian Institution,
Washington, D.C. 20560.

Appendix

Collection localities, numbered as in Figs. 2-7. Data
include name of site, state, county, topographic sheet,
township and range coordinates, site elevation, and
date of visitation (for negative sites only).

1. Stream in Sage Canyon. CA: Kern; Horse Canyon,
CA (7.5), 7.0 km SW of NE corner of quadrangle, 1342
m. 2. Boulder Spring. CA: Kern; Horse Canyon, CA,
5.6 km S-SW of NE corner of quadrangle, 1251 m,
3-26-87. 3. Stream in Indian Wells Canyon. CA: Kern;
Inyokern, CA, NW '4 sec. 17, T 26S, R 38E, 1068 m,
3-26-87. 4. Spring in SW corner of Short Canyon. CA:
Kern; Inyokern, CA, NW 4 sec. 5, T 26S, R 38E, 1129
m, 3-26-87. 5. Stream in Grapevine Canyon. CA: Kem;
Inyokern, CA, center of sec. 29, T 25S, R 38E, 946 m,
3-26-87. 6. Stream in Sand Canyon. CA: Kern; Little
Lake, CA, center of sec. 7, T 25S, R 38E, 1068 m. 7.
Stream in Noname Canyon. CA: Kern; Little Lake,
CA, 10.0 km NE of SW corner of quadrangle, 976 m,
3-26-87. 8. Stream in Ninemile Canyon. CA: Inyo;
Little Lake, CA, 12.2 km NE of SW corner of quad-
rangle, 976 m, 3-26-87. 9. Spring 0.8 km §S of Little
Lake, W of HW 395. CA: Inyo; Little Lake, CA, SE
sec. 18, T 23S, R 38E, 946 m, 4-1-87. 10. Spring at
Little Lake, E of HW 395. CA: Inyo; Little Lake, CA,
NW 4 sec. 17, T 23S, R 38E, 946 m. 11. Stream in
Little Lake Canyon. CA: Inyo; Little Lake, CA, NE %
sec. 12, T 23S, R 37E, 1129 m, 4-1-87. 12. Springs ca.
1.0 km N of Little Lake Canyon. CA: Inyo; Little Lake,
CA, SW sec. 1, T 23S, R 37E, 1159 m, 4-30-87. 13.
Stream in canyon ca. 3.0 km N of Little Lake Canyon.
CA: Inyo; Little Lake, CA, 4.4 km SE of NW corner
of quadrangle, 1220 m, 4-30-87. 14. Stream in Por-
tugese Canyon. CA: Inyo; Haiwee Reservoir, CA, 2.19
km NE of SW corner of quadrangle, 1342 m, 4-1-87.
15. Springs on Portugese Bench. CA: Inyo; Haiwee
Reservoir, SW corners of secs. 3, 10, T 22S, R 37E,
1160-1220 m, 4-30-87. 16. Lower spring in Tunawee
Canyon. CA: Inyo; Haiwee Reservoir, CA, SW ' sec.
33, T 21S, R 37E, 1373 m, 4-25-87. 17. Upper spring
in Tunawee Canyon. CA: Inyo; Monache Mtn., CA,
7.0 km N-NW of SE corner of quadrangle, 1525 m,
4-30-87. 18. Haiwee Creek. CA: Inyo; Monache Min..,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

CA, 12.6 km S-SW of NE corner of quadrangle, 1586
m, 4-30-87. 19. Springs in (2) unnamed canyons N of
Haiwee Canyon. CA: Inyo; Monache Mtn., CA, NW
corners secs. 30, 31, T 20S, R 37E, 1556 m, 4-30-87.
20. Hogback Creek. CA: Inyo; Monache Mtn., CA, 7.3
km SW of NE corner of quadrangle, 1586 m. 21. Sum-
mit Creek. CA: Inyo; Monache Mtn., CA, NE 4 sec.
7, T 20S, R 37E, 1373 m. 22. Spring ca. 3.0 km N of
Summit Creek. CA: Inyo; Monache Mtn, CA, center
of sec. 6, T 20S, R 37E, 1281 m. 23. Walker Creek.
CA: Inyo; Monache Mtn., CA, NE 4% sec. 34, T 19S,
R 37E, 1769 m, 4-16-87. 24. Cartago Creek. CA: Inyo;
Olancha, CA, NE ‘4 sec. 11, T 19S, R 37E, 1159 m,
4-16-87. 25. Spring at Cabin Bar Ranch, ca. 2.5 km N
of Olancha. CA: Inyo; Olancha, CA, SW 4 sec. 6, T
19S, R 37E, 1098 m. 26. Braley Creek and springs just
to S. CA: Inyo; Olancha, CA, 10.0 km NW of SEcorner
of quadrangle, 1190 m, 4-16-87. 27. Springs on edge
of Owens Lake at Permanente. CA: Inyo; Olancha, CA,
13.8 km N of SE corner of quadrangle, 1068 m, 2-9-
85. 28. Ash Creek. CA: Inyo; Olancha, CA, 15.6 km
NW of SE corner of quadrangle, 1068 m, 4-16-87. 29.
Cottonwood Creek. CA: Inyo; Olancha, CA, 9.6 km
SW of NE corner of quadrangle, 1037 m, 4-16-87. 30.
Lower Centennial Spring. CA: Inyo; Keeler, CA, 2.3
km W of SE corner of quadrangle, 1769 m, 3-31-87.
31. Dirty Socks (Hot Spring). CA: Inyo; Keeler, CA,
NE 4 sec. 34, T 18S, R 37E, 1098 m, 2-9-85. 32.
Springs at S end of Owens Lake, ca. 3.5 km NW of
Dirty Socks. CA: Inyo; Keeler, CA, NW 4 sec. 17, T
18S, R 38E, 1098 m, 2-9-85. 33. Springs on edge of
Owens Lake, ca. 3.0 km S of Keeler. CA: Inyo; Keeler,
CA, NW 4 sec. 22, T 17S, R 38E, 1098 m, 4-25-87.
34. Lubkin Creek and spring feeding creek from south.
CA: Inyo; Lone Pine, CA, SE 4 sec. 16, T 16S, R 36E,
1220 m. 35. Spring along E side of Tuttle Creek. CA:
Inyo; Lone Pine, CA, NE 4 sec. 6, T 16S, R 36E, 1281
m. 36. Hogback Creek. CA: Inyo; Lone Pine, CA, NW
%, sec. 2, T 15S, R 35E, 1159 m. 37. Spring at NE end
of Alabama Hills, ca. 4.2 km N-NW of Lone Pine. CA:
Inyo; Lone Pine, CA, NE 4 sec. 31, T 14S, R 36E,
1159 m, 4-25-87. 38. George Creek. CA: Inyo; Lone
Pine, CA, NE 4 sec. 27, T 14S, R 35E, 1251 m, 4-25-
87. 39. Independence Creek. CA: Inyo; Independence,
CA, SE 4 sec. 23, T 13S, R 34E, 1342 m, 4-17-87. 40.
Boron Springs. CA: Inyo; Mt. Pinchot, CA, NW '% sec.
22, T 13S, R 34E, 1556 m. 41. Oak Creek, south fork.
CA: Inyo; Mt. Pinchot, CA, SW 4 sec. 10, T 13S, R
34E, 1525 m, 4-17-87. 42. Springs ca. 1.0 km W of
Mt. Whitney Fish Hatchery. CA: Inyo; Mt. Pinchot,
CA, SE 4 sec. 3, T 13S, R 34E, 1342 m, 4-18-87. 43.
Stream in Charlie Canyon. CA: Inyo; Mt. Pinchot, CA,
SW sec. 3, T 13S, R 34E, 1617 m. 44. Springs feeding
N fork Oak Creek. CA: Inyo; Mt. Pinchot, CA, SW
sec. 3, T 13S, R 34E, 1586 m, 4-25-87. 45. Oak Creek,
north fork. CA: Inyo; Mt. Pinchot, CA, center of sec.
3, T 13S, R 34E, 1525 m, 4-17-87. 46. Grover Anton
Spring. CA: Inyo; Mt. Pinchot, CA, SW 4 sec. 20, T

VOLUME 102, NUMBER 1

12S, R 34E, 1586 m, 4-18-87. 47. Harry Birch Springs.
CA: Inyo; Mt. Pinchot, CA, NW '%4 sec. 17, T 12S, R
34E, 1464 m, 4-18-87. 48. Sawmill Creek. CA: Inyo;
Mt. Pinchot, CA, SW '% sec. 9, T 12S, R 34E, 1251
m, 4-17-87. 49. Division Creek and Scotty Springs just
to N. CA: Inyo; Mt. Pinchot, CA, SW % sec. 1, T 12S,
R 33E, 1739 m, 4-17-87. 50. Goodale Creek. CA: Inyo:
Mt. Pinchot, CA, SE 4 sec. 17, T 11S, R 34E, 1251
m, 4-18-87. 51. Taboose Creek. CA: Inyo; Big Pine,
CA, SE % sec. 8, T 11S, R 34E, 1281 m, 4-18-87. 52.
Tinnemaha Creek. CA: Inyo; Big Pine, CA, NW 's sec.
28. T 10s, R 34E, 1281 m, 4-18-87. 53. Springs N of
Tinnemaha Creek and W of Poverty Hills. CA: Inyo;
Big Pine. CA, NE '% sec. 30, T 10S, R 34E, 1403 m,
4-18-87. 54. Big Pine Creek. CA: Inyo: Big Pine, CA,
SE % sec. 24, T 9S, R 33E, 1373 m, 4-18-87. 55. Baker
Creek. CA: Inyo; Big Pine, CA, NE % sec. 13, T 9S, R
33E, 1312 m, 4-18-87. 56. Big Pine Spring. CA: Inyo:
Big Pine, CA, NE 4 sec. 16, T 9S, R 33E, 1891 m,
4-18-87. 57. Spring on hill S of Warren Lake. CA: Inyo:
Big Pine, CA NE 4 sec. 16, T 9S, R 33E, 1220 m. 58.
Springs in canyon W of Warren Lake. CA: Inyo; Big
Pine, CA, SW 4 sec. 2. T 9S, R 33E, 1251 m, 4-18-
87. 59. Springs in canyon S of Shannon Canyon. CA:
Inyo; Big Pine, CA, SW 4 sec. 33, T 8S, R 33E, 1373
m. 60. Springs N of Klondike Lake. CA: Inyo: Big Pine,
CA, SW ¥s sec. 23, T 8S, R 34E, 1208 m. 5-1-87. 61.
Freeman Creek. CA: Inyo: Big Pine, CA, NE 4 sec.
20. T 8S, R 33E, 1434 m, 4-18-87. 62. Spring S of
Rawson Creek. CA: Inyo; Bishop. CA, center of sec.
6, T 8S, R 33E, 1434 m, 4-18-87. 63. Spring in Chip-
munk Canyon. CA: Inyo; Bishop, CA, SW 's sec. 28.
T 7S, R 32E, 1708 m, 4-20-87. 64. Stream in Butter-
milk Country. CA: Inyo; Mt. Tom, CA, NE 4 sec. 31,
T 7S, R 31E, 2349 m, 5-1-87. 65. Stream in McGee
Meadow. CA: Inyo: Mt. Tom, CA, NE 4 sec. 22, T
7S, R 31E, 1800 m, 4-24-87. 66. Spring in Deep Can-
yon. Mt. Tom, CA, center of sec. 12, T 7S, R 31E,
1555 m, 4-29-87. 67. Horton Creek. CA: Inyo; Mt.
Tom, CA, center of sec. 33, T 6S, R 31E, 1495 m,
4-28-87. 68. Springs N of Horton Creek. CA: Inyo; Mt.
Tom, CA, SW ‘4 sec. 32, T 6S, R 31E, 1586 m, 4-28-
87. 69. Spring along N side of upper Pine Creek. CA:
Inyo; Mt. Tom, CA, NE % sec. 26, T 6S, R 30E, 1830
m. 70. Springs entering Pine Creek in Birchim Canyon.
CA: Inyo: Mt. Tom, CA, SE '% sec. 9, T 6S, R 31E,
1373 m. 71. Spring in Wells Meadow. CA: Inyo; Mt.
Tom, CA, NW % sec. 2, T6S, R 31E, 1617 m, 4-24-
87. 72. Spring W of Sierra Paradise. CA: Mono; Mt.
Tom, CA, NW '% sec. 25, T 5S, R 30E, 1769 m, 4-24-
87. 73. Stream in Swall Meadow. CA: Mono: Casa
Diablo Mtn., CA, NW 1s sec. 14, T 5S, R 30E, 2196
m, 4-24-87. 74. Spring in Owens Gorge. CA: Inyo; Casa
Diablo Mtn., CA, SW % sec. 31, T 4S, R 31E, 1830
m. 75. Fish Slough, “BLM Spring.” CA: Mono: Bishop,
CA, SW \% sec. 30, T 5S, R 33E, 1251 m. 76. Fish
Slough, ““Northwest Springs.*”* CA: Mono: White Mtn.
Peak, CA, SE % sec. 13, T 5S, R 32E, 1281 m. 77. Fish

245

Slough, ““NE Spring.” CA: Mono; White Mtn. Peak,
CA, NW '4 sec. 18, T 5S, R 33E, 1281 m. 78. Antelope
Spring. CA: Mono; Casa Diablo Mtn., CA, SW 's sec.
9, T 4S, R 31E, 1830 m, 5-1-87. 79. Moran Spring.
CA: Mono; Casa Diablo Mtn., CA, SW '%4 sec. 29, T
3S, R 31E, 2104 m, 5-1-87. 80. Banner Springs. CA:
Mono: Casa Diablo Mtn., CA, SE % sec. 18, T 3S, R
31E, 2196 m, 5-1-87. 81. Spring by Tower Mine. CA:
Mono: Casa Diablo Mtn., CA, SE % sec. 3, T 3S, R
31E, 1952 m, 4-21-87. 82. Springs entering Marble
Creek. CA: Mono: White Mtn. Peak, CA, SE ' sec 28,
T 2S, R 32E, 1525 m. 83. Marble Creek at HW 6. CA:
Mono; White Mtn. Peak, CA, NE 4 sec. 28, T 2S, R
32E, 1525 m, 4-21-87. 84. Springs ca. 3.0 km S of
Benton, W of HW 6. CA: Mono: Benton, NV-CA, SE
Vy sec. 8, T 2S, R 32E, 1617 m, 4-21-87. 85. Benton
Hot Springs. CA: Mono: Glass Mountain, CA-NV, SW
Ys sec. 2, T 2S, R 31E, 1708 m, 4-21-87. 86. Springs
at N end of Blind Spring Valley. CA: Mono; Glass
Mountain, CA-NV, SW 14 sec. 36, T 1S, R 31E, 1708
m. 87. Springs at Bramlette Ranch. CA: Mono; Benton,
NV-CA, SW ‘4 sec. 6 (springsnail positive), SE 4 sec.
6 (negative), T 1S, R 32E, 1678 m, 4-21-87. 88. Stream
in Long John Canyon. CA: Inyo: New York Butte, CA,
14.4 km N-NE of SW corner of quadrangle, 1769 m,
5-4-87. 89. French Spring. CA: Inyo, New York Butte,
CA, 8.8 km S of NW corner of quadrangle, 1617 m.
90. Spring in Willow Springs Canyon. CA: Inyo: In-
dependence, CA, 6.5 km NW of SE comer of quad-
rangle, 1861 m, 8-18-87. 91. Barrel Springs. CA: Inyo,
Independence. CA, 13.6 km SW of NE corner of quad-
rangle, 1952 m. 92. Spring ca. 1.6 km SE of Mule
Spring. CA: Inyo; Waucoba Mtn., CA, 11.7 km N-NE
of SW corner of quadrangle. 1586 m. 93. Mule Spring.
CA: Inyo, Waucoba Mtn., CA, SE 4 sec. 1. T 10S, R
34E, 1312 m. 94. Spring at Graham Ranch. CA: Inyo,
Waucoba Mtn., CA, NE % sec. 12. T9S, R 34E, 1373
m. 95. Ulymeyer Spring. CA: Inyo; Waucoba Min.,
CA, NE ' sec. 10, T 9S, R 34E, 1235 m, 4-19-87. 96.
Wilkerson Springs. CA: Inyo, Waucoba Mtn., CA, NW
Ys sec. 10. T 9S, R 34E, 1220 m. 97. Spring at Toll
House. CA: Inyo, Waucoba Mtn., CA, NE 4 sec. 24,
T 8S, R 34E, 1861 m. 98. Warm Springs. CA: Inyo,
Bishop, CA, SW ‘4 sec. 8, T 8S, R 34E, 1220 m. 99.
Spring S of Poleta Canyon. CA: Inyo; Bishop, CA, SE
Vs sec. 18, T 7S, R 34E, 1373 m, 5-4-87. 100. Spring
S of Silver Canyon. CA: Inyo; Bishop, CA, SE % sec.
26, T6S, R 33E, 1281 m, 4-26-87. 101. Stream in Silver
Canyon. CA: Inyo; Bishop, CA, SE 4 sec. 24, T 6S, R
33E, 1434 m, 4-26-87. 102. Stream in Coldwater Can-
yon. CA: Mono; Bishop, CA, SE "% sec. 26, T 5S, R
33E, 1494 m, 5-3-87. 103. Springs on bench S of Piute
Creek. CA: Mono, Bishop, CA, SE % sec. 22, T 5S, R
33E, 1342 m. 104. Stream in canyon S of Piute Creek.
CA: Mono, Bishop, CA, NE ¥% sec. 23, T 5S, R 33E,
1617 m. 105. Spring S of Millner Creek at Copper
Queen Mine. CA: Mono; White Mtn. Peak, CA, NE
Ys sec. 21, T 4S, R 33E, 1586 m, 4-28-87. 106. Spring

246

S of Lone Tree Creek at Hill Ranch. CA: Mono; White
Mtn. Peak, CA, NE % sec. 5, T 4S, R 33E, 1617 m,
4-22-87. 107. Birch Creek. CA: Mono; White Mtn.
Peak, CA, SE % sec. 7, T 3S, R 33E, 1617 m, 4-22-
87. 108. Orchard Spring. NV: Mineral; Benton, NV-
CA, SW % sec. 35, T IN, R 32E, 2135 m, 5-10-87.
109. Spring in Queen Canyon. NV: Mineral; Benton,
NV-CA, SE % sec. 25, T 1N, R 32E, 2166 m, 5-10-
87. 110. Stream in Queen Canyon (above spring). NV:
Mineral; Benton, NV-CA, NE 4 sec. 36, T 1N, R 32E,
2288 m, 5-10-87. 111. Springs just downslope from
Long Valley Dam. CA: Mono; Casa Diablo Mtn., CA,
NW 4 sec. 20, T 4S, R 30E, 1922 m, 5-3-87. 112.
Stream on W side of Little Round Valley. CA: Mono;
Casa Diablo Mtn., CA, SE %4 sec. 36, T 4S, R 29E,
2105 m, 4-23-87. 113. McGee Creek. CA: Mono; Mt.
Morrison, CA, 7.9 km NW of SE corner of quadrangle,
2501 m, 4-23-87. 114. Stream (W of Convict Creek)
feeding Convict Lake. CA: Mono; Mt. Morrison, CA,
SW \% sec. 22, T 4S, R 28E, 2562 m, 4-27-87. 115.
Springs on S side of HW 395 W of Crowley Lake. CA:
Mono; Mt. Morrison, CA, SW 4 secs. 1 (4-23-87) and
2 (4-29-87), T 4S, R 28E, 2172 m. 116. Spring E of
Laurel Creek. CA: Mono; Mt. Morrison, CA, NE %4
sec. 9, T 4S, R 28E, 2196 m, 4-29-87. 117. Whitmore
Hot Springs. CA: Mono; Mt. Morrison, CA, NE % sec.
6, T 4S, R 29E, 2105 m, 4-29-87. 118. Springs at Hot
Creek Hatchery. CA: Mono; Mt. Morrison, CA, SW 4
sec. 35, T 3S, R 28E, 2166 m, 4-23-87. 119. Springs
at Hot Creek Ranch. CA: Mono; Mt. Morrison, CA,
SE %4 sec. 35, T 3S, R 28E, 2166 m, 4-23-87. 120. Casa
Diablo Hot Springs (cool seeps). CA: Mono; Mt. Mor-
rison, CA, NW % sec. 32, T 3S, R 28E, 2257 m, 4-23-
87. 121. Hot Creek. CA: Mono, Mt. Morrison, CA,
NE % sec. 25, T 3S, R 28E, 2196 m. 122. Layton
Springs (unnamed on topographic sheet). CA: Mono,
Casa Diablo Mtn., CA, SE % sec. 36, T 3S, R 29E,
2074 m. 123. Springs feeding Little Alkali Lake. CA:
Mono; Mt. Morrison, CA, SW 4 sec. 20, T 2S, R 29E,
2104 m, 4-29-87. 124. Little Hot Creek (source). CA:
Mono; Mt. Morrison, CA, NW 4 sec. 13, T 3S, R 28E,
2074 m, 5-6-87. 125. Deadman Creek. CA: Mono; Mt.
Morrison, CA, NE 4 sec. 27, T2S, R 27E, 2257 m,
4-29-87. 126. Big Springs. CA: Mono; Cowtrack Mtn..,
CA, NE 4 sec. 25, T 2S, R 27E, 2166 m, 4-23-87. 127.
Spring ca. 0.5 km E-NE of Arcularius Ranch. CA: Mono;
Cowtrack Mtn., CA, SE %4 sec. 20, T 2S, R 28E, 2135
m, 5-6-87. 128. Springs along lower section of Mc-
Laughlin Creek. CA: Mono; Cowtrack Mtn., CA, NE
% sec. 15, T 2S, R 28E, 2318 m, 5-6-87. 129. Mc-
Laughlin Spring. CA: Mono; Cowtrack Mtn., CA, NE
Y, sec. 12, T 2S, R 28E, 2654 m, 5-6-87. 130. Springs
at S end of Black Lake. CA: Mono; Glass Mountain,
CA-NV, SW %4 sec. 4, T 2S, R 31E, 1922 m, 4-21-87.
131. Springs at N end of Black Lake. CA: Mono; Glass
Mountain, CA-NV, SE %4 sec. 29, T 1S, R 31E, 1952
m, 4-21-87. 132. River Spring. CA: Mono, Glass
Mountain, CA-NV, NE 4 sec. 24, T 1N, R 30E, 1983

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

m. 133. Stream in Crooked Meadows. CA: Mono;
Cowtrack Mtn., CA, NW 4 sec. 36, T 1S, R 28E, 2654
m, 5-6-87. 134. Pilot Spring. CA: Mono; Cowtrack
Mtn., CA, SE % sec. 27, T 1S, R 28E, 2623 m, 5-6-
87. 135. Baxter Spring. CA: Mono; Cowtrack Mtn.,
CA, SE % sec. 1, T 1S, R 28E, 2623 m, 5-6-87. 136.
Gaspipe Spring. CA: Mono; Cowtrack Mtn., CA, NW
¥% sec. 1, T 1S, R 28E, 2440 m, 5-6-87. 137. Indian
Spring. CA: Mono; Cowtrack Mtn., CA, 13.4 km SE
of NW corner of quadrangle, 2288 m, 5-6-87. 138.
Simons Spring (and others in large marshy area). CA:
Mono; Cowtrack Mtn., CA, 6.9 km SE of NW corner
of quadrangle, 1968 m, 5-5-87. 139. Willow Spring
(not shown on topographic sheet). CA: Mono; Cow-
track Mtn., CA, NE % sec. 15, T IN, R 27E, 1952 m,
5-5-87. 140. Spring along Lee Vining Creek. CA: Mono;
Mono Craters, CA, NW 4 sec. 19, T IN, R 26E, 2227
m, 5-5-87. 141. Springs feeding lower section of Lee
Vining Creek. CA: Mono; Mono Craters, CA, NW 4
sec. 9, T IN, R 26E, 2000 m, 5-5-87. 142. Springs
along HW 395 ca. 5.0 km NW of Lee Vining. CA:
Mono; Mono Craters, CA, NE 4 sec. 31, T 2N, R 26E,
1950-2075 m, 5-6-87. 143. Lower Dechambeau Creek
and associated springs. CA: Mono; Bodie, CA, SE %4
sec. 19, T 2N, R 26E, 1922 m, 5-6-87. 144. Spring ca.
2.0 km S of Dechambeau Creek along HW 395. CA:
Mono; Bodie, CA, SE % sec. 30, T 2N, R 26E, 1922
m, 5-6-87. 145. Mill Creek. CA: Mono; Bodie, CA,
NW 4% sec. 15, T 2N, R 25E, 2379 m, 5-6-87. 146.
Stream N of Mill Creek. CA: Mono; Bodie, CA, SE
sec. 1, T 2N, R 25E, 2105 m, 5-6-87. 147. Stream in
Rattlesnake Canyon. CA: Mono; Bodie, CA, NW 4
sec. 30, T 3N, R 26E, 2288 m, 5-6-87. 148. Murphy
Spring. CA: Mono; Bodie, CA, SE % sec. 24, T 4N, R
26E, 2501 m, 5-6-87. 149. Stream S of Murphy Spring.
CA: Mono; Bodie, CA, NE 4 sec. 25, T 4N, R 26E,
2410 m, 5-6-87. 150. Stream in Cottonwood Canyon.
CA: Mono; Bodie, CA, SE % sec. 32, T 4N, R 27E,
2227 m, 5-6-87. 151. Sulphur Pond and pool to E. CA:
Mono; Trench Canyon, CA-NV, SE 4 sec. 26, T 3N,
R 27E, 1952 m, 5-6-87. 152. Saline Marsh. CA: Inyo;
New York Butte, CA, center of sec. 27, T 14S, R 28E,
335 m. 153. Stream in Hunter Canyon. CA: Inyo; New
York Butte, CA, 14.5 km SE of NW corner of quad-
rangle, 549 m, 3-28-87. 154. Stream in Beveridge Can-
yon. CA: Inyo; New York Butte, CA, 12.3 km E-SE of
NW corner of quadrangle, 580 m, 3-28-87. 155. Bad-
water Spring. CA: Inyo; Waucoba Wash, CA, 10.0 km
NE of SW corner of quadrangle, 488 m, 3-28-87. 156.
Springs S of Lower Warm Springs. CA: Inyo; Waucoba
Wash, CA, NE % sec. 30, T 13S, R 39E, 381 m, 3-29-
87. 157. Springs NW of Upper Warm Springs. CA:
Inyo; Waucoba Wash, CA, SE 4 sec. 5, T 13S, R 39E,
549 m, 3-29-87. 158. Upper Warm Spring. CA: Inyo;
Dry Mountain, CA, NE 4 sec. 9, T 13S, R 39E, 488
m, 2-11-85. 159. Willow Creek. CA: Inyo; Waucoba
Wash, CA, 12.3 km N-NE of SW corner of quadrangle,
915 m, 3-28-87. 160. Corral Springs. CA: Inyo; Blanco

VOLUME 102, NUMBER 1

Mtn., CA, SW % sec. 3, T 8S, R 36E, 1525 m. 161.
Bog Mounds Springs. CA: Inyo; Blanco Mtn., CA, SW
Ys sec. 32, T 7S, R 36E, 1508 m, 4-26-87. 162. Antelope
Spring. CA: Inyo; Blanco Mountain, CA, NW % sec.
24, T 7S, R 35E, 1708 m. 163. Samms Spring. CA:
Inyo; Blanco Mountain, CA, SW 4 sec. 13, T 78, R
35E, 1739 m, 3-29-87. 164. Willow Spring. CA: Inyo;
Magruder Mtn., NV-CA, 10.0 km N-NE of SW corner
of quadrangle, 1739 m, 3-30-87. 165. Wheelbarrow
Spring (not shown on topographic sheet). CA: Inyo;
Soldier Pass, CA-NV, NE '4 sec. 5, T 7S, R 37E, 1708
m, 3-30-87. 166. Wyman Creek. CA: Inyo; Soldier
Pass, CA-NV, SW 4 sec. 23, T 6S, R 36E, 1769 m,
3-29-87. 167. Stream in Indian Joe Canyon. CA: Inyo;
Trona, CA, NW 4 sec. 24, T 24S, R 42E, 732 m, 4-13-
87. 168. Allen Spring (not shown on topographic
sheet). CA: Inyo; Trona, CA, NE 4 sec. 24, T 24S, R
42E, 854 m, 4-13-87. 169. Stream in Great Falls Basin.
CA: Inyo; Trona, CA, SW 4 sec. 12, T 24S, R 42E,
854 m, 4-13-87. 170. Stream in Canyon N of Great
Falls Basin. CA: Inyo; Trona, CA, NW 4 sec 7, T 24S,
R 43E, 915 m, 3-25-87. 171. Stream in S fork Home-
wood Canyon. CA: Inyo; Trona, CA, NW 4 sec 2, T
24S, R 42E, 1159 m, 3-25-87. 172. Benko Spring. CA:
Inyo; Trona, CA, NE 4 sec. 34, T 23S, R 42E, 1037
m, 3-25-87. 173. Stream in N fork Homewood Can-
yon. CA: Inyo; Trona, CA, SW \4 sec. 25, T 23S, R
42E, 1098 m, 3-25-87. 174. Bircham Springs. CA: Inyo;
Trona, CA, NE 4 sec. 20, T 23S, R 42E, 1708 m, 3-
4-85. 175. Wild Rose Spring. CA: Inyo; Mountain
Springs Canyon, CA, SW 4 sec. 11, T 23S, R 41E,
1525 m, 3-4-85. 176. LaMotte Spring. CA: Inyo; Trona,
CA, NE % sec. 31, T 22S, R 42E, 1525 m, 3-4-85. 177.
Stream in N fork Water Canyon. CA: Inyo; Trona, CA,
NE 4 sec. 36, T 22S, R 42E, 1098 m, 4-11-87. 178.
Tennessee Spring. CA: Inyo; Coso Peak, CA, SW 4
sec. 13, T 21S, R 41E, 1830 m. 179. Haiwee Spring.
CA: Inyo; Haiwee Reservoir, CA, SW 4 sec. 10, T 21S,
R 39E, 1403 m, 3-5-85. 180. Stream in Knight Canyon.
CA: Inyo; Maturango Peak, CA, SE 4 sec. 14, T 21S,
R 42E, 1037 m. 181. Stream in Revenue Canyon. CA:
Inyo; Maturango Peak, CA, NW '% sec. 3, T 21S, R
42E, 976 m, 4-14-87. 182. Stream in Snow Canyon.
CA: Inyo; Maturango Peak, CA, 5.1 km S-SE of NW
corner of quadrangle, 1342 m. 183. Black Spring. CA:
Inyo; Coso Peak, CA, 1.6 km E of NW corner of quad-
rangle, 1769 m, 3-31-87. 184. Darwin Wash. CA: Inyo;
Darwin, CA, China Garden Spring, NE % sec. 4, T
19S, R 41E, 1037 m; spring above Darwin Falls, NW
% sec. 3, T 19S, R 41E, 854 m; stream below falls, SW
Ys sec. 34, T 18S, R 41E, 732 m. 185. Stream in Grape-
vine Canyon. CA: Inyo; Ubehebe Peak, CA, 8.4 km
NW of SE corner of quadrangle, 1525 m, 3-28-87. 186.
Big Dodd Spring. CA: Inyo; Ubehebe Peak, CA, 11.5
km NW of SE corner of quadrangle, 1159 m, 3-12-87.
187. Stream in Goler Canyon. CA: Inyo; Manley Peak,
CA, 15.3 km NE of SW corner of quadrangle, 549 m,
3-31-87. 188. Anvil Spring. CA: Inyo; Manley Peak,

247

CA, 11.3 km SW of NE corner of quadrangle, 1373 m,
2-18-85. 189. Seepage on SW side of Panamint Lake
bed. CA: Inyo; Manley Peak, CA, E sides of secs. 28
and 34, T 22S, R 44E, 317 m, 3-31-87. 190. Stream
in Pleasant Canyon. CA: Inyo; Telescope Peak, CA,
6.9 km NE of SW corner of quadrangle, 793 m. 191.
Post Office Spring. CA: Inyo; Telescope Peak, CA, NE
4 sec. 9, T 22S, R 44E, 317 m, 2-14-85. 192. Stream
in Happy Canyon. CA: Inyo; Telescope Peak, CA, 11.3
km NE of SW corner of quadrangle, 1098 m, 4-11-87.
193. Spring mounds on Panamint Lake bed NW of
Ballarat. CA: Inyo; Maturango Peak, CA, NW ' sec.
20, T 21S, R 44E, 323 m, 4-14-87. 194. Stream in
Surprise Canyon. CA: Inyo; Telescope Peak, CA, 15.3
km NE of SW corner of quadrangle, 1098 m. 195.
Warm Sulphur Springs. CA: Inyo; Telescope Peak, CA,
SE 4 sec. 10, T 21S, R 44E, 305 m, 2-13-85. 196.
Stream in Jail Canyon. CA: Inyo; Telescope Peak, CA,
10.0 km SE of NW corner of quadrangle, 1220 m. 197.
Stream in Tuber Canyon. CA: Inyo; Telescope Peak,
CA, 5.6 km SE of NW corner of quadrangle, 1098 m,
4-11-87. 198. Wildrose Spring. CA: Inyo; Emigrant
Canyon, CA, 3.8 km NE of SW corner of quadrangle,
1080 m, 2-15-85. 199. Springs in upper Wildrose Can-
yon. CA: Inyo; Emigrant Canyon, CA, 5.3 km NE of
SW corner of quadrangle, 1220 m, 2-15-85. 200. Cave
Spring. CA: San Bernardino; Avawatz Pass, CA, 7.7
km NE of SW corner of quadrangle, 1129 m, 4-9-87.
201. Sheep Creek Spring. CA: San Bernardino; Ava-
watz Pass, CA, 3.8 km NW of SE corner of quadrangle,
427 m, 4-10-87. 202. Salt Creek E of HW 127. CA:
San Bernardino; Avawatz Pass, CA, 14.1 km S-SW of
NE corner of quadrangle, 92 m, 3-15-87. 203. Owl
Hole Springs. CA: San Bernardino; Leach Lake, CA,
NW 1% sec. 22, T 19N, R 3E, 641 m, 3-16-85. 204.
Saratoga Spring. CA: San Bernardino; Avawatz Pass,
CA, NW 4sec. 2, T 18N, R 5E, 61 m. 205. Ibex Spring.
CA: San Bernardino; Shoshone, CA, 8.5 km E-NE of
SW corner of quadrangle, 366 m, 3-22-87. 206. Horse
Thief Springs. CA: San Bernardino; Horse Thief Springs,
CA-NV, 10.4 km E-NE of SW corner of quadrangle,
1403 m, 4-10-87. 207. Willow Spring. CA: Inyo; Te-
copa, CA, SW 4 sec. 25, T 20N, R 7E, 427 m, 3-22-
87. 208. Springs in Amargosa Gorge, S of Tecopa. CA:
Inyo; Tecopa, CA, NW '% sec. 15, T 20N, R 7E, 397
m. 209. Spring in marsh E of Grimshaw Lake. CA:
Inyo; Tecopa, CA, NE 4 sec. 9, T 20N, R 7E, 427 m.
210. Northernmost of Tecopa Hot Springs. CA: Inyo;
Tecopa, CA, NW 4 sec. 33, T 21N, R 7E, 397 m. 211.
Shoshone Spring. CA: Inyo; Shoshone, CA, NW 4 sec.
30, T 22 N, R 7E, 519 m. 212. Resting Spring. CA:
Inyo; Tecopa, CA, SE % sec. 31, T 21N, R 8E, 549 m.
213. Lost Spring (unnamed on quadrangle). CA: Inyo;
Wingate Wash, CA, 12.8 km SE of NW corner of quad-
rangle, 702 m, 3-10-87. 214. Salsberry Spring. CA:
Inyo; Shoshone, CA, 10.6 km SW of NE corner of
quadrangle, 366 m, 3-14-85. 215. Warm Spring. CA:
Inyo; Wingate Wash, CA, 7.2 km SE of NW corner of

248

quadrangle, 732 m, 2-18-85. 216. Lower Talc Mine
Spring. CA: Inyo; Bennetts Well, CA, 6.4 km E-NE of
SW corner of quadrangle, 915 m, 3-10-87. 217. Willow
Spring. CA: Inyo; Funeral Peak, CA, 7.8 km NE of SW
corner of quadrangle, 732 m, 3-20-87. 218. Hidden
Spring. CA: Inyo; Funeral Peak, CA, 15.3 km NE of
SW corner of quadrangle, 1342 m, 3-17-87. 219. Springs
at Mormon Point. CA: Inyo; Bennetts Well, CA, 6.7
km N-NW of SE corner of quadrangle, —64 m, 3-9-
85. 220. Spring (lower) in Johnson Canyon. CA: Inyo;
Telescope Peak, CA, SW 4 sec. 22, T 21S, R 46E, 1190
m. 221. Eagle Borax Spring. CA: Inyo; Bennetts Well,
CA, 13.3 km SE of NW corner of quadrangle, —79 m,
2-16-85. 222. Hanaupah Spring. CA: Inyo; Telescope
Peak, CA, 7.6 km S-SW of NE corner of quadrangle,
1281 m. 223. Badwater Spring. CA: Inyo; Bennetts
Well, CA, 2.7 km SW on NE corner of quadrangle,
—79 m, 2-16-85. 224. Tule Spring. CA: Inyo; Bennetts
Well, CA, 10.8 km E-SE of NW corner of quadrangle,
—79 m, 2-28-85. 225. Grapevine Springs. NV: Nye;
Ash Meadows, NV-CA, NE 4 sec. 2, T 19S, R SOE,
732 m, 3-14-85. 226. Blackwater Spring. CA: Inyo;
Emigrant Canyon, CA, 12.9 km S-SW of NE corner of
quadrangle, 946 m, 3-10-87. 227. Navel Spring. CA:
Inyo; Ryan, CA-NV, NW 4 sec. 13, T 26N, R 2E, 640
m, 3-17-87. 228. Travertine Springs. CA: Inyo; Fur-
nace Creek, CA, NW ‘4 sec. 25, T 27N, R 1E, 122 m.
229. Salt Springs, NW of Cow Creek. CA: Inyo; Chlo-
ride Cliff, CA-NV, SW % sec. 21, T 28N, R 1E, —79
m, 2-3-85. 230. Nevares Springs. CA: Inyo; Chloride
Cliff CA-NV, NE 4 sec. 36, T 28N, R 1E, 275 m. 231.
Cottonball Marsh. CA: Inyo; Chloride Cliff, CA-NV,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

2.8 km NE of SW corner of quadrangle, —79 m. 232.
Salt Springs, W of Cottonball Marsh. CA: Inyo; Chlo-
ride Cliff, CA-NV, 1.0 km NE of SW corner cf quad-
rangle, —73 m, 2-22-85. 233. Salt Springs S of Cow
Creek. CA: Inyo; Furnace Creek, CA, SE % sec. 3, T
27N, R 1E, 31 m, 2-5-85. 234. Stream in Cottonwood
Canyon. CA: Inyo; Marble Canyon, CA, stream 10.0
km NW of SE corner of quadrangle, 732 m, 2-15-85,
stream 11.2 km E-NE of SW corner of quadrangle,
1037 m, 3-12-87. 235. McLean Spring (Salt Creek).
CA: Inyo; Stovepipe Wells, CA, SW 4 sec. 31, T 15S,
R 46E, 37 m, 3-8-85. 236. Springs NW of Keane Won-
der Mine. CA: Inyo; Chloride Cliff, CA-NV, SE % sec.
1,T 15S, R 46E, 366 m, 2-13-85. 237. Monarch Spring.
CA: Inyo; Chloride Cliff, CA-NV, SE 4 sec. 24, T 14S,
R 46E, 915 m, 2-4-85. 238. Keane Spring. CA: Inyo;
Chloride Cliff, CA-NV, NW ‘4 sec. 8, T 30W, R 1E,
1159 m, 2-4-85. 239. Cane Spring. NV: Nye; Cane
Spring, NV, 6.0 km N-NE of SW corner of quadrangle,
1238 m. 240. Spring in Oasis Valley, 13.2 km N of
Beatty. NV: Nye County; Thirsty Canyon, NV, SE 4
sec. 32, T 10S, R 47E, 1128 m. 241. Grapevine Springs.
CA: Inyo; Ubehebe Crater, CA-NV, NE % sec. 3, T
11S, R 42E, 824 m. 242. Spring 1.6 km E of Scotty’s
Castle. CA: Inyo; Ubehebe Crater, CA-NV, NW 4 sec
5, T 11S, R 42E, 946 m. 243. Little Sand Spring. CA:
Inyo; Last Chance Range, CA-NV, SE % sec. 17, T 9S,
R 41E, 915 m, 387. 244. Sand Spring. CA: Inyo; Last
Chance Range, CA-NV, SE 4 sec. 7, T 9S, R 41E, 946
m, 3-30-87. 245. Last Chance Spring. CA: Inyo; Ma-
gruder Mtn., NV-CA, SE % sec. 2, T 8S, R 39E, 1739
m, 3-30-87.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 249-263

FOUR NEW SPECIES OF FREE-LIVING MARINE
NEMATODES IN THE GENUS PAREURYSTOMINA
(NEMATODA: ENOPLIDA) WITH OBSERVATIONS ON
OTHER MEMBERS OF THE GENUS

Edwin J. Keppner

Abstract.—Four new species of Pareurystomina and the male of P. flagelli-
caudata are described for the first time from sediments in St. Andrew Bay and
the Gulf of Mexico, Bay County, Florida, U.S.A. Additional information is
provided for P. bissonettei, P. floridensis, and P. acuminata. P. americana n.
sp. and P. vaughtae n. sp. differ from one another in the shape of the spicules
and gubernaculum, and from P. atypica in the location of the excretory pore
and shape of the spicules and gubernaculum. P. parafloridensis n. sp. differs
from P. floridensis in the number of circles of denticles in the stoma and in
the size and arrangement of the denticles. P. alima n. sp. differs from P.
parafloridensis n. sp. in the number of circles of denticles in the stoma, in the
absence of a gubernaculum, and in the shape of the spicules. It differs from P.
agubernacula in the number of circles of denticles in the stoma and the length
and shape of the spicules. The genus Megeurystomina is placed in synonymy
with Pareurystomina, and M. combesi becomes P. combesi n. comb. A key to

the species of the genus Pareurystomina is given.

The genus Pareurystomina Micoletzky &
Kreis, 1930, was erected for those species
of free-living marine nematodes of the
subfamily Eurystomininae Filipjev, 1921,
in which the spinneret and caudal glands
are absent, and the terminus of the tail is
acute. Wieser (1959) described Pareurys-
tomina pugetensis, described the cervical
setae as flattened in this species, and re-
viewed the known species in the genus. He
recognized as valid only those species for
which males had been described. He ex-
pressed doubt as to the inclusion of Pareu-
rystomina tenuicauda Stekhoven, 1950, in
the genus because circles of denticles are
absent from the stoma in this species where-
as they are present in all other species in the
genus. Luc & De Coninck (1959) described
Pareurystomina armorica and provided a
key to six species of the genus.

Chitwood (1960) described Pareurystom-
ina atypica and provided a key to the nine

species then in the genus and included P.
tenuicauda. The key used a number of char-
acters to differentiate the species including
the number of circles of denticles in the sto-
ma, tail length, length of spicules, and pres-
ence or absence of flattened cervical setae.
He also recognized the problem of the ab-
sence of male descriptions for some species.

Hopper (1963) discussed the presence of
large hypodermal gland cells in Pareurys-
tomina. Hopper (1970) described Pareurys-
tomina bissonettei, discussed the presence
of flattened cervical setae, the arrangement
of the denticles in the stoma in Pareurys-
tomina, and emended the description of P.
atypica to include flattened cervical setae.
Lambshead & Platt (1979) described a new
genus Batheurystomina, provided a tabular
key to the genera of the Eurystomininae,
and discussed each genus in the subfamily.
They considered male specimens to be crit-
ical to an understanding of the Eurystomi-

250

ninae. They considered Eurystomina re-
panda Wieser, 1959, to be a synonym of P.
pugetensis based on the presence of flat-
tened cervical setae and similarity of the
male genital apparatus. They considered all
species of Pareurystomina for which males
are unknown as species dubiae, and P. ten-
uicauda was considered species dubia due
to the absence of information pertaining to
the stoma.

Collections of free-living marine nema-
todes from various habitats in St. Andrew
Bay, and the Gulf of Mexico, Bay County,
Florida, U.S.A. from 1981-1987 yielded 174
specimens of nine species in the genus Par-
eurystomina. The purpose of this paper is
to describe the species collected, provide
some additional information for known
species and present a key to the species of
the genus Pareurystomina.

Specimens were recovered from sediment
from a number of subtidal sites in the above-
mentioned waterbodies with core samplers
of variable diameters. Nematodes were ex-
tracted from the samples by repeated wash-
ing of the sediment in water from the site.
Nematodes were removed alive from the
settled material of the final washing, fixed
in hot alcohol-formalin-acetic acid or hot
4% formalin in sea water for 24 h and de-
hydrated and mounted in anhydrous glyc-
erine on Cobb slides.

All measurements are given in um unless
otherwise stated, and the mean is followed
by the range in parentheses. The spicular
lengths are given as the chord of the arc
unless otherwise indicated. The taxonomic
hierarchy is from Gerlach & Riemann
(1974). Type specimens and specimens of
known species have been deposited in the
U.S. National Museum of Natural History
(USNM) and the Florida Nematode Col-
lection, University of Florida, Gainesville
(FNC).

Taxonomic Account

The specimens of Pareurystomina col-
lected during this study presented a variety

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

of spicular and gubernacular shapes and
sizes, a variable number of circles of den-
ticles in the stoma, and a variety of denticu-
lar shapes and arrangements. Particular em-
phasis was given to placing males collected
during this study with previously described
species in which the males are unknown.
The characters used to place males with fe-
males were the number of circles of stomatal
denticles, shape and size of the denticles in
each circle, tail length, size of amphid, and
location of the excretory pore. Comparison
of males with males of recognized species
included the above characters as well as the
shape of the male genital apparatus.

All specimens examined during this study
have the following features in common (Figs.
1-3). Body size and shape is variable within
a species as evidenced by the wide range of
lengths and wide range of the demanian “a”
values in some species. The cuticle is thin
and smooth, and the terminus of the tail is
acute. Caudal glands and spinneret were not
observed. Hypodermal gland cells are pres-
ent and arranged as described by Hopper
(1963, 1970). The gland cells are more
prominent and abundant in some species
than in others. The head is retracted in some
specimens which gives the appearance of an
anterior displacement of the amphid and
cephalic setae. The head has six small lips
each with a papilla, and a single circle of 10
(6 + 4) cephalic setae. Four of the subme-
dian cephalic setae are shorter than the oth-
er six cephalic setae. The lip region is de-
lineated from the remainder of the head by
a cephalic groove. The amphid is large, wid-
er than long, and located over the stoma.
Stoma is large with heavily cuticularized
walls and is divided into an anterior and
posterior chamber by a stomatal suture at
about midlevel. The stoma has a single,
large, right subventral tooth and a variable
number of circles of denticles. The circles
of denticles can be complete or interrupted
on the right subventral wall of the stoma as
described by Hopper (1970). The excretory
pore is anterior to the level of the cephalic

VOLUME 102, NUMBER 1

setae. Cervical setae are compressed and ap-
pear flattened with a broad base and ter-
minus. The width and length of these setae
vary somewhat among species. The female
reproductive system is amphidelphic and
the ovaries are antidromous. Males have
two spicules, and two cuticularized, cup-
shaped, pre-anal supplements. Two excep-
tions to these statements were observed
during this study; a single male of P. flori-
densis with only one pre-anal supplement
and a male of P. floridensis in which the
gubernaculum was absent. Gerlach (1952)
described a male of Pareurystomina acu-
minata (De Man, 1889) with a single pre-
anal supplement. A gubernaculum is absent
or reduced in some species of Pareurysto-
mina.

The characters of most value in differ-
entiating the species of Pareurystomina are
the shape and size of the spicules and the
presence or absence and shape of the gu-
bernaculum. The number of circles of den-
ticles in the stoma, the shape of the denticles
in each circle, tail length and position of the
excretory pore were also of value in differ-
entiating both males and females of the var-
ious species. The use of the number of cir-
cles of denticles in the stoma can be a
problem because the posterior circles are
often incomplete. Hopper (1970), in the de-
scription of P. bissonettei, noted that the
fewest rows of denticles occur on the wall
of the stoma from which the large subven-
tral tooth originates, and the largest number
of rows of denticles occurs on the stomatal
wall opposite the origin of the subventral
tooth. The presence of partial circles of den-
ticles and the location of the interruption
requires examination from lateral aspects in
order to determine accurately the maxi-
mum number of denticular circles in the
stoma. Some descriptions of Pareurysto-
mina species are not clear as to whether the
maximum or minimum number of denticu-
lar circles is stated. It is assumed, for pur-
poses of this paper, that the stated or figured
number of circles of denticles for each pre-

251

Figs. 1-3. Pareurystomina floridensis: 1, Male, head,
lateral view; 2, Entire male, lateral view; 3, Male, an-
terior pre-anal supplement, lateral view. Scale bars in
um. Legend: a = amphid. aa = anterior apophysis. cg
= cervical groove. cs = cephalic setae. d = denticles.
ep = excretory pore. es = esophagus. fs = flat cervical
setae. ga = gubernacular apophysis. hg = hypodermal
gland cell. in = intestine. lp = labial papilla. nr = nerve
ring. p = papilla. pa = posterior apophysis. ps = pre-
anal supplement. r = mm of cup. s = spicule. ss =
stomatal suture. st = subventral tooth. t = testis.

viously described species is the maximum
number.

Wieser (1959) recognized that the size and
shape of the denticles can vary between cir-
cles in Pareurystomina and used these dif-
ferences with other characters to distinguish
the species. In the specimens examined dur-
ing this study, the size and shape of the
denticles varied between circles in some
species but were of the same size and shape
within a given circle in the same species.

Pareurystomina americana, new species
Figs. 4-12

Diagnosis. —(Ten males, 3 females and 5
juveniles from St. Andrew Bay.) Body long,
slender. Four submedian cephalic setae

252

Figs. 4-12. Pareurystomina americana: 4, Male
paratype, head, lateral view; 5, Male holotype, head,
lateral view; 6, Male paratype, head, lateral view; 7,
Male holotype, posterior end, lateral view; 8, Male
holotype, pre-anal supplements, lateral view (anterior,
left; posterior, right); 9, Male holotype, right spicule
and gubernaculum, lateral view; 10, Male paratype,
right spicule and gubernaculum, lateral view; 11, Male
paratype, pre-anal supplements, lateral view (anterior,
left; posterior, right); 12, Female, posterior end, lateral
view. Scale bars in um.

about 4 length of other 6. Flat cervical setae
with narrow base present from base of sto-
ma to about 2 distance to nerve ring; one
male with additional setae for about '2 dis-
tance from nerve ring to base of esophagus.
Somatic setae not observed; caudal setae
sparse in male, not observed in female. Am-
phids over midlevel of stoma. Single com-
plete circle triangular denticles and one in-
complete circle small, round denticles
anterior to stomatal suture; single incom-
plete circle large, triangular denticles pos-
terior to suture. Excretory pore anterior to
cephalic setae. Tail conical with acute ter-
minus.

Males (n = 6): Length 4.52 mm (4.24—

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

4.94), width at midbody 35.8 (30-43). Head
diameter 27 (25-29) at level of cephalic se-
tae; cephalic setae 15 (14-17) and 5.3 (5-6)
long. Amphid 13.7 (13-16) wide. Stoma 24.2
(23-26) long, 17.5 (16-19) wide. Esophagus
0.964 mm (0.730-1.10) long; nerve ring
188.2 (179-195) from anterior end. Tail
140.2 (138-163) long, 34.2 (32-37) wide at
anus. Two midventral, cuticularized, cup-
shaped, pre-anal supplements present 119
(107-136) and 228 (208-248) anterior to
anus. Posterior apophysis of each supple-
ment larger than anterior; rim of cup striat-
ed. Two pairs subventral papillae between
anus and first pre-anal supplement; two pairs
subventral papillae between first and second
pre-anal supplements. Two pairs subventral
setae immediately anterior to anus; subven-
tral, lateral, subdorsal setae present poste-
rior to anus. Spicules gently curved 41.8
(37-45) long; tips blunt, barb absent. Gu-
bernaculum with narrow corpus surround-
ing spicules; gubernacular apophysis 14.7
(14-16) long, bent dorsally at midpoint. a
= 125.9 (114.9-147.8); b = 4.77 (3.96-6.03);
c = 30.8 (27.0-33.6).

Females (n = 3): Length 4.19 mm (3.36-
4.65), width at midbody 42.3 (42-45). Head
diameter 28.7 (28-30) at level of cephalic
setae; cephalic setae 15 (14-17) and 5.3 (5-
6) long. Amphid 12.3 (11-13) wide. Stoma
24 (22-26) long and 18.7 (18-19) wide.
Esophagus 848 (789-896) long, nerve ring
182.3 (171-200) from anterior end. Tail
186.7 (165-203) long, 33.7 (32-35) wide at
anus. Vulva 2.76 mm (2.15-—3.11) from an-
terior end. a = 98.7 (84.0-110.7); b = 4.92
(4.26—-5.31); c = 23.7 (17.5—31.2); V = 65.7%
(64-68).

Type specimens. — Holotype male, USNM
77137; paratype males, USNM 77138-
77140; allotype female, USNM 77141;
paratype female, USNM 77142. Paratype
males, FNC A-144 and A-146.

Type locality. —Holotype male from non-
vegetated sediment from the north shore of
St. Andrew Bay, Bay County, Florida 0.3
km west of the inlet to Lake Caroline

VOLUME 102, NUMBER 1

(85°41'10”W, 30°09’45’”N). Also in vegetat-
ed and nonvegetated sediment from Fresh-
water Bayou, St. Andrew Bay, Bay County,
Florida, (85°39'00”W, 30°07'30’N).

Etymology. —Named for the place of col-
lection, America.

Remarks. —Pareurystomina americana 1s
most similar to Pareurystomina atypica
Chitwood, 1960, from which it differs in the
location of the excretory pore (anterior to
cephalic setae vs. 104 from anterior end),
in the width of the amphid (11-16 vs. 7),
in the shape of the spicules (blunt tip vs.
narrow, pointed tip) and in the shape of the
gubernacular apophysis (bent dorsally vs.
bent ventrally). P. americana is also similar
to P. vaughtae described next but differs
from P. vaughtae in the shape of the spicules
(absent of barb at tip), the corpus of the
gubernaculum is not conical and the gu-
bernacular apophysis is bent dorsally rather
than straight.

Pareurystomina vaughtae, new species
Figs. 13-20

Diagnosis. —(Five males, 3 females, and
3 juveniles from mouth of Freshwater Bay-
ou and Grand Lagoon, St. Andrew Bay.)
Body long, slender. Four submedian ce-
phalic setae about 28% length of other 6.
Flat cervical setae narrow; present from base
of stoma almost to nerve ring. Somatic setae
sparse; caudal setae abundant in males,
sparse in females. Amphids over midlevel
of stoma. Single complete circle large tri-
angular denticles anterior to stomatal su-
ture, single incomplete circle large, trian-
gular denticles posterior to suture. Excretory
pore anterior to cephalic setae. Tail conical
with acute terminus.

Males (n = 5): Length 3.24 mm (2.63-
4.70), width at midbody 33.8 (30-37). Head
diameter 26.6 (26-29) at level of cephalic
setae; cephalic setae 18.1 (16-20) and 4.8
(4—5) long. Amphid 11.4 (11-12) wide. Sto-
ma 24.6 (24-27) long, 17.2 (16-18) wide.
Esophagus 713.2 (606—936) long; nerve ring
179 (168-200) from anterior end. Tail 138

253

Figs. 13-20. Pareurystomina vaughtae: 13, Male
holotype, head, ventral view; 14, Male paratype, head,
lateral view; 15, Male holotype, posterior end, lateral
view; 16, Male, anterior pre-anal supplement, lateral
view; 17, Male holotype, left spicule and gubernacu-
lum, lateral view; 18, Male holotype, anterior pre-anal
supplement, lateral view; 19, Male paratype, left spic-
ule and gubernaculum, lateral view; 20, Female, tail,
lateral view. Scale bars in um.

(126-144) long, 31.6 (29-35) wide at anus.
Two cuticularized, cup-shaped, midventral,
pre-anal supplements present 118.8 (114—-
136) and 200.4 (184-216) anterior to anus.
Posterior apophysis of each supplement
larger than the anterior; rim of cup not
striated. Two pairs subventral papillae be-
tween anus and first pre-anal supplement.
Two pairs subventral setae immediately an-
terior to anus. Spicules 41.8 (38-45) long,
gently curved, tip with small barb. Corpus
of gubernaculum 7.2 (6-8) long, conical,
surrounds distal end of spicules; gubernacu-
lar apophysis straight, 11 (8-13) long, di-
rected dorsally. Post-anal setae abundant. a
= 95.0 (80.0—127.0); b = 4.50 (3.95-5.02);
c = 23.4 (18.7-32.6).

254

Females (n = 3): Length 3.32 mm (3.07—
3.54), width at midbody 45.3 (43-48). Head
diameter 28.3 (27-29) at level of cephalic
setae; cephalic setae 19.3 (18-21), 4.7 (4-5)
long. Amphid 11.7 (1 1-13) wide. Stoma 23.3
(22-24) long, 19 (18-20) wide. Tail 166.3
(163-168) long, 34.7 (32-37) wide at anus.
Vulva 2.05 mm (1.91—2.16) from anterior
end. a = 73.4 (63.9-78.7); b = 3.83 (3.69-
4.02); c = 19.9 (18.8-21.1); V = 61.7% (61-
62).

Type specimens. — Holotype male, USNM
77143; paratype males, USNM 77144—-
77145; allotype female, USNM 77146.
Paratype male, FNC A-155; paratype fe-
male, FNC A-156.

Locality. —Holotype from sediments in a
bed of Shoalgrass (Halodule wrightii) at the
mouth of Freshwater Bayou, St. Andrew
Bay, Bay County, Florida (85°39'00’W,
30°07'30’”N). Other site, Grand Lagoon off
St. Andrew Bay, Bay County, Florida
(85°43'15”W, 30°07'50’N).

Etymology. —Named for Ms. Rosalie
Vaught, librarian. This and other studies
could not have been completed without her
voluntary dedication to obtaining the re-
quired literature.

Remarks.—Pareurystomina vaughtae is
most similar to P. americana but differs from
it in the presence of the large cone-shaped
corpus of the gubernaculum, straight rather
than bent gubernacular apophysis, and the
presence of two rather than three circles of
denticles in the stoma. P. vaughtae is also
similar to P. armorica in the presence of
two circles of denticles in the stoma. It dif-
fers from that species in that the excretory
pore is anterior to the cephalic setae rather
than at the cephalic groove, the amphid is
larger (50% of head diameter vs. 31% of
head diameter), and the tail is shorter (4.8
anal diameters vs. 6 anal diameters).

Pareurystomina parafloridensis, new species
Figs. 21-28

Diagnosis. —(Eleven males, 4 females and
6 juveniles from various sites in St. Andrew

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Bay). Body long, broad. Four submedian
cephalic setae about '4 length of other 6.
Flat cervical setae broad; present from base
of stoma almost to nerve ring. Somatic setae
and caudal setae sparse. Amphids over mid-
level of stoma. Stoma with 4 circles of den-
ticles anterior to stomatal suture, one circle
posterior to suture. Last circle anterior to
suture and circle posterior to suture incom-
plete. Small triangular denticles in anterior-
most 4 circles uniform in size within each
circle and between circles. Posterior-most
circle with larger triangular denticles. Ex-
cretory pore anterior to cephalic setae. Male
tail, short, conical with acute terminus. Fe-
male tail longer, conical then narrow to acute
terminus.

Males (n = 6): Length 4.59 mm (3.95—
5.10), width at midbody 70 (61-75). Head
diameter 37.3 (37-38) at level of cephalic
setae; cephalic setae 16.3 (15-18) and 5 (4—
6) long. Amphid 20.3 (19-21) wide. Stoma
30.2 (29-32) long, 24.6 (24-26) wide.
Esophagus 1.02 mm (0.971-1.08) long;
nerve ring 230.7 (224-242) from anterior
end. Tail 188.8 (157-222) long, 57.5 (56—-
61) wide at anus. Two cuticularized, cup-
shaped, midventral, pre-anal supplements
present 179.5 (152-208) and 295 (240-360)
anterior to anus. Posterior apophysis of each
supplement about twice size of anterior
apophysis; rim of cup striated. Two pairs
small subventral papillae immediately an-
terior to anus and another more lateral pair
posterior to anus. Spicules 78.3 (75-82) long,
gently curved then recurved at tip; tip broad
with large barb. Gubernaculum with small
corpus and narrow apophysis 34.3 (32-38)
long; lateral parts of apophysis more heavily
cuticularized; apophysis directed almost
dorsally. a = 65.3 (60.3-70.8); b = 4.51
(3.91-4.95); c = 24.5 (20.6—27.7).

Females (n = 4): Length 4.39 mm (3.63-—
4.96), width at midbody 104.8 (99-109).
Head diameter 37.5 (34—40) at level of ce-
phalic setae; cephalic setae 17 (16-18) and
5.5 (5-6) long. Amphid 18.8 (18-19) wide.
Stoma 32 (30-34) long, 27.3 (26-29) wide.

VOLUME 102, NUMBER 1

Esophagus 1.06 mm (0.846-1.12) long;
nerve ring 240.8 (213-254) from anterior
end. Tail 283.5 (258-323) long, 52 (48-54)
wide at anus. Caudal setae restricted to anal
region. Vulva 2.68 mm (2.03-3.12) from
anterior end. a = 43.7 (36.7-51.7); b = 4.15
(3.94—4.31); c = 15.5 (14.0-17.1); V = 62%
(60-64).

Type-specimens. — Male holotype, USNM
77147; male paratypes, USNM 77148-
77149; female allotype, USNM 77150; fe-
male paratype, USNM 77151. Paratype
males, FNC A-152, A-153; female para-
type, FNC A-154.

Locality. —Holotype male from sedi-
ments in a bed of shoalgrass (Halodule
wrightii) in Grand Lagoon, St. Andrew Bay,
Bay County, Florida (85°43'15’W,
30°07'50”N). Other locality, sediments in a
bed of manatee grass (Syringodium fili-
forme) at mouth of Freshwater Bayou, Bay
County, Florida (85°39’00” W, 30°07'30’N).

Etymology. —From the Greek para
meaning beside and floridensis referring to
Pareurystomina floridensis.

Remarks.—Pareurystomina paraflori-
densis 1s similar to P. floridensis in the length
and shape of the spicules and gubernaculum
in males and most characters in females. P.
parafloridensis differs from P. floridensis
primarily in the number of circles of den-
ticles in the stoma and the shape and size
of the denticles in each circle. P. paraflori-
densis has five circles of stomatal denticles
and the anterior four circles are of equal size
and shape; each denticle is a small triangular
structure. The posterior-most circle has
larger triangular denticles. In P. floridensis
there are three to four circles of denticles of
variable size and shape; the denticles of the
anterior-most circle are large elongate trian-
gules, the following two circles are small and
rounded and the posterior circle is large and
rounded. P. parafloridensis is also similar
to Pareurystomina combesi (Luc & De Con-
inck, 1959) n. comb. in the number of rows
of denticles in the stoma. It differs from P.
combesi in that the excretory pore is ante-

255

Figs. 21-28. Pareurystomina parafloridensis: 21,
Male holotype, head, lateral view; 22, Male holotype,
head, lateral view; 23, Male holotype, posterior end,
lateral view; 24, Male holotype, left spicule and gu-
bernaculum, lateral view; 25, Male holotype, pre-anal
supplements, lateral view (posterior, top right; anterior,
bottom left); 26, Female, posterior end, lateral view;
27, Male paratype, left spicule and gubernaculum, lat-
eral view; 28, Female, head, lateral view. Scale bars in
yum.

rior to the cephalic setae rather than pos-
terior to the cephalic groove and the female
tail is longer (c = 14—-17.1 vs. c = 47.8).

Pareurystomina alima, new species
Figs. 29-36

Diagnosis. —(Four males, 3 females, and
5 juveniles from mouth of Freshwater Bay-
ou and Grand Lagoon, St. Andrew Bay).
Body short, broad. Four submedian ce-
phalic setae about %4 length of other 6. Flat
cervical setae broad; extend from base of
stoma to nerve ring; somatic setae not ob-
served; caudal setae present. Amphids over
level of stomatal suture when relaxed, an-
terior to stomatal suture when contracted.
Denticles in 2 broad bands of 3 circles each;

Ol

Figs. 29-36. Pareurystomina alima: 29, Male ho-
lotype, head, right lateral view (contracted); 30, Male
holotype, posterior end, lateral view; 31, Male holotype
pre-anal supplements, lateral view (anterior, top; pos-
terior, bottom); 32, Male paratype, right spicule, lateral
view; 33, Female, tail, lateral view; 34, Male holotype,
right spicule, lateral view; 35, Female, head, lateral
view; 36, Male paratype, head, lateral view. Scale bars
in um.

first 4 circles anterior to suture complete
with small, triangular denticles; next 2 cir-
cles posterior to suture incomplete; first with
small, round denticles; second with large,
triangular denticles. Excretory pore anterior
to cephalic setae. Tail conical, terminus
acute.

Males (n = 4): Length 3.22 mm (3.05-—
3.57), width at midbody 75.5 (53-86). Head
diameter 38 (32-40) at level of cephalic se-
tae; cephalic setae 13.5 (13-14), 3.7 (3-4)
long. Amphid 16.5 (14-19) wide. Stoma 26
(24-27) long, 23.8 (21-25) wide. Esophagus
709.8 (598-792) long; nerve ring 210.5 (194—

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

219) from anterior end. Tail 168.5 (128-
184) long, 46 (38-50) wide at anus. Two
midventral, cuticularized, cup-shaped, pre-
anal supplements present 130.5 (70-160)
and 211.8 (101-272) anterior to anus. Pos-
terior apophysis of each supplement about
twice size of anterior apophysis; rim of cup
not striated. Two pairs circular, subventral,
plaque-like papillae between anus and first
pre-anal supplement. Two pairs small, sub-
ventral setae immediately anterior to anus.
Spicules straight, broad, 50 (45-54) long.
Gubernaculum absent. a = 44.0 (36.6—57.4);
b = 4.57 (4.36-5.10); c = 19.5 (17.1—23.8).

Females (n = 3): Length 4.19 mm (3.24—
4.90), width at midbody 90 (80-104). Head
diameter 42.3 (40-45) at level of cephalic
setae; cephalic setae 14.7 (14-16), 3.7 (3-4)
long. Amphid 17.3 (16-19) wide. Stoma 30.3
(27-32), 27.7 (26-30) wide. Esophagus 1.03
mm (0.843-1.15) long; nerve ring 236.7
(213-259) from anterior end. Tail 282.6
(248-328) long, 50 (48-51) wide at anus.
Caudal setae sparse. Vulva 2.46 mm (1.83-
2.94) from anterior end. a = 47.8 (31.2-
57.0); b = 4.04 (3.84-4.26); c = 14.8 (13.1-
16.3); V = 58.3% (56-60).

Type-specimens.— Holotype male, USNM
77152; paratype males, USNM 77153-
77154; allotype female, USNM 77155;
paratype female, USNM 77156. Paratype
male, FNC A-143; paratype female, FNC
A-145.

Locality.—Holotype male nonvegetated
sediments from the mouth of Freshwater
Bayou, St. Andrew Bay, Bay County, Flor-
ida (85°39'00”W, 30°07'30’N). Other site,
nonvegetated sediments in Grand Lagoon
off St. Andrew Bay, Bay County, Florida
(85°43'15”W, 30°07'50’N).

Etymology.—From the Greek halimos
meaning pertaining to the sea.

Remarks.— Pareurystomina alima is most
similar to P. parafloridensis in the stomatal
characters. It differs in the greater number
of circles of denticles (6 vs. 5), in the shape
and size of the denticles in the posterior-
most two circles, and the denticles in each

VOLUME 102, NUMBER 1

circle are smaller. P. alima males differ from
P. parafloridensis in the absence of a gu-
bernaculum and in the short straight spic-
ule. P. alima is similar to P. agubernacula
Keppner, 1986, in the absence of a guber-
naculum and the presence ofa short straight
spicule. It differs from P. agubernacula in
the greater number of circles of denticles (6
vs. 2) and shape and length of the spicules
(45-54 vs. 25-27).

Pareurystomina flagellicaudata
Stekhoven, 1946
Figs. 37-41

Diagnosis. —(One male from site in Gulf
of Mexico, Bay County, Florida.) Body very
long, slender. Four submedian cephalic se-
tae about '4 length of other 6. Flat cervical
setae short, broad; present from base of sto-
ma to about *3 distance to nerve ring. So-
matic setae and caudal setae sparse. Am-
phid just posterior to midlevel of stoma.
Denticles in 2 circles, one complete circle
large, triangular denticles anterior to suture,
one incomplete circle large, triangular den-
ticles posterior to suture. Excretory pore an-
terior to cephalic setae. Tail conical then
flagellate with acute terminus.

Male (n = 1): Length 7.83 mm, width at
midbody 48. Head diameter 30 at level of
cephalic setae; cephalic setae 22 and 7 long.
Amphid 14 wide. Stoma 29 long, 18 wide.
Esophagus 782 long; nerve ring 248 from
anterior end. Tail 600 long and 45 wide at
anus. Two cuticularized, cup-shaped, mid-
ventral supplements present 139 and 304
anterior to anus. Anterior and posterior
apophyses of each supplement about equal;
rim of cup not striated. Two pairs subven-
tral setae immediately anterior to anus; sub-
ventral and dorsolateral setae present from
anus to just anterior to anterior supplement.
Conical portion of tail with numerous small
setae. Spicules almost straight, 48 long with
blunt tip. Gubernaculum with short, weakly
cuticularized apophysis 6 long, corpus a thin
distal extension. a = 163; b = 10.0; c = 13.5

Figs. 37-41. Pareurystomina flagellicaudata: 37,
Male, anterior end, lateral view; 38, Male, posterior
end, lateral view; 39, Male, pre-anal supplements, lat-
eral view (posterior, top; anterior, bottom); 40, Male,
head, lateral view; 41, Male, right spicule and guber-
naculum, lateral view. Scale bars in wm.

Specimen. —One male, USNM 77157.

Locality.—Coarse sand and shell near
ledge in water 24 m deep in the Gulf of
Mexico, Bay County, Florida (85°49'30’"W,
30°01'30’N).

Remarks.—Pareurystomina flagellicau-
data was described from a single female
specimen. Hopper (1963) observed a spec-
imen of P. flagellicaudata from Woods Hole,
Massachusetts U.S.A., but did not mention
the sex or provide figures of the specimen.
To the writer’s knowledge, the male of this
species has not been described. The speci-
men described above is considered to be P.
flagellicaudata based upon the number of
rows of denticles in the stoma and the length
of the tail. It differs from the original de-
scription in the length of the cephalic setae
and in having distinctly flattened cervical
setae. However, the original description did
not figure or state that the cervical setae

258

Figs. 42-46. Pareurystomina acuminata: 42, Male,
head, right lateral view; 43, Male, head, left lateral
view; 44, Male, posterior end, lateral view; 45, Male,
pre-anal supplements, lateral view (posterior, left; an-
terior, right); 46, Male, right spicule and gubernaculum,
lateral view. Scale bars in pm.

were flat or not, and the position of the ex-
cretory pore was not mentioned.

Pareurystomina acuminata
(De Man, 1889)
Figs. 42—46

Diagnosis. —(one male from sediment in
St. Andrew Bay.) Body long, slender. Four
submedian cephalic setae about '3 length of
other 6. Fiat cervical setae broad, restricted
to region just posterior to stoma. Somatic
and caudal setae sparse. Amphid wide, an-
terior to stomatal suture. Two circles tri-
angular denticles present anterior to sto-
matal suture, posterior circle incomplete
with slightly smaller denticles; denticles ab-
sent posterior to suture. Excretory pore an-
terior to cephalic setae. Tail conical, tapers
rapidly to acute terminus.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Male (n = 1): Length 3.50 mm, width at
midbody 37. Head diameter at level of ce-
phalic setae 24; cephalic setae 15 and 5 long.
Amphid 13 wide. Stoma 28 long, 14 wide.
Esophagus 651 long; nerve ring 194 from
anterior end. Tail 235 long, 37 wide at anus.
Two midventral, cuticularized, cup-shaped,
pre-anal supplements present 165 and 309
anterior to anus. Anterior and posterior
apophyses of each supplement about equal.
Two pairs subventral papillae between anus
and first pre-anal supplement. Single pair
subventral setae immediately anterior to
anus. Spicules 55 (chord), 72 (arc) long,
strongly arcuate, tip with narrow velum and
small lateral barb. Corpus of gubernaculum
14 long, does not surround spicules; gu-
bernacular apophysis 16 long, straight,
broad, heavily cuticularized, directed pos-
tero-dorsally. a = 94.4; b = 5.38; c = 14.9.

Specimen. —Male, USNM 77158.

Locality.—Coarse sand and shell from
water about 9 m deep in St. Andrew Bay off
the mouth of Freshwater Bayou, Bay Coun-
ty, Florida (85°39'17”W, 30°07'45’N).

Remarks. —Gerlach (1952) described a
male of P. acuminata and figured the am-
phid as spiral in shape. Blome (1974) de-
scribed two males of P. acuminata and fig-
ured the amphid as a loose spiral. Neither
author mentioned or figured the position of
the excretory pore, and both figured the cer-
vical setae as not flattened. The specimen
described herein as P. acuminata 1s similar
in all important characters to those de-
scribed above with the exception of the shape
of the amphid and the presence of flat cer-
vical setae.

Hopper (1963) described a female nema-
tode collected at Gulf Shores, Alabama,
U.S.A. as P. acuminata. However, this
specimen has three circles of denticles in the
stoma, a long tail, an oblong amphid, and
the cephalic setae are shorter than those de-
scribed for P. acuminata. The description
and figures given by Hopper (1963) appear
to be that of P. americana n. sp. with the
exception of the absence of flat cervical se-

VOLUME 102, NUMBER 1

tae. The cervical setae in P. americana are
narrow and could be overlooked. Therefore,
P. acuminata sensu Hopper (1963) is con-
sidered to be a synonym of P. americana.

Pareurystomina bissonettei Hopper, 1970
Figs. 47-50

This species was the second most abun-
dant Pareurystomina collected (20 males,
14 females, and 9 juveniles). It was present
in nonvegetated and vegetated, shallow
water, estuarine sediments and one male was
obtained from nonvegetated sediments in
the Gulf of Mexico. Specimens agree with
the description given by Hopper (1970). The
notable exception is the male collected in
the Gulf of Mexico. This male is appreciably
larger but morphologically similar to the
males from shallow, estuarine sediments.
The body measurements are about equally
proportional to the smaller estuarine males.
A description of this male follows.

Diagnosis.—Body long, slender. Large,
vacuolated, subcuticular cells present in ad-
dition to the hypodermal gland cells. Four
submedian cephalic setae about 4 length of
other 6. Flat cervical setae short, narrow,
extend about '2 distance to nerve ring. So-
matic setae short, sparse; caudal setae nar-
row, flattened in males. Amphid wide, at
midlevel of stoma. Denticles in 5 circles;
those in 2 posterior circles slightly larger
than those in anterior circles. Tail short,
with narrow terminal spike with minute
spines.

Male (n = 1): Length 7.56 mm, width at
midbody 107. Head diameter 85 at level of
cephalic setae; cephalic setae 40 and 11 long.
Stoma 66 long, 56 wide. Amphid 35 wide.
Esophagus 1.70 mm long; nerve ring 320
from anterior end. Tail 145 long, 88 wide
at anus; caudal setae numerous. Two cuti-
cularized, cup-shaped, midventral, pre-anal
supplements present 221 and 422 anterior
to anus. Cup of each supplement with 4
arms each with striated distal terminus; arms
directed anteriorly, posteriorly, and later-
ally. Posterior apophysis of each supple-

259

Za

Figs. 47-54. Pareurystomina bissonettei: 47, Male,
head, sublateral view; 48, Male, pre-anal supplements,
lateral view (anterior, left; posterior, right); 49, Male,
posterior end, lateral view; 50, Female, tail, lateral view.
Pareurystomina floridensis: 51, Male, head, lateral view;
52, Male, left spicule and gubernaculum, lateral view.
Pareurystomina agubernacula: 53, Male, head, lateral
view; 54, Male, left spicule, lateral view. Scale bars in
um for all figs.

ment very large, anterior apophysis small.
Two pairs subventral papillae between anus
and first pre-anal supplement. Single pair
subventral setae immediately anterior to
anus. Spicules 128 long, arcuate with small
barb at tip. Corpus of gubernaculum does
not surround spicules; gubernacular apoph-
ysis 26 long, directed dorsally. a = 70.7; b
= 4.45: c = 52.1

Specimen.—One male, USNM 77165;
other males, 77159-77163; females, 77164
and 77166. Two males, FNC, A-147, A-148,
A-149: female, A-150.

Locality.—Coarse sand and shell near
ledge in water about 24 m deep, Gulf of
Mexico, Bay County, Florida (85°49'30’W,
30°01'30”N). Other specimens from various
sites in St. Andrew Bay, Florida.

260

Pareurystomina floridensis Keppner, 1986
Figs. 51-52

This species was the most abundant en-
countered during this study (33 males, 16
females, and 6 juveniles). It was recovered
only from shallow water, nonvegetated sed-
iments at a variety of sites in St. Andrew
Bay and Lake Powell, Bay County, Florida.
Examination of specimens recovered after
the original description reveals that in some
of the specimens there is an additional, par-
tial circle of very small, round denticles
present just anterior to the stomatal suture
in addition to the three circles originally
described. The apophysis of the gubernac-
ulum in the specimens examine curves
slightly ventrally at the tip. Figures are in-
cluded for comparative purposes.

Pareurystomina agubernacula
Keppner, 1986
Figs. 53-54
This species has not been recovered since
the original description, and the figures are
included for comparative purposes.

Megeurystomina combesi
Luc & De Coninck, 1959

Luc & De Coninck (1959) described a
new genus and species of Eurystomininae,
Megeurystomina combesi. They differen-
tiated the genus Megeurystomina from Par-
eurystomina on the basis of the short length
of the tail, the structure of the head partic-
ularly its ability to retract and protract, and
body size (10 mm and greater) in Megeu-
rystomina. Lambshead & Platt (1979) con-
sidered Megeurystomina to be a dubious
genus due to the absence of a description of
the male. The tail length (2 anal diameters),
as a generic character for Megeurystomina,
is contradicted by P. bissonettei that also
has a short tail (1.5—2.5 anal diameters). The
large male of P. bissonettei collected during
this study from the Gulf of Mexico is about
1.6 times larger than the average P. bisso-
nettei male collected from inshore waters.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

This reduces the importance of body size as
a generic character for Megeurystomina. The
retracted head observed in some of the spec-
imens examined during this study reduces
the importance of this as a generic character.
M. combesi, therefore, may be a large form
of a species of Pareurystomina. The struc-
ture of the head is similar to that of the
species of Pareurystomina with five or six
circles of stomatal denticles. For these rea-
sons, Megeurystomina is considered to be
a synonym of Pareurystomina and the
species Pareurystomina combesi (Luc & De
Coninck, 1959) n. comb. is included in the
key. It can be differentiated from the other
species on the basis of the number of circles
of denticles in the stoma, the position of the
excretory pore, and the tail length.

Key to Species of
Adult Pareurystomina

The taxonomic history of the genus Par-
eurystomina 1s similar to that of the genus
Eurystomina Filipjev, 1921, with regard to
the characters of importance in differen-
tiating species. Wieser (1953) stressed the
importance of the characters of the head in
distinguishing the species of Eurystomina.
Inglis (1962) stressed the importance of the
male genital apparatus, particularly the
shape of the gubernaculum, in distinguish-
ing the species of Eurystomina, and Yeates
(1967) agreed with this approach. Wieser &
Hopper (1967) recognized the value of using
a combination of characters involving the
head and male genital apparatus. As in the
Eurystomina, the identification of the
species of Pareurystomina is best achieved
when males are available for study.

In the following key, the maximum num-
ber of circles of denticles in the stoma, po-
sition of the excretory pore, and tail length
are used to differentiate those species for
which only females are known. The use of
these characters allowed all species previ-
ously described to be placed in the key with
the exception of P. parapugetensis Vitiello,
1970, described from one juvenile and P.

VOLUME 102, NUMBER 1

tenuissima (Filipjev, 1927) Filipjev 1946
described from one female. Both species
could not be placed in the key because the
position of the excretory pore is unknown.
Therefore, these two species remain species
dubiae in accordance with Lambshead &
Platt (1979). The presence or absence of flat
cevical setae was not used in the key because
their presence on the species described prior
to Hopper (1970) is uncertain.

1. Circles of denticles present in
StORDE) eee D

SiC RE eee
_..P. tenuicauda Stekhoven, 1950
2(1). One to two circles of denticles

HUAMSPOMNAS oie c. cas wyoue epee oe at 3

— Three or more circles of den-
GIGIES IM SCOMA ...5.....4. 6. 11

3(2). One circle of denticles in sto-
I ee a cP atts Oe 4 rk: 4

— Twocircles or denticles in sto-
eee ht Oe, GUA), 5

4(3). Tail 18 anal diameters long .
eee P. filicaudata Allgen, 1934

— Tail 4.5 anal diameters long
...P. typica Micoletzky & Kreis,

1930

5(3). Excretory pore posterior to
ASE OF StOMA. 42. ean: s me ne 6

— Excretory pore anterior to base
Ommstomiauxts! |. corey le aor! i

6(5). Stoma twice as long as wide;
tail 10-11 anal diameters long
(male unknown) ...........

... P. biserialis Stekhoven, 1946
— Stoma about 1.5 times as long
as wide; tail less than 10 anal
diameters long; spicules slen-
der with recurved tip .......

Se « P. atypica Chitwood, 1960
7(5). Excretory pore at cephalic
groove; amphid 31% of head
diameter; tail 6 anal diameters
long (male unknown) .......

..P. armorica Luc & De Coninck,

1959

10(9).

11(2).

12(11).

Excretory pore anterior to ce-
phalic groove and setae; am-
phid greater than 40% of head
diameter; tail less than 6 anal
diameters long .2:j/om05.:..- 8
Gubernaculum absent; spic-
miesSiraight evn. dikekrn 2.
.. P. agubernacula Keppner, 1986
Gubernaculum present; spic-
ules straight to curved ...... 9
Gubernacular apophysis re-
duced; spicules straight; tail 15
anal diameters long ........
_.. P. flagellicaudata Stekhoven,
1946
Gubernacular apophysis dis-
tinct, elongate; spicules curved;
tail less than 6 anal diameters
lomo Sethe SERIA BOR: 10
Spicules thin, strongly arcuate,
tip not recurved, with small
lateral barb; corpus of guber-
naculum elongate, does not
surround spicules; gubernac-
ular apophysis straight, direct-
ed postero-dorsally .........
.. P. acuminata (De Man, 1889)
Spicules broad, gently curved,
tip recurved, small lateral barb
absent; corpus of gubernacu-
lum cone-shaped, surrounds
spicules; gubernacular apoph-
ysis straight, directed dorsally
CHO ers Te: P. vaughtae, n. sp.
Tail short, cylindrical, abrupt-
ly narrows to terminal spike
with minute spines in male,
terminal spines absent in fe-
male P. bissonettei Hopper, 1970
Tail conical, tip without mi-
nute spines in males and fe-
males Siew Saline. Ls i?
Stoma with 5-6 circles of den-
ticles, denticles in circles an-
terior to stomatal suture small
andiof equalisized 5 &. iil. «.: 13
Stoma with 3-4 circles of den-
ticles, denticles in circles an-

262

1312):

14(13).

15(12).

16(15).

17(16).

18(17).

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

terior to stomatal suture equal
orunequaloinsizes Gijon... ES
Excretory pore posterior to ce-
phalic groove; tail short (2 anal
diameters) .. P. combesi, n. comb.
Excretory pore anterior to ce-
phalic groove and setae; tail
longer (4 anal diameters or
MOLO}e st. HU AGR ee 14
Stoma with 5 circles of denti-
cles; spicules curved, tip re-
curved with terminal barb; gu-
bernaculum present; female
cephalic setae 45% and amphid
50% of head diameter ......

Baths) etic P. parafloridensis, n. sp.
Stoma with 6 circles of denti-
cles; spicules straight, tip with-
out terminal barb; gubernac-
ulum absent; female cephalic
setae 35% and amphid 40% of
head diameter ...P. alima, n. sp.
Tail 14 anal diameters long ..

.... P. micoletzkyii Filipjev, 1946
Tail less than 10 anal diame-
ters ONS IE O=rri a0 Fee. 16
Spicules without terminal
barb; gubernacular apophysis
bent dorsally at midpoint; male
post-anal papillae absent

ae 2 eee P. americana, Nn. sp.
Spicules with terminal barb;
gubernacular apophysis not
bent dorsally at midpoint; male
post-anal papillae present or
absentrio).2) Serie hes 4 by/
Male post-anal papillae ab-
sent; spicules with broad, cup-
shaped tip with large barb

....P. floridensis Keppner, 1986
Male post-anal papillae pres-
ent; spicules with narrow tip
and:small bathtins 2no.%. .44 18
Male post-anal papillae about
4 distance from anus to tail
tip; gubernacular apophysis
short, directed dorsally .....

.. P. scillionensis Warwick, 1977

— Male post-anal papillae im-
mediately post-anal; guber-
nacular apophysis long, di-
rected postero-dorsally .....

Gitar P. pugetensis Wieser, 1959

Acknowledgments

Sincere appreciation is expressed to Dr.
W. Duane Hope, National Museum of Nat-
ural History, for his review of the manu-
script, examination of the specimens, and
continued encouragement. Appreciation is
also expressed to Dr. Armen C. Tarjan,
University of Florida, for his review of the
manuscript and his encouragement. Mr. and
Mrs. William Fable, National Marine Fish-
eries Service, collected the sediment sam-
ples from the Gulf of Mexico.

Literature Cited

Blome, D. 1974. Zur Systematik von Nematoden aus
dem Sandstrand der Nordseeinsel Sylt.—Mik-
rofauna Meeresboden 33:1-25.

Chitwood, B. G. 1960. A preliminary contribution
on the marine nemas (Adenophorea) of North-
ern California.— Transactions of the American
Microscopical Society 79:347-384.

Gerlach, S. A. 1952. Nematoden aus dem Kiusten-
grundwasser. — Abhandlungen der Akademie der
Wissenschaften und der Literature, Mainz 6:315—
S72:

Hopper, B. E. 1963. Marine nematodes from the coast

line of the Gulf of Mexico III. Additional species

from Gulf Shores, Alabama.—Canadian Jour-

nal of Zoology 4:841-863.

1970. Free-living marine nematodes from
Biscayne Bay, Florida, III. Eurystominidae:
Pareurystomina bissonettei sp. n. from Biscayne
Bay and other locations.— Proceedings of the
Helminthological Society of Washington 37:175—
178.

Inglis, W.G. 1962. Marine nematodes from Banyuls-
sur-Mer: With a review of the genus Eurysto-
mina. — Bulletin of the British Museum (Natural
History) Zoology 10:531-552.

Lambshead, P. J. D., & H. M. Platt. 1979. Batheu-
rystomina, a new genus of free-living marine
nematodes (Enchelidiidae) from the Rockall
Trough.—Cahiers de Biologie Marine 20:371-
380.

Luc, M., & L. A. P. De Coninck. 1959. Nematodes
libres marins de la region de Roscoff. — Archives

VOLUME 102, NUMBER 1

de Zoologie Expérimentale et Générale 98:103-
165.
Wieser. W. 1953. Free-living marine nematodes. I.
Enoploidea.—Acta Universitatis Lundensis,
Medica, Mathematica, Scientiae Rerum Natur-
alium (N. F. 2) 49:1-155.
. 1959. Free-living nematodes and other small
invertebrates of Puget Sound beaches. Univer-
sity of Washington Press, Seattle, Wash. 179 pp.
, & B. E. Hopper. 1967. Marine nematodes of
the east coast of North America. I. Florida.—

263

Bulletin of the Museum of Comparative Zool-
ogy, Harvard University 135:239-344.

Yeates,G. W. 1967. Studies on nematodes from dune
sands 3. Oncholaimidae, Ironidae, Alaimi-
dae.— New Zealand Journal of Science 10:299-
Va

306 Hibiscus Avenue, Panama City
Beach, Florida 32407.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 264-271

NEOTROPICAL MONOGENEA. 14. REVISION OF
CALLORHYNCHOCOTYLE SURIANO AND INCORVAIA,
1982 (HEXABOTHRIIDAE) WITH THE
DESCRIPTION OF C. AMATOI

Walter A. Boeger, Delane C. Kritsky, and Joaber Pereira, Jr.

Abstract. —The generic diagnosis of Callorhynchocotyle Suriano & Incorvaia,
1982, is emended based on new information obtained from specimens of the
type species, C. marplatensis, collected from Callorhinchus callorynchus from
the coasts of Uruguay and Argentina. Callorhynchocotyle marplatensis is re-
described; Squalonchocotyle callorhynchi is redescribed and transferred to Cal-
lorhynchocotyle; and C. amatoi is proposed for hexabothriids reported previ-
ously from Callorhinchus milii by Manter (1955) and Dillon & Hargis (1968).

All known Hexabothriidae are parasitic
on gills of elasmobranch or holocephalan
fishes (Class Chondrichthyes). Approxi-
mately 60 species are described from elas-
mobranchs; two are known from holo-
cephalans. Manter (1955) described
Squalonchocotyle callorhynchi from the
holocephalans, Callorhinchus capensis Du-
meril and C. milii Bory, from marine waters
off South Africa and New Zealand, respec-
tively. Dillon & Hargis (1968) redescribed
this species as Erpocotyle callorhynchi
(Manter, 1955) Yamaguti, 1963, based on
specimens from C. milii in New Zealand
waters. Lebedev & Parukhin (1969) report-
ed the parasite under the latter name from
C. capensis in Wallfish Bay (southwestern
Africa); and in 1970, Kuznetsova recorded
S. callorhynchi from Callorhinchus antarc-
ticus Fleming (=C. callorynchus) from the
Patagonian Shelf off South America. Suri-
ano and Incorvaia (1982) proposed Callo-
rhynchocotyle (Hexabothriidae) for their new
species, C. marplatensis, from Callorhin-
chus callorynchus (Linnaeus) off Argentina
(38°S, 57°W). In the present study, Callo-
rhynchocotyle Suriano & Incorvaia, 1982,
is revised; C. marplatensis is redescribed;
S. callorhynchi is redescribed and trans-
ferred to Callorhynchocotyle; and C. ama-
tol, N. sp., is proposed for hexabothriids re-

ported from Callorhinchus milli by Manter
(1955) and Dillon & Hargis (1968).

Materials and Methods

Hosts (Callorhinchus callorynchus) were
collected by trawl off the coasts of Uruguay
and Argentina during May 1985 and June
1986. Methods of parasite collection, prep-
aration, measurement, and illustration are
as described by Kritsky et al. (1986). Mea-
surements are in micrometers; averages are
followed by ranges in parentheses. Mea-
surements of curved structures represent
straight-line distances between extreme
points; see Fig. 4 for those obtained from
each sclerite. Vouchers (specimens collected
during the present study) are deposited in
helminthological collections of the National
Museum of Natural History (USNM), which
is housed at the USDA in Beltsville, Mary-
land; the University of Nebraska State Mu-
seum (HWML), Lincoln, Nebraska; the
Fundacao Instituto Oswaldo Cruz (FIOQ),
Rio de Janeiro, Brazil; and the Museo de
La Plata (MP), La Plata, Argentina.

Callorhynchocotyle
Suriano & Incorvaia, 1982

Emended diagnosis. —Hexabothriidae.
Body elongate. Tegument thin, smooth.

VOLUME 102, NUMBER 1

Caeca confluent in peduncle posterior to
testes. Testes numerous, irregular; vas def-
erens sinuous, surrounded by small gland
cells along most of its length, with small
loop proximal to entrance into cirrus. Cirrus
comprising two parts: distal part ovate, bul-
bous; proximal part a thick-walled nonex-
panded tube. Genital pore at level of gut
bifurcation. Ovary lobate anteriorly, coiled
posteriorly; oviduct originating from pos-
terior end of ovary; seminal receptacle sac-
like, reduced; ootype smooth; uterus dorsal
to ovary, ventral to vas deferens. Vagina
parallel, comprising two portions; distal
portion glandular, proximal portion deli-
cate. Vaginal pores on ventral surface about
midway between body midline and lateral
margin on each side of cirrus. Vitellaria
comprising 2 bilateral bands extending from
level of vaginal pores into peduncle; vitel-
line commissure, genitointestinal canal dor-
sal to ovary. Eggs with two polar filaments,
forming continuous chain by united fila-
ments. Excretory pores marginai at level of
genital pore. Haptor asymmetrical; with 2
small, 4 large sucker-sclerite complexes;
small complexes lying on same side of lon-
gitudinal axis of haptor; appendix originat-
ing from dorsal haptoral surface lateral to
body midline, armed with 2 anchors, 2 ter-
minal suckers. Parasites of Callorhinchidae.

Type species.—Callorhynchocotyle mar-
platensis Suriano and Incorvaia, 1982, from
Callorhinchus callorynchus.

Other species.—Callorhynchocotyle cal-
lorhynchi (Manter, 1955), n. comb., from
Callorhinchus capensis; Callorhynchocotyle
amatoi, n. sp., from Callorhinchus milii.

Callorhynchocotyle marplatensis
Suriano & Incorvaia, 1982
Figs. 1-9, 10a, lla, 12a

Synonyms.—Callorhynchocotyle callo-
rhynchi Suriano & Incorvaia, 1982; C. cal-
lorhynchy Suriano & Incorvaia, 1982.

Host.—Callorhinchus callorynchus (Lin-
naeus).

265

Distributions. —Coasts of Uruguay, Ar-
gentina (holotype).

Specimens studied.—Holotype, MP 12;
17 vouchers, USNM 80279, HWML 20705,
20706, FIOC 32.442, MP 1613 D.

Description (measurements in Table 1).—
Weak genital sucker, 94—110 testes. Papillae
lacking on rim and inner wall of oral, hap-
toral suckers. Sucker sclerite 2, 3 with elon-
gate point, evenly curved shaft and point.
Anchor with wide base, short roots, medi-
ally curved shaft, short point.

Remarks.—Suriano & Incorvaia (1982)
treat this species under three names: C.
marplatensis, C. callorhynchi, and C. cal-
lorhynchy. The latter two apparently rep-
resent typographic and/or editing errors. It
iS apparent that these authors were not re-
ferring to other species of Callorhynchoco-
tyle since they consider the genus monotyp-
ic. Clearly, the authors intended the name
of their new species to be C. marplatensis
which is considered herein as the valid name
of the taxon.

Based on the holotype and voucher spec-
imens, it is evident that Suriano & Incorvaia
(1982) confused polarity of the dorsoventral
axis in their specimens. This is supported
by their illustration of the holotype pre-
sented as a dorsal rather than a ventral view
as the specimen is mounted. Consequently,
the sinistrodextral orientations of features
of the parasite are reversed in the original
description. Also contrary to the original
description, the haptoral appendix origi-
nates from the dorsal surface of the haptor,
and the genitointestinal canal, uterus and
vitelline commissure are dorsal to the ovary.

The holotype is a damaged, excessively
flattened, and contracted specimen. One of
the haptoral suckers and its sclerite are torn
away (not shown in fig. 1 of Suriano & In-
corvaia 1982). Apparently, these original
authors added the missing sucker to their
drawing of the holotype based on the pres-
ence of six suckers in other specimens of
their collection. This, however, resulted in
their apparent confusion of a damaged mus-

266

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Measurements (in micrometers) of Callorhynchocotyle species.

Body length
Body width
Haptor
Length
Width
Oral sucker
Width

Pharynx
Length
Width
Genital sucker
Diameter
Egg
Length
Width
Ovary
Length

Distal vagina

Appendix
Length
Width
Appendix sucker
Length
Width
Anchor
Length

Base width

Diameter of haptoral sucker

Pair |
Pair 2
Pair 3
Sclerite #1
Total
Shaft
Point
Sclerite #2
Total
Shaft
Point
Sclerite #3

Total
Shaft
Point

C. callorhynchi
(n = 2)

6052 (4995-7109)
845 (669-1021)

2007

284 (248-322)

102
64 (56-72)

152 (136-168)
58 (53-62)

639 (459-818)
276

2104 (1844-2363)
997 (972-1021)

289
109

194 (186-203)

274 (258-290)
362
319

283 (281-285)
31-32
42 (39-45)

362
88
106

35
84 (83-86)
123-124

C. marplatensis
(n = 17)

9989 (7200-12,430)
939 (649-1217)

2719 (2051-3324)
1596 (1331-1918)

302 (231-362)

71 (58-83)
71 (60-78)

205 (153-262)

145 (125-193)
59 (52-66)

1147 (816-1505)
180 (136-203)

1397 (1275-1637)
256 (149-350)

181 (164-204)
195 (154-190)

190 (144-210)
87 (72-98)

305 (243-346)
365 (270-419)
345 (267-393)

353 (289-385)
33 (27-44)
50 (42-57)

460 (362-499)
76 (59-92)
117 (95-141)

454 (382-512)
80 (65-92)
129 (99-145)

C. amatoi
(n = 8)

7091 (5329-8474)
989 (706-1316)

1959 (1451-2482)
1201 (755-1639)

252 (191-356)

79 (69-91)
70 (62-89)

156 (144-169)
54 (48-60)

859 (651-971)
350 (269-432)

1108 (674-1429)
850 (541-1206)

272 (236-320)
144 (102-185)

182 (172-195)
85 (83-87)

267 (188-362)
332 (232-435)
310 (217-415)

297 (239-355)
20 (18-22)
37 (36-39)

412 (347-452)
54 (46-70)
83 (70-98)

408 (332-469)
56 (45-66)
92 (86-98)

VOLUME 102, NUMBER 1 267

002

0002

5 =
6

Figs. 1-6. Callorhynchocotyle marplatensis: 1, Whole mount (ventral); 2, Sucker-sclerite complex; 3, Sclerite
of complex 1; 4, Sclerite of complex 2; 5, Sclerite of complex 3; 6, Anchor. All figures are reproduced to respective
scales except Figs. 3, 4, 5 are drawn to the 250-micrometer scale. Circled numbers identify sucker-sclerite pairs

in the haptor. Fig. 4 shows measurements taken from each sclerite (Table 1): a = total length; b = shaft length;
c = point length.

cle as a haptoral extension of the gut. In the the esophagus has lateral diverticulae, the
holotype the gut is not visible in the haptor, caeca are confluent posterior to the testes,
although the structure depicted as gut (the and the intestine extends into the haptor

muscle which apparently served the lost with branches, one of which occurs in the
sucker) is evident. Our specimens show that haptoral appendix.

268 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 7-9.

Callorhynchocotyle marplatensis: 7, Ventral view of anterior end; 8, Egg chain from uterus; 9,

Female reproductive system (ventral). Figures are to respective 200-micrometer scales.

Callorhynchocotyle amatoi, new species
Figs. 10c, llc, 12c

Synonyms.—Squalonchocotyle_ callo-
rhynchi Manter, 1955 (part); Erpocotyle
callorhynchi (Manter, 1955) Yamaguti, 1963
(part).

Host.—Callorhinchus milii Bory.

Type locality. —Coast of New Zealand.

Specimens studied.—Holotype, USNM
37448; seven paratypes, USNM 71197,
HWML 1442.

Description (measurements in Table 1).—
Genital sucker absent; 94-120 testes. Pa-
pillae present on rim and inner wall of oral,
haptoral suckers. Sucker sclerite 2, 3 with
short point; indistinct angle formed at union
of shaft and point of sclerite 3; sclerite 1
with delicate point, short shaft. Anchor with
wide base, short roots, medially curved shaft,
short point.

Etymology. —This species is named for
Dr. Jose Felipe R. Amato, Universidade
Federal Rural do Rio de Janeiro, in recog-
nition of his contributions in marine par-
asitology.

Remarks. —Our study of the type speci-
mens of Squalonchocotyle callorhynchi

Manter, 1955, and specimens identified as
Erpocotyle callorhynchi (Manter, 1955) by
Dillon & Hargis (1968) revealed that those
from Callorhinchus milii from New Zea-
land represent a species distinct from that
occurring on C. capensis. Although Dillon
& Hargis (1968) recognized the morpholog-
ic differences between these specimens, they
did not question their conspecificity.

Since the holotype of Squalonchocotyle
callorhynchi is a specimen collected from
the gills of Callorhinchus capensis captured
off South Africa, Callorhynchocotyle amatoi
is proposed for those specimens from Cal-
lorhinchus milii from New Zealand. Cal-
lorhynchocotyle amatoi differs from C. cal-
lorhynchi and C. marplatensis by having
shorter shafts and points of all sucker scler-
ites (Fig. 13) and an indistinct angle between
the point and shaft of sclerite 3 (Fig. 12c).

Callorhynchocotyle callorhynchi
(Manter, 1955), new combination
Figs. 10b, 11b, 12b

Synonyms.—Squalonchocotyle callo-
rhynchi Manter, 1955 (part); Erpocotyle cal-

VOLUME 102, NUMBER 1

269

CA
\ i,
: \ | } 4
| \ WOE +
: a. | | Soa /
Se" | pny b
| ( /
b/ / shy Se / —e
Ne i Se
Se

Figs. 10-12. Sclerites of Callorhynchocotyle spp.: 10, Sclerite of complex 1; 11, Sclerite of complex 2; 12,
Sclerite of complex 3. a = C. marplatensis; b = C. callorhynchi; c = C. amatoi. All figures are drawn to the

same scale.

lorhynchi (Manter, 1955) Yamaguti, 1963
(part).

Host.—Callorhinchus capensis Dumeril.

Type locality. —Coast of South Africa.

Specimens studied. —Holotype, paratype,
USNM 37447.

Description (measurements in Table 1).—
Genital sucker absent, 96—98 testes. Pa-
pillae present on rim and inner wall of oral,
haptoral suckers. Sucker sclerite 1, 2, 3 with
short point, moderate shaft, evenly curved
shaft and point. Anchor with wide base,
short roots, medially curved shaft, short
point.

Remarks.—Callorhynchocotyle callo-
rhynchi is most similar to C. marplatensis,
from which it differs by possessing papillae
on the rim and inner walls of the oral and
haptoral suckers, a comparatively long dis-
tal portion of the vaginae, absence of a gen-
ital sucker, and by the comparative mor-
phology of the shaft and point of the sucker
sclerites (Figs. 10-12).

Discussion

Callorhynchocotyle was proposed by
Suriano & Incorvaia (1982) to accommo-
date C. marplatensis. Among characters
used to define the genus was the presence
of marginal vaginal pores. Examination of
the holotype (MP 12) of C. marplatensis and
vouchers collected during the present study

shows that the vaginae actually open sub-
marginally as they do in members of all
other hexabothriid genera. Apparently Suri-
ano and Incorvaia (1982) confused the lat-
eral excretory pores as the vaginal apertures.

Other characters used by Suriano and In-
corvaia (1982) for diagnosis of the genus
include: 1) an asymmetric haptor with 6
suckers armed with sclerites of different
sizes, 2) a sac-like seminal receptacle, 3) par-
allel bilateral vaginae, 4) an unarmed cirrus,
5) a smooth ootype, and 6) the absence of
a glandular region of the cirrus. Of these,
only the absence of a glandular region of the
cirrus is unique for the genus. In Callo-

= 160

3 140

~~"

I= 120

ne

© 100

QO,

+4 80

O

1 ‘60 8 06 o e C. marplatensis

=) 4 « C. amatoi

a 40 ¢ © C. callorhynchi

4 20 ,
15 30 45 60 45) 90 105

Length of shaft (um) 13

Fig. 13. Scatter diagram of length of point versus
length of shaft of the sclerite for three species of Ca/lo-
rhynchocotyle. The left column of symbols of the leg-
end refers to sclerite 1 of respective species; the second
column refers to sclerites 2, 3 of the respective species.

270

rhynchocotyle, the glandular region of the
cirrus is apparently replaced by an extensive
glandular segment of the vas deferens. These
characters may represent an example of
compensatory change (evolution) as defined
by Brooks and Wiley (1986).

Other hexabothriid genera with asym-
metric haptors include Heteronchocotyle
Brooks, 1934, Paraheteronchocotyle Mayes,
Brooks and Thorson, 1981, Epicotyle Euzet
& Maillard, 1974, Rhinobatonchocotyle
Doran, 1953, and Neonchocotyle Ktari &
Maillard, 1972. The haptor of Callorhyn-
chocotyle most closely resembles that of
Neonchocotyle in that the asymmetry re-
sults from the apparent migration of one
sucker-sclerite complex to the opposite side
of the haptor. Further, the appendix does
not originate from the midline which adds
to haptoral asymmetry.

In addition to the above, Callorhyncho-
cotyle is distinguished from other hexa-
bothriid genera by the combined presence
of the following characters: presence of a
bulbous distal cirrus, a dorsal instead of
terminal haptoral appendix, and a terminal
glandulomuscular portion of the vagina.

Hargis (1955) reported that the genitoin-
testinal canal of Heteronchocotyle leucas
Hargis, 1955 (Hexabothriidae), apparently
joined the left intestinal caecum and sug-
gested that this condition was rare among
the higher Monogenea. In individual spec-
imens of Callorhynchocotyle, the genitoin-
testinal canal was found to be united to the
intestinal caecum which corresponded to the
respective position of the ovary, 1.e., if the
Ovary occurred to the right of the body mid-
line (Fig. 9) the genitointestinal canal united
with the right caecum and conversely with
the left caecum if the ovary occurred to the
left of the body midline (Fig. 1). Similarly,
the haptoral appendix and the migrated
sucker-sclerite complex occur either to the
left or right of the longitudinal haptoral axis.
However, there was no correlation between
the asymmetry of the haptor and the loca-
tion of the ovary. The sinistral or dextral

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

position of the ovary and genitointestinal
canal and the position of the appendix and
sucker-sclerite in the haptor are examples
of intraspecific variation and do not con-
stitute generic characteristics for the Hexa-
bothrudae.

Acknowledgments

The authors wish to thank Drs. O. M.
Blanco (MP), J. R. Lichtenfels (USDA), and
M. Pritchard (HWML) for allowing us to
examine specimens in their care. The Con-
selho Nacional de Desenvolvimento Cien-
tifico e Tecnologico provided a study grant
(20.0115/84) to WAB.

Literature Cited

Brooks, D. R., & E. O. Wiley. 1986. Evolution as
entropy. Toward a unified theory of biology.
University of Chicago Press, Chicago, Illinois.
335 pp.

Dillon, W. A., & W. J. Hargis. 1968. Monogenetic
trematodes from the southern Pacific Ocean. Part
IV. Polyopisthocotyleids from New Zealand
fishes: The families Mazocraeidae, Diclidopho-
ridae and Hexabothriidae. — Proceedings of the
Biological Society of Washington 81:351-366.

Hargis, W. J., Jr. 1955. Monogenetic trematodes of
Gulf of Mexico fishes. Part VI. The superfam-
ilies Polystomatoidea Price, 1936 and Dicli-
dophoroidea Price, 1936.—Transactions of the
American Microscopical Society 74:361-377.

Kritsky, D. C., V. E. Thatcher & W. A. Boeger. 1986.
Neotropical Monogenea.. 8. Revision of Uro-
cleidoides (Dactylogyridae, Ancyrocephali-
nae). — Proceedings of the Helminthological So-
ciety of Washington 53:1-37.

Kuznetsova, I. G. 1970. Parasitofauna of cartilagi-
nous fishes of the Patagonian Shelf. Pp. 52-54
in Issues on marine parasitology, materials from
the Ist annual All-Union Symposium on Par-
asites and Diseases of Marine Life, Sevastopol.
Publ. Naukova Dumka, Kiev.

Lebedev, B. I., & A. M. Parukhin. 1969. Monogenea
of some fish from Wallfish Bay (south-western
Africa). —Gidrobiologicheskii Zhurnal 5:70-8 1.

Manter, H. W. 1955. Two new monogenetic tre-
matodes from elephant fishes (Callorhynchus)
from South Africa and New Zealand. Pp. 211-
220 in Essays in the natural sciences in honor
of Captain Allan Hancock. University of South-
ern California Press, Los Angeles.

Suriano, D. M., & I. S. Incorvaia. 1982. Sistematica

VOLUME 102, NUMBER 1

y biologia de Callorhynchocotyle marplatensis
gen. et sp. n. (Monogenea: Polyopisthocotylea)
parasita de las branquias de Callorhynchus cal-
lorhynchus (Linne, 1758) Garman, 1904 (Pisces:
Holocephali) de la region costera de Mar del
Plata. —Comunicaciones del Museo Argentino
de Ciencias Naturales Bernardino Rivadavia e
Instituto Nacional de Investigacion de las Cien-
cias Naturales 2:19-32.

(WAB) Department of Biological Sci-
ences, Idaho State University, Pocatello,

271

Idaho 83209, and Instituto Nacional de
Pesquisas da Amazonia, Manaus, Amazon-
as, Brazil; (DCK) Department of Allied
Health Professions and Idaho Museum of
Natural History, Idaho State University,
Pocatello, Idaho 83209; (JP) Departamento
de Ciéncias Morfobidlogicas, Fundacao
Universidade do Rio Grande, Rio Grande,
Brazil.

PROC. BIOL. SOC. WASH.
102(1), 1989, pp. 272-279

PHYTOPLANKTON COMPOSITION IN A
BORROW PIT LAKE IN VIRGINIA

Seba B. Sheavly and Harold G. Marshall

Abstract.—The phytoplankton assemblages in Lake Trashmore, Virginia, a
borrow pit, were dominated by centric diatoms and cyanobacteria, with sea-
sonal pulses of cryptomonads, euglenoids, and chlorophyceans. Ninety species
were identified and their abundance levels noted for a 12-month period.

Borrow pit lakes are generally associated
with highway construction and residential
landscaping. In contrast, this borrow pit lake
was formed as part of an above-ground
landfill operation. Alternate layers of soil
and refuse were laid down to eventually form
a mound that was capped with soil, then
landscaped. The pit formed in this process
was gradually filled with water by 1971. The
lake is located in Virginia Beach, Virginia,
where it is now a part of a recreational park
complex. The lake has a mean depth of 4
m, a maximum depth of 7.3 m, a surface
area of 21 ha, and a volume of 8.4 x 10°
m? (Virginia State Water Control Board
1982). The surrounding area is highly de-
veloped. The lake is flanked to the north
and south by major highways and is adja-
cent to the disposal mound (Mount Trash-
more) on the west, with residential devel-
opment along the eastern section.

The purpose of this study was to identify
the seasonal phytoplankton composition and
concentrations of this lake and to make
comparisons with a phytoplankton survey
of the lake made over a decade ago (Cocke
1973).

Other phytoplankton studies in Virginia
have concentrated on the two natural lakes
in the state, Lake Drummond and Moun-
tain Lake (Simmons & Neff 1974, Marshall
1979). In addition, the National Eutrophi-
cation Survey (Anonymous 1972) included
eight other Virginia lakes and reservoirs that

were of greater size than Lake Trashmore,
and all were found to be eutrophic. Phyto-
plankton were included in a limited water
quality study during the early formation of
Lake Trashmore by Beck (1973). He indi-
cated an abundance of diatoms, chlorophy-
ceans, cyanobacteria and phytoflagellates.
Later Cocke (1973) completed a one year
study of the lake and reported 45 taxa, with
chlorophyceans and diatoms dominant.

In general, borrow pit lake studies are rare.
In Nebraska, McCarraher et al. (1974) sur-
veyed 41 borrow pit lakes in the Platte River
Valley, with Adrian et al. (1970) conducting
a primary productivity study on one lake.
Seven Illinois borrow pit ponds were stud-
ied by Lipsey (1980). In all of these studies,
the formation of borrow pits was associated
with a major highway development. Dia-
toms, cyanobacteria and chlorophyceans
were generally the dominant forms in these
ponds or lakes.

Methods. —One collection station was es-
tablished at the center and deepest (7.3 m)
portion of the lake. Samples were collected
monthly from the upper and lower 0.5 m
of the euphotic zone from Mar 1985 to Feb
1986. The lower sampling depth was deter-
mined from transparency measurements
with a Secchi disk (15 cm diam), according
to Holmes (1970). A Kemmerer water bot-
tle (2 liter) was used to obtain 500 ml water
samples from both depths. These were pre-
served immediately with Lugol’s solution

VOLUME 102, NUMBER 1

and returned to the laboratory, where a set-
tling and siphoning procedure was followed
to obtain a 40 ml concentrate. Aliquots were
taken from this concentrate and placed in
settling chambers for examination with a
Zeiss inverted plankton microscope. A ran-
dom field and minimum count procedure
was followed at magnifications of 312 x and
500 x for micro-, nano-, and picoplankton
for a precision estimate of 85%. In addition,
the net phytoplankton were counted by
scanning the entire chamber at 125.

Results. -—Mean water temperatures
ranged from 4.0°C in January to 27.3°C in
August, with only small differences gener-
ally found with depth in the euphotic zone.
The mean monthly pH value was 7.52, with
a range from 6.5 (Dec) to 8.6 (Sep). Secchi
disk readings ranged from 0.47 m (Mar) to
1.18 m (Aug), averaging 0.84 m.

Ninety phytoplankton species were iden-
tified in this study (Table 1), including 32
Chlorophyceae, 28 Bacillariophyceae, 16
Cyanobacteria, 8 Euglenophyceae, 5 Di-
nophyceae, and 1 Cryptophyceae. Uniden-
tified picoplankton and nanoplankton cells
were counted by size groups <3, 3-5 and
5-10 wm. They were present throughout the
year, with lowest concentrations in May (3.7
x 10° cells/liter), gradually increasing to a
December high (61.5 x 10° cells/liter). The
majority of these cells were <3 wm in size
and under epifluorescence microscopy
proved to be cyanobacteria. In contrast,
larger and identifiable cyanobacteria had one
major pulse in summer and early fall, reach-
ing 37.0 x 10° cells/liter in September. Eu-
glenoids and dinoflagellates also had high
summer concentrations. In contrast, the
cryptomonads were in lowest concentrations
in summer and early fall and became more
abundant during winter and spring. The
diatoms had low concentrations in spring
and summer, gradually increased into fall
and early winter and peaked in December
(9.4 x 10° cells/liter). The dominant species
were Cyclotella spp. and Melosira spp., with
representatives from these genera most

273

common in the unidentified category of cen-
tric diatoms <20 um size.

Discussion. —In comparison to early data
on phytoplankton populations in Lake
Trashmore (Beck 1973, Cocke 1973), there
has been a shift in composition and domi-
nant species. Cocke identified 45 species in
a seven-month study (Aug—Feb), with the
two dominant groups: pennate diatoms (e.g.,
Pleurosigma normanii, P. strigosum) and
chlorophyceans (Pediastrum simplex) the
major species. He found cyanobacteria
(blue-green algae) to be common, but not
abundant. Other dominant species included
the desmids Closterium lunuae and Cos-
marium circulare, the diatoms Fragilaria
crotonensis, Melosira spp., Navicula spp..,
and the dinoflagellate Gymnodinium sim-
plex. Secchi disk readings at this time ranged
between 24.0 and 54.0 cm, averaging 40.9
cm. The range of the surface pH values was
6.6 to 7.1, averaging 6.8. The present study
indicated a more basic pH mean of 7.52,
ranging between 6.5 and 8.6. However, the
Secchi disk readings were higher, averaging
84 cm, with readings between 47 cm (Mar)
and 118 cm (Aug).

In addition to an increased diversity of
species, there has been a change over the
past decade in the phytoplankton popula-
tions in Lake Trashmore. The transition has
been from a dominance of pennate diatoms,
chlorophyceans, and a filamentous-coccoid
assemblage of cyanobacteria to the current
status, where cyanobacteria, centric dia-
toms (e.g., Cyclotella spp., Melosira spp.),
plus a seasonal abundance of cryptomo-
nads, euglenoids, and chlorophyceans are
dominant. It is impossible to evaluate the
significance of the high picoplankton and
nanoplankton (<10 um) concentrations be-
cause Cocke’s collection procedure by tow
net would not have collected many of these
cells. However, these changes in phyto-
plankton composition and abundance are
assumed to be in association with the chang-
ing and advancing eutrophic state of the lake
over the past decade.

274

Table 1.—Mean monthly abundance for each identified taxon (cells/liter).

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Mar Apr May Jun
Bacillariophyceae

Achnanthes sp. 47,268 639 5373 1S
Amphora costata W. Smith 0 0 0 213
Amphora sp. 0 21 0 0
Biddulphia alternans (Bailey) Van Heurck 0 0 5 0
Cocconeis distans Gregory 21 0 0 0
Cyclotella meneghiniana Kitzing 0 77,867 29,373 18,131
Cyclotella sp. 273,843 27,084 28,213 4578
C. striata (Kiitzing) Grunow 0 0 18,056 0
Cylindrotheca closterium (Ehrenberg)

Reimann & Lewin DTT 0 0 53
Cymbella sp. 9515 602 123 85
Diploneis sp. 0 0 16 12
Fragilaria sp. 0 0 0 10,579
Gomphonema sp. 0 0 16 53
Gyrosigma sp. 0 0 0 21
Melosira distans (Ehrenberg) Kiitzing 33,824 0 0 0
M. granulata (Ehrenberg) Ralfs 221,898 9649 299 1664
M. islandica Muller 5195 0 4557 85
M. moniliformis (Muller) Agardh 0 0 32 0
Melosira sp. 0 37 107 427
Navicula spp. 9813 37 32 117
Nitzschia clausii Hantzsch 0 0 0 64
N. pungens Grunow 43 0 0 75
N. seriata Cleve 43 0 0 0
Nitzschia sp. 14,411 16 0 32
Rhizosolenia eriensis H. L. Smith 0 0 0 0
Synedra acus Kaiitzing 0 0 0 0
Synedra spp. 0 21 85 587
Thalassionema nitzschioides (Grunow)

Grunow & Hustedt 0 16 0 384
centrics (unid.) <20 wm diam. 285,376 293,413 178,303 81,607
centrics (unid.) 20-100 um diam. 109,243 0 0 14,374
pennates (unid.) >20 um in length 51,857 56,426 28,341 22,072
pennates (unid.) <20 um in length 26,076 112 101 21

subtotal 1,088,703 465,940 294,032 156,000
Dinophyceae
Ceratium hirundinella (O. F. Muller)

Dujardin 0 0 368 19,040
Glenodinium gymnodinium Penard 0 0 0 43
Glenodinium sp. 0 3 0 0
Gymnodinium sp. 0 0 0 36,113
Protoperidinium sp. 0 0 0 96

subtotal 0 3 368 55,292
Cyanobacteria
Anabaena sp. IP) 23,138 0 576
Chroococcus limnecticus Lemmerman 0 0 0) 477,533
Dactylococcopsis raphidiodes Hansgirg 2,607,829 920,865 478,484 101,230
Gomphosphaeria aponina Kutzing 120,610 lay 2294 9028
Johannesbaptistia pellucida (Dickie)

Taylor & Drouet 0 0 0 0

Lyngbya controta Lemmermann 0 0 0 0

VOLUME 102, NUMBER 1 275
Table 1.—Continued.
Jul Aug Sep Oct Nov Dec Jan Feb
1984 0 0 0 0 256 0 0
469 64 107 0 0 0 0 64
0 0 0 0 0 247 0 0
0 0 0 0 0 0 0 0
0 0 0 0 0 0 0 0
18,057 18,078 36.113 0 36,455 180,907 J9,297, 483,016
9028 P5525 52 270,850 81,255 81,255 424,328 40,626 731,316
0 0 22.9571 0 0 0 0 40,628
0 32,154 46,187 0 128 213 208 725
43 64 207 128 128 640 52 139
43 0 0 0 0 0 0 0
533 853 36,849 0 341 768 0 0
0 64 0) 0 0 0 0 0
0 0 0 0 0 0 0 0
0) 0 0 36,415 1,363,278 3,403,654 ES7.677 153,780
28557 31,864 248,919 768,108 806,756 3,710,612 343,064 690,668
320 1472 427 198,623 0 0 0 0
0 0 0 0 0 0 0 0
9220 0 0 40 0 0 0 0
768 939 2965 1493 256 1963 64 18,099
9114 4557 0 0 0 256 0 0
0 0 0 0 0 0 0 0
0 0 0 0 0 0 0 0
128 0 Se) 0 0 0 0 0
0 117,432 63,198 0 0 0 0 0
43 875 532675 128 469 700 S17 789
597 51,789 124,323 49,878 3029 3212 171 299
299 18,505 2T2Z2NS 0 0 180,799 0 0
117,368 153,482 329,534 135,425 261,864 1,426,466 279,190 1;923;035
oA 21 18,227 0 213 640 38,369 31,599
22,571 9028 995312 54,170 939 232,435 58,682 90,282
85 40,734 341 85 768 1244 8028 533
219,248 617,507 1,626,645 1,325,748 2,555,879 9,569,435 961,930 4,164,983
5589 3392 23,467 13,227 1408 537 16 0
85 1835 1771 981 0 0 0 0
213 0 1600 18,185 0) 0 0 0
64 0 0 0 0 0 22,570 0
43 0 0 0 0 0 0 0
5994 3227 26,838 32,393 1408 ae We 22,586 0
1,173,683 9,619,689 6,238,578 157605525 2987 356,250 0 0
537,186 64,009 643,401 451,417 504,161 2,644,782 205,091 175,549
94,947 0 225,708 370,162 353,897 839,628 815,906 1,047,287
0 128 100,784 397,247 91,862 58,479 21,479 32,815
341 0 0 0 0 0 0 0
0 794,493 174,098 64,350 0 0 0 0

SSS

276

Table 1.—Continued.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Mar Apr May Jun
Merismopedia glauca (Ehrenberg)

Naegeli 0 0 0 0
Merismopedia punctata Meyen 0 0 8038 18,526
Merismopedia sp. 0 0 0 0
Merismopedia tenuissima Lemmermann 0 0 0 0)
Microcystis aeruginosa Kiutzing 408,516 2,471,603 2,059,101 2,719,878
Nostoc commune Vaucher DAT 15,738 432 779
Oscillatoria limnetica Lemmermann 0 0 0 0
Oscillatoria sp. 0 32 208 4274
Spirulina laxa G. M. Smith 0 0 0 53
Spirulina subsalsa Oersted 0 0 0 52
blue green spheres (unid) 0) 0 54,168 0
blue green trichomes (unid) 64 1129 6782 23,010

subtotal 3,138,021 3,432,657 2,609,507 3,354,919
Euglenophyceae
Euglena acus Ehrenberg 0 0 0 0
Euglena sp. 0 0 0 1821
Eutreptia lanowii Steuer py syl 52,529 D591 87,704
Eutreptia viridis Perty 85 0 5) 43
Phacus longicaudus (Ehrenberg) Dujardin 0 0 0 0
Phacus curvicaudus Swirenko 0 0 0 0
Trachelomonas hispida (Perty) Stein 0 0 0 0
Trachelomonas volvocina Ehrenberg 0 0 0 76,608
subtotal 2342 52,529 22,591 182,572
Chlorophyceae
Ankistrodesmus falcatus Beijerinck 156,672 27,655 31,316 31,445
Ankistrodesmus fractus (West & West)

Brunnthaler 0 0 0 0
Chlorella sp. 9028 0 0 0
Chlorella vulgaris Beijerinck 0 0 0 4104
Cosmarium botrytis Meneghini 0 0 0 0
Crucigenia apiculata (Lemmermann)

Schmidle 0 0 0 0
Crucigenia fenestrata Schmidle 92,219 0 0 18,057
Crucigenia quadrata Morren 0 0 9113 744,539
Crucigenia sp. 0 0) 4514 0
Crucigenia tetrapedia (Kirchner) West

& West 1,831,853 1435152 0 207,908
Dictyosphaerium pulcellum Wood 1365 309,316 143,839 2,013,046
Euastrum denticulatum (Kirchner) Gay 0 0 0 21
Franceia droescheri (Lemmermann) 0 0 0 0
Kirchneriella contorta (Schmidle) Bohlin STL WS) 293,977 302,438 283,177
Lagerheimia ciliata (Lagerheim) Chodat 0 0 0 0
Lagerheimia quadriseta Lemmermann

(G. M. Smith) 0 0 0 0
Micractinium pusillum Frensenius 0 0 0 0
Oedogonium sp. 0 0 421 0
Oocystis borgei Snow 114,091 129,214 105,047 250,682
Pediastrum duplex Meyen 0 0 0 0
Pediastrum simplex (Meyen)

Lemmermann 36,147 421 1259 5845
Scenedesmus armatus (Chodat) G. M. Smith 363 0 0 0

VOLUME 102, NUMBER 1 ZIT
Table 1.—Continued.

Jul Aug Sep Oct Nov Dec Jan Feb

0 S225 0 341 0 0 0 0

0 eA | ZAZS 1024 308,329 704 0 0

0 0 0 0 288,907 0 0 0

0 OF» 1559241935 5,673,395 870,724 928 18,056 0

£251,540 75031,505 8,564,855 9,331,530 6,593,405 4,709,592 82,145 251,908

2,902,609 6,414,631 4,441,940 5,254,490 3,891,212 2,146,677 928,350 853

0 993,117 672,611 3615133 0 0 0 0

22,912 132756 0 18,057 0 427 52 0

0 18,675 85 0 ii | 0 0 0

0 9071 0 0 0 0 0 0

0 0 0 0 0 0 0 0

4514 9028 90,283 18,057 43 0 48 213

14,987,732 25,083,530 37,079,401 23,701,728 1,290,568 10,757,467 2,071,107 150,862

0 192 299 0 0) 0 0 0

395953 12,634 186,958 55,493 768 341 0 0

1263397 54,234 288,907 180,567 63,284 208,333 80 4099

0 21 0 0 0 0 0 0

0 0 1664 0 128 0 0 0

0 256 e709 95,446 1323 0 2 0

448 832 832 46,282 118,307 2603 16,992 49,763

525218 0 0 27,085 TT, 54,170 16,928 63,433

198,568 67,785 355,751 404,873 256,037 265,447 34,032 154,186

66,646 9600 46,780 317,562 712,133 8021 437 405

0 0 0 0 0 112,487 6485 2112

0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0

0 0 $6,113 0 384 0 0 0

0 0 0 0 583,616 15295370 252,869 597

0 0 0 0 63,198 0 341 77,082

T2376 162,851 110,900 580,928 361,133 O15 07 DHT 1280

18,057 0 0 0 0 0 0 235

341 0 144,453 306,963 184,151 1,594,264 188,460 217;399

0 0 0 0 0 0 0 0

128 9178 135,425 S97 299 1024 0 0

18,057 0 0 0 384 0 0 0

18,057 0 0 4949 383,115 1,119,844 125,264 112,876

0 Joa 0 0 0 0 0 0

63,198 58,684 18,057 7d 0 341 0 0

0 0 0 4779 0 0 0 0

0 Nig 0 0 0 0 0 0

176,053 361,389 487,701 397,247 297,935 2,058,442 164,761 es er eS

0 0 0 0 0 0 0 0

10,197 10,603 G3, 95d 88,832 153,583 55,808 599 1792

0 0 0 0 0 0 0 0

278 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

Mar Apr May Jun
Scenedesmus bijugus Turpin Lager. 20,096 2343 0 37,685
Scenedesmus dimorphus (Turpin) Kutzing 0 0 0 85
Scenedesmus quadricaudus (Turpin)

Brébisson 9625 40,405 27,276 196,406
Scenedesmus sp. 85 11,285 4530 18,398
Selenastrum gracile Reinsch 0 0 0 0
Staurastrum americanum (West & West) G. M.

Smith 0 0 11 0
Staurastrum leptocladum var. insigne West

& West 0 0 80 3008
Staurastrum paradoxum Meyen 0 0 11 341
Staurastrum sp. 0 a7 43 117
Tetraedron minimum (Braun) Hansgirg 64 580 21 102,122
Chlorophyceans (unid.) 171 16,928 136,549 650,038

subtotal 2,462,904 975,313 766,468 4,567,024
Cryptophyceae
Cryptomonas sp. 825,700 1,816,905 1,271,836 554,242
subtotal 825,700 1,816,905 1,271,836 554,242
Other taxa
micro-phytoflagellates <10 um 47,637 0 0 175,407
micro-phytoflagellates > 10 wm 0 0 8215 43,810
small green spheres (<3 um) 18,320,895 3,792,191 3,348,778 9,835,356
small green spheres (3—5 wm) 4,786,425 470,241 262,864 675,405
small green spheres (5—10 um) 997,156 131,192 84.883 115,001
subtotal 24,152,113 4,393,624 3,704,740 10,844,979
Total 31,669,783 11,136,971 869,542 19,715,028

Literature Cited

Adrian, G. L., C. Throckmorton, & R. McDonald.
1970. A study of primary production in a Ne-
braska Interstate 80 lake.— Transactions of the
Kansas Academy of Science 73(2):227—236.

Anonymous. 1972. Water quality criteria. A report
of the Committee on Water Quality Criteria. —
National Academy of Science and National
Academy of Engineering. Washington, D.C., 219
pp.

Beck, W. M., Jr. 1973. Building an amphitheater and
costing ramp of municipal solid waste. U.S. En-
vironmental Protection Agency. SW-52d.of. Of-
fice of Solid Waste Programs Vol. I and II.
Washington, D.C., 265 pp.

Cocke, G. R. 1973. A comparative limnological sur-
vey of a brackish water lake and freshwater lake
at Mount Trashmore, Virginia Beach, Virginia.
Masters Thesis. Old Dominion University,
Norfolk, Virginia, 120 pp.

Holmes, R. W. 1970. The secchi disk in turbid coastal
waters. — Limnology and Oceanography 15:688-
694.

Lipsey, L.L. 1980. Phytoplankton of selected borrow
pit ponds in northern Illinois.— Ohio Journal of
Science 80(3):108-113.

Marshall, H.G. 1979. Lake Drummond: With a dis-
cussion regarding its plankton composition. Pp.
169-182 in P. W. Kirk, ed., The Great Dismal
Swamp. University Press, Charlottesville, Vir-
ginia.

McCarraher, D. B., R. McDonald, & G. Adrian. 1974.
Some hydrobiological characteristics of Inter-
state-80 highway lakes in Nebraska.—Trans-
actions of the Kansas Academy of Science 77(2):
93-102.

Simmons, G. M., & S. E. Neff. 1974. Observations
on limnetic carbon assimilation rates in Moun-
tain Lake, Virginia during its thermal stratifi-
cation periods.— Virginia Journal of Science
24(4):206—-211.

VOLUME 102, NUMBER 1

Table 1.—Continued.

Jul Aug Sep Oct Nov Dec Jan Feb
388,218 173,006 439,333 507,080 257,401 15,104 299 1365
128 341 72,483 683 0 1024 0 0
193,623 55,109 218,088 361-133" ~ 352505202 326,042 84,234 198,623
49,656 0 0 153,482 35,455 270,847 0 0
0 0 21,662 128 0 0 0 0
0 0 0 0 0 981 0 64
555 2688 1536 1621 1579 512 16 0
128 1152 1314 724 384 256 0 64
13,628 85 0 0 0 0 32 0
(Peps | 148,968 2305223 252-193 99,696 622,950 ZIV SS ZAG?
1,060,829 S72 34" 3.954 410) 213975150. ~. 1,932,063). 3,186,975 354,349 442,388
Pee NO2 [887,651 5,982,435 . 24,376,822 4,752,529 11,915,805 1,232,874 1,315,247
306,963 446,903 979,574 451,417 486,558 1,733,426 822,677 1,584,473
306,963 446,903 979,574 451,417 486,558 1,733,426 822,677 1,584,473
81,255 279,878 419,818 343,077 379,190 442,385 144,448 18,057
1535 1535393 0 0 591,435 219,050 240,959 503,816
14,763,970 14,238,250 18,641,155 13,708,148 16,143,985 56,011,283 5,542,049 12,748,730
1,171,918 104,048 3,066,790 1,533,350 1,555,255 3,986,723 3933223 6Go75151
328,575 394,290 6,900,008 635,245 240,955 876,203 35,596 120,478
16,361,053 16,106,241 22,817,681 16,219,820 18,910,820 61,535,644 6,316,275 14,048,232
34,306,660 44,214,844 69,068,298 66,512,801 39,870,929 95,777,761 11,461,481 22,775,746

Virginia State Water Control Board. 1982. Classifi- Department of Biological Sciences, Old

cation and Priority Listing of Virginia Lakes.
U.S. Environmental Protection Agency. S-
003219-010. Richmond, Virginia, 727 pp.

29

Dominion University, Norfolk, Virginia

23529.

280 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

INTERNATIONAL COMMISSION ON ZOOLOGICAL
NOMENCLATURE

% British Museum (Natural History)
Cromwell Road
London, SW7 5BD
23 September 1988

The following applications were published on 23 September 1988 in Vol. 45, Part
3 of the Bulletin of Zoological Nomenclature. Comment or advice on these appli-
cations is invited for publication in the Bulletin and should be sent to the Executive
Secretary, ICZN, British Museum (Natural History), Cromwell Road, London SW7
SBD, U.K.

Case No.

2449 Septotrochammina Zheng, 1979 (Foraminiferida): proposed designation of
Remaneica gonzalezi Seiglie, 1964 as the type species.
2622 Pleuromma princeps Scott, 1894 (currently Gaussia princeps; Crustacea, Co-
pepoda): proposed conservation of the specific name.
2568 ACRIDIDAE Karny, 1907, OEDIPODIDAE Walker, 1870 and LOCUSTIDAE La-
treille, 1802 (Insecta, Orthoptera): proposed order of precedence.
2618 Bruchus Linnaeus, 1767, Ptinus Linnaeus, 1767 and Mylabris Fabricius, 1775
(Insecta, Coleoptera): proposed conservation.
2627 Coryphium angusticolle Stephens, 1834 (Insecta, Coleoptera): proposed con-
servation of both the generic and specific names.
2632 Tachina orbata Wiedemann, 1830 (currently Peribaea orbata; Insecta, Dip-
tera): proposed confirmation of neotype designation.
2628 Tenthredo zonula Klug, 1817 (Insecta, Hymenoptera): proposed conservation
of the specific name.
2625 Saccopharynx Mitchill, 1824 (Osteichthyes, Saccopharyngiformes): proposed
conservation.
2616 ICHTHYOPHIIDAE Taylor, 1968 (Amphibia, Gymnophiona): proposed con-
servation.
2650 Thorius pennatulus Cope, 1869 (Amphibia, Caudata): proposed conservation
of the specific name.
2441 Semioptera wallacii Gray, 1859 (Aves, Paradisaeidae): proposed confirma-
tion as the correct spelling.
2640 Mus musculus domesticus Schwarz & Schwarz, 1943 (Mammalia, Rodentia):
proposed conservation.
J. D. D. SMITH
Scientific Administrator

VOLUME 102, NUMBER 1 281

INTERNATIONAL COMMISSION OF ZOOLOGICAL
NOMENCLATURE

% British Museum (Natural History)
Cromwell Road
London, SW7 5BD
23 September 1988

The following Opinions were published on 23 September 1988 in Vol. 45, Part
3 of the Bulletin of Zoological Nomenclature.

Opinion No.

1501 Alveolina d@Orbigny, 1826 (Foraminiferida): Oryzaria boscii Defrance in
Bronn, 1825 designated as the type species.

1502 Conus fergusoni G. B. Sowerby III, 1873 (Mollusca, Gastropoda): specific
name conserved.

1503 PSEUDOCALANIDAE Sars, 1901 (Crustacea, Copepoda): not to be given pre-
cedence over CLAUSOCALANIDAE Giesbrecht, 1892.

1504 BERYTIDAE Fieber, [1851] and Berytinus Kirkaldy, 1900 (Insecta, Heterop-
tera): conserved.

1505 Sigara scholtzi Fieber, [1860] (currently Micronecta (Dichaetonecta) scholtzi;
Insecta, Heteroptera): specific name conserved.

1506 Oncomera Stephens, 1829 (Insecta, Coleoptera): Dryops femorata Fabricius,
1792 designated as the type species.

1507 Musca marginalis Wiedemann, 1830 (currently Chrysomya marginalis; In-
secta, Diptera): specific name conserved.

1508 Simulium austeni Edwards, 1915 (Insecta, Diptera): not to be given prece-
dence over Simulium posticatum Meigen, 1838.

1509 Paraphytomyza Enderlein, 1936 (Insecta, Diptera): Phytagromyza luteo-
scutellata de Meiere, 1924 designated as the type species.

1510 Microgaster Latreille, 1804 (Insects, Hymenoptera): Microgaster australis
Thomson, 1895 deisgnated at the type species.

1511 Halictus costulatus Kriechbaumer, 1873 (currently Lasioglossum costulatum;
Insecta, Hymenoptera): specific name conserved.

1512 Desorella Cotteau, 1855 (Echinodermata, Echinoidea): confirmation of Hy-
boclypus elatus Desor, 1847 as the type species.

1513. Synapturanus Carvalho, 1954 (Amphibia, Anura): Synapturanus miranda-
riberoi Nelson & Lescure, 1975 designated as the type species.

1514 Liasis Gray, 1842 (Reptilia, Serpentes): Liasis mackloti Duméril & Bibron,
1844 designated as the type species.

1515 LARIDAE Rafinesque Schmaltz, 1815 (Aves) and LARINI LeConte, 1861 (In-
secta, Coleoptera): homonymy removed.

1516 Taeniolabis Cope, 1882 (Mammalia, Multituberculata): Polymastodon taoensis
Cope, 1882 designated as the type species.

1517 Viverravus gracilis Marsh, 1872 (Mammalia, Carnivora): generic and specific
names conserved.

282 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

REVIEWERS

The following people reviewed manuscripts for the Proceedings in 1988. A. Alvarino, M. V. Angel,
F. M. Bayer, B. Bieri, R. Birdsong, J. A. Blake, T. Bongers, J. Bouillon, T. E. Bowman, R. O.
Brinkhurst, B. Brown, S. D. Cairns, D. R. Calder, R. Castro, F. L. Chace, Jr., L. L. Cole, B. B. Collette,
P. F. S. Cornelius, R. F. Cressey, A. Crosnier, D. Davis, G. M. Davis, H. DeWitt, M. Dojiri, S.
Edmonds, D. J. Eernisse, C. Ehrlich, K. Fauchald, K. Fitzhugh, V. A. Funk, A. L. Gardner, S. Gelder,
I. M. Goodbody, T. M. Gosliner, G. R. Graves, K. Green, M. Harasewych, E. Harrison-Nelson, C.
W. Hart, Jr., W. R. Heyer, H. H. Hobbs, Jr., W. D. Hope, J. Iverson, S. C. Jameson, B. F. Kensley,
L. S. Kornicker, J. D. Kudenov, K. Kuroda, D. B. Lellinger, R. J. L. Lester, M. S. Loden, J. Lynch,
W. G. Lyons, N. J. Maciolek, R. B. Manning, J. W. Martin, R. W. McDiarmid, J. H. McLean, M.
Milligan, G. C. Mitchell, R. F. Modlin, R. J. Mooi, T. A. Munroe, A. T. Newberry, R. T. O’Grady,
S. L. Olson, L. Parenti, K. Parks, D. L. Pawson, T. H. Perkins, M. H. Pettibone, W. L. Pratt, J. W.
Reid, M. E. Rice, M. Robbins, C. R. Robins, W. Rudman, D. E. Russell, J. M. Savage, V. G. Springer,
W. C. Starnes, M. Stiassny, F. G. Thompson, D. A. Thoney, R. P. Vari, M. Vecchione, T. C. Walter,
L. Watling, S. H. Weitzman, M. J. Wetzel, A. B. Williams, S. Williams, R. Winterbottom, J. J.
Wurdack, R. Zottoli, G. R. Zug, R. L. Zusi.

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CONTENTS

Paleontological type specimens in the Santa Barbara Museum of Natural History
Scott E. Miller
A new generic name for Tate’s (1933) Microtarsus group of South American mouse opossums
(Marsupialia: Didelphidae) Alfred L. Gardner and G. Ken Creighton
Description of a rock-dwelling cichlid (Teleostei: Cichlidae) from Lake Malawi, Africa
J. R. Stauffer, Jr. and J. M. Boltz
Ascidians collected around the Galapogos Islands using the Johnson-Sea-Link research sub-

mersible Claude Monniot and Francoise Monniot
Eukrohnia calliops, a new species of Chaetognatha from the northern Gulf of Mexico with
notes on related species Jerry A. McLelland

Smalleyus tricristatus, new genus, new species, and Pseudothelphusa parabelliana, new species
(Brachyura: Pseudothelphusidae) from Los Tuxtlas, Veracruz, México Fernando Alvarez
Two new genera and nine new species of geryonid crabs (Crustacea, Decapoda, Geryonidae)
Raymond B. Manning and L. B. Holthuis
Synalpheus arostris and Philocheras lapillus, two new species of caridean shrimp (Crustacea)
from the tropical eastern Pacific Mary K. Wicksten
The holotype of Heterocarpus alexandri A. Milne-Edwards (Crustacea: Decapoda: Pandalidae)
Fenner A. Chace, Jr.
Gammaropsis Ee a new species of marine Amphipoda (Crustacea) from Jamaica
James Darwin Thomas and J. L. Barnard
Natatolana pastorei (Giambiagi, 1925) (Crustacea, Isopoda, Cirolanidae) from the Straits of
Magellan, South America: Redescription and notes on functional morphology
Johann Wolfgang Wagele and Niel L. Bruce
Neobrachiella anisotremi (Copepoda: Lernaeopodidae), a new species parasitic on an inshore
fish, Anisotremus scapularis, off the Chilean coast
Raul Castro Romero and Hernan Baeza Kuroki
The First Annual Riser Lecture: Meiofauna and Microbes—the interactive relations of annelid

hosts with their symbiotic bacteria Olav Giere
Revision of the family Lysaretidae, and recognition of the family Oenonidae Kinberg, 1865
(Eunicida: Polychaeta) G. Kent Colbath
Redescription of Nephtys squamosa Ehlers (Polychaeta: Nephtyidae) Takashi Ohwada
Phallodrilus hessleri, new species (Oligochaeta: Tubificidae) from abyssal depths in the western
Pacific Ocean Christer Erséus

Phallodrilus vescus, new species (Oligochaeta, Tubificidae) from the Gulf of Mexico
Christer Erséus
New species of scale-worms (Polychaeta: Polynoidae) from ee hydrothermal rift-area of the
Mariana Back-Arc Basin in the western central Pacific Marian H. Pettibone
Polynoidae and Sigalionidae (Polychaeta) from the Guaymas Basin, with descriptions of two
new species, and additional records from hydrothermal vents of the Galapagos Rift, 21°N,
and seep-sites in the Gulf of Mexico (Florida and Louisiana) Marian H. Pettibone
A new eulimid gastropod, Trochostilifer eucidaricola, parasitic on the pencil urchin Eucidaris
tribuloides from the southern Caribbean Anders Warén and Robert Moolenbeek
Springsnails (Gastropoda: Hydrobiidae) of Owens and Amargosa River (exclusive of Ash
Meadows) drainages, Death Valley System, California-Nevada Robert Hershler
Four new species of free-living marine nematodes in the genus Pareurystomina (Nematoda:
Enoplida) with observations on other members of the genus Edwin J. Keppner
Neotropical Monogenea. 14. Revision of Callorhynchocotyle Suriano and Incorvaia, 1982
(Hexabothriidae) with the description of C. amatoi
Walter A. Boeger, Delane C. Kritsky, and Joaber Pereira, Jr.
Phytoplankton composition in a borrow pit lake in Virginia
Seba B. Sheavly and Harold G. Marshall
International Commission on Zoological Nomenclature: Applications and Opinions

Reviewers— 1988

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THE BIOLOGICAL SOCIETY OF WASHINGTON

1988-1989
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PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 283-299

A REVISION OF ARABELLA MUTANS
(CHAMBERLIN, 1919) AND RELATED SPECIES
(POLYCHAETA: ARABELLIDAE)

G. Kent Colbath

Abstract. — Arabella mutans (Chamberlin, 1919), previously considered cir-
cumtropical in distribution, is revised using a combination of soft-part, setal,
and maxillary characters. Emphasis is placed on adopting a population ap-
proach to evaluating character states. Poorly preserved, isolated, or juvenile
individuals may not be identifiable to species based on the criteria advocated
here. Seven species with non-overlapping geographic ranges are recognized.
Arabella mutans s. s. is restricted to tropical Polynesia. Arabella novecrinita
Crossland, 1924 is recorded from the western Indian Ocean. Arabella logani
Crossland, 1924 and A. atlantica Crossland, 1924 are recognized as distinct
species, and recorded from the Red Sea and Cape Verde Islands, respectively.
Arabella multidentata (Ehlers, 1887) is recognized to encompass western At-
lantic and Carribean forms. Two new species are erected: Arabella panamensis

for specimens from Pacific Panama, and A. monroi from the Galapagos.

The genus Arabella is distributed
throughout the world ocean, and includes
20 named species (Fauchald 1977b). Many
of these species are relatively rare, and have
not been satisfactorily circumscribed in
terms of either morphology or distribution.

Arabella mutans (Chamberlin, 1919) is
one such species. Although A. mutans is re-
ported to have a circumtropical distribution
(Monro 1928, 1933a, b; Day 1967; Reish
1968; Gardiner 1976), this assertion has not
been supported by detailed morphological
comparisons of specimens (Fauchald 1970).
The present study attempts to partially re-
dress this situation by presenting informa-
tion compiled from the detailed examina-
tion of numerous specimens, including the
holotypes of both A. mutans and the similar
A. novecrinita Crossland, 1924.

From a paleontological perspective, Ar-
abella mutans is of interest because of the
wide range of jaw morphologies which have
been recorded for the species. Establishing
constraints on the amount of variation in
jaw morphology that can be expected within

modern species should aid in the interpre-
tation of fossil jaw assemblages.

History of Study

Chamberlin (1919:329-—332) described the
new genus and species Cenothrix mutans
based on a single specimen collected from
Easter Island by the Albatross Expedition.
He distinguished the genus Cenothrix from
Arabella Grube, 1850 on the presence of a
ventral, hooded seta in medial and posterior
parapodia in the former, and the supposed
absence of such setae in the latter. The new
species was distinguished on the same taxo-
nomic character.

Crossland (1924:70) considered the pres-
ence ofa single hooded seta per parapodium
insufficient grounds for recognition of a new
genus, and transferred C. mutans into Ar-
abella. He described the new species Ara-
bella novecrinita based on 16 specimens
from the Atlantic and Indian Oceans, and
noted that A. novecrinita “*... is like, and
probably identical with A. (Cenothrix) mu-

284

tans,’ (Crossland 1924:71). Crossland ap-
parently considered the jaws of the type
specimen of A. mutans atypical, and justi-
fied the erection of a new species on that
basis. He recognized four varieties of A. no-
vecrinita: the type material from Zanzibar
and the Maldive Islands, A. novecrinita var.
logani from the Red Sea, and A. n. var.
atlantica and A. n. var. asymmetrica from
the Cape Verde Islands.

Monro (1928:91; 1933b:88) considered
A. novecrinita a junior synonym of A. mu-
tans, and considered the species circum-
tropical in distribution. Monro essentially
used Chamberlin’s specific name for spec-
imens included within the range of variation
described by Crossland. Subsequent work-
ers prior to Orensanz (1974) followed Mon-
ro, although Fauchald (1970:129) noted that
Monro’s revision did not involve exami-
nation of the type material of either species,
and suggested that A. mutans, as considered
by Monro, might actually include several
species with more restricted distributions.

Orensanz (1974:388) used Cenothrix as a
subgenus of Arabella, and recognized three
species within the subgenus, which he dis-
tinguished based on their maxillary mor-
phology. A. mutans was recognized for spec-
imens with asymmetrical maxillae with
falcate MI elements, A. asymmetrica for
those with asymmetrical maxillae with a bi-
fid left MI, and A. novecrinita for those with
symmetrical maxillae.

Perkins (1979:445—450) followed this ar-
rangement and placed three species in Ar-
abella (Cenothrix). He made the important
observation that the type specimens of both
Arabella multidentata (Ehlers, 1887) and A.
maculosa Verrill, 1900 have ““hooded”’ ven-
tral setae, and therefore should not be con-
sidered junior synonyms of Arabella iricolor
(Montagu, 1804) as proposed by Hartman
(1944:173). Perkins distinguished A. mul-
tidentata from A. mutans based on jaw sym-
metry, and considered the two species sym-
patric in the Florida region. He recognized

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

A. maculosa based on the extreme asym-
metry of the “hoods” on the ventral setae
on the type specimen from Bermuda.

Materials and Methods

The extensive collection of specimens
catalogued as Arabella mutans in the U.S.
National Museum of Natural History
(USNM) was examined in its entirety, as
were samples identified as Arabella novecri-
nita. Additional material was obtained from
the British Museum (Natural History)
(=ZK), including Crossland’s type material
for A. novecrinita. Selected specimens of Ar-
abella iricolor from the National Museum
collection and the Florida Department of
Natural Resources (FSBC, EJ) were also ex-
amined for comparative purposes.

On each dissected specimen the lengths
of the first maxillae (MIs), length and width
of the prostomium, and length of the peri-
stomium were measured. Measurements
were made using an ocular micrometer on
a Wild M5 wide-field microscope. Denticles
were counted (where possible) for all max-
illary jaws.

Setae were prepared for scanning electron
microscopy by allowing parapodia suspend-
ed in alcohol to dry onto glass cover slips,
which were in turn glued onto stubs. Jaws
were mounted by leaving a flange of muscle
tissue around the jaw margins which, when
dried, fixed the jaws firmly onto glass cover
slips. All specimens were coated with a thin
film of carbon, then sputter coated with gold
palladium. Observations were made on a
Cambridge scanning electron microscope at
the National Museum of Natural History,
Smithsonian Institution.

Taxonomic Characters

Arabella mutans has been recognized by
most authors based on the presence of a
single morphologic character (the “hooded”
ventral seta). Consideration of additional
characters coupled with a closer examina-

VOLUME 102, NUMBER 2

285

Table 1.— Distribution of taxonomic characters in species of Arabella under consideration.

Nuchal Anal Postsetal

Species papillae cirri lobe Ventral setae Maxillary symmetry Robust MI
iricolor no 2 short ridged, gradual invariant bifid-falcate
atlantica no 2 short ridged, abrupt invariant falcate
logani no 0 short ridged, gradual ? absent
monrol no 0 short ridged, abrupt invariant falcate
multidentata no 4 short ridged, abrupt polymorphic bifid
mutans no 4 short smooth, abrupt polymorphic dentate
novecrinita no 4 short smooth, abrupt invariant absent
panamensis no y, long ridged, abrupt polymorphic falcate
sp. A yes @ short ridged, abrupt polymorphic bifid

tion of the setae has allowed separation of
discrete morphological subgroups, here rec-
ognized as distinct species with discrete geo-
graphic ranges. The taxonomic characters
are discussed below, and the character states
exhibited by each of the species are listed
in Table 1.

Soft-part morphology. —Soft-part mor-
phology within the genus Arabella is noto-
riously uniform and relatively simple, which
accounts in part for the difficulty encoun-
tered in treating these worms taxonomical-
ly. The relative proportions of the prosto-
mium are fairly uniform throughout the A.
mutans species complex, and the ventral
furrowing of the peristomium, although
variable, does not seem to vary in a system-
atic fashion.

There are three features which are useful
in differentiating these species, however. The
postsetal lobe in parapodia from A. pana-
mensis, N. sp. is markedly longer than that
in the other species.

Nuchal papillae are present in some large
individuals from Guayanilla Bay, Puerto
Rico, a unique feature in the arabellids. The
specimens are from a population of other-
wise identical worms which lack papillae,
however, and the taxonomic significance of
this character is unclear.

The morphology of the pygidium is an
extremely useful character, limited only by
the difficulty of obtaining complete speci-

mens. Species are characterized by having
two anal cirri (Fig. 3b, f), four anal cirri (Fig.
3c, d), or two swollen pads lacking cirri (Fig.
Base):

Setae.—The most important point to
emerge from SEM examination of the setae
from the specimens studied here is that the
“*hooded”’ ventral seta in medial and pos-
terior parapodia used to define A. mutans
is not hooded (in the sense of having a three
dimensional structure enclosing the distal
end of the seta). In all specimens examined,
the distal end of the ventral seta consists of
a relatively flat extension of the setal shaft.

Although the diameter of the ventral seta
is generally twice that of the other setae in
these forms, it originates in the same mus-
cular bundle, and is not an acicula as re-
corded by some authors. The modification
of the distal end of the ventral seta is dif-
ferent only in degree, not in structure, from
that observed in the ventral setae of Ara-
bella iricolor. This observation strongly sup-
ports Crossland’s (1924) contention that
Cenothrix Chamberlin should be regarded
as a junior synonym of Arabella.

The terminology adopted here to describe
the ventral setae is illustrated in Fig. 1. Three
distinct morphologies are recognized. The
shaft of the seta may be smooth with an
abrupt taper from the shaft to the guard,
ridged with an abrupt taper from the shaft
to the guard, or ridged with a gradual taper

286
shaft
Pee SE
ridge
Fig. 1.

from the shaft to the guard. Arabella pan-
amensis, n. sp. has ventral setae which are
somewhat modified from the second type
above in having an additional ridge near the
tip of the shaft (Fig. 3h).

SEM examination of the other setae re-
veals that the distinction between smooth
and “‘toothed”’ geniculate setae made by both
Chamberlin (1919) and Crossland (1924) is
indeed significant. The “‘toothed”’ setae are
not actually toothed, however, but are in-
stead ringed with loose rows of large fibrils,
in contrast to the tight, irregular arrange-
ment of fibrils in the smooth setae (Fig. 4k,
1).

Jaws. — Maxillary jaws in eunicoid poly-
chaetes are numbered, by convention, from
posterior to anterior with roman numerals.
Left and right jaws are indicated by a capital
L or R respectively. Thus MIR refers to the
posterior maxilla on the right side of the
apparatus. Denticle formulas are indicated
in sequence, with the left jaw in a given pair
listed first. For populations, the low end of
the range of denticles for a given jaw is listed
first, with the high end of the range in pa-
rentheses.

Maxillae within some of the species are
here considered polymorphic, but variation
within single populations is limited to two
or three morphs. The first and second max-
illae (MI and MII) were found to be the most
significant in differentiating species. The
morphs recognized here are illustrated in
Fig. 2. The MII jaws may be either “‘long,”’

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ao ert

guard

Terminology used to describe morphology of the ventral seta in medial and posterior parapodia.

extending posteriorly to the base of the MI,
or “‘short,’’ extending posteriorly only half
of the distance to the base of the MI (Fig.
2a, b). A long MII is invariably associated
with a relatively slender MI (here termed
“‘sracile’’) with a single falx (or hook) and a
pronounced gap between the falx and the
basal denticle row (Fig. 2e). MIs adjacent
to short MIIs are wider and generally slight-
ly longer than the gracile MIs, and are here
termed “‘robust.’’ They may take one of three
forms in adult worms. These jaws may have
a single falx with pronounced gap (=“‘fal-
cate,’ Fig. 2c), a bifid falx with pronounced
gap (=“‘bifid,’’ Fig. 2d), or no distinct falx
and no gap (=“‘dentate,”’ Fig. 2f). Dentate
MIs are found in juvenile specimens of a
number of different species, but in the ju-
veniles with dentate MIs the anterior den-
ticles project laterally rather than dorsally
as in the adults (compare Fig. 4c, d).

The statistical frequency of these jaw
morphologies varies systematically among
different populations of worms, and in most
cases only one of the three morphologies of
robust MIs is represented (if any). The pres-
ence or absence of jaw symmetry polymor-
phism within a population is also a useful
taxonomic character.

Within the species considered here, the
lengths of MIs are highly correlated with the
sizes of individual worms as measured by
prostomial width (Fig. 7). A plot of MI length
against prostomial width is similar for most
of these species, with the exception of Ar-

VOLUME 102, NUMBER 2

4
ial

287

e f

Fig. 2. Schematic representation of morphologies observed in first and second maxillae. a, short MII; b, long
MII; c, falcate robust MI; d, bifid robust MI; e, gracile falcate MI; f, dentate robust MI.

abella panamensis. In A. panamensis indi-
viduals have relatively large jaws for their
size.

The number of denticles on a given max-
illa is variable, and has generally been dis-
counted as a useful taxonomic character in
eunicoid polychaetes (Kielan-Jaworowska
1966, Wolf 1980). With sufficient speci-
mens, however, it may be possible to use
the number of denticles on a statistical basis
(see below under Arabella multidentata).

The shape of the mandibles exhibits with-
in-sample variation in these forms (Cross-
land 1924), but proved of no value in dis-
tinguishing species. A typical mandible is
illustrated in Fig. 5b.

Order Eunicida Fauchald, 1977
Family Arabellidae Hartman, 1944
Genus Arabella Grube, 1850

Arabella Grube, 1850:17.
Notopsilus Ehlers, 1868:406.
Maclovia Grube, 1871:58.
Cenothrix Chamberlin, 1919:329.

Type species. — Arabella iricolor (Monta-
gu, 1804).

Remarks. —Fauchald (1977b:110-11)
noted that generic diagnoses within the Ar-

abellidae are inconsistent. In forms with
projecting acicular spines, the shape of the
ML is used to separate the genus Drilonereis
Claparede (falcate MIs) from WNotocirrus
Schmarda (dentate MIs), while the genus
Arabella (which lacks emergent acicular
spines) includes forms with both maxillary
morphologies. Fauchald suggested that
either the genus Notopsilus Ehlers be rec-
ognized to include species with dentate MIs
which are presently placed in Arabella, or
that Drilonereis and Notocirrus be consid-
ered synonymous.

The report here of species in which the
jaws are polymorphic (including both den-
tate and falcate MIs) strongly suggests that
Notopsilus should be considered a junior
synonym of Arabella. This does not imply,
however, that Drilonereis and Notocirrus
should also be considered synonymous. The
basic arrangement of jaws is quite different
in Drilonereis than it is in Notocirrus, and
intermediate or hybrid maxillae have not
been recorded.

Arabella iricolor (Montagu, 1804)
Figs. 3b, 4j, k, 5a, 6a

Nereis iricolor Montagu, 1804.
Arabella iricolor. — Pettibone, 1963:269, fig.

288

ee

Big. 3.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

al

h

a, Arabella logani, holotype, ZK 1924.3.1.15, pygidium, left-lateral view; b, Arabella iricolor, USNM

61732, pygidium, right-lateral view; c, Arabella novecrinita, lectotype, ZK 1924.3.1.15, pygidium, dorsal view;
d, Arabella mutans, USNM 57434, pygidium, oblique right-lateral view; e, Arabella monroi, holotype, ZK
1932.12.24.595, pygidium, right-lateral view; f-—h, Arabella panamensis, USNM 98237, holotype; f, pygidium,
dorsal oblique right-lateral view; g, posterior parapodium, anterior view, note elongate post-setal lobe; h, ventral
seta of parapodium in g, note 2 ridges, abrupt taper of shaft; i, Arabella sp. A, USNM 54915, anterior end with
pharynx extended, lateral view, note nuchal papillae. Scale bars on a-f, i = 0.5 mm; g = 0.2 mm; h = 20 um.

7la—e; Gardiner, 1976:206-—207, figs.

27r—u, 28a; Uebelacker, 1984:42-5, fig.

42-2.

Materials examined. —USNM 30058,
FSBC I 24159, Seashore Key, Florida (6);
FSBCI 13436, Terra Ceia Bay, Florida (1).—
FSBC I 13102, Lower Tampa Bay, Florida
(1).—EJ-72-66, EJ-73-414, Hutchinson Is-
land, Florida (3).—USNM 61732, USNM
53026, Cape Lookout, North Carolina
(11).—USNM 4676, Woods Hole, Massa-
chusetts (7).—USNM 33787, Province-
town, Massachusetts (8).

Remarks.—The specimens of Arabella
iricolor examined here are characterized by
having ventral setae in medial and posterior
parapodia which are ridged, with a gradu-
ally tapering shaft (Fig. 4j). The pygidium
has two cirri which arise dorsally (Fig. 3b).
No variation in jaw symmetry was observed
among adult specimens. In each individual
the MIL is robust with a short MIIL, the
MIR gracile with a long MIIR.

The Florida specimens differ from those
from North Carolina and Massachusetts in
that all adult worms have falcate robust MIs

VOLUME 102, NUMBER 2 289

Fig. 4. Setae, all in anterior view. a, b, Arabella monroi, ZK 1932.12.24.337-340, paratype, large anterior
fragment, ventral seta, setiger 143, note ridge, abrupt taper of shaft. c, Arabella novecrinita, ZK 1924.3.1.15,
lectotype, ventral seta, setiger 200, note smooth, abruptly tapered shaft. d, Arabella mutans, USNM 23933,
ventral seta, posterior setiger, note smooth, abruptly tapered shaft. e, Arabella mutans, USNM 57434, ventral
seta, posterior setiger, dorsal to bottom of page, note smooth, abruptly tapered shaft. f, Arabella atlantica, ZK
1923.11.5.1-2, large syntype, ventral seta, setiger 122, note ridge, abrupt taper. g, h, Arabella logani, ZK
1924.3.1.15, holotype, ventral seta, setiger 125, note ridge, gradually tapered shaft. i, Arabella multidentata,
USNM 98154, ventral seta, posterior setiger, note ridge, abruptly tapered shaft. j, k, Arabella iricolor, USNM
30058 (Florida); j, ventral seta, posterior setiger, note ridge, gradually tapered shaft; k, ““smooth” seta, posterior
setiger. 1, Arabella sp. A, USNM 54915, posterior setiger, smooth seta above, toothed seta below. All scale bars
= 5 um.

290

(n = 8), in contrast to the bifid robust mor-
phology (Fig. 5a) observed in the northern
populations (25 of 26 adult specimens).

Jaw morphology in these populations is
quite stable, as the jaws are essentially iden-
tical in specimens collected from Woods
Hole in 1883 (USNM 4676), Provincetown
in 1954 (USNM 33787), and North Caro-
lina in 1974 (USNM 53026) and 1975
(USNM 61732). Thus although the differ-
ence between the Florida and northern
populations is small, the stability of the
character suggests that the difference is he-
reditary, and may be used to subdivide east
coast populations of A. iricolor. As presently
recognized, A. iricolor is probably a species
complex.

Two juvenile specimens of A. iricolor from
Florida have dentate MILs which resemble
those in several other species of Arabella,
including A. mutans (Fig. 5d).

Arabella atlantica Crossland, 1924,
new rank
Fig. 4f

Arabella novecrinita var. atlantica Cross-
land, 1924:78—80, text-figs. 99-101, 105.

Arabella novecrinita var. asymmetrica
Crossland, 1924:80-83, text-figs. 102-
104.

Syntypes. —ZK 1923.11.5.1-2, Cape
Verde Islands, two anterior fragments.

Materials examined.—ZK 1923.11.5.1-
2, Cape Verde Islands (two anterior frag-
ments).—jaw mounts from six specimens
from the Cape Verde Islands as follows: ZK
1923.11.8.12-14, three unregistered slides
labelled CV Isl. W9; three unregistered jaw
mounts of Arabella novecrinita var. asym-
metrica, Cape Verde Islands.

Remarks. —Crossland (1924:78) record-
ed 11 specimens from the Cape Verde Is-
lands. Three small specimens which exhibit
maxillae typical of juveniles in the genus
Arabella were assigned to Arabella nove-
crinita var. asymmetrica (Crossland, 1924,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

text-fig. 103). These specimens are here re-
garded as juveniles of Arabella atlantica.

Of the remaining eight specimens, two
anterior fragments and six sets of jaws are
represented in the collection of the British
Museum (Natural History). At present it is
impossible to determine which sets of jaws
belong to the two preserved specimens, and
thus no lectotype is designated.

The soft-part morphology of this species
is adequately described by Crossland (1924:
78-80, text-fig. 99). The ventral setae are
ridged, with an abrupt taper from the guard
to the shaft (Fig. 4f). In the specimens avail-
able for examination, the MIL is robust fal-
cate, the MIIL short, and the MIR gracile
falcate with a long MIIR. No symmetry
polymorphism observed (n = 6).

This species is distinguished from Ara-
bella novecrinita by its ridged ventral setae
and robust MIL. It differs from A. multi-
dentata in having falcate rather than bifid
robust MIs, and in not exhibiting symmetry
polymorphism. A. atlantica most closely re-
sembles A. monroi n. sp. from the Gala-
pagos, but because neither preserved spec-
imen of A. atlantica has an intact pygidium,
it cannot be compared based on this im-
portant character.

Arabella logani Crossland, 1924, new rank
Figs. 3a, 4g, h

Arabella novecrinita var. logani Crossland,
1924:75-78, text-figs. 96-98.

Holotype. —ZK 1924.3.1.138, Red Sea.

Materials examined. —ZK 1924.3.1.138,
Red Sea (1).

Remarks. —The soft-part morphology of
the type and only specimen is accurately
described by Crossland (1924:75-76).
Noteworthy is the presence of two inflated
pads on the pygidium (Fig. 3a) which lack
anal cirri. No evidence for incomplete re-
generation is apparent, and this may rep-
resent the fully developed character state.

The ventral setae in posterior parapodia

VOLUME 102, NUMBER 2 291

Fig. 5. a, Arabella iricolor, USNM 33787, maxillae, dorsal view, note bifid MIL. b—-d, Arabella mutans,
USNM 23933; b, mandibles, ventral view; c, maxillae of adult specimen, dorsal view, note dentate MIR, falcate
tip of MIL broken during mounting; d, maxillae of juvenile specimen, note dentate MIL with anterior denticles
which project laterally. e, Arabella sp. A, USNM 54915, maxillae, dorsal view, example of symmetrical apparatus
with 2 long MIIs, 2 gracile falcate MIs. f, Arabella panamensis, USNM 66043, paratype, maxillae, dorsal view,
note falcate MIL, falcate tip of MIR broken during mounting. Scale bars on a-c, e, f = 100 um; d = 50 um.

292

are ridged and gradually tapering (Fig. 4g,
h), and are thus unlike those in other species
within the Arabella mutans complex.

The MIL is robust bifid, the MIIL short,
and the MIR gracile falcate with a long MIIR
(note that Crossland reversed the image in
text-fig. 96, and incorrectly labeled the spec-
imen as a “Cape Verde specimen’’). The
bifid MIL serves to distinguish this species
from Arabella iricolor, which in the eastern
Mediterranean and Red Sea has a falcate
robust MIL (Ben-Eliahu 1976:172). No
statement concerning jaw variability can be
made based on a single specimen, and full
characterization of the species must await
further collecting.

Arabella monroi, new species
Figs. 3e, 4a, b

Arabella mutans. —Monro 1933b:88-89 (in
part).

Holotype.—ZK 1932.12.24.595, washed
from peat, Isla Santa Cruz (Indefatigable),
Galapagos Islands.

Etymology. —In honor of C. C. A. Monro.

Materials examined. —Holotype; ZK
1932.12.337—340 (paratypes), washed from
peat, Isla Santa Cruz (Indefatigable), Ga-
lapagos Islands (26).—ZK 1932.12.336, fine
gravel in pools, Isla Isabela (Albermarle),
Galapagos Islands (1).—ZK 1932.12.341,
Post Office Bay, Isla Santa Maria (Charles),
Galapagos Islands (3).

Description. —Prostomium bullet-shaped,
slightly longer than wide to slightly wider
than long, V-shaped furrow on ventral sur-
face, four eyes embedded in posterior dorsal
margin. Peristomium divided into two rings,
a shallow ventral median furrow in anterior
ring only.

Setigers numerous, exceeding 200 in large,
adult specimens. Notopodia represented by
small tubercle, neuropodia with blunt, sub-
triangular postsetal lobe. Pygidium with two
swollen pads (Fig. 3e), no anal cirri ob-
served.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Notoacicula small, 2—3 per setiger. Neu-
roacicula large, amber colored, 3—4 per se-
tiger with tips protruding slightly through
parapodial wall. Anterior setigers with 5-6
setae, 3 being smooth, 2—3 toothed. Medial
and posterior setigers with 2 smooth, 1
toothed, and 1 ventral seta. Ventral setae
ridged, shaft tapering abruptly to distal guard
(Fig. 4a, b).

Mandibles H-shaped, typical of Arabella.
Maxillae with 2 long carriers and unpaired
ventral ligament (=“‘third carrier” of some
authors). Maxillary apparatus asymmetri-
cal, no symmetry polymorphism observed.
MIL robust falcate, MIIL short, MIR gracile
falcate, MIIR long. Denticle formula: 6(9)-
6(8); 7(8)-12(16); 5(6)-5(6); 4(5)-4; 1-1.

Holotype large, complete individual with
208 setigers, prostomium 0.68 mm long,
0.55 mm wide, peristomium 0.54 mm long,
MIL 0.36 mm long, MIR 0.35 mm long,
denticle formula 9-8; 8-16; 6-6; 4-4; 1-1.

Remarks.—The collection from the Ga-
lapagos includes 13 complete specimens.
None have anal cirri, and it seems likely
that the presence of two pygidial lobes with
no cirri represents the fully developed adult
character state.

Arabella monroi is distinguished from A.
mutans based on its asymmetrical maxillae
which are not polymorphic, falcate rather
than dentate robust MIs, ridged rather than
smooth ventral setae, and by the absence of
anal cirri. It differs from A. multidentata in
having falcate rather than bifid robust MIs,
in not exhibiting symmetry polymorphism
of the maxillae, and in lacking anal cirri.

The maxillary apparatus in A. monroi is
identical to that in A. atlantica. The pygid-
ium in the latter species is unknown, how-
ever, and I consider it prudent to describe
the Galapagos specimens as a separate
species pending collection of additional ma-
terial from the type locality of A. atlantica.

The geographic range of A. monrolis pres-
ently restricted to the islands of the Gala-
pagos.

VOLUME 102, NUMBER 2

Arabella multidentata (Ehlers, 1887)
Figs. 41, 6b, 7

Aracoda multidentata Ehlers, 1887:112, pl.
34, figs. 8-10, pl. 35, figs. 1-4.

Arabella multidentata. —Hartman, 1938:12;
Perkins, 1979:447-449, fig. 13; Uebe-
lacker, 1984:42-9, figs. 42-5, 6a-g.

Arabella maculosa Verrill, 1900:651.—
Treadwell, 1921:114, pl. 9, figs. 12-13,
text-figs. 424-428; Perkins, 1979:450, fig.
14.

Arabella mutans. —Gardiner, 1976:206, figs.
270—q; Fauchald, 1977a:45 (in part); Per-
kins, 1979:445-447, fig. 12; ?Duenas,
1981:193.

Arabella novecrinita. —Monro, 1933a:260-
261:

?Arabella asymmetrica.—Orensanz, 1974:
368-388, fig. III-6.

Holotype. —MCZ 825, Florida.

Materials examined. -USNM 55882,
Gulf of Mexico (1).—USNM 54694-54696,
Hutchinson Island, Florida (3).—USNM
59526, N Atlantic off Florida (2); USNM
59528, N Atlantic off N Carolina (1).—
USNM 56844, N Atlantic off Virginia (1).—
USNM 98142-98156, Bermuda (22).—
USNM 53297, 54215, 54220, 54221, 54223,
54818, 54820, 98238, Guayanilla Bay,
Puerto Rico (22).—USNM 54217, 54218,
54222, 54837-54840, Jobos Bay, Puerto
Rico (8).—USNM 66044-66046, 73653,
Galeta Reef, Panama (5).—?USNM 58273,
Bay of Cartagena, Colombia (1).

Remarks.—Perkins (1979:448-449) ac-
curately described the soft-part morphology
of A. multidentata. To his description of the
setae I add that the ventral setae are ridged
and abruptly tapered (Fig. 41).

Perkins (1979:450) distinguished Arabel-
la maculosa Verrill, 1900 from A. multi-
dentata based on the “more obviously
asymmetrical’’ ventral setae in the syntype
of the former. I found the degree of asym-
metry variable on a single specimen, and
was unable to separate the species on this

293

Table 2.— Distribution of specimens of Arabella with
particular MI morphologies in polymorphic species.

1 gracile,

Species 2 robust 1 robust 2 gracile
multidentata
Bermuda 1 9 8
Others 5 15 12
mutans 4 4 2
panamensis 3 2, _
sp. A — 4 2

basis. The syntype of A. maculosa has a
regenerated pygidium with two poorly de-
veloped lobes and no cirri (Perkins 1979,
fig. 14a). Eight complete specimens from
Bermuda, with no indication of posterior
regeneration, have four anal cirri as 1s typ-
ical of A. multidentata. The jaws of the Ber-
muda specimens are indistinguishable from
those of other west Atlantic and Caribbean
populations of A. multidentata (Table 2),
and I therefore consider A. maculosa a ju-
nior synonym of A. multidentata.

Perkins (1979:447) distinguished sym-
patric Florida populations of A. multiden-
tata from A. mutans based on jaw sym-
metry. Instead, I recognize a single species
(A. multidentata) with polymorphic jaw
symmetry which is distinct from A. mutans.
A. multidentata has bifid (25 of 27 adult
specimens with robust elements, remaining
2 falcate) rather than dentate robust MI ele-
ments (where present), and further differs
from A. mutans in having ridged rather than
smooth ventral setae.

A comparison between A. multidentata
and A. iricolor sensu Pettibone (1963) is par-
ticularly instructive, as the geographic ranges
of the two species overlap along the east
coast of the United States. A. multidentata
has abruptly tapering shafts on the ventral
setae (Fig. 41), rather than gradually tapering
shafts as in A. iricolor (Fig. 4j), 1s further
characterized by having four rather than two
anal cirri, and by having polymorphic max-
illary symmetry. The number of denticles

294

30:..=

7s

20 a

US: =

Number of Jaws

@) — Saco aeecccenaeaaaeoe renee =

Gn? BLL oulO LAL 11s 1 S16

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

6 1728

Number of Denticles

Fig. 6. Histograms of denticle number in MII jaws in Arabella iricolor (a) and Arabella multidentata (b).
Note that the distinction between short and long MII elements is evident from the diagram. Means are indicated

for long MIIs.

in long MII elements overlaps between the
two species (Fig. 6a, b), but the difference
in means is highly significant (P < 0.001, t
= 6.14).

Dentate jaws were observed in four ju-
venile specimens of A. multidentata. The
juvenile dentition is indistinguishable from
that in other species of Arabella.

As defined here, Arabella multidentata is
a west Atlantic species recorded from the
east coast of the United States as far north
as Virginia, from Bermuda, Puerto Rico, the
Gulf of Mexico, Atlantic Panama, and ques-
tionably from the Bay of Cartagena, Colom-
bia (single specimen with falcate rather than
bifid robust MIL). The specimens described
by Orensanz (1974) as A. asymmetrica have
maxillae identical to the present material,
and ventral setae with abruptly tapered
shafts. It’s unclear whether or not the ven-
tral setae are ridged, however, and the py-
gidium was undescribed. This attribution
must therefore be considered tentative.

Arabella mutans (Chamberlin, 1919)
Figs. 3d, 4d, e, 5b—-d

Cenothrix mutans Chamberlin, 1919:330-
332, pl. 61, figs. 2-9, pl. 62, fig. 1.

Arabella mutans. —Crossland, 1924:71;
Hartman, 1954:624; Reish, 1968:220;
Kohn and Lloyd, 1973:699.

Arabella novecrinita. —Hartman, 1954:625.

Holotype. —USNM 19740, Easter Island.

Materials examined. —-USNM 19740,
57434, Easter Island (2).—USNM 23933-
23935, Bikini Island (14).

Remarks. —Chamberlin’s (1919:330, pl.
61, fig. 4) description of the anterior end of
the holotype is accurate. The pygidium is
poorly developed, and represents incom-
plete regeneration. A second specimen from
Easter Island (USNM 57434) has four anal
cirri (Fig. 3d), as do two specimens from
Bikini, and this is considered the fully de-
veloped character state.

As noted above, the hooded acicula (or

VOLUME 102, NUMBER 2

‘““crochets”) which Chamberlin (1919:331)
originally described for this species are neli-
ther hooded nor acicula, but rather are setae
typical of the genus Arabella which have an
abruptly tapering shaft. The shaft is smooth
on ventral setae in the holotype, and on the
other specimens examined (Fig. 4d, e).

Chamberlin (1919:61, fig. 2) inaccurately
figured the MIs in the holotype. The MIL
is robust dentate, with two upturned ante-
rior denticles (as in MIR in Fig. 5c) which
were not depicted by Chamberlin. The MIR
is gracile falcate with the anterior tip broken
off. Examination of additional specimens
reveals that maxillary symmetry is poly-
morphic (Table 2), and that if robust MIs
are present they are dentate (Fig. 5c). Two
juvenile specimens were observed with den-
tate MIs of the type illustrated in Fig. 5d.
Mandibles (Fig. 5b) are as described by
Chamberlin (1919:332, pl. 61, fig. 3).

Arabella mutans most closely resembles
A. novecrinita as restricted here. In contrast
to the symmetry polymorphism exhibited
by A. mutans, A. novecrinita is characterized
by symmetrical maxillae with gracile MIs
and long MIIs. In view of the small number
of A. novecrinita included here (n = 4), this
distinction must be considered tentative. A.
mutans has more denticles on long MIIs on
average (13.3, n = 12) than does A. nove-
crinita (10.6, n = 5), but again the small
sample size makes use of this distinction
tentative.

As restricted here, A. mutans has been
recorded from Easter Island and the Mar-
shall Islands in the tropical Pacific. A spec-
imen from Guam (USNM 54426), which
had been identified as A. mutans, 1s now
Missing its jaws, and the identification could
not be confirmed.

Day’s (1975:196-197) report of A. mu-
tans from Western Australia was based on
a single juvenile specimen, and must be con-
sidered questionable on that basis. The jaws
illustrated by Fauvel (1953:275, fig. 1401)
from a small specimen collected from Cey-

295

lon are also those of a juvenile, and the
record cannot be accepted. Tampi & Ran-
garajan (1964:11, figs. 29-31) did not de-
scribe the jaws in the anterior fragment they
identified as A. mutans from Car Nicobar,
Indian Ocean, and their record must also
be considered questionable. Silva (1961:182)
described but did not illustrate a specimen
from Ceylon as A. mutans which has two
“hooded acicula’” per parapodium, and
asymmetrical maxillae with MIL robust fal-
cate, MIIL short, MIR gracile falcate, and
MIIR long. The presence of two “‘hooded”’
setae per parapodium is unusual, and the
maxillae are unlike those in either A. mu-
tans or A. novecrinita.

Arabella novecrinita Crossland, 1924
Figs. 3c, 4c

Arabella novecrinita Crossland, 1924:71-75,
text-figs. 89-95.

Lectotype.—ZK 1924.3.1.15, Zanzibar;
designated herein.

Materials examined.—ZK 1924.3.1.15,
Zanzibar (1).—ZK 1924.3.1.292, Minikoi
Island, Maldives, mounted jaws and para-

podia (1?).—uncatalogued anterior frag-

ment, Aldabra.

Remarks. —Crossland (1924:71) record-
ed two specimens (with maxillary formulas)
from the Maldives. The British Museum
(Natural History) collection includes only
mounted jaws and parapodia from this lo-
cation which may be from a single speci-
men. In light of the fragmentary nature of
the Maldive material, and the wide geo-
graphic separation between Crossland’s
original type localities, the Zanzibar speci-
men is herein designated the lectotype.

Crossland’s (1924:7 1-75, text-fig. 89) de-
scription of the soft-part morphology in this
species is accurate. Noteworthy is the pres-
ence of four anal cirri (Fig. 3c) on the lec-
totype (Crossland 1924:75). The ventral se-
tae in medial and posterior parapodia have

296

a smooth shaft which tapers abruptly to the
guard (Fig. 4c). The maxillae on the lecto-
type are symmetrical, with gracile MIs and
long MIIs (Crossland 1924:72), as are the
jaws on the two specimens Crossland (1924:
73) recorded from the Maldives, and the
anterior fragment recorded here from Al-
dabra. The jaw formula for the lectotype is
7-7; 7+-10; 5-5; 5-4; 1-1. For the species it
is 6(7)-7; 10(11)-10(1 1); 4(6)-4(5); 4(5)-3(4);
1-1. The mandibles are typical of Arabella,
as illustrated by Crossland (1924, text-fig.
91).

As noted above, the setae and soft-part
characteristics of this species are similar to
those in Arabella mutans, and the species
are distinguished based on the maxillae.

As restricted here, A. novecrinita is known
only from islands in the Indian Ocean. Day
(1967:446, fig. 17.18fh) recorded speci-
mens (as A. mutans) from Mozambique and
South Africa which may belong in A. no-
vecrinita based on the illustrated maxillary
apparatus. It’s not clear whether the ventral
setae are ridged based on Day’s (1967, fig.
17.18f) illustration, and thus the report can-
not be securely credited to A. novecrinita.
A South African specimen identified as A.
mutans in the British Museum collection
(ZK 1932.11.24.38) proved to be a mis-
identified specimen of Arabella iricolor sen-
su lato, and thus this question remains un-
resolved.

Arabella panamensis, new species
Figs. 3f—g, 5f, 7

Arabella mutans. —Fauchald, 1977a:45 (Gn
part).

Holotype. —USNM 98237, Paitilla Beach,
Panama.

Etymology. —From Panama, the type lo-
cality.

Materials examined. —Holotype; USNM
66042, 66043, 66365 (paratypes), Paitilla
Beach, Panama (9).—?USNM 80225, Golfo
de Nicoya, Costa Rica (1).

Description. —Prostomium bullet-shaped,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

slightly wider than long, V-shaped mark on
ventral surface, four eyes embedded in pos-
terior dorsal margin. Peristomium divided
into two rings, ventral median furrow in
anterior ring only.

Setigers numerous, up to 177 in complete
adults. Notopodia not expressed on surface,
neuropodia with elongate postsetal lobe (Fig.
3g) which is commonly pigmented. Pygid-
ium with two ventral anal cirri (Fig. 3f).

Notoacicula fine, from two to three per
setiger. Neuroacicula large, amber colored,
three per setiger, tips commonly protruding
slightly through parapodial wall. Anterior
setigers with three smooth and from two to
three toothed setae, medial and posterior
setigers with two smooth, from one to two
toothed, and one ventral seta. Shafts on
ventral setae taper abruptly to guard, two
ridges present (Fig. 3h).

Mandibles H-shaped. Maxillae with two
long carriers and unpaired ventral ligament.
Maxillae exhibit symmetry polymorphism.
One or both MI elements robust falcate (Fig.
5f). Denticle formula: 5(8)-6(9); 6(7) or
9-5(7) or 11(12); 4(6)-4(5); 3(4)-3(4); 1-1.

Holotype large, complete specimen with
177 setigers, prostomium 0.39 mm long,
0.70 mm wide, peristomium 0.39 mm long,
MIL 0.41 mm long, MIR 0.41 mm long,
denticle formula 7-9; 6-11; 6-5; 4-4; 1-1.

Remarks. — This species differs from oth-
ers in the A. mutans complex in having long-
er postsetal lobes on medial and posterior
parapodia, two anal cirri which arise ven-
trally, and in having two ridges on the ven-
tral setae. It further differs in having some-
what longer MIs in individuals of a given
size (Fig. 7). The post-setal lobe in Arabella
semimaculata (Moore, 1911) resembles that
in A. panamensis (compare Fig. 3g; Fau-
chald 1970, pl. 20, fig. g), but the former
species has smoothly bilimbate ventral se-
tae (Fauchald 1970:133), unlike the ridged,
abruptly tapering ventral setae in A. pana-
menslis.

A juvenile specimen from the Pacific side
of Costa Rica (USNM 80225) with dentate

VOLUME 102, NUMBER 2

Length of Robust MI (mm)

0 01 0-2 0-3
Width of Prostomium (mm)

Da |

0-4 0-5 0-6 0-7 0:8

Fig. 7. Plot of robust MI length against width of prostomium for Arabella multidentata (@), Arabella pana-
mensis (x), and Arabella sp. A (*). Prostomial width is less subject to distortion by fixation than is total length,
and is used here as a more reliable measure of specimen size (and, by inference, maturity). The spread of points
for A. multidentata is typical of that observed for the other species considered here with the exception of A.
panamensis. The regression line for A. multidentata is of the form y = 0.55x — 0.44 (r = 0.94). Note that all
of the points for A. panamensis plot above the regression line, suggesting that individuals of A. panamensis have
larger jaws for their size than do individuals of A. multidentata. The slope of a linear regression for A. panamensis
(not shown) is significantly different from the slope for A. multidentata (P = 0.005, t = 3.04, d.f. = 35). Note
also that plots of points for papillate individuals (Arabella sp. A) fall within the large end of the range of variation

for A. multidentata.

MIR has setae comparable to A. panamen-
sis, but lacks the elongate postsetal lobe in
posterior parapodia. No data is available on
ontogenetic changes to be expected in this
character, and the specimen is only tenta-
tively attributed to A. panamensis.

The species 1s presently known only from
the Pacific coast of Panama.

Arabella sp. A
Figs. 31, 41, S5e

Materials examined. —USNM 54216,
54915, 98239, Guayanilla Bay, Puerto Rico
(8).

Remarks. —Three collections of Arabella
multidentata from Guayanilla Bay included
several relatively large individuals with two
distinctive papillae arising from the nuchal

fold (Fig. 31). The jaws (Figs. 5e, 7) and setae
are comparable to those in A. multidentata.
No papillate specimens were recorded from
any other locality, and the significance of
the papillae is presently unclear. The char-
acter is sufficiently distinctive to warrant
segregation of these forms (informally as sp.
A) pending further investigation.

Discussion

The systematic treatment here afforded
members of the Arabella mutans species
complex should be regarded as a working
hypothesis. The stability of the taxonomic
characters employed can be evaluated by
the collection of additional specimens, and
data on additional characters (using electro-
phoresis, for example) can be collected for

298

comparative purposes. Studies on the
breeding interactions in living populations
would, of course, provide the most powerful
test of the systematics, but at the same time
may be the most challenging to implement.

Particular emphasis is placed here on
characterizing species based on populations
rather than single individuals. Thus, poly-
morphism of jaw symmetry is considered a
useful taxonomic character, even though it
is Meaningless in regard to a single individ-
ual. A single individual collected from a new
location may thus not be identifiable to the
species level.

This restriction especially applies to ju-
venile individuals. Similar maxillae have
been recorded from juveniles in four (ques-
tionably five) of the species of Arabella con-
sidered here, although the adult maxillae in
these forms exhibit significant differences.

Studies of other species of Arabella should
consider the maturity of specimens before
maxillary form is included in the species
diagnosis. In particular, the distinction
commonly made between Arabella iricolor
and A. geniculata (Claparede, 1868) might
be worth re-evaluation in this light (Fauvel
1923:439-440, fig. 1751-1; Fauchald 1970:
226; Ben-Eliahu 1976:172).

As restricted here, two of the species con-
sidered are incompletely characterized (Ar-
abella atlantica, A. logani). The remaining
five species are separated geographically as
well as morphologically. The concept of A.
mutans as a single circumtropical species 1s
clearly incorrect. This observation serves to
underline the futility inherent in attempting
to use the distribution of rare, poorly under-
stood species for the study of biogeography.

The chief lessons for paleontologists
studying fossil polychaete jaws are as fol-
lows: 1) jaw symmetry may vary within a
single population, and should be used as a
taxonomic character with caution, 2) the
morphologies of the MI and MII maxillary
elements may be useful for species level di-
agnoses, 3) the denticles on jaws may be
worth counting after all, but denticle num-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ber should only be used as a taxonomic
character on a population basis. The second
point has already been deduced from the
study of fossil material (Kielan-Jaworowska
1966; Edgar 1984), suggesting that classifi-
cation of recent and fossil material may be
roughly comparable. The first point has been
less fully appreciated, and some revision of
generic and family level fossil taxa may be
necessary on that basis.

Acknowledgments

I thank Kristian Fauchald and Marian H.
Pettibone for informative discussions con-
cerning polychaete systematics, and for crit-
ically reading the manuscript. Alex I. Muir,
British Museum (Natural History), and
Thomas H. Perkins, Florida Department of
Natural Resources, generously arranged the
loan of type and comparative material. I
also thank numerous members of the staff
of the U.S. National Museum of Natural
History, Smithsonian Institution, for their
cooperation in the completion of this study.
Joanne Simpson drafted Figs. 1, 2, 6, and
7. This research was supported by a post-
doctoral fellowship, Smithsonian Institu-
tion.

Literature Cited

Ben-Eliahu, M. N. 1976. Errant polychaete crypto-
fauna (excluding Syllidae and Nereidae) from
rims of similar intertidal vermetid reefs on the
Mediterranean coast of Israel and the Gulf of
Elat.—Israel Journal of Zoology 25:156-177.

Chamberlin, R. V. 1919. The Annelida Polychaeta. —
Memoirs of the Museum of Comparative Zo-
ology, Harvard 48:1-514.

Crossland, C. 1924. Polychaeta of tropical East Af-
rica, the Red Sea and Cape Verde Islands. The
Lumbriconereidae and Staurocephalidae.—
Proceedings of the Zoological Society of London
1924:1-106.

Day. J. H. 1967. A monograph on the Polychaeta of

southern Africa, part I. Errantia.— British Mu-

seum (Natural History), London, 458 p.

1975. On a collection of Polychaeta from
intertidal and shallow reefs near Perth, Western
Australia.—Records of the Western Australia
Museum (1975) 3:167—208.

VOLUME 102, NUMBER 2

Duenas, P. R. 1981. Inventario preliminar de los
poliquetos (Annelida) de aguas de la bahia de
Cartagena y areas adyacentes.—Boletin Museo
del Mar 10:82-138.

Edgar, D.R. 1984. Polychaetes of the lower and mid-
dle Paleozoic: a multi-element analysis and a
phylogenetic outline. — Review of Palaeobotany
and Palynology 43:255-—284.

Ehlers, E. 1864-1868. Die Bornstenwurmer (Anneli-

da, Chaetopoda). Leipzig, 748 p.

1887. Reports on the results of dredging in
the U.S. Coast steamer ‘Blake.’ XX XI. Report
on the annelids.—Memoirs of the Museum of
Comparative Zoology, Harvard 15:1-335.
Fauchald, K. 1970. Polychaetous annelids of the fam-
ilies Eunicidae, Lumbrineridae, Iphitimidae,
Arabellidae, Lysaretidae and Dorvilleidae from
western Mexico.— Allan Hancock Monographs
in Marine Biology 5:1-335.

1977a. Polychaetes from intertidal areas in
Panama, with a review of previous shallow-water
records.—Smithsonian Contributions in Zool-
ogy 221:1-81.

1977b. The polychaete worms.— Natural
History Museum of Los Angeles, Science Series
28:1-188.

Fauvel, P. 1923. Faune de France 5, polychétes er-

rantes. Office Central de Faunistique, Paris,

488 p.

. 1953. The fauna of India including Pakistan,

Ceylon, Burma and Malaya. Annelida Poly-

chaeta. Allahabad, 507 p.

Gardiner, S. L. 1976. Errant polychaete annelids from
North Carolina. — Journal of the Elisha Mitchell
Scientific Society 91:77—220.

Grube, A.-E. 1850. Die Familien der Anneliden.—

Archiv fur Naturgesch. Berlin 16.1:249-364.

1871. [untitled].—Jahresberichte der Schle-
siche Gesellschaft ftir Vaterlandische Kultur,
Breslau 49:56-58.

Hartman, O. 1938. Annotated list of the types of

polychaetous annelids in the Museum of Com-

parative Zoology.—Bulletin of the Museum of

Comparative Zoology, Harvard 85:1-31.

1944. Polychaetous annelids. Part V. Euni-
cea.—Allan Hancock Pacific Expedition 10:1-
339.

1954. Marine annelids from the northern
Marshall Islands.—U.S. Geological Survey
Professional Paper 260Q:615-644.
Kielan-Jaworowska, Z. 1966. Polychaete jaw appa-

ratuses from the Ordovician and Silurian of Po-
land and a comparison with modern forms.—
Paleontologica Polonica 16:1—152.

Kohn, A. J.,& M. C. Lloyd. 1973. Marine polychaete
annelids of Easter Island.—Internationale Re-
vue der Gestamten Hydrobiologie 58:69 1-712.

299

Monro, C. C. A. 1928. On the Polychaeta collected
by Dr. Th. Mortensen off the coast of Panama. —
Videnskabelige Meddelelser fra den Naturhis-
toriske Forening 1 Kjobenhavn 85:75-103.
1933a. On a collection of Polychaeta from

Dry Tortugas, Florida.—Annals and Magazine

of Natural History, London, series 10, 12:244—

269.

1933b. The Polychaeta Errantia collected by

Dr. C. Crossland at Colon, in the Panama Re-

gion, and the Galapagos Islands during the ex-

pedition of the S.Y. “‘St. George’’.— Proceedings

of the Zoological Society of London 1933:1-96.

Orensanz, J. M. 1974. Los anélidos poliquetos de la
Provincia Biogeografica Argentina. VI. Arabel-
lidae.—Physis, Seccion A 33:381-408.

Perkins, T. H. 1979. Lumbrineridae, Arabellidae, and
Dorvilleidae (Polychaeta), principally from
Florida, with descriptions of six new species. —
Proceedings of the Biological Society of Wash-
ington. 92:415-465.

Pettibone, M. H. 1963. Marine polychaete worms of
the New England Region. 1. Aphroditidae
through Trochochaetidae.— U.S. National Mu-
seum Bulletin 227:1-356.

Reish, D. J. 1968. The polychaetous annelids of the
Marshall Islands. — Pacific Science 22:208-231.

Silva, P. H. D. H. de. 1961. Contributions to the
knowledge of the polychaete fauna of Ceylon.
Part I. Five new species, two new varieties and
several new records principally from the south-
ern coast.—Spolia Zeylan 29:164-194.

Tampi, P. R. S., & K. Rangarajan. 1964. Some poly-
chaetous annelids from the Andaman waters. —
Journal of the Marine Biological Association of
India 6:98-121.

Treadwell, A. L. 1921. Leodicidae of the West Indian
region.— Publications of the Carnegie Institu-
tion, Washington 15:1-131.

Uebelacker, J. M. 1984. Family Arabellidae Hart-
man, 1944. Pp. 42-1—42-29 in J. M. Uebelacker
and P. G. Johnson, eds., Polychaetes of the
Northern Gulf of Mexico, Volume 6, Barry A.
Vittor & Associates, Mobile, Alabama.

Verrill, A. E. 1900. Additions to the Turbellaria,
Nemertina and Annelida of the Bermudas, with
revisions of some New England genera and
species.— Transactions of the Connecticut
Academy of Arts and Sciences 10:595-671.

Wolf, G. 1980. Morphologische Untersuchungen an
den Kieferapparaten einiger rezenter und fos-
siler Eunicoidea (Polychaeta). —Senckenber-
giana Maritima 12:1-182.

2195 Deborah Way, Upland, California
91786.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 300-304

A NEW SPECIES OF BENHAMIPOLYNOE
(POLYCHAETA: POLY NOIDAE: LEPIDASTHENIINAE)
FROM AUSTRALIA, ASSOCIATED WITH THE
UNATTACHED STYLASTERID CORAL
CONOPORA ADETA

Marian H. Pettibone

Abstract. — Benhamipolynoe cairnsi, n. sp. from deep-water (398 m) off
Queensland, Australia, was found in close symbiotic relationship with the
unusual, unattached stylasterid coral Conopora adeta Cairns. The coral grows
around the polynoid, enclosing it in a coiled tube. The species is compared
with the type species of the genus, B. antipathicola (Benham), from off New
Zealand, found in latticed tunnels formed by the antipatharian coral Paranti-
pathes tenuispina (Silberfeld). Both polynoid species are referred to the new

subfamily Lepidastheninae.

While working on his description of the
new species of stylasterid coral from Aus-
tralia, the unique unattached Conopora ad-
eta, Cairns (1987) noted the presence of a
polychaete enclosed in each of his ten coral
specimens. He brought them to my atten-
tion and allowed me to extract the polynoids
from two of the corals, which was no small
task, since the coral paratypes had to be cut
with the minimum of damage. On exami-
nation, the commensal polynoids appeared
to belong to an undescribed genus and
species (Cairns 1987:143). After further
study, they are described below as a new
species of Benhamipolynoe Pettibone, 1970,
with type species B. antipathicola, described
by Benham (1927, under Lepidasthenia) and
Pettibone (1970, under the new genus), from
off New Zealand, associated with the anti-
patharian coral Parantipathes tenuispina.

The types are deposited in the Northern
Territory Museum, Darwin, Australia
(NTM) and the National Museum of Nat-
ural History, Smithsonian Institution
(USNM).

Family Polynoidae
Lepidastheniinae, new subfamily

Types genus: Lepidasthenia Malmgren,
1867, with type species L. elegans (Grube,
1840), by monotypy.

Diagnosis. —Prostomium bilobed, with 3
antennae and 2 palps; ceratophore of me-
dian antenna in anterior notch, lateral an-
tennae inserted terminally on anterior ex-
tensions of prostomium, on same level as
median antenna (or slightly subterminally);
two pairs of eyes on posterior half of pro-
stomium. First or tentacular segment not
distinct dorsally; tentaculophores lateral to
prostomium, each with aciculum and pair
of dorsal and ventral tentacular cirri, with-
out setae. Second or buccal segment with
first pair of elytrophores and elytra, setig-
erous parapodia and ventral or buccal cirri
lateral to ventral mouth. Elytra smooth,
without fringes of papillae, without tuber-
cles, with or without scattered micropapil-
lae. Parapodia subbiramous, with small no-
topodium, notoaciculum, and without, or

VOLUME 102, NUMBER 2

with very few, notosetae. Neuropodia with
anterior and posterior lobes deeply cut dor-
sally and ventrally, without projecting acic-
ular lobes. Dorsal cirri on non-elytrigerous
segments with basal cirrophores and distal
styles; dorsal tubercles indistinct. Pharnyx
with 9-13 pairs of border papillae and 2
pairs of chitinous jaws.

Remarks. —Lepidasthenia has been in-
cluded in the Subfamily Lepidonotinae Wil-
ley, 1902, chiefly on the basis of the simi-
larity of the prostomium to Lepidonotus
Leach, 1816. It is separated herein based on
the different types of parapodia and elytra.
Also included in Lepidastheniinae are the
following polynoid genera: Perolepis Ehlers,
1908; Parahalosydna Horst, 1915; Hyper-
halosydna Augener, 1922; Alentiana Hart-
man, 1942; Telolepidasthenia Augener &
Pettibone, 1970; Benhamipolynoe Petti-
bone, 1970. A new species is described be-
low under the last named genus.

Benhamipolynoe Pettibone, 1970

Type species: Lepidasthenia antipathicola
Benham, 1927, by original designation.

Diagnosis. —Body elongate, slender, flat-
tened, segments numerous (more than 50,
up to 200 or more). Elytra 10 or more pairs,
on segments 2, 4, 5, 7, alternate segments
to 15, then variable in arrangement, with
either long posterior region without elytra
or with elytra continuing posteriorly. Elytra
without fringes of papillae, smooth except
for scattered micropapillae. Prostomium
lepidasthenoid, bilobed, with two palps and
three antennae; ceratophore of median an-
tenna in anterior notch of prostomium, lat-
eral antennae inserted terminally on ante-
rior extensions of prostomium, lateral to
median antenna, with two pairs of eyes on
posterior half of prostomium. Tentaculo-
phores of tentacular segment lateral to pro-
stomium, without setae, with two pairs of
dorsal and ventral tentacular cirri. Second
or buccal segment with first pair of elytra

301

and long ventral buccal cirri; without nuchal
lobe. Parapodia subbiramous, with small
conical notopodia on anterodorsal faces of
neuropodia, without notosetae. Neuropo-
dia deeply cut dorsally and ventrally, with
anterior and posterior subequal rounded
lobes, without projecting acicular lobes.
Neurosetae relatively few in number (5-12),
of single type, rather stout, smooth or with
slight indication of spinous rows, with fal-
cate entire tips. Dorsal cirri with short cy-
lindrical cirrophores and long, smooth, ta-
pering styles; dorsal tubercles indistinct.
Ventral cirri short, subulate. Pygidium with
paired long anal cirri. Nephiridial papillae
short, cylindrical, beginning on segment 8.

Remarks. — Benhamipolynoe differs from
Lepidasthenia Malmgren in regard to the
number and arrangement of the elytra and
the types of neurosetae. In Lepidasthenia,
the neurosetae have well-developed short
and wide spinous regions and bifurcate tips;
in Benhamipolynoe, the neurosetae have
spinous regions scarcely developed or lack-
ing and entire falcate tips.

Benhamipolynoe antipathicola (Benham,
1927), originally described as Lepidasthen-
ia, from off New Zealand in 128 meters, was
associated with Parantipathes tenuispina
(Silberfeld), where it was found in latticed
tunnels formed by the serrated branchlets
of the antipatharian. Additional specimens
referred to B. antipathicola by Pettibone
(1970) include one from the Gulf Stream off
Key West, Florida, in 237 meters, associ-
ated with Antipathes columnaris Duchas-
saing, and one from Indonesia, in 520 me-
ters, found in an empty tube of a eunicid
polychaete.

Benhamipolynoe cairnsi, new species
Figs. 1, 2

Material. —Queensland, Australia, Mar-
ion Plateau off MacKay, 20°46.2’S,
152°51.8'E, 398-399 m, F.R.V. Soela sta
25, 22 Nov 1985, A. Bruce, collector, re-
moved from coiled galleries in stylasterid

302

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Po 1.

Benhamipolynoe cairnsi, A, C-H, Holotype, NIM W4900; B, Paratype, USNM 118373: A, Dorsal

view of anterior end, styles of median and might lateral antennae, left dorsal and ventral tentacular cirri all
missing; right parapodium and dorsal cirrus of segment 3 not shown; B, Ventral view of anterior end, styles of
median and left lateral antennae, left palp, and all tentacular cirri missing; C, Right elytrigerous parapodium
of segment 2, posterior view, acicula and small notopodium dotted, elytrophore not shown; D, Right cirrigerous
parapodium of segment 3, anterior view, acicula dotted; E, Neurosetae from same; F, Right Ist elytron from
segment 2, with few micropapillae (not to scale); G, Right 2nd elytron from segment 4; H, Right 9th elytron
from segment 19. Scales = 1.0 mm for A, B; 0.5 mm for C, D, F—H; 0.1 mm for E.

coral Conopora adeta Cairns, holotype
(NIM W4900) and paratype (USNM
118373).

Description. —Holotype (in two pieces,
incomplete posteriorly) with 38 segments,
16 mm long, 2 mm wide; paratype (in three
pieces, complete) with 63 segments, 33 mm
long, 3 mm wide; both specimens females,
full of large yolky eggs.

Body elongate, flattened ventrally, arched
dorsally, widest in anterior half, tapering
gradually posteriorly; without color except
for black eyes. Prostomium oval, bilobed;
ceratophore of median antenna large, oval,
in anterior notch, style missing; lateral an-
tennae inserted terminally on anterior con-
tinuations of prostomium, lateral to median
antenna, with long styles about equal to stout

VOLUME 102, NUMBER 2

A B

303

Fig. 2. Benhamipolynoe cairnsi, Holotype, NIM W4900: A, Right cirrigerous parapodium from segment
12, posterior view, extension on anterior side of cirrophore dotted; B, Right cirrigerous parapodium from segment
21, posterior view; C, Upper and middle neurosetae from same, latter with tip worn; D, Right cirrigerous
parapodium from segment 35, anterior view, acicula dotted; E, Upper and middle neurosetae from same. Scales

= 0.5 mm for A, B, D; 0.1 mm for C, E.

tapering palps; two pairs of subequal black
eyes on posterior half of prostomium (Fig.
1A, B). First or tentacular segment not vis-
ible dorsally; tentaculophores lateral to pro-
stomium, without setae, each with pair of
tentacular cirri, ventral one about as long
as lateral antennae, dorsal one longer than
palps; ventrally with bulbous facial tubercle
(Fig. 1A, B). Second or buccal segment with
slightly developed nuchal lobe, first pair of
elytrophores, subbiramous parapodia with
few neurosetae, and ventrally with long buc-
cal cirri lateral to ventral mouth (Fig. 1A—
C). Segment 3 with first pair of dorsal cirri;
cirrophores longer than neuropodia, cylin-
drical with distal bulbous extension and long
style; ventral cirri almost as long as buccal
cirri of segment 2 (Fig. 1A, B, D).

Elytra and elytrophores 10 pairs on seg-
ments 2-4, 5—7—9-1 1—13-15—19— 23, with
dorsal cirri on all segments from 24 on. Ely-
tra oval, attached eccentrically near lateral
side, delicate, with “‘veins’’ and scattered
round micropapillae on surface (Fig. 1 F—H).

Parapodia subbiramous; notopodia in
form of small conical acicular lobes on

anterodorsal faces of larger neuropodia,
without notosetae; neuropodia rather deep-
ly cut dorsally and ventrally, forming
rounded presetal and postsetal lobes, with
row of 5—10 short neurosetae (Figs. 1C, 2A,
B, D). Neurosetae stout, with faint spinous
rows and slightly hooked entire tips, some-
times slightly worn (Figs. 1E, 2C, E). Cir-
rophores of dorsal cirri cylindrical, extend-
ing beyond neuropodia, with triangular
extension on lower side, with long, smooth
styles tapering to slender tips and extending
far beyond parapodia; ventral cirri wider
basally, with long slender tips, extending
slightly beyond tips of neuropodia (Fig. 2A,
BaD):

Nephridial papillae beginning on segment
8, small at first, becoming longer about seg-
ment 15, cylindrical, directed dorsally be-
tween parapodia. Pharynx not extended and
not examined. Pygidium forming small lobe
with dorsal anus between last pair of small
parapodia and pair of long anal cirri, similar
to dorsal cirri.

Biology.—The two symbiont polynoids
were removed from two paratypes of the

304

colonial stylasterid coral Conopora adeta,
well figured by Cairns (1987, figs. 1-6). The
coral is unusual in that it is unattached,
formed of a massive globose solid base with
several (3-6) stout, radiating, stabilizing
branches and two main ascending porous
branches. The polynoid occupied a coiled
runway in the globose base connected to a
U-shaped tube in the vertical branches. The
close symbiotic relationship of the polynoid
with the coral begins at an early stage in
development and determines the morphol-
ogy of the colony. One of the openings near
the base of the colony was large enough for
the polynoid to be able to protrude from the
tube. All of the ten coral colonies examined
by Cairns contained a polynoid symbiont.
Both specimens of Benhamipolynoe cairnsi
were females, full of large yolky eggs.
Remarks. — Benhamipolynoe cairnsi dif-
fers from B. antipathicola in having only 10
pairs of elytra and only dorsal cirri from
segment 24 on, compared to numerous pairs
of elytra continuing to the posterior end. In
B. cairnsi, the cirrophores of the dorsal cirri
have unique extra extensions: bulbous on
segment 3 and triangular on the following
cirrophores; this condition was not found
on B. antipathicola and has not been noted
previously on other polynoids. The two
species differ in their positions with their
respective coral hosts. As indicated above,
the coiled runway of B. cairnsi is enclosed
in the stylasterid coral Conopora adeta, al-
lowing only limited movement to the ex-
terior. For B. antipathicola, the antipathar-
ian coral host, Parantipathes tenuispina, is
a bottle-brush shaped colony where the in-
terlacing branchlets form a complicated lat-
tice-work tube that serves as an elongated

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

“‘worm-run,” described by Totton (1923:
106, fig. 9), thus allowing much greater ex-
ternal movement for the polynoid com-
mensal.

Etymology.—The species is named for
Stephen D. Cairns, who described and
named the stylasterid coral host of the po-
lynoid symbiont.

Acknowledgments

My thanks go to Stephen Cairns for bring-
ing the polynoid commensal to my atten-
tion and for allowing me to extract them
from the stylasterid coral. The manuscript
benefited from the reviews of Russel Hanley
and Stephen Cairns.

Literature Cited

Benham, W.B. 1972. Polychaeta.— British Antarctic
“Terra Nova” Expedition, 19106, Natural His-
tory Reports, Zoology 7(2):47-182.

Cairns,S.D. 1987. Conopora adeta, new species (Hy-
drozoa: Stylasteridae) from Australia, the first
known unattached stylasterid.— Proceedings of
the Biological Society of Washington 100(1):141-
146.

Pettibone, M. H. 1970. Polychaeta errantia of the
Siboga-Expedition. Part 4: Some additional
polychaetes of the Polynoidae, Hesionidae, Ne-
reidae, Goniadidae, Eunicidae, and Onuphidae,
selected as new species by the late Dr. Hermann
Augener with remarks on other related species. —
Siboga-Expedite 24,1d:199-270.

Totton, A. K. 1923. Coelenterata. Part III. Antipa-
tharia (and their Cirripede Commensals). — Brit-
ish Antarctic (“Terra Nova’’) Expedition, 1910,
Natural History Report. Zoology 5(3):97—120.

Department of Invertebrate Zoology, Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 305-310

TWO NEW SPECIES OF HARMOTHOINAE
(POLYCHAETA: POLY NOIDAE) FROM
THE EAST PACIFIC RISE, COLLECTED BY
ALVIN DIVES 2000 AND 2003

Marian H. Pettibone

Abstract.— Alvin dives in the East Pacific Rise in 2700 meters near 11°N
collected two polynoid species of Harmothoinae: Harmothoe hollisi, n. sp. from
dive 2000, associated with a “‘wood island,” and Eunoe alvinella n. sp. from
dive 2003, collected from an inactive sulfide chimney.

As a part of the dive series of the DSRV
Alvin on the East Pacific Rise, during Dive
2000 in March 1988, near 11°51'N in 2750
meters, the pilot picked up a piece of wood
that was densely covered with biota, in-
cluding serpulid worms, gastropods, gala-
theids, ophiuroids, and additional fauna
(Van Dover 1988). Among the latter were
two polynoid polychaetes sent to me for
study by Cindy Lee Van Dover of the Woods
Hole Oceanographic Institution. The col-
lection site was in a field of lightly sedi-
mented pillow basalt, within a few kilo-
meters of known hydrothermal vent
communities. The fauna of this ““wood is-
land” is of particular interest in comparing
it with the fauna associated with the hydro-
thermal vents.

The polynoid is described below as a new
species of Harmothoe, H. hollisi, a widely
distributed genus in Harmothoinae. A large
polynoid, collected by Dive 2003 on an in-
active sufide chimney, is described as a new
species of Eunoe, E. alvinella, also a widely
distributed genus is the same subfamily.
Previously reported species of Harmothoe,
associated with the “wood island” and panel
experiments by Turner (1973, 1981) in the
deep water of the North Atlantic, included
two species described by Pettibone (1985a):
H. ingolfiana Ditlevsen and H. vagabunda
Pettibone. Harmothoe macnabi Pettibone
was described by Pettibone (1985b) from

Alvin dive 895 in the Galapagos Rift. The
above three species of Harmothoe, as well
as the new species, lack eyes, as character-
istic of most deep sea polynoids. The new
species of Eunoe differs from other species
of Eunoe in having very large eyes, occu-
pying the lateral borders of the prostomium.

Subfamily Harmothoinae Horst, 1917
Genus Harmothoe Kinberg, 1856
Harmothoe hollisi, new species
Pigs) 2

Material. —East Pacific Rise, Alvin dive
200022 Mar 1988) hS2"Ng .103°S 1 W,
2750 m, “wood island,” holotype (USNM
118867) and paratype (USNM 118868).

Description. —Length of complete holo-
type 24 mm, width with setae 15 mm, seg-
ments 37, last one minute. Length of in-
complete paratype 15 mm, width 7 mm,
segments 21. Without color except for gold-
en setae.

Body flattened, subrectangular, tapered
slightly anteriorly and posteriorly. Elytra 15
pairs, on large prominent elytrophores on
segments 2, 4, 5, 7, alternate segments to
23, 26, 29, and 32 (Figs. 1A, 2A). Elytra
large, overlapping, covering dorsum, oval
to subreniform in shape, delicate (tear eas-
ily), densely covered with conical micro-
tubercles, smaller on anterior part, becom-
ing gradually larger on posterior and lateral

306 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Pips:

Harmothoe hollisi, A, B, D, holotype (USNM 118867); C, paratype (USNM 118868): A, Anterior

end, dorsal view, palps missing; B, Same, ventral view; C, Anterior end, dorsal view, pharynx partially extended,
right palp and right ventral tentacular cirrus shorter, regenerating, right dorsal tentacular cirrus missing; D, Right
5th elytron from segment 9, with detail of microtubercles and papillae (not to scale). Scales = 1.0 mm for A—

C; 2.0 mm for D.

borders; larger tubercles cylindrical or ovoid,
with tips entire or with 2—8 nodular exten-
sions; scattered filiform papillae on surface
and lateral and posterior borders (Fig. 1D).
Non-elytra-bearing segments with dorsal
cirri and nodular to digitiform dorsal tu-
bercles (Figs. 1A, 2B).

Prostomium deeply bilobed, with distinct
cephalic peaks, tending to curl inward, with-
out eyes; median antenna with large bul-
bous ceratophore in anterior notch and long
papillate style; lateral antennae with short
bulbous ceratophores inserted ventrally,
with styles short, bulbous and papillate ba-
sally, and long filamentous tips; palps (miss-
ing on holotype) stout, tapered, smooth (Fig.

1A-C). Tentaculophores lateral to prosto-
mium, each with small acicular process and
2-3 short notosetae on inner side, and dor-
sal and ventral tentacular cirri similar to
median antenna (Fig. 1A—C). Second or
buccal segment with first pair of large elytro-
phores, biramous parapodia, and long ven-
tral buccal cirri inserted basally on neuro-
podia lateral to ventral mouth, similar to
tentacular cirri (Fig. 1A, B).

Parapodia biramous, with golden setae.
Notopodia rounded basally, with projecting
acicular lobe on lower side, almost as long
as neuropodia (Fig. 2A, B). Notosetae nu-
merous, forming radiating bundle, slightly
curved, about four different lengths, shorter

VOLUME 102, NUMBER 2

307

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wey W if / 4
we 4 j i j
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ay wl | 4 /
way al } ; |
wily me i y !
: 4
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0) ] | | y
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C D

re

Fig. 2. Harmothoe hollisi, holotype (USNM 118867): A, Right elytrigerous parapodium from segment 11,
anterior view, acicula dotted; B, Right cirrigerous parapodium from segment 12, posterior view; C, Long and
short notosetae; D, Subacicular neurosetae, with detail of tips; E, Supraacicular neuroseta, with detail of tip.

Scales = 1.0 mm for A, B; 0.1 mm for C—E.

than and much stouter than neurosetae; no-
tosetae with numerous spinous rows and
rather long, tapered bare tips (Fig. 2C). Cy-
lindrical cirrophores of dorsal cirri on dor-
soposterior sides of notopodia, with slender
papillate styles extending beyond neurose-
tae (Fig. 2B). Neuropodia with conical pre-
setal lobe and projecting acicular process
with small supraacicular extension, and
shorter rounded postsetal lobe (Fig. 2A, B).
Neurosetae numerous, forming fan-shaped
bundle, with numerous spinous rows. Su-

praacicular neurosetae more slender, with
long spinous regions and bare split tips (Fig.
2E). Subacicular neurosetae stouter, with
shorter spinous regions, and slightly hooked
bare tips, all with rather long secondary tooth
(Fig. 2D). Ventral cirri short, tapered, with
abruptly narrowed distal tip (Fig. 2A, B).
Pharynx not extended and not examined.
Inflated ventral nephridial areas with small
papilla on posterior side, beginning on seg-
ment 6 (Fig. 2B). Pygidium forming small
lobe between small parapodia of 2 posterior

Table 1.—Comparison of Harmothoe ingolfiana and Harmothoe hollisi.

H. ingolfiana

H. hollisi

Cephalic peaks of prostomium
Elytral macrotubercles
Neurosetae

small
present

upper ones with bifid tips, middle
and lower ones with entire tips

prominent, tending to curl inward
absent

all with bifid tips

308

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Eunoe alvinella, holotype (USNM 118866): A, Anterior end, dorsal view, pharynx partially extended,
styles of median antenna, left lateral antenna, right and left dorsal and ventral tentacular cirri missing; B, Left
2nd elytron from segment 4, with detail of microtubercles and papillae (not to scale); C, Left middle elytron,
with detail of microtubercles and papillae (not to scale). Scale = 1.0 mm for A-C.

segments, with pair of anal cirri (styles miss-
ing).

Etymology.—The species is named for
Ralph Hollis, the pilot of Alvin dive 2000,
who collected the interesting ““wood island”’
with its associated biota.

Comparison. —Harmothoe hollisi agrees
in most respects with H. ingolfiana Ditlev-
sen, reported from the North Atlantic south
of Iceland to off New England in 1830-3506
meters and found in burrows of wood-bor-
ing bivalves (Ditlevsen 1917:32, Pettibone
1985a:143). The two species are compared
in Table 1.

Genus Eunoe Malmgren, 1865
Eunoe alvinella, new species
Figs. 3, 4

Material.—East Pacific Rise, Alvin dive
2003, 25 Mar 1988, 11°46’N, 104°47’'w,
2725 m, inactive sulfide chimney, sample
no. 9, holotype (USNM 118866).

Description. —Length of complete holo-
type 41 mm, width with setae 14 mm, seg-

ments 34, last one minute. Dorsum light
brown, with colorless transverse bands.

Body elongate-oval, slightly tapered an-
teriorly and more so posteriorly, flattened
ventrally, arched dorsally. Elytra 15 pairs,
on large bulbous elytrophores on segments
2, 4, 5, 7, alternate segments to 23, 26, 29,
and 32 (Figs. 3A, 4A, C). Elytra large, over-
lapping, covering dorsum, oval to subreni-
form in shape, thickly covered with rusty
yellow and brown foreign material; elytral
surface with microtubercles on most of sur-
faces large globular macronodules on pos-
terior third of elytra, variable in number
and size, and filiform papillae on lateral bor-
der and scattered on surface (Fig. 3B, C).
Non-elytra-bearing segments with dorsal
cirri and nodular dorsal tubercles (Fig. 4B,
D).

Prostomium deeply bilobed, wider than
long, without distinct cephalic peaks, with
two pairs of very large eyes occupying lateral
sides of prostomium; median antenna with
large bulbous ceratophore in anterior notch,
style missing (probably long); lateral anten-

VOLUME 102, NUMBER 2

309

Fig. 4. Eunoe alvinella, holotype (USNM 118868): A, Right elytrigerous parapodium from segment 2,
anterior view, acicula dotted, style of ventral buccal cirrus missing; B, Right cirrigerous parapodium from
segment 3, posterior view, style of dorsal cirrus missing; C, Right elytrigerous parapodium from segment 13,
anterior view, acicula dotted, style of ventral cirrus hidden from view; D, Right cirrigerous parapodium from
segment 14, posterior view; E, Short and long notosetae; F, Supraacicular neuroseta; G, Middle and lower
subacicular neurosetae. Scales = 1.0 mm for A—D; 0.1 mm for E-G.

nae with short bulbous ceratophores in-
serted ventrally, styles short, tapered; palps
stout, tapered, about one and a half times
length of prostomium (Fig. 3A). Tentacu-
lophores lateral to prostomium, each with
projecting aciculum and one or two noto-
setae on inner side, and dorsal and ventral
tentacular cirri, styles missing (probably
long); bulbous facial tubercle medial to cer-

atophores of lateral antennae (Fig. 3A). Sec-
ond or buccal segment with first pair of large
elytrophores, biramous parapodia; cerato-
phores of ventral buccal cirri inserted ba-
sally on neuropodia lateral to ventral mouth,
styles missing (probably longer than follow-
ing ventral cirri) (Figs. 3A, 4A).

Parapodia biramous, with golden setae.
Notopodia smaller and shorter than neu-

310

ropodia (Fig. 4C, D). Notopodium wider
basally, tapering to digitiform acicular lobe
on lower side, tip of notoaciculum project-
ing (Fig. 4C). Notosetae moderate in num-
ber, much stouter than neurosetae, upper
ones shorter, curved, lower ones longer,
nearly straight, both with numerous spinous
rows and rather short bare blunt tips (Fig.
4E). Cylindrical cirrophores of dorsal cirri
on dorsoposterior sides of notopodia, with
long papillate styles extending far beyond
neurosetae (Fig. 4D). Neuropodium with
conical presetal acicular lobe and projecting
acicular process, tip of neuroaciculum pro-
jecting; postsetal lobe shorter, rounded (Fig.
4A-D). Neurosetae numerous, with spinous
rows. Supraacicular neurosetae slender, with
long spinous regions, tapering to slender
straight bare tips (Fig. 4F); middle and low-
er subacicular neurosetae stouter, with
shorter spinous regions and slightly hooked
bare tips (Fig. 4G). Ventral cirri short, ta-
pered (Fig. 4D).

Pharynx not extended and not examined.
Nephridial papillae small, beginning on seg-
ment 4. Pygidium rectangular lobe between
small parapodia of last segment, with pair
of anal cirri (styles missing).

Etymology.—The species is named for the
collecting vessel, the DSRV Alvin.

Comparisons. —E. alvinella differs from
other species of Eunoe in having very large
eyes occupying the lateral sides of the pro-
stomium. The elytra, with their unique type
of numerous globular macronodules, also
differ from other species of the genus.

Acknowledgments

I wish to thank Cindy Lee Van Dover of
the Woods Hole Oceanographic Institution,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

as well as the pilot and observers of the
submersible A/vin, on their celebration of
dives 2000 and 2003, for the material on
which this study is based. The manuscript
benefited from the reviews of James A.
Blake, J. Fred Grassle, and Cindy Lee Van
Dover.

Literature Cited

Ditlevsen, Hj. 1917. Annelides I.—The Danish In-
golf-Expedition, Copenhagen 4(4):1-71.
Pettibone, M. H. 1985a. Polychaete worms from a
cave in the Bahamas and from experimental
wood panels in deep water of the North Atlantic
(Polynoidae: Macellicephalinae, Harmothoi-
nae).— Proceedings of the Biological Society of

Washington 98(1):127-149.

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(3):740-757.
Turner, R. D. 1973. Wood-boring bivalves, oppor-
tunistic species in the deep sea.—Science 180:
1377-1379.
. 1981. “Wood islands” and “thermal vents”
as centers of diverse communities in the deep
sea. — The Soviet Journal of Marine Biology 7(1):
1-9.
Van Dover, C. L. 1988. Alvin Dive 2000.—Sea Fron-

tiers, 34(6):326-—331.

Department of Invertebrate Zoology, Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 311-312

VARICHAETADRILUS ANGUSTIPENIS
(BRINKHURST AND COOK, 1966),
NEW COMBINATION FOR LIMNODRILUS
ANGUSTIPENIS (OLIGOCHAETA: TUBIFICIDAE)

Ralph O. Brinkhurst

Abstract. —Limnodrilus angustipenis Brinkhurst & Cook, 1966 is recognized
as a member of the genus Varichaetadrilus Brinkhurst & Kathman, 1983.

Re-examination of some of the type spec-
imens of Limnodrilus angustipenis Brink-
hurst & Cook, 1966 reveals that this taxon
should be attributed to the genus Varichae-
tadrilus Brinkhurst & Kathman, 1983. Five
specimens of the original set collected from
Saginaw Bay, Lake Huron in 1956 by the
former United States Bureau of Commer-
cial Fisheries Laboratory of Ann Arbor,
Michigan, were examined. These were orig-
inally donated to the author by J. K. Hil-
tunen, now of Sugar Island, Michigan.

As the species is usually identified by ref-
erence to the shape of the somewhat elon-
gate penis sheaths, very few if any dissec-
tions or sections appear to have been
prepared from this species. Furthermore, the
male ducts have never been fully illustrated.
The species was originally described in the
third part of a series of publications on the
North American aquatic Oligochaeta
(Brinkhurst & Cook 1966) in which a few
tubificid descriptions were added to a paper
largely directed to the Family Lumbriculi-
dae. No follow-up description has been
published.

The male ducts of one of the five speci-
mens at hand are visible as a series of frag-
ments from a dissection.

The union of the vas deferens and atrium
and the origin of the prostate gland close to
this point is evident on both male ducts.
The form of these could well be mistaken
for those of Limnodrilus. Beyond this point,
however, the atrium does not narrow to form

an ejaculatory duct, but remains wide all
the way to the base of the penis sheath on
each side (Fig. 1). This unique construction
of the male ducts is typical of Varichaeta-
drilus. Neither of these male ducts is com-
plete, and so the total length cannot be es-
timated, but they would appear to be more
like those of V. psammophilus (Loden, 1977)
and V. minutus (Brinkhurst, 1965) than
those of V. pacificus (Brinkhurst, 1981) or

Pig. 1.
Varichaetadrilus angustipenis drawn with the aid of a
drawing tube from a paratype in the author’s collection.
Scale in mm. a—atrium, p—prostate stalk, s—penis
sheath, sp—spermatheca, sz—spermatozeugmata, v—
vas deferens.

Fragmented male ducts and spermatheca of

312

Fig. 2. Chaetae from segments II and VI of the
dissected specimen, posterior chaetae from another
whole mount. Scale in mm. dIJ—dorsal chaetae of II,
dV1I—dorsal chaetae of VI, vVI— ventral chaetae of VI,
pv—posterior ventral chaetae.

V. fulleri Brinkhurst & Kathman, 1983, as
illustrated by Brinkhurst (1986).

In V. angustipenis the penis sac enclosing
the long cuticular penis sheath lacks the spi-
ral muscles diagnostic of the genus Lim-
nodrilus s.s. (the positions of Limnodrilus
grandisetosus Nomura, 1932 and Limnod-
rilus silvani Eisen, 1879, might be chal-
lenged in regard to this feature). The usual
erectile penes of Varichaetadrilus appear to
be reduced to the short conical structures in
the bases of the penis sheaths in this species.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

In the original account, no mention was
made of penial chaetae, and these do not
seem to be modified on the specimens ex-
amined here. The penial chaetae are mod-
ified in V. pacificus and V. minutus (=V.
nevadanus) but not in V. fulleri or V. psam-
mophilus. Varichaetadrilus israelis Brink-
hurst, 1971 appears to lack them, too, but
this requires confirmation.

The somatic chaetae have reduced upper
teeth in anterior bundles (Fig. 2) but the
upper teeth are longer than the lower in most
bundles, though they may vary within a
bundle. There are mostly five chaetae in an-
terior bundles, but only one or two poste-
riorly. The upper teeth are especially short
in chaetae from II to V.

The form of the penis sheath, and in par-
ticular its length, is diagnostic for this
species.

Literature Cited

Brinkhurst, R. O. 1986. Guide to the freshwater
aquatic microdrile oligochaetes of North Amer-
ica. — Canadian Special Publication of Fisheries
and Aquatic Sciences 84:1—259.

—, & D. G. Cook. 1966. Studies on the North
American aquatic Oligochaeta. III: Lumbricu-
lidae and additional notes and records of other
families.— Proceedings of the Academy of Nat-
ural Sciences of Philadelphia 118:1-33.

Institute of Ocean Sciences, P.O. Box
6000, Sidney, British Columbia V8L 4B2,
Canada.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 313-323

THE ADULT MALE OF THE TROGLOBITIC OSTRACODE
SPELAEOECIA BERMUDENSIS ANGEL AND ILIFFE, 1987,
FROM AN ANCHIALINE CAVE IN BERMUDA
(CRUSTACEA: OSTRACODA: HALOCYPRIDOIDEA)

Louis S. Kornicker

Abstract.—The previously unknown adult male of Spelaeoecia bermudensis
Angel & Iliffe, 1987, the type species of the monotypic genus Spelaeoecia Angel
& Iliffe, 1987, is described and illustrated from a specimen collected in Won-
derland Cave, Bermuda. The fifth limb bears a large sensory organ that is
absent on the limbs of the adult female as well as the juvenile male, and has
not been reported previously in the suborder Halocypridina. The rod-shaped
posterior branch of the copulatory organ is not inside the canal of the broad
anterior branch as in other genera of the superfamily Halocypridoidea. The
separated anterior and posterior branches resemble those of the copulatory
organ of adult males in the superfamily Thaumatocypridoidea, which together
with the Halocypridoidea form the suborder Halocypridina. A brief supple-

mentary description is given of the adult female.

The species Spelaeoecia bermudensis was
described by Angel & Iliffe (1987:545) from
56 specimens (22 adults, but 2 of the adults
represented by carapaces only) collected in
8 anchialine caves in Bermuda. Angel &
Iliffe (p. 552) stated, ““The absence of males
in the collections could indicate that the
species breeds by parthenogenesis, but it
seems more likely that breeding occurs in
parts of the cave inaccessible to divers.”
Kornicker & Iliffe (1989) reported from 2
anchialine caves in Bermuda 10 additional
specimens including 2 A-1 males, which they
described briefly. They mentioned an adult
male collected in Wonderland Cave, Ber-
muda, which was encountered too late to
conveniently include in their paper. That
male is described herein. It was collected
along with one juvenile and nine adult fe-
males in a baited trap by Jan H. Stock in
1984. Angel & Iliffe (1987:tab. 2) reported
three adults (two represented by carapaces
only) from the same cave. Unless the ratio
of adult females to males in the species is
unusually high it seems likely, as suggested

by Angel & Iliffe (1987), that adult females
are more widespread than adult males. Per-
haps the adult male described herein was
attracted from some distance by the bait in
the trap, and trapping rather than netting
may be a more efficient method for collect-
ing males. A brief supplementary descrip-
tion is presented based on a study of the
holotype.

Superorder Myodocopa Sars, 1866

Composition. — This superorder compris-
es the orders Myodocopida Sars, 1866, and
Halocyprida Dana, 1853.

Order Halocyprida Dana, 1853

Composition. —This order comprises the
suborders Cladocopina Sars, 1866, and Hal-
ocypridina Dana, 1853.

Suborder Halocypridina Dana, 1853

Composition. —This suborder comprises
the superfamilies Halocypridoidea Dana,

314

1853, and Thaumatocypridoidea Muller,
1906.

Superfamily Halocypridoidea
Dana, 1853

Composition. —This superfamily con-
tains the family Halocyprididae Dana, 1853.

Family Halocyprididae Dana, 1853

Composition. —This family comprises the
subfamilies Halocypridinae Dana, 1853,
Conchoecinae Claus, 1891, Archiconchoe-
cinae Poulsen, 1969, Euconchoecinae Poul-
sen, 1969, and Deeveyinae Kornicker & II-
iffe, 1985.

Subfamily Deeveyinae
Kornicker & Iliffe, 1985

Composition. —The subfamily comprises
the genera Deeveya Kornicker & Iliffe, 1985,
and Spelaeoecia Angel & Iliffe, 1987.

Spelaeoecia Angel & Iliffe, 1987

Type species.—Spelaeoecia bermudensis
Angel & Iliffe, 1987.

Composition. —The genus is monotypic.

Terminology. — The lettering of bristles on
the first and second antennae is the same as
for the genus Deeveya explained in Kor-
nicker and Palmer (1987:610). On the eighth
joint of the first antenna of Deeveya the base
of the d-bristle is slightly more lateral than
on Spelaeoecia. Therefore, it is proposed
that the anterior bristle regardless of wheth-
er it is lateral or medial be designated the
d-bristle; the definitely lateral bristle near

Fig. 1.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

midwidth of the joint and posterior to the
d-bristle and stouter than other bristles on
the joint be designated the e-bristle; the def-
initely medial bristle near midwidth of the
joint and with proximal part slightly angled
ventrally be designated the f-bristle, and the
posterior bristle that is lateral to the f-bristle
and either slightly medial to, or at same
level, as the e-bristle be designated the
g-bristle (Fig. 2a). The e-bristle is the “‘prin-
cipal bristle.”’ The terminology of the sixth
limb is that suggested by Angel & Iliffe (1987:
551) in which the joint previously inter-
preted to be the first exopodial joint is now
designated the basale, and a terminal dorsal
process on that joint is now designated the
endopodite.

Spelaeoecia bermudensis
Angel & Iliffe, 1987
Fig. 1-5

Spelaeoecia bermudensis Angel & Iliffe,
1987:545, figs. 2-6.—Kornicker & Iliffe,
1989, fig. 29.

Holotype. —USNM 228468, adult female
on five slides.

Type locality.—Green Bay Cave, Ber-
muda.

Material. —Wonderland Cave, Bermuda:
USNM 193483, one adult male on slide and
in alcohol; USNM 193484A, one adult fe-
male in alcohol; USNM 193484B, one ju-
venile (length 1.27 mm, height 0.74 mm)
plus nine adult females, all in alcohol; all
collected 4 Oct 1984 in baited trap at 0-1
m depth by Dr. Jan H. Stock. Also exam-
ined: holotype, and 2 A-1 males (USNM
193404A, 193405A) from Jane’s Cave, Ber-

—

Spelaeoecia bermudensis, adult male, USNM 193483: a, Lateral view of specimen from left side

showing representative surface lineations and some internal characters, length 1.37 mm; b, Dorsal view of
carapace with valves partly open (dashed lines indicate ventral edges of valves); c, Detail of posterodorsal corner
of carapace, from “‘a’’; d, Detail of posterior of right valve; e, Specimen showing relationship of appendages
(Bellonci Organ, furca, copulatory organ, and appendages of left side; not all bristles of appendages shown); f,
Bellonci Organ and joints | and 2 of left 2nd antenna; g, Anterior of body and upper lip from right side, anterior
to right; h, Upper lip from left side, anterior to left; i, }, Upper lip, anterior and posterior views, respectively;

VOLUME 102, NUMBER 2 315

gland

a,

lig,

i,

k, Ventral view of lower lip, anterior towards bottom. Abbreviations as follows: a.m. = adductor muscle

attachment area of shell; B.O. = Bellonci Organ; c.o. = copulatory organ; endop. = endopodite; epip. = epipodite;
exop. = exopodite; i.m. = inner margin of infold; mnd. = mandible; mx. = maxilla; r.v. = right valve; v.e. =
valve edge. Ist = First antenna, 2nd = second antenna, 5th = fifth limb, 6th = sixth limb, 7th = seventh limb.

Arabic numerals without a prefix refer to joints of endopodite or exopodite with | indicating proximal joint.
Roman numerals identify endites with I indicating proximal endite.

316

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

“

Fig.2. Spelaeoecia bermudensis, adult male, USNM 193483: a, Left 1st antenna, lateral view; b, c, Endopodite
of right 2nd antenna, medial and lateral views, respectively; d, Endopodite of left 2nd antenna, lateral view; e,
Exopodite of 2nd antenna; f, Right 6th limb (only proximal part of all except dorsal epipodial bristles shown),
lateral view; g, Left 7th limb, lateral view, ventral to left; h, Left lamella of furca, apron, and bifurcate unpaired

bristle; i, Copulatory organ, anterior to left. For abbreviations, see Fig. 1.
—

Fig. 3. Spelaeoecia bermudensis, adult male, USNM 193483: a-d, Left mandible: a, Dorsal view of coxale
endite, anterior to right; b, Lateral view of mandible (not all bristles of basale and none of 3rd endopodial joint

VOLUME 102, NUMBER 2 317

Moy

[(
yl KW wl NT

ha wht f

shown); c, Detail of proximal set of teeth of coxale endite, from “‘a’’; d, Joints 2 and 3 of endopodite, medial view.
e-g, Right mandible: e, Ventral view of outer distal set of teeth and part of proximal teeth of coxale endite (inner
distal set not shown), anterior to right; f, Ventral view of proximal set of teeth, anterior to right; g, Lateral view
of basale and joint | of endopodite, lateral view. Adult female, holotype, USNM 228468: h, i, bristles between
proximal and distal sets of teeth of coxale endite: h, Ventral view of endite of right limb, anterior to right; 1,
Dorsal view of endite of left limb, anterior to right. For abbreviations, see Fig. 1.

318 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

baa
Z Z<--., /coxale ;
recoxal Tt
ie Nae basale /
| coxale / pasale | \
XX \ III |
II \

precoxale,

I

a a

; I
\)
|
:

Fig. 4. Spelaeoecia bermudensis, adult male, USNM 193483, maxilla: a, Right limb (bristles of endites not
shown); b, Left limb (not all endite bristles shown); c—-e, Endites I-III of left limb (only proximal part of dashed
bristle of endite II shown). For abbreviations, see Fig. 1.

—=

Fig. 5. Spelaeoecia bermudensis, adult male, USNM 193483, 5th limb: a, Lateral view of right limb (only
proximal part of epipodial bristles shown); b, Lateral view of sensory organ of protopodite of left limb, anterior
to left (epipodial bristles not shown); c, Medial view of endopodite of right limb, anterior to left; d, Ventral

VOLUME 102, NUMBER 2 319

oblique view of sensory organ of protopodite of right limb, distal end of limb to lower right. For abbreviations,
see Fig. 1.

320

muda, both discussed in Kornicker & Iliffe
(1989).

Description of adult male (Figs. 1—3g, 4,
5).—Carapace uncalcified, elongate; dorsal
margin straight, ventral margin broadly
convex; anterior incisur just dorsal to mid-
height (Fig. la); in dorsal view carapace
broadest near middle and narrowing pos-
teriorly (Fig. 1b). Anterior edge of valve sin-
uate and posterior to anterior end of ros-
trum.

Ornamentation: Surface with thin linea-
tions (representative lineations near poste-
rior of left valve illustrated in Fig. la), and
few single hairs.

Infold: Broad infold except along hinge,
narrower just posterior to inner end of in-
cisur (Fig. la). Narrow list distal to mid-
width of infold extending from anteroven-
tral to posteroventral corners of valve (Fig.
la); a second list forming narrow bar pos-
terior to posterior juncture of hinge, then
extending ventrally as narrow list intersect-
ing posterior edge of valve at midheight (Fig.
lc, d); at intersection second list joins la-
mellar prolongation of selvage suggesting
that second list represents edge of valve and
that segment of shell posterior to list is outer
valve surface and similar in nature to outer
surface of valve forming inner side of ros-
trum.

Glands (Fig. la—d): Posteroventral corner
of right valve with about 10 minute glan-
dular pores along dorsal edge. In lateral view
posterodorsal corner (part bearing glands)
of right valve extending dorsally past more
rounded corner of left valve (Fig. 1c). Outer
edge of that part of infold extending from
incisur to midheight of posterior end of valve
with about 18 minute broad tube-like glan-
dular openings, mostly along anteroventral
margin and anterior three-fourths of ventral
margin (last opening just ventral to inter-
section of valve margin and second list);
minute bristles along valve edge, generally
one or two between adjacent tube-like glan-
dular openings.

Selvage: With narrow unfringed prolon-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

gation extending from inner corner of in-
cisur to midheight of posterior end of each
valve. Posterior end of prolongation ends
at intersection between valve edge and ven-
tral end of second list.

Size: USNM 193483, length 1.37 mm,
height 0.78 mm.

First antenna (Figs. le, f, 2a): With eight
distinct joints (Fig. 2a). First joint with ter-
minal ventral extension with long spines.
Second joint with distinct dorsal bristle with
indistinct rings; medial surface of joint with
distal spines. Third joint with ventral bristle
with base on lateral surface. Fourth joint
about half length of third, with dorsal bristle
reaching past eighth joint, and very long
ventral bristle. Fifth joint shorter than
fourth, with long ventral bristle about three-
times length of a-bristle of seventh joint.
Sixth joint about same length as fifth joint,
bare. Seventh joint longer than sixth, with
short spinous a-bristle, long ventral b-bris-
tle, and longer ventral c-bristle. Eighth joint
with d-bristle anterior and slightly medial
to e-bristle and about '4 longer than a-bristle
of seventh joint; e-bristle about twice length
of d-bristle with indistinct proximal rings,
f-bristle medial, bent ventrally, slightly
shorter and narrower than e-bristle, g-bris-
tle about same length as f-bristle. Long bris-
tles of fourth, fifth, seventh, and eighth joints
with indistinct minute marginal spines and
longer terminal spine.

Second antenna (Figs. le, 2b—e): Proto-
podite bare (Fig. le). Endopodite three-
jointed (Fig. 2b—d): First joint elongate with
two slender spinous a- and b-bristles (a-
bristle about two-thirds length of b-bristle);
second joint with short c- and d-bristles with
distal spines, one short lateral e-bristle, one
long f-bristle, and one long g-bristle (g-bris-
tle longer, stouter, and medial to f-bristle);
third joint with long equilength h-, i-, and
j-bristles, and terminal club-like process
(process of right limb longer and stouter than
that of left limb); long bristles of second and
third joints with terminal papilla. Exopod-
ite with nine joints (Figs. le, 2e): First joint

VOLUME 102, NUMBER 2

divided into long proximal and short distal
segments (well developed separation of seg-
ments in sclerotized dorsal (concave) edge
but not in ventral (convex) edge), and with
long terminal bristle (with minute ventral
spines) reaching well past ninth joint; bris-
tles of joints two to eight stouter than bristle
of first joint and with natatory hairs; ninth
joint with four bristles (1 minute bare, one
short with slender ventral spines, two long
with natatory hairs).

Mandible (Figs. le, 3a—g): Coxale endite
with proximal and distal sets of teeth sep-
arated by space (Fig. 3a—c, e, f): proximal
set comprising four stout cusps and with
indistinct spinous bristle at each end; sur-
face between cusps and proximal to cusps
with slender spines; stout tooth and four
spinous bristles between proximal and dis-
tal sets of teeth. Distal set of teeth com-
prising two flat teeth, each with seven cusps;
one stout curved process and one small bris-
tle proximal to flat teeth. Basale (Fig. 3b,
g): distal edge with five terminal triangular
cusps, one sharper triangular anterior cusp,
and a smaller posterior cusp; lateral surface
near distal edge with sharp tooth near mid-
width; lateral surface at or distal to mid-
length with one small bristle and five longer
bristles; anterior margin with one long bris-
tle distal to midlength; posterior margin hir-
sute, with two distal ringed bristles (proxi-
mal with slender tip, distal tubeformed);
proximal medial surface with transparent
plumose bristle on protuberance, and one
short bristle near endopodite (Fig. 3g); two
transparent plumose bristles present on or
close to dorsal margin; lateral surface near
insertion of endopodite with one long bare
bristle. Endopodite (Figs. le, 3b, d, g): First
joint with three bristles (one short, ventral,
one long, medial, and one long, terminal,
dorsal); second joint widening distally, with
three terminal dorsal bristles (one claw-like,
unringed, two ringed), one long unringed
subterminal ventral bristle, and minute
spines on dorsal margin and lateral surface
near dorsal margin; third joint with two long

321

stout claw-like spinous terminal bristles, four
short ringed bristles forming medial row
along terminal edge, and one longer ringed
bristle on terminal lateral edge; anterior
margin and medial surface of third joint hir-
sute.

Maxilla (Figs. le, 4): Endite I with two
proximal and nine terminal bristles (three
tubeformed) (Fig. 4c); endite II with two
proximal and eight terminal bristles (two
tubeformed) (Fig. 4d); endite III with one
long proximal bristle and six terminal bris-
tles (two tubeformed) (Fig. 4e); some bris-
tles on each endite stout pectinate. Basale
with long stout dorsal bristle with long mar-
ginal spines and one long ventral bristle with
long marginal spines (Fig. 4a, b); division
of basale and first endopodial joint indicat-
ed only by separation at dorsal edge, else-
where joints appearing fused. Endopodite
(Fig. 4a, b): First joint with five anterior
bristles (four proximal, one distal), two
proximal bristles near posterior margin, and
six distal bristles on or near posterior mar-
gin (because of orientation of limb on slide,
the six distal bristles of the illustrated left
maxilla are not close to the posterior mar-
gin); end joint with two stout claw-like bris-
tles and five slender ringed bristles, and hairs
along surface anterior to claws and bristles.

Fifth limb (Figs. le, 5): Epipodite with
bristles forming three groups, each with four
(dorsal group), six, and five plumose bristles
(Fig. 5a). Protopodite with long striate sen-
sory organ with many minute papillae at tip,
and two endites (Fig. 5a, b, d): endite I with
three bristles with long spines; endite II with
four bristles (two with spines, two tube-
formed). Basale with medial hairs, one long
anterior bristle with long spines, two prox-
imal medial bristles with short spines, and
six ventral bristles (two pectinate claw-like,
three tubeformed and either bare or with
short spines, one with long spines) (Fig. 5a).
Endopodite with medial hairs, one proxi-
mal medial bristle (with short spines) and
eight additional bristles (one short tooth-
like medial, two tubeformed, two claw-like,

322

two plumose, one long bare lateral) (Fig. 5a,
c). First exopodial joint weakly divided into
broad proximal and narrower distal parts
(Fig. 5a): proximal part with eight bristles
(three slender bare ventral bristles, one long
plumose lateral bristle near ventral margin,
one spinous medial distal bristle near ven-
tral margin, one very long lateral plumose
distal bristle at midheight of joint, one spi-
nous lateral bristle near dorsal margin, and
One spinous distal dorsal bristle); distal part
with four bristles (one long subterminal bare
dorsal bristle and three shorter bristles on
or near ventral margin). Second exopodial
joint: dorsal margin with one bare distal
bristle; ventral margin with four slender
bristles (bare or with short marginal spines)
at midlength. Third exopodial joint with two
stout claw-like unringed bristles (longer with
indistinct minute ventral spines, other bare)
and one slender bare ringed ventral bristle.

Sixth limb (Figs. le, 2f): Epipodite with
bristles forming three groups, each with six,
six, and five plumose bristles (Fig. 2f). Pro-
topodite separated from basale by distinct
suture and separated into two shallow lobes
by ventral indentation and suture not reach-
ing dorsal margin: proximal part with four
ventral bristles, all with long spines; distal
part with five bristles (four with long spines,
One with short spines). Basale with six plu-
mose bristles on or near ventral margin, and
one distal lateral bristle at about midwidth
ofjoint. Endopodite single-jointed with four
long bristles (three with long spines, one
bare). Exopodite three-jointed: first joint
with four bare ventral bristles; second joint
with three bare bristles (two ventral, one
dorsal); third joint with three bristles (mid-
dle bristle claw-like with short ventral spines,
dorsal bristle bare, tending to be claw-like,
with oblique rings, ventral bristle bare).
Protopodite and basale with long medial
hairs.

Seventh limb (Figs. le, 2g): Elongate with
three long terminal bristles.

Furca (Figs. le, 2h): Each lamella with

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

eight claws; claw 1 with two weak transverse
sutures, claw 3 with three, claw 4 with two,
claw 5 with one; claw 2 broken off near base;
left lamella slightly anterior to right. Un-
paired bristle bifurcate.

Bellonci organ (Fig. le, f): Elongate with
suture at proximal third, bifurcating at about
two-thirds length; each branch with round-
ed tip just reaching third joint of first an-
tenna.

Lips (Fig. 1g—-k): Anterior face with two
small triangular processes (one on each side)
and small glandular processes forming row
on each side (four—six processes in each row)
(Fig. 1g—i). Terminal posterior edge with a
minute spine-like process and slender spines
at each side of shallow median concavity
(Fig. 1h, 1). Lower lip with triangular process
on each side of mouth (Fig. 1k).

Copulatory organ (Figs. le, 21): On left
side of body and consisting of two parts:
posterior rod-shaped organ with very long
styliform process with hirsute tip; anterior
part broad with slight knee-like bend at
about distal two-thirds and terminating in
foot-shaped process with rows of minute
teeth in “‘heel”’ area.

Apron (Fig. 2h): Reaching about three-
fourths length of anterior margin of furcal
lamellae.

Supplementary description of adult fe-
male (Fig. 3h, 1).—Size (mm): USNM
193484A, length 1.52, height 0.86; USNM
193484B, six specimens (length:height):
1.54:0.91, 1.49:0.85, 1.49:0.82, 1.43:0.83,
1.53:0.85, 1.46:0.83. Range: length 1.43-
1.52 mm, height 0.82—0.91 mm. Because
some specimens were measured with valves
slightly open, height measurement is less
accurate than that of length. (Angel & Iliffe
(1987:545) gave the dimensions of three fe-
males: range of lengths 1.58—1.64 mm. They
also presented (Fig. 7) a shell length-width
graph with dimensions of 21 females having
a length range of about 1.41—1.64 mm. Kor-
nicker & Iliffe (1989) gave the dimensions
of two females with lengths of 1.54 mm and

VOLUME 102, NUMBER 2

1.57 mm. The lengths of the females in the
present collection fall within the range of
previous collections.)

First antenna: Ventral bristle of fourth
joint about same length as a-bristle of sev-
enth joint; limb otherwise similar to that of
adult male.

Mandible (Fig. 3h, i): Similar to that of
male. The exact number of bristles adjacent
to the stout tooth between proximal and
distal sets of teeth of the coxale endite is
difficult to resolve; it is interpreted to be
four herein (Fig. 3h, i), but could be only
three. Angel & Iliffe (1987:547, fig. 4B) in
describing the coxale stated, ““Between outer
and middle list single long curved spine.”
The spine is present on one limb of the ho-
lotype but not on the other, and examina-
tion of the “‘spine”’ under oil immersion re-
vealed it to be debris.

Sexual dimorphism. —The carapace of the
single adult male (length 1.37 mm) is short-
er than known females (range of length 1.41-
1.64 mm). The ventral bristle of the fourth
joint of the first antenna is about the same
length as the a-bristle of the seventh joint
on the adult female and almost three-times
the length on the adult male. The fifth limb
of the adult male bears a well developed
sensory organ not previously reported in the
halocyprids. The fifth limbs of two adult
females (Holotype and USNM 193484A)
and 2 A-1l males (USNM_ 193404A,
193405A) were examined and found not to
have a sensory organ; the A-1 males are
from Jane’s Cave, Bermuda, and were dis-
cussed in Kornicker & Iliffe (1989).

Discussion. — Although classed in the
Halocypridoidea, the Deeveyinae in many
nonsexual characters appear to be a link be-
tween the Thaumatocypridoidea and Hal-
ocypridoidea. This relationship is also ev-
ident when comparing the copulatory organs
of adult males in the two superfamilies. The
copulatory organ of the adult male S. ber-
mudensis has the rod-shaped process sep-

323

arate from the broader anterior lobe, similar
to the copulatory organ of thaumatocyprids.
In halocyprids other than S. bermudensis
the rod-shaped process of the adult male lies
inside a canal within the anterior lobe. Ju-
venile males of both thaumatocyprids and
halocyprids have the rod-shaped process
separate from the anterior lobe suggesting
that having the rod-shaped process inside a
canal within the anterior lobe is an apo-
morphic character state. The endopodite of
the adult male second antenna of S.
bermudensis differs from those generally
present in both thaumatocyprids and hal-
ocyprids in having a straight rather than
hook-shaped clasper. Not all halocyprids
have a clasper.

Acknowledgments

I thank Dr. Jan H. Stock, Institut voor
Taxonomische Zoology, Zodlogisch Mu-
seum, Amsterdam, Nederland, for the os-
tracodes from Wonderland Cave, Mr. Jack
Schroeder for inking my camera lucida
drawings, and Ms. Elizabeth Harrison-Nel-
son for general assistance.

Literature Cited

Angel, M. V., & T. M. Iliffe. 1987. Spelaeoecia ber-
mudensis, new genus, new species, a halocyprid
ostracod from marine caves in Bermuda.—
Journal of Crustacean Biology, 7(3):541-553.

Kornicker, L. S., & T. M. Iliffe. 1989. Ostracoda

(Myodocopina, Cladocopina, Halocypridina)

mainly from Anchialine Caves in Bermuda.—

Smithsonian Contributions to Zoology 470:

1-88.

,& R. J. Palmer. 1987. Deeveya bransoni, a

new species of troglobitic halocyprid ostracode

from anchialine caves on South Andros Island,

Bahamas (Crustacea: Ostracoda). — Proceedings

of the Biological Society of Washington, 100(3):

610-623.

Department of Invertebrate Zoology, Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 324-330

NEW RECORDS OF ENTOCYTHERID OSTRACODS
INFESTING BURROWING CRAYFISHES, WITH
THE DESCRIPTION OF A NEW SPECIES,
ASCETOCYTHERE STOCKERI

Horton H. Hobbs, Jr., and Daniel J. Peters

Abstract.—New records of the occurrence of entocytherids on burrowing
crayfishes are reported from Kentucky, North Carolina, Virginia, and West
Virginia. These include members of the genera Ascetocythere (3), Dactylocythere
(3), Donnaldsoncythere (1), Entocythere (2), and Uncinocythere (2). Ascetocyth-
ere Stockeri, a close relative of A. cosmeta, is described from four localities in
the Ohio Basin in Mason and Cabell counties, West Virginia, and a single

locality in Carter County, Kentucky.

In their study of burrowing crayfishes,
Raymond F. Jezerinac and G. Whitney
Stocker, of Ohio State University, Newark,
saved the detritus from containers in which
the crayfishes were preserved and have
kindly permitted us to study the ostracods
that were entrapped in it. Of the 20 samples
received, 15 from the Ohio, Potomac, and
Tennessee watersheds contained entocyth-
erids among which were representatives of
11 species belonging to 5 genera. These were
infesting six species of crayfishes (the iden-
tities of which were furnished to us by
Messrs. Jezerinac and Stocker), five of which
are largely restricted to burrows. The as-
sociations of the ostracods and their crayfish
hosts in the 15 localities are recorded below.

For each species of ostracod, the original
description and the first use of the currently
accepted orthography are cited. These are
followed by a list of the new localities, cray-
fish hosts, and remarks on the ecological
and geographic distribution of the species.

Except as noted otherwise, the ostracods
are deposited in the National Museum of
Natural History, Smithsonian Institution.

Ascetocythere cosmeta Hobbs & Hart

Ascetocythere cosmeta Hobbs & Hart, 1966:
46, figs. 24, 25. [Type locality: 4 mi N of

jct of St. Rte. 89 and Blue Ridge Parkway,
Grayson Co., Virginia. Host: Cambarus
dubius Faxon.]

New record.—North Carolina: Ashe
County [New Basin], burrows near un-
named trib of North Fork of New River on
county road, 0.1 mi E of Crumpler, 8.8 mi
NE of West Jefferson, 11 Aug 1984, G. W.
Stocker, R. F. Jezerinac, and R. F. Thoma,
host: Cambarus (Jugicambarus) dubius
Faxon, associates: Dactylocythere daph-
nioides (Hobbs, 1955), Entocythere harrisi
Peters, 1975, and Uncinocythere simondsi
(Hobbs and Walton, 1960).

Habitat. —Crayfish burrows.

Distribution. —The Catawba and New ba-
sins in Alleghany, Ashe, and Avery coun-
ties, North Carolina, and Grayson County,
Virginia.

Ascetocythere hyperoche Hobbs & Hart

Ascetocythere hyperoche Hobbs & Hart,
1966:41, figs. 12, 13. [Type locality: trib.
of Cherokee Reservoir, 0.5 mi W of ject
of U.S. Hwys. 11E and 25E, Grainger Co.,
Tennessee. Host: Orconectes spinosus
(Bundy), Cambarus (Hiaticambarus) lon-
girostris Faxon, and Cambarus (C.) bar-
tonii cavatus Hay.]|

VOLUME 102, NUMBER 2

325

Pig. |.

Ascetocythere stockeri, new sp.: a, Copulatory complex of holotypic male; b, Right valve of shell of

holotype; c, Same of paratypic female (two irregular, overlapping masses in interior Ova).

New records.— Virginia: Russell County
[Holston Basin], floodplain of tributary to
Big Moccasin Creek on County Rd 657, 0.9
air mi W of Hansonville, 15.9 air mi N of
Bristol, 9 Aug 1986, G.W.S. and R.F.J., host:
Cambarus (Jugicambarus) dubius Faxon,
associate, Donnaldsoncythere donnaldson-
ensis (Klie 1931).

Habitat.—Probably largely confined to
crayfish burrows.

Distribution.—This ostracod was previ-
ously known only from the type locality in
the Holston River basin in Grainger Coun-
ty, Tennessee, where Hobbs and Hart re-
ported the hosts to be three stream-dwelling
crayfishes.

Ascetocythere stockeri, new species
Figs

Diagnosis. —Eye pigmented and located
about one-sixth shell length from anterior
margin. Shell (Fig. 1b) ovate with greatest

height about one-third shell length from
posterior end where about 1.5 times height
at level of eye; margin entire, lacking emar-
ginations and prominences. Submarginal
setae present except dorsally.

Copulatory complex (Fig. la) with peni-
ferum expanded ventrally in bubiform en-
largement following conspicuously slender
constricted section about one-fourth of
length of peniferum from distal extremity;
anterior process lacking; ventral process
digitiform extending ventrally much be-
yond other processes; posterior process rep-
resented by two acute teeth. Penis very
prominent, spermatic and prostatic ele-
ments not contiguous throughout their
lengths, both C-shaped and emerging ven-
trally at base of ventral process between it
and small cephaloventrally directed lobe.
Clasping apparatus broadly arched, obliter-
ating junction of vertical and horizontal
rami; postaxial border entire; preaxial bor-

326

Table 1—Measurements (in mm) of Ascetocythere
stockeri, n. sp.

Holotype Males (14) Females (5)

Length (range) 0.39 0.35-0.43 0.39-0.42
Average 0.38 0.41

Height (range) 0:23 0.21-0.25 0.22-0.25
Average 0.23 0.24

der with three teeth grouped on thickened
distal fifth. Dorsal finger moderately heavy
and about one-half as long as curved, more
slender ventral finger.

Triunguis female.—Eye located about
one-sixth shell length from anterior end.
Shell (Fig. 1c) subovate, weakly concave in
anterior three-fifths. Greatest height dis-
tinctly posterior to midlength where about
1.4 times that at level of eye. Submarginal
setae present except dorsally. Genital ap-
paratus supported by pair of ribs extending
dorsomesially from complex of supports at
base of third leg. Apices of ribs in contact
with small, subspherical, homogeneous,
hyaline body embraced dorsally and pos-
teriorly by larger, heterogeneous mass. For
measurements, see Table 1.

Type locality: Chief Cornstalk Hunting
Grounds Campground, 2.1 miles southwest
of Beech Hill, 10.1 mules southeast of
Gallipolis, Mason County, West Virginia
[Kanawha Basin], where it was found in a
collection of crayfishes containing represen-
tatives of three species: Cambarus (Jugi-
cambarus) dubius Faxon, Cambarus (La-
cunicambarus) diogenes Girard, and
Cambarus (C.) bartonii cavatus Hay. These
specimens were collected on 20 Apr 1986
by G. Whitney Stocker, Dave Chrisman,
and R. F. Jezerinac.

Disposition of types: The holotypic male
is deposited in the National Museum of
Natural History (Smithsonian Institution),
USNM 240114. Paratypes are in the col-
lections of H. H. Hobbs III and the Smith-
sonian Institution.

Hosts: Cambarus (C.) bartonii cavatus

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Hay, C. (Jugicambarus) dubius Faxon, and
C. (Lacunicambarus) diogenes Girard. C.
(J.) dubius was present in all of the collec-
tions containing specimens of Ascetocythere
stockeri, and we think it unlikely that this
ostracod occurs on the first and last named
crayfishes.

Entocytherid associates: Dactylocythere
macroholca Hobbs & Hobbs, 1970; Don-
naldsoncythere donnaldsonensis (Klie,
1931); Entocythere elliptica Hoff, 1944; and
Uncinocythere zancla Hobbs & Walton,
1963.

Range and specimens examined: Nine-
teen specimens from the following locali-
ties. Kentucky: Carter County [Little Sandy
Basin], (1) roadside ditch along Dry Fork
1.5 air mi S of Willard, 1 Sep 1986, Dave
Hile, G.W.S., and R.F.J., host: C. (/.) du-
bius, associates: Dactylocythere macroholca
Hobbs & Hobbs, Entocythere elliptica Hoff,
and Uncinocythere zancla Hobbs & Walton.
West Virginia: Mason County [Kanawha
Basin], (2) Type locality; (3) 3 mi S of type
locality and 2.7 mi SE of Arlee, 2 May 1987,
G. Lombardo, R. French, and G.W.S., host:
C. J.) dubius, no ostracod associates. Cabell
County [Ohio Basin], (4) Spurlock Creek on
Union Ridge Rd, 1 mi S of St. Rte. 2, 1.8
mi N of Union Ridge, 3 May 1987, R.F.,
D. Chrisman, G.L., R.F.J., and G.W:S., host:
Cambarus (J.) dubius, associate, Dn. don-
naldsonensis; (5) along Brian Creek Rd, S
of Union Ridge Rd, 4.4 mi S of Glenwood,
3.May 1987, G.L., R.F., D.G., Roe eae
G.W.S., host: C. (J.) dubius, associate: Dn.
donnaldsonensis.

Relationships: Ascetocythere stockeri is a
member of the Coryphodes Group (Hobbs
& Hart 1966:44), characterized by the teeth
on the preaxial margin of the clasping ap-
paratus being grouped distally. Its closest
relative is probably A. cosmeta Hobbs &
Hart, 1966, from which it differs most con-
spicuously in lacking an anterior process on
the peniferum. It differs from the other
members of the group, A. coryphodes Hobbs
& Hart, 1966, and A. myxoides Hobbs &

VOLUME 102, NUMBER 2

Hart, 1966, in lacking a snoutlike promi-
nence on the anterior surface of the bulbous
part of the peniferum.

Etymology: This ostracod is named in
honor of G. Whitney Stocker, who collected
or participated in collecting most of the ma-
terials on which this report is based.

Dactylocythere crawfordi Hart

Dactylocythere crawfordi Hart, 1965:255,
figs. 1, 2. [Type locality: Macacheek Creek
on St. Rte. 287, Logan Co., Ohio. Host:
Cambarus (Lacunicambarus) diogenes
Girard. ]

New record. —West Virginia: Mason
County [Ohio Basin], wet field along St. Rte.
2, 2.0 mi S of U.S. Hwy. 35, 1.5 air mi E
of Gallipolis, 19 Apr 1986, D.C., R.F.J.,
G.W.S., host: Fallicambarus (Creaserinus)
fodiens (Cottle).

Habitat. —Crayfish burrows.

Distribution.—Ohio River Basin: White
and Wabash drainages in Indiana, Little
Miami, Scioto, and Symmes Creek in Ohio,
and in Mason County, West Virginia.

Dactylocythere daphnioides (Hobbs)

Entocythere daphnioides Hobbs 1955:325,
figs. 1, 2, 5, 6-9.

Dactylocythere daphnioides.—WHart, 1962:
130.

New record.—North Carolina: Ashe
County [New Basin], burrows near trib to
North Fork of New River, 0.1 mi E of
Crumpler; 8.8 mi NE of West Jefferson, 11
Aug 1984, R.F.J., G.W.D., R.F.T., host: C.
(J.) dubius, associates: As. cosmeta, E. har-
risi, and U. simondsi.

Habitat.—Except for the record cited
above, all known specimens have been found
on crayfishes living in lotic habitats.

Distribution. —“‘From headwaters of the
Little Tennessee and Pee Dee drainage sys-
tems in North Carolina northward to the
New and Big Sandy systems in Virginia and
West Virginia. The Kentucky and Missouri

327

localities cited by Hart and Hart (1974:56)
should be confirmed’ (Hobbs & Peters,
1977:28).

Dactylocythere macroholca
Hobbs & Hobbs

Dactylocythere macroholca Hobbs & Hobbs
1970:6, fig. 3a-d.

New record.—Kentucky: Carter County
[Little Sandy Basin], roadside ditch 1.5 air
mi S of Willard, near Lawrence Co line, 1
Sep 1986, D.H., R.F.J., G.W.S., host: C. (/.)
dubius, associates, As. stockeri, E. elliptica,
and U. zancla.

Habitat. — Burrows and streams.

Distribution. —Ohio River Basin in Ken-
tucky: Barren, Licking, and Little Sandy wa-
tersheds.

Donnaldsoncythere donnaldsonensis (Klie)

Entocythere donnaldsonensis Klie, 1931:
334-341, figs. 1-9. [Type locality: Don-
ald’s Cave, Lawrence County, Indiana.
Host/s: uncertain; however, probably Or-
conectes inermis inermis Cope and/or
Cambarus (Erebicambarus) tenebrosus
Hay.

Donnaldsoncythere donnaldsonensis. — Hart,
1962:131.

New record. — Kentucky: Lawrence Coun-
ty [Big Sandy Basin], (1) roadside ditch 1.3
air mi NW of Webbyville, 1.4 air mi S of
Willard, 1 Sep 1986, D.H., G.W.S., and
R.F.J., host: C. (J.) dubius, associates: none.
Lewis County [Licking Basin], (2) seep on
St. Rte. 24, 3.8 mi E of county line, 2 air
mi E of Foxport, 9 Jun 1984, G.W.S., Diane
M. Williams, R.F.J., host: C. (J.) dubius, no
associates. Virginia: Russell County [Hol-
ston Basin], (3) floodplain along trib of Big
Moccasin Creek on Co Rd 657, 0.9 air mi
W of Hansonville, 15.8 air mi N of Bristol,
9 Aug 1986, G.W.S. and R.F.J., host: C. (J.)
dubius, associate: As. hyperoche. West Vir-
ginia: Barbour County [Tygart Basin], (4)
roadside ditch 0.8 mi E of U.S. Hwy. 119

328

on Arden Rd, 1.3 air mi NE of Hopewell,
R.F.J. and G.W:S., hosts: C. (J.) dubius, C.
(J.) diogenes, no associates. Cabell County
[Ohio Basin], (5) burrows along Brian Creek
Rd, S of Union Ridge Rd, 4.4 mi S of Glen-
wood, 3 May 1987, Greg Lombardo, Randy
French, Dave Chrisman, R.F.J.,and G.W.S..,
host: C. (J.) dubius, associate: As. stockert;
(6) burrows on Union Ridge Rd, 1 mi S of
St. Rte. 2, 1.75 mi N of Union Ridge, G.L.,
RF) D:.Ga RES and Giw'8:vhest €. Gs)
dubius, associate: As. stockeri. Grant Coun-
ty [Potomac Basin], (7) roadside ditch on
Forest Rd 75, 10.2 air mi NW of Peters-
burg, 28 Jun 1986, D.C. and G.W.S., host:
C. (J.) monongalensis, no associates. Ma-
son County [Kanawha Basin], (8) see type
locality of As. stockeri, (9) burrows along
dirt road 3 mi S of Chief Cornstalk Hunting
Ground Campground, 2.7 mi SE of Arlee,
2 May 1987, G.L., R.F., and G.W:S. (8) See
Locality “2” for As. stockeri. Pocahontas
County [Greenbrier Basin], (10) seep in Falls
of Hills Creek Scenic Area off St. Rte. 39,
1.6 mi E of Greenbrier Co line, 11.1 air mi
ESE of Richwood, 7 Jul 1985, V.L. Stocker
and G.W.S., host: C. (J.) monongalensis, no
associates.

Habitat. — All of the records cited above
are based on specimens collected from bur-
rowing crayfishes. This ostracod, however,
inhabits a wide range of epigean and hy-
pogean habitats.

Distribution. —‘‘From northern Georgia
to Indiana and Maine”? (Hobbs & Peters
1977:44). All of the localities cited since this
summary statement appeared (Hobbs & Pe-
ters, 1982:312; and Hobbs & McClure 1983:
772-778) lie within these limits.

Entocythere elliptica Hoff

Entocythere elliptica Hoff, 1944:328, figs.
15-21. [Type locality: north of Fargo,
Clinch County, Georgia. Host: Procam-
barus (Ortmannicus) seminolae Hobbs.]

New record.—Kentucky: Carter County

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

[Little Sandy Basin], roadside ditch 1.5 air
mi S of Willard near Lawrence County line,
1 Sep 1986, D.H., G.W.S., R.F.J., host: C.
(J.) dubius, associates: As. stockeri, Dt. ma-
croholca, and U. zancla.

Habitat.— Burrows and lentic and lotic
habitats.

Distribution. —“‘Hart & Hart (1974:[87-—
88], 90) recorded the presence of E. elliptica
and/or E. internotalus from Texas to North
Carolina and from Kentucky to Florida, and
Peters (1975:34) reported it from a number
of localities in the James River watershed
in Virginia’ (Andolshek & Hobbs, 1986:
31). The locality cited above extends the
range diagonally across the entire state of
Kentucky.

Entocythere harrisi Peters

Entocythere harrisi Peters, 1975:32, figs.
5-7.

New record.—North Carolina: Ashe
County [New Basin], burrows near trib of
North Fork of New River, 0.1 mi E of
Crumpler; 8.8 mi NE of West Jefferson, 11
Aug 1984, R.F.J., G.W.S., R.F.T., host: C.
(J.) dubius, associates: As. cosmeta, Dt.
daphnioides, and U. simondsi.

Habitat. —Burrows and lotic habitats.

Distribution. —On the Atlantic versant
from the York River Basin in Virginia
southward through the Catawba Basin in
North Carolina, and the New River Basin
in Ashe County, North Carolina.

Uncinocythere simondsi (Hobbs & Walton)

Entocythere simondsi Hobbs & Walton,
1960:17, figs. 1-10.
Uncinocythere simondsi. — Hart, 1962:138.

New record. —North Carolina: Ashe
County [New Basin], burrows near trib of
North Fork of New River, 0.1 mi E of
Crumpler; 8.8 mi NE of West Jefferson, 11
Aug 1984, R.F.J., G.W.S., R.F.T., host: C.

VOLUME 102, NUMBER 2

(J.) dubius, associates: As. cosmeta, Dt.
daphnioides, and E. harrisi.

Habitat. — Burrows and a broad range of
epigean lotic and lentic habitats.

Distribution. —From Illinois, Kentucky,
and North Carolina southward to Missis-
sippi and northern Florida (Hart & Hart
1974; Hobbs & Peters 1977, 1982; Andol-
shek & Hobbs 1986).

Uncinocythere zancla Hobbs & Walton

Uncinocythere zancla Hobbs & Walton
1963:456, figs. 1-3.

New record.—Kentucky: Adair County
[Green Basin], burrows along St. Rte. 205,
0.8 mi NE of Neatsville, 10 mi N of Russell
Springs, 15 Mar 1987, G.W.S. and R.F-.J.,
host: Cambarus (Jugicambarus) sp.?; no as-
sociates; Carter County [Little Sandy Ba-
sin], roadside ditch 1.5 air mi S of Willard
near Lawrence County line, 1 Sep 1986,
D.H., G.W.S., R.F.J., host: C. (J.) dubius,
associates: As. stockeri, Dt. macroholca, and
E. elliptica.

Habitat.—This ostracod has been found
most frequently infesting stream-dwelling
crayfishes, but has also been found associ-
ated with a troglobitic species in Hart Coun-
ty, Kentucky, and with the burrower cited
above.

Distribution. —Known only from the
Kentucky, Green, and Little Sandy wa-
tersheds in Kentucky, and the Cumberland
and Tennessee basins in Tennessee.

Acknowledgments

We are grateful to Raymond F. Jezerinac
and G. Whitney Stocker for making possible
our study of the material gleaned from their
collections of burrowing crayfishes. Our
thanks are also extended to C. W. Hart, Jr.,
of the Smithsonian Institution, and H. H.
Hobbs III, of Wittenburg University for their
criticisms of the manuscript.

329

Literature Cited

Andolshek, Margaret D., & Horton H. Hobbs, Jr. 1986.
The entocytherid ostracod faunas of southeast-
ern Georgia.—Smithsonian Contributions to
Zoology 424:1-—43.

Hart, C. W., Jr. 1962. A revision of the ostracods of

the family Entocytheridae.— Proceedings of the

Academy of Natural Sciences of Philadelphia

114(3):121-147.

1965. New entocytherid ostracods and dis-
tribution records for five midwestern states. —
Transactions of the American Microscopical
Society 84(2):255-259.

Hart, Dabney G., & C. W. Hart, Jr. 1974. The os-
tracod family Entocytheridae.—Academy of
Natural Sciences of Philadelphia Monograph 18:
ix + 239 pages.

Hobbs, Horton H., Jr. 1955. Ostracods of the genus
Entocythere from the New River system in North

Carolina, Virginia, and West Virginia. — Trans-

actions of the American Microscopical Society

74(4):325-333.

, & 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.

——, & H. H. Hobbs, III. 1970. New entocytherid
ostracods with a key to the genera of the subfam-
ily Entocytherinae.— Smithsonian Contribu-
tions to Zoology 47:1-19.

—., & Auden C. McClure. 1983. Ona small col-
lection of entocytherid ostracods with the de-
scriptions of three new species. — Proceedings of
the Biological Society of Washington 96(4):770—
WIS).

——., & Daniel J. Peters. 1977. The entocytherid

ostracods of North Carolina.—Smithsonian

Contributions to Zoology 147:1-73.

, and . 1982. The entocytherid ostracod

fauna of northern Georgia. — Proceedings of the

Biological Society of Washington 95(2):297-318.

, & Margaret Walton. 1960. Three new ostra-

cods of the genus Entocythere from the Hiwas-

see drainage system in Georgia and Tennes-
see.—Journal of the Tennessee Academy of

Science 35(1):17—23.

1963. Three new ostracods (Ostracoda, En-
tocytheridae) from the Duck River drainage in
Tennessee.—American Midland Naturalist
69(2):456-461.

Hoff, C. Clayton. 1944. New American species of the
ostracod genus Entocythere.—American Mid-
land Naturalist 32(2):327-357.

330 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Klie, W. 1931. Campagne spéologique de C. Bolivar
et R. Jeannel dans l’Amérique du Nord (1928)
3: Crustacés Ostracodes.—Biospeologica: Ar-
chives de Zoologie Expérimentale et Générale
71(3):333-344.

Peters, Daniel J. 1975. The entocytherid ostracod
fauna of the James and York River basins with
a description of a new member of the genus
Entocythere. — Virginia Polytechnic Institute and
University, Research Division Bulletin 93:i11 +
50.

(HHH) Department of Invertebrate Zo-
ology, National Museum of Natural His-
tory, Smithsonian Institution, Washington,
D.C. 20560; (DJP) York High School, 9300
George Washington Highway, Yorktown,
Virginia 23692.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 331-338

PSEUDOCYCLOPS LEPIDOTUS, A NEW SPECIES
OF DEMERSAL COPEPOD
(CALANOIDA: PSEUDOCYCLOPIDAE) FROM
THE NORTHWESTERN PACIFIC

Douglas J. Barr and Susumu Ohtsuka

Abstract. —Pseudocyclops lepidotus, is described from Kuchinoerabu Island,
off Kyushu, Japan. The main diagnostic characters are the lateral clefts in
pediger 5, foliaceous urosomal scales, and structure of the fifth legs in both

Sexes.

Although most oceanic calanoid cope-
pods are planktonic, many coastal species
are demersal and occupy restricted habitats
on or near the substrate. Such species may
be overlooked or misidentified owing to their
small size and resemblance to cyclopoid co-
pepods. Several families of calanoids are
comprised entirely of demersal species, the
largest and most widely distributed of these
being the family Pseudocyclopidae Gies-
brecht, 1893. This family is monogeneric,
containing only the genus Pseudocyclops
Brady, 1872.

Pseudocyclops at present contains 26 de-
scribed species, 15 of which have been col-
lected only from the North Atlantic. Only
three species have been described from the
Pacific: P. australis Nicholls, 1944, from
southern Australia; P. pacificus Vervoort,
1964, from the Caroline Islands; and P. bi-
lobatus Dawson, 1977, from southern Cal-
ifornia. P. australis was subsequently re-
ported by Tanaka (1966) from Kyushu,
Japan, close to the type locality of the new
species described below. Our specimens were
obtained with a hand-held plankton net
while using SCUBA over a coral sandy bot-
tom at a depth of 9 m off Kuchinoerabu
Island, Kyushu, Japan.

Pseudocyclops lepidotus, new species
Figs. 1-5

Material. —Dissected female holotype
(USNM 229956), and dissected male allo-

type (USNM 229957) collected 10 Sep 1986
from Nishino-Hama, Kuchinoerabu Island,
Kagoshima Prefecture, Japan. Two female
paratypes (USNM 229958), one dissected
on slides and one whole, collected on same
date from same locality.

Description. — Female (Fig. 1A, B): length
0.90 mm, body compact, prosome oval in
dorsal view; cephalosome partly fused with
pediger 1, produced anteroventrally into
acute rostrum (Fig. 1C); pediger 5 separate
from pediger 4, with dorsolateral cleft bear-
ing small seta (Figs. 1B, D; 5A). Urosome
(Fig. 1E) 4-segmented, segments bearing
transverse rows of subtriangular foliaceous
scales, many of which have fallen off during
dissection (Fig. 5B); genital segment longer
than following segments combined, open-
ings widely separate; segment 3 (Fig. 5C)
with two dorsal processes reaching mid-
length of caudal rami; anal segment re-
duced, telescoped into segment 3; caudal
ramus serrate posterodorsally, bearing one
dorsal, one lateral and four apical setae.

First antenna (Fig. 1F, G) 20-segmented,
reaching posterior end of first pediger; seg-
ment 1 with rows of anterior surface spi-
nules, three large esthetascs; suture between
segments 17 and 18 incomplete.

Second antenna (Fig. 2A) with basipodal
segments each bearing one seta on disto-
medial margin; endopod three-segmented,
segment | with two setae on medial margin
and small spinules on outer distal margin,

332

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

yy
i
WH
° 9

0.2 mm

Fig. 1. Pseudocyclops lepidotus, female: A, Habitus, dorsal; B, Habitus, lateral; C, Rostrum, ventral; D,
Pedigers 4—5, lateral; E, Urosome, dorsal; F, First antenna; G, First antenna, segment 1.

VOLUME 102, NUMBER 2 333

WAGE

WY |

AWM ANN ‘a

Fig. 2. Pseudocyclops lepidotus, female: A, Second antenna; B, Mandible; C, First maxilla; D, Second maxilla;
E, Maxilliped, posterior; F, First leg, anterior.

334 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

4
Vie vyy
y

ry
‘

,
"tre Wry ayt

yoy a
y ve YANN SS
E ayy yw
- v
awe Y. v

Fig. 3. Pseudocyclops lepidotus, female: A, Second leg, anterior; B, Third leg, anterior; C, Fourth leg, anterior;
D, Fifth leg, anterior.

VOLUME 102, NUMBER 2 335

Se —)
sy x
&

f

Vt ae
V7. S ee
OG,

op
ni
AIS
\ :
WAN)
\ VAN
I ZZNZW

YN’

Fig. 4. Pseudocyclops lepidotus, male: A, Habitus, dorsal; B, Urosome, dorsal; C, Right first antenna; D, Left
fifth leg, posterior; E, Right fifth leg, posterior.

D

0.05 mm

-————_+4 By, Dye

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Pseudocyclops lepidotus, female: A, Pediger 5, lateral; B, Scales on urosome segment 2, dorsal; C,
Urosome, posterodorsal. Male: D, Right first antenna, segments 14—18; E, Fifth legs, posterior; F, Right endopod,
fifth leg, posterior. Scale bars = 10 microns.

VOLUME 102, NUMBER 2

segment 2 with five medial and three ter-
minal setae, segment 3 with seven terminal
setae; exopod seven-segmented, segments
1—6 each with one terminal seta, segment 7
with one medial and four terminal setae.

Mandible (Fig. 2B) with 8 heavily scler-
otized teeth on gnathobase; basis with 3
medial setae; endopod with 2 segments,
bearing 3 and 10 setae, respectively; exopod
4-segmented, bearing 1, 1, 1, and 3 setae,
respectively.

First maxilla (fig. 2C) with first inner lobe
bearing 9 strong terminal spines, 4 setae on
posterior surface, 1 seta on anterior surface;
second and third inner lobes with 3 and 4
setae, respectively; outer lobes 1 and 2 with
8 and | setae, respectively; basis 2 bearing
4 distomedial setae; endopod 2 segmented,
bearing 8 medial and 7 terminal setae; ex-
opod one-segmented, with 10 setae.

Second maxilla (Fig. 2D) with five inner
lobes bearing five, three, three, three, and
seven setae, respectively; endopod reduced,
indistinctly segmented, bearing seven setae.

Maxilliped (Fig. 2E) with coxa carrying
four lobes bearing one, two, three, and
three setae, respectively, lobes three and four
with surface spinules; basis with two lobes
bearing three and two setae, respectively;
endopod five-segmented, bearing four, four,
three, four, and four setae, respectively.

Legs 1—4 (Figs. 2F; 3A—C) with three-seg-
mented rami, all bearing anterior and pos-
terior surface spinules; leg 1 (Fig. 2F) with
acute medial process and inner seta on ba-
sis, distolateral margins of endopod seg-
ments 2 and 3 acutely produced, distolateral
margins of exopod segments | and 2 pro-
duced into blunt serrate processes; leg 2 (Fig.
3A) with serrate lateral margins of exopod
segments 2 and 3; leg 4 (Fig. 3C) with short
external seta on basis, endopod segment 3
bearing 7 setae.

Fifth leg (Fig. 3D) smaller than fourth,
basis with short outer posterior seta; en-
dopod three-segmented, segments 1 and 2
produced distolaterally into bifurcate pro-

337

cess, segment 3 bearing six setae; exopod
segment 3 with four inner setae and four
flanged spines, innermost spine narrow, only
flanged at base.

Male (Fig. 4A): length 0.73 mm, prosome
as in female. Urosome (Fig. 4B) 5-seg-
mented, adorned with rows of foliaceous
““scales,’’ anal segment and caudal rami as
in female.

Left first antenna 20-segmented, seg-
ments 4 and 5 partly fused, otherwise as in
female. Other cephalic appendages, rostrum
and legs 1—4 as in female.

Right first antenna (Fig. 4C) 19-segment-
ed, geniculate between segments 15 and 16
(Fig. 5D); segments 14 and 15 partly fused,
bearing a sinuous process on anterior mar-
gin; segment 16 with acute lateral process;
segment 17 produced distally into sharp tri-
angular process curving medially, almost
reaching end of segment 18.

Fifth legs (Figs. 4D—-E, 5E) asymmetrical,
biramous, highly modified. Basipodal seg-
ments of left leg (Fig. 4D) fused, those of
right leg (Fig. 4E) separate, all with posterior
surface spinules. Left endopod one-seg-
mented, reaching tip of left exopod, bearing
five plumose setae; left endopodal process
slender, narrowing distally, confluent with
basis, reaching end of endopod. Left exopod
two-segmented, segment 1 with terminal
spine flanged laterally, segment 2 complex,
membranous, bearing four lobes: innermost
lobe spatulate, medial margin serrate; sec-
ond innermost lobe elongate, narrowed at
tip; second outermost lobe half as wide as
long, bearing proximal seta, distal margin
with v-shaped cleft; outermost lobe longest,
three times as long as wide, tip dissected
into three acute processes, bearing small seta
at base. Right endopod (Fig. 5F) one-seg-
mented, narrow; small posterior sensory pit
on lateral margin; distal margin blunt, with
patch of small setae. Right exopod one-seg-
mented, with flanged outer spine and two
long equal spines curving medially, inner-
most spine narrowed to acute apex, tip

338

curved laterally, with two setae and naked
process at base.

Etymology.—The specific name, /epido-
tus (G.), meaning “‘with scales,” refers to
the foliaceous scales on the urosome of both
SEXES.

Discussion.—Known species of Pseudo-
cyclops are demersal, spending the greater
part of the diel cycle on or near the sub-
strate. The nearly spherical shape and short
first antennae limit the ability of these co-
pepods to remain in the water column. P.
lepidotus, however, is unique among mem-
bers of the genus in possessing relatively
long first antennae. In previously known
species, the first antennae have from 15 to
18 segments and scarcely extend beyond the
cephalosome. In P. /epidotus the first an-
tennae have 20 segments and extend be-
yond the end of the first pediger. This fea-
ture, combined with its comparatively large
size and light purple coloration often as-
sociated with planktonic species, suggests
that P. lepidotus may spend more time in
the water column than other species in the
genus.

The unique characteristics of urosomal
scales, lateral clefts in pediger 5 and elongate
first antennae, separate this species from all
known species of the genus. At least one
undescribed species of Pseudocyclops oc-
curring in the western Pacific is closely re-
lated to this new species (Barr, unpub-
lished). This undescribed species, collected
from coastal waters off Guam and the Phil-
ippines, possesses all of the above-men-
tioned characters. It differs from P. /epidotus
in details of the fifth legs in both sexes and
will be described in a forthcoming paper.

Acknowledgments

We thank Drs. Thomas E. Bowman
(USNM), Taisoo Park (Texas A&M Uni-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

versity) and Merrill H. Sweet II (Texas A&M
University) for their critical reviews of the
manuscript. We also extend appreciation to
Dr. Robert C. Burghardt (Texas A&M Uni-
versity) for allowing access to the Electron
Microscopy Center, and Dr. K. Gushima
and Mr. M. Noda (Hiroshima University)
for cooperation at sea.

Literature Cited

Brady, G. S. 1872. Contributions to the study of the
Entomostraca, VII. A list of the non-parasitic
marine Copepoda of the northeast coast of En-
gland. — Natural History Transactions of North-
umberland and Durham 4:423-445, pl. X VII-
XX.

Dawson, J. K. 1977. A new species of Pseudocyclops
(Copepoda: Calanoida) from the southern Cal-
ifornia coast.—Transactions of the American
Microscopical Society 96:247-253.

Giesbrecht, W. 1893. Mitteilungen uber Copepoden.
1-6.— Mitteilungen Zoologischen Station zu
Neapel 11:56-106, pl. 5-7.

Nicholls, A. G. 1944. Littoral Copepoda from South
Australia (II). Calanoida, Cyclopoida, Notodel-
phyoida, Monstrilloida and Caligoida.—Rec-
ords of the South Australian Museum 8:1-62.

Tanaka, O. 1966. Neritic Copepoda Calanoida from
the north-west coast of Kyushu.— Proceedings
of the Symposium on Crustacea, held at Erna-
kulam from January 12 to 15, 1965, 1:38—50.
Marine Biological Association of India, Man-
dapam Camp.

Vervoort, W. 1964. Free-living Copepoda from Ifa-
luk Atoll in the Caroline Islands with notes on
related species.— Bulletin of the United States
National Museum 236:1-431.

(DJB) Department of Biology, Texas
A&M University, College Station, Texas
77843-3258; (SO) Hiroshima University,
Faculty of Applied Biological Science, Si-
tami, Saijo-cho, Higashi-Hiroshima 724,
Japan.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 339-361

A REVISION OF THE GENUS MICROPROTUS
RICHARDSON WITH DESCRIPTIONS OF
TWO NEW SPECIES, M. ACUTISPINATUS

AND M. LOBISPINATUS
(ASELLOTA, ISOPODA, CRUSTACEA)

George D. F. Wilson, Oleg G. Kussakin, and Galina S. Vasina

Russian Abstract.—

PE3IOME

Pesu3ua poga Microprotus Richardson c ommcaHuem
JIByX HOBbIX BUO0B (Asellota, Isopoda, Crustacea)
Yuscou J[x.J1.®., Kycakuu O.T., Bacuna I.C.

B pe3yjIbTaTe MpofesaHHOM peBu3uu poya Microprotus, paHee OTHOCHMOTrO K
IIPHMUTUBHOMy cpegu Asellota cemelictTBy Janiridae, omMcaHbI 1Ba HOBbIX Bua
u3 OaTHMasM pawona KypusibcKux OCTpoBos, Buy Storthyngura paradoxa Birstein
TlepeHeceH B 3TOT Xe por. Iloka3aHo, 4TO, HECMOTPA Ha OTCYTCTBHE IlaBaTe-
JIbHbIX TlepeonoyzoB, pox Microprotus ou2%KeH ObITb TepeHeceH B OHO 3 Hal-
Oosee BbICOKO CllellMaJIM3HMPOBaHHbIX CeMencTB Asellota— Munnopsidae s.1., a1
IIpeqcTaBuTesIeh KOTOPOrO XapaKTepHbI IJlaBaTeJIbHble 3aHMe Mapbl MJICOMO OB.
IIpusogzatca JIuarHo3bl cemeticTBa Munnopsidae s.]. B o6beMe, BKJIIOUaIOLIeM
ceMenicTBa Eurycopidae, Ilyarachnidae 1 Munnopsidae s. s., poga Microprotus
WM BCeX NATH M3BECTHbIX BHOB 3TOTO poga. ObcyxTaIoTCA POACTBeEHHbIe CBA3N
Microprotus c qpyruMu pozamu Munnopsidae B uacTHOCTH c pogzoM Storthyn-
gura, K KOTOpOMy OH HanOosiee O1u30K, xoTa Storthyngura—Tunuyunasa Eury-
copidae c llaBaTesIbHbIMHM MepeonogqamMu, OTCyTCTBy1olWjuMU y Microprotus.
OOcyxaeTCA BOMpPOC, ABJIA€TCA JIM OTCYTCTBMe IlaBaTeJIbHbIX HepeOMOOB y
Microprotus nie3HOMOp@HbIM TpH3HaKOM WIM 3TO—pe3yJIbTaT BTOPHYHOTO
yMpomeHus.

Abstract.—The genus Microprotus Richardson, referred earlier to the rela-
tively plesiomorphic family Janiridae, is revised. Two new species from the
bathyal zone of the Kurile Islands, M. acutispinatus and M. lobispinatus, and
Storthyngura paradoxa Birstein are also placed in this genus. Microprotus should
be included in the Munnopsidae s.1., in spite of its lack of the highly specialized
posterior natatory pereopods that characterize this family. Diagnoses are given
for the family Munnopsidae s.l., for the genus Microprotus, and for all five
species referred to this genus. The relationships between Microprotus and other
munnopsid genera are discussed, and it is found to be most closely related to
Storthyngura. Whether the absence of natatory pereopods in Microprotus is a
plesiomorphic or an apomorphic reversion is discussed; the latter hypothesis
is favored.

340

The status of the janiroidean genus Mi-
croprotus has been suspect since its incep-
tion. Richardson (1910) described two in-
dividuals from the South Bering Sea which
she put in the family Janiridae, where the
genus remained until now. Her remarks on
the genus include the statement (Richard-
son 1910:116) “This genus has affinities with
the Munnopsidae”’ (under Sars’ (1899) def-
inition, this family also included Eurycope
and I/yarachna), but later in the text she
writes ““The absence of natatory legs, the
general form of the body and the style of
the uropods, however, are characters refer-
able to the Janiridae.”’ The Munnopsidae
Sars s.1. (Wilson 1989) is defined by unique
natatory modifications, including paddle-
shaped posterior legs with many fringing
plumose setae. Richardson’s (1910) species,
M. caecus, closely resembles some species
in the munnopsid genus Storthyngura, ex-
cept that its posterior legs are not in the least
paddle-shaped and they have no plumose
swimming setae.

Vanhoffen (1914), the author of the genus
Storthyngura, described a new species,
Microprotus antarcticus from pleotelson
fragments. Apparently Vanhoffen did not
regard this species as a member of Stor-
thyngura and included it in his currently
unused and misspelled family Jolellidae.
Storthyngura, however, has come to include
a great variety of forms (see reviews by Bir-
stein 1957, George & Menzies 1968a, b).
Birstein (1970) included the species para-
doxa in Storthyngura despite its lack of na-
tatory posterior pereopods. His comments
(translated) are revealing: ““This inexplica-
ble trait [lack of swimming limbs] contra-
dicts not only the diagnosis of the genus but
even that of the family, although in all the
other characteristics the new species can be
regarded as a typical form of the genus Stor-
thyngura.”’ Birstein (1970) undoubtedly
overlooked the obscure genus Microprotus
because all authors, including Wolff (1962),
had put it in the Janiridae.

In this paper, we remove the veil of ob-
scurity from Microprotus by providing new

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

descriptions of the its members, and by dis-
cussing the meaning of its lack of natatory
limbs, at least from a phylogenetic point of
view. M. caecus is redescribed, two new
species are fully described, and diagnoses
and a key to all species are provided. We
argue that Microprotus is a derived member
of the Munnopsidae related to the hetero-
geneous genus Storthyngura, and that the
absence of natatory limbs is a reversion to
a primitive condition, rather than a plesio-
morphy.

The Systematic Position of Microprotus

We present here a formal diagnosis of the
Munnopsidae Sars, 1899 (sensu Wilson
1989), to aid the discussion of the system-
atic position of Microprotus.

Family Munnopsidae Sars, 1899 s.1.

Diagnosis (derived from Wilson, 1989).—
Janiroidea with a distinct natasome: pere-
onites 5—7 enlarged, muscular, with artic-
ulations often broadly joined or fused; pos-
terior ventral nerve cord ganglia fused into
a single mass in pereonite 5; pleotelson gen-
erally tapering posteriorly. Pereopods V—VII
natatory (in most genera) with many long,
fully plumose setae on margins of broad,
paddle-like carpi and propodi. Pereopodal
dactylar claws with trough-like hollow be-
tween superior and inferior claws enclosing
distal sensillae. Antennular article 1 broad-
ened and laterally flattened.

Discussion. —The above definition of the
Munnopsidae subsumes the smaller fami-
lies Eurycopidae, Ilyarachnidae, and Mun-
nopsidae s.s. The diagnosis includes fea-
tures that are found in no other family of
the Janiroidea in this combination. For a
more detailed discussion, see Wilson 1989).
The best corroborated phylogenetic trees of
the Munnopsidae (Wilson 1989) suggest that
the ancestral munnopsid had the following
characters in addition to the diagnostic
characters above: rostrum (a distinct frontal
projection between the antennulae) present,
natasomal pereonites and pleotelson freely

VOLUME 102, NUMBER 2

articulated, pleopod III rami with many
plumose setae.

The diagnosis of the Munnopsidae is pre-
sented here because Microprotus is a Ssig-
nificant exception in its lack of natatory legs
(Figs. 3D, 7, 10). This genus nevertheless
has all other diagnostic characters including
a completely fused natasome, an unusual
feature for an ambulatory animal (Fig. 1A-
B). Given this distribution of characters, two
possible hypotheses emerge for the classi-
fication of this genus: (1) the non-natatory
legs are plesiomorphic within the Munnop-
sidae, and Microprotus diverged early in the
evolution of the family before swimming
was acquired, i.e., it is the sister group of
the remaining munnopsids; or (2) the non-
natatory legs are a complete reversion, are
a distinctive autapomorphy for the genus,
and other characters must be used to estab-
lish the affinities of the genus within the
Munnopsidae. Using the non-natatory pe-
reopod character alone, one would be forced
to accept hypothesis 1 (H/) over hypothesis
2 (H2) because the former is more parsi-
monious. Several other character com-
plexes are examined here to establish which
hypothesis is most likely. The parsimony
values are counted globally for the Mun-
nopsidae.

Cephalon (Fig. 1E; Wilson 1989, fig. 38):
The ancestral munnopsid probably had a
distinct rostrum, while Microprotus does
not. H/ predicts either that the presence of
a rostrum in other munnopsids is a rever-
sion or that Microprotus lost the rostrum
independently of other munnopsids (two
evolutionary steps), while H2 predicts that
only the immediate ancestor of Microprotus
(and taxa of its clade) lost the rostrum in its
evolution (one evolutionary step).

The cephalons of some munnopsids bear
a synapomorphy called the frontal ridge, a
supporting bridge between the mandibular
articulations on both sides of the head. Be-
cause Microprotus has a frontal ridge, H/
predicts that the frontal ridge is a synapo-
morphy of all munnopsids and was subse-
quently lost in several taxa (e.g., Euryco-

341

Table 1.—A parsimony analysis of two evolutionary
hypotheses for the non-natatory pereopods V-VII in
Microprotus. Hl = The ancestral munnopsid did not
have natapods, and Microprotus diverged before na-
tapods were evolved. H2 = The natapods in Micro-
protus reverted to an ambulatory state. Values are the
number of evolutionary steps required in the character
for the phylogeny of the Munnopsidae (Wilson, 1989)
given that a hypothesis is true.

Character H1 H2

Pereopods V—VII not natatory
Rostrum absent

Frontal arch present
Mandibular notch in cephalon
Natasome completely fused

ONNNN KS
OK — — —& LO

Total parsimony values

pinae, Munnopsinae) or was developed at
least twice (two steps). H2, on the other hand,
only requires that the frontal ridge is a syn-
apomorphy of the group to which Micro-
protus belongs (one step).

In the sister group Acanthaspidiidae and
in some munnopsid genera, the mandible
articulates with the head by a rounded pos-
terolateral border, while other taxa, includ-
ing Microprotus and Storthyngura, have a
posterior projection of the mandible that
articulates with the cephalon in a narrow
posteriorly-directed slot. HJ suggests that
this feature either evolved twice, or evolved
once and then was lost in more derived taxa
(two steps); H2 requires only that it is a
apomorphy of some group of the munnop-
sids (one step).

Natasome (Fig. 1A—C): The posterior pe-
reonites and the pleotelson of the ancestral
munnopsid, although integrated into a unit,
were probably fully articulated because both
the Acanthaspidiidae, and several munnop-
sid genera, such as Amuletta Wilson and
Thistle, Munnopsurus Richardson, and
Munnicope Menzies & George are fully ar-
ticulated both ventrally and dorsally. The
flexion of the segments, however, is restrict-
ed and the integration of the natasome is
obvious in these taxa. Because Microprotus
has a fully fused natasome, H/ requires that
this was achieved independently from the

342

remaining munnopsids (two steps), while H2
allows that complete fusion could be a syn-
apomorphy ofa subset of the Munnopsidae.
Table 1 shows parsimony values for mun-
nopsid phylogenetic estimates using the
characters discussed above under H/ or H72.
Although these characters are only a subset
of the possible features that are useful in
munnopsid systematics (Wilson 1989), they
are highly derived features that are unlikely
to appear more than once in the evolution
of this family. HJ adds nine steps to the
munnopsid phylogenetic tree, while H2 only
adds six steps and is therefore more prob-
able. The ambulatory posterior pereopods
of Microprotus are best explained as a re-
version from a swimming form, and must
be an autapomorphy of this genus alone.
Furthermore, the natapod to pereopod re-
version is not simply a reduction of features:
no natatory landmark is present on pereo-
pods V—VII of the adult (juveniles are un-
known at present). There are no natatory
setae, the carpi and propodi are much longer
than in any other munnopsid, and these seg-
ments are tubular rather than flattened.
Other taxa, such as Syneurycope where the
pereopodal swimming function may be
greatly reduced, typically retain the rem-
nants of these landmarks (e.g., Haugsness &
Hessler 1979, fig. 31). The detailed similar-
ity between the anterior and posterior legs
(Fig. 3B, D) suggests that a genetic transfer
in pereopodal developmental processes from
anterior to posterior may underlie this re-
version (first author’s opinion). In the cur-
rent absence of information on munnopsid
genetics, further evidence could be obtained
by examining the pre- and postmarsupial
development in Microprotus species, once
juveniles and embryos are collected.

Microprotus Richardson, 1910

Microprotus Richardson, 1910:116.
Storthyngura (pars) Birstein, 1970:352.

Type species. —Microprotus caecus Rich-
ardson, 1910.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Species included. —Microprotus caecus
Richardson, 1910; M. antarcticus Vanhof-
fen, 1914; M. paradoxus (Birstein, 1970);
M. lobispinatus, n. sp.; M. acutispinatus n.
sp.

Diagnosis. —Munnopsidae with ambula-
tory pereopods V—VII: carpus and propodus
elongate and tubular, not flattened, with op-
posing margins with rows of unequally-bifid
setae; all segments lacking plumose swim-
ming setae. Cephalon with frontal ridge and
without rostrum. Pereonites 5—7 and pleo-
telson fully fused, with no free articulations
either dorsally or ventrally; natasomal pere-
onites collectively shorter than ambuloso-
mal pereonites. Body dorsal surface, lateral
margins of pereonites and pleotelson, coxae
of pereonites 5—7, and basal articles of an-
tenna with elongate spines; pleotelson ter-
minating in pair of elongate spines. Pleo-
telson with pronounced preanal ridge; ridge
and anus not covered by opercular pleo-
pods. Antennular article 1 elongate and sub-
triangular. Mandible with cuticular projec-
tion on posterolateral margin articulating
with cephalon in elongate notch. Uropods
elongate with tubular protopods and rami.

Additional description. — Body with elon-
gate, anteriorly curved spines in following
pattern: dorsal midline with one spine on
each pereonite and two on pleotelson; dor-
solateral spines on pereonites 2-7; lateral
spines on pereonites 5—7; two pairs of spines
on lateral margins of pleotelson; one pair of
spines on posterior tip of pleotelson; one or
two spines on articles 1—3 of antenna; single
spines on coxae of pereopods I; paired spines
on coxae of pereopods II-IV.

Cephalon without eyes or elongate spines,
broader than long, with robust laterally pro-
jecting cheeks at mandibular articulation
point. Pleotelson broad, in most species
broader than long; dorsally trilobed, with
central section separated by troughs from
dorsally projecting lateral lobes.

Antennular flagellum elongate, with many
articles, each article with one or several
aesthetascs; articles 2, 3 and 5 near same

VOLUME 102, NUMBER 2

length, distinctly longer than article 4; fla-
gellar articles of male very short, wider than
long, shorter than article 4. Antenna elon-
gate, length more than two body lengths.
Mandible normal but body abbreviated
proximal to dorsal condyle and palp; molar
process triturative but distally tapering; palp
functional, longer than mandibular body;
teeth on incisor process not distinct; lacinia
mobilis normal with distinct teeth; spine
row normal with many spines. Maxillipedal
basis with many coupling hooks; epipod
elongate, reaching to articulation between
palp articles 2 and 3, with small rounded
dorsolateral spine.

Pereopod I much smaller than posterior
pereopods, with thin carpus and propodus
having only simple setae. Pereopods II-VII
robust, elongate, with strong unequally-bi-
fid setae on opposing margins of carpus and
propodus.

Male pleopod I elongate, much longer than
wide, with distinct waist midlength, often
with short thick spines or tubercles on ven-
tral surface; distal tip with distinct medial
and lateral lobes; lateral lobe more elongate,
curving medially posterior to medial lobe.
Male pleopod II protopod broad, laterally
rounded, distally curving to short pointed
tip; protopodal setae on lateral margin short,
simple, not hemiplumose; endopodal stylet
shorter than protopod length; exopod very
short, rounded, with nonprojecting distal
hook. Female pleopod II broader than long,
without setae on keel. Pleopod III endopod
broad and rounded; exopod long, thin, dis-
tally rounded; exopod and endopod with
numerous plumose setae.

Remarks.—The inclusion of M. antarc-
ticus Vanhoffen is rather dubious because
the species is known only from fragments.
Its pleotelson, however, is more similar to
other species of Microprotus than to Stor-
thyngura so it must remain in this genus
until redescribed from new material. We
(GSV & OGK) have examined the paratype
of Storthyngura paradox Birstein (1970): this
species has a strong resemblance to Micro-

343

protus and is transferred herein to this ge-
nus. Two new species, M. lobispinatus and
M. acutispinatus, extend the known mor-
phological range of this genus.

Microprotus has clear affinities with the
heterogeneous genus Storthyngura, which
does have well-developed natapods. The two
genera share the following characters, often
to small details: cephalon shape and man-
dibular articulation; dorsal and lateral spi-
nation; form and orientation of the anten-
nula; uropod shape; fusion of natasomal
segments (in some species); shape of venter
of pleotelson; pleopod shape. Such a long
list of similar features practically guarantees
that these two genera form a monophyletic
taxon within the Munnopsidae. Formal rec-
ognition of this group as a subfamily, how-
ever, must wait until the genus Storthyngura
is revised, and its relationships to Acantho-
cope are clarified.

The subgeneric division of Storthyngura
provided by George & Menzies (1968b) falls
short of being useful because, although they
say their analysis was based on 138 char-
acters, only 3 or 4 characters were given in
the descriptions of the subgroups; analysis
of their groupings is therefore extremely dif-
ficult. Furthermore, the subgroups may have
dubious meaning because they separate
species with truncate or forked pleotelson
tips (as in Microprotus) into different groups,
even though this may be an important char-
acter at the generic level. Birstein (1970)
also found the classification of George &
Menzies (1968b) to be ineffective. Stor-
thyngura should be subdivided because the
type species S. elegans Vanhoffen is flat-
bodied and broad while S. pul/chra Hansen
(for example) is deep-bodied and has a mus-
cular natasome. The strongly natatory at-
tributes of the latter species are more like
other Munnopsidae, indicating it may have
diverged early in the evolution of the Stor-
thyngura group. For the moment, Micro-
protus 1s considered to be among a complex
of genera represented by the polyphyletic or
paraphyletic genus Storthyngura.

344

Geographic distribution of Micropro-
tus.— This genus is an Indo-Pacific bathyal
to upper-abyssal genus with 4 species in the
northern Pacific: from 40°N to 52°14’N in
the Japan and Kurile-Kamchatka Trenches,
and off the South Aleutian Islands. A single
Antarctic species has been found in the Da-
vis Sea. This genus inhabits depths in the
range of 550-3400 m.

Key to the Species of Microprotus

la. Posterior spines on pleotelson ex-
tending to tip of uropods; uropodal
exopod distinctly shorter than en-
dopod ....... Microprotus antarcticus
Vanhoffen, 1914
lb. Posterior spines on pleotelson not
extending to tip of uropods; uro-
podal exopod subequal to endo-
pod
2a. Lateral spines on pleotelson broad,
nearly as broad as long; male pleo-
pod II stylet not tapering to thin
hair-like tip
2b. Lateral spines on pleotelson thin,
much longer than broad; male
pleopod II stylet tapering to thin
hair-like tip
3a. Dorsal and lateral spines on body
distally thick and typically round-
ed; pleotelson width (excluding
spines) distinctly greater than
length; male pleopod II stylet long,
distinctly longer than half proto-
pod length
Microprotus lobispinatus, n. sp.
(Figs. 5-7)
3b. Dorsal and lateral spines on body
distally thin and pointed; pleotel-
son width (excluding spines) sub-
equal to length; male pleopod II
stylet short, distinctly less than half
protopod length
.... Microprotus acutispinatus, n. sp.
(Figs. 8-10)
4a. Dorsal and lateral spines on body
strongly denticulate; posterior
spines on pleotelson not recurved

es © © © © © © © © © © © ee ee ee le

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

dorsally; pleotelson dorsal surface
without pair of low bumps anterior
to uropods; pleotelson width sub-
equal to length
vommde: Vhor: Microprotus paradoxus
(Birstein, 1970)
4b. Dorsal and lateral spines on body
finely denticulate; posterior spines
on pleotelson strongly recurved
dorsally; pleotelson dorsal surface
with pair of low bumps anterior to
uropods; pleotelson width greater
than length
tia ae Microprotus caecus Richardson,
1910 (Figs. 1-4)

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The Species of Microprotus

Microprotus caecus Richardson, 1910
Figs. 1—4

Types. — Holotype copulatory male, pleo-
telson damaged, some legs missing, others
loose in vial, body length 12.0 mm, USNM
(United States National Museum no.)
39521. Paratype copulatory male, dam-
aged, partly dissected, some limbs missing,
estimated length 11.2 mm, USNM 39917.

Type locality.—Albatross station 4781,
52°14.5’N, 174°13’E, off Cape Sabak, Agat-
tu Island, Aleutian Archipelago, depth 544
m, bottom consisting of fine gray sand and
pebbles.

Diagnosis.—Cephalon width 1.1 times
pereonite 1 width. Dorsal and lateral spines
on body distinctly pointed distally, not
rounded, finely denticulate. Pleotelson width
1.3 times length; dorsal surface with pair of
low bumps anterior to uropods; lateral spines
narrow, length 2.8 times basal width. Male
pleopod II stylet short, length 0.45 of pro-
topod length, with curved hair-like distal
part. Uropods much longer than posterior
spines of pleotelson; endopod length sub-
equal to exopod length.

Additional description of males. —Body
(Fig. 1A—D): Cuticle heavily calcified. Pres-
ervation color white. All margins and spines
rugose, with many fine denticles. Body wid-
est at pereonite 5, body length 2.7 times

Fig. 1. Microprotus caecus, holotype male: A—B, Dorsal and lateral views of body; C, Ventral view of
natasome; D, Ventral oblique view of cephalon and ambulosome, pereopods on right side omitted. Paratype
male: E, Lateral oblique view of cephalon. Scale bars: 1.0 mm, all.

346

width. Articular margins of anterior per-
eonites set in shallow transverse grooves.
Dorsal-most parts of pereonites 1-4 also
with shallow transverse groove. Pleotelson
with pair of short broad spines or bumps
posterior to last median spine; bumps not
visible in lateral view; lateral spines on pleo-
telson flattened dorsoventrally, postero-
lateral spines anterior to uropods triangular
in cross-section; posterior spines strongly
curving dorsally.

Antennula (Fig. 2H, I): Left antennula of
holotype approximately 10 mm long; length
0.83 of body length. Surface of articles 1
and 2 denticulate. Article 1 length 1.9 times
width; medial margin with dense group of
denticles adjacent to insertion of article 2;
dense patch of broom setae proximal to
group of denticles on medial margin. Article
2 length 0.39 of article 1 length (including
distal lobe). Article 3 subequal to article 2.

Antenna (Fig. 2H): Length 2.8 times body
length. Articles 2 and 3 with broad flattened
spines projecting ventrally. Article 3 with
spine in approximate position of scale. Ar-
ticles 5 and 6 subequal, length of either 0.22
of total antennal length.

Left mandible (Fig. 2A—D): Incisor pro-
cess with one distinct cusp ventrally, with
remaining incisive margin sinuous and
lacking distinct cusps. Lacinia mobilis with
four cusps, decreasing in length dorsally;
ventral surface of lacinia mobilis with tuft
of spinules or cuticular hairs. Spine row with
15 members, distal spines with spinules or
cuticular hairs on basal ventral surface. Mo-
lar process tapering distally, distal width
roughly half proximal width; posterior mar-
gin of triturative surface with 8 doubly set-
ulose setae and row of basal denticles, ven-
tral tooth present.

Maxillula (Fig. 2E): Outer lobe with 12
large spine-like toothed setae. Inner lobe
distal tip broadly rounded, not extending
beyond medial end of outer lobe setal row.

Maxilliped (Fig. 2G): Epipod length 0.93
of basis length; width 0.37 of length; distally
rounded. Basal endite with 13 coupling

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

hooks and approximately 5 apical fan setae
distally. Palp article 2 width subequal endite
width.

Pereopods (Fig. 3A—E): Total length in-
creasing posteriorly with pereopod I dispro-
portionately smaller than others: pereopod
I approximately half body length while pe-
reopods II-VI increasing from slightly less
(approximately 5% shorter) to near body
length. Pereopodal bases: II-IV length sub-
equal; I length 0.71 length of II; V—VII long-
er than IJ-IV, increasing in length poste-
riorly, length ratios with H-IV 1.1, 1.3, 1.5.
Ischia of all pereopods distinctly shorter than
corresponding bases: ischia I and VI less
than half bases length, ischium II length 0.69
of basis II length. Pereopod I propodus with
many fine, blunt-tipped, aesthetasc-like se-
tae; opposing margins of propodus and car-
pus with only fine setae. Setae of pereopods
II-VI: single row of unequally-bifid setae on
opposing margins of propodus and carpus;
propodus with numerous fine setae both
dorsally and ventrally. Superior dactylar
claws II-VI with triangular or blunt projec-
tion on posterior margin. (Pereopod VII not
known.)

Male pleopod I (Fig. 4A—E): Sympod wid-
est at proximal insertion, tapering to less
than half proximal width at midlength, and
expanding to 0.75 of proximal width dis-
tally. Sympod with two broad irregular rows
of low spines or bumps on ventral surface.
Distal tip of in situ sympod resting exactly
at tip of pleopod II protopod. Lateral lobes
elongate, pointed, proximally broad, distal
parts curving medially. Medial lobes short,
rounded, approximately 0.25 of lateral lobe
length, with numerous long curved simple
setae reaching to tip of lateral lobe on distal
margin.

Male pleopod II (Fig. 4F, G): Protopod
broad, laterally rounded, with fringe of sim-
ple setae laterally and distally; length 1.4
times width; ventral surface with concavity
lateral to internal musculature for exopod.
Endopod inserting 0.65 of protopod length
from pleopod insertion; exopod short, distal

VOLUME 102, NUMBER 2 347

Fig. 2. Microprotus caecus, paratype male, mouthparts: A-D, Left mandible: A, Mandible, dorsal view; B,
Incisor process and lacinia mobilis, anterior view; C, Molar process, anteromedial view; D, Distal part of
mandible showing spine row; E, Right maxillula, ventral view; F, Right maxilla, ventral view; G, Right maxilliped,
ventral view. Holotype male: H, Cephalon, dorsal view, showing antennula and antenna; I, Antennular basal
articles, in situ. Scale bars: 0.2 mm, all except for H, 2.0 mm long.

348 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Microprotus caecus, paratype male, pereopods: A, Pereopod I, lateral view; B, Pereopod II, lateral
view; C, Enlargement of adjacent parts of carpus and propodus of pereopod VI; D, Pereopod VI, medial view;
E, Pereopod VI, enlargement of dactylar claw, lateral view. Scale bars: 1.0 mm.

VOLUME 102, NUMBER 2

hook not protruding, with small distal tuft
of fine simple setae; exopodal extrinsic mus-
culature small compared to size of proto-
pod, extending only half way to lateral mar-
gin of protopod.

Pleopod III: Exopod distally rounded,
with approximately 17 plumose setae. En-
dopod distally broad, rounded, very thin,
fringed with numerous setae.

Uropod (Fig. 4H): Long, thin, with short
setae, length 0.69 of pleotelson length. En-
dopod and exopod subequal, length of either
0.87 of protopod length.

Remarks. — Microprotus caecus is known
only from two adult male specimens; fe-
males or juveniles have not been collected.
This species may be distinguished from oth-
er species of the genus by the following char-
acters: cephalon only slightly wider than
pereonite 1; spines distinctly pointed, not
rounded distally; pleotelson much wider
than long; lateral pleotelson spines that are
much longer than their basal width; and
subequal uropodal rami. Overall, the other
species are similar to M. caecus, perhaps
with the exception of VM. antarcticus whose
pleotelson shape differs rather more.

Richardson (1910: fig. 38) illustrated the
holotype of Microprotus caecus with distinct
articulations between the natasomal seg-
ments. Study of the type material reveals
that the natasome ts fully fused with no free
articulations. The dorsal surface of the na-
tasome, however, does have slight ridges
that are either remnants of the articulations
or exterior expressions of apodemes. These
ridges further corroborate the contention
that pereonites 5—7 are natasomal in origin
because each ridge extends well into the seg-
ment anterior to it (Fig. 1A), as is typical
with most munnopsids.

Microprotus lobispinatus, new species
Figs. 5-7

Types.—Holotype male, 12 mm long,
(Zoologicheskogo Instituta Akademiya

349

NAUK (ZIN) No. 1/81502). Paratype male,
11 mm, (ZIN No. 2/81503).

Type locality. —Pacific Ocean, near Itu-
rup Island, 44°52’N, 149°27’E, depth 910-
920 m, habitat: muddy sand with stones,
25 Jul 1984, coll. B. Sirenko.

Diagnosis. — Dorsal and lateral spines on
body thick and rounded distally; spines fine-
ly denticulate. Pleotelson width (excluding
spines) 1.2 times length; lateral spines broad,
length subequal to basal width. Male pleo-
pod II stylet 0.66 of protopod length, with-
out curved hair-like distal part. Uropods
much longer than posterior spines of pleo-
telson; endopod length subequal to exopod
length.

Additional description of adult male (Fig.
5). — Body somewhat robust, front half sub-
equal in width to natasome. Body length
approximately 2.2 times body width with-
out lateral spines; body widest at pereonite
4. All dorsal spines distinctly flattened, often
with concave front surface, usually curved
forwards, sometimes widened in middle
part. All dorsal spines with finely spinulous,
rasp-like surface.

Cephalon broad, approximately three
times wider than long; frontal margin
broadly concave.

Pereonite 1 considerably narrower than
cephalon, with single medial spine only.
Pereonite 2 approximately 1.5 times wider
than pereonite 1, slightly wider than pere-
onite 4 and slightly narrower than pereonite
3. Dorsal surface of pereonites 2—7 with three
spines. Medial spines gradually decreasing
in length from pereonite 2 to 6, medial spine
on pereonite 7 longer than that on pereonite
6. Dorsolateral spines nearly as long as me-
dial spines; on natasomal pereonites, much
nearer to medial spines than on pereonites
2-4. Lateral margins of pereonites 1 and 4
rounded, pereonites 2 and 3 nearly truncate,
with produced posterolateral angles. Coxal
plates of pereonites 2—4 long, strongly pro-
duced, each plate with long spine-like lobes,
anterior lobe longer than posterior. Coxal

350 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Microprotus caecus, paratype male, pleopods: A—C, Pleopod I, ventral, dorsal, and lateral views
respectively; D, E, Enlargements of pleopod I distal tip, ventral and dorsal views respectively; F, G, Right
pleopod II, ventral and enlarged dorsal view respectively. Holotype male: H, Left uropod, in situ ventral view.
Scale bars: 1.0 mm.

plates of pereonite 1 with single anterior gles of pereonites 5-7 produced outwards
spine-like lobe. into stout conical spine-like processes. Cox-

Segments of natasome fused medially but al plates of pereonites 5—7 small, with
distinct on lateral parts. Anterolateral an- rounded posterolateral angles.

VOLUME 102, NUMBER 2

Pleon length not including posterior spines
0.3 of total body length. Anterodorsal mar-
gin of first pleonite slightly marked by shal-
low transverse depression, posterior margin
somewhat more distinct. Dorsal surface of
anterior pleonite convex, with median spine
strongly curved forwards. Pleotelson broad,
1.3 times wider than long; lateral margins
each with two long stout conical spines; an-
terior spine thicker, located nearly at mid-
length of pleotelson lateral margin. Dorsal
surface of pleotelson with flattened spoon-
like median spine curving forwards.

Antennula (Figs. 5, 6F) when bent back-
wards slightly exceeding middle of body;
basal article more than two times longer
than broad, its outer margin irregularly con-
vex, inner margin roughly concave; second
article inserting near midlength of first ar-
ticle, approximately one third as long as first
article, slightly increasing in width distally.
Antennal second article with two curved
spines, one spine directed outwards, and
second inwards. Third article also with two
spines, outer spine longer than inner one.

Left mandible (Fig. 6C): Incisor process
with one tooth. Lacinia mobilis with two
teeth. Spine row with 12 spines, posterior
spines longer than anterior ones. Molar pro-
cess forming truncate cone, cut off distally,
with small tooth on inner side and shallow
apical excavation in middle. Third article
of mandibular palp broad, oval, twisted,
with numerous marginal setae, longer than
palp article 1.

Maxilliped (Fig. 6A, B): Endite with 10
coupling hooks. Epipod approximately three
times longer than broad, distally rounded,
with triangular lateral projection.

Pereopods (Fig. 7A, E, F): Pereopod I
simple, little differentiated; carpus approx-
imately 2 times longer than propodus; both
articles with numerous short setae. Carpus
1.3 times longer than ischium and merus
together. Propodus of pereopod II slightly
longer than carpus, ventral margin with
dense row of bifid spine-like setae, dorsal
margin with numerous fine setae; merus

Fig. 5. Microprotus lobispinatus, male holotype,
body in dorsal view.

short, bearing setae on both margins. Car-
pus and propodus of pereopods IIJ—VII ob-
long, narrow, subequal in length, ventral
margins with dense row of short spine-like
bifid setae; on propodus all setae subequally
long, on carpus small setae alternate with
larger setae (not shown in Fig. 7E, F); pro-
podus with one distal plumose seta. Per-
eopod VI propodus slightly longer and near-
ly twice as narrow as carpus.

Male pleopod I (Fig. 7B): Pair complex
narrowest at midlength, length 2.8 times
proximal width. Lateral lobe longer than
medial lobe, curving medially; medial lobe
with distal tip rounded, lobe-like, bearing
numerous setae.

Male pleopod II (Fig. 7C): Somewhat
broad; protopod suboval, length nearly 2
times width, inner distal tip triangular,
pointed; distal part of outer margin and pos-
terior margin with setae. As compared with
M. acutispinatus, endopod relatively long.

352 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Microprotus lobispinatus, male holotype, mouthparts and antennula: A, Maxilliped, dorsal view; B,
Maxillipedal epipod, dorsal view; C, Right mandible, dorsal view; D, Maxillula, ventral view; E, Maxilla, ventral
view; F, Antennula. Scale bars: 0.2 mm, all except F, 0.3 mm.

=

Fig. 7. Microprotus lobispinatus, male holotype, pereopods and uropod: A, Pereopod I; B, Pleopod I;
C, Pleopod II; D, Uropod; E, Pereopod II; F, Pereopod VII. Scale bars: 0.2 mm—B—D; 0.3 mm—A; 0.5 mm—

BSE:

354

Fig. 8. Microprotus acutispinatus, male holotype,
body in dorsal view.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Stylet extending slightly beyond distal tip of
protopod, length 0.7 of total protopod
length. Exopod small, with dense group of
fine simple setae on posterior curve.

Uropod (Fig. 7D): Very long and slender,
extending much beyond posterior spines of
pleotelson. Endopod slightly shorter than
exopod; latter ramus nearly as long as pro-
topod.

Remarks. — Microprotus lobispinatus is
represented by only two adult males from
one sample. This species can be distin-
guished from all Pacific species by the wide,
apically flattened and rounded dorsal spines.
M. lobispinatus also differs distinctly from
M. acutispinatus and M. paradoxus by its
relatively wider pleotelson; the species is
similar in this respect to M. caecus. M. lo-
bispinatus is easily distinguished from the
latter species by having thicker and shorter
lateral spines, especially on the posterior
pereonites and the pleotelson, by lacking
dorsal lumps on the pleotelson anterior to
the uropods, and by uropodal protopods that
do not extend beyond the posterior spines
of the pleotelson (as in M. caecus). This new
species, like all other boreal species, dis-
tinctly differs from M. antarcticus, a single
species from the southern hemisphere con-
ventionally assigned to this genus, by pos-
terior spines on the pleotelson considerably
shorter than the uropods, and by uropodal
rami that are nearly equal in length.

Etymology. —“‘Lobispinatus,”’ the adjec-
tival form of two classical nouns, means
““provided with lobe-spines.”’

Microprotus acutispinatus, new species
Figs. 8-11

Types. — Holotype, male 13 mm (ZIN No.
1/81500). Paratypes, 2 males, 2 females, all
fragmented (ZIN No. 2/81501).

Type locality. — Pacific Ocean, near Itu-
rup Island, 44°48'N, 149°31’E, depth 1100-
1200 m; habitat: sand with pebbles; 25 Jul
1984, coll. B. Sirenko.

VOLUME 102, NUMBER 2

Diagnosis. — Dorsal and lateral spines on
body thin and pointed distally; spines finely
denticulate. Pleotelson width (excluding
spines) subequal to length; lateral spines
broad, length subequal to basal width. Male
pleopod II stylet 0.4 protopod length, with-
out curved hair-like distal part. Uropods
much longer than posterior spines of pleo-
telson.

Additional description of adult males (Figs.
8, 11B, E).— Body relatively slender, slight-
ly widening posteriorly, so that natasome
distinctly wider than pereonites 1-4 and ce-
phalon; length slightly more than three times
body width across pereonite 5 without lat-
eral projections. All dorsal spines rather
slender, narrow-conical, pointed, their sur-
faces covered with minute spinules. Dorsal
spines directed forwards and upwards on
pereonites and backwards and upwards on
pleon.

Cephalon broad, width approximately
three times length; frontal margin broadly
concave.

Pereonite 1 distinctly narrower than ce-
phalon, with single median dorsal spine.
Pereonites each with three spines; median
spines subequal in length. Dorsolateral
spines on natatory pereonites placed much
nearer to medial spines than on pereonites
2—4. Lateral margins of pereonites 1 and 4
rounded, those of pereonites 2 and 3 nearly
truncate, posterolateral angles produced
outwards. Coxal plates of pereonites 2—4
with two produced, long, spine-like lobes
(damaged on pereopod III in Figs. 8, 11A—
C); anterior lobe longer than posterior one.
Segments of natasome lacking dorsal artic-
ulations; pereonites 5—7 and pleon not clear-
ly delimited. Anterolateral corners of pere-
onites 5—7 produced outwards into stout,
conical spine-like projections. Coxal plates
of pereonites 5-7 in dorsal view relatively
short, suboval in form.

Pleon length excluding posterior spines
approximately one third total body length.
First pleonite small, narrow, convex dor-

355

sally, with medial dorsal spine curved back-
wards. Pleotelson (without projections) only
slightly broader than long, somewhat pro-
duced posterior part with pair of long spine-
like pointed projections directed backwards
and slightly outwards and two pairs of lat-
eral long stout conical spines: anterior spine
considerably thicker and slightly longer, lo-
cated approximately at midlength of lateral
margin of pleotelson, posterior spine smaller
and placed near posterolateral angle of pleo-
telson. Low broad dorsomedial keel with
recurved spine-like process in middle part.

Antennula (Fig. 8) when bent backwards,
reaching midlength of body; basal article
two times longer than broad, its outer mar-
gin irregularly convex, inner margin con-
cave; second article inserting at midlength
of basal article, approximately one third
length of basal article, slightly widening dis-
tally.

Antennal (Figs. 8, 11B) second article with
two stout spines, one on inner and another
on outer margin, third article also with two
subequal spines.

Left mandible (Fig. 9D): Incisor process
with one tooth. Lacinia mobilis with three
teeth. Spine row with eight setae, posterior
spines longer than anterior ones. Molar pro-
cess conical, obliquely cut off distally, with
four setae on distal posterior edge; distal
triturating surface with conical tooth. Third
article of mandibular palp longer than first,
broad, twisted, with setal row.

Maxilliped (Fig. 9A): Endite with 11 cou-
pling hook on inner margin. Epipod three
times longer than broad; distally rounded,
with triangular lateral projection.

Pereopod I (Fig. 10C) simple, little dif-
ferentiated; carpus slender and very long,
length 1.4 times ischium and merus length
together and 1.6 times propodus length.
Ventral and dorsal margins of propodus and
ventral margin of carpus with rows of short
stout spine-like bifid setae.

Pereopod II (Fig. 10D): Propodus some-
what longer, slightly longer than carpus, with

356 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

\\ y
Nh). Y

LAs Z Wh,

ee D

Fig. 9. Microprotus acutispinatus, male holotype, mouthparts: A, Maxilliped and epipod, ventral view; B,
Maxillula, ventral view; C, Maxilla, ventral view; D, Left mandible, dorsal view. Scale bars: 0.2 mm.

VOLUME 102, NUMBER 2

setae only along ventral margin. Carpus with
four bifid setae on ventral margin. Dorsal
margin of merus serrate distally.

Pereopods V—VII (Fig. 10E): Carpus and
propodus long, slender, linear, subequal in
length, without natatory setae, but ventral
margins with row of short stout spine-like
bifid setae. Dactylus short.

Male pleopod I (Fig. 10A): Pair complex
much narrower in middle part; length ap-
proximately 2.5 times proximal width. Lat-
eral lobe much longer than endopod; distal
tip of medial lobe narrowly rounded, with
short row of simple setae arranged in fan-
like order.

Male pleopod II (Fig. 10B): Protopod very
broad, length approximately 1.5 times long-
er than wide, semicircular, distomedial cor-
ner slightly produced, pointed; posterior
margin and distal part of outer margin with
row of thin simple setae. Endopod and ex-
opod small, reaching well short of distal tip
of protopod. Stylet relatively short, extend-
ing only slightly beyond distal margin of
exopod and not reaching tip of protopod.
Exopod with dense group of long, fine, sim-
ple setae on posterolateral margin.

Uropods (Fig. 8) long and slender, ex-
tending beyond posterior spines of pleotel-
son. Exopod slightly shorter than endopod;
latter nearly as long as protopod.

Description of female.—In general, out-
line (Fig. 11A, C, D, F) similar to male;
body length 12 mm. Dorsum somewhat
wider than in male. Pleotelson bearing two
tiny low lumps posterior to central dorsal
median spine and anterior to uropods, sim-
ilar to those in M. caecus (lumps not visible
in lateral view, hidden by swelling of dorsal
surface of pleotelson lateral to depressed
medial part). Pleopod II (Fig. 10G) very
broad, 1.3 times broader than long, ventro-
medial keel without setae.

Remarks.—Microprotus acutispinatus is
represented by five specimens: three adult
males and two females, of which only the
holotype male is preserved in relatively good
condition. The other individuals are badly

357

damaged, their bodies having broken in half.
This species is easily distinguished from M.
caecus and M. lobispinatus by its narrow
pleotelson, which is nearly as wide as long
and similar in this respect to M. paradoxus.
It differs, however, from the latter species
by the narrowly conical (not flattened) dor-
sal spines on the pleotelson which are con-
siderably shorter and stouter than those of
M. paradoxus.

Etymology. —“‘Acutispinatus,” contain-
ing the adjectival form of “‘spinus” and
modified by “acutus,’’ means “‘provided
with pointed spines.” ~

>

Microprotus paradoxus (Birstein, 1970)

Storthyngura paradoxa Birstein, 1970, p.
334, Figs. 19-20.

Syntypes. —Types were not found by Dr.
Mezhov, who looked for them at our re-
quest in the deep-sea isopod collection stud-
ied by the late Professor Ya. Birstein. The
collection was first deposited in the De-
partment of Invertebrate Zoology of Mos-
cow University, was later handed over to
the Institute of Oceanology, and finally was
deposited in the Zoological Museum of
Moscow University.

Type locality. —Kurile-Kamchatka
Trench, Vityaz Station 5601, 46°26’N,
152°07'E, 2770-2820 m.

Additional material. —Japan Trench, Vi-
tyaz Station 6671, 40°15'N, 143°35’E, 2500
m. Three adult males (14.8, 14.6, and 12.9
mm long), one male fragment, one damaged
female with oostegites (13.8 mm long), two
fragments of females. This material was
identified by B. Mezhov who kindly showed
it to one of the authors (GSV).

Diagnosis. — Dorsal and lateral spines on
body thin and pointed distally; spines
coarsely denticulate. Pleotelson width (ex-
cluding spines) subequal to length; lateral
spines narrow, length much greater than
width. Male pleopod II stylet short, 0.44 of
protopod length, with curved hair-like dis-
tal part. Uropods much longer than poste-

VOLUME 102, NUMBER 2

rior spines of pleotelson; endopod length
subequal to exopod length.

Remarks.—The specimens from the Ja-
pan Trench in general correspond well to
descriptions and drawings of syntypes given
by Birstein. This species is clearly distin-
guished by considerably thinner and longer
acute spines on the posterior part of the
body; these spines are not bent dorsally and
extend far beyond the distal ends of the uro-
podal protopodites. The dorsal surface of
the pleotelson lacks lumps anterior to the
uropods and the pleotelson is relatively nar-
row. This species differs from M. Jobispi-
natus and M. acutispinatus by considerably
longer and thinner spines on the pleotelson,
from M. lobispinatus by thin dorsal spines,
and from M. acutispinatus by a flattened
body.

Microprotus antarcticus Vanhoffen, 1914

Microprotus antarcticus Vanhoffen, 1914:
545-546, Fig. 71la-d.

Type. —Only a 2 mm long abdomen with
a portion of the posterior pereonal segment
and a pereopod basis are known. This ma-
terial (not examined by the authors) is prob-
ably at the Berlin State Museum, East Ger-
many, where other Vanhoffen material has
been found.

Type locality.— Antarctic Indian Ocean,
Davis Sea, Gauss station 30.11.1903,
65°27'S, 80°33’E, 3398 m.

Diagnosis. —Lateral and posterolateral
spines on pleotelson robust and elongate,
posterior spines extending to distal tips of
uropods. Uropodal exopod distinctly short-
er than endopod.

Remarks. —Microprotus antarcticus may
actually belong in another genus of the Stor-
thyngura complex, but until more speci-

——

359

mens are discovered, this species must be
retained in Microprotus.

With such scarce material at his disposal,
Vanhoffen (1914) could not describe this
Antarctic species adequately, and could only
give comments, comparing the specimen
with M. caecus, the only species of the genus
Microprotus known at that time. According
to Vanhoffen, Pacific (Arctic in Vanhoffen’s
term) and Antarctic species differ mostly in
that the latter has a low tubercle on the lon-
gitudinal thickening of the caudal segment,
while in the Antarctic species, the middle
spine between both lateral spines looks
abrupt. In addition, there are constrictions
over the two posterolateral spines, and the
uropods are equal in length to the distal
spines, with the outer uropodal ramus being
only one third of the inner ramus length.
In M. caecus, all spines are considerably
smaller in comparison with the abdomen,
so that uropodal rami, which are subequal
in length, extend beyond the distal spines.
Judging from the rest of the posterior pe-
reonal segment, the lateral spines in M. ant-
arcticus are directed backwards, while in
M. caecus, they are directed forwards and
curved, and the dorsal surface of these seg-
ments bears spines. The operculum and oth-
er pleopods of the female, which are sche-
matically drawn by Vanhoffen in his figure
71d, do not seem useful in his opinion and
cannot be used in comparison because the
male was known only for the northern
species.

Vanhoffen (1914) put the genus Micro-
protus in his family Jolellidae (sic), where
he also assigned the genera Jolella (sic, cor-
rectly spelled Jolella Richardson), Janthop-
sis (sic, correctly spelled Janthopsis Bed-
dard), Acanthaspidia Stebbing, Jolanthe (sic,
correctly spelled Jolanthe Beddard), Rhac-

Fig. 10. Microprotus acutispinatus, male holotype: A, Pleopod I, ventral view; B, Pleopod II, dorsal view;
C, Pereopod I; D, Pereopod II; E, Pereopod VII; F, Uropod. Female paratype: G, Pleopod II. Scale bars: 0.2

mm—A, B, F; 0.3 mm—C; 0.5 mm—D, E, G.

360

ee}

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. it.

Microprotus acutispinatus, paratypes, body fragments: A, Female, cephalon and pereonites 1-3,
dorsal view; B, Male, cephalon and pereonites 1-3, lateral view; C, Female, cephalon and pereonites 1-3, lateral

view; D, Female, pleotelson and pereonites 6—7, lateral view; E, Male, pleotelson and pereonites 5-7, lateral
view; F, Female, pleotelson and pereonites 6—7, dorsal view.

VOLUME 102, NUMBER 2

ura Richardson, and Jaerella Richardson.
These genera were earlier referred to the
family Janiridae, owing to the presence of
a more or less distinct rostrum, elongate
lateral lobes on the body segments, and two
or more lateral spines on the abdomen. In
Vanhoffen’s opinion, the anterior body part
of Microprotus is most similar to that of
Tolanthe, but in the last case, the abdomen
lacks distinct distal lobes. At the same time,
Vanhoffen noted certain similarities be-
tween Microprotus and the Munnopsidae,
to which he also assigned the Eurycopidae:
similar structure of long antennae, which he
attributed to the deep-sea mode of life. Van-
hoffen’s comments are important, because
although Microprotus does not belong to his
family “‘Jolellidae,’’ his family concept
clearly indicates that certain of the deep-sea
*‘Janiridae,”’ Jolella at least, may need to be
recognized as belonging to a separate family
with its name corrected to Iolellidae.

Acknowledgments

This paper had its inception when Kus-
sakin asked Wilson to examine Microprotus
caecus to clarify its relationships with the
new species collected during Russian ex-
peditions that are described in this paper.
We would like to thank Dr. Thomas E. Bow-
man, National Museum of Natural History,
for loaning the type material of Microprotus
caecus, carefully reading the manuscript, and
offering suggestions for its improvement. We
are also grateful to Dr. Boris Mezhov, Zoo-
logical Museum of the University of Mos-
cow, for giving us an opportunity to observe
Storthyngura paradoxa specimens identi-
fied by him. This research was partially sup-
ported by National Science Foundation Sys-
tematic Biology Grants BSR 82-15942 and
BSR 86-04573 to Wilson.

361

Literature Cited

Birstein, Ya. A. 1957. Certain peculiarities of the
ultra-abyssal fauna with the example of the ge-
nus Storthyngura. —Zoologicheskii Zhurnal,
36(7):961-985.

1970. Additions to the fauna of Isopods
(Crustacea, Isopoda) of the Kurile-Kamchatka
Trench. Part I. Jn Fauna of the Kurile-Kam-
chatka Trench and its environment.—Akade-
miya Nauk CCCP, Trudy Instituta Okeanologii
Im. P. P. Schirshova 86:292-—340. (In Russian,
English Translation: Israel Program for Scien-
tific Translations, Jerusalem, 1972)
Haugsness, J. A., & R. R. Hessler. 1979. A revision

of the subfamily Syneurycopinae (Isopoda:
Asellota: Eurycopidae) with a new genus and
species (Bellibos buzwilsoni).—Transactions of
the San Diego Society of Natural History 19(10):
121-151.

George, R. Y., & R. J. Menzies. 1968a. Species of
Storthyngura (Isopoda) from the antarctic with
descriptions of six new species.—Crustaceana
14(3):275-301.

——-;, & . 1968b. Distribution and probable
origin of the deep-sea isopod genus Storthyn-
gura. —Crustaceana 15(2):171-187.

Richardson, H. 1910. Isopods collected in the North-
west Pacific by the U.S. Bureau of Fisheries
steamer ‘Albatross’ in 1906.— Proceedings of the
United States National Museum 37(1701):75—
129.

Vanhoffen, E. 1914. Die Isopoden der Deutschen
Sudpolar-Expedition 1901-1903.— Deutschen
Sudpolar-Expedition 15 Zoology, 7:447-598.

Wilson, G. 1989. Deep-sea Lipomerinae and the
Munnopsidae (Crustacea: Isopoda: Asellota): a
systematic revision. — Bulletin of the Scripps In-
stitution of Oceanography 27:i—xiii, 1-138.

Wolff, T. 1962. The systematics and biology of bathy-
al and abyssal Isopoda Asellota.—Galathea Re-
port 6:1-320.

(GDFW) A-002, Scripps Institution of
Oceanography, La Jolla, California 92093,
USA; (OGK & GSV) Institute of Marine
Biology, Far East Science Center, Academy
of Sciences of the USSR, Vladivostok
690022, USSR.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 362-374

FOUR SPECIES OF SYNOPIIDAE FROM THE
CARIBBEAN REGION (CRUSTACEA: AMPHIPODA)

J. L. Barnard and James Darwin Thomas

Abstract.— Synopia ultramarina is redescribed from the Gulf Stream and the
Florida Keys and comments are made on the tangled taxonomy in Synopia.
Tiron bellairsi, originally described from Barbados, is reported from Belize, at
the opposite side of the Caribbean Basin. Garosyrrhoe bigarra, a Pacific species,
is also reported from Belize, but no clear subspecific differences are detected
as yet between Pacific and Atlantic populations.

Specimens of Synopia ultramarina Dana
(1853) are described and illustrated from
the Gulf Stream and Synopia scheeleana
Bovallius (1886) is also reported. A key to
seven species of the genus Synopia was giv-
en by J. L. Barnard (1972); Andres (1984)
described 2 additional species. Four of the
nine species remain poorly described: S. an-
gustifrons Dana (1853, tropical Pacific), S.
caraibica Bovallius (1886, Venezuela), S.
gracilis Dana (1853, tropical Atlantic) and
S. orientalis Kossmann (1880, Red Sea).
Species of this genus are rarely reported in
modern times, perhaps because they occur
mainly in neritic waters or in epipelagic
waters of the high seas. Collections made in
neritic and epipelagic waters rarely are ex-
amined by gammaridean taxonomists, who
appear to concentrate on benthic samples.
Two of the species come from the tropical
Pacific where little neritic collecting has been
accomplished this century. Individuals of
Synopia may actually be mostly nighttime
emergents into neritic waters though they
appear to occur in daytime in epipelagic
waters particularly in or near sargasso-like
flotsam. We have searched for S. caraibica
in neritic waters in the daytime in Florida,
Belize and Trinidad, to no avail. Pereopods
5-7 of that species have ovate article 2 of
pereopods 5-7 but otherwise the species is
well enough described by Bovallius to sug-
gest 1t is not a species of Jiron, a better
known genus than Synopia.

Synopiidae
Synopia Dana

Synopia Dana, 1852:315 (Synopia ultra-
marina Dana, 1853, selected by J. L. Bar-
nard, 1969b).—J. L. Barnard, 1972:50.

Diagnosis. — Forehead protuberant (pos-
sible exception, S. orientalis); lateral ce-
phalic lobe not sharp. Eyes and accessory
eyes present. Antenna | of female shorter
than pereon, of male much longer than pere-
on. Mandible with huge palp (mainly article
2), molar large, triturative or swollen, pil-
lowlike in shape and poorly triturative (S.
variabilis). Outer plate of maxilla 1 with eight
or nine spines. Maxilliped inner plate lack-
ing large smooth spines; outer plate fur-
nished medially only with plumose setae
(no inflexible spines).

Coxae 1-2 small, similar, truncate dis-
tally; coxae 3-4 pelagont, coxa 3 very large
and enfolding small coxa 4. Gnathopods
simple (propodus of gnathopod 1 often
tumid), defining spines absent; gnathopod
2 very slender, dactyl tiny. Pereopods 3—4
slightly to strongly diversified, articles 4—5
stout; pereopods 5-7 elongate.

Pleonites 1-6 neither denticulate nor
toothed. Uropod 3 exceeding uropods 1-2,
peduncle short. Telson relatively short for
family.

Relationship. — Differing from TJiron in the
shorter peduncle of antenna 1, thick man-

VOLUME 102, NUMBER 2

dibular palp, presence of only 8—9 spines on
the outer plate of maxilla 1, the absence of
large smooth apical spines on the inner plate
of the maxilliped, the outer plate bearing
only medial plumose setae, no smooth
spines; the much enlarged coxa 3; inflated
propodus of gnathopod 1; tiny dactyl of
gnathopod 2; diversity and tumidity of pe-
reopods 3-4; lack of dorsal body teeth; and
relatively shorter telson.

Remarks. —We believe that Barnard
(1965) misrepresented the mandibular mo-
lar of S. variabilis as being non-triturative;
he apparently did not turn the molar far
enough upward to see the triturative sur-
face.

Key to the Species of Synopia

_. JES LS i 2
= 2117 00S ee 4
2. Apex of telson broad and armed with

3 widely spread spines ...... rotunda

— Apex of telson narrow and armed
with 2 or fewer appressed spines .. 3
3. Telson with 2 notches separated by
subtruncate protrusion, bearing only
setules variabilis
— Telson with single apical notch

armed with 2 spines ....... triangula
4. Forehead not protuberant .. orientalis
—eenoneag protuberant ........... 5

5. Article 2 of pereopods 5-6 rectan-
gular or with quadrate posteroven-

i Sto SE ae 6
— Article 2 of pereopods 5-6 ovate or

with rounded posteroventral cor-

SES |. a 7

6. Article 2 of pereopod 5 narrowly
rectangular, scarcely twice as broad
as article 3 and with scarcely pro-
truding posteroventral corner ....
_ 2 fe angustifrons
— Article 2 of pereopod 5 widely rect-
angular, nearly twice as broad as ar-
ticle 3 and with strongly protruding
posteroveniral Corner ........ gracilis

363

7. Flagellar articles 2, 3,4...n ofan-
tenna | with long “‘hairs’’* .. caraibica
— Flagellar articles 2, 3,4...nofan-
tenna | lacking long “‘hairs”’
8. Telson scarcely longer than broad,
apicolateral margins smooth, each
apex with one stout spine .. scheeleana
— Telson much longer than broad,
apicolateral margins crenulate, each
apex with one stout spine and one
Femeiseed ys oes. 28002 ultramarina

Synopia ultramarina Dana
Fig. 1-4, part

Synopia ultramarina Dana, 1853:995, pl.
68, fig. 6a—h.— Bovallius, 1886:613, pl. 1,
figs. 1-21.

Diagnosis.—Telson much longer than
broad, deeply cleft, apicolateral margins
scalloped-serrate, each apex with one long
seta, one short setule, medial subapex with
spine in notch. Forehead protuberant, with
anterodorsal keel. Article 2 of pereopods 5—
6 broadly rounded posteroventrally, of pe-
reopod 7 subrectangular and quadrate pos-
teroventrally. Flagellar articles 1, 2, 3 on
antenna 1 of both sexes with at least one
stout spine, short in male, elongate in fe-
male.

Description of female “u’’ 4.53 mm.—
Head with low anterior keel: rostrum ex-
tending downward to shallow keel connect-
ed to epistome, upper lip weakly bilobate
from lateral view and bilobate from anterior
view. Eyes purple in alcohol, pigment dif-
fuse (omitted in drawings), accessory eye of
2 ommatidia each stained purple on inner
apices.

Accessory flagellum 2-articulate, basal ar-
ticle elongate, armed with stiff spines; pri-
mary flagellum short, 8-articulate, articles
1-5 and 7 each with stiff spine, spines longer
on articles 1—3 than on 5 and 7. Each incisor

* Structure unknown, possibly setae, aesthetascs, or
straw-like scales.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

364

x
‘en rt ft

625

aaa

VOLUME 102, NUMBER 2

and left lacinia mobilis with 4 teeth, mght
lacinia mobilis bifid and weakly hook-
shaped, larger rakers about 4 left, 7 right,
first 2 or 3 when turned to plane view bifid;
molar heavily triturative, armed basally with
large brush of apically feathered setae. Inner
plate of maxilla 1 with 5 large medial setae
and 2 short apical setae, outer plate with 8
spines, apex of palp with 5 tooth-spines (one
illustration showing 4 as aberrancy), medial
oral face with groups of 3 and 4 apical setae.
Two principal apical spines on inner plate
of maxilliped plumose, with two other api-
cal setae, one apicoventral seta, two medial
setae and one ventral locking spine; medial
margin of outer plate with marginal and
ventral submarginal rows of plumose setae,
lateral margin with two long setae, one ap-
icad, the other strongly subapical, lateral
base raggedly serrate; article 2 of palp evenly
and densely setose medially; dactyl short,
nail elongate.

Many setae on gnathopod 2 apically
forked; dactyl with shriveled appearance (but
this appearance normal). Pereopods 3-4
with three unlocking setae each, pereopods
5-7 with two unlocking spines each, short
and simple on pereopod 7, on pereopods 5-—
6 one spine elongate and with apical hooded
hook. Gills large, unpleated, on coxae 2-7;
oostegites, thin, strap-shaped, sparsely se-
tose, on coxae 2-5.

Epimera naked below, epimeron | sharp-
ly quadrate, epimeron 2 with small tooth
and epimeron 3 round-quadrate postero-
ventrally. Each pleopod with two coupling
hooks and one plumose accessory spine, pe-
duncles with 1-3 setae only, rami with 12-
14 articles, inner rami with one fewer article

—_

Fig. 1.

365

than outer rami and with forked basomedial
spines, generally two.

Urosomite 3 with posterodorsal spine.
Outer rami of uropods 1-2 with dorsolateral
comb, similar comb on inner rami ventro-
lateral. Telson scarcely longer than pedun-
cle of uropod 3, cleft about two-thirds, cu-
ticle covered with notch-like lunes.

Male “q’’ 5.84 mm.—Antenna 1 about
4.0 mm long, antenna 2 about 5.9 mm long.
Eyes slightly larger than in female. Basal
article of primary flagellum slightly longer
than in female and callynophore denser; ar-
ticles of primary flagellum with extra mid-
ventral male sensory setules, large spines of
female reduced in size. Dorsal margins of
articles 4—5 of peduncle of antenna 2 plus
articles of flagellum bearing dense male sen-
sory setules. Epimera and pleopods larger
than in female, epimeron 2 lacking tooth.
Uropod 3 more setose than in female.

Intermediate male “‘s’’ 4.33 mm.—Eyes
scarcely enlarged. Antenna 1 elongate but
basal spines of flagellum elongate as in fe-
male. Posterior setules on flagellum of an-
tenna 1 and anterior setules on antenna 2
poorly developed. Tooth of epimeron 2 in-
termediate between male and female. Setae
of uropod 3 moderately developed.

Illustrations. —Maxilla 1 magnified more
than maxilla 2; telson magnified more than
uropod 3.

Material. —MFP-1, in Gulf Stream, off
Grand Bahama Island, 27°26'N, 78°57’'W,
219 m, 8 Jul 1986, coll. Jack Morton, at-
tracted to night light, female “‘p” 4.35 mm,
male “‘q’’ 5.84 mm (illustrated), female “‘r”’
4.40 mm, male “s’’ 4.33 mm, male “‘t”’ 4.79
mm, female “‘u’’ 4.53 mm (illustrated) (and

Synopia ultramarina, unattributed figures = female “‘u’’; q = male “‘q.”’ Capital letters as follows refer

to parts; lower case letters to the left of capital letters refer to specimens noted in legends; lower case letters to
right of capitals refer to adjectival modifications in list below: B, body; D, dactyl; G, gnathopod; H, head; I,
inner plate or ramus; J, prebuccal; L, labium; M, mandible; O, outer plate or ramus; P, pereopod; R, uropod;
S, maxilliped; T, telson; U, labrum; V, palp; W, pleon; X, maxilla; a, anterior; d, dorsal; 0, opposite; r, right;

s, setae removed; t, left; x, apex.

366

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ee

Fig. 2. Synopia ultramarina, unattributed figures = female “u”; q = male “q.” Letter codes, see Fig. 1.

VOLUME 102, NUMBER 2 367

eee 99.

u’’; q = male “q.”’ Letter codes, see Fig. 1.

368

6 other specimens). LKFR-5, Florida Keys,
Looe Key Reef, 7 m, sand plain in front of
fore reef zone, 9 Oct 1983, 2300 h, night
light, full moon, coll. J. D. Thomas (35 spec-
imens).

Remarks. —Said by Dana (1853) to be
“rich blue, to nearly colorless, with tinge of
rich blue along the venter or about the ar-
ticulations’’; our specimens and those of the
first redescriber, Bovallius (1886), however
were hyaline with yellowish cast when caught
alive, mouthparts, especially maxillipeds
with red cast under microscopy seen to be
represented by tiny red granular inclusions
below the chitinous exoskeleton. One may
wonder if Dana might not also have col-
lected S. scheeleana; he called the “‘male
specimen”’ of S. u/tramarina ““ultramarine”
but labeled it on his plate as S. gracilis and
changed its identification in the published
text; S. gracilis has since been recognized as
a distinct species but has never been clari-
fied adequately; unfortunately Dana’s ma-
terial is presumed to have been lost in the
great fire of Chicago, 1871; see other notes
below.

Despite the color, Bovallius chose this
species to represent Dana’s u/tramarina on
the basis of serrate telson and conformity
in pereopods 5-7; we agree that this solution
is correct on morphological grounds.

Relationship. — Differing from the well
defined S. scheeleana Bovallius (1886) in
the longer telson bearing apicolateral ser-
rations and at least one long apical seta on
each lobe.

Synopia caraibica Bovallius (1886) has
article 2 of pereopod 7 ovate like that of
pereopods 5-6. Synopia angustifrons Dana
(1853) and S. gracilis Dana (1853) have ar-
ticle 2 of pereopods 5-7 rectangular or with
quadrate posteroventral corners. There is
the possibility that Dana misinterpreted
these legs as there is no indication he dis-
sected them off the animal and pressed them
flat. Synopia orientalis Kossmann (1880)
does not have a protuberant forehead. The
telson of S. variabilis Spandl (1923, see J.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

L. Barnard, 1965), S. triangula Andres
(1984) and S. rotunda Andres (1984) is un-
cleft.

Distribution. —Tropical western Atlantic;
specific localities include Cuba, Bahamas,
Florida Keys and Dana’s 8-12°S, 11l-
144%4°W, 4-7°S, 21—25°W; neritic and pre-
sumably epipelagic on the high seas; attract-
ed to night lights.

Synopia scheeleana Bovallius

Synopia scheeleana Bovallius, 1886:16-18,
pl. 2, figs. 22—-29.—Stebbing, 1888:799-—
804, pl. 52.—Chevreux, 1900:54.— Reid,
1951:233.

Material.—Gulf Stream, 40 miles off
Florida coast between Miami and Fort
Pierce, 10 Dec 1987, R/S Sea Diver, male
“a” 2.52 mm, specimen “‘b”’ 1.80 mm.

Remarks. —Live color deep blue to s-blue,
nos. 178 to 179, as matched to ISCC-NBS
Centroid Color Chart 2106; we described
this in the laboratory as cobalt blue; spec-
imens courtesy of Dr. Mary E. Rice, Direc-
tor, Smithsonian Research Station at Link-
port.

Distribution. — Atlantic Ocean, off Ft.
Pierce, Florida to Barbados; tropical Pacific
Ocean; assumed to be epipelagic.

Tiron bellairsi Just
Fig. 4, part

Tiron bellairsi Just, 1981:259-263, figs. 14.

Diagnosis. — Accessory eye composed of
2 and | separated ommatidia; accessory fla-
gellum 2-articulate; labrum densely spi-
nose; palp of mandible replaced by seta; in-
ner plate of maxilla 1 long and thin, fully
setose medially but setae small and widely
spread, outer plate with 10 spines forming
elongate multi-sigmoid cluster, palp re-
duced, about half as long as outer plate; in-
ner plate of maxilla 2 with row of submar-
ginal setae; inner plate of maxilliped lacking
thick simple spines; outer plate of maxil-

VOLUME 102, NUMBER 2 369

ST OX1

Fig. 4. Left upper = Synopia ultramarina, male “‘q.”’ Right, Tiron bellairsi, male “‘o,”’ note 3 renderings of
outer plate on maxilliped. Letter codes, see Fig. 1.

370

liped abnormal, lacking major spines, bear-
ing plumose and tube-setae; dactyls of pe-
reopods 3-7 stubby, nails scarcely curved;
article 2 of pereopods 6—7 without special
setal row, article 2 of pereopod 7 weakly
crenulate and setulose posteriorly; pereon-
ites 5—7 and pleonites 1-3 dorsally crenu-
late, pleonite 4 with medium dorsal tooth,
pleonite 5 with large dorsal tooth, pleonite
6 lacking tooth; outer ramus of uropod 1 as
long as inner ramus but outer ramus of uro-
pod 2 slightly shortened; rami of uropod 3
apically blunt; telson with one apical spine
and 3 setae on each side, no dorsal or medial
spines.

Remarks. —We have redrawn minute de-
tails of several mouthparts and the telsonic
apex; we assume these are congruent with
the Barbados population. The diagnostic
format above is revised to include new char-
acters not used by Barnard (1972).

Material.—JDT Bel-53, Belize, Carrie
Bow Key, 34 m, sand trough near fore reef
drop-off, in coralgal mud, 11 Jun 1980, coll.
J. D. Thomas, male “o” 2.49 mm (illus-
trated), 2juvenile “‘p” 1.96 mm.

Distribution. — Barbados, Holetown (type-
locality), 5—? m (‘SCUBA”’ depths); Belize,
Carrie Bow Cay, 34 m.

Garosyrrhoe bigarra (J. L. Barnard),
new synonymy
Figs. 5-6

Syrrhoites bigarra J. L. Barnard, 1962:73-
1S USN

Garosyrrhoe disjuncta J. L. Barnard, 1969a:
224-225, fig. 30.

Taxonomy.—Syrrhoites bigarra and Ga-
rosyrrhoe disjuncta are the male and female
of the same species. The male is character-
ized by the larger head with blunter ros-
trum, elongate antenna 2 and more aes-
thetascs and male setae on antennae 1-2.
Barnard (1962) overlooked the complex na-
ture of the dorsal teeth on pereonite 7 to
pleonite 3.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

The Caribbean specimens are smaller than
those from the eastern Pacific, where the
species has been found between about 25°
32°N. As yet no specimens from the Pan-
amanian isthmus region have been found
and until comparisons between specimens
from exactly similar latitudes and thermal
regimes from Pacific and Atlantic waters can
be compared, the small differences we find
in the Atlantic specimens cannot be attrib-
uted to speciation.

Both sexes of material from Belize have
fewer serrations on the epimera and the fe-
male lacks serrations on article 6 of pereo-
pod 4. Other differences between bigarra
and disjuncta cited by Barnard (1969a) are
inconsistencies lacking specific value as
demonstrated in the present material and
reexamination of the earlier materials. For
example, the ventral spine on the inner plate
of the maxilliped and the dorsally promi-
nent pleonal teeth were overlooked in var-
ious specimens; differences among pereo-
pods are simply variables. Any of the
differences, albeit minor, between individ-
uals from different oceans, could be attrib-
uted to the smaller adult size of Caribbean
specimens. Small potential distinctions of
specific value between Atlantic and Pacific
populations may be present in articular pro-
portions and armament patterns but con-
firmation of these requires study of larger
suites of Pacific material than are now avail-
able. The following descriptions contain only
new information and points we judge to be
necessary to amplify and rectify the former
depiction of this genus and species; illustra-
tions are provided for characters which are
difficult to visualize from descriptions.

Description of male “‘x’’ 3.12 mm.—Pe-
duncular articles 1—2 of antenna 1 with nu-
merous dorsal aesthetascs, multiple aesth-
etascs also present on articles 1-2 of
accessory flagellum and article 1 of primary
flagellum; on antenna 2 similar aesthetascs
on article 4 of peduncle, aesthetascs on ar-
ticle 5 apically curled, basal articles of fla-
gellum also with multiple aesthetascs.

VOLUME 102, NUMBER 2 a7

Fig. 5. Garosyrrhoe bigarra, unattributed figures = male “x”; w = female “‘w.” Letter codes, see Fig. 1.

S72

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Fig. 6.

Garosyrrhoe bigarra, male “*x.”” Letter codes, see Fig. 1.

VOLUME 102, NUMBER 2

Right and left mandibular incisors with
three large and one small teeth, each lacinia
mobilis with five teeth, first raker on each
side broadened, more so on left than on
right mandible, right and left mandibles each
with total of two and three rakers. Outer
plate of maxilla 1 with 11 spines (versus 9
in Pacific material), inner plate with 6 me-
dial setae, palpar apex with 4 spines, one
apicolateral seta, 2 mediofacial setae. Inner
plate of maxilliped with 5 subapical setae,
2 medial setae, one apicomedial spine, one
ventral hooked locking spine; outer plate
with 8 marginal spines and one apical spine-
seta.

Articles 4-5 of pereopod 5 more sparsely
armed than in Pacific male (pereopods 6-7
in Pacific male unknown). Dactyl of pereo-
pod 7 lacking comb present on pereopods
3-6. Serrations on epimera 1,2,3 = 2-4-9
(versus 4-7-12 on Pacific male). Details of
one dorsal pleonite plus uropod 3 and telson
are shown in illustrations.

Female “w’” 3.40 mm (body slightly con-
tracted). —Serrations on epimera 1,2,3 =
2-2-9 (versus 5-6-13 in Pacific female).

Material.—JDT BEL-53, Belize, Carrie
Bow Cay, 11 Jun 1980, forereef sand trough,
29 m, coralgal sand-mud, coll. J. D. Thom-
as, female “w”’ 3.40 mm, male “x”? 3.12 mm
and 4 other specimens. Also found in sim-
ilar samples: JOT BEL-9B,C,D; JDT BEL-
14; -19; -53. Associated in amphipod com-
munity on coralagal sediments at depths of
27 to 33 m with Netamelita and Hornellia.

Distribution. —Southern California at
border with Mexico and in Gulf of Califor-
nia, 24-44 m; Caribbean Sea, Belize, 27-
33 m.

Acknowledgments

We thank Captain Jack Morton and stu-
dents of the Marine Field Projects class of
1986, Florida Institute of Technology, Mel-
bourne, Florida, for collecting amphipods
while at sea. The second author was sup-
ported by NSF grant BSR-8515186.

373

Literature Cited

Andres, H. G. 1984. Zwei neue Synopiiden (Crus-
tacea: Amphipoda: Gammaridea) aus dem war-
men zentralen Nordatlantik. — Mittheilungen aus
den Hamburg Zoologisch Museum und Institut
81:109-116.

Barnard, J. L. 1962. Benthic marine Amphipoda of

southern California: Families Tironidae to

Gammaridae— Pacific Naturalist 3:70-115.

1965. Marine Amphipoda of atolls in Mi-
cronesia. — Proceedings of the United States Na-
tional Museum 117:459-552.

1969a. A biological survey of Bahia de los
Angeles Gulf of California, Mexico, IV. Benthic
Amphipoda (Crustacea).— Transactions of the
San Diego Society of Natural History 15:175-
228.

1969b. The families and genera of marine
gammaridean Amphipoda.— United States Na-
tional Museum Bulletin 271:535 pp. 173 figs.

1972. A review of the family Synopiidae

(=Tironidae), mainly distributed in the deep sea

(Crustacea: Amphipoda).—Smithsonian Con-

tributions to Zoology 124:1-94.

Bovallius, C. 1886. Amphipoda Synopidea.— Nova
Acta Regiae Societatis Scientiarum Upsaliensis
13:1-36.

Chevreux, E. 1900. Amphipodes provenant des cam-
pagnes de /’Hirondelle (1885-1888). — Résultats
de Campagnes Scientifiques Accomplies par le
Prince Albert I Monaco 16:iv and 195 pp., 18
pls.

Dana, J. D. 1852. On the classification of the Crus-

tacea Choristopoda or Tetradecapoda. — Amer-

ican Journal of Science and Arts (2) 14 [Appen-

dix]:297-316.

. 1853. Crustacea. Part II.— United States Ex-

ploring Expedition 14:689-1618, atlas of 94 pls.

published in 1855.

1981. Tiron bellairsi sp. n. (Amphipoda, Sy-
noplidae) from coral sand in Barbados, with
notes on behaviour.— Zoologica Scripta 19:259-
263.

Kossmann, R. 1880. Malacostraca. Zoologische Er-
gebnisse einer im Auftrage KOniglichen Aca-
demie der Wissenschaften zu Berlin ausgefuhr-
ten Reise in die Kustengebiete des Rothen
Meeres. Zweite Halfte, Erste Lieferung. III. Ma-
lacostraca 67-140, pls. 4-15. Leipzig: Wilhelm
Engelmann.

Reid, D. M. 1951. Report on Amphipoda (Gam-
maridea and Caprellidea) of the Coast of Trop-
ical West Africa. — Atlantide Report 2:189-291,
58 figs.

Spandl, H. 1923. Amphipoden der ‘Pola’-Expediti-
onen in das Rote Meer.—Akademie der Wis-

Just. Ff

374 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

senschaftlichen in Wien, Anzeiger, Jahrgang 60:
17-20, 87-89, 111-112.

Stebbing, T. R. R. 1888. Report on the Amphipoda
collected by H.M.S. Challenger during the years
1873-76. Report on the scientific results of the
voyage of H.M.S. Challenger during the years
1873-1876, Zoology 29: xxiv and 1737 pp.,
210 pls.

(JLB) NHB-163, Division of Crustacea,
Smithsonian Institution, Washington, D.C.
20560; (JDT) Newfound Harbor Marine In-.
stitute, Rt. 3, Box 170, Big Pine Key, Flor-
ida 33043.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 375-384

A NEW SPECIES, AMPELISCA BURKET,
(CRUSTACEA, AMPHIPODA) FROM FLORIDA

J. L. Barnard and James Darwin Thomas

Abstract. — A new species Ampelisca burkei, from Florida, is described. This
species appears to be very close to A. /obata Holmes, from the eastern Pacific
Ocean, but differs in the shape and setation of article 4 on pereopod 7, and the
presence of an anteroventral spine-seta on article 5 of pereopod 7.

The new species, Ampelisca burkei de-
scribed herein is a twin of A. Jobata Holmes
from the eastern Pacific. The differences be-
tween the two species are very small but
recognition of both is justified by the trend
for taxonomists to find and validate at the
specific level small differences between Pa-
cific and Atlantic amphipods. Small mor-
phological characters are extremely consis-
tent over wide geographic areas in the family
Ampeliscidae. This consistency is not widely
shared by amphipod groups, and care must
be taken to document variation in particular
characters before incorporating them into a
taxonomic treatment. For example, Bar-
nard (1980) separated the Pacific Methar-
pinia oripacifica from the Atlantic M. flor-
idana by the presence or absence of a single
spine on the inner plate of the maxillipeds
(2 spines in Pacific, one in Atlantic popu-
lations). Dickinson (1982) separated the Pa-
cific Ampelisca fageri from the Atlantic A.
schellenbergi on subtle shapes and setation
patterns of coxa 1, pereopod 7 and uropod 3.

Ampelisca burkei, new species
Figs. 1-5

?Ampelisca lobata.—J. L. Barnard, 1954b: 2
(not Holmes, 1908, possible misidenti-
fied specimens from Colombia and Aru-
ba).

Description of holotype, male, 4.92 mm.—
Body smooth; urosomite | elevated, with

thick keel, urosomites 2—3 coalesced, with
strong dorsal saddle.

Head as long as first three pereonites com-
bined, slightly longer than tall, with short
rostrum; head with lobe below attachment
of antenna 1, then head sloping down and
posteriorwards in two steps, no ventral
tooth. One corneal lens present on each side
strongly removed from anterior margin,
second pair on ventral margin far below lat-
eral lobe. Brown (unusual) pigmentary mass
present for each lens, similar mass behind
upper pair.

Antenna | slender, much shorter than an-
tenna 2, reaching 25 percent along flagellum
of antenna 2; peduncle short, articles 1—2
equally long, article 3 short; base of flagel-
lum forming callynophore; accessory fla-
gellum absent; main flagellum 5 times as
long as peduncle, first 2 articles with aes-
thetascs attached ventrally. Antenna 2 about
2.4 times as long as antenna 1, about 1.4
times as long as body, peduncular article 5
almost as long as 4, anterior margins of ar-
ticles 3—5 with male setular tufts, flagellum
3 times as long as peduncle.

Epistome weakly and obtusely projecting
anteriorly, labrum incised distally, broader
than long (tall). Labium normal, with well-
developed inner lobes.

Mandible with well-developed strong tri-
turative molar, incisor toothed; palp 3-ar-
ticulate, article 1 short, article 2 longer than
3, setose, article 3 non-falciform, not dilated
distally, with 3 D setae, 4 E setae.

Maxilla 1: inner lobe with 2 short simple

376 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

7 et yn ee
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VOLUME 102, NUMBER 2

setae, outer lobe with 11 spines, three of
these provided with 2-4 lateral teeth each;
palp 2-articulate, symmetrical on both sides,
second article barely dilated distally, pro-
vided with 4 distal sharp cusps accom-
panied by 4 spines, 4 subdistal setae and 2
apicomedial setae. Maxilla 2 with narrow
lobes, inner lobe shorter than outer, without
oblique facial row of setae, all medial setae
marginal. Maxilliped: inner lobe of ordinary
length, with 2 distal spines intermixed with
several plumose setae; outer lobe reaching
apex of palp article 2, bearing row of thick
spines along medial margin and 2-3 apical
setae; palp 4-articulate, article 3 not lobed,
article 4 with main nail about as long as
remaining part of article.

Coxae 1—4 much longer than broad, coxa
1 dilated distally, with convex distal (=ven-
tral) margin provided with one row of tiny
setae, lacking posteroventral notch; coxae
2—4 with increasingly truncate distal mar-
gins, coxae 2—4 with increasingly sparse set-
ules, coxa 4 with small posteroproximal lobe
produced bluntly in proximal part, other-
wise apposite margins almost parallel.

Gnathopods 1-2 linear, simple, gnatho-
pod | shorter than gnathopod 2; gnathopod
1: article 5 scarcely longer than 6, densely
setose along posterior margins, article 6 ta-
pering distally with dactyl shorter than ar-
ticle 6, bearing 5 setules along inferior mar-
gin and | seta on outer margin. Gnathopod
2: article 5 linear, long, moderately setose;
article 6 much shorter than 5, tapering dis-
tally, dactyl like that of gnathopod 1.

Pereopod 3 of medium stoutness, with
articles 4-6 bearing long plumose setae along
both margins, except posteriorly on article
6 and anteriorly on article 5; article 4 not
inflated; dactyl slender, almost straight,

, eel
Fig. 1:

377

slightly longer than article 6. Pereopod 4
like that of pereopod 3, but much longer,
article 4 setose on both sides to base.

Pereopod 5: article 2 ovoid, with large
posterior flangehump, anterior margin with
one row of plumose setae, article 5 scarcely
produced distoposteriorly, bearing row of
spines on apex; article 6 with few long distal
setae, dactyl short, unguiform, with 2 outer
cusps. Pereopod 6: article 2 subquadrate,
with strong posterior lobe and poorly setose
anterior margin; articles 3-7 like those of
pereopod 5. Pereopod 7: almost as long as
pereopod 6 but article 2 large, scarcely ex-
panding distalwards, lobe reaching middle
of article 4, bearing sparse plumose setae
along ventral margin in two disjunct groups;
article 3 short; article 4 slightly longer than
3, weakly produced at posterodistal apex
and bearing one short spine and posterior
setae; article 5 narrow, with weak postero-
distal spination and one anterior subdistal
spine-seta, article 6 weakly tumid, scarcely
longer than 5, dactyl tumid, tapering rapidly
and apically pointed, much shorter than ar-
ticle 6.

Strongly plaited large gills on coxae 1-6,
gills generally with 20+ pleats. Pleopods well
developed, normal, with 2 retinacula and
one simple accessory retinaculum each.

Epimera 1-2 weakly convex behind,
epimeron 1 with posteroventral setule at
subsharp corner, epimeron | with 3 ventral
setae; epimeron 3 not larger than 2, poste-
rior margin almost straight, posteroventral
corner sharply protuberant or obtuse (vari-
able).

Uropods 1-2 of ordinary length, uropod
1: peduncle slender, outer face of peduncle
with 5 spines, dorsolateral margin with 3
spines, apex with cusp, medial corner with

Ampelisca burkei, n. sp., unattributed figures, holotype male “h”’ 4.92 mm; k = male “k” 4.66 mm.

Capital letters denote main parts in following list; lower case letters to left of capital letters or in body of figure
indicate modifications as per following list; lower case letters to right of capital letters indicate specimens described
in captions: A, antenna; C, coxa; D, dactyl; E, epimera; G, gnathopod; H, head; L, lower lip; M, mandible; P,
pereopod; R, uropod; S, maxilliped; T, telson; U, upper lip; W, pleon; X, maxilla; m, medial.

378

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Ampelisca burkei, n. sp., holotype male “h” 4.92 mm. Letter codes, see Fig. 1.

VOLUME 102, NUMBER 2 379

Fig. 3. Ampelisca burkei, n. sp. Upper, holotype male “‘h” 4.92 mm. Lower, female “‘o” 5.66 mm. Letter
codes, see Fig. 1.

380 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Ampelisca burkei, n. sp., unattributed figures = female ““m” 4.40 mm; h = holotype male “h” 4.92
mm. Arrows on hA2 and mH point to lines denoting undrawn lengths of antennae. Letter codes, see Fig. 1.

Wie at

Noyiratesar inurl) 6)

> roy

"Waray: 7

Fig. 5. Ampelisca burkei, n. sp. Upper, female ““m’ 4.40 mm. Lower, female “‘o” 5.66 mm. Letter codes,
see Fig. 1.

382

2 spines, medial margin with 5 other spines,
rami slender, as long as peduncle, curved
and pointed distally, outer with 2 tiny lat-
eral spines, medial margin with 4 spines,
inner ramus lacking lateral spines, with 7
medial spines. Uropod 2: peduncle slender,
slightly shorter than outer ramus, with 3
moderately spread dorsolateral and 3 tightly
packed dorsomedial spines, with apicolat-
eral cusp, outer ramus smaller than inner,
of triangular cross-section, with 4—5 dorsal
spines, 2 basomedial spines and one tiny
lateral spine, inner ramus with 1 lateral
spine, and 7—8 medial spines. Uropod 3 long,
subfoliaceous, strongly exceeding apex of
uropod 2: peduncle stout, short, with 2 me-
dial spines; rami extending subequally, in-
ner foliaceous, with sharp apices, outer much
narrower than inner, setose laterally, and
apicomedially, inner lined with stout me-
dial spines paired with plumose setae, extra
setae in gaps between spines.

Telson reaching one third along rami of
uropod 3, longer than broad, incised 90 per-
cent of its length, each lobe expanding in
middle and then tapering distally, notch
narrowly beveled apically, each with apical
spinule, 2 dorsal axial spines and subbasal
pair plus satellite of penicillate setules (one
spine occasionally substituted by pair of set-
ules).

Male “‘k.’’—Antenna 2 only 0.8 times as
long as in holotype.

Female ““m’’ 4.40 mm.—Oostegites on
coxae 2—5 strap-shaped, narrow, weakly se-
tose. Antenna | slightly exceeding peduncle
of antenna 2, peduncle short, article 2 as
long as 1, flagellum with few basal aesthe-
tascs. Antenna 2 elongate, article 5 of pe-
duncle slightly shorter than article 4. Gills
much smaller than in male and with about
5—6 pleats. Hump on pleonite 4 lower than
in male. Dorsolateral margin on peduncle
of uropod 1 with 3 spines, inner apex with
spine pair, medial margin with 3 other
spines, outer ramus with 2 lateral and 3 me-
dial spines, inner ramus with 3 medial
spines; peduncle of uropod 2 with 2 dor-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

solateral spines, inner apex with triad of
spines, medial margin with one other spine,
outer ramus with 3 dorsal and one medial
spines, inner ramus with 4 medial spines,
other spine positions noted for male with-
out spines. Outer ramus of uropod 3 slightly
shorter than inner, with 5 lateral setal po-
sitions bearing 1—2 setae each, apex notched
and with 2 setae; inner ramus with 4 api-
colateral setules, inner margin with 5 spine-
notch positions. Telson with only one dorsal
spine on each lobe besides one apical spine.

Notes on the redescription of Ampelisca
lobata by Dickinson, 1982.—Dickinson’s
figure 5 of a female from North Bank Island,
British Columbia, does not show details of
pereopod 7 but in the small view presented
lacks posterior setae on article 6 (a pre-
sumed oversight) and lacks the anteroven-
tral seta-spine on article 5 which is present
in the Floridian material and was shown to
be absent by Barnard on males (1954a).
Dickinson does not show the basal aesthe-
tascs on the flagellum of female antenna 1.
Both inner rami of uropods 1—2 of Dick-
inson’s specimen bear one lateral spine but
none is present on Floridian female mate-
rial; we assume this may have connection
with size differences because the British Co-
lumbian material also has many more pe-
duncular and inner marginal rami spines
than our females.

Illustrations. —Left male uropod 3 drawn
ventral side up.

Etymology. —Named for the late William
W. Burke, III, an original ““Son of Bennett”
(honorary society of biologists at Louisiana
State University), who first worked with
Ampelisca in the marshes of Louisiana.

Holotype. —USNM_ 195153, male ‘“h”
4.92 mm, illustrated.

Type locality. —Florida Keys, Looe Key
Reef, forereef, 8 m, Apr 1982, full moon
2200 hours, night diving, attracted to night
light, coll. J. D. Thomas.

Material.—Type locality, male “1” 4.72
mm, male “‘j” 4.84 mm, male “k” 4.66 mm
and 29 other specimens. Females collected

VOLUME 102, NUMBER 2

at same site, LKFR-1C, 18 Apr 1982, from
algal turf community on Acropora cervicor-
nis forereef, 8 m, coll. J. D. Thomas, female
“1”? 4.69 mm, female ““m” 4.40 mm (illus-
trated), female “‘o” 5.66 mm (illustrated)
and 10 other specimens.

Relationship. —Ampelisca burkei is easily
distinguished from such sympatric species
as A. abdita Mills (1964), A. vadorum Mills
(1963) and A. declivititis Mills (1967) by the
short article 3 of pereopod 7. The following
species of Ampelisca bear similarities to A.
burkei: A. agassizi Judd (see Dickinson
1982), A. holmesi Pearse (1908) (see Goeke
& Gathof 1983), A. panamensis J. L. Bar-
nard (1954a), A. parapanamensis J. L. Bar-
nard (1954b), A. verrilli Mills (see Bousfield
1973), A. lobata Holmes (see Dickinson
1982), A. hancocki J. L. Barnard (1954a),
A. milleri J. L. Barnard (1954a), A. cucullata
J. L. Barnard (1954a), and A. romigi = is-
ocornea J. L. Barnard (1954a).

Ampelisca burkei differs: from eastern Pa-
cific A. /Jobata in the weaker expansion and
fewer setae on article 4 of pereopod 7, the
presence of an anteroventral spine-seta on
article 5 of pereopod 7 and the lack of a
protrusion apically on article 5 of pereopod
5 in the female; from A. verri//iin the slender
pereopod 7, short article 2 of antenna 1, lack
of tooth on epimeron 3 and widely expand-
ed head; from A. cucullata and A. agassizi
in the slender articles 2-6 of pereopod 7,
and short article 2 of antenna 1; from A.
romigi in the smaller posteroventral lobe on
article 4 of pereopod 7, less beveled article
2 of pereopod 7, short article 2 of antenna
1 and narrower posterior flange-lobe on ar-
ticle 2 of pereopod 5; and from the male A.
isocornea form by the large spines on female
uropod 3, much larger process of pleonite
4 and lack of facial spines on article 6 of
pereopod 7; from A. hancocki in the shape
of pereopod 7, numerous spines of uropods
1-2, longer antenna | and shape of coxa 4;
from A. milleri in the short article 3 and
general shape of pereopod 7, dense spina-
tion of uropods 1-2, and shape of coxa 4;

383

from A. panamensis and A. parapanamensis
in the broad head and narrow articles 4—5
of pereopod 7; and from A. ho/lmesi in the
short article 2 of antenna 1, narrower pereo-
pod 7 and broader head.

As far as we know, this species differs
from all other American species of Ampe-
lisca in the brown (versus red) pigment of
the eyes; the species therefore looks super-
ficially like a species of Bybiis.

Distribution. —Florida Keys, Looe Key
Reef, 0-8 m.

Acknowledgments

We thank Linda B. Lutz of Vicksburg, for
inking our drawings; Janice B. Clark of
Smithsonian for laboratory assistance; and
the following referees who helped improve
this study: Dr. James K. Lowry of the Aus-
tralian Museum, and Dr. Thomas E. Bow-
man of Smithsonian Institution; and anon-
ymous referees. The junior author was
supported by Grant BSR 8515186 from the
National Science Foundation.

Literature Cited

Barnard, J. L. 1954a. Amphipoda of the family Am-

peliscidae collected in the Eastern Pacific Ocean

by the Velero III and Velero IV.—Allan Han-

cock Pacific Expeditions 18:1—137, pls. 1-38.

. 1954b. Marine amphipoda of Oregon.—Or-

egon State Monographs, Studies on Zoology 8:

103.

1980. Revision of Metharpinia and Micro-
phoxus (marine Phoxocephalidae Amphipoda
from the Americas).— Proceedings of the Bio-
logical Society of Washington 93:104—135.

Bousfield, E. L. 1973. Shallow-water Gammaridean
amphipoda of New England. Ithaca & London,
Cornell University Press, vii—xii, 312 pp., 13
figs., 69 pls.

Dickinson, J. J. 1982. The systematics and distri-
butional ecology of the family Ampeliscidae
(Amphipod: Gammaridea) in the Northeastern
Pacific region I. The genus Ampelisca. —Publi-
cations in Biological Oceanography, National
Museum of Canada 10:1-39.

Goeke, G. D., & J. M. Gathof. 1983. Amphipods of
the family Ampeliscidae (Gammaridea). II.
Notes on the occurrence of Ampelisca holmesi
in the northern Gulf of Mexico. — Gulf Research
Reports 7:289-291.

384 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Holmes, S. J. 1908. The amphipoda collected by the
United States Bureau of Fisheries Steamer, “‘Al-
batross,”’ off the west coast of North America,
in 1903 and 1904, with descriptions of a new
family and several new genera and species. —
Proceedings of the United States National Mu-
seum 35:489-543.

Mills, E. L. 1963. A new species of Ampelisca (Crus-
tacea: Amphipoda) from Eastern North Amer-
ica, with notes on other species of the genus. —
Canadian Journal of Zoology 41:971-989.

——. 1964. Ampelisca abdita, a new amphipod

crustacean from Eastern North America.—Ca-

nadian Journal of Zoology 42:559-575.

1967. The biology of an ampeliscid amphi-

pod crustacean sibling species pair.— Journal of
the Fisheries Research Board of Canada 24:305-—
355:

Pearse, A. S. 1908. Descriptions of four new species
of amphipodous Crustacea from the Gulf of
Mexico. — Proceedings of the U.S. National Mu-
seum 34 (1594):27-32.

(JLB) NHB-163, Department of Inver-
tebrate Zoology, Smithsonian Institution,
Washington, D.C. 20560; (JDT) Newfound
Harbor Marine Institute, Rt. 3, Box 170,
Big Pine Key, Florida 33043.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 385-393

AEGLA DENTICULATA LACUSTRIS, NEW
SUBSPECIES, FROM LAKE RUPANCO, CHILE
(CRUSTACEA: DECAPODA: ANOMURA: AEGLIDAE)

Carlos G. Jara

Abstract.—A subspecies of Aegla denticulata Nicolet, 1849, is described as
A. d. lacustris. The new subspecies is restricted to the preandean Lake Rupanco,
in the River Bueno basin, Osorno Province, Chile, while the nominate sub-
species is widespread in small, moderately fast running streams in the Central
Valley of Valdivia, Osorno, and Llanquihue provinces. A. d. /acustris differs
from A. d. denticulata in having markedly more prominent spinulation along
the branchial margins of the carapace and upon the chelipeds.

Aegla denticulata Nicolet, 1849, is one of
the best known Chilean aeglids. Its mor-
photype is easily distinguished from that of
all other species of Aegla Leach (Schmitt
1942). Extensive collections made by the
author in central southern Chile have shown
that A. denticulata ranges between Cautin
and Chiloé (Jara 1980 and unpublished
data), living in small to medium-sized rivers
along the Chilean Central Valley.

A large population of A. denticulata was
found occupying the sublittoral of Lake Ru-
panco (Osorno Province). When I com-
pared the specimens collected in the lake
with those collected in rivers, it became ap-
parent that the lake specimens differed
markedly in the expression of the carapace
ornamentation. The spininess of the lacus-
trine specimens is remarkably more prom-
inent than that of the riverine animals.

Since this highly ornamented morpho-
type was found to be restricted to Lake Ru-
panco, this variant population was consid-
ered to represent a geographic isolate of A.
denticulata, thus deserving a description as
a new subspecies (as defined by Mayr 1969).

Aegla denticulata lacustris, new subspecies
igs la;cje; ge, 1; komo

Holotype.—Instituto de Zoologia, Uni-
versidad Austral de Chile (IZUA), IZUA

C-686, adult male, Huillin Cove on north-
western shore of Lake Rupanco (40°49’S;
72°28'W), Chile, 25 m depth, baited crab
trap, 30 Apr 1984, coll. C. G. Jara.

Allotype.—Adult female, 8 Sep 1984,
IZUA C-686, same locality, collecting de-
vice and depth as for holotype.

Paratypes. -IZUA C-687; eight adult
males (P1 to P8), 30 Apr 1984; eight adult
females (P10 to P16), 8 Sep 1984; one ovig-
erous female (P9), 3 Apr 1984; same local-
ity, collecting device and depth as for ho-
lotype. Four juveniles (P17 to P20), 7 Sep
1984, 18 m depth, Scuba hand collection.

Diagnosis. —External margin of antero-
lateral lobe of carapace with row of tiny
spinules; summit of middorsal carina with
irregular narrow band of minute scales; dis-
todorsal margin of merus of chelipeds with
transverse row of small but well defined
spines; posterior branchial margin of cara-
pace definitely denticulate; summit of dor-
solateral carina on second abdominal epi-
meron with irregular row of. scales;
laterodorsal area of tergum of second ab-
dominal segment markedly protuberant, its
apex scaly; dorsum of merus and carpus of
second to fourth pereiopods clearly dentic-
ulate; denticles on anterior branchial mar-
gin conical; palmar crest of chelipeds deeply
incised.

Description. —Carapace ovoid, markedly

386

elevated along middorsal line forming well
defined keel extending from rostrum to pos-
terior margin of carapace. Dorsal surface of
carapace and abdomen finely punctate; when
dry, polished and glossy brilliant. Short stiff
setae protrude from punctae, some together
with minute lenslike scales.

Rostrum prominent, straight, broad-
based, extended in horizontal plane but
sometimes slightly recurved at tip. Form
varying, from strictly triangular to ligulate.
Rostral margins well defined and slightly
upturned, fringed with row of closely packed
minute conical scales pointing towards ros-
tral apex. Distal third flattened, merging with
rostral carina into acute conical apex. Ros-
tral carina narrow, sharp, with row of mi-
nute scales not reaching rostral apex. Prox-
imal end of rostral carina marked by two
small pits just in front of protogastric lobes.
Epigastric prominences blunt, slightly pro-
tuberant. Protogastric lobes pronounced,
their uppermost part with small field of
lenslike scales. Orbits wide, deep, limited at
external angle by acute conical orbital spine;
short row of acute scales just posterior to
orbital spine on orbital margin. Anterolat-
eral angle of carapace produced as strong
acute conical spine extending from broad
flattened anterolateral lobe. Tip of spine
reaching at least middle of cornea. Extraor-
bital sinus a deep V-shaped notch. External
margin of anterolateral lobe with row of
small acute spines variable in number and
size. Free angle of first hepatic lobe pro-
duced as strong, acute, dorsally recurved
spine bearing from two to five scales at base.
Second and third hepatic lobes also well
marked and bearing spiniform scale at fron-
tal angle and row of minute scales along free
margin. Gastric area quite elevated at mid-
line, then sloping down both sides of cara-
pace, reaching broad concavity at dorsum
of hepatic lobes. Uppermost part of mid-
dorsal carina rugose and scaly throughout.
Cervical groove distinctly marked but not
deeply incised.

Anterior branchial area raised towards

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

midline but leveling into horizontal plane
towards free margin. Frontal angle pro-
duced as one or two coalescent, strong,
sharply acute conical spines. Anterior bran-
chial margin laterally expanded in row of
from seven to nine single or multiple tipped
prominent sharply acute denticles that
scarcely decrease in size from anterior to
posterior. Row of 9-11 tiny conical scales
mounted atop small tubercles along external
branchial suture.

Posterior branchial area with complicat-
ed topography; inner part raised and leveled
with cardiac area, then sloping towards lat-
eral margin, which rises again above mar-
ginal zone leaving behind wide longitudinal
trough. Margin markedly expanded in row
of 9-11 clear-cut conical, single or multiple
tipped, markedly recurved spinelike denti-
cles. Anteriormost and posteriormost den-
ticles smallest. At limit between anterior and
posterior branchial margin, first posterior
denticle overlaps last anterior denticle. Pos-
terior margin of carapace thick, more pro-
tuberant at middle section, which occasion-
ally forms broad based tubercle.

Abdominal terga deeply sculptured. Me-
dian section markedly protuberant espe-
cially at frontal edge, which forms moder-
ately sharp transverse carina. On pleural
area, near laterodorsal pit on second ab-
dominal segment, dorsal surface raised
frontally forming rotund broad-based
prominence bearing at summit from two to
five tiny scales. Anterolateral angle of sec-
ond abdominal epimeron produced as strong
spine tipped with sharp conical scale. It
stands at frontal end of narrow oblique ca-
rina extending over anterior two thirds of
epimeral dorsum. Crest of carina with row
of closely juxtaposed minute scales decreas-
ing in size posteriorly. Lateroventral angle
of epimeron acute; if rounded off, bearing
two or three acute scales. Lateral angle of
third and fourth epimera sharply acute. Tel-
son plate dimerous, subpentagonal.

Third thoracic sternum broad, its ventral
surface protuberant, rugose, bearing tiny

VOLUME 102, NUMBER 2

scales at frontal tip and sometimes one or
two scales along median line. Fourth ster-
num with long robust spinelike tubercle
markedly compressed into sagittal plane,
tipped with one major and two to three mi-
nor scales. Fifth sternum slightly swollen
but not forming tubercle. Frontal margin of
sixth sternum bearing one or more tiny scales
in transverse row.

Chelae subtriangular to ovoid, dorsoven-
trally depressed. Dorsal surface quite punc-
tate and covered by numerous small scales
atop minute tubercles. Scales on proximal
external surface lenticular and flat, then be-
coming more acute and prominent towards
finger tips. Two low, relatively narrow ridges
on dorsal surface of chelae, one starting from
upper articulating condylus at carpus-prop-
odus joint and ending at condylus of prop-
odus-dactylus joint. Uppermost part of this
ridge with irregular row of scales larger on
proximal end. Second ridge along median
line of chela extending from carpus-propo-
dus joint to proximal end of cutting edge of
fixed finger; it is marked by straight row of
acute scales. Finger cutting edges closely fit-
ted along distal two thirds but with small
gap at proximal third. Dorsal edge of dac-
tylus with sharply prominent conical tuber-
cle at proximal end, followed distally by row
of smaller acute tubercles or enlarged scales.
Palmar crest semicircular in outline, deeply
incised, formed by from two to three stout
single or multiple tipped acute spines arising
from thick bases. Smaller spines or spinules
attached to basis of body of main spines.
Frontal margin of propodus above propo-
dus-dactylus joint with acuminate tubercle;
spinules are found on both sides of it. Ven-
tral surface of chela with low but well de-
fined carina between lower condylus of car-
pus-propodus joint and tip of fixed finger.

Carpus strongly armed. Carpal crest with
two or three sharp, long, robust spines; one
or two small tubercles behind proximal
spine. Carpal lobe spiniform, as long as sec-
ond spine of carpal crest. Row of from five
to seven spiniform tubercles external to and

387

parallel to carpal crest. Second row of small-
er but acute tubercles external to above row.
Distodorsal margin of carpus fringed by
acute scales. Ventral surface with acute con-
ical spine as long as second spine in carpal
crest. Distoventral angle below carpus-pro-
podus condylus with small spiniform tu-
bercle.

Dorsal margin of merus with from five to
eight single or multiple tipped distinct spines
decreasing in size proximally. Spinules oc-
casionally present among bases of spines.
External face roughened by scales, mostly
on upper half. Distodorsal vertex with large,
somewhat flattened procurved spine longer
than the longest in row behind. Margin ex-
ternal to it with row of sharp conical spi-
nules. Inner ventral margin with from two
to five spiniform tubercles in row behind
single or double tipped spine at distal end.
External ventral margin scabrous or tuber-
culate. Distal end with spine as large as that
on inner ventral margin. One to three small
acute tubercles variable in size in front of
latter. Articular lobe at distalmost end of
same margin spiniform. Inner ventral mar-
gin of ischium with low broad nodule on
proximal end. Distal end with conical spine.
Between both ends from one to four acu-
minate tubercles variable in size. Distodor-
sal angle with short but acute spine slightly
displaced over outer surface.

Distoventral margin of ischium of pe-
reiopods fringed with minute scales; one of
these scales occasionally larger just behind
ischiomeral joint. Dorsal margin of merus
with row of small but clear-cut acute den-
ticles decreasing in size proximally; disto-
dorsal angle noticeably spiniform:; ventral
margin scabrous bearing minute scales in-
termingled with short stiff setae. Dorsal
margin of carpus also with small denticles;
distodorsal angle spiniform. Dorsal and
ventral margin of propodus with dense nar-
row band of scales and setae. Dorsal surface
of all pereiopodal articles except dactyl mi-
cronodulose and squamous.

Color (in life).—Dorsum of cephalotho-

388

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1.
male 25.7 mm CL from River Puquitre (IZUA C-686), b, d, f, h, j, 1, n, p. a, b, Dorsal view of carapace; c, d,
Lateral view of carapace; e, f, Left margin of carapace (ventral); g, h, Second right pereiopod; i, j, Inner view of
ischium and merus of left cheliped, carpus seen from above; k, 1, Ventral view of right cheliped; m, n, Chela
and carpus (dorsal); 0, p, Lateral view of left second abdominal epimeron.

rax and abdomen highly variable among in-
dividuals, ranging from orange-yellow to al-
most black. Though variation between both
extremes is continuous, an attempt was
made to quantify the frequency with which
different patterns appear in the population.
The following patterns were defined and
their frequency recorded (Table 1):

Pattern A: Dorsum of cephalothorax and
abdomen uniformly orange-yellow to ter-
racotta, occasionally scattered with small,
irregular, dark brownish spots.

Pattern B: Dorsum of cephalothorax and
abdomen variegate; extensive reticulate dark
green or dark brown design on a khaki or
light brown background.

Pattern C: Dorsum of cephalothorax and
abdomen uniformly dark greenish brown or
dark reddish brown.

Aegla denticulata lacustris (holotype), a, c, e, g, i, k, m, 0; A. denticulata denticulata Nicolet, 1849,

Pattern D: Dorsum of cephalothorax and
abdomen variegate; small to extensive dark
green patches on a lighter blue-green or
brown-green background.

Pattern E: Dorsum of cephalothorax and
abdomen variegate; small but numerous
yellowish or light greenish brown spots on
a uniform dark brown background; in some
specimens dorsal carina bluish tinged.

Pattern F: Dorsum of cephalothorax and
abdomen variegate; ample irregular blue-
black or brown-green-black patches on a
lighter blue-gray or gray-violet background.

In all specimens, the dorsal surface of the
pereiopods has the same color pattern as
the dorsum of the carapace but with darker
stripes across the carpus and merus. The
distal half of the chelae is always white or
yellowish-white, contrasting markedly with

VOLUME 102, NUMBER 2

the darker proximal half. The ventral sur-
face of the body and appendages is milky
white or slightly yellowish. The articular
membrane of the appendages is bright pur-
ple-red.

In formalin fixed specimens, the dorsum
of the carapace is wine-red in color, whereas
in alcohol preserved specimens the color
changes to light brown or tan.

Variation and measurements. —Making
allowances for minor variation in the size
and number of spines along the upper mar-
gin of the merus and dorsum of the carpus
of the chelipeds, adults in the type-series are
quite similar. Juveniles (P17—20) have many
features of the adult morphotype only
slightly developed (spines, tubercles, etc.)
but are still recognizable. The major differ-
ence between adults and juveniles is that in
the latter the tegumental scales are replaced
by short fine setae, most notably on the dor-
sum of the chelae and on the abdominal
terga.

Somatometric data of specimens in the
type-series are give in Table 2. Measure-
ments were made with calipers to the near-
est 0.1 mm; symbols for the morphometric
characters in the first column are given by
Jara & Lopez (1981).

Distribution. —Known only from the type
locality. Attempts to collect A. denticulata
lacustris in Lake Puyehue, 12 km to the north
of Lake Rupanco, failed. In July 1984, dur-
ing the winter high-water period of Lake
Rupanco, two small specimens (LC about
10 mm) were collected in a protected site at
the bank of River Rahue, 150 m from the
outlet of the lake. These specimens were
most probably washed out from the lake by
the effluent current.

Habitat and natural history.—Lake Ru-
panco is the southernmost of three major
preandean piedmont lakes included in the
River Bueno basin, and is the fifth in size
among Chilean lakes. It has a maximum
depth of 274 m and an area of 233.4 km”.
Limnologically, it is an oligotrophic, cold
temperate monomictic lake that circulates
at 9.6°C in midwinter and reaches 19°C at

389

Table 1.—Frequency of color patterns in 4. dentic-
ulata lacustris, from Lake Rupanco. Color patterns de-
fined in text. Sample size: 116 adult specimens col-
lected at 25 m depth on 30 Apr 1984, with baited traps.
Sex unrecorded.

Frequency
Pattern Absolute Relative (%)
A 5 4.31
B 11 9.48
ee 3 2.59
D 39 33.62
E 29 25.00
EF 29 25.00

epilimnion in summer. Dissolved oxygen is
high throughout the year (9.1 to 12.2 mg/
liter); conductivity is low (40.6 to 54.0 uS)
and calcium is scarce (2.1 to 4.8 mg/liter)
(Campos et al. 1985). Lake Rupanco lies at
118 m above sea level. Its eastern half is
surrounded by massive mountains and vol-
canoes (Puntiagudo and Casablanca) of the
Andes Cordillera. Its affluents are small to
medium-size rivers of pluvial or pluvio-gla-
cial regime. Its effluent (River Rahue) dis-
charges about 100 m?/sec on the average
(Campos et al. 1985). Lake Rupanco is of
glacial origin. The inferred chronology of
the regional glacial events suggests that Lake
Rupanco attained its present form about
11,000 years B.P. (Mercer 1972).
Underwater observations, made by the
author between 0 and 30 m depth at Huillin
Cove, showed that A. d. lacustris occupies
the sublittoral zone from 8 m depth down-
ward. Its vertical distribution appears closely
related to that of the soft substrate on which
crabs alternatively walk and collect particles
with their maxillipeds. Activity is markedly
higher at night, while during the day a great
proportion of the crabs rest buried in the
substrate. No data are available on the diet
of A. d. lacustris, but it can be presumed
that they feed on fine-grained detrital mat-
ter, though they were seen consuming a dead
fish lying on the bottom of the lake. Berried
females appear at the end of the summer
period (March), and the release of newborn
crabs from the brooding abdominal cham-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

390

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VOLUME 102, NUMBER 2

ber of the females occurs at the beginning
of the subsequent spring (October).

At Lake Rupanco, A. d. lacustris coexists
with Aegla abtao Schmitt and the crayfish
Samastacus spinifrons (Philippi). A. abtao
occupies the littoral zone and the upper sub-
littoral to 15 or 20 m depth. However, be-
low 4 m only large adult males are found.
The bulk of A. abtao’s population lives
among and under big stones and boulders
in the littoral. S. spinifrons is found in the
sublittoral at least down to 80 m depth, only
on soft substrate. Biological interactions
among these three species of decapod crus-
taceans are suspected but the only one ob-
served so far is predation by large A. abtao
males upon juveniles of A. d. /acustris. Oth-
er faunal components of the benthic com-
munity at Lake Rupanco are freshwater
mussels of the genus Diplodon and Pisidi-
um, aquatic snails of the genus Chilina, the
minute janirid isopod Heterias exul (Bow-
man et al. 1987), amphipods, acari, oligo-
chaetes, chironomid larvae and caenid
ephemeropteran nymphs.

Etymology. —The trivial name /acustris,
from Latin, alludes to the fact that this sub-
species of A. denticulata is apparently re-
stricted to a lentic environment.

Comparison. —To compare lacustrine and
riverine specimens of A. denticulata, four
crabs (two males and two females) from the
series from which the neotype of A. dentic-
ulata was drawn, National Museum of Nat-
ural History (USNM 169096 = USNM
80021) (Schmitt 1942) were examined. Ad-
ditionally, 30 males (LC 18 to 26 mm) col-
lected in rivers Puquitre and Remehue, 8
km north of Osorno city, and 50 males (LC
23 to 33 mm) from Rupanco were examined
and measured. One male from the Puquitre
(IZUA C-686) was employed for preparing
Fig. 1b, d, f, h, j, 1 and p.

A. denticulata lacustris is very similar to
the nominate subspecies A. denticulata den-
ticulata Nicolet, 1849, from which it de-
parts by the morphological features already
mentioned in the Diagnosis. Additionally,

Rupanco —s n=50
Puquitre ch n=30
Valdivia es n=29
1.30 1.40 1.50 1.60 CW/ PCW
Rupanco ch
Puquitre ch
Valdivia ch
1.30 1.40 1.50 1.60 CL/ PCL

Fig. 2. Somatometric comparison of males of A.
denticulata lacustris (Rupanco) and males of A. den-
ticulata denticulata (Puquitre and Valdivia). Sample
labeled Valdivia is a composite made of individuals
collected all over the Valdivia River Basin (Jara 1980).
Vertical line: mean value of somatometric proportion;
open rectangle: confidence interval for the mean (P =
0.05). Above: proportion between maximum carapace
width (CW) and maximum precervical width (PCW).
Below: proportion between carapace length (CL) and
precervical length (PCL).

A. d. lacustris differs from A. d. denticulata
in the following respects: ridge of dorsal ca-
rina sharper; dorsal profile of carapace less
inflated (cf. Fig. lc, d); external margin of
anterolateral lobe of carapace minutely den-
ticulate; denticles along ridge of merus of
chelipeds longer and sharper; tubercles of
middorsal row of carpus definitely spini-
form; spine of carpal lobe longer and more
slender (cf. Fig. 11, j); carpal ventral surface
less hairy; denticles of palmar crest longer
and slender; ventral angle of second abdom-
inal epimeron often produced into short
acute spine (cf. Fig. lo, p).

Because of the slender and clear-cut ap-
pearance of the main spines of its carapace,
together with the cleanness of the carapace
surface, A. d. lacustris is reminiscent of
another lacustrine species, A. rostrata Jara,
but differs from it in that A. rostrata lacks
the dorsal carina and the median tubercle
on the fourth thoracic sternum. On the other
hand, A. d. l/acustris lacks the distinctive
stylet-like rostrum of A. rostrata.

In Fig. 2 the average and confidence in-

392

tervals (P = 0.05) of two somatometric pro-
portions for riverine (Puquitre and Valdi-
via) and lacustrine (Rupanco) samples are
compared.

Remarks. — Though at first sight the Lake
Rupanco morphotype is clearly differentia-
ble from the riverine morphotype found
elsewhere in southern Chile, key characters
that differentiate the two forms appear elu-
sive. With the exception of the row of tiny
spinules along the external margin of the
anterolateral lobe of the carapace, the band
of tiny scales on the ridge of the dorsal ca-
rina, and some other minor details, every
morphological attribute in_A. d. lacustris has
its counterpart in A. d. denticulata. The dif-
ference is a matter of degree of expression
rather than of difference of attributes. The
Rupanco’s morphotype appears to be an
elaborate manifestation of the attributes
found in riverine specimens. The decision
to assign the Lake Rupanco population a
subspecific status 1s supported by the fact
that Lake Rupanco’s population is, on geo-
graphical and ecological grounds, an outlier
as compared to the remaining populations
of the species. In fact, Lake Rupanco is the
easternmost locality where A. denticulata has
been reported in the River Bueno basin, and
the only one in which the species inhabits
a lentic environment. The possibility that
Lake Rupanco’s population has already at-
tained reproductive isolation with respect
to the riverine populations, and therefore
qualifies as a different species, remains an
open question.

As mentioned elsewhere (Jara 1986), the
trend towards profuse spinulation in Aegla
seems to be correlated with living in lentic
environments. The prominent ornamenta-
tion seen in A. d. lacustris lends support to
this hypothesis. However, no explanation is
available for this overgrown spinulation,
neither from the ecological nor from the
genetical point of view. Profuse spinulation
in A. d. lacustris may well be an adaptive
response to 1ts immediate biotic or abiotic
environment. Otherwise, spinulation could

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

be a by-product of genetic changes related
to presently unknown factors. Comparison
of A. d. lacustris and A. d. denticulata life
histories could be illuminating in these re-
spects.

If the geographical and chronological re-
lationship between rivers and lakes in
southern Chile is considered (see comments
in Jara 1977, 1982, 1986), it appears evi-
dent that lacustrine populations of aeglids
descended from riverine stocks. These last
would have withstood the Quaternary gla-
cial events in non-glaciated areas, located
along the western border of the Chilean
Central Valley and in the Coastal Cordillera
(Illies 1960). That scenario implies that ae-
glids could have colonized the lakes rather
recently (in geological time). Mercer (1972)
estimates that lakes Llanquihue and Ru-
panco became free of ice about 11,000 years
B.P. Therefore, to colonize the Andean lakes,
aeglids had to advance from west to east,
along the rivers that drained the newly
formed lakes. Most probably, the forerun-
ners of A. d. lacustris were specimens of A.
d. denticulata, which could have entered the
young Lake Rupanco from rivers of the
Central Valley of Osorno Province. If spi-
nulation of A. d. lacustris is a derived con-
dition related to modifications of the former
riverine invader’s genome, one could expect
to find evidence of these changes by com-
paring population samples of both subspe-
cies by suitable techniques (protein electro-
phoresis, caryological analysis, etc.) (Wiley
1981).

Acknowledgments

The author thanks René Navarro and
Gonzalo Aguilar for their assistance during
the field work at Lake Rupanco; Dr. Ray-
mond E. Manning of the National Museum
of Natural History for the loan of four spec-
imens of A. denticulata; Mrs. Patricia Araya
and an anonymous reviewer for correcting
the MS; Dr. Thomas E. Bowman for making
available to the author his paper on Heterias

VOLUME 102, NUMBER 2

exul; and Direccion de Investigacion y De-
sarrollo, Universidad Austral de Chile, for
funding field work through grant RS-83-39.

Literature Cited

Bowman, T. E., R. Prins, & J. Arenas. 1987. The
occurrence of the freshwater isopod Heterias
(Fritzianira) exul in the Lakes Region of Chile,
with notes on the genus Heterias (Asellota: Jani-
ridae).— Hydrobiologia 146:275-281.

Campos, H., J. Arenas, W. Steffen, G. Agiiero, L. Vil-
lalobos, & G. Gonzalez. 1985. Investigacion
de la capacidad de cultivo de salmonidos de las
principales hoyas hidrograficas del pais. III An-
tecedentes limnologicos hoya Lago Rupanco. —
Corporacion de Fomento de la Produccion
(CORFO), Santiago, Chile, 404 pp.

Illies, H. 1960. Geologie der Gegend von Valdivia
(Chile). —Neues Jahrbuch fiir Geologie und Pa-
laeontologie 111:30-110. (Translated into
Spanish by J. Walper, in Publicacion Ocasional
de la Universidad Austral de Chile, Valdivia,
1970.)

Jara, C. 1977. Aegla rostrata n. sp., (Decapoda, Ae-

glidae), nuevo crustaceo dulceafuicola del Sur

de Chile.—Studies on Neotropical Fauna and

Environment 12:165-176.

1980. Taxonomia y distribucion del género

Aegla Leach (Crustacea, Decapoda, Anomura)

ete le

en el sistema hidrografico del rio Valdivia
(Chile).— Universidad Austral de Chile, Facul-
tad de Ciencias, Thesis, 126 pp.

1982. Aegla bahamondei, new species (Crus-
tacea: Decapoda: Anomura) from the coastal
mountain range of Nahuelbuta, Chile. — Journal
of Crustacean Biology 2(2):232-238.

1986. Aegla spectabilis, a new species of
freshwater crab from the eastern slope of the
Nahuelbuta Cordillera, Chile.— Proceedings of
the Biological Society of Washington 99(1):34—-
41.

,& M. T. Lopez. 1981. Anew species of fresh-
water crab (Crustacea: Anomura: Aeglidae) from
insular South Chile.— Proceedings of the Bio-
logical Society of Washington 94(1):88-93.
Mayr, E. 1969. Principles of systematic zoology.
McGraw-Hill Book Co., New York, 428 pp.
Mercer, J. H. 1972. Chilean glacial chronology 20,000
to 11,000 carbon-14 years ago: some global
comparisons.—Science 176:1118-—1120.
Schmitt, W. L. 1942. The species of Aegla, endemic
South American freshwater crustaceans.— Pro-
ceedings of the United States National Museum
91:431-520, pls. 25-28.
Wiley, E. O. 1981. Phylogenetics. The theory and
practice of phylogenetic systematics. John Wiley
& Sons, New York, 439 pp.

Instituto de Zoologia, Universidad Aus-
tral de Chile, Casilla 567, Valdivia, Chile.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 394-400

FRESHWATER CRABS ASSOCIATED WITH CAVES
IN SOUTHERN MEXICO AND BELIZE, WITH
DESCRIPTIONS OF THREE NEW SPECIES
(CRUSTACEA: DECAPODA)

Gilberto Rodriguez and Horton H. Hobbs, Jr.

Abstract. — Records of nine freshwater crabs frequenting subterranean hab-
itats in Belize and Mexico are reported. Three of them are described as new:
Potamocarcinus leptomelus, a troglophile from Veracruz; Odontothelphusa
monodontis, also a troglophilic species, from Tabasco; and Typhlopseudo-
thelphusa hyba, a troglobite, from Chiapas. The remaining crabs are troglo-
philes, one of which occurs in Belize, and the others, two of which because of
inadequate material cannot be fully identified, were found in caves in the states
of Chiapas, Guerrero, Oaxaca, and Tabasco, Mexico.

A study of several collections of cave-
dwelling crabs, kindly donated to the
Smithsonian Institution by James R. Red-
dell, has prompted us to record additions
to his list of Brachyura known to occur in
subterranean habitats in Mexico, Guate-
mala, and Belize (Reddell 1981). His list
includes three troglobites and six others that
are probable troglophiles. Subsequently, one
troglobitic form, Typhlopseudothelphusa
acanthochela Hobbs (1986), was described
from Blind Crab Cave, 12 km SW Millio-
nario, Cayo District, Belize, and a troglo-
philic one, Pseudothelphusa mexicana A\-
varez-Noguera (1987), from La Jolla [sic,
Joya?] Cave in the State of Guerrero, Mex-
ico. Records of the occurrence of eight ad-
ditional species, two of which cannot be ful-
ly identified because of inadequate material,
are reported herein from spelean waters of
Belize and Mexico.

All of the ““Material’’ cited herein is de-
posited in the National Museum of Natural
History, Smithsonian Institution (USNM).
Other abbreviations employed are: cl = car-
apace length, and cb = carapace width.

Pseudothelphusidae Ortmann, 1893

Potamocarcinus leptomelus, new species
Fig. la—h

Material. — Mexico: Cueva del Tuknel,
Mahoilca, Porvenir Zongolica, Veracruz, 31
Dec 1986, Steve and Lori Robertson, male
holotype, cl. 13.3 mm, cb. 21.1 mm (USNM
230080).

Description. —Cervical groove narrow,
almost straight, very shallow, reaching mar-
gin of carapace. Postfrontal lobes obsolete,
their position marked by slight depressions;
median groove absent except for notch on
upper margin of front. Surface of carapace
between postfrontal lobes and front sloping
gently ventromesially. Front low, of equal
height throughout, bilobed in dorsal view,
upper margin defined by series of irregularly
placed papillae; lower margin V-shaped in
median part, straight in lateral parts. Dorsal
margin of orbits sinuous. Lateral orbital an-
gle forming papillated tooth, followed by
deep notch; rest of anterolateral margin cov-
ered by small, irregularly placed, conical pa-
pillae. Surface of carapace covered by very

VOLUME 102, NUMBER 2

Fig. la-o.

4

(Gee jlet kag
nse

395

}
\

a-h, Potamocarcinus leptomelus, male holotype: a, Left gonopod, caudal view; b, Same, cephalic

view; c, Same, lateral view; d, Right cheliped; e, Distal part of chela; f Opening of efferent channel; g, Left third
maxilliped; h, Dorsal view of carapace. i-o, Pseudothelphusa mexicana, male, from Resumidor de la Joya: /,
Left gonopod, mesial view; j, Same, cephalic view; k, Same, subapical view; /, Same, lateral view; m, Same,
caudal view; n, Opening of efferent channel; 0, Left third maxilliped.

small papillae, invisible to naked eye, par-
ticularly in branchial region.

Eyes small, with distinct faceted cornea
and pigment, of usual shape, not filling or-
bit. Third maxilliped with conspicuous
notch on external margin of merus, near
insertion of palp; external margin of is-
chium convex; exognath 0.59 times length
of ischium. As name suggests, all pereio-

pods conspicuously slender. Chelae only
slightly dissimilar in size and shape; palm
of larger one very elongate with straight up-
per and lower margins; fingers not gaping;
carpus with strong internal spine preceded
by four spinules; merus elongate with row
of spinules on internodorsal and interno-
ventral ridges and smaller spinules on outer
ridge. Walking legs very long, length of third

396

pair 1.6 times breadth of carapace, and me-
rus 6.3 times as long as broad; relations be-
tween podomeres as follows: merus 1, car-
pus 0.33, propodus 0.54, dactylus 0.54.
Dactylus with seven spines in upper row
and four in two lower ones.

Gonopod subcylindrical, straight; apical
part bearing triangular marginal process
overreaching apex, spiniform mesial pro-
cess directed distomesially, and bifid ce-
phalic process with apices pointing mesial-
ly. Field of spines very narrow, with few
setae placed on distal surface of apex.

Remarks.—The gonopod of this species
resembles that of Potamocarcinus magnus
(Rathbun 1896:377); the mesial process is
similar but directed distomesially instead of
mesially; the bifid process is present in both,
although distinctly reduced in P. leptome-
lus; and the marginal process of the latter
is rudimentary rather than well developed
and cup-shaped as in P. magnus.

Potamocarcinus aspoekorum (Pretzmann)

Pseudothelphusa (Zilchia) aspoekorum
Pretzmann, 1968a:12.

Potamocarcinus (Zilchia) mexicanus Pretz-
mann, 1968a:12 [nomen nudum].

Potamocarcinus (Zilchia) aspoekorum. —
Pretzmann, 1971:20; 1972:75, figs. 405-
407, 417-420.

Potamocarcinus aspoekorum. — Rodriguez,
1982:115, figs. 69d; 70f, m; 71le, 1; 74a—c.

Material. —Belize: Balam’s Cave (Uchen
Balam), Cayo District, 22—26 Mar 1979, L.
McNatt, 1 male cl. 18.6 mm, cb. 30.1 mm
(USNM 230082).

Remarks. — Troglobitic characters are not
conspicuously developed in this species. The
walking legs are moderately slender (third
right pereiopod of largest male with total
length 34.0 mm, merus 3.3 times as long as
broad), but the dorsal surface of the cara-
pace is not pale, and the eyes are normally
developed.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Pseudothelphusa mexicana
Alvarez-Noguera, 1987
Fig. li-o

Pseudothelphusa mexicana Alvarez-No-
guera,1987:1.

Material.—Mexico: Resumidero de la
Joya, San Gregorio, Guerrero, 23 Feb 1981,
Steve Robertson, 2 males, cl. 18.6 and 16.5
mm, cb. 28.7 and 25.8 mm; 1 female, cl.
18.0 mm, cb. 28.2 mm (USNM 230084).

Remarks.—Our specimens are from the
type locality. Troglobitic characters are not
obvious in this crab, although the walking
legs are moderately slender (third right pe-
reiopod of largest male with total length 29.5
mm, merus width/length = 0.35). The dor-
sal surface of the carapace, however, is dark
and the eyes are well developed.

Odontothelphusa monodontis, new species
Fig. 2a—f

Material. —Mexico: Grutas del Cocona,
Tabasco, 26 Aug 1972, R. W. Mitchell and
W. H. Russell, 1 male holotype, cl. 17.0
mm, cb. 25.2 mm (USNM 230078).

Description. —Cervical groove very shal-
low and wide, not reaching margin of car-
apace. Postfrontal lobes wide, flat, clearly
delimited anteriorly, surface covered by flat
papillae not visible to naked eye; median
groove well marked between lobes, but not
anterior to them. Surface of carapace be-
tween frontal lobes and front flat. Upper
border of front well marked by overhanging
papillae and conspicuous notch in middle;
lower margin slightly sinuous and lying
slightly anterior to upper one; front high and
excavate. Anterior margin of carapace with
notch behind orbit, about 5 small papillae
situated between notch and cervical groove,
and approximately 15 papilliform teeth on
rest of lateral margin. Third maxilliped con-
spicuously wide; merus 1.43 times as wide
as long; exognath 0.93 length of ischium of
endognath.

VOLUME 102, NUMBER 2

30

Fig. 2a-k.

a-f, Odontothelphusa monodontis, male holotype: a, Right gonopod, caudal view; b, Same, mesial

view; c, Same, cephalic view; d, Same, subapical view; e, Left third maxilliped; {, Opening of efferent channel.
g—k, Typhlopseudothelphusa hyba, male holotype: g, Opening of efferent channel; 4, Left third maxilliped; 7, Left
gonopod, caudal view; j, Same, lateral view; k, Same, cephalic view.

Gonopod slender with apical part bent
laterally at angle of 45 degrees; apex con-
sisting of flat quadrangular lobe with long
spur on proximomesial angle and papilli-
form tubercle and small finger-like tubercle
on cephalic surface; field of spines narrow,
slitlike.

Remarks. — This crab differs from its con-
gener in possessing two tubercles on the ce-
phalic surface of the gonopod and lacking a
spiniform process on the distal angle of the
mesial border. Some of the characteristics
that have been associated with an obligate
spelean existence are lacking in this species,

but the dorsal surface of the carapace is pale,
and the walking legs moderately slender
(third right pereiopod with total length of
30.3 mm, merus 2.87 times as long as broad);
the eyes, however, are well pigmented.

?Raddus sp.

Material. —Mexico: Cueva El Chorrea-
dero, Chiapas, 24 Dec 1973, R. Syme, |
juvenile male, cl. 9.9 mm, cb. 13.8 mm
(USNM 230090).

The only specimen available is a distinct-
ly juvenile male; however, the gonopod ex-

398

hibits the flat quadrangular form character-
istic of Raddus, and the features of the
carapace are those of other members of the
genus. This cave is located 12 km E Tuxtla
Gutiérrez (Reddell 1981:292), near the range
of Raddus bocourti (A. Milne-Edwards,
1866).

Typhlopseudothelphusa hyba, new species
Fig. 2¢-k

Potamocarcinus (Typhlopseudothelphusa)
mocinoi. — Hobbs et al. 1977:145 [in part :
localities 2 and 3].

Material. —Mexico: Cueva de Los Lla-
nos, 15 km ESE San Cristobal de Las Casas,
Chiapas, 29 Aug 1972, J. Cooke, W. Rus-
sell, and R. Mitchell, male holotype, cl. 12.4
mm, cb. 18.1 mm (USNM 150633), 1 fe-
male paratype, cl. 15.4 mm, cb. 22.8 mm
(USNM 230081).—same locality, 5 Feb
1972, D. McKenzie and J. R. Reddell, 1
female, cl. 1.2 mm, cb. 24.6 mm (USNM
143613).—Cueva de los Murciélagos, 15 km
ESE of San Cristobal de Las Casas, Chiapas,
29 Aug 1972, J. Cooke and R. Mitchell, 1
male, cl. 12.6 mm, cb. 19.9 mm, 2 females,
cl. 14.7 and 11.6 mm, cb. 20.9 and 16.8
mm (USNM 150629).

Description. —Carapace narrow (cb/cl =
1.46), strongly convex in anterior part. Cer-
vical grooves forming paired, shallow
depressions in holotype but absent in topo-
typic female. Postfrontal lobes and median
groove obsolete, but carapace with slight
depression immediately posterior to front;
latter, lacking upper margin, rounded and
projecting forward, its lower margin bilobed
in dorsal view. Lateral orbital angle forming
well developed triangular tooth, followed by
shallow notch. Margins of orbits and front
with row of small granules. Lateral margin
of carapace with larger, irregularly placed
granules. Eyes reduced, lacking distinct fac-
eted cornea and pigment. Third maxilliped
with noticeable impression on external
margin of merus, near insertion of palp; ex-
ognath 0.61 times length of ischium. All

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

pereiopods extremely slender. Chelipeds
subequal in size and shape; palm cylindri-
cal; fingers also very slender, approximately
twice as long as palm, and armed with mi-
nute teeth; carpus with small, hooked spine
on internal margin; merus very elongate,
overreaching carapace by halfits length, and
bearing rows of granules on its three ridges.
Third pair of walking legs 1.6 times width
of carapace; merus 5.87 times as long as
broad, and relations of its podomeres as fol-
lows: merus 1, carpus 0.35, propodus 0.66,
dactyl 0.74; dactylus with six or seven spines
in five rows.

Gonopod wide in lateromesial plane, nar-
row in cephalocaudal plane; distal part
strongly bent; gonopore and field of spines
directed cephalad; distal margin rounded
and armed with prominent conical spines;
marginal process cup-shaped and directed
cephalad; and strong, triangular mesial pro-
cess disposed in same plane as field of spines.

Remarks. — Typhlopseudothelphusa hyba,
like its congeners, exhibits a combination
of advanced characters associated with its
adaptations to a spelean environment. It
may be distinguished from them by the tri-
angular mesial process of the gonopod with
a single apex.

Pseudothelphusidae indet. species

Material. — Mexico: Sumidero de Citla-
lapa, Zongolica, Veracruz, 23 Mar 1981,
Steve Robertson and Phillipe Ackamann, 1
fragmented female, cl. approx. 19.5 mm, cb.
approx. 33.5 mm (USNM 240112).

Description. —Cervical groove very shal-
low and wide, not reaching margin of car-
apace. Postfrontal lobes rounded, flat, not
well delimited anteriorly; median groove
well marked between lobes and narrowly
linear anterior to them. Surface of carapace
between frontal lobes and over front flat;
upper border of latter well marked by over-
hanging papillae and bearing conspicuous
median notch; lower margin slightly sin-
uous and lying slightly anterior to upper.

VOLUME 102, NUMBER 2

Front high and excavate. Anterior margin
of carapace with notch behind orbit, about
5 small papillae between notch and level of
cervical groove, and approximately 15 pa-
pilliform teeth on rest of lateral margin. Eyes
normally developed, but unpigmented.

Trichodactylidae

Avotrichodactylus constrictus
(Pearse, 1911)

Trichodactylus constrictus Pearse, 1911:111,
fig. 4.—Rodriguez & Manrique, 37:183.
Trichodactylus (Trichodactylus) constric-

tus. —Coifmann, 1939:111.
Trichodactylus (Avotrichodactylus) constric-
tus. —Pretzmann, 1968b:71.
Trichodactylus (Rodriguezia) constrictus. —
Bott, 1969:26.—Cottarelli & Argano,
19772210.

Material. —Mexico: | km N of Palenque,
Chiapas, 25 Jul 1973, J. Reddell, N. Ka-
wakatsu, D. Denson, and S. R. Mitchell, 2
males, cl. 19.8 and 12.5 mm, cb. 17.9 and
12.9 mm (USNM 230085).— Cueva de Juan
Sanchez, NW Acatlan, Oaxaca, 1 Jul 1976,
T. Byrd, A. G. Grubbs, and M. Cossey, 1
female, cl. 16.5 mm, cb. 15.8 mm (USNM
230086).

Remarks.—The female specimen from
Cueva de Juan Sanchez is assigned to this
species with doubt. It has five lateral teeth
(not including the external orbital angle) on
the right side and four on the left, whereas
A. constrictus usually has two or three teeth.

Avotrichodactylus bidens (Bott, 1969)

Trichodactylus (Rodriguezia) bidens Bott,
1969:25, pl. 24: figs. 68, 69.—Cottarelli
& Argano, 1977:207, figs. 3, 4.

Material.— Mexico: Cuevita de la Cas-
cada Azufre, 3 km N Tapiulapa, Tabasco,
15 Jun 1975, A. G. Grubbs, 1 juvenile male
(USNM 230083).

Remarks.—Bott (1969) designated the
type locality ““Arroyo del Solpho’’ near

399

Tapiyulapa, which is almost certainly the
same as, or perhaps an extension of, that
from which our specimen was taken. This
juvenile is very small, and the lateral teeth
on the carapace are not well formed, but
even at this size, the gonopod is clearly typ-
ical of A. bidens.

Acknowledgments

Thanks are extended to James R. Reddell
for donating the material cited herein to the
Smithsonian Institution, thereby making it
available to us. We are also grateful to Brian
F. Kensley and Raymond B. Manning for
their criticisms of the manuscript.

Literature Cited

Alvarez-Noguera, Fernando. 1987. Pseudothelphusa
mexicana, a new freshwater crab from the state
of Guerrero, Mexico (Brachyura: Pseudothel-
phusidae).— Proceedings of the Biological So-
ciety of Washington 100(1):1-3.

Bott, Richard. 1969. Die Siisswasserkraben Siid-
Amerikas und ihre Stammesgeschichte. — Ab-
handlungen der Senckenbergischen Natur-
forschenden Gesellschaft (Frankfurt am Main)
518:1-94.

Coifmann, Isabel. 1939. Potamonidi della Guiana
Inglesi.—Archivo Zoologico Italiano 27:93-1 16.

Cottarelli, Vezio, & Roberto Argano. 1977. Tricho-
dactylus (Rodriguezia) mensabaki n. sp. (Crus-
tacea, Decapoda, Brachyura) granchio cieco delle
acque sotterranee del Chiapas (Messico). —
Quaderni Problemi Attuali di Scienza e di Cul-
tura, Academia Nazionale dei Lincei 171(3):207-
Die

Hobbs, Horton H., Jr. 1986. A new troglobitic crab
(Crustacea: Decapoda: Pseudothelphusidae)
from Belize.—Texas Memorial Museum, Spe-
leological Monographs 1:1-4.

—., H. H. Hobbs II], & Margaret A. Daniel. 1977.
A review of the troglobitic decapod crustaceans
of the Americas.—Smithsonian Contributions
to Zoology 244:v + 183 pages.

Milne-Edwards, A. 1866. Description de trois nou-
velles espéces du genre Bosica, Crustacés Brach-
yures de la tribu des Telphusiens.— Annales de
la Société Entomologique de France (4)6:203-
205.

Pearse, A.S. 1911. Report on the Crustacea collected
by the University of Michigan-Walker Expedi-
tion in the State of Vera Cruz, Mexico. —Thir-

400

teenth Annual Report of the Michigan Academy
of Science 13:108-113.
Pretzmann, Gerhard. 1968a. Neue sudamerikanische
Susswasserkraben (Vorlaufige Mitteilung).—
Entomologisches Nachrichtenblatt (Wien) 15:1-
20.
. 1968b. Die Familie Trichodactylidae (Milne
Edwards 1835) Smth [sic] (Vorlaufige Mitteil-
ung). —Entomologishce Nachrichtenblatt (Wien)
15:70-76.
1971. Fortschritte in der Klassifizierung der
Pseudothelphusidae. —Sitzungsberichten der
Osterreichische Akademie der Wissenschaften,
Mathematisch-Naturwissenschaftliche Klasse
(1)179(1-4):14-24.
1972. Die Pseudothelphusidae (Crustacea,
Brachyura).— Zoologica 42(120), part 1:1-182.
Rathbun, Mary J. 1896. Descriptions of two new
species of fresh-water crabs from Costa Rica.—
Proceedings of the United States National Mu-
seum 18:(377-—389).

Reddell, James R. 1981. A review of the cavernicole

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

fauna of Mexico, Guatemala, and Belize. — Bul-

letin of the Texas Memorial Museum 27:1-327.
Rodriguez, Gilberto. 1982. Les crabes d’eau douce
d’Amérique. Famille des Pseudothelphusi-
dae.—Office de la Recherche Scientifique et
Technique Outre-Mer, Faune Tropicale 22:1-
223.
, & Fernando Manrique. 1967. El género 7ri-
chodactylus en México (Brachyura, Potamoni-
dae).—Anales del Instituto de Biologia, Uni-
versidad Nacional Autonoma de México 37:183-
186.

(GR) Instituto Venezolano de Investiga-
ciones Cientificas, Apartado 21827, Cara-
cas 1010A, Venezuela; (HHH) Department
of Invertebrate Zoology, National Museum
of Natural History, Smithsonian Institu-
tion, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 401-404

A NEW DEEP-SEA CRAB OF THE GENUS
CHACEON FROM CHILE
(CRUSTACEA, DECAPODA, GERYONIDAE)

Luis A. Chirino-Galvez and Raymond B. Manning

Abstract.—Chaceon chilensis, a large species with five anterolateral teeth on
the carapace and narrow, compressed dactyli on the walking legs, is described
from localities off Chile, the Juan Fernandez Islands, and Isla San Félix. It is

_ the only Recent East Pacific species of the genus.

Until recently, carcinologists working with
deep-sea crabs of the genus Chaceon from
localities around the world often have iden-
tified their material with the West Atlantic
C. quinquedens (Smith, 1879), or the East
Atlantic C. affinis (A. Milne Edwards &
Bouvier, 1894); both of these species for-
merly had been assigned to the genus Ger-
yon (see Manning & Holthuis 1989). A
species of Chaceon from off Chile has been
identified erroneously in the past with both
of these species. We describe the Chilean
species as new herein.

The holotype and several paratypes are
in the Museo Nacional de Historia Natural,
Santiago (MNHN). Other paratypes are in
the collections of the Museo Instituto de
Zoologia, Universidad Austral de Chile,
Valdivia (MIZUA), the National Museum
of Natural History, Smithsonian Institu-
tion, Washington (USNM), the Instituto de
Oceanologia, Universidad Catolica de Val-
paraiso, Valparaiso (IOUV), the Museo de
Zoologia, Universidad de Concepcion
(MZUC), and the Museo de Historia Nat-
ural, Valparaiso (MHNV).

We use cl for carapace length, measured
on the midline, and cb for carapace breadth,
measured at and including the fifth antero-
lateral spines, m for meter(s), and mm for
millimeters.

Chaceon chilensis, new species
Figs. 1-2

Geryon affinis. —Dupré, 1975:34.—Baez &
Andrade, 1977:215.—Andrade & Baez,

1980:263, 264.—Chirino-Galvez, 1985:
R126.—Baez & Ruiz, 1985:103. [Not
Geryon affinis A. Milne Edwards & Bou-
vier, 1894.]

Geryon quinquedens. —Retamal, 1977:249,
250, fig. 1; 1981:33, fig. 179. [Not Geryon
quinquedens Smith, 1879.]

Previous records. —Chile: Zapallar,
33°34'S, 71°38'W, 380-450 m (Baez & An-
drade 1977).—Central Chile, taken in fish-
ery for Heterocarpus reedi Bahamonde (An-
drade & Baez 1980).

Juan Fernandez Islands: Juan Fernandez
Islands [33°00’S, 80°00’W] (Retamal
1977, Baez & Ruiz 1985).—Isla Alejandro
Selkirk [33°45’S, 80°46’ W] (Chirino-Galvez
1985).—East of Isla Robinson Crusoe and
near Isla Santa Clara [33°42’S, 79°00’W]
(Dupré 1975).

Isla San Félix (Baez & Ruiz 1985).

Material. —Chile: Off Chile, 33°35'S,
/TA2Z2W, 370m: 3 males (in 3° lots,
MNHN D-10824, D-10825, D-10875).—
Isla San Félix, 26°16’S, 80°00’W, 27 May
1969, in lobster traps: one male (cl 126.8
mm, cb 145.5 mm, holotype, MNHN
D-10821).—Off Quintero [32°47’'S, 71°32’W],
400 m: one male IOUV C-005).— Off Zapa-
lar [32°33’S, 71°29’W], 400 m: one female
(LOUV C-003).— Off Zapallar, 360-400 m:
one male (MHNYV).—Islas Juan Fernandez:
Isla Robinson Crusoe [33°38’S, 78°52'W]:
three males IOUV C-001, C-002, C-004). —
Isla Robinson Crusoe, 270-300 m: one
specimen (MZUC).—Off Caleta Sanchez,
Isla Alejandro Selkirk, 150 m, in lobster

402

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Chaceon chilensis, new species. Male holotype, cl 127 mm, Isla San Félix: a, dorsal view: b, ventral
view. Male paratype, cl 120.5 mm, off Chile: c, front. Male paratype, cl 115 mm, off Caleta Sanchez: d, dorsal
view; é, anterior part of carapace; f, cheliped.

traps, 1985, associated with Jasus frontalis
(H. Milne Edwards) and Paramola rathbuni
Porter: one male, one female (USNM
205697), one male (MIZUA C-688).

All specimens other than the holotype are
paratypes.

Description. —A large Chaceon, cl to 152
mm, cb to 175 mm in adults, with five an-

terolateral teeth on the carapace and later-
ally compressed dactyli on the walking legs.
Carapace broader than long, breadth 1.0-
1.2 times length, inflated anteriorly, espe-
cially at protogastric regions. Median pair
of frontal teeth large, separated by broad,
V-shaped sinus, overreaching similar lateral
frontal teeth. Distance between submedian

VOLUME 102, NUMBER 2

frontal teeth less than distance between them
and lateral frontal teeth. Second and fourth
anterolateral teeth more obtuse and lower
than first, third, and fifth teeth. Distance
between first and third teeth subequal to
distance from third to fifth, distance be-
tween first and second teeth much less than
from second to third. Suborbital tooth vis-
ible in dorsal view, not extending to apex
of lateral frontal tooth. Carapace with curved
ridge extending mesially from each fifth lat-
eral tooth, surface, except for gastric, and
branchial regions, especially granular. Che-
liped rough dorsally, with blunt subdistal
tooth dorsally on merus, carpus lacking out-
er spine but with well-developed triangular
inner spine, propodus lacking distal dorsal
spine. Meri of walking legs lacking distal
dorsal spine. Fifth leg: merus broad, length
five times height, about two-thirds cb (0.61-
0.65); propodus broad, length about four
times height; dactylus about four-fifths as
long as propodus. Dactyli of walking legs
compressed, narrow, height at midlength
much greater than width.

Size. —Males, cl 107 to 152 mm, cb 124
to 174.5 mm; females, cl 102 to 124.5 mm,
cb 112 to 151 mm. Baez & Ruiz (1985)
recorded males with cl ranging from 105.8
to 124 mm, cb from 120.8 to 144.3 mm.
Baez & Andrade (1977) reported a specimen
with cl 152.1 mm, cb 174.5 mm, and Re-
tamal (1977) studied a specimen measuring
cl 120 mm, cb 140 mm.

Remarks.—This new species resembles
C. fenneri (Manning & Holthuis, 1984) in
general appearance, but differs in numerous
features. It is a longer-legged species, with
the merus of the fifth leg greater than 0.6 of
the carapace breadth (less than 0.6 in C.
fenneri), and, although the legs are longer,
the dactylus is shorter. The dorsal ridge on
the carpus of the walking legs is much
smoother. The carapace is more inflated
dorsally and more rounded posterolaterally,
and is much more granular. The median
frontal teeth extend further forward. The
suborbital margin is shaped differently: it is

403

Fig. 2. Chaceon chilensis, new species. Male para-
type, cl 115 mm, off Caleta Sanchez. a, orbit, ventral
view; b, propodus of fifth leg, posterior view; c, same,
dorsal view.

evenly curved in C. chilensis, flattened for
much of its length in C. fenneri.

One of the specimens (MNHN D-10875)
has but three frontal teeth; it is shown in
Fig. bf:

N. Bahamonde N., in a letter of 8 August
1973 to John S. Garth, commented that the
fishermen called this species “‘jaiba blanca,”
because of its light color in life.

Retamal (1981) called this species “‘can-
grejo de profundidad.”’ There are records in
the literature from the following depths:
270-300 m (Dupré 1975, Retamal 1977);
380-450 m (Baez & Andrade 1977); 200-
2000 m, in fishery for Heterocarpus reedi
Bahamonde (Andrade & Baez 1980).

Etymology.—The specific name reflects
the known geographic range of the species,
from localities off central Chile.

Distribution. —Known only from locali-
ties off central Chile, including the Juan Fer-
nandez Islands and Isla San Félix.

404

Acknowledgments

We thank our colleagues who facilitated
our study of material of this species in the
collections under their care. Roy K. Kropp
photographed the specimens from USNM
205697. Lilly King Manning prepared all of
the figures for publication.

Literature Cited

Andrade V., H., & P. Baez R. 1980. Crustaceos de-
capodos asociados a la pesqueria de Heterocar-
pus reedi Bahamonde 1955 en la zona central
de Chile.—Boletin de Museo Nacional de His-
toria Natural (Chile) 37:261-267.

Baez R., P., & H. Andrade V. 1977. Geryon affinis

Milne Edwards y Bouvier 1894 frente a las cos-

tas de Chile (Crustacea, Decapoda, Brachyura,

Geryonidae).—Boletin de Museo de Historia

Natural, Valparaiso 10:215-—219.

, & R. Ruiz L. 1985. Crustaceos de las islas

oceanicas de Chile depositados en el Museo Na-

cional de Historia Natural de Santiago. Pp. 93-

108 in P. Arana, ed., Investigaciones Marinas

en el Archipiélago de Juan Fernandez. Univer-

sidad Catolica de Valparaiso.

Chirino-Galvez, L. 1985. Stenotermia y biogeografia
de gerionidos (Crustacea, Decapoda, Brachy-
ura).—Archivos de Biologia y Medicina Exper-
imentales 18(2):R126 (abstract).

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Dupré M., E. 1975. Lista de Crustaceos Decapodos
citados para el Archipiélago de Juan Fernan-
dez.—CIMAR Documentos Técnicos 8/75:1—46.
Centro de Investigaciones del Mar, Universidad
Catolica de Valparaiso.

Manning, R. B., & L. B. Holthuis. 1984. Geryon fen-
neri, anew deep-water crab from Florida (Crus-
tacea: Decapoda: Geryonidae).— Proceedings of
the Biological Society of Washington 97(3):666—
673.

——., & 1989. Two new genera and nine
new species of geryonid crabs (Crustacea, De-
capoda, Geryonidae).— Proceedings of the Bi-
ological Society of Washington 102(1):50-77.

Retamal R., M. A. 1977. Sobre Geryon quinquedens

Smith, 1870, en el Archipiélago de Juan Fer-

nandez.—Boletin de Sociedad de Biologia de

Concepcion 51(1):249-251.

1981. Catalogo ilustrado de los crustaceos

decapodos de Chile. —Gayana (Zoologia) 44:110

pp.

(LAC-G) Museo Sociedad Fonck, 4 Norte,
784 Vina del Mar, Chile (Present address)
Department of Geology, Kent State Uni-
versity, Kent, Ohio 44242; (RBM) Depart-
ment of Invertebrate Zoology, National
Museum of Natural History, Smithsonian
Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 405-410

UPOGEBIA CORALLIFORA, A NEW SPECIES OF
CORAL-BORING SHRIMP FROM THE WEST INDIES
(DECAPODA: UPOGEBIIDAE)

Austin B. Williams and P. J. B. Scott

Abstract. — Upogebia corallifora, a new species of shrimp from the West In-
dies, lives in boreholes that it excavates in CaCO, substrates, dead coral and
Pleistocene limestone. The species, here described and illustrated, is similar in
morphology to U. noronhensis from Arquipélago de Fernando de Noronha off
northeastern Brazil. The chelae of the new species differ in having a longer
fixed finger and lacking a row of spines on the lower mesial surface of the palm.

Recent ecological studies on thalassinid
shrimps, Upogebia, in the West Indies have
shown that an undescribed species of this
genus constructs boreholes in “‘skeletons of
dead coral or the dead bases of living corals”’
(Scott, Reiswig & Marcotte 1988), and that
juveniles, here provisionally included with
it, bore into limestone (Scott, Moser & Risk
1988). We describe the new species in this

paper.

Upogebia corallifora, new species
Fig. 1

Material examined.—Jamaica: USNM
230075. Holotype 2; Port Royal, Drunken-
mans Cay, 3—4 m, area of dead Siderastrea
siderea (Ellis & Solander 1786), P. J. B. Scott,
Oct 1986.—USNM 230076. Paratypes, 2 6,
22(1 ovig.); Port Royal, Drunkenmans Cay,
approx. 2—4 m, P. J. B. Scott, 10 Oct 1986.
One ¢ and one 2 from this lot of specimens
are to be deposited in the National Museum
of Canada.

Upogebia corallifora new species ? USNM
230077. 2 2 Guv.), 2 juvs.; Discovery Bay,
boring in Pleistocene CaCO, of an artificial
reef, about 5 m depth, P. J. B. Scott, Oct
1986 (specimens softened in Aerosol OT
solution after being dried).

Diagnosis. — Projections to either side of
rostrum ending in acute spine. Postocular
spine present. First and second abdominal

segment lacking ventral spines; sixth seg-
ment lacking hooked anterolateral spine.
Telson subrectangular. Carpus of cheliped
with one small spine on mesiodistal margin.
Leg 2 with proximal mesioventral spine on
merus; merus of leg 4 spineless.

Description. —Rostrum triangular, short,
slightly downturned; tip exceeding eye-
stalks; dorsal pair of strong subapical spines
followed on each side by two spines suc-
cessively shorter in length, and separated
from first by wide interval; ornamented dor-
sal two-thirds of carapace surface anterior
to cervical groove bearing pilose tufts,
changing from spiny anteriorly to tubercu-
late posteriorly, and angling toward sides
posteriorly; gastric surface posterior to this
smooth; ornamented anterior part separat-
ed from and flanked on each side by pos-
teriorly divergent ridge bearing crest of about
12 spines, strongest on lateral rostral pro-
cess and decreasing to obsolescence poste-
riorly. Cervical groove deep and continu-
ous, shoulder lateral to it bearing one
obsolescent tubercle below intersection with
thalassinidean line; latter continuing to pos-
terior margin of carapace. Postocular spine
present.

Abdomen broadly and smoothly arched
dorsally; pleura of segment 1 narrowly
rounded posterolaterally, those of 2—5
broadly rounded, with dense fine plumose

406

TV
A~

uC a A. 5
alee

Fig. A.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Upogebia corallifora, 2 holotype: a, Cephalic region, lateral; b, Anterior carapace, dorsal; c, Right

cheliped, lateral; d, Same, mesial; e, Leg 2; h, Leg 5; i, Parts of abdominal segment 6, telson and uropods.
Paratypes: dissociated legs 3-4, f, g. Scales: 1, a—h; 2, 1; each 1 mm.

setae on posterolateral corner of terga 3-4,
and in tracts on pleura of segments 3-5;
segment 6 subrectangular, wider than long,
lateral margin sinuous, adapted for inter-
locking with extended uropodal exopod,
notch anterior to posterolateral lobe contin-
uous with oblique groove and lunate dorsal
impression.

Telson subrectangular, a little wider than

long, broadest proximally, angle on antero-
lateral margin interlocking with groove on
central rib of uropodal endopod; posterior
margin shallowly biarcuate and densely
fringed with setae; transverse proximal ridge
fairly prominent and continuous with low
lateral ridges at each side; median groove
obsolescent.

Eyestalk stout, deepest at about mid-

VOLUME 102, NUMBER 2

length, convex ventrally; eye narrower than
diameter of stalk and directed anterolater-
ally.

Antennular peduncle reaching to about
proximal one-fourth of terminal article of
antennal peduncle, its proximal two articles
together slightly longer than terminal arti-
cle; flagella unequal, lower thinner ramus
somewhat longer than thicker upper one.

Antennal peduncle with about one-half
its length extending beyond tip of rostrum;
article 2 bearing tiny subdistal ventral spine;
scale moderate, oval, sometimes with mi-
nute terminal spine.

Mouthparts as figured for U. jonesi (Wil-
liams 1986); maxilliped 3 with epipod.

Epistomial projection rather broad in lat-
eral view, bearing small apical spine.

Chelipeds essentially equal. Ventral mar-
gin of ischium bearing one tiny spine. Merus
with single subdistal dorsal spine reaching
level of postocular spine, row of five small
to moderate spines on ventral margin. Car-
pus trigonal, shallow longitudinal groove
jaterally and slender spine at anterior ven-
trolateral corner; mesiodorsal crest of al-
most uniform small spines remote behind
prominent spine on anterior margin and
partly obscured by setae in proximal part
of row, all directed distad, and three very
small spines on anterodorsal margin mesial
to articulation of carpus; one small spine
near middle of distomesial margin, and an
obsolescent spine dorsal to it. Palm oval in
cross section, length including fixed finger
about three times maximal height; bearing
mixed prominent plumose and long setae
in one sparse dorsal row, plus lateral oblique
row of setae ending anteriorly in patch of
long setae near base of fixed finger; mesio-
dorsal row of small somewhat irregular
spines ending distally in small acute spine
above condyle of dactyl; lower mesial sur-
face spineless but bearing low transversely
arcuate ridge proximally. Fixed finger
slightly shorter than dactyl and more slen-
der, slightly downcurved in middle and ta-

407

pering to slender tip, four small teeth on
proximal prehensile edge. Dactyl much
stouter than fixed finger, its curved upper
surface bearing about three small, proximal
tubercles; tip preceded on prehensile edge
by more or less straight edge growing more
elevated proximally and capped by obscure
small teeth, then a toothless notch basally.

Leg 2 reaching about to distal one-fourth
of palm; carpus with obsolescent distodor-
sal spine and tiny, acute, subdistal ventral
spine; merus with small subdistal dorsal
spine and moderate proximal mesioventral
spine; coxa with strong proximal and small-
er distal spine mesially. Leg 3 with slender
ventral spines on proximal half. Leg 4 with
spineless merus and ischium. Subchelate leg
5 reaching beyond ischium of cheliped.

Two arthrobranchs arranged in biserial
rows of undivided (entire) lamellae on max-
illiped 3 and legs 1-4.

Uropod with obsolescent spine on pro-
topod above base of endopod; nearly straight
distal margin of endopod exceeded by
curved margin of exopod, both rami with
dense fringe of setae and minute scattered
spines or granules on distal margin; endo-
pod with low, median longitudinal rib, and
less prominent lateral rib having shallowly
concave margin except for rounded, prox-
imal shoulder; exopod with three ribs, me-
sial sturdiest one bearing obsolescent spine
proximally, intermediate rib longer, lateral
rib least evident of all, slightly curved out-
ward but shallowly concave proximally.

Measurements in mm.—Holotype 2,
length anterior carapace 4.6, length cara-
pace 7.3, length chela including fixed finger
4.5, greatest height chela 1.2. Paratype 4,
length anterior carapace 2.4, length cara-
pace 3.3, chelae missing.

Known range. —Specimens examined are
limited to Jamaican localities listed above,
although one additional specimen of what
appeared to be U. corallifora was collected
from dead coral on the reef of Bellairs Re-
search Institute, Barbados in 1981. The lat-

408 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig.2. a, Mud-lined borehole ofjuvenile U. corallifora? exposed ina slabbed section of Pleistocene limestone,
with part of one individual intact; b, SEM micrograph x 560 of burrow lining composed of organic and inorganic
constituents.

VOLUME 102, NUMBER 2

ter specimen was lost and cannot now be
verified. It is possible that the species is
widely distributed, though rare, in other
parts of the Caribbean.

Known distribution of U. corallifora is
patchy. No specimens were found in recent
reef coral on the north coast of Jamaica,
where 188 kg of coral rubble and 97 kg of
living coral were surveyed quantitatively.
All specimens boring in recent reef rock were
found in much smaller, qualitative samples
taken at Drunkenmans Cay on the south
coast, near Kingston. Bioeroders are gen-
erally much more abundant in that area than
on the north coast, including a species of
boring mussel Lithophaga aristata (Dillwyn
1817) which does not occur at all in Dis-
covery Bay on the north coast of the island
(Scott 1988). Juveniles questionably re-
ferred to U. corallifora were, however, found
in Pleistocene limestone placed in Discov-
ery Bay during the building of an artificial
reef (Fig. 2). This limestone is much harder
and denser than natural, recent reef rock.
These juveniles have more robust chelipeds
than the adults described above, but resem-
ble the adults in other respects. We ques-
tionably refer them to the new species until
more material is available for comparative
study.

Habitat.—The habitat of U. corallifora
partly overlaps that of the much more abun-
dant U. operculata Schmitt 1924. Both
species inhabit dead coral skeletons in Ja-
maica, but occurrence of the former is much
restricted. It has not been found with bore-
holes opening from living corals, which are
a common habitat for U. operculata (Scott,
Reiswig & Marcotte 1988); see Warme
(1975) for distinction between boring in hard
and burrowing in soft substrates. A coral-
boring species, U. trypeta Sakai (1970, 1982),
has also been reported from Japan.

Boreholes of U. corallifora are similar in
appearance to those of U. operculata (Klee-
mann 1984; Scott, Reiswig & Marcotte
1988). Pairs of individuals (one 6, one 9)
live in complex, multi-chambered excava-

409

tions which are just wide enough to allow
the occupants to somersault and reverse di-
rection. Excavations are lined with a mix-
ture of mud, coral fragments and organic
material. Energy dispersive X-ray analysis
(EDAX) of borehole lining material re-
vealed the presence of Ca, Si, Mg, Al, Cl, K
& S. Openings to borings are difficult to
detect on the irregular surface of dead coral;
each opening lies a few millimeters below
the surface in a smooth conical pit and is
much narrower than the remainder of the
hole to which it gives access. Adult shrimps
are trapped in the borings by the restrictive
openings.

Remarks. — Upogebia corallifora is simi-
lar to U. noronhensis Fausto-Filho 1969
(comparison made with paratype °°, No.
163, carcinological collection, LABOMAR,
Universidade Federal do Ceara, Brazil, Fer-
nando de Noronha, Baia de Sueste, Fausto-
Filho, 7/8/68), which in turn has some sim-
ilarity to U. spinigera (Smith 1871) of the
eastern Pacific, but U. corallifora is appar-
ently a smaller and less spinose species than
either of the latter. Upogebia corallifora keys
out to U. noronhensis in Williams (1986:
10), but the two can be separated by means
of the following emendation to the con-
cluding couplet of his key to Atlantic species
of the genus.

— Chelae with no spines proximal to
fixed finger on ventral keel of palm

7. Chela with fixed finger about equal
in length to dactyl; palm spineless
on lower mesial surface

eee eee U. corallifora, n. sp.

— Chela with fixed finger about %
length of dactyl; palm with row of
more or less distinct spines on lower
mesial surface

lpg U. noronhensis Fausto-Filho
Etymology.—The name, a noun form
from the Latin “corallum,” coral, and

‘foro,’ bore or pierce, alludes to the coral-

boring capabilities of the species.

e, 0) lapresrette: tw) (oe) Gs) -el fe! 0) .a) ep 6 .e ce

410

Acknowledgments

We thank Dr. Jeremy Woodley and staff
of the Discovery Bay Marine Laboratory,
University of the West Indies, and Dr. Dun-
bar Steele, Chairman, Department of Zo-
ology, University of the West Indies, Mona
Campus, for facilities and assistance in the
field. Many thanks also to dive buddy and
cheerful coral basher Ian Sandeman. Con-
tribution No. 437, Discovery Bay Marine
Laboratory, University of the West Indies.

Literature Cited

Kleemann, K. 1984. Lebensspuren von Upogebia
operculata (Crustacea, Decapoda) in karibisch-
en Steinkorallen (Madreporaria, Anthozoa).—
Beitrage zur Palaontologie von Osterreich, In-
stitut flir Palaontologie der Universitat Wien 11:
35-49.

Sakai, K. 1970. A new coral burrower, Upogebia try-
peta sp. nov. (Crustacea, Thalassinidea) col-

lected from Amami-Oshima, Japan.— Publica-

tions of the Seto Marine Biological Laboratory
18(1):49-56.
1982. Revision of Upogebiidae (Decapoda,

Thalasssinidea) in the Indo-West Pacific re-
gion.— Researches on Crustacea, The Carcino-
logical Society of Japan, Special Number 1:1-
106, plates A-C.

Scott, P. J. B. 1988. Distribution, habitat and mor-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

phology of the Caribbean coral- and rock-boring
bivalve, Lithophaga bisulcata (d’Orbigny) (My-
tilidae: Lithophaginae).— Journal of Molluscan
Studies 54:83-95.

—., K.A. Moser, & M. J. Risk. 1988. Bioerosion
of concrete and limestone by marine organisms:
A 13 year experiment from Jamaica.— Marine
Pollution Bulletin 19(5):219-222.

—., H. M. Reiswig, & B. M. Marcotte. 1988. Ecol-
ogy, functional morphology, behaviour, and
feeding in coral- and sponge-boring species of
Upogebia (Crustacea: Decapoda: Thalassini-
dae).—Canadian Journal of Zoology 66:483-
495.

Warme, J. E. 1975. Borings as trace fossils, and the
processes of marine bioerosion. Pp. 181—227 in
R. W. Frey, ed., The study of trace fossils, a
synthesis of principles, problems, and proce-
dures in ichnology. Springer-Verlag, New York.

Williams, A. B. 1986. Mud shrimps Upogebia, from
the eastern Pacific (Thalassinoidea: Upogebi-
idae).—San Diego Society of Natural History,
Memoir 14:1-60.

(ABW) National Marine Fisheries Ser-
vice Systematics Laboratory, U.S. National
Museum of Natural History, Smithsonian
Institution, Washington, D.C. 20560; (PJBS)
Department of Geology, McMaster Uni-
versity, Hamilton, Ontario L8S 4M1, Can-
ada.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 411-417

ADDITIONAL RECORDS FOR AN ATLANTIC REEF
LOBSTER, ENOPLOMETOPUS ANTILLENSIS
LUTKEN, 1865
(CRUSTACEA, DECAPODA, ENOPLOMETOPIDAE)

Raymond B. Manning and David K. Camp

Abstract. — Additional records from the Gulf of Guinea, the Caribbean Sea,
Bermuda, and Florida are provided for E. antillensis Litken, 1865. The syn-
onymy of E. dentatus Miers, 1880, with E. antillensis is confirmed.

The astacidean genus Enoplometopus A.
Milne Edwards, 1862, now includes two
subgenera and nine species (Holthuis 1983:
282; Kensley & Child 1986:520; de Saint
Laurent 1988:61,62). Two representatives
of the subgenus Hoplometopus Holthuis,
1983, occur in the Atlantic: E. callistus Intés
& Le Loeuff (1970:1442), with its junior
synonym E. biafri Burukovsky (1972:180)
(see Holthuis 1983:281), from the East At-
lantic, and E. antillensis Litken, 1865, from
amphi-Atlantic localities. Another member
of this subgenus, EF. holthuisi Gordon, 1968,
is known from several localities in the Indo-
West Pacific (Holthuis 1983:297), and de
Saint Laurent (1988) recently described a
fifth species, Hoplometopus gracilipes, from
the Loyalty Islands. De Saint Laurent ele-
vated the subgenus Hoplometopus to ge-
neric status. All five representatives of the
nominal subgenus occur only in the Indo-
West Pacific (Holthuis 1983; Debelius 1984,
1986; Kensley & Child 1986).

De Saint Laurent (1988) also established
the family Enoplometopidae and _trans-
ferred the family from the Thalassinidea to
the Astacidea.

Holthuis (1983:281-282) enumerated
morphological differences between the two
subgenera, and it appears that these are
partly accompanied by differences in color
pattern also. Members of the nominal sub-
genus show color patterns dominated by
scattered spots or lines of pigment on the

abdomen and carapace, whereas two of the
four members of Hoplometopus each have
distinctive irregular circles of pigment, one
white, one reddish, around a central white
spot on each side of the red to reddish-or-
ange carapace. Remnants of this pattern are
visible in recently preserved specimens of
E. antillensis reported here. The pattern is
clearly shown for specimens of E. antillensis
and E. holthuisi by the following authors:
Cousteau (1958, color figure of E. antillensis
on p. 393); Forest (1959, black and white
print of E. antillensis; same figure as in
Cousteau 1958); Debelius (1984, color fig-
ure of E. holthuisi on p. 37); Debelius (1986,
color figures of EF. antillensis and E. hol-
thuisi on p. 13); Kosaki (1987, color figure
of E. holthuisi on p. 14); Brady (1987, black
and white figure of E. antillensis on p. 26);
and Hunziker (1988, color figures of E. hol-
thuisi on pp. 83 and 84). The pattern also
was described by Holthuis (1983:297) based
on color figures of E. holthuisi published by
Daum (1982) and George & George (1979).
In contrast, E. callistus lacks this distinctive
color pattern (P. Le Loeuff, in litt.), as does
H. gracilipes (M. de Saint Laurent, in litt.).

Until recently, Enoplometopus antillensis
had been recorded only once in the West
Atlantic north of Brazil (1.e., ““West Indies,”
Litken 1865). Another nominal species, EF.
dentatus Miers, 1880, from St. Helena Is-
land, had been questionably synonymized
with E. antillensis by Holthuis (1946:79),

412

then more positively by Chace (1966:634)
and Gordon (1968:80, footnote), although
none of those authors had had an oppor-
tunity to compare the types of the two
species. Gordon (1968) published the first
detailed description of the type of E. den-
tatus, compared it with two unpublished
drawings of the type of E. antillensis, and
concluded that the two species were con-
specific. Examination by one of us (D.K.C.)
of material sent by Mr. John Brady for iden-
tification after the discovery of a population
of E. antillensis off Florida (Brady 1987)
revealed some apparent discrepancies be-
tween his material and Gordon’s descrip-
tion of the type of E. dentatus. Our finding
of the apparent discrepancies prompted us
to compare the types of the two species, that
of E. dentatus in the British Museum (Nat-
ural History), London (BM), and that of E.
antillensis in the Zoological Museum, Co-
penhagen (ZMC), and to examine other At-
lantic specimens of the genus in the collec-
tions of the National Museum of Natural
History, Smithsonian Institution (USNM),
the Florida Department of Natural Re-
sources (FSBC I), and the Indian River
Coastal Zone Museum (IRCZM). The re-
sults are presented here.

Enoplometopus antillensis Liitken, 1865
Figs. 1-4

Enoplometopus antillensis Litken, 1865:
265.—De Man, 1916:96.—Holthuis,
1946:72, pl. 5, fig. h, pl. 7, fig. b.—Chace,
1966:634—Gordon, 1968:80 [foot-
note].— Fausto Filho, 1970:55, figs. 1-2.-
Burukovsky, 1972:188, fig. 2.—Fausto
Filho, 1976:222, fig. 1.—Holthuis, 1983:
281, 282.—Debelius, 1986: lower right
fig, on p.-1 3.

Enoplometopus dentatus Miers, 1880:381,
pl. 15, fig. 7.-—De Man, 1916:96.—Hol-
thuis, 1946:72.—Gordon, 1968:80, figs.
lL, 3-/3—Intes and ke Locult..19 70:
1442.—Holthuis, 1983:281.

Enoplometopussp.—Gurney, 1938:296

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

[part; origin of figured specimens un-
known].— Forest, 1959: pl. 3, fig. 4.
nephropsid.— Cousteau, 1958: color fig. on
p. 393.
Enoplometopus Antillensis.—Brady, 1987:
26, unnumbered fig.

Previous records. —East Atlantic: 1°
25'10’S, 5°36'10”E, Sao Tomé Island, Gulf
of Guinea (Cousteau 1958, Forest 1959).

Central Atlantic: St. Helena Island
[15°58’S, 05°43'W] (Miers 1880, Chace
1966, Gordon 1968).—Off Brazil, 03°17’S,
29°57'W (late larva; Gurney 1938).

West Atlantic: Off Rio Grande do Norte
State, Brazil (Fausto Filho 1970).—Off
northeast coast of Brazil (Fausto Filho
1976).— West Indies (Lutken 1865, Gordon
1968).—About | mile NE of Lake Worth
Inlet, Florida (Brady 1987).

Material. —East Atlantic: Gulf of Guinea,
off Gabon, 3°30’S, 8°53’E, 110 fm (=201
m), Geronimo Sta. 227/2, 7 Sep 1963: 1
juvenile, cl 6.5 mm (USNM 136692).

Central Atlantic: St. Helena Island: | fe-
male, cl 19.5 mm (BM 68.57; holotype of
E.. dentatus Miers).

West Atlantic: Bermuda: Half way be-
tween Kitcheners and N.E. marker, from
gut of red hind caught in 10 fm (=18 m),
W. Soares, 24 Feb 1985: 1 male, cl 38 mm
(USNM).

East coast of Florida: Spanish Anchor
Reef, about 1 mile N of Lake Worth Inlet,
90 ft (=27 m), John Brady, 15 Dec 1985:
exuvium of 1 male, cl 30 mm (USNM
231216).—Same locality, 27 Aug 1985: 1
damaged female, tl about 110 mm (FSBC I
31885).—Larsen’s Valley Reef, about 1 mile
NE of Lake Worth Inlet, 65 ft (=20 m), John
Brady, 15 Feb 1986: 1 female, cl 43 mm
(IRCZM 89:6349).

West Indies: 1 female, cl 20 mm (ZMC;
holotype of E. antillensis Lutken).

Bahama Islands: Grand Bahama Island,
SE of Settlement Point, D. de Sylva et al.,
Sta..355, 27 Jul 1961: 1 female, cl 1525 mama
(USNM 126226).

Panama: Golfo de Mesquites, Bahia de

VOLUME 102, NUMBER 2

413

Pip. 1.

Holotype of Enoplometopus antillensis Liitken. a, Carapace, Lateral view; b, Carapace, dorsal view;

c, Third maxilliped: d, Cheliped: e, Abdomen, lateral view; f, Posterior margin of sixth abdominal somite and

tail fan.

Almirante, 9°25'30”N, 82°20’W, lobster
trap, 3-11 fm (=5-—20 m), Pelican Sta. 930,
23-25 May 1963: 1 male, cl 31 mm (USNM
170659).

Netherlands West Indies: Bonaire, in
caves, 30 ft (=9 m), R. V. Harrison, 21 Apr
1975: 1 ovigerous female, cl 31 mm (USNM
155655).

Remarks. —In June 1986, one of us
(R.B.M.) was able to compare the holotypes
of E. dentatus Miers and D. antillensis Lit-
ken at the British Museum (Natural His-
tory), London. As a result of this compar-
ison, we agree that these species are
conspecific, as already suggested by others.
Apparent discrepancies between Gordon’s
account of the type of E. dentatus and ac-
counts in the literature of E. antillensis are
largely attributable to the poor condition of
Miers’ type specimen, which apparently had
dried out at one time. We provide here some

sketches of the type of E. antillensis (Fig.
1).

In her account of the type of FE. dentatus,
Gordon noted that there was no inferodistal
spine on the merus of the second pereopod.
The spine is in fact present on all but one
of the legs still loose with the holotype (Fig.
2a); on the second pereopod it appears to
have been worn off. Gordon’s figure of the
posterior margin of the sixth abdominal so-
mite (see our Fig. 25) is a little misleading;
the margin is not nearly so smooth as she
shows it. Numbers of spines on the outer
surface of the dactyli of the third and fourth
pereopods could not be verified on the type
of E. dentatus, as the ends of the dactyli
were missing. The terminal spine of the an-
tennal scale is not rounded anteriorly as de-
scribed by Gordon; the tip appears to have
been broken off, possibly since she exam-
ined it.

414

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

a b
ip
i

Fig. 2. Holotype of Enoplometopus dentatus Miers
b, Telson.

The color pattern of this species 1s quite
distinctive. Overall, the body has an orange
to orange red cast, and the long setae fring-
ing the chelipeds are reddish; marginal spines
on chelipeds are red proximally and have a
white tip. Walking legs and third maxil-
lipeds are banded with white, as are meri
and fingers of the chelipeds; propodi of the
chelipeds are red with darker red granules.
In dorsal view, there is a darker red stripe

. a, Distal part of merus and carpus of second pereopod;

down the center of the body. On the cara-
pace each dorsal spine bears a white spot,
and on the abdomen there are white spots
on the darker median stripe, as well as two
prominent white spots laterally on each ab-
dominal somite. Much of the lateral surface
of the carapace is marked with wavy white
lines, which, anteriorly, are replaced by a
large, irregular, reddish circle surrounding
a white spot (Fig. 3).

Fig. 3. Enoplometopus antillensis Liitken. Color pattern of carapace of exuvium of male from Florida.

VOLUME 102, NUMBER 2

415

Fig.4. Enoplometopus antillensis Litken. Juvenile from Gabon. a, Carapace, dorsal view; b, Carapace, lateral
view; c, Cheliped: d, Abdomen, lateral view; e, Posterior margin of sixth abdominal somite and telson.

Records for E. antillensis given here ex-
tend the known range of the species in the
West Atlantic northward to continental
North America and Bermuda. Off Florida,
specimens have been captured or observed
(John Brady, in litt.) at the following local-
ities between West Palm Beach and Fort
Pierce: 26°41'22”N, 80°00'09”W, depth 60
ft (=18 m), four individuals seen at night
only; 26°47'75"N, 79°59'28”W, depth 100
ft (=30 m), 18 individuals seen during day
and night; 27°28'02”N, 80°09'23”W, depth
50 ft (=15 m), two specimens seen during
daytime. All specimens were seen on lime-
stone ridges that parallel the shoreline in
those areas. The lobsters are very cryptic,
usually hiding in crevices within the ledges
during day and emerging at night, except in
late March and early April in West Palm
Beach. Numerous specimens, as many as
five on one dive, were seen during the day.
and they appeared to display aggressive ter-
ritorial behavior. In contrast, few or none
were observed during the day at the same
location in May.

The immature specimen from the Gulf of
Guinea (Fig. 4) agrees well with other ma-
terial of this species, in spite of its unusually
deep occurrence in 201 m, and differs from
E. callistus in the characters listed by Intés
and Le Loeuff (1970). There are five rather
than four median spines and three rather
than two lateral spines on the carapace, and
there are two rather than three posterior
spines on the sixth abdominal segment. The
anterior telson spines are close together and
the claws are long and slender.

Gurney (1938) reported a late larva of an
Enoplometopus that may be identifiable with
this species.

According to M. de Saint Laurent (in litt.)
there is a previously unreported specimen
of this species from Madeira in the Muséum
National d’Histoire Naturelle in Paris.

Acknowledgments

Torben Wolff, Zoological Museum, Co-
penhagen, made possible the direct com-
parison of the holotype of E. antillensis Lut-

416

ken with that of E. dentatus Miers by sending
Liitken’s type to the British Museum (Nat-
ural History), London, where one of us
(R.B.M.) could examine both types togeth-
er. Both Dr. Wolff and R. W. Ingle of the
British Museum also provided working
space and access to their collections. We
thank Paula Mikkelsen for allowing us to
work with material in the Indian River
Coastal Zone Museum at the Harbor Branch
Oceanographic Institution, Link Fort, Flor-
ida. L. B. Holthuis made available a copy
of the 1972 paper by Burukovsky, for which
we thank him. John Brady of Sebastian,
Florida, first brought the occurrence of E.
antillensis off the east coast of Florida to
our attention, and subsequently made spe-
cial efforts to secure material for us; he also
provided us with color photographs of this
species. We thank M. de Saint Laurent, Mu-
seum National d’ Histoire Naturelle, Paris,
for telling us about the references to E. an-
tillensis from the Gulf of Guinea and Gur-
ney’s paper, which we otherwise would have
overlooked, and for her comments on a draft
of our manuscript. We thank Thomas H.
Perkins and William G. Lyons, Florida Ma-
rine Research Institute, for their comments
on the manuscript. The figures were pre-
pared by Lilly King Manning.

This is contribution number 198 from the
Smithsonian Marine Station at Link Port,
Florida.

Literature Cited

Brady, J. 1987. Rare reef lobster identified. — Florida
Scuba News, January 87:26.

Burukovsky, R. N. 1972. Enoplometopus biafri—
new lobster species of the family Nephropidae
(Decapoda, Crustacea).— Trudy Atlantniro 42:
180-189 [in Russian].

Chace, F. A., Jr. 1966. Decapod crustaceans from St.
Helena Island, South Atlantic.— Proceedings of
the United States National Museum 118(3536):
623-661, pls. 1-2.

Cousteau, J.-Y. 1958. Calypso explores an undersea
canyon.—The National Geographic Magazine
113(3):373-396.

Daum, W. 1982. Ueber die Gattung Enoplometopus.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Grossekrebse im Aquarium, 3.— Die Aquarien-
und Terrarien-Zeitschrift, 35(7):265—269 [not
seen].

Debelius, H. 1984. Armoured knights of the sea. 120

pp. Kernen Verlag, Essen.

1986. Reef lobsters: Genus Enoplometo-
pus.—Freshwater and Marine Aquarium 9(3):
12-17.

Fausto Filho, J. 1970. On the occurrence of Enoplo-

metopus antillensis Litken, 1865 (Decapoda,

Nephropidae) on the Brazilian coast.—Crusta-

ceana 18(1):55-59.

. 1976. Description of male of Enoplometopus

antillensis Litken, 1865 (Decapoda, Nephro-

pidae).—Crustaceana 30(2):222-224.

Forest, J. 1959. Introduction, 1. Campagne de la Ca-
lypso dans le golfe de Guinée et aux iles Principe,
Sao Tomé, Annobon (1956), X. Résultats scien-
tifiques des campagnes de la “Calypso,” 4.—
Annales de |’Institut Océanographique 37:4—36,
pls. 1-3.

George, J. D., & J. J. George. 1979. Marine life. An
illustrated encyclopedia of invertebrates of the
sea. 288 pp, pls. 1-128 [not seen].

Gordon, I. 1968. Description of the holotype of En-
oplometopus dentatus Miers, with notes on other
species of the genus (Decapoda).—Crustaceana
15(1):79-97.

Gurney, R. 1938. Nephropsidea and Thalassinidea.
Larvae of Decapod Crustacea, part V.— Discov-
ery Reports 17:291-344.

Holthuis, L. B. 1946. The Decapoda Macrura of the

Snellius Expedition, I.—Temminckia 7:1-178,

pls. 1-11.

1983. Notes on the genus Enoplometopus,
with descriptions of a new subgenus and two
new species (Crustacea Decapoda, Axiidae).—
Zoologische Mededelingen, Leiden 56(22):28 1-
298.

Hunziker, R. 1988. Curious crustaceans—the reef
lobsters.—-Tropical Fish Hobbyist 36(5):80-84.

Intés, A., & P. Le Loeuff. 1970. Sur une nouvelle
espéce du genre Enoplometopus A. Milne Ed-
wards du Golfe de Guinée: Enoplometopus cal-
listus nov. sp. (Crustacea, Decapoda, Homari-
dea). — Bulletin du Muséum National d’ Histoire
Naturelle, Paris (2)41(6):1442-1447, pl. 1.

Kensley, B., & C. A. Child. 1986. A new species of
Enoplometopus (Thalassinidea: Axiidae) from
the northern Philippines.—Journal of Crusta-
cean Biology 6(3):520-524.

Kosaki, R. K. 1987. Hawaiian cave crustaceans. —
Freshwater and Marine Aquarium 10(4):12-15.

Liitken, Chr. 1865. Enoplometopus antillensis Ltk.,
en ny vestindisk Hummer-Art.— Videnskabe-
lige Meddelelser fra Dansk Naturhistorisk For-
ening, Copenhagen 6:265-268.

VOLUME 102, NUMBER 2

de Man, J. G. 1916. Families Eryonidae, Palinuri-
dae, Scyllaridae and Nephropsidae. The Deca-
poda of the Siboga Expedition, Part III.—Si-
boga-Expeditie 39a?:1—122, pls. 1—4.

Miers, E. J. 1880. Crustacea Anomura and Macrura
(except Penaeidea). On a collection of Crustacea
from the Malaysian region, IJJ.—Annals and
Magazine of Natural History (5)5:370-384, pls.
14-15.

de Saint Laurent, M. 1988. Enoplometopoidea, nou-
velle superfamille de Crustacés Décapodes As-

417

tacidea.—Compte Rendu Hebdomedaire des
Seances de l’Académie des Sciences, Paris, Zoo-
logie (3)307:59-62.

(RBM) Department of Invertebrate Zo-
ology, National Museum of Natural His-
tory, Smithsonian Institution, Washington,
D.C. 20560; (DKC) Florida Marine Re-
search Institute, 100 Eighth Avenue, S.E.,
St. Petersburg, Florida 33701.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 418-433

STUDIES OF NEOTROPICAL CADDISFLIES, XL:
NEW SPECIES OF SMICRIDEA (SMICRIDEA)
FROM MIDDLE AMERICA AND THE WEST INDIES
(TRICHOPTERA: HYDROPSYCHIDAE)

Oliver S. Flint, Jr. and D. G. Denning

Abstract. —Twelve species are described in the genus Smicridea, subgenus
Smicridea: tobada (Tobago, Trinidad), anomala (Trinidad, Tobago), aurimac-
ula (St. Vincent), cartiensis (Panama), multidens (Panama), corralita (Mexico),
bulara (Trinidad), mirama (Panama), filicata (Costa Rica, Panama), circinata
(Panama), holzenthali (Costa Rica), and /atipala (Panama). Abdominal pher-
omone glands, found in segments 6 and 7 of the male, are described and
illustrated for S. breviuncata Flint and S. cuna Flint. Variations in male genitalia
are shown for these two species and S. turrialbana Flint.

The nominal subgenus of Smicridea, con-
taining over 60 species, is limited to the
New World. Its closest relationships are with
several similar genera from Tasmania and
Australia (Flint 1989). The New World
species are found from southwestern United
States to southern South America, with
species on all the larger Antillean islands.

Flint revised the systematics of the species
in North and Central America (1974a), in
the Lesser Antilles (1968), in central, north-
ern Venezuela (1981), and in Suriname
(1974b). Although the majority of species
from these regions are now namable, un-
described species continue to be collected,
and species descriptions and names are nec-
essary for impending faunal studies.

We have also obtained from the Univer-
sity of California, Berkeley those unnamed
specimens reported by McElravy et al.
(1981). The genitalia were missing from the
only example of “A,”’ so its identity remains
unknown. Species “‘B” is described as S.
filicata.

All species described here belong to the
subgenus Smicridea as defined by Flint
(1974a). Males of species in this subgenus
are easily distinguished form those in sub-
genus Rhyacophylax by a spur count of 1,

4, 4 (rather than 1, 4, 2), and by two pairs
of pheromone glands in the abdomen (lack-
ing in Rhyacophylax). These glands open
externally through the membranes between
segments 7 and 8 and segments 6 and 7, but
lie within segments 7 and 6 when not ex-
truded.

Within the subgenus Smicridea, the new
species tobada is placed in the fasciatella
group, and the remaining new species are
placed in the nigripennis group (Flint 1974a).
Smicridea anomala is quite unusual, be-
cause its phallus is very long, open and
membranous ventrally and contains a series
of heavy spines. These characteristics are
lacking in the fasciatella group. Species of
the nigripennis group, however, have some-
what similar phalli: generally open and
membranous apicad and bearing a wide as-
sortment of spines and processes. Therefore
we place anomala in the latter group and
recognize that when the Neotropical fauna
is better known it may be transferred to a
newly recognized species group.

Acronyms used in the text indicate depos-
itories of specimens as follows: CAS (Cali-
fornia Academy of Sciences, San Francisco,
California), DGD (D. G. Denning collec-
tion, Moraga, California), NMNH (Nation-

VOLUME 102, NUMBER 2

al Museum of Natural History, Smithson-
ian Institution, Washington, D.C.), UCD
(University of California, Davis, Califor-
nia), UMSP (University of Minnesota, St.
Paul, Minnesota), UPP (Fairchild Museo de
Invertebrados, University of Panama, Pan-
ama), WSU (Washington State University,
Pullman, Washington).

Smicridea (Smicridea) tobada, new species
Figs. 1-5

This species is related to S. bulbosa Flint
(Flint 1974b, figs. 191-193) on the basis of
the general structure of the genitalia and the
presence of apical sclerites in the phallus.
Diagnostic differences are a less bulbous
apex of the phallus, and especially, the basal
angulation of the phallotheca, which is very
prominent on the ventral margin, and the
broader apex of the clasper.

Adult.—Length of forewing, 4-5 mm.
Color fuscous, legs and antennae paler; fore-
wing fuscous with two white bands, apical
fringe white. Male genitalia (Figs. 1-2):
Ninth segment with anterior margin slightly
expanded. Tenth tergite elongate, narrow-
ing apicad in lateral aspect; in dorsal aspect
elongate, apex obliquely truncate, with small
apicomesal lobe. Clasper with basal seg-
ment elongate, parallel-sided; apical seg-
ment third length of basal segment, dorsal
margin with apex acute, enlarged ventrad
and nearly truncate, with cluster of small
setae (Fig. 3). Phallus (Figs. 4—5) tubular,
angled posteriad from base, with a distinct
angle near base which is most pronounced
on ventral margin; apex with a pair of dor-
solateral sclerites with apices curved mesad;
with a recurved internal sclerite in lateral
aspect, dorsal aspect of this sclerite with me-
sobasal section divided apically into two
lateral, bandlike sclerites that curve ventrad
and then basad.

Type material. —Holotype, ¢, Tobago: St.
John Province; Charlotteville, 14-21 Mar
1979, D. Hardy and W. Rowe (NMNHB).
Paratypes: Same data, 1 6, 1 2 (NMNH).

419

Trinidad: Arima Valley, Spring Hill Estate,
1500’, 1 Feb 1983, W. G. Downs, 24 4
(DGD, NMNH).

Smicridea (S.) breviuncata Flint
Figs. 26-29

Smicridea (S.) breviuncata Flint 1974a:18.

Minor differences are present between a
specimen from southern Costa Rica (Figs.
26-29) and figures in Flint (1974a, figs. 60-
63) prepared from a specimen from central
Costa Rica.

Pheromone glands of abdominal segment
7 are shown in Figs. 28, 29. The reticulated
capsule with glands extruded is translucent
and covered with dense silver colored spi-
nules (Fig. 28). The reticulated capsule with
glands not extruded is shown in Fig. 29.

Material examined.—Costa Rica: Las
Cruces, 20 Jul 1977, Evan A. Sugden, 1 6
(UCD).

Smicridea (S.) anomala, new species

This peculiar species is placed in the m-
gripennis group, although it differs radically
from all other described species in the ge-
nus. It alone has a phallus that is sclerotized
and elongate dorsally. The similarity be-
tween S. anomala and the nigripennis group
is the membranous, ventral opening of the
phallus which bears a series of large spines
internally.

Adult. —Length of forewing 4.5-5.0 mm.
Wings uniformly black in alcohol, heavily
setose; head, thorax black; antennae, legs
ochraceous. Spurs small on first and third
legs, large on second pair of legs. Male gen-
italia (Figs. 6—7): Ninth segment with an-
terior margin rounded, slightly produced.
Tenth tergite in lateral and dorsal aspects
wide, lightly sclerotized, tapering to a
narrow, rounded apex. Clasper basal seg-
ment expanded distally; apical segment
short, apex acute and curved ventromesad.
Phallus (Figs. 8, 9) semierect, sclerotized
dorsally; apex and ventral area membra-

~

—

—

420 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-9. Male genitalia of Smicridea. 1-5, tobada. 1, Lateral view. 2, Ninth and tenth terga and clasper,
dorsal view. 3, Apical segment of clasper, posteroventral view. 4, Apex of phallus, dorsal view. 5, Phallus, lateral
view. 6-9, anomala. 6, Lateral view. 7, Ninth and tenth terga and clasper, dorsal view. 8, Apical half of phallus,
ventral view. 9, Phallus, lateral view.

VOLUME 102, NUMBER 2

nous; 5—6 large, dark brown spines arise from
mesal surface and project to or beyond ven-
tral margin, a slightly darkened phallotre-
mal sclerite near apex.

Type material.—Holotype, 6, Trinidad
Island: Simla Research Station, 2-15 Jun
1981, W. J. Hanson and C. L. Clemons
(DGD, to be deposited in CAS). Paratypes:
Same data, 1 6 (DGD). Tobago: St. John
Province; Charlotteville, 14-21 Mar 1979,
D. Hardy and W. Rowe, 1 6 (NMNH).

Dr. W. H. Hanson informs us that the
specimens were collected in a light trap, with
“‘a few ponds and a stream about one-eighth
mile away in the heavily forested area.”’

Smicridea (S.) aurimacula, new species
Figs. 18-21

This species is a member of the nigripen-
nis group of Smicridea and is in the cariba
section where it is closely related to S. gren-
adensis Flint (1968, figs. 61-63). Smicridea
aurimacula shares with S. grenadensis the
slightly upturned and truncate apex of the
tenth tergum and six pairs of long internal
phallic spines. However, it differs conspic-
uously by the golden banding on the fore-
wings and the presence of straight lateral
spine and an apicolateral plate bearing sev-
eral small teeth on the phallus.

Adult. — Length of forewing, 4.5 mm. Col-
or golden brown, head and thorax with gold-
en hair; forewing with stripe of golden hair
in costal cell widening at stigma and apicad,
another broad golden stripe along posterior
margin almost touching anterior stripe at
chord, thus leaving a central, fuscus stripe
in center of wing which is almost bisected
at chord. Male genitalia (Figs. 18, 19): Ninth
segment with anterior margin vertical, mak-
ing a right angle posteriad subdorsally. Tenth
tergite short, apex truncate and slightly up-
turned in lateral aspect; in dorsal aspect
short, apex truncate, broad. Clasper with
basal segment long, extending beyond apex
of segment 10, expanded distally; apical seg-

421

ment elongate, apex acute. Phallus (Figs. 20,
21) elongated basally, open apically; apex
with ventral lobe entire, produced into a
slightly flattened mesal lobe, with dorsal
margin inrolled apically; with a large, paired,
basolateral spine, endotheca with five to six
pairs of large, apical spines and a paired,
apicolateral, lightly sclerotized sclerite bear-
ing several small spines, and a ventrobasal
Y-shaped sclerite.

Type material. — Holotype, 6, St. Vincent:
Hermitage, 14 Nov 1975, C. L. de Freitas
(NMNH). Paratypes: Same data, 16 6, 14 9
(NMNB).

Smicridea (S.) cartiensis, new species
Figs. 10-13

This species and S. multidens are the first
discovered continental species of the cariba
section of the nigripennis group. S. cartien-
sis and multidens are closely related, differ-
ing only in the armature of the endotheca.
In cartiensis the endotheca bears a few scat-
tered sclerotized small spines on its surface,
but in multidens there is a large basolateral
plate with many acute spines.

Adult. — Length of forewing, 4 mm. Color
fuscous, head with silvery hair mesally;
forewing fuscous with scattered silvery hair
basally, with two white bands, more distal
band arcuate and not quite reaching pos-
terior margin, apical fringe white. Male gen-
italia (Figs. 10, 11): Ninth segment with
small lateral lobe on anterior margin. Tenth
tergite short, apex divided into lateral and
mesal lobes, in lateral aspect apicodorsal
angle acute, mesal lobe mostly hidden; in
dorsal aspect short, lateral lobe truncate,
broad, mesal lobe shorter and darker than
lateral lobe. Clasper with basal segment long,
inflated distally; apical segment elongate,
apex acute. Phallus (Figs. 12, 13) elongated
basally, open apically; apex with ventral lobe
entire, produced into a tapered mesal pro-
jection; endotheca with dorsal margin light-

422 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

16

Figs. 10-17. Male genitalia of Smicridea. 10-13, cartiensis. 10, Lateral view. 11, Ninth and tenth terga and
clasper, dorsal view. 12, Phallus, ventral view. 13, Phallus, lateral view. 14-17, multidens. 14, Lateral view. 15,
Ninth and tenth terga and clasper, dorsal view. 16, Phallus, ventral view. 17, Phallus, lateral view.

ly sclerotized, with a large, paired, basolat- Type material.—Holotype, ¢, Panama:
eral spine, a pair of large, internal spines Intendency of San Blas; Rio Carti Grande,
apically and broad, short spines irregularly 2 km W. Nusagandi (9°20'N; 78°56’W), 5
located apicolaterally. Mar 1985, Flint and Louton (NMNH).

VOLUME 102, NUMBER 2

Paratypes: Same data, 7 6, 3 2? (NMNH,
UPP).

Smicridea (S.) multidens, new species
Figs. 14-17

This species is closely related to S. car-
tiensis. Diagnostic differences are in the ar-
mature of the membranous central lobe of
the endotheca. In multidens this lobe bears
a large basolateral plate armed with many
spines; in cartiensis the plate is broken up
and the spines are fewer and concentrated
on the lateral surface of the endotheca.

Adult. —Length of forewing, 4.5 mm. Col-
or brown in alcohol, denuded; forewing
membrane with pale band at level of stigma.
Male genitalia (Figs. 14, 15): Ninth segment
with small lateral lobe on anterior margin.
Tenth tergite short, apex divided into lateral
and mesal lobes, in lateral aspect, apico-
dorsal angle acute, mesal lobe mostly hid-
den; in dorsal aspect lateral lobe truncate,
broad, mesal lobe shorter and darker than
lateral. Clasper with basal segment long, in-
flated distally; apical segment elongate, apex
acute. Phallus (Figs. 16, 17) elongated ba-
sally, open apically; apex with ventral lobe
entire, produced into a mesal process; en-
dotheca with large, paired, basolateral spine,
a pair of large, internal spines apically, a
basolateral plate with many short spines and
an apicomesal membranous lobe bearing
sclerotized projections.

Type material.—Holotype, 6, Panama:
Bocas del Toro Province; Miramar [9°N;
82°15’W], 21-27 Feb 1979, H. Wolda
(DGD, to be deposited in UCD). Paratypes:
Same data, but 15-21 Nov 1978, 1 4
(NMNH); 28 Feb-6 Mar 1979, 1 6 (UPP).

Smicridea (S.) corralita, new species
Figs. 22-25

The species is related to S. cholta Flint
(1974a, figs. 90-93) and is distinguished by
the male genitalia. In corralita the tenth ter-
gite is shorter and broader in dorsal aspect
than in cholta, and the apex is shaped dif-

423

ferently in lateral aspect. The base of the
apicolateral lobe of the phallus is angled
sharply ventrad in lateral aspect, and the
dorsal arm is long and slender, as opposed
to being nearly straight ventrally with the
dorsal arm almost lost in cholta.

Adult. — Length of forewing, 4.0—4.5 mm.
Color fuscous, head with silvery hair mes-
ally; forewing fuscous with scattered silvery
hair basally, with two white bands, distal
band arcuate and not quite reaching pos-
terior margin, apical fringe white. Male gen-
italia (Figs. 22, 23): Ninth segment with an-
terior margin produced into a rounded lobe.
Tenth tergite narrow, apex upturned in lat-
eral aspect; in dorsal aspect lateral margin
expanded basally, apex broadly rounded.
Clasper with basal segment long, expanded
distally; apical segment elongate, apex acute.
Phallus (Figs. 24, 25) elongate basally, open
apically, with small dorsomesal lobe; apical
portion divided mesally into broad lateral
plates that are sharply angled ventrad at base,
apicodorsal arm long and slender bearing
small ventral tooth, ventral arm very short
and ending in 2 laterally directed teeth; in-
ternally with a pair of long, slender spines.

Type material.—Holotype, 6, Mexico:
Chiapas State; Corralito, 7 km W. Aba-
solo, 21 May 1981, C. M. and O. S. Flint,
Jr. (NMNH). Paratype: Same data, 1 °
(NMNH).

Smicridea (S.) bulara, new species
Figs. 30-33

This species is a member of the S. nigri-
pennis group and is similar to the Mexican
S. nahuatl Flint (1974a, figs. 86-89). S. bu-
lara is distinguished from nahuatl by the
upturned apex of the tenth tergite, by the
upturned apex of the ventral arm of each
ventrolateral phallic lobe, and by the longer
spines in the phallotheca.

Adult.—Length of forewing, 4.5 mm.
General color of body and wings fulvous in
alcohol; forewings with an indication of faint
pale band. Spurs of second and third pair
of legs heavily setose. Fourth segment of

424 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 18-25. Male genitalia of Smicridea. 18-21, aurimacula. 18, Lateral view. 19, Ninth and tenth terga
and clasper, dorsal view. 20, Phallus, ventral view. 21, Phallus, lateral view. 22-25, corralita. 22, Lateral view.
23, Ninth and tenth terga and clasper, dorsal view. 24, Phallus, ventral view. 25, Phallus, lateral view.

maxillary palpus about twice length of sec- _tergite slender, apex curved dorsad; in dor-
ond or third segment. Male genitalia (Figs. sal aspect (Fig. 31), posterior margin of ninth
30, 31): Ninth segment with anterior margin tergum dark brown and deeply incised; tenth
expanded near base, sternum narrow. Tenth tergites rounded apically; a brown pig-

VOLUME 102, NUMBER 2

425

Figs. 26-39. Male genitalia and pheromone glands of Smicridea. 26-29, breviuncata. 26, Lateral view, with
ventral view of apex of phallus. 27, Ninth and tenth terga, dorsal view. 28, Pheromone capsule of segment 7,
glands extruded. 29, Same, glands not extruded. 30-33, bulara. 30, Lateral view. 31, Ninth and tenth terga,
dorsal view. 32, Phallus, ventral view. 33, Phallus, lateral view. 34-39, cuna. 34, Phallus, lateral view. 35,
Phallus, lateral view with lateral portion of phallus detached to show translucent process (a), paired endothecal
spines (sp), and large membranous mesal lobe. 36, Lateral plate of phallus (b) and mesal tubular process (c).
37, Segment 7 pheromone capsule, gland extruded, ventral view. 38, Segment 7 pheromone glands extruded,
lateral view. 39, Segment 6 pheromone capsule, glands extruded, ventral view to left, dorsal view to right.

mented line divides tergites into fulvous and
translucent portions. Basal clasper segment
expanded distally; apical segment slender,
obtuse. Phallus (Figs. 32, 33) with dorsal
margin expanded, giving rise to a short

translucent ventrally curved lobe; ventral
lobe divided mesally into broad lateral
plates, dorsal arm produced into laterally
directed point, ventral arm a large, apically
acute lobe curved dorsad, in ventral aspect

426

apex of lobe acute and curved laterad; with
pair of long, dark internal spines, each bear-
ing a small mesal tooth near midlength.

Type material.—Holotype, 6, Trinidad
Island: Simla Research Station, 2-15 Jun
1981, W. J. dlansom’ and C."Yf.. Clemons
(DGD, to be deposited in CAS).

Smicridea (S.) mirama, new species
Figs. 40-43

This species is distantly related to S. cuna
Flint (1974a, figs. 106-109) and is easily
distinguished from it by the male genitalia.
The distinctive phallus has long, slender,
ventrolateral plate with reflexed apices in
both lateral and ventral aspects.

Adult. — Length of forewing, 4.0—4.5 mm.
Color brown in alcohol; forewing denuded,
membrane with pale band from stigma.
Male genitalia (Figs. 40, 41): Ninth segment
with anterior margin produced into a
rounded lobe. Tenth tergite narrow, apex
slightly upturned in lateral aspect; in dorsal
aspect base expanded laterally, apex broad-
ly rounded with small mesal indentation.
Clasper with basal segment long, expanded
distally; apical segment elongate, apex acute.
Phallus (Figs. 42, 43) elongate basally, open
apically, with a dorsally sclerotized, dor-
somesal appendage; ventrolateral process
long and slender, with apex reflexed dor-
solaterally; endotheca with 2 long and near-
ly straight internal spines, and centrally an
irregular sclerotized band, bifurcate apical-
ly.

Type material.—Holotype, 6, Panama:
Bocas del Toro Province; Miramar [9°N;
82°15'W], 14-20 Feb 1979, H. Wolda
(DGD, to be deposited in UCD). Paratypes:
Same Locality: 3 6; 18-24 Oct 1978, 1 4;
15-21 Nov 1978, 3 6; 20-26 Dec 1978, 8
6; 10-16 Jan 1979, 3 6; 7-13 Feb 1979, 4
6; 21-27 Feb 1979, 12 6; 28 Feb 6—Mar
1979, 11 6; 7-13 Mar 1979, 19 6; 14—20
Mar 1979, 8 8; 9-15 May 1979, 13 4; 16—-
22 May 1979, 11 6; 23-29 May 1979, 7 6;

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

22-28 Aug 1979, 2 4; 10-16 Oct 1979, 16
(DGD, NMNH, UCD, UPP).

Smicridea (S.) filicata, new species
Figs. 44-47

Smicridea undescribed sp. ““B”’: McElravy,
etal. LOST SS:

Smicridea filicata, S. mirama and S. ulva
Flint (1974a, figs. 102-105) are closely re-
lated. Each has a phallus with an elongate
dorsomesal structure, a pair of ventrolateral
spines, and the ventrolateral margins pro-
duced into one or two long processes. In
both filicata and ulva the ventrolateral mar-
gins of the phallus are produced into two
pairs of processes; in mirama there is only
one pair. In filicata the dorsalmost process
of each pair is much longer than the ven-
tralmost and its apex is sharply hooked lat-
erad. In u/va the ventralmost process slight-
ly surpasses the dorsalmost, and neither are
sharply hooked apicad.

Adult. — Length of forewing, 4 mm. Color
fuscous, appendages paler, frons with white
setae; forewing mostly denuded, a white
band at stigma, possibly a pale band basally.
Male genitalia (Figs. 44, 45): Ninth segment
with anterior margin produced into a
rounded lobe. Tenth tergite in lateral aspect
narrow, apex angled dorsad; in dorsal aspect
with apex rounded. Clasper with basal seg-
ment long, expanded distally; apical seg-
ment elongate, apex acute. Phallus (Figs. 46,
47) elongate basally, open apically, with a
declivous, dorsomesal appendage; with two
ventrolateral processes on each side, ven-
tralmost short and acute, dorsalmost longer
with apex sharply hooked laterad; endo-
theca with two long, nearly straight internal
spines, and centrally with long, lightly
sclerotized lobe, weakly divided on the mid-
line.

Type material. — Holotype, 3, Costa Rica:
Puntarenas Province; Rio Singri (9.057°N;
83.082°W), ca. 2 km (air) S. Finca Hele-
chales, 720 m, 21 Feb 1986, Holzenthal,

VOLUME 102, NUMBER 2 427

46

Figs. 40-47. Male genitalia of Smicridea. 40-43, mirama. 40, Lateral view. 41, Ninth and tenth terga and
clasper, dorsal view. 42, Phallus, ventral view. 43, Phallus, lateral view. 4447, filicata. 44, Lateral view. 45,
Ninth and tenth terga and clasper, dorsal view. 46, Phallus, ventral view. 47, Phallus, lateral view.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

428

“MOTA [RIJUDA “ATUO / JUSUISES JO soINjon.s sUIeS “Gh “MOTA
[e191] “/ PUL 9 S]USWdas jo ‘spue]s popniixe puke sofnsdes suoWIOIOY ‘Bp “7 OOT = SUT] [eS ‘DUND VaplUgIWG Jo sydeIBOIOTUN UOI}99J9 SUTUULIS “6 p—-8p “SBI

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VOLUME 102, NUMBER 2

Morse and Fasth (NMNBH). Paratype: Same
data, 1 6 (UMSP). Panama: Chiriqui Prov-
ince; Fortuna Dam Site [8°44’N, 82°16’ W],
23-29 Nov 1977 H. Wolda, 1 é(NMNH);
same, but 28 Dec 1977-3 Jan 1978, 1 4

(OPP):

Smicridea (S.) cuna Flint
Figs. 34-39, 48, 49

Smicridea (S.) cuna Flint 1974a:23.

This somewhat variable species is abun-
dant in central Panama. Minor differences
in the male genitalia, especially the phallus
(Flint 1974a, figs. 108, 109), are figured here.
The pheromone glands (Figs. 37-39, 48, 49)
are referred to as “reticulated bodies” in the
literature.

The phallus (Figs. 34, 35) has an enlarged
base and a dorsal margin that is produced
into a ventral curved translucent process (a).
Ventrad to this process is a small structure
with only the acute apex discernable. Be-
neath it are a pair of darkly pigmented, api-
cally curved endothecal spines directed pos-
terioly (sp). The ventral portion of the
endotheca (Figs. 35, 36) consists of a large
membranous lobe containing small, dark
pigmented sclerites (possibly the phallotre-
mal sclerites), which are hidden in lateral
aspect by the brown pigmented lateral plate
(b). The mesal surface of plate (b) gives rise
to a slender, tubular, opaque lobe (c).

Figs. 37-39, 48, 49 show the paired pher-
omone glands of segment 6 and segment 7
extruded through inter-segmental mem-
branes. Segment 7 pheromone glands (Figs.
37, 38, 49) are covered with minute silver-
colored spinules. The glands originate from
internal surface of a vase-like capsule with
a lacelike and irregular external membrane.
Segment 6 pheromone glands, extruded
through intersegmental membrane 6-7, are
smaller, but are also densely covered with
silver spinules (Fig. 39).

Material examined. —Panama: Canal
Zone; Lion Hill Island, 27 Nov 1981, R. B.
Kimsey, | 6; 27 Oct 1981, 1 6; Puma Island,

429

20 Nov 1981, R. B. Kimsey, 1 6; Juan Gal-
legos Island, 27 Nov 1981, R. B. Kimsey,
11 6; Barro Colorado Island, Mar 1963, R.
D. Akre, 16; 13 Mar 1963, Malaise Trap,
R. D. Akre 2 6; Mar 1963, 7 4, 1 9; 21 May
1980, Henk Wolda, 1 ¢(DGD, UCD, WSU).

Smicridea (S.) circinata, new species
Figs. 50-53

The species is closely related to S. tur-
rialbana Flint (1974a, figs. 94-97) with
which it co-exists at the Miramar site. Di-
agnostic characters are recognized only in
the phallus. In circinata a pair of large dorsal
processes arise from the phallus. The apex
of the ventral process is not hooked laterad,
and the basolateral spine is larger and more
curved than in S. turrialbana.

Adult. —Length of forewing, 3.5—-4.0 mm.
Color brown in alcohol; forewing denuded,
membrane with a pale transverse mark at
stigma. Male genitalia (Figs. 50, 51): Ninth
segment with anterior margin produced into
a rounded lobe. Tenth tergite in lateral as-
pect short, broad, with dorsal, subapical
point; in dorsal aspect expanded laterally at
base, narrowed apically with mesal inden-
tation. Clasper with basal segment long, ex-
panded distally; apical segment elongate,
apex acute. Phallus (Figs. 52, 53) elongate
basally, open apically; apex with a pair of
large dorsal spines gradually curving later-
ad, ventral lobe produced into a pair of pro-
cesses curving dorsad, pair of curved baso-
lateral spines ventrally, endotheca
membranous with indistinct mesal sclero-
tization.

Type material.—Holotype, ¢, Panama:
Bocas del Toro Province; Miramar [9°N;
82°15’'W], 14-20 Feb 1979, H. Wolda
(DGD, to be deposited in UCD). Paratypes:
Same data 18-24 Oct 1978, 1 6; 15-21 Oct
1978, 14 6; 29 Nov—5 Dec 1978, 3 6; 20-
26 Dec 1978, 1 6; 10-16 Jan 1979, 2 6; 7-
13 Feb 1979, 1 6; 21-27 Feb 1979, 11 6; 28
Feb—6 Mar 1979, 31 6; 7-13 Mar 1979, 4
6; 14-20 Mar 1979, 3 6; 16-22 May 1979,

430 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 50-56. Male genitalia of Smicridea. 50-53, circinata. 50, Lateral view. 51, Ninth and tenth terga and
clasper, dorsal view. 52, Phallus, ventral view. 53, Phallus, lateral view. 54-56, turrialbana. 54, Phallus of
holotype, lateral view. 55, Phallus of specimen from Miramar, Panama. 56, Phallus of specimen from Quebrada
Grande, Costa Rica.

VOLUME 102, NUMBER 2

5 6; 23-29 May 1979, 8 6; 4-10 Jul 1979,
3 4; 22-28 Aug 1979, 2 6; 10-16 Oct 1979,
3 6 (DGD, NMNH, UCD, UPP).

Smicridea (S.) turrialbana Flint
Figs. 54—56

Smicridea (S.) turrialbana Flint 1974a:22.

We have seen material of this species from
four widely separated localities in Costa Rica
and Panama. There are three different forms
of the dorsolateral point of the phallus. In
the holotype from Turrialba, Costa Rica,
the point is barely developed (Fig. 54). In
the material from Miramar, Panama, the
point is developed into a strong spine (Fig.
55), and in the examples from Quebrada
Grande and Reserva Forestal San Ramon,
Costa Rica, the point is produced into a
hook-like process directed vertically (Fig.
56). The remainder of the genitalia do not
appear to be significantly different. We be-
lieve this to be variation in a single species
and predict that examples from other lo-
calites will also differ in the development of
the dorsolateral point.

Material examined. —Costa Rica: Carta-
go Province; 3 mi W Turrialba, 18-21 Jun
1967, Flint and Ortiz, holotype 6 (NMNH);
Guanacaste Province; Quebrada Garcia
(10.862°N; 85.428°W), 10.6 km ENE Que-
brada Grande, 470 m, 8 Mar 1986, Hol-
zenthal and Fasth, 1 6 (NMNH); Alajuela
Province; Rio San Lorenzo and tribs.
(10.192°N; 84.510°W), Reserva Forestal San
Ramon, 1090 m, 2-4 Jul 1986, Holzenthal,
Heyn and Armitage, 6 6 (NMNH, UMSP).
Panama: Bocas del Toro Province; Mira-
mar [9°N; 82°15’W], 14-20 Feb 1979, H.
Wolda, 2 6 (UCD, NMNH).

Smicridea (S.) holzenthali, new species
Figs. 57-64

This unusual species appears to be dis-
tantly related to S. turrialbana. In holzen-
thali the internal plate and spine of the phal-
lus are greatly reduced to a small dorsal tooth

431

and a slightly curved ventral spine, neither
extending beyond the margin of the phallus.
The rodlike ventrolateral processes of the
phallus are flat and broad in holzenthali.

Adult. —Length of forewing, 4 mm. Color
fuscous, appendages paler; frons white;
forewing fuscous, with two white bands one
at stigma, the other halfway between first
band and wing base. Male genitalia (Figs.
57, 58): Ninth segment with anterior margin
produced into a rounded lobe. Tenth tergite
narrow, with upturned apex in lateral as-
pect; in dorsal aspect, apex obliquely trun-
cate. Clasper with basal segment long, ex-
panded distally; apical segment elongate,
apex acute. Phallus (Figs. 59, 60) elongate
basally, open apically, with a small dorso-
mesal appendage; ventrolateral plates long
and broadened distally, each bearing a short
hook apicodorsally; internal plate small,
terminating in a slightly curved ventral spine
and a short, erect, dorsal spine.

Type material. —Holotype, 6, Costa Rica:
Guanacaste Province; Rio Tizate (10.773°N;
85.449°W), 7.2 km NE Canas Dulces, 275
m, 28 Jun 1986, Holzenthal, Heyn and Ar-
mitage (NMNH). Paratype: Same data, 1 4
(NMNH).

Smicridea (S.) latipala, new species
Figs. 61-64

This species is related to both S. campana
Flint (1974a, Figs. 128-131) and S. sirena
Bueno (1986, figs. 7-9) on the basis of the
undivided, apicoventral lobe of the phallus.
It is distinguished from both species in the
details of the phallic structure. Only in /a-
tipala does the phallus bear internally two
pairs of long, slender spines, a mesal, un-
paired structure, and a broadly rounded apex
on the phallus, in ventral aspect.

Adult. —Length of forewing, 5.5 mm. Col-
or uniformly fuscous in alcohol. Male gen-
italia (Figs. 61, 62): Ninth segment with an-
terior margin produced into a narrowly
rounded lobe. Tenth tergite narrow, apex
upturned in lateral aspect; in dorsal aspect

432 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

64

Figs. 57-64. Male genitalia of Smicridea. 57-60, holzenthali. 57, Lateral view. 58, Ninth and tenth terga
and clasper, dorsal view. 59, Phallus, ventral view. 60, Phallus, lateral view. 61-64, Jatipala. 61, Lateral view.
62, Ninth and tenth terga and clasper, dorsal view. 63, Phallus, lateral view. 64, Phallus, ventral view.

VOLUME 102, NUMBER 2

apex broadly rounded. Clasper with basal
segment long, expanded distally; apical seg-
ment elongate, apex acute. Phallus (Figs. 63,
64) elongate basally, open apically; apex
produced into single broadly rounded lobe
beneath endotheca; internally bearing pair
of long, dark spines dorsally, another pair
of pale spines ventrally, and a single, shorter
mesal process.

Type material.—Holotype, ¢, Panama:
Chirigqui Province; Guadalupe Arrriba
(8°52'26"N; 82°33'13”W), 2200 m, 18-24
Apr 1984, H. Wolda, in light trap 20 m
above ground (NMNH). Paratypes: Same
data, 2 6, 1 2(NMNH); same, but 2-8 May
1984, 1 6, 1 2 (UPP); same, but 6-12 Feb
1985, 1 2 (NMNH); same, but 17-23 Apr
1985, 1 6 (NMNH); same, but 24-30 Apr
1985, 1 2 (NMNH); same, but 8-14 May
1985, 1 6, 1 2 (UPP); same, but 22—28 May
1985, 1 6 (NMNH); same, but 12-18 Jun
1985, 1 2 (NMNH); same, but 19-25 Jun
1985, 1 é (NMNH).

Acknowledgments

Thanks are given for loan of specimens
by R. D. Akre, Washington State Univer-
sity; E. H. Sugden, R. D. Schuster and R.
B. Kimsey, University of California, Davis:
W. J. Hanson, Utah State University; and
R. W. Holzenthal, University of Minnesota.
We also thank Eric P. McElravy, University
of California, Berkeley for his work locating
the material of Smicridea “A” and “‘B.”
Henk Wolda, Smithsonian Tropical Re-
search Institute, Panama made his light trap
collections from Fortuna, Guadalupe Ar-
riba, Miramar and Barro Colorado Island
available to us for study. Discussions of the
site and collecting methods at Fortuna are
found in McElravy et al. (1981), at Miramar
in Wolda & Flowers (1985) and Barro Col-
orado Island in Wolda (1982). Thanks are
extended to R. D. Schuster for the SEM
photographs.

433

Literature Cited

Bueno Soria, J. 1986. Estudios en insectos acuaticos
VII: Cinco nuevas especies de Trichopteros de
Mexico y Costa Rica (Trichoptera: Hydro-
psychidae).— Folia Entomologica Mexicana 68:
53-65.

Flint, O. S., Jr. 1968. Bredin-Archbold-Smithsonian

Biological Survey of Dominica, 9. The trichop-

tera of the Lesser Antilles.— Proceedings of the

United States National Museum 125 (3665):1-

60.

1974a. Studies of neotropical caddisflies,
XVII: The genus Smicridea from North and
Central America (Trichoptera: Hydropsychi-
dae).—Smithsonian Contributions to Zoology
167:1-65.

. 1974b. The trichoptera of Surinam. — Studies

on the Fauna of Suriname and other Guyanas

14 (55):1-151.

1981. Studies of neotropical caddisflies,
XXVIII: The Trichoptera of the Rio Limon Ba-
sin, Venezuela.—Smithsonian Contribution to
Zoology 330:1-61.

1989. Studies of neotropical caddisflies,
XXXIX: The genus Smicridea in the Chilean
subregion (Trichoptera: Hydropsychidae).—
Smithsonian Contributions to Zoology 472:1-
45.

McElravy, E. P., V. H. Resh, H. Wolda, & O. S. Flint,
Jr. 1981. Diversity of adult Trichoptera in a
““non-seasonal”” tropical environment.—Pro-
ceedings of the 3rd International Symposium on
Trichoptera: 149-156.

Wolda, H. 1982. Seasonality of Homoptera at Barro
Colorado Island. Pp. 319-330 in E. G. Leigh,
Jr., A. S. Rand, & D. M. Windsor, ed., The
ecology of a tropical forest: seasonal rhythms
and long-term changes. Smithsonian Institution
Press, Washington, D.C.

—. & R. W. Flowers. 1985. Seasonality and di-
versity of Mayfly adults (Ephemeroptera) in a
““nonseasonal” tropical environment.— Biotro-
pica 17: 330-335.

(OSF) Department of Entomology, NHB
105, Smithsonian Institution, Washington,
D.C. 20560; (DGD) 2016 Donald Drive,
Moraga, California 94556. Dr. Denning died
on February 7, 1988 while this paper was
in the final review stages.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 434-446

A REVIEW OF THE SHORE-FLY GENUS
POLYTRICHOPHORA CRESSON FROM ASIA
(DIPTERA: EPHYDRIDAE)

Wayne N. Mathis and Jin Zuyin

Abstract.— Asian shore flies of the genus Polytrichophora Cresson are re-
viewed. Included are four previously described species (P. brunneifrons (Mei-
jere), P. canora Cresson, P. duplosetosa (Becker), and P. pollinosa Miyagi). The
tribe Discocerinini is characterized and a key to Asian discocerinine genera is
provided; the monophyly of Polytrichophora is established, and a key to species
occurring in the Palearctic and Oriental regions is presented, including P. or-
bitalis (Loew), which was reported to occur in north Africa (we have not seen
specimens of this species from the Old World, however, and doubt its occur-
rence there). Polytrichophora luteicornis Cresson is determined to be a junior
synonym of P. canora Cresson. All Asian species treated are redescribed, and
characters of the male genitalia are illustrated.

This is the second paper of a series on the
Ephydridae or shore flies of Asia with em-
phasis on species that occur in China. The
first paper was on the genus Lamproscatella
Hendel (Mathis & Zuyin 1988).

The purposes of this paper are twofold:
First, we present an overview of the tribe
Discocerinini, including a key to genera that
are found or are reported to occur in Asia.
Second, we review the genus Polytricho-
phora Cresson, one of the genera of Dis-
cocerinini, from the same geographic area.
As part of the latter study, we have exam-
ined all primary types of species that have
been reported to occur in Asia, and for each
species, illustrations of the male genitalia
are provided. With the exceptions to be not-
ed below, these are the first illustrations of
these species.

Previous papers on Asian species of Poly-
trichophora are mostly of limited scope,
usually isolated descriptions as part of fau-
nistic reviews. No comprehensive treat-
ment exists for the Asian species, although
Cresson (1945) did provide a list of species
occurring in the Indoaustralian region. That
list included a complete synonymy, and for

some species, brief remarks, alluding to the
generic placement of the species or a no-
menclatural issue. More recently, Miyagi
(1977) treated the Japanese fauna, which
included two species of Polytrichophora, and
the figures he produced were the first avail-
able for these taxa. Two other important
works are the fairly recent catalogs of the
Oriental (Cogan & Wirth 1977) and the
Palearctic (Cogan 1984) components of the
Asian fauna. The catalogs provide a listing
of species, primary references, and an out-
line of their distributions.

In this paper, we adhere generally to the
methods and descriptive terminology that
were used in the first paper of this series
(Mathis & Jin 1988). For each species treat-
ed in this study, we have noted, as appro-
priate, its descriptive and nomenclatural
history as part of the synonymy. Specimens
for this study came primarily from the col-
lections of the Academy of Natural Sciences
of Philadelphia (ANSP), the Smithsonian
Institution (USNM), Washington, D.C., and
the Northwestern Teachers College (NTC),
Lanzhou, Gansu Province, China. Sexes are
similar, so descriptions are based on both

VOLUME 102, NUMBER 2

sexes except for male genitalia. The termi-
nology we have used for structures of the
genitalia is as indicated on Figs. 6-9. Five
cephalic and two venational ratios are used
commonly in the descriptions; these are de-
fined here for the convenience of the user.

Head ratio: Head height/head width. Both
measurements are the greatest dimensions
of the head from an anterior view.

Frontal ratio: Frons height/frons width.
The width measurement is the straight line
distance between the median margins of the
compound eyes at the level of the anterior
ocellus; the height measurement is the dis-
tance from the anterior margin of the frons
to a “‘line”’ through the posterior ocelli.

Facial ratio: Face height/face width. The
height measurement is the distance between
the oral margin and the dorsum of the fron-
tal suture; the width measurement is the
narrowest distance between the compound
eye across the face.

Eye ratio: Eye height/eye width. Both
measurements are the greatest dimensions
as taken from a lateral view.

Eye-to-cheek ratio: genal height/eye
height. Both measurements are the greatest
dimensions as taken from the head in lateral
view.

Costal vein ratio: The straight line dis-
tance between the apices of R,,3; and R,,;/
distance between the apices of R, and R,, 3.

M vein ratio: The straight line distance
along vein M between crossveins (dm-cu
and r-m)/distance apicad of dm-cu.

Tribe Discocerinini

Diagnosis.—Small to moderately small
shore flies, length 1.25 to 3.5 mm; usually
invested with considerable microtomen-
tum, especially frons and mesonotum.

Head: Ocellar setae inserted anterior to
level of anterior ocellus, sometimes only
slightly so; reclinate fronto-orbital seta in-
serted anterior to proclinate fronto-orbital
(if two proclinate fronto-orbitals present,
inserted in front of the larger one); posto-
cellar setae well developed, proclinate, and

435

slightly divergent, usually at least one-half
length of ocellar setae; arista with from five
to seven dorsally branching rays; face not
conspicuously pitted or rugose; gena, in-
cluding midportion, bearing setulae, its pos-
terior (postgenal) margin rounded.

Thorax: Mesonotum generally microto-
mentose, usually densely so, although with
some variation; supra-alar seta usually ev-
ident although sometimes reduced; acros-
tichal setae arranged in about 8 irregular
rows; prescutellar acrostichal setae approx-
imate and inserted behind level of poste-
riormost dorsocentral setae; scutellum usu-
ally densely setulose; both anterior and
posterior notopleural setae inserted at about
the same level.

Abdomen: Male genitalia: surstyli not ev-
ident, apparently lost or fused indistin-
guishably with the ventral margin of the
epandrium.

Key to Asian Genera of the
Tribe Discocerinini

1. Notopleuron with scattered setulae

in addition to two larger setae .... 2
— Notopleuron setulae absent, with
only the two larger setae ......... 3

2. Facial setae arranged in two dor-
soventral series, secondary series of
dorsolateroclinate setae laterad of
inclinate primary series; eye densely
invested with microscopic setulae
eV aLIge TH OBIS Polytrichophora Cresson

— Facial setae arranged in one series;
eye at most with sparse, scattered
microscopic setulae
Ee RS POETS Discocerina Macquart

3. Supra-alar seta well developed,
longer than posterior notopleural
seta; face with an upcurved seta at
lower lateral extremity
AP OB F) he SEIS Diclasiopa Hendel

— Supra-alar seta weak or absent, if
present, distinctly shorter than pos-
terior notopleural seta; face lacking
upcurved seta at lower lateral ex-
tremity

436

4. Hind tibia bearing spurlike, preapi-
cal seta ventrally; facial series com-
prised of from two to three large se-
tae; dorsal seta inserted slightly
medially and arising from distinct,
shining papilla, with a small, slightly
anaclinate seta laterad of dorsal seta;
generally microtomentose, cinere-
ous species, appearing dull

side ee ra tah te Hecamedoides Hendel

— Hind tibia lacking spurlike, preapi-
cal seta; facial series comprised of
two large setae, dorsal seta not aris-
ing from a shining papilla and lack-
ing a smaller seta laterad of dorsal
seta; mostly sparsely microtomen-
tose, shining to subshining species

Ditrichophora Cresson

ee © © ce ©

eo 8 © © © © © © © © © © ©

Polytrichophora Cresson

Polytrichophora Cresson, 1924:161. Type
species: Polytrichophora agens Cresson,
1924:161, orig. des. —Cogan and Wirth,
1977:327-—238 [Oriental catalog].—Co-
gan, 1984:137-138 [Palearctic catalog].

Diagnosis. —Small to moderately small
shore flies, length 1.25—3.0 mm; generally
densely microtomentose, grayish species.

Head: Eye conspicuously microsetulose;
facial setae usually comprising eight setae,
these generally decreasing in size from top
to bottom, appearing as two series due to
divergent orientation of setae in series; setae
of primary series inclinate (setae 1, 2, 5, and
7), generally larger than setae of secondary
series except for seta 2, which is much re-
duced and inserted laterad and sometimes
slightly ventrad of seta 1, seta 1 largest of
all facials, inclinate (cruciate with opposite
seta), but not arising from shining papilla;
setae of secondary series oriented dorsolat-
erally to laterally (setae 3, 4, 6, and 8), usu-
ally smaller than setae of primary series;
face lacking a distinctly anaclinate seta at
ventral extremity; parafacial narrow to
moderately wide, with or without setulae;
gena generally low.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Thorax: Notopleuron bearing numerous
setulae in addition to two larger setae; an-
terior notopleural seta inserted conspicu-
ously closer to posterior notopleural seta
than to postpronotal seta; supra-alar seta
reduced, about one-half length of postalar
seta.

Abdomen: Fourth tergum of male only
slightly longer than third. Male genitalia with
cerci fused ventrolaterally to median mar-
gin of epandrium.

Distribution. — This is one of the few gen-
era of Ephydridae that occurs worldwide,
and with approximately the same species
diversity in each region, whether temperate
or tropical. A dozen new species have been
segregated from the neotropics (Mathis, in
preparation), however, which, would make
that region by far the richest in species.

Natural history. —Specimens of Poly-
trichophora are usually associated with mud-
shore and sand-shore habitats (Deonier
1965) or rarely with a marsh-reed habitat
(Scheiring & Foote 1973). We usually found
the greatest diversity and abundance when
collecting on mud and sandy shores that had
a matlike covering of algae.

Discussion. — Polytrichophora, along with
Discocerina and Hydrochasma Hendel (New
World), form a monophyletic lineage within
the Discocerinini. An apomorphy to sub-
stantiate the monophyly of this group is the
setulose notopleuron. In other genera of the
subfamily Psilopinae, the notopleuron is
bare except for an anterior and a posterior
seta, both of which are inserted near the
ventral margin. In Polytrichophora, Hydro-
chasma, and Discocerina, however, the
notopleuron bears a few additional setulae
that are usually inserted slightly dorsad and
toward the anterior portion of the pleuron,
usually around the anterior notopleural seta.
Polytrichophora may be distinguished from
Discocerina or Hydrochasma by the sec-
ondary “‘series’”’ of facial setae (see generic
description), a character that is unique to
this genus and establishes its monophyly.
Its relationship to Discocerina and Hydro-

VOLUME 102, NUMBER 2

437

Figs. 1-5.
lateral view. 3. Fifth sternum and hypandrium, ventral view. 4. Aedeagal apodeme and aedeagus, lateral view.
5. Aedeagus, dorsal view.

chasma is not resolved, however; either or
both of these genera could be paraphyletic
and include Polytrichophora as a sublineage.

In the following key, we have included all
species that are reported to occur in the
Palearctic and Oriental Regions. One
species, P. orbitalis (Loew), which was de-
scribed from specimens collected in North
America, was reported to occur in North
Africa (Cogan 1984). We have not con-
firmed this, having studied no specimens
that are conspecific with P. orbitalis from
North Africa. In the event that this species

Polytrichophora orbitalis. 1. Epandrium and cerci, posterior view. 2. Epandrium and left cercus,

does occur there, we have included it in the
key and have also provided figures of the
male genitalia (Figs. 1-5) based on speci-
mens from North America.

Key to Palearctic and Oriental Species of
Polytrichophora

1. Mesonotum mostly brown, but
brown coloration extended laterally
only to level of posterior dorsocen-
tral setae, thereafter with gray-col-
ored microtomentum in a broad lat-

438

eral stripe; membranous aedeagal
flap closely appressed to aedeagus;
ventral margin of epandrium short
and broad in lateral view (Europe
and’ ASia)i).2).4 - P. duplosetosa (Becker)
— Mesonotum more extensively brown
colored, brown coloration extended
laterally to level of supra-alar setae,
gray-colored areas restricted to ex-
treme lateral and anterior margins;
membranous aedeagal flap, if pres-
ent, separated from aedeagus except
at base; ventral margin of epan-
drium long and narrow in lateral
view
2. Face narrow, facial ratio averaging
0.75; fore femur with ventral flexor
setulae longer than width of fore tib-
ia at midlength; aedeagus lacking
membranous flap at apex (North
America, ? North Africa) (Figs. 1-
2D) he a) yer 2” dom P. orbitalis (Loew)
— Face narrow, facial ratio averaging
less than 0.60; fore femur with ven-
tral flexor setulae shorter than width
of fore tibia at midlength; aedeagus
with apical, membranous flap .... 3
3. Fore coxa extensively invested with
whitish gray to gray microtomen-
tum, yellowish coloration faint; fore
femur with a posteroventral row of
6—7 spine-like setae (Japan)
kiione nea el ee P. pollinosa Miyagi
— Fore coxa mostly yellow; fore femur
with a posteroventral row of only 4—
5 spine-like setulae
4. Length of ventral process of epan-
drium short, equal to one-fifth width
of epandrium; aedeagus 3.5 x as long
as wide; aedeagal apodeme sub-
triangular ... P. brunneifrons (Meijere)
— Length of ventral process of epan-
drium long, narrow, digitiform,
equal to 2 width of epandrium; ae-
deagus 5 x as long as wide; aedeagal
apodeme subrectangular

P. canora Cresson

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Polytrichophora brunneifrons (Meijere)
Figs. 6-10

Discocerina brunneifrons Meijere, 1916:270.

Polytrichophora brunneifrons. —Cogan &
Wirth, 1977:327-—328 [generic placement;
Oriental catalog].—Cogan 1984:137
[Palearctic catalog].

Description.—Small shore flies, length
1.25 to 1.75 mm (holotype 1.72 mm).

Head: Head ratio averaging 0.85. Frons
rectangular, frontal ratio averaging 0.70,
generally microtomentose, microtomentum
tan; ocellar triangle more densely microto-
mentose than remainder of frons; parafrons
black, slightly orangish red along anterior
margin; anterior fronto-orbital plate with
very narrow white band immediately ad-
jacent to eye; ocellar setae inserted slightly
in front of anterior ocellus. Antennae most-
ly orange, but dorsal surface of pedicel and
lst flagellomere partially blackish; arista
with 5 dorsally branching rays. Face densely
invested with grayish to slightly bluish white
microtomentum; parafacial setulae hair-like;
parafacials lightly yellowish but becoming
whitish; gena white and densely microto-
mentose. Eye ratio averaging 0.70; eye-to-
cheek ratio averaging 0.11. Palpus yellow.

Thorax: Mesonotum densely microto-
mentose, mostly light brown but postpro-
notum, notopleuron, pleural sclerites whit-
ish to silvery gray. Fore coxa yellow; femora
yellow, invested with grayish microtomen-
tum except for base and apex; tibiae and
tarsi yellowish, apical tarsomere usually
slightly darker, brownish; fore femur with
a posteroventral row of 4 stout but acutely
pointed setae. Wing hyaline; costal vein ra-
tio averaging 0.80; M vein ratio averaging
0.52. Halter yellowish.

Abdomen: Dorsum generally unicolor-
ous, brown, darker than coloration of meso-
notum and very slightly shining. Length of
5th tergum of male nearly 1.5 x that of 4th;
5th sternum (Fig. 8) with anterior margin
rounded, lateral arms connected broadly at

VOLUME 102, NUMBER 2

oe epandrium
f\

é
o
°

aedeagal apodeme

aedeagus

on
SS
9 membranous flap

439

5th sternum

0.045mm

ee |

10

Figs. 6-10. Polytrichophora brunneifrons. 6. Epandrium and cerci, posterior view. 7. Epandrium and left
cercus, lateral view. 8. Fifth sternum and hypandrium, ventral view. 9. Aedeagal apodeme and aedeagus, lateral

view. 10. Aedeagus, dorsal view.

base, base not interrupted medially. Male
genitalia (Figs. 6—10) as follows: Epandrium
(Figs. 6—7) more or less elliptical in posterior
view, dorsal margin rounded, ventral mar-
gin reduced as 2 short, sinuate (in posterior
view), pointed processes, the length of which
is Slightly less than greatest width of a cercus
in posterior view; aedeagus (Figs. 9-10)
about 5 x as long as width at middle, broad-
er basally and with distinct taper at base
and apex, apex pointed and bearing a nar-
row, membranous flap that generally folds
back under aedeagus, flap about *%4 length of
aedeagus; aedeagal apodeme somewhat tri-
angular, dorsal process short; posterior arms

of hypandrium (Fig. 8) widely separated and
nearly 74 total length of hypandrium, ante-
rior margin with shallow but wide emargi-
nation.

Type material. —The holotype female of
Discocerina brunneifrons is labeled “K. Ja-
cobson Wonosobo Java [Indonesia] Mei
1909/9 29/927-929 op modderig grindban-
ken langs stroomen[?] water. [handwritten
and barely legible, = on mud and gravel banks
along flowing water]/Clasiopa brunneifrons
det. de Meijere. Type [black submargin;
species name and “‘type”’ handwritten]/HO-
LOTYPE [red with black border]/HOLO-
TYPE 2 Discocerina brunneifrons de

440

M[eijere]. det. B. H. Cogan 1971 [all but
“det. B. H. Cogan 197” handwritten]/Poly-
trichophora brunneifrons meij[ere] det. B.
H. Cogan 1973 [all but “det. B. H. Cogan
197° handwritten]/Polytrichophora brun-
neifrons (Meij[ere].) WWirth 74 [black sub-
margin; all but ““WWirth” handwritten].”
The holotype is double mounted (minute
nadel in rectangular block of foam plastic),
is in good condition, and is deposited in the
Instituut voor Taxonomische Zodlogie,
Universiteit van Amsterdam, The Nether-
lands.

Distribution. —Oriental: India (Assam),
Malaysia (Selangor), Pakistan (Dhamyal),
and Thailand (Bangkok, Chiengmai, Chol-
buri, Khon Kaen, Udon Thani). Several
other records listed in Cogan and Wirth
(1977), mostly from countries not listed
above, have not been confirmed.

Remarks. —The description above is
based primarily on the holotype of P. brun-
neifrons, which is a female. The illustrations
of the male genitalia are from a specimen
that was collected in Thailand (Khon Kaen
Province, Meung District).

Polytrichophora canora Cresson,
revised status
Figs. 11-15

Polytrichophora canora Cresson, 1929:165;
1945:62 [review of Indoaustralian
species]|.—Cogan & Wirth, 1977:328
[Oriental catalog; as a synonym of P.
brunneifrons}.

Polytrichophora luteicornis Cresson, 1929:
166; 1945:62 [review of Indoaustralian
species]|— Miyagi, 1977:20 [review, Ja-
pan].—Cogan & Wirth, 1977:328 [Ori-
ental catalog; as a synonym of P. brun-
neifrons|. New Synonym

Description. —Small shore flies, length of
males averaging 1.15 mm (1.0-1.30 mm),
females averaging 1.35 mm (1.15—1.55 mm);
body mostly bronze colored but with some
whitish gray microtomentum laterally; fore
femur with 4—5 short, stout setae along api-
cal half of posteroventral surface.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Head: Head ratio averaging 0.84. Frons
rectangular, frontal ratio averaging 0.71
(males) or 0.73 (females), subshining with
a sparse whitish gray microtomentum;
parafrons black, partially golden to yellow-
ish or slightly greenish dorsally; mesofrons,
fronto-orbital plate, and ocellar triangle
golden brown, sometimes slightly greenish;
ocellar setae inserted slightly in front of or
aligned immediately laterad of anterior
ocellus. Antennae mostly yellow, dorsal sur-
face of pedicel and 1st flagellomere black;
arista with five dorsally branching rays. Face
bearing three pairs of primary and four pairs
of secondary facial setae; parafacial setulae
hair-like; parafacials and gena ochraceous,
with silvery microtomentum, microtomen-
tum becoming denser on gena. Eye ratio
averaging 0.74; eye-to-cheek ratio averag-
ing 0.097. Palpus yellow.

Thorax: Mesonotum mostly bronze col-
ored, but with whitish gray microtomentum
anteriorly and laterally; pleura brown, with
densely whitish gray microtomentum. Fore
coxa yellow, brownish basally; femora
brown except base of fore femur yellow; tib-
iae mostly yellow but brownish in median
part; tarsomeres 1-4 yellow, Sth brownish;
fore femur with 4—5 short, stout, black setae
along apical half of posteroventral surface.
Wing hyaline; costal vein ratio averaging
0.84; M vein ratio averaging 0.50. Halter
yellowish.

Abdomen: Dorsum generally concolor-
ous with mesonotum but with whitish gray
microtomentum laterally and posteriorly.
Length of 5th tergum of male nearly 1.5 x
that of 4th; 5th sternum (Fig. 13) with an-
terior margin truncate. Male genitalia (Figs.
11-15) as follows: Epandrium (Figs. 11—12)
in posterior view oval except for ventral
processes; dorsal margin of epandrium
rounded; ventral process of epandrium
elongate, length greater than width of cer-
cus, parallel sided; aedeagus (Figs. 14-15)
5x longer than wide, broader basally, ta-
pered irregularly to apex; aedeagal flap ex-
tended laterally from apex, length about two-
thirds that of aedeagus; aedeagal apodeme

VOLUME 102, NUMBER 2

441

Figs. 11-15.

Polytrichophora canora. 11. Epandrium and cerci, posterior view. 12. Epandrium and left cercus,

lateral view. 13. Fifth sternum and hypandrium, ventral view. 14. Aedeagal apodeme and aedeagus, lateral view.

15. Aedeagus, dorsal view.

(Fig. 14) subrectangular, dorsal process
short; posterior arms of hypandrium (Fig.
13) about one-half total hypandrial length,
anterior margin mucronate.

Type material.—The holotype female of
Polytrichophora canora is labeled ‘‘Tai-
nan[,] 909.II./Formosa Sauter/1115/TYPE
No. 6330[,] Polytrichophora Canora 2 E. T.
Cresson, Jr, [pink; number and species name
handwritten].’> The holotype is double
mounted (minute nadel in rectangular foam
block), is in good condition, and is depos-
ited in the ANSP (6330).

The holotype female of Polytrichophora
luteicornis is labeled ‘‘Singapore[,] BirOf[,]
1898/6 [sic]/Property of Hung [handwrit-

ten] Loaned [pink]/Type No. Polytricho-
phora Luteicornis E. T. Cresson, Jr, [dark
pink; species name handwritten].’’ The ho-
lotype is double mounted (minute nadel in
rectangular form block), is in poor condition
(pin obscures several structures), and is
presently deposited in the ANSP.

Distribution. —Oceania: Belau (Pelau);
Federated States of Micronesia (Yap Is-
lands, Caroline Islands); Solomon Islands
(Rondova). Palearctic: China (Guang Xi);
Japan (Honshu, Ryukyu Islands, Iriomote-
jima, Okinawa). Oriental: Malaysia (Sara-
wak, Selangor, Singapore); Taiwan (Tai-
nan); Thailand (Bangkok).

Remarks. —This species is similar to P.

442

brunneifrons, but the genitalia of the male,
especially the long, slender processes at the
ventral margin of the epandrium, are con-
sistently different.

We could not identify the holotype of
either the senior or junior synonym with
complete assurance and have primarily
based our assignment of names to this
species on its distribution. The holotype of
P. canora was collected on Taiwan; that of
P. luteicornis in Malaysia. We have exam-
ined male specimens from Malaysia, and
Taiwan is near the center of the known dis-
tribution of this species. This also follows
the precedent of Miyagi (1977), although he
used the junior synonym as the name for
the Japanese specimens he studied. The male
specimen we illustrated is from China
(Guang Xi).

Polytrichophora duplosetosa (Becker)
Figs. 16-20

Clasiopa duplosetosa Becker, 1896:162.

Diclasiopa duplosetosa. — Becker, 1926:45.

Polytrichophora duplosetosa. —Cresson,
1929:166 [generic placement].— Cogan,
1984:137 [Palearctic catalog].

Description.—Small shore flies, length
1.30 to 1.85 mm.

Head: Head ratio averaging 0.80. Frons
rectangular, frontal ratio averaging 0.68,
generally brown microtomentose; ocellar
triangle more densely microtomentose than
remainder of frons, usually bearing from two
to four setulae near anterior margin; para-
frons black, slightly orange red along ante-
rior margin; anterior fronto-orbital plate
with narrow band of sparsely white micro-
tomentum immediately adjacent to eye;
ocellar setae inserted slightly in front of an-
terior ocellus. Antennae mostly orange, but
dorsal surface of pedicel and Ist flagello-
mere partially blackish; arista with 5 dor-
sally branching rays. Face densely invested
with grayish to yellowish gray microtomen-
tum; parafacial setulae hair-like; parafacials
lightly yellowish, concolorous with face but
becoming whitish; gena white and densely

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

microtomentose. Eye ratio averaging 0.67;
eye-to-cheek ratio averaging 0.12. Palpus
yellow.

Thorax: Mesonotum distinctly two toned,
with broad longitudinal brown vittae ex-
tended laterally to level of posterior dor-
socentral seta, thereafter lateral margin gray;
pleural areas mostly gray to whitish gray,
becoming lighter colored, more whitish,
ventrally. Fore coxa mostly yellow, at most
very lightly brownish basally but exten-
sively invested with silvery gray microto-
mentum; femora brownish gray to gray; fore
and mid tibiae mostly gray, hind tibia most-
ly yellow; tarsomeres 1-4 yellow, 5th, oc-
casionally 4th, brownish; fore femur with
four short, stout, black setae along apical
half of posteroventral surface. Wing hya-
line; costal vein ratio averaging 0.86; M vein
ratio averaging 0.51. Halter yellowish.

Abdomen: Dorsum generally concolor-
ous with mesonotum but with whitish gray
microtomentum laterally and posteriorly;
posterior margin of 5th tergum of male with
some gray microtomentum. Length of 5th
tergum of male nearly 1.5 x that of 4th; 5th
sternum (Fig. 18) with anterior margin near-
ly straight and with posterior cleft extended
to anterior margin, forming two triangular
sternites. Male genitalia (Figs. 16-20) as fol-
lows: Epandrium (Figs. 16-17) more or less
elliptical in posterior view, dorsal margin
rounded, ventral margin pointed in poste-
rior view but lacking a process, ventral mar-
gin blunt in lateral view; aedeagus (Figs. 19—
20) about 5 x as long as wide, an irregularly
shaped tube, membranous flap weakly de-
veloped, closely appressed to ventral sur-
face, about 0.4 length of aedeagus; aedeagal
apodeme (Fig. 19) subtriangular; posterior
arms of hypandrium (Fig. 18) moderately
close together and comparatively short,
slightly more than one-half total length, an-
terior margin rounded.

Type material.—The lectotype male of
Clasiopa duplosetosa, designated here, is la-
beled ““Oderwald [Poland. Silesia] 19/5.
37059 [handwritten]/lecto Typus Mathis
1984 [red; “‘lecto,”’ ““Mathis,”’ “1984” hand-

VOLUME 102, NUMBER 2

19

443

0.045mm

—— oo

Figs. 16-20. Polytrichophora duplosetosa. 16. Epandrium and cerci, posterior view. 17. Epandrium and left
cercus, lateral view. 18. Fifth sternum and hypandrium, ventral view. 19. Aedeagal apodeme and aedeagus,

lateral view. 20. Aedeagus, dorsal view.

written]. There are three other specimens
labeled ““Typus’’ from Oderwald; these are
designated as paralectotypes. A fourth para-
lectotype is labeled “‘duplosetosa Beck.
[handwritten]/Orsova V.37681. [number
handwritten; black submargins]/Typus
[red].’’ The lectotype male is double mount-
ed (minute nadel in rectangular block of
pith), is in good condition, and is in the
Humboldt Universitat.

Distribution. — Widespread in Europe and
extending into Asia as far east as Iran (Co-
gan 1984).

Remarks.—Except for records of P. or-
bitalis in North Africa, specimens of which
we have not studied, this is the only species
of Polytrichophora in the western Palearctic
Region.

There is considerable variation in the col-
oration of the face and parafacials in this
species. The color varies from a silvery
cream to a rusty gold. Moreover, the para-
facials are not always concolorous with the
face, although they usually are.

Polytrichophora pollinosa Miyagi
Figs. 21-25

Polytrichophora pollinosa Miyagi, 1977:
20.—Cogan, 1984:138 [Palearctic cata-

log].

Description.—Small shore flies, length
1.83 mm (holotype, the only specimen with
an intact abdomen).

Head: Head ratio averaging 0.84. Frons
rectangular, frontal ratio averaging 0.66,

ttt PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 21-25.

generally brown microtomentose; ocellar
triangle more densely microtomentose than
remainder of frons, usually bearing from two
to four setulae near anterior margin; para-
frons black, slightly orangish red along an-
terior margin; anterior fronto-orbital plate
with narrow band of sparsely white micro-
tomentum immediately adjacent to eye;
ocellar setae inserted slightly in front of an-
terior ocellus. Antennae mostly orange, but
dorsal surface of pedicel and Ist flagello-
mere partially blackish; arista with 5 dor-
sally branching rays. Face densely invested
with grayish to yellowish gray microtomen-

Polytrichophora pollinosa. 21. Epandrium and cerci, posterior view. 22. Epandrium and left
cercus, lateral view. 23. Fifth sternum and hypandrium, ventral view. 24. Aedeagal apodeme and aedeagus,
lateral view. 25. Aedeagus, dorsal view.

tum; parafacial setulae hair-like; parafacials
lightly yellowish, concolorous with face but
becoming whitish; gena white and densely
microtomentose. Eye ratio averaging 0.65;
eye-to-cheek ratio averaging 0.17. Palpus
yellow.

Thorax: Mesonotum densely microto-
mentose, mostly brown but postpronotum
and notopleuron silvery gray to gray; pleural
sclerites whitish to silvery gray, becoming
more light colored ventrally. Fore coxa
mostly yellow, at most very lightly brown-
ish basally, but extensively invested with
silvery gray microtomentum; femora

VOLUME 102, NUMBER 2

brownish gray to gray; fore and mid tibiae
mostly gray, hind tibia mostly with more
yellow exposed on base and apex; tarso-
meres 1-4 yellow, 5th, occasionally 4th,
brownish; fore femur with 4 short, stout,
black setae along apical half of posteroven-
tral surface. Wing hyaline; costal vein ratio
averaging 0.87; M vein ratio averaging 0.56.
Halter yellowish.

Abdomen: Dorsum generally concolor-
ous with mesonotum but with whitish gray
microtomentum laterally and posteriorly;
posterior margin of 5th tergum of male with
some gray microtomentum. Length of 5th
tergum of male nearly 1.5 x that of 4th; Sth
sternum (Fig. 23) divided into 2 diamond
shaped sternites. Male genitalia (Figs. 21-
25) as follows: Dorsal margin of epandrium
(Fig. 21) rounded in posterior view, pointed
ventrally in lateral and posterior views (Figs.
21-22), median margin about one-third dis-
tance from ventral margin, with small
pointed spur in posterior view; aedeagus
(Figs. 24-25) between 4—5x longer than
wide, basal one-half parallel sided, apical
one-half evenly tapered to apex; aedeagal
flap attached subapically, elongate, about
twice length of aedeagus; aedeagal apodeme
(Fig. 24) subtriangular; posterior arms of
hypandrium (Fig. 23) widely separated and
over two-thirds total length of hypandrium,
anterior margin with a distinct U-shaped
emargination.

Type material.—The holotype female is
labeled “‘Hiraizumi [Iwate-ken, Honshu]
Aug 25 1962/Japonia tohoku Prov. I. Mi-
yagi/Discocerina-type (Polytrichophora)
pollinosa I. Miyagi [except for “‘-type’’ the
data on this label are handwritten; red].”
The holotype is double mounted (minute
nadel in cardbord rectangle), is in good con-
dition, and is in the collection of I. Miyagi
(University of the Ryukyus, Okinawa, Ja-
pan). In the original publication, this spec-
imen is stated to be a male, but it is a female.

Distribution. —Palearctic. Japan: Hon-
shu: Iwate-ken (Hiraizumi); Aomori-ken
(Towada). Hokkaido: Horonobe; Barato.

445

Remarks. —This species is distinguished
from Asian congeners by the following char-
acters: fore femur with flexor row of from
six to seven stout setulae along posteroven-
tral margin; male genitalia as illustrated
(Figs. 21-25) and described above.

We found that separation of this species
from P. canora is virtually impossible on
external characters alone. The characters of
the male genitalia, however, are very dis-
tinctive and can be used to distinguish these
species.

Acknowledgments

For reviewing a draft of this paper and
offering constructive criticism, we thank T.
Pape and R. V. Peterson.

We also thank the following curators and
their respective institutions for the loan of
specimens, especially primary type mate-
rial: Academy of Natural Sciences of Phila-
delphia (Mr. Donald Azuma); Humboldt
Universitat, Berlin, DDR (Dr. H. Schu-
mann); Instituut voor Taxonomische Zoo-
logie, Universiteit van Amsterdam, Am-
sterdam, The Netherlands (Dr. Pjotr
Oosterbroek and Mr. Ben Brugge); School
of Health Science, University of the Ryu-
kyus, Okinawa, Japan (Dr. Ichiro Miyagi).

Literature Cited

Becker, Th. 1896. Dipterologische Studien IV:
Ephydridae.— Berliner Entomologische Zeit-
schrift 41(2):91-276.
. 1926. 56a, Ephydridae und 56b, Canaceidae.
In. Lindner, ed., Die Fliegen der palaarktisch-
en Region 6(1):1-115.
Cogan, B.H. 1984. Family Ephydridae. Pp. 126-176
in A. Soos and L. Papp, eds., Catalogue of Pale-
arctic Diptera. Volume 10. Clusiidae—Chloropi-
dae. Akad. Kiado. Budapest, 402 pp.
, & W. W. Wirth. 1977. Family Ephydridae.
Pp. 321-339 in M. D. Delfinado and D. E. Hardy,
eds., A catalogue of the Diptera of the Oriental
Region, Vol. III: Suborder Cyclorrhapha (ex-
cluding Division Aschiza). University Press of
Hawaii, Honolulu.
Cresson, E. T., Jr. 1924. Descriptions of new genera
and species of the dipterous family Ephydridae.
VI.—Entomological News 35:159-164.

446

. 1929. Studies in the dipterous family Ephyd-
ridae. Paper IJ.— Transactions of the American
Entomological Society 55:165-195.

1945. A systematic annotated arrangement
of the genera and species of the Indoaustralian
Ephydridae (Diptera). 1. The subfamily Psilo-
pinae.—Transactions of the American Ento-
mological Society 71:47-75.

Deonier, D. L. 1965. Ecological observations on Iowa
shore flies (Diptera, Ephydridae).—Iowa Acad-
emy of Science 71:496-510.

Mathis, W. N., & Jin Zuyin. 1988. A review of the
Asian species of the genus Lamproscatella Hen-
del (Diptera: Ephydridae).— Proceedings of the
Biological Society of Washington 101:540—-548.

Meijere, J. C. H. de. 1916. Studien iiber siidostasia-

tische Dipteren. XII. Javanische Dolichopodi-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

den und Ephydriden.— Tijdschrift voor Ento-
mologie 59:225-273.

Miyagi, I. 1977. Ephydridae (Insecta: Diptera). Jn
Fauna Japonica. Keigaku Publishing Company.
Tokyo, 113 pp.

Scheiring, J. F., & B. A. Foote. 1973. Habitat distri-
bution of the shore flies of northeastern Ohio
(Diptera: Ephydridae).—The Ohio Journal of
Science 73(3):152-166.

(WNM) Department of Entomology,
NHB 169, Smithsonian Institution, Wash-
ington, D.C. 20560; (JZ) Department of Bi-
ology, Northwestern Teachers College, Lan-
zhou, Gansu Province, China.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 447-457

ADELPHYDRAENA, NEW GENUS, AND TWO NEW
SPECIES FROM VENEZUELA, AND REMARKS
ON PHYLOGENETIC RELATIONSHIPS WITHIN

THE SUBTRIBE HYDRAENINA
(COLEOPTERA: HYDRAENIDAE)

Philip D. Perkins

Abstract.—A new genus of aquatic beetle, Adelphydraena (Coleoptera: Hy-
draenidae) and two new species are described. The new species are from south-
ern Venezuela: A. spangleri was collected from a stream at the base of the tepui
Cerro de la Neblina; A. orchymonti was collected from a stream near Puerto
Ayacucho. Synapotypic and plesiotypic character states of the genera Adelphy-
draena and Hydraena are discussed. Phylogenetic relationships within the genus
Hydraena, and evolution of the lineage leading to Hydraena s.s. are also dis-
cussed. Taxonomically and phylogenetically significant structures of the two

new species are illustrated.

The sandstone mesas in northern South
America, known locally as tepuis, have per-
haps a greater concentration of floral and
faunal endemics than any other area in the
Americas. The tepuis are of great age, ap-
parently having undergone relatively little
of the massive geological and climatic
changes which greatly modified South
America after separation from Africa. Some
of the endemic species found on or near the
base of tepuis are thought to represent phy-
logenetically relictual taxa. The new genus
Adelphydraena described below appears to
be such a phylogenetic relict.

A biological survey of one of the tepui
areas in Venezuela, Cerro de la Neblina Park,
was conducted in 1985 by several institu-
tions under the organization and direction
of the Venezuelan Government (Spangler
1985). A team from the Smithsonian Insti-
tution, comprised of Paul J. Spangler, Phyl-
lis M. Spangler, Robin A. Faitoute, and
Warren E. Steiner, surveyed the aquatic in-
sects. Collecting on the tepui and in streams
near the base, the team collected over 55,000
insects, including a total of 16,694 speci-
mens of aquatic Coleoptera, representing

many genera and families. Despite the re-
markably high number of specimens, with
collections from many habitat types, the
material included only a single specimen of
the new genus Adelphydraena.

In 1986, Paul J. Spangler collected aquat-
ic insects near Puerto Ayacucho, which is
about 400 miles north of Cerro de la Neb-
lina Park. Among 2419 specimens of aquat-
ic Coleoptera collected by Spangler was a
second specimen, representing a second
species, of Adelphydraena.

Members of the new genus Adelphy-
draena are phylogenetically revealing in that
they retain several ancestral characteristics
which are found in the derived state in the
putative sister-group, the widespread and
successful genus Hydraena. The combina-
tion of plesiotypic versus apotypic character
states of members of Adelphydraena reveal
part of the sequential development of de-
rived characters in the lineage leading to
Hydraena s.s.

Based on the extreme rarity of the spec-
imens, and remoteness of the habitat, it
seems unlikely that new material will soon
be forthcoming. Therefore, the new genus

448

and two new species are described at this
time.

Adelphydraena, new genus

Type species.—Adelphydraena spangleri,
new species.

Description. —Form: Body oblong, elytra
sometimes ovate. Size: Length 1.24—-1.72
mm, width 0.58—0.84 mm. Color: Brown;
pronotum lighter than head and elytra.
Head: Frons with a low carina at rear, ar-
cuate to posterior. Fronto-clypeal suture well
developed. Labrum long, projecting ante-
riorly, with deep median emargination to
form two lobes, each lobe approximately
parallel sided; labrum not expanded on sides
near base. Mentum long, sides slightly con-
vergent from posterior to anterior; apico-
median emargination well developed. Max-
illary palpi elongate, combined lengths of
articles longer than ventral surface of head;
ratios of articles 2, 3, and 4 (last) about 3:
1:2. Antennomeres 10 (5+5) (Fig. 11).
Mandibles with lateral surface visible in
dorsal aspect; not interlocking with labrum.
Pronotum: Sides produced, sometimes sub-
cordiform, margin weakly denticulate. Disc
with shallow transverse depression across
base, lacking posterointernal foveolae; a
weakly developed, incomplete transverse
depression slightly anterior to midlength,
depression slightly arcuate to posterior and
interrupted in middle. Elytra: Nine com-
plete rows of punctures, row ten reduced to
a few punctures in anterior. Explanate mar-
gin moderate to well developed. Proster-
num: Intercoxal process broad, flat, ex-
panded laterally behind coxae, coxal cavities
closed; postcoxal process with a low carina
on each side extended posteriorly from me-
dian margins of procoxae. A median lon-
gitudinal, low carina in front of coxae; ca-
rina extended beyond anterior margin to
form very small apicomedian process. An-
terior margin on each side of apicomedian
process with small acute process slightly
larger than apicomedian process. Trochan-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

tin concealed. Proepisternum wide, not
emarginate anteriorly, hence antennal cav-
ities without ventral opening. Low carina
separating hypomeron and proepisternum
becoming effaced in anterior 0.33. Antennal
cavities well developed. Mesosternum:
Mesosternum with low median longitudinal
carina and two carinae on each side, one
carina at mesosternal-mesepisternal suture
and one carina about midway between the
latter and median carina. Sometimes with
weakly developed carina extended ante-
riorly from each side of mesosternal inter-
coxal process. Mesosternal intercoxal pro-
cess short, broad. Metasternum: Low
median longitudinal carina in anterior 0.5.
Plaque size and shape varying (Figs. 2, 3).
Each side with two longitudinal low ridges
or cuticular thickenings, visible when in
fluid, one extended from middle of margin
of mesocoxal cavity and one extended from
near posterior limit of lateral carina of
mesosternum. Abdomen: Intercoxal ster-
nite slightly wider than mesosternal inter-
coxal process, fused with first sternum to
form plate, plate delimited on each side by
a low ridge extended posteriorly from me-
dian margin of metacoxal cavity. Suture be-
tween intercoxal sternite and first sternum
visible through cuticle, sometimes not ex-
pressed externally. Legs: Tarsi with five ar-
ticles. All femora moderately narrowed in
basal 0.33. Ventral Vestiture: Vestiture ex-
tremely short and dense.

Etymology. —Greek adelphe (sister), plus
Hydraena. Adelphydraena is the putative
sister-group of Hydraena.

Adelphydraena spangleri, new species

Type data.— Venezuela, Territorio Fed-
eral Amazonas, Cerro de la Neblina, 1 km
SE. Basecamp, 0°50'N., 66°10’W., 140 m,
21 Feb 1985, P. J. and P. M. Spangler, R.
A. Faitoute, W. E. Steiner. Holotype female
deposited in the National Museum of Nat-
ural History, Smithsonian Institution.

Description.—Size: Body 1.72 x 0.84

VOLUME 102, NUMBER 2

mm; pronotum 0.40 x 0.68; elytra 0.96 x
0.84. Color: Brown-testaceous. Head: Frons
finely punctate, punctures smaller than eye
facets; shiny in middle, microreticulate and
dull laterally; posterior line not extended
forward at juncture of shiny and microre-
ticulate areas. Clypeus strongly microreticu-
late. Labrum slightly longer than clypeus;
median emargination setose, length nearly
0.66 labrum length. Eyes small in dorsal
aspect, width 0.3 interocular distance.
Mentum elongate, narrowed anteriorly, lobe
on each side of apicomedian emargination
curved ventrally and toward midline; apical
width 0.5X basal width. Genae without
posterior ridge. Pronotum: Pronotum sub-
cordiform. Posterior 0.5 of disc with raised,
rounded relief on each side of midline, re-
liefs separated anteriorly by two large punc-
tures side-by-side, followed by a single
puncture in midline; reliefs very shiny, very
finely, sparsely punctate, punctures similar
to those of frons. Punctures posterior to re-
liefs very coarse and dense, forming two
transverse rows, rows separated from one
another by confluent interstices which form
transverse line slightly less raised than dis-
cal reliefs, transverse raised line connected
to each discal relief by similarly raised, short
longitudinal line on each side of midline.
Punctures anterior to discal reliefs coarse,
slightly smaller than posterior punctures,
shallow, bottoms apparently flat with cen-
tral seta, interstices shiny, 0.51—1.0 x punc-
ture diameter. All interstices anterior and
posterior to discal reliefs with extremely
sparse, very fine punctures similar in size
to those of discal reliefs. Lateral depressions
with punctures less developed and interstic-
es less shiny than punctate discal areas. Lat-
eral margin denticulate, more distinctly so
anterior to middle. Anterior margin weakly
trisinuate. Posterior margin weakly bisin-
uate. Elytra: Punctures on disc 1 x punc-
tures on anterior area of pronotal disc. In-
tervals raised slightly, shiny, 2—3 x puncture
diameter. Punctures in anterior 0.5 of rows
1-5 larger and more widely spaced than on

449

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tus.

Adelphydraena spangleri, new species, habi-

remainder of elytron, interstices 2—4 x
puncture diameter. Raised humeri con-
nected with slightly produced anterior an-
gles. Lateral margin strongly arcuate, ex-
planate margin well developed, at midlength
slightly wider than distal width of metatibia.
Anterior margin with extremely small, pos-
teriorly produced acute process at base of
second interval. Prosternum: Prosternal in-
tercoxal process as wide as mesosternal in-
tercoxal process; prosternal midline length
2.75 X prosternal intercoxal process width.
Prosternal postcoxal process flat between
submedian carinae. Mesosternum: Meso-
sternal intercoxal process length 1 x width,
apex slightly arcuate. Submedian carinae at-
taining thickened anterior border of meta-

450

sternum. Metasternum: Plaques well de-
veloped (Fig. 2). When in fluid, a dark
longitudinal line of strength visible, extend-
ed posteriorly from each mesosternal suture
to near posterior margin of metasternum.
Abdomen: Distance separating coxal cavi-
ties 2X mesosternal intercoxal process
width. Submedian carinae of first sternum
terminated slightly before posterior border
of first sternum, distance separating carinae
at posterior extreme slightly greater than
midline length of first sternum plus fused
intercoxal sternite. Sterna 1—5 with hydro-
fuge, sterna 6 and 7 shiny, sparsely pubes-
cent. Legs: All legs without apparent mod-
ifications. Flight wings: Fully developed.
Spermatheca: Fig. 13 (male unknown).

Habitat. —The holotype was seined from
rocks and leaf packs in the rapids of a white-
water stream. The stream was 5 m wide and
30 cm deep, with sand/gravel margins and
bottom; oxygen, 9 ppm; hardness, 0; pH,
5.0; water temperature 29°C The air tem-
perature was a sweltering 42°C (Spangler,
pers. comm.).

Etymology.—It is with pleasure that I
dedicate this new species to Paul J. Span-
gler, whose dynamic fieldwork has provided
a wealth of aquatic Coleoptera for system-
atic studies.

Adelphydraena orchymonti, new species

Type data.—Venezuela, Territorio Fed-
eral Amazonas, Puerto Ayacucho (40 km
S.) at Tobogan, 24 Feb 1986, P. J. Spangler.
Holotype female deposited in the National
Museum of Natural History, Smithsonian
Institution.

Diagnosis.—Adelphydraena orchymonti
and A. spangleri differ markedly in many
characteristics. A. orchymonti lacks the sub-
cordiform shape of the pronotum and
broadly explanate elytra seen in A. spangleri
(Figs. 1, 4), and is much smaller, 1.24 vs.
1.72 mm. The metasternal plaques are very
small in A. orchymonti and very large in A.
spangleri (Figs. 2, 3). The shape of the men-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

tum differs in the two species (Figs. 9, 10).
Additional differences detailed in the species
descriptions include sculpture of the prono-
tum and head, relative widths of elytral
punctures and intervals, and relative widths
of the intercoxal processes.

Description. —Size: Body 1.24 x 0.58;
pronotum 0.32 x 0.54; elytra 0.76 x 0.58.
Color: Brown, pronotum lighter than head
and elytra. Head: Frons finely punctate,
punctures smaller than eye facets; posterior
arcuate line with lateral area extended for-
ward. Clypeus finely punctate. Labrum as
long as clypeus; median emargination se-
tose, length slightly greater than 0.5 x la-
brum length. Eyes small in dorsal aspect,
width 0.2 interocular distance. Genae
without posterior ridge. Pronotum: Disc
with two slightly depressed, transverse areas
with large, well impressed punctures 4-5 x
eye facets, separated by thin walls: one area
occupying basal 0.25 of disc, other area an
ill-defined line, arcuate to posterior, across
disc slightly anterior to middle; relief be-
tween coarsely punctate areas very finely,
sparsely punctate, punctures | x frons punc-
tures, interstices shiny; area anterior to
transverse line of coarse punctures with
punctures becoming increasingly smaller
toward anterior, those at margin | x frons
punctures. Lateral depression finely sparse-
ly punctate, moderately produced slightly
anterior to midlength. Elytra: Punctures on
disc 1x largest pronotal punctures. Inter-
vals raised slightly, shiny, 1 x puncture di-
ameter on disc, slightly narrower at inter-
vals 4—6. Interstices slightly less than 1 Xx
puncture diameter. Anterior margin with
extremely small, posteriorly produced acute
process at base of second interval. Proster-
num: Prosternal intercoxal process as wide
as mesosternal intercoxal process; proster-
nal midline length 2.66 x prosternal inter-
coxal process width. Prosternal postcoxal
process slightly concave between subme-
dian ridges (Fig. 5). Mesosternum: Meso-
sternal intercoxal process length 0.66
width, apex slightly rounded. Submedian

VOLUME 102, NUMBER 2

4

Figs. 2-5.

451

2, Adelphydraena spangleri, metasternum and anterior abdominal sterna; 3, A. orchymonti, same;

4, A. orchymonti, body outline; 5, A. orchymonti, prosternum. (Scale lines equal 0.1 mm.)

Carinae not attaining thickened anterior
border of mesosternum. Metasternum:
Plaques extremely small, located at border
of median, triangular, shallow depression.
Apex of triangular depression joining low
median carina to form inverted Y (Fig. 3).
Abdomen: Distance separating coxal cavi-
ties 1.25 mesosternal intercoxal process
width. Submedian carinae of first sternum
diverging posteriorly, distance separating
carinae posteriorly equal to length of first
sternum plus fused intercoxal sternite. Legs:
Legs without apparent modifications. Flight

wings: Fully developed. Spermatheca: Fig.
14 (male unknown).

Habitat.—The holotype was collected
among rootlets along the shallow, slightly
sandy margin of the clear stream as it tum-
bled over the polished granite bedrock. The
stream was partially shaded, 3 m wide, with
a maxinuney depth of ca.50.cm: pH, 5.0:
oxygen, 12 ppm; hardness, 0; velocity, 46
cm/sec; water temperature, 25°C; and air
temperature, 33°C at the time of collection.

Etymology.—This new species is dedi-
cated to Armand d’Orchymont in recogni-

452

tion of his contributions to the systematic
literature of the family Hydraenidae.

Phylogenetic Relationships of the
Genus Adelphydraena

Synapotypic Characters of the
Subtribe Hydraenina

All members of the subtribe Hydraenina
share two distinctive derived characters: the
prosternal intercoxal process is expanded
laterally behind the procoxae, closing the
procoxal cavities, and the second article of
the maxillary palpus is elongate and slender
(Perkins 1981: 415, Fig. 148 C; 431). Mem-
bers of Adelphydraena also have these apo-
typic characters (Figs. 5, 12), and are, there-
fore, members of the Hydraenina.

Synapotypic Characters of Hydraena s.1.

In addition to possessing the apotypic
characters of the subtribe Hydraenina,
members of Hydraena s.|. have the follow-
ing, previously undiscussed, synapotypic
characters of the labrum, mandibles, and
mentum.

All members of Hydraena s.l. have the
labrum abruptly widened, slightly de-
pressed and thickened near the base. Each
mandible has a dorsolateral process which,
when the mandible is closed, fits tightly
against and over the widened area of the
labrum (fig. 7). The labral and mandibular
structures function together to form an in-
terlocking device. Interlocking may, in ad-
dition to adding rigidity to the labrum, re-
sult in better integration of movement of
the labrum and mandibles during feeding
and/or function to transfer to the labrum
some of the opening and closing power of
the strong mandibular muscles.

In all members of Hydraena s.\. the men-
tum has a prominent apicomedian process
(Fig. 8). This process is generally lobate in
profile and directed obliquely toward the
dorsal surface of the head. When the head
is seen in dorsal aspect the process is visible

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

because of the large median emargination
between the lobes of the labrum.

Synapotypic and Plesiotypic Characters of
Adelphydraena

The derived characters of the labrum,
mandibles, and mentum present in Hy-
draena s.\. are absent in members of Adel-
phydraena. The labrum is straight sided, and
the mandible lacks the anteriorly projecting
dorsolateral process (Fig. 6). The mentum
is emarginate apicomedially (Figs. 9, 10);
this probably represents a derived condi-
tion, but in a direction opposite that taken
in the lineage leading to Hydraena. The api-
comedian emargination of the mentum re-
lates to other modifications in the labium;
these labial structures will be described and
compared with other genera in a separate
paper.

The ancestral number of antennal articles
for the family Hydraenidae is almost cer-
tainly eleven (by outgroup comparison with
other Coleoptera). Members of Adelphy-
draena have ten articles (Fig. 11), whereas
members of Hydraena s.|. represent a fur-
ther derived step in reduction, having nine
articles.

The ancestral number of elytral striae for
the order Coleoptera is almost certainly ten
(Crowson 1981:42), as is also true for the
family Hydraenidae. Members of Adelphy-
draena have the reduced condition of nine
complete striae, with stria ten reduced to a
few basal punctures (Fig. 1). The elytra of
all members of Hydraena s.l. show at least
some increase in the number of striae (see
below). The reduced striae of Adelphy-
draena members is a distinctive dorsal apo-
typic character.

The prosternal intercoxal process of Adel-
phydraena is unique among Hydraenina in
the possession of a longitudinal ridge on
each side which extends posteriorly from
the median margin of the procoxal cavity.
The middle portion of the postcoxal pro-
cess, between the ridges, is flat or slightly

VOLUME 102, NUMBER 2

7
her 1
7

—_-+--
= -.

453

14

Figs. 6-14. 6, Adelphydraena orchymonti, labrum, mandible, and anterior region of head; 7, Hydraena riparia
Kugelann, same. 8, H. riparia, mentum; 9, A. orchymonti, mentum; 10, A. spangleri, mentum; 11, A. spangleri,
antenna; 12, A. orchymonti, maxillary palpus; 13, A. spangleri, spermatheca; 14, A. orchymonti, spermatheca.

(Scale line equals 0.1 mm.)

concave, and contiguous with the flat inter-
coxal area (Fig. 5). In members of Hydraena
the postcoxal process is raised in the middle,
varying from slightly rounded to strongly,
longitudinally carinate.

In members of Hydraena the proepister-
num is emarginate anteriorly, the emargi-

nation providing a ventral opening to the
pocket in which the antenna is held (cf. Per-
kins 1981, figs. 63F, 148C). This emargi-
nation is a synapotypic character for the ge-
nus Hydraena; variation in degree of
development of the episternal emargination
will be discussed in a separate paper. Mem-

454

bers of Adelphydraena retain the plesiotypic
condition, having the proepisternum non-
emarginate anteriorly, hence the well de-
veloped antennal pockets are not open ven-
trally (Fig. 5).

Nonpubescent, smooth and shiny sub-
median areas of the metasternum, termed
plaques, are characteristic for the genus Hy-
draena, and frequently provide species-spe-
cific characters. The presence of plaques in
Adelphydraena spangleri (large) and A. or-
chymonti (small) (Figs. 2, 3), suggests that
plaques were present in the common ances-
tor of Hydraena and Adelphydraena.

In all other Hydraenidae, a portion of the
true second sternum is visible between the
metacoxae. This can be referred to as the
intercoxal sternite (Perkins 1981:21). The
intercoxal sternite takes many shapes in the
family, but is always separated from the first
complete sternum (true third) by a suture.
In members of Adelphydraena the intercox-
al sternite and the first complete sternum
are fused, on the same plane, and together
form a plate. This plate is delimited on each
side by a raised line which originates near
the lateral margin of the intercoxal area, and
extends diagonally across the first sternum
(Figs. 2, 3). In A. spangleri the suture be-
tween the intercoxal sternite and the first
sternum is visible through the cuticle (tem-
porary slide mount), but no surface mani-
festation of the suture was detected, whereas
the suture is visible externally in A. orchy-
monti. Additionally, in both species the first
sternum is more tightly joined to the meta-
sternum than is usually seen in members of
Hydraena, adding to the rigidity of the area
surrounding the metacoxae. The discal plate
of the first sternum, which is similar to
structures found in the family Elmidae, is a
distinctive ventral apotypic character.

Phylogenetic Relationships within
the Genus Hydraena

Hydraena is a large genus found on all
continents. It is quite diverse, and has many

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

clusters of obviously closely related species.
Various, apparently monophyletic, sub-
groups of Hydraena were given formal,
subgeneric status by Rey (1886) and Kuwert
(1888). Of these subgenera, d’Orchymont
(1936) recognized only Hydraena s.s.,
Haenydra Rey, and Kuwert’s Phothohy-
draena, Holcohydraena, and Taenhy-
draena. Perkins (1981:61) compared Amer-
ican Hydraena with these subgenera and
found that the characters upon which Ku-
wert’s subgenera Holcohydraena and Taen-
hydraena were based appeared in only some
members of separate monophyletic clusters
of species, and concluded that Holcohy-
draena and Taenhydraena were not valid.
Species with the strial configuration found
in members of Haenydra and Phothohy-
draena are not known from the New World
(Perkins 1981).

In contrast to the views of d’Orchymont
(1936) and Perkins (1981), Ienistea (1968,
1978, 1982) considered the subgenera of Rey
and Kuwert worthy of full generic status.

In a revealing paper, Berthélemy (1986)
compared the elytral striae of several species
(principally Palearctic) representing the main
groups of Hydraena s.l. Using ten striae as
the ancestral condition (by outgroup com-
parison), Berthelemy reasoned that all species
with a duplication of striae resulting in the
maximum configuration of 3a4b4c2d2e (a-e
being nervures), are monophyletic and were
derived after the divergence of species with
less duplication. Since Hydraena riparia
Kugelann has this strial configuration, and
is the type-species of Hydraena, Berthélemy
concludes, correctly I think, that all species
with the derived condition of 3,4,4,2,2 are
rightly members of Hydraena s.s.

Perkins (1981:212) erected the genus
Spanglerina for four species of Central
American hydraenines. These four species
share several unusual, derived characters,
but also have the 3,4,4,2,2 strial configu-
ration. If the nonduplicated condition is
considered ancestral for Hydraena, and if
one assumes that strial duplications have

VOLUME 102, NUMBER 2

not been independently derived in different
lineages, then one would conclude that the
common ancestor of these four species must
have arisen within Hydraena s.s. Spangler-
ina, therefore, cannot be considered a taxon
of equal rank with Hydraena, and perhaps
for the time being can be treated as a sub-
genus until the elytra and other structures
of the diverse genus Hydraena are more
completely known.

The strial configurations illustrated by
Berthelemy (1986) are summarized below,
where the number of striae between the ner-
vures are given for 16 species. Also included
are six hypothetical intermediates, each in-
dicated by a (?). Species having primarily
two striae between nervures, but with some
areas with three punctures, are assigned a
value of 2+. Similarly, a value of 3— in-
dicates that there are primarily three striae
between the nervures, but with some areas
having two punctures.

2 2 2+ 2 2. --gracilis, truncata, bicuspidata,

polita
2 2 2+ 2— 2- -- minutissima
eee et 2 = sharpi
Pee ae bay 2 ==?
Rap La Dae ie OY at,
Dee oo lk 62) C-- pyemaea
ieee l|6|6CD == Servilia
Dae SVS 8) Rr a 4
3— 3-— 3-— 1 2. --pulchella
Soo 2— 2 =- stussinerni
Seow 4, lL) .2),.) =-stestacea
SMe ot 2 2 aH?
Semone ot 2. 2 --?
Seen 4" 282) ==?
3 4 3 2 #2 -- rugosa (reversal)
3 4 4 2 2. --exarata, quadricollis
3 4 4 2. 2+ -- riparia, brachymera

No species illustrated by Berthélemy has
the ancestral condition of ten simple striae
with no indication whatsoever of duplica-
tion. Species traditionally placed in the sub-
genus Haenydra (gracilis, etc.) have the least
amount of strial duplication, with duplica-
tion limited to a portion of stria six. Species
sometimes placed in the subgenus Hadren-
ya also have this configuration. Species in

455

Hydraena s.s. have the greatest amount of
duplication.

The clinical progression seen in the elytral
data can be interpreted as reflecting many
phylogenetic divergences from the line
which eventually resulted in the maximum
expression of strial duplication now known
(3,4,4,2,2+). Although further study may
reveal that some of the transitional config-
urations are actually reversals from a more
derived condition (Berthéelemy 1986), for
the sake of discussion we can assume that
transitional configurations did exist in some
species (or are present in as yet undiscov-
ered ones). This position would be consis-
tent with the uncontested transformational
nature of some extant configurations.

Although Berthélemy concludes that striae
2 through 6 are duplicated in Hydraenas.s.,
it would seem that in any instance where
there is a whole number increase without
intermediate steps known (for example the
increase from 3 to 4 striae between nervures
a and b, and the similar increase between
nervures b and c) we do not know for certain
which of the striae became duplicated. But
when transitional numbers exist, we can as-
sume that they truly represent retention of
the ancestral elytral condition for the mono-
phyletic group of which they are a part, and
from their configuration infer which stria
was duplicated to form the next higher whole
number configuration. For example, it is
from the partial duplication of stria 6 pres-
ent in gracilis (etc.), minutissima and sharpi
that we infer that the extra stria between
nervures b and c in testacea derives from
stria 6 rather than stria 5.

With respect to reversions, Berthélemy
suggests that perhaps the pulchella-pyg-
maea configuration (2+,2+,3—,1,2) isa re-
version from the Hydraena s.s. configura-
tion (3,4,4,2,2). However, this would skip
the intermediate configuration of 3,3,3,1,2.
H. testacea has this configuration and also
shares with pulchella-pygmaea the derived
state of only one stria between nervures c
and d. According to the subgeneric divisions

456

recognized by Berthélemy, the strial reduc-
tion from two to one between nervures c
and d occurred independently in at least
three separate lineages.

Based on retention of the presumed an-
cestral elytral configuration of Hydraena s.1.,
Berthélemy recognizes Haenydra as a valid
subgenus. he also recognizes Hadrenya, al-
though its members have the same config-
uration as Haenydra. This decision is ap-
parently based on the lack of known derived
characters shared by all species in these two
groups. Similarly, Berthélemy recognizes as
valid the subgenus Phothohydraena, the
members of which have a 3,3,3,2,2 config-
uration (e.g. festacea).

Therefore, formal subgeneric ranking is
based upon two criteria: (1) elytra with less
duplication than the maximum number
(<3,4,4,2,2) and (2) absence of any synapo-
typy with species which have the same ely-
tral configuration. If these criteria are ac-
cepted, then every species found to have a
truly intermediate strial configuration (..e.,
not resulting from reversals) and no clear
synapomorphy with members of the rec-
ognized subgenera, would be treated as rep-
resenting a new subgenus.

Interestingly, the formal categories of
subgenera as now recognized in Hydraena
are therefore not simply monophyletic clus-
ters of species which are easily recognized
(the “sorting categories” of taxonomists),
nor do they meet the criteria of sister-groups.

Evolution of the Lineage Leading to
Hydraena s.s.

Based on the apotypic characters dis-
cussed above, it can be said that in the sub-
tribe Hydraenina, the following structural
modifications were among those that
evolved in the lineage leading to Hydraena
S.S.:

1. Postcoxal process of prosternum de-
velops lateral extensions, closing
procoxal cavities

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

— Antenna becomes reduced from ten
to nine articles

— Labrum-mandible interlocking
structure develops

— Mental apicomedian process devel-
ops

— Proepisternal anterior emargination
develops

6. Elytral strial duplications develop

We can be fairly certain that modification
of the postcoxal process was the first of these
innovations in the Hydraenina because
members of Adelphydraena have this struc-
ture but retain the primitive condition for
the antennae, labrum, mandibles, and pro-
episterna. These four characters are thought
to be truly plesiotypic in Adelphydraena and
not a result of reversals from a derived con-
dition. The following considerations seem
to support this position: (A) Reduction in
number of antennomeres has occurred in
several lineages in the family, but no in-
stance of increase in number of antenno-
meres is known (Perkins, unpublished). (B)
The labrum-mandible interlocking struc-
ture 1s complex, involving corresponding
modifications to each structure. The form
of the labrum and mandibles in Adelphy-
draena do not suggest reversal from the de-
rived condition.

The mentum of Adelphydraena, based
upon comparison with other genera (Per-
kins, unpublished), probably does not rep-
resent the ancestral condition for the Hy-
draenina. The form of the mentum 1s derived
in both genera, but in opposite directions,
the apical area being enlarged in Hydraena
and emarginate in Adelphydraena.

The primitive number of elytral striae for
the Hydraenina is 10. In Adelphydraena
none of the striae show any degree of du-
plication, and stria 10 is reduced to an an-
terior remnant consisting of a few punc-
tures; consequently, there are only nine
complete rows. In known Hydraena there
is always some degree of strial duplication,
and no species show a loss of stria 10.

VOLUME 102, NUMBER 2

Of the six structural modifications, the
duplication of elytral striae is almost cer-
tainly the most recent development. All
members of Hydraena have the first five
derived characters but some members re-
tain the ancestral condition for the striae
(except stria six). Further, a transformation
series of increasing strial duplication is seen
in extant species of Hydraena.

The sequence of appearance of the four
other structural modifications, if indeed they
developed sequentially and not concurrent-
ly, is problematic. If they did develop se-
quentially, their relative positions along the
phylogenetic line leading to Hydraena s.s.
can only be resolved with the discovery of
as yet unknown taxa which have more of
these derived states than Adelphydraena, and
less than Hydraena.

Acknowledgements

I thank Paul J. Spangler for providing in-
formation about the habitats of the new
species, and for reviewing the manuscript.
I also thank Young T. Sohn for drawing the
habitus illustration.

Literature Cited

Berthelemy, C. 1986. Remarks on the genus Hy-
draena and revision of the subgenus Phothohy-
draena (Coleoptera: Hydraenidae).— Annales de
Limnologie 22:181-193.

457

Crowson, R.A. 1981. The biology of the Coleoptera.
Academic Press, London, 802 pp.

Ienistea, M. A. 1968. Die Hydraeniden Rumaniens

(Coleoptera, Hydraenidae).—Travaux du Mu-

séum d’Histoire Naturelle ““Gr. Antipa.”’ Bucu-

resti 8:759-795.

1978. Hydradephaga und Palpicornia, in J.
Illies ed., Limnofauna Europaea, 2nd edit., G.
Fischer, Stuttgart. XVII + 532 pp.

. 1982. Hydraenoidea de la Suisse (Coleoptera)

(Catalogue). — Archives des Sciences, Genéve 35:

303-316.

Kuwert, A. 1888. Generalubersicht der Hydraenen
de europaischen Fauna.— Deutsch Entomolo-
gische Zeitschrift. Berlin 32:113-123.

Orchymont, A.d’ 1936. Au sujet de la phylogenie du
genre Hydraena (Col. Palpicornia Fam. Hy-
draenidae).— Memoires du Musée Royal d’His-
toire Naturelle de Belgique (2nd ser.) 3:61-67.

Perkins, P. D. 1981. Aquatic beetles of the family
Hydraenidae in the Western Hemisphere: clas-
sification, biogeography and inferred phylogeny
(Insecta: Coleoptera).—Quaestiones Entomo-
logicae 16 (1980):3-554.

Rey, C. 1886. Histoire naturelle des coléoptéres de
France.—Annales de la Société Linnéenne de
Lyon 32(1885):1-190.

Spangler, P.J. 1985. A new genus and species of riffle
beetle, Neblinagena prima, from the Venezuelan
tepui, Cerro de la Neblina (Coleoptera, Elmidae,
Larinae).— Proceedings of the Entomological
Society of Washington 87:538-544.

U.S. Dept. of Agriculture, APHIS, Suite
518, 10 Causeway Street, Boston, Massa-
chusetts 02222.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 458-467

SYMPHURUS CALLOPTERUS
(CYNOGLOSSIDAE, PLEURONECTIFORMES), A
NEW DEEPWATER TONGUEFISH FROM
THE EASTERN PACIFIC

Thomas A. Munroe and Madhu N. Mahadeva

Abstract.—A new species of eastern Pacific tonguefish, Symphurus callop-
terus, is described from moderately deep continental shelf waters from Mexico
south to northern Peru. The species is characterized by a unique interdigitation
pattern (1-3-4) of dorsal pterygiophores and neural spines and the combination
of a well-developed pupillary operculum, high counts (105-114 dorsal-fin rays,
91-98 anal-fin rays, 57-61 total vertebrae), and four hypurals. It is further
characterized by a unique combination of pigmentation features including an
unpigmented peritoneum, the presence of a dark brown or black blotch on the
ventral portion of the ocular-side operculum, banding on the posterior head
and along the body, and especially by the occurrence of an alternating series
of distinct, black rectangular blotches and unpigmented areas on the posterior
two-thirds of the dorsal fin and throughout the anal fin. Comments on overall
size, size at sexual maturity, and co-occurrence with sympatric species of Sym-
phurus are provided. Examination of radiographs revealed an interesting anom-
aly in the number of neural and/or haemal spines occurring on preural centra.
Of 180 specimens examined, 27 (15%) had double neural and/or haemal spines
on the first and/or second preural centra. This anomaly is shown to result most

likely from the fusion of two centra.

Tonguefishes (family Cynoglossidae) are
small to medium-sized, sinistral flatfishes.
In the eastern Pacific, they range from
southern Oregon (Eschmeyer et al. 1983) to
northern Peru (Hildebrand 1946, Chiri-
chigno 1974) and inhabit diverse sedimen-
tary and bathymetric environments. Thir-
teen nominal species of Symphurus have
been described from the eastern Pacific and
there are at least four additional species to
be described. Knowledge of the eastern Pa-
cific species of this genus is inadequate, al-
though symphurine tonguefishes comprise
significant components of the benthic ich-
thyofauna in certain habitats (Bartels et al.
1983, 1984; Love et al. 1984). Bartels et al.
(1984) found that tonguefishes, including
several undescribed species (one of which
was S. callopterus), were a numerically

dominant component of the demersal fish
fauna in the Gulf of Nicoya, Costa Rica.
Since the earlier, mostly regional, studies
by Jordan & Evermann (1898), Meek & Hil-
debrand (1928) and Hildebrand (1946),
there has been no published comprehensive
revision of eastern Pacific Symphurus. The
most recent, and only comprehensive, treat-
ment of eastern Pacific tonguefishes is an
unpublished dissertation by Mahadeva
(1956) in which several new species were
recognized. Mahadeva recognized the new
species described herein based on ten spec-
imens. Since Mahadeva’s study, trawling on
the continental shelf off Mexico, Central
America and South America has resulted in
the collection of many additional tonguefish
specimens, including 292 specimens of this
species. Inclusion of this new material al-

VOLUME 102, NUMBER 2

lows us to present a more complete and ex-
panded description of the new species.

A detailed systematic revision of eastern
Pacific tonguefishes is in progress by the first
author, but final results are not expected
until a later date. Therefore, in order to make
the name of this species available to other
researchers conducting ecological and early
life history studies on eastern Pacific
tonguefishes, a formal description is pro-
vided at this time.

This paper is the first in a projected series
of papers on symphurine tonguefishes. Re-
visionary studies on eastern and western At-
lantic tonguefishes, based largely on Mun-
roe (1987) are forthcoming. Additionally,
results ofa survey of interdigitation patterns
of dorsal pterygiophores and neural spines
in the genus Symphurus is in preparation.
Completing this series will be a revision of
eastern Pacific and Indo-Pacific Symphu-
rus. A phylogenetic analysis of intrageneric
relationships based on detailed osteological
studies is also in progress and will comple-
ment these revisionary studies.

Type specimens of the new species are
deposited in the following institutions: Cal-
ifornia Academy of Sciences (CAS); Na-
tional Museum of Natural History (USNM);
Natural History Museum of Los Angeles
County (LACM); Scripps Institution of
Oceanography, Marine Vertebrate Collec-
tion (SIO); University of Miami, Rosenstiel
School of Marine and Atmospheric Sciences
(UMML); Florida State Museum (UF); and
the British Museum (Natural History)
(BMNH).

Methods. —Counts and measurements
were modified from Ginsburg (1951), Ma-
hadeva (1956), Menezes & Benvegnu (1976),
and Munroe (1987). Counts and measure-
ments are outlined briefly below. Meristic
data, exclusive of scale counts, were taken
from radiographs.

Interdigitation pattern (ID). — Patterns of
interdigitation of proximal dorsal pteryg-
iophores and neural spines were counted
and recorded for the first three, or in un-

459

usual cases, the first four interneural spaces.
The number of dorsal pterygiophores in-
serted into interneural spaces 1—3 was found
to be diagnostic for species or groups of
species of Symphurus (Munroe 1987). ID
patterns are indicated by a pterygiophore
formula such as 1-3-4. The 1-3-4 ID pattern
indicates one pterygiophore inserts in inter-
neural space one, three in interneural space
two, and four in interneural space three. The
first neural spine abuts directly against the
cranium so there is no obvious space be-
tween it and the cranium. Therefore, the
first interneural space reflected in the for-
mula is that between the first and second
neural spines.

Caudal-fin rays. —Previous authors
(Ginsburg 1951, Mahadeva 1956, Menezes
& Benvegnu 1976, Munroe 1987) have
found that caudal-fin-ray count is extremely
conservative within the genus. Previous
studies have included the ultimate dorsal-
and anal-fin rays in the caudal-ray counts.
Because the ultimate rays of the median fins
are supported by caudal elements and are
aligned along the same vertical plane as the
caudal-fin rays they are also included in cau-
dal-ray counts reported here.

Anal-fin rays.—Includes all rays and is
exclusive of the thick, muscular gonadal duct
preceding the first anal-fin ray.

Vertebral counts. —There are consistently
nine abdominal vertebrae, three without and
six with haemopophyses; abdominal ver-
tebrae counts are thus reported as (3+6).
Counts of total vertebrae include the urosty-
lar centrum (see Remarks section below).
Specimens with obviously damaged verte-
bral skeletons, underlying support bones, or
caudal fins were excluded from summaries
and analyses.

Scale counts. — Accurate, repeatable scale
counts are difficult to make in Symphurus,
especially for species in which the scales are
often abraded and lost during capture when
specimens are trawled from any appreciable
depths. Approximate counts were based on
partial scale counts, counts of scale pockets,

460

or a combination of the two. Longitudinal
scale row count includes the total number
of complete diagonal rows of scales along a
hypothetical line starting immediately above
the opercular angle to the end of the hypural;
the few rows of scales along the caudal-fin
base are not included and the last scale must
be at least halfin front of the hypural. Trans-
verse scale count is the number of scales in
a diagonal row from the base of the dorsal
fin at a point directly above the posterior
margin of the operculum to the base of the
anal fin. Scales extending out onto the dor-
sal- and anal-fin rays are not included. Head
scale row count includes all the oblique rows
of scales on the head counted posteriorly
from the first complete row of scales passing
the posterior border of the lower eye; it in-
cludes the last complete row of scales im-
mediately anterior to the mid-point emar-
gination on the posterior border of the
operculum, but the few small rows of scales
present on either the dorsal or ventral fleshy
lobes of the operculum are not included.
All measurements were made on the ocu-
lar surface except where noted. Measure-
ments were taken to the nearest 0.1 mm
with dial calipers (to 150 mm) or ocular
micrometer. Measurements are expressed
either as thousandths of standard length or
thousandths of head length. Standard length:
distance from tip of fleshy snout to posterior
end of hypural plate. Body depth: distance
across body at the anus, exclusive of fins;
measured on blind side. Preanal length: tip
of fleshy snout to origin of anal fin; mea-
sured on blind side. Caudal-fin length: base
of articulations of middle caudal-fin rays to
tip of longest middle rays. Pelvic-fin length
(only blind side pelvic fin present in adults):
basal articulation to distal tip of the longest
ray. Pelvic to anal length: shortest horizon-
tal distance from base of most posterior pel-
vic-fin ray to anal-fin origin. Head length:
tip of fleshy snout to most posterior exten-
sion of upper fleshy lobe of operculum. Head
width: greatest distance across head at pos-
terior portion of operculum. Postorbital

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

head length: posterior margin of lower eye
to posterior extent of upper fleshy lobe of
operculum. Upper head lobe width: dis-
tance at operculum from dorsal margin of
body to dorsal origin of operculum. Lower
head lobe width: distance at operculum from
dorsal origin of operculum to most ventral
part of operculum. Snout length: anterior
rim of lower eye to tip of snout. Upper jaw
length: shortest horizontal distance from
bony tip of premaxilla to angle of mouth.
Eye diameter: greatest horizontal diameter
of the cornea of the lower eye; does not
include fleshy tissue surrounding eye. Chin
depth: vertical distance from angle of the
mouth to most ventral aspect of the head.

Size at maturity was estimated by mac-
roscopic examination of the extent of pos-
terior elongation of the ovaries (ovaries of
mature females are conspicuous through the
body wall when light is transmitted from
beneath the specimen; in immature females,
developing ovaries are best observed by dis-
section). Since no obvious differences in tes-
ticular size were apparent in males, esti-
mates of maturity were based entirely on
females.

Depth of capture (in feet or fathoms) was
transformed to the nearest meter. If depth
included a range of depths over which the
nets were towed, an average depth for that
particular trawl was calculated.

Symphurus callopterus, new species
Figs. 1-2

Holotype. —CAS 64023 (male, 116.9 mm
SL); Peru; 05°59'S, 81°)2°W tomO0Se56 ss.
81°11’W, 65-70 m, 4 Jun 1966. Collected
with 70 ft Gulf of Mexico semi-balloon
trawl, R/V Anton Bruun Cruise 16, Station
631 B.

Paratypes.—(Measured and counted 20
specimens; 93.2—-127.6 mm SL): Costa
Rica.—USNM 164492, 2(93.2—96.1), 6 mi
off Cabo Velas, Papagayos, 101 m, 01 Jul
1932.—UF 47590, (117.1), Puntarenas
Province, Gulf of Nicoya, 317 m, 29 Jun

VOLUME 102, NUMBER 2

461

Fig. 1.

1973. Panama Bay.—UMML 26069,
15(97.3-127.6), 8°00.4—00.7'N, 79°48.8-
41.5'W, 78 m, 4 May 1967.—UMML
34317, 2(104.1-121.7), 7°40.3'N, 79°50.9'W;
78 m, 2 May 1967. Counts were also taken
from the following 46 paratypes (88.5-—145.7
mm SL): Mexico, Gulf of Tehuantepec. —
SIO 63-518, 9(99.5-136.6), 15°57.5'N,
95°22.5'W, 65 m, 12 Jul 1963.—SIO 63-
521, 2(96.5—105.6), 15°41'N, 96°07.5’W, 55
m, 14 Jul 1963. Costa Rica. —BMNH
1956.3.1:3-4, 2(88.5-94.5), 6 mi off Cabo
Velas, Papagayos (ca. 10°22’N, 85°53’W),
101 m, 1 Jul 1932.—LACM 33827-55,
4(96.6-119.9), Gulf of Nicoya (ca. 9.5°N,
85°W), 29 Jun 1973. Panama Bay.— UMML
26069, 18(92.4-128.3), 8°00.4—-00.7'N,
79°48 .8-41.5'W, 78 m, 4 May 1967. Peru. —
CAS 24189, 11(109.7-145.7), (same as ho-
lotype).

Diagnosis.—A medium-sized Symphu-
rus, reaching 162 mm SL (most specimens
90-140 mm SL), distinguished from all oth-
er species in the genus by a unique 1-3-4
ID pattern and the following combination
of characters: 105-114 dorsal-fin rays; 91-—
98 anal-fin rays; 57-61 (57 and 61 rare) ver-
tebrae; 12 caudal-fin rays; 104-113 scales
in a longitudinal series; head length equal
to or nearly equal to head width in all but
the largest specimens; a well-developed
pupillary operculum; a distinct series of
rectangular black blotches on the posterior
three-fourths of the dorsal fin and along the
entire length of the anal fin; an unpigmented
peritoneum; a prominent black blotch across
the lower half of the operculum; and 9-11
faint crossbars on the head and body begin-

Symphurus callopterus, n.sp., holotype, CAS 64023, male 116.9 mm SL; Peru.

ning from just behind the eyes and ending
just anterior to the base of the caudal fin.

Description. —Frequency distributions of
meristic data are given in Table 1. ID pat-
tern typically 1-3-4 (123 of 191 specimens),
less frequently 1-4-3 (23/191), rarely 1-3-
3-3 or 1-4-4. Caudal-fin rays 12 (160 of 165),
rarely 11 or 13. Dorsal-fin rays 105-114,
usually 109-113, xX = 110.8. Anal-fin rays
91-98, usually 93-97, x = 94.7. Pelvic-fin
rays 4. Total vertebrae 57-61, usually 58-
60, rarely 57 or 61, X = 58.9; abdominal
vertebrae 9 (3+6). Hypurals 4. Longitudi-
nal scale rows 97-114, usually 100-111.
Scale rows on head posterior to lower orbit
20-26, usually 22-24. Transverse scales 35—
45, usually 37-41.

Body relatively deep, proportionately
deeper in larger specimens, 233-296 SL, x
= 248; with greatest depth in anterior third
of body; body width gradually tapering pos-
teriorly giving body lanceolate shape.
Preanal length 216-265 SL, x = 236; slightly
less than body depth. Head relatively wide
(216-278 SL, x = 237), nearly equal to body
depth in specimens smaller than 150 mm
SL, proportionately smaller than body depth
in specimens larger than 150 mm SL. Head
length nearly equal to head width (218-261
SL, x = 236). Postorbital length 146-189,
X = 163. Lower head lobe (117-158 SL, x
= 139) slightly larger than upper head lobe
(101-136 SL, x = 118). Snout relatively
short, 171-203 HL, xX = 185; covered with
small ctenoid scales. Dermal papillae well-
developed on blind side of snout. Anterior
nostril relatively short, not reaching ante-
rior margin of eye when extended poste-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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VOLUME 102, NUMBER 2

riorly. Mouth large, 222-271 HL, x = 245;
posterior extension of maxilla reaches usu-
ally to rear margin of lower eye, less fre-
quently extending posteriorly only to rear
margin of pupil of lower eye. Chin depth
179-272 HL, X = 207; slightly larger than
snout length. Lower eye relatively small, 79-
118 HL, x = 103; eyes usually slightly sub-
equal in position with upper slightly in ad-
vance of lower eye. Anterior and medial
surfaces of eyes partially covered with 3-5
rows of small ctenoid scales; 1—3 small cte-
noid scales in narrow interorbital region.
Pupillary operculum triangular and well-de-
veloped. Length of dorsal-fin base 941-955
SL, x = 948. Dorsal-fin origin far forward;
dorsal fin usually reaches anteriorly to a
point directly above anterior margin of mi-
grating eye, less frequently to anterior mar-
gin of pupil and in a few specimens, only
to middle of migrating eye; predorsal length
46-59 SL, x = 52. Length of anal-fin base
736-821 SL, X = 760. Blind sides of dorsal
and anal fins without scales; 3-5 rows of
small scales present on ocular-side bases of
dorsal- and anal-fin rays. Pelvic fin with 4
rays; relatively short, 38-63 SL, x = 55;
longest pelvic-fin ray reaches to first anal-
fin ray. Caudal fin relatively short, 75-94
SL, x = 83.

Teeth well-developed on blind side jaws.
Teeth on ocular side of premaxilla slender,
in single row, usually covering only anterior
half or three-fourths of jaw; extending pos-
teriorly to point directly below base of an-
terior nostril. Teeth on ocular side dentary
in single row, usually only extending along
anterior three-fourths of dentary, some-
times along entire dentary, or only anterior
half.

Pigmentation. —Specimens in alcohol
with a chocolate brown ground color with
series of 9-11 faint crossbars of variable
width (usually 4—8 scale rows wide) from
behind nape to base of caudal fin (holotype
with 12 bars); and 10-19 well-defined, rect-
angular, or sometimes slightly oval, black
blotches alternating with very light areas on

463

median fins. Specimens from southern
Mexico, Costa Rica and Panama generally
darker than those from other locations. Blind
side typically uniformly unpigmented,
probably creamy white in life. Some indi-
viduals of both sexes also have darkly pig-
mented patches concentrated in caudal re-
gion and sometimes scattered at various
places along bases of dorsal- and anal-fins
along posterior half of blind side of body.

Head usually with two faint crossbars.
Anterior bar about 3-4 scales in width,
barely reaching ventral portion of head, and
located posteriorly (about 4—5 scales width)
to eyes. Well-defined posterior bar about 6-
8 scales wide, crossing opercular region, and
extending ventrally slightly onto blind side
of head; sometimes somewhat faint and dif-
fuse dorsally. Inner lining of eyed-side oper-
culum only lightly pigmented, most often
only sprinkled with melanophores; blind-
side opercular lining usually unpigmented.
Isthmus mostly unpigmented except for
small patch of melanophores on dorsal sur-
face.

Posterior two-thirds of dorsal fin and en-
tire length of anal fin bear clearly-defined
blotches, not extending to either tips or bas-
es of rays. Blotches on both fins start ap-
proximately equidistant from snout; ante-
riormost blotches typically lighter than
posterior ones; each blotch usually covers
about 5-7 rays. Lighter areas between
blotches cover about 3—5 rays and are stip-
pled with tiny melanophores producing a
faint duskiness barely discernible to naked
eye. Median fins on blind side without me-
lanophores. Number of black blotches on
dorsal fin varies between 10-19. Anterior
to point above anal-fin origin, dorsal fin
without distinct blotches. Anal-fin blotches
usually parallel in position to dorsal-fin
blotches and with similar intensity of pig-
mentation; usually between 12-15 (one
specimen had 8, and another 19 blotches).
Holotype with 14 blotches on dorsal and 11
on anal fin. Most posterior portions of me-
dian fins heavily pigmented in melanistic

464

Fig. 2. Geographic distribution of Symphurus cal-
lopterus based on available study material. Dots in-
dicating collection localities may represent more than
one occurrence and more than a single specimen from
each locality.

specimens, obscuring blotches to naked eye
(but still perceptible under magnification).

Caudal fin generally with two ill-defined
blotches, one on the base and the other at
distal tip, with lighter space between. Cau-
dal-fin blotches may be faded and diffuse
giving entire caudal uniformly dark color-
ation. In some specimens of both sexes,
blind side of caudal fin darkly pigmented.

Size and sexual maturity. —Sexual ma-
turity occurs at a large size relative to other
Symphurus species. Of 53 females, 52, rang-
ing in size from 90.8—162 mm SL, were ma-
ture with elongate, gravid ovaries. The
smallest of these measured 90.8, 93.0, 97.3
and 97.9 mm SL. The only immature fe-
male examined was slightly smaller (87.3
mm SL) with ovaries just undergoing pos-
terior elongation. The 66 males were only
slightly smaller than the females (76.5-151
mm SL).

Etymology. — Callopterus meaning ““pret-
ty fin” (Greek kallos, beauty; and pterygion,
fin, a diminutive form of pteron, wing); in
reference to the striking black blotches on
the median fins in an otherwise mundanely
pigmented group of fishes.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Distribution.—Throughout the tropical
eastern Pacific (Fig. 2) from Mazatlan and
southern Baja California, Mexico, to north-
ern Peru (10°36’S).

Symphurus callopterus occurs primarily
in moderate depths (18-317 m; SIO 63-522
and UF 47590, respectively) throughout its
geographic range. The bathymetric center of
abundance, based on both frequency of cap-
ture and numerical abundance, occurs on
the continental shelf between 60-110 m.
Over 85% (244 of 285) of the specimens
studied were collected at these depths. Cap-
ture of this species at depths less than 60 m
is unusual; only five specimens were col-
lected between 40-60 m and only three
specimens were taken shallower than 40 m
(18 and 20 m). Symphurus callopterus has
been collected rarely at depths greater than
110 m; only 10 of 285 specimens (4%) were
collected deeper than 110 m with the deep-
est occurrences reported for single speci-
mens being 146, 196, 206 and 317 m.

Geographical variation.—No latitudinal
variation in meristic characters was ob-
served in specimens collected between the
northern and southern parts of the geo-
graphic range.

Remarks.—Two haemal and/or neural
spines occurred on the first preural (PU 1)
or the second preural centra (PU 2) in 27
of 180 (15%) specimens examined. Two hy-
potheses may explain the high frequency of
doubling of spines in these centra: 1) PU 1
or PU 2 frequently possess double neural
and/or haemal spines, or 2) double spines
result from fusion of two caudal vertebrae.
The data support the latter hypothesis. In
some specimens with two spines on PU 1,
that centrum was slightly longer than nor-
mal, suggesting that it might be the result
of fusion of two centra. Additionally, when
two-spined preural centra were counted as
a single vertebra (Table 2), specimens with
these centra had, as a group, one fewer ver-
tebra than the normal specimens (x = 58.0
for double-spined specimens; x = 58.9 for

VOLUME 102, NUMBER 2

465

Table 2.—Number of vertebrae in Symphurus callopterus. T-test comparing specimens possessing double
neural and/or haemal spines on the PU 1 or PU 2 to those with single spines.

Number of vertebrae

PU 1/PU 2 spines 57 58 59 60 61 n x SD t jz
Double (counted as one centrum) = 19 4 O7.3.0 Zi 53:0. 90:56 6.30 <0.000
Single (normal condition) SS Oe, Da 1 RSS) 758.910.7971
Double (counted as two centra) 0 rms TS 4 0 2h ODO, SOS6i F065 >0.90
Single (normal condition) ERIE S lies: 27 | 1 1532 S829 > 0.71

normal specimens; t = 6.30; P < Q.000).
However, when two-spined centra were
counted as two vertebrae, there were no sig-
nificant differences in total vertebrae be-
tween the two groups (x = 59.0 for double-
spined group versus 58.9 for “normal”
group; t = 0.63, P > 0.90). Because the data
support the hypothesis that double spines
are the result of vertebral fusion, vertebrae
bearing them were counted as two.
Co-occurring Symphurus species. —In the
shallower portions of its bathymetric range
along the coast of Mexico and Central
America (at least as far south as Panama),
S. callopterus has been collected with S. gor-
gonae Chabanaud and S. atramentatus Jor-
dan & Bollman; at greater depths in this
region, S. callopterus is partially syntopic
with an undescribed species.
Comparisons.— Other tonguefishes with
counts comparable to those of S. callopterus
are S. maldivensis Chabanaud, S. marmor-
atus Fowler and S. strictus Gilbert from the
Indo-West Pacific including Hawaii, S. mi-
crolepis Garman from the eastern Pacific,
S. ligulatus (Cocco) from the eastern Atlan-
tic, and S. jenynsi Evermann & Kendall and
S. nebulosus (Goode & Bean) from the west-
ern Atlantic. Symphurus callopterus differs
from S. microlepis principally in ID pattern
(1-3-4 versus 1-3-2) and pigmentation
(peritoneum unpigmented and dorsal and
anal fins with blotches in S. callopterus ver-
sus dark black peritoneum and unpigment-
ed fins in S. microlepis). Symphurus callop-
terus differs from the other species in ID

pattern and number of caudal-fin rays (1-3-
4, 12 versus 1-2-2 and 14, respectively, in
S. strictus, S. marmoratus, S. ligulatus and
S. nebulosus; 1-4-3 and 10 in S. jenynsi;
1-2-2 and unknown number of caudal-fin
rays in S. maldivensis), as well as in features
of its pigmentation (peritoneum unpig-
mented in S. callopterus versus darkly pig-
mented in all of the above species, except
S. jenynsi; median fins highly ornate with
alternating series of blotches and unpig-
mented areas in S. callopterus versus uni-
formly pigmented fins in the other species).

Other species of Symphurus with similar
pigmentation on the median fins are S. atra-
mentatus Jordan & Bollman, and an un-
described species from the eastern Pacific,
as well as S. diomedeanus (Goode & Bean)
from the western Atlantic. Symphurus cal-
lopterus differs from them in its extremely
high number of dorsal- and anal-fin rays and
vertebrae and in the larger number of
blotches on the fins. In addition, in S. cal-
lopterus, head length is nearly equal to head
width in all except the largest adults, where-
as in the other species, head length is usually
less than head width. Throughout its range,
S. callopterus is occasionally collected with
an undescribed species that has a similar
body shape and pigmentation pattern. Sym-
phurus callopterus differs from this unde-
scribed species in several major features in-
cluding ID pattern (1-3-4 versus 1-3-2),
hypural number (4 versus 5), presence of a
well-developed pupillary operculum versus
its absence, pigmentation of the peritoneum

466

(unpigmented versus dark black showing
through the abdominal wall), and modal dif-
ferences in meristic characters.

Material Examined

Counts were taken from the following 138
non-type specimens (71.0-162 mm SL):
Mexico.—UCLA W61-198, 2(116.0-
131.8), Sinaloa, 3—4 miles southwest of Ma-
zatlan (ca. 23.5°N, TOGSSO0AW)2 29 ct
1961.—CAS 4785-4786, 2(93.0-101.9),
Michoacan, South of Punta San Telmo off
Michoacan coast (ca. 18.3°N, 103°40’W), 19
Jul 1932.—UCLA W56-80, (104.1), Oa-
xaca, off Port Angeles Light (ca. 15.5°N,
96°40’W), 14 Jul 1932. Guatemala: —SIO
73-265, 14(71.0—90.0), 13°55.6-53.5'N,
92°02.5'W, 101 m, 13 Apr 1973. Nicara-
gua.—SIO 73-279, 3(107.4—116.1), 12°09-
11.5’N, 87°32-35.5'W, 113 m, 17 Apr 1973.
Costa Rica.—CAS 5304, (94.2), off Cabo
Velas (ca. 10°22'"N, 85°53'W), FOL m. f Jul
1932.—CAS 11710, (119.3), 6 mi off Cabo
Velas (ca. 10°22’N, $5°53'W), 1LOb met jul
1932.—LACM 33590-2, 3(105.0—130.0), 9
mi from Isla Herraderra, 17 May 1973.—
LACM 33827-11, (124.4), Gulf of Nicoya,
central Puntarenas (ca. 9.5°N, 85°W), 29 Jun
1973.—SIO 79-5, 42(76.6—-116.8), Gulf of
Papagayo, 10°42.4’N, 85°55’W, 88 m, 02
Apr 1978. Panama Bay.—UMML 26003,
11(76.5-117.6), 7°49.3-49.8'’N, 80°00-
00.7'W, 46 m, 2 May 1967.—UMML
34318, 30(85.8-133.8), 7°40.3’N, 79°
50.9'W, 78 m, 2 May 1967. Colombia. —
UMML 34319, (124.5), 4°00’N, 77°30'W,
92 m, 17 Sep 1961. Ecuador. —CAS 24195,
3(140.6—-162), 2°29’'S, 80°59'W, 93 m, 11
Sep 1966.—CAS 24198, (131.2), 3°35’S,
78°35'W, 79 m, 16 Sep 1966. Peru.—CAS
24188, 10(114.9-140.1), 5°59’S, 81°12’W,
100 m, 4 Jun 1966.—CAS 24190, (112.2),
6°20’S, 81°01’ W, 146 m, 4 Jun 1966.—CAS
2AITST SCiS1 35-1513). 62S. st 36 Ww.
100 m, 5 Jun 1966.—CAS 24957, (129.2),
6°21’S, 80°59’W, 140 m, 4 Jun 1966.—CAS
24192, 2(122.2—139.2), 6°24’S, 80°44’W, 20

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

m, 5 Jun 1966.—CAS 24194, 2(111.3-
132.6), 4°55’S, 81°19'W, 70 m, 8 Sep 1966.—
UCLA W58-395, 3(106.5—123.5), 10°36’S,
77°54'W, 16 Oct 1958.

Additional material examined but not
counted or measured (97 specimens; 52.2—
143.1 mm SL): Mexico.—SIO 64-878,
2(55.9-58.7), Baja California, 24°18.9-
19.0'N, 111°47.6-48.6'W, 78 m, 13 Nov
1964.—SIO 62-75, (52.2), Sinaloa, 22°15’N,
106°03’W, 55 m, 25 Aug 1961.—SIO 62-
76, 23 (73.1-116.0), 22°13’N, 106°10’W, 79
m, 25 Aug 1961.—SIO 73-230, (89.9),
21°59'N, 106°20’W, 102 m, 30 Mar 1973.—
SIO 73-232, 8(82.8-103.6), 21°57—-S6’N,
106°10-11'W, 52 m, 31 Mar 1973. Mexico,
Gulf of Tehuantepec. —SIO 63-504, (113.0),
15°57.5'N, 95°00’W, 61 m; 8 Jul 1963.—
SIO 65-165, (62.0), 15°55’N, 95°24’W, 102
m, 6-7 Jun 1965.—SIO 63-525, (104.2),
15°52’N, 95°43’W, 55 m, 15 Jul 1963.—SIO
63-522, (89.5), 15°44’N, 96°06.5’W, 18 m,
15 Jul 1963.—SIO 63-523, 2(70.5-89.1),
15°41'N, 96°07.5'W, 55 m, 15 Jul 1963.—
SIO 63-513, (96.5), 15°07.5'N, 93°25’W, 102
m, 10 Jul 1963. Guatemala.—SIO 73-265,
4(81.0-92.0), 13°55.6-53.5'N, 92°02.5'W,
100 m, 13 Apr 1973. Costa Rica.—CAS
20965, (82.5), 6 mi off Cabo Velas (ca.
10°22'N, 85°53'W),; 101 m,; Igelies2
CAS 43880, (114.5), Cabo Blanco (ca. 9.5°N,
85°W), 206 m, 9 Mar 1974.—CAS 44140,
(111.6), Gulf of Nicoya (ca. 9.5°N, 85°W),
83 m, 14 Mar 1974.—SIO 73-281, (103.1),
10°50.2—53.2'N, 86°20.0—24.3’W, 196 m, 18
Apr 1973.—SIO 73-296, 26(83-126.5),
9°37.4—37.7'N, 84°49.0-51.8'W, 87 m, 22
Apr 1973. Gulf of Panama.—CAS 24958,
2(115.5-117.2), 8°11'N, 79°36'W, 88 m, 23
Jul 1967.—CAS 24959, 3(127.5-143.1),
6°34'’N, 77°21'20”W, 60 m, 21 Apr 1967.
Colombia.—CAS 24960, 6(121.5-132.0),
6°44'N, 77°33'30’W, 82 m, 10 Aug 1967.

Acknowledgments

We thank the following people who as-
sisted with this study: R. Gibbons (photog-

VOLUME 102, NUMBER 2

raphy and map preparation) and M. Nizin-
ski (radiographs and technical support); the
following curators and support staff who
generously provided space and arranged for
loan of specimens: S. Jewett (USNM); A.
Wheeler (BMNH); R. Rosenblatt, D. Gib-
bons, H. J. Walker (SIO); R. Lavenberg, J.
Seigel, R. Feeney (LACM); C. Gilbert, G.
Burgess (FSU); C. R. Robins (UMML); W.
Eschmeyer, M. E. Anderson, D. Catania
(CAS); and D. Buth (UCLA). The manu-
script was much improved through critical
reviews by B. B. Collette, G. D. Johnson,
N. Menezes, R. Rosenblatt and R. P. Vari.
Portions of this study were initiated while
the senior author was a postdoctoral fellow
in the Division of Fishes at the USNM.
Travel funds enabling Munroe to work at
UMML in 1982 were provided by the De-
partment of Fisheries, Virginia Institute of
Marine Science.

A special thanks to W. Bussing, Univer-
sity of Costa Rica, for allowing us to use
information contained in his unpublished
manuscript on the tonguefishes of Costa
Rica.

Literature Cited

Bartels, C. E., K. S. Price, M. I. Lopez, & W. A. Bussing.
1983. Occurrence, distribution, abundance and
diversity of fishes in the Gulf of Nicoya, Costa
Rica.— Revista De Biologia Tropical 31(1):75-
101.

—_. : , & 1984. Ecological
assessment of finfish as indicators of habitats in
the Gulf of Nicoya, Costa Rica.— Hydrobiologia
112:197-207.

Chirichigno, F. N. 1974. Clave para identificar los
peces marinos del Peru. —Informes Instituto del
Mar del Pert: 44: 1-387.

467

Eschmeyer, W. N., E.S. Herald, & H. Hammann. 1983.
A field guide to Pacific coast fishes from the Gulf
of Alaska to Baja California. Boston, Massa-
chusetts, Houghton Mifflin Company, 336 pp.

Ginsburg, I. 1951. Western Atlantic tonguefishes with
descriptions of six new species.— Zoologica,
N.Y., 36(3):185-201.

Hildebrand, S. F. 1946. A descriptive catalog of the
shore fishes of Peru.— United States National
Museum Bulletin 189:1-—530.

Jordan, D. S., & B. W. Evermann. 1898. The fishes
of North and Middle America.— United States
National Museum Bulletin 47(3):2183-3136.

Love, M.S., G. E. McGowen, W. Westphal, R. J. Lav-
enberg, & L. Martin. 1984. Aspects of the life
history and fishery of the white croaker, Geny-
onemus lineatus (Sciaenidae), off California. —
Fishery Bulletin, U.S. 82(1):179-198.

Mahadeva, N. 1956. A review of the tonguefishes of
the eastern Pacific, with descriptions of six new
species. Unpublished dissertation, University of
California at Los Angeles. 272 pp.

Meek, S. E., & S. F. Hildebrand. 1928. The marine
fishes of Panama.—Field Museum of Natural
History, Publication 249, Zoological Series Vol-
ume 15 (Part 3):709-1045.

Menezes, N. A., & G. de Q. Benvegnu. 1976. On the
species of the genus Symphurus from the Bra-
zilian coast, with descriptions of two new species
(Osteichthyes, Pleuronectiformes, Cynoglossi-
dae).—Papeis Avulsos de Zoologia, Sao Paulo
30(11):137-170.

Munroe, T. A. 1987. A systematic revision of Atlan-
tic tonguefishes (Symphurus: Cynoglossidae:
Pleuronectiformes) with a preliminary hypoth-
esis of species group relationships. Unpublished
dissertation, College of William and Mary, Wil-
liamsburg, Virginia. 598 pp.

(TAM) National Marine Fisheries Ser-
vice, Systematics Laboratory, National Mu-
seum of Natural History, Washington, D.C.
20560; (MNM) Department of Biology,
Wisconsin State University, Oshkosh, Wis-
consin 54901.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 468-482

SYSTEMATICS OF THE STEINDACHNERINA HYPOSTOMA

COMPLEX (PISCES, OSTARIPHYSI, CURIMATIDAE), WITH THE

DESCRIPTION OF THREE NEW SPECIES
Richard P. Vari and Ann Williams Vari

Abstract. — The Steindachnerina hyposoma complex is revised and four species
are recognized: S. hypostoma (Boulenger) widely distributed through the Rio
Amazonas system upstream of Manaus; S. planiventris, new species, from the
Rio Negro and Rio Madeira basins, and various localities along the Rio Ama-
zonas upstream of Manaus; S. gracilis, new species, apparently endemic to the
Rio Tocantins system of eastern Brazil; and S. quasimodoi, new species, of the
western portions of the Amazon basin in Brazil and Peru. The four species
have in common derived expansions of the mesethmoid and frontals that result
in a reduction in the size of the cranial fontanel, and unite them as a mono-
phyletic assemblage. A key to the species of the complex is provided.

Resumo.—O complexo Steindachnerina hypostoma € revisto e quatro es-
pécies reconhecidas: S. hypostoma (Boulenger) amplamente distribuida no sis-
tema do Rio Amazonas, acima de Manaus; S. planiventris, espécie nova, das
bacias do Rio Negro e do Rio Madeira e varias localidades ao longo do Rio
Solimoes acima de Manaus; S. gracilis, espécie nova, aparentemente endémica
da bacia do Rio Tocantins no leste do Brasil; e S. guasimodoi, espécie nova,
da parte oeste da Bacia AmazoOnica, no Brasil e Peru. As quatro espécies tém
em commum expansoes derivadas do mesetmoide e frontais que causam a
reducao em tamanho da fontanela craniana e unem estas quatro espécies em
um grupo monofilético. Uma chave para as espécies do grupo é apresentada.

Boulenger (1887:172) described Curi-
matus hypostoma on the basis of four spec-
imens collected by “Mr. W. Davis’ in the
‘““Ucayali River’? of Peru. At the end of his
brief description, he noted the very shallow
body depth characteristic of the species. That
feature is indeed striking when one com-
pares this species to other curimatids, most
of which have moderately to very deep bod-
les (see figures in Vari, 1982, 1984, in press
a, b, c). Although this unusually shaped
species was subsequently referred to by Ei-
genmann & Eigenmann (1891), Eigenmann
(1910), Fowler (1942, 1950, 1975), and Fer-
nandez-Yépez (1948), among others, those
citations were based on the original descrip-
tion without examination of other speci-

mens. The only subsequent locality record
of Curimatus hypostoma based on addi-
tional material was that of Eigenmann and
Allen (1942:296) who reported on a series
of specimens from various localities in the
Amazon basin in Peru. Most recently Vari
(1989) assigned Curimatus hypostoma to the
speciose genus Steindachnerina Fowler. The
limited number of references in the litera-
ture to Steindachnerina hypostoma are rath-
er surprising given that the species is rela-
tively common in many of the major river
systems of the Amazon basin, and has a
geographic distribution much wider than re-
ported in the literature.

Collecting efforts in recent years through-
out the Amazon basin have yielded large

VOLUME 102, NUMBER 2

series of specimens typically identified as
Curimatus hypostoma. The examination of
these collections undertaken as part of more
encompassing studies within the Curima-
tidae has revealed that this material actually
encompasses four distinct, albeit similar,
species that form a monophyletic subunit
of the Curimatidae. The known distribution
of the species of the clade encompasses both
the main Rio Amazonas basin and the as-
sociated Rio Tocantins system of eastern
Brazil. The shared derived features of these
species are discussed, and a key to, and de-
scriptions of the species are provided.

Materials and methods. —Counts and
measurements were taken using the meth-
ods outlined in Vari (1982, 1984, in press
a, b, c). In the case of new species the pre-
sented ranges of counts and measurements
include values of all type specimens. The
ranges of these values for Steindachnerina
hypostoma include all specimens cited for
which standard lengths (SL) are given. Val-
ues in square brackets in species descrip-
tions are those of the holotype or lectotype.
Information on localities and specimens ex-
amined in detail is arranged as in Vari (1982,
1984, in press a, b, c). Measurements are
presented as proportions of standard length
(SL) or head length (HL).

The following abbreviations for institu-
tions are used: American Museum of Nat-
ural History, New York (AMNH); British
Museum (Natural History), London
(BMNH); California Academy of Sciences,
San Francisco (CAS); Stanford University,
now deposited at CAS (CAS-SU); Jacques
Gery, personal collection (GC); Indiana
University, collections now at various re-
positories (IU); Los Angeles County Mu-
seum of Natural History (LACM); Museum
of Comparative Zoology, Cambridge (MCZ);
Museu de Zoologia da Universidade de Sao
Paulo (MZUSP); Naturhistoriska Riksmu-
seet, Stockholm (NRM); Naturhistorisches
Museum, Vienna (NMW); Museo de His-
toria Natural de la Universidad Nacional
Mayor de San Marcos, Lima (MHN-USM);

469

and National Museum of Natural History,
Smithsonian Institution, Washington, D.C.
(USNM).

Steindachnerina hypostoma complex

Vari (1989) redefined Steindachnerina
Fowler (1906) to include 32 nominal species
or subspecies. Vari and Castro (1988) sub-
sequently noted that Prochilodus stigma-
turus Fowler is also a curimatid of the genus
Steindachnerina rather than a prochilodon-
tid. Steindachnerina has a wide distribution
in lowland tropical South America on both
sides of the Andean Cordilleras, but is by
far most speciose in the Amazon basin, the
region to which the Steindachnerina hypos-
toma complex is endemic. The members of
Steindachnerina share a series of synapo-
morphous features of the first and second
infraphyarnygobranchials, the mode of at-
tachment of the ligament joining the second
and third hypobranchials, and the form of
the basihyal and its associated tooth-plate
(see Vari [1989] synapomorphies 97 to 100).
The four species of the Steindachnerina hy-
postoma complex are, in turn, characterized
by derived modifications of the meseth-
moid and frontals of lesser universality that
are reflected in the relative size and form of
the cranial fontanel.

Species of Steindachnerina outside of the
hypostoma complex and curimatids in gen-
eral have relatively extensive dorsomedian
cranial fontanels extending from the supra-
occipital to the mesethmoid (e.g., S. bi-
maculata, Fig. 1A). In those outgroup taxa
the anterior two-thirds of the relatively large
opening is delimited laterally by widely sep-
arated frontals. The medial margins of the
frontals anterior to the epiphyseal bar are
nearly parallel or converge very gradually
anteriorly. Anteriorly the fontanel is bor-
dered by a distinct notch situated in the
middle of the posterior border of the mes-
ethmoid. This arrangement of bones along
an extensive fontanel is hypothesized as
primitive within Steindachnerina given its

470

D

Fig. 1. Dorsal view of anterior portion of cranial
fontanel and proximate bones of A) Steindachnerina
bimaculata, USNM 261450, 76.4 mm SL; B) Stein-
dachnerina quasimodoi, USNM 293042, 76.0 mm SL;
C) Steindachnerina planiventris, USNM 267987, 62.1
mm SL; D) Steindachnerina gracilis, USNM 293034,
63.4 mm SL; and E) Steindachnerina hypostoma, CAS
63192 (formerly IU 15825), 82.00 mm SL. Abbrevi-
ations: EB—epiphyseal bar; FR—frontal; ME—mes-
ethmoid; P—Parietals. Mesethmoid and parietals trun-
cated, horizontal lines indicate cranial fontanel.

general distribution in that genus and other
curimatids.

The species of the S. hypostoma complex
demonstrate marked changes in the mor-
phology of those portions of the meseth-
moid and frontals bordering the cranial fon-
tanel. All four species lack the extension of
the fontanel into the posterior portion of
the mesethmoid. The posteromedial region
of the mesethmoid is instead developed
posteriorly into a triangular process (Fig. 1B
to E). More striking is the expansion in all
four species of the anteromedial portions of
the frontals and the associated pronounced
reduction of the portion of the cranial fon-
tanel anterior of the epiphyseal bar. As a
consequence of this expansion the antero-
medial margins of the frontals no longer
extend roughly in parallel. The margins of
the bones instead converge and are either
separated by a small gap (Steindachnerina
quasimodoi, Fig. 1B), or are in contact me-
dially to varying degrees (S. planiventris, S.
gracilis, S. hypostoma, Fig. 1C-E).

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Both the extension of the posteromedial
portion of the mesethmoid and the medial
expansion of the frontals in the four species
of the Steindachnerina hypostoma complex
are derived relative to the conditions of these
bones in outgroups. These synapomorphies
support a hypothesis that S. hypostoma, S.
gracilis, S. quasimodoi, and S. planiventris
are a monophyletic lineage within Stein-
dachnerina.

Key to the species of the
Steindachnerina hypostoma complex

1. Dorsal fin plain, without dark spot
at base of middle rays; snout length
0.28—0.31 of HL; lateral-line scales
to hypural joint 46 to 50 ........
S. hypostoma (Boulenger)
— Dorsal fin with dark spot at base of
middle rays, spot sometimes faint;
snout length 0.31—0.34 of HL; lat-
eral line scales to hypural joint 40
to 46 or 50 to 54
2. Lateral-line scales to hypural joint
SOHOmas . ae. S. gracilis, new species
— Lateral line scales to hypural joint
40 to 46
3. Greatest body depth in specimens
over 50 mm SL 0.30-0.34 of SL,
body with distinctly humped dorsal
profile (Figs. 6, 7); frontals separated
by slight gap anteriorly (Fig. 1B)
esheets S. quasimodoi, new species
— Greatest body depth in specimens
over 50 mm SL 0.27-0.34 of SL,
dorsal profile of body without dis-
tinct hump (Fig. 11); frontals in con-
tact anteriorly (Fig. 1\C) eee
a Gikas Pass S. planiventris, new species

Steindachnerina hypostoma
(Boulenger, 1887)
Figs. 1-4, Table 1

Curimatus hypostoma Boulenger, 1887:172
[type locality: Ucayali River ].— Vari,
1989:tabs. 2, 3 [phylogenetic relation-
ships].

VOLUME 102, NUMBER 2

7
> 7?

a8
as a es
~~ = ~ s
: +o>04
=a ss
a 4
= .

"
E>.

*Sess ~3RBar
>

a Pf = Li

bee 53.3
DEVILS

471

a* 8480995 > %:

—

Fig. 2. Steindachnerina hypostoma, USNM 229190, 93.2 mm SL; Brazil, Amazonas, Rio Solimoes, Ilha

Machantaria.

Curimatus hypostomus, Eigenmann & Ei-
genmann, 1889:426 [reference].—Eigen-
mann & Eigenmann, 1891:47 [refer-
ence].—Eigenmann, 1910:422 [reference].

Curimata hypostoma, Fowler, 1942:208
[reference].— Fowler, 1945:116 [refer-
ence].—Eigenmann & Allen, 1942:296
[Peru: Rio Huallaga, Yurimaguas; Rio
Morona; Rio Paranapura, Yurimaguas (in
part)].— Fowler, 1945:116 [reference].—
Géry, 1977:235 [reference].—Ortega &
Vari, 1986:11 [Peru, Amazon basin; com-
mon name].

Curimata hypostoma hypostoma, Fowler,
1950:285 [literature compilation].

Steindachnerina hypostomus, Fernandez-
Yépez, 1948:58 [assignment to Stein-
dachnerina].

Steindachnerina hypostoma, Fowler, 1975:
375 [reference].

Diagnosis. —The absence of a discrete
patch of dark pigmentation at the base of
the middle rays of the dorsal fin distinguish-
es S. hypostoma from the other members
of the complex, all of which have such pig-
mentation developed to some degree. The
46 to 50 lateral line scales of S. hypostoma
further differentiate the species from S. gra-
cilis which has 50 to 54 scales in that series
and from S. guasimodoi and S. planiventris
which have 40 to 46. Differences in other
meristic and morphometric features also
discriminate S. hypostoma from the re-
maining members of the complex (Table 1).

Description.—Body elongate, relatively
wide, particularly in ripe females. Dorsal
profile of head straight or very slightly con-
vex. Dorsal profile of body smoothly curved,
slightly convex from rear of head to origin
of dorsal fin; straighter and slightly postero-

Fig. 3. Steindachnerina hypostoma, USNM 278579, 48.9 mm SL; Bolivia, Pando, Rio Madre de Dios,

Laguna San Luis.

472

Fig. 4. Map of northern South America showing
geographic distribution of Steindachnerina hypostoma
(dots) and Steindachnerina gracilis (stars). Type local-
ities: 1, Rio Aripuana, Aruana; 2, Rio Tocantins, Tu-
curui; and 3, Rio Ucayali (exact collection site of syn-
types of Curimatus hypostoma unknown). Some
symbols represent more than one lot and/or collecting
locality.

ventrally slanted at base of dorsal fin, straight
or very gently convex from base of last dor-
sal-fin ray to caudal peduncle. Dorsal sur-
face of body with distinct median keel an-
terior to dorsal fin, keel more pronounced
proximate to fin; body surface smoothly
rounded transversely posterior to fin. Ven-
tral profile of body very gently curved from
tip of lower jaw to caudal peduncle. Pre-
pelvic region wide, distinctly flattened, with
five series of scales in transverse series across
flattened surface. Pelvic fins distinctly sep-
arated medially. Post-pelvic region some-
what flattened proximate to pelvic fin, grad-
ually becoming transversely rounded
posteriorly.

Greatest body depth at origin of dorsal
fin, depth 0.24—0.27 [0.25]; snout tip to or-
igin of dorsal fin 0.44—0.48 [0.45]; snout tip
to origin of anal fin 0.80-0.85 [0.82]; snout
tip to insertion of pelvic fin 0.48-0.53 [0.48];
snout tip to anus 0.70-0.75 [0.72]; origin of
dorsal fin to hypural joint 0.56—0.60 [0.59].
Dorsal-fin margin pointed; anteriormost
rays 2.8-3.5 times length of ultimate ray.
Pectoral-fin margin pointed; length of pec-
toral fin 0.17-0.20 [0.18], extends slightly

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

over three-quarters of distance to vertical
through insertion of pelvic fin. Pelvic-fin
margin obtusely pointed, length of pelvic
fin 0.19-0.22 [0.20], reaches about three-
fifths of distance to origin of anal fin. Caudal
fin distinctly forked. Adipose dorsal fin well
developed. Anal fin emarginate, anterior-
most branched rays approximately two and
one-half times length of ultimate ray. Cau-
dal peduncle depth 0.11-—0.12 [0.11].

Head distinctly pointed in profile, more
so in larger specimens, head length 0.24—
0.28 [0.27]; upper jaw distinctly longer than
lower, mouth inferior; snout length 0.28-
0.31 [0.28]; nostrils very close, anterior cir-
cular, posterior crescent-shaped with aper-
ture closed by thin flap of skin separating
nares; orbital diameter 0.28-0.34 [0.29];
adipose eyelid present, poorly developed,
with rotund opening over center of eye;
length of postorbital portion of head 0.39-
0.45 [0.43]; gape width 0.28—0.34; interor-
bital width 0.38—-0.42 [0.39].

Pored lateral line scales from supraclei-
thrum to hypural joint 46 to 50 [49]; all
scales of lateral line pored, canals in scales
straight; 4 to 6 series of scales extend beyond
hypural joint onto caudal fin base; 8'2 to 914
[814] scales in transverse series from origin
of dorsal fin to lateral line; 642 to 7% [7]
scales in transverse series from lateral line
to origin of anal fin.

Dorsal-fin rays 11,9 or 111,9 (111,9 rare, when
three unbranched rays present, first very
short) [11,9]; anal-fin rays 1,7 or i11,7 (when
three unbranched rays present, first very
short) [11,7]; pectoral-fin rays 14 to 16 [14];
pelvic-fin rays 1,8 [1,8].

Total vertebrae 35 (8), 36 (86), 37 (2).

Color in alcohol. —Overall coloration of
specimens retaining guanine on scales sil-
very to silvery-golden, darker on dorsal por-
tions of head and body. Specimens fixed in
formalin and lacking guanine on scales with
overall ground coloration tan to light brown,
darker on dorsal portions of head and body
(Fig. 2). Pores of lateral line scales outlined
by dark pigmentation in some specimens.

VOLUME 102, NUMBER 2

473

Table 1.—Range of morphometrics and meristic features of species useful in discriminating species of Stein-
dachnerina hypostoma complex. Measurements | to 4 are proportions of standard length; 5 to 7 are proportions

of head length.

hypostoma gracilis quasimodoi planiventris
Morphometrics
1. Greatest body depth 0.24-0.27 0.25-0.27 0.27-0.34* 0.27-0.30
2. Snout to pelvic-fin origin 0.48-0.53 0.53-0.55 0.50-0.54 0.52-0.56
3. Snout to anus 0.70-0.75 0.73-0.75 0.75-0.76 0.72-0.76
4. Pelvic-fin length 0.19-0.22 0.19-0.21 0.23-0.26 0.19-0.23
5. Snout length 0.28-0.31 0.31-0.34 0.31-0.34 0.33-0.36
6. Orbital diameter 0.28-0.34 0.32-0.35 0.31-0.37 0.29-0.33
7. Postorbital length 0.39-0.45 0.39-0.42 0.36-0.40 0.36-0.41
Meristics

Lateral line scales 45-50 50-54 40-46 40-46
Scale rows between dorsal-fin

origin and lateral line 814-9 10-10% Th-9 614-8
Scale rows between anal-fin

origin and lateral line 612-72 6'h-7'h Sh-6"2 Sh-6%/

* 0.30-0.34 in specimens of 50 mm SL or greater.

Lateral line pigmentation more intense,
when present, in smaller individuals, some-
times forming distinct mid-lateral stripe
(Fig. 3). Deeper lying dusky mid-lateral band
extends from supracleithrum to caudal pe-
duncle; typically more obvious in smaller
individuals; masked in specimens retaining
guanine on scales. Caudal-fin rays outlined
by series of small chromatophores, lower
lobe of caudal fin dusky. Anterior margin
of dorsal fin sometimes dusky, but without
spot of dark pigmentation at base of middle
rays; other fins hyaline.

Distribution.—Amazon basin upstream
of Manaus, Brazil (Fig. 4).

Common name.—Chio—chio (Ortega &
Vari, 1986:11, Peru).

Remarks. —Boulenger (1887a) described
Curimatus hypostoma on the basis of four
specimens from an unspecified location
along the Rio Ucayali of Peru. An 87.5 mm
SL syntype (BMNH 1881.5.13:105), which
is in the best overall condition, is designated
as the lectotype. The remaining specimens
in the type series (BMNH 1881.5.13:106-
108) thus become paralectotypes.

Allen Gn Eigenmann & Allen 1942:297)
described a nominal subspecies Curimata

hypostoma hastata on the basis of a single
specimen from Puerto Bermudez, on the Rio
Pichis, Peru. A re-examination of the ho-
lotype (CAS 19881, formerly IU 17859) re-
veals that although it is a member of the
Steindachnerina clade, it is not conspecific
with S. hypostoma, and indeed is not even
a member of the S. hypostoma complex, but
rather a specimen of S. dobula (Gunther).
The specimens identified by Eigenmann &
Allen (1942:296) as Curimatus hypostoma
are an admixture of S. hypostoma and S.
dobula (e.g., IU 17858 from Yurimaguas,
Rio Paranapura). The complexity of that
series of specimens may account, at least in
part, for Allen’s assignment of hastata as a
subspecies of hypostoma.

Material examined. —286 specimens (70,
55.1—98.3).

PERU: Ucayali River, BMNH 1881.5.13:
105, 1 (87.5, lectotype of Curimatus hypo-
stoma); BMNH 1881.5.13:106-108, 3
(85.5—90.0, paralectotypes of Curimatus hy-
postoma). Loreto; Shansho Cano, USNM
175867, 1. Beirut, near mouth of Rio Am-
biyacu, USNM 261452, 2 (76.3-77.8). Rio
Javari, near Petropolis, USNM 261438, 1
(78.3). Rio Amazonas, Ramon Castilla,

474

USNM 261512, 1 (79.6). Rio Javari system,
Rio Galvez near mouth, NRM SOK/
1984314.4058, 3. Rio Huallaga, Yurima-
guas, USNM 293094, 1 (98.3); CAS 63191,
2 (84.0-86.8, formerly IU 17858); USNM
167802, 5 (83.0—-91.3); CAS 63192, 3 (83.8-
92.0; formerly IU 15825). Yurimaguas,
Rio Paranapura, IU 17858, 2 (part of
lot). Huanuco; Rio Pachitea, Tournavista,
USNM 293092, 1. Ucayali; Rio Ucayali,
Pucallpa, USNM 261513, 5 (80.1-91.2);
USNM 261489, 5 (65.8—96.5); USNM
261465, 6 (3, 55.3-59.8); USNM 261477,
5 (3, 70.4-83.4); AMNH 35686, 1 (70.0);
USNM 261493, 6; USNM 261496, 2;
USNM 261488, 6; USNM 293087, 1. Rio
Neshuyo where crossed by Pucallpa-Huan-
uco Road, USNM 261401, 1. Rio Ucayali,
Masisea, USNM 243237, 3 (60.4—63.4);
USNM 293090, 5; USNM 293095, 10. Rio
Ucayali, Nuevo San Juan near Masisea,
USNM 293089, 3. Rio Ucayali, Utuquinia,
USNM 293091, 1. BRAZIL: Amazonas;
Ilha da Machantaria, USNM 293096, 1
(89.0); GC, 3 (86.3-—92.0); USNM 229190,
1 (94.1); USNM 229187, 1; USNM 229185,
2. Lago Janauaca and vicinity, MZUSP
21699, 2. Rio Solimoes, Fonte Boa, MZUSP
2094756) (7 TS=92/8) MZAUSPH 2095712
(81.5-84.9). Rio Jurua, BMNH 1897.12.1:
88-100, 8 (4, 55.1-60.7). Ilha Xibeco, Rio
Solimoes above mouth of Rio Jutai, MZUSP
2VOLTSE1(94.4) = MZUSP) 2 10172683 7933=
83.7). Rio Solimoes, Ilha Sorubim above
Coari, MZUSP 20926, 161 (6, 68.5-84.5).
Rio Solim6ées, Tefé, mouth of Rio Japura,
MZUSP 27360, 1. Rio Solimoes, Benjamin
Constant, ‘Costa do*Capacétes MZUSP
27379, 1. Santo Antonio do Ica, mouth of
Rio Ica, MZUSP 21000, 4 (75.0-82.8).

Steindachnerina gracilis, new species
Figs. 1, 4-5, Table 1

Diagnosis.—The 50 to 54 scales in the
lateral line to the hypural joint and 10 or
104 scales above the lateral line to the or-
igin of the dorsal fin distinguish S. gracilis

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

from S. planiventris and S. quasimodoi
which have 40 to 46 lateral line scales and
7'2 to 8% and 7% to 9% scales above the
lateral line respectively, and from S. hy-
postoma which has 46 to 50 lateral line scales
and 8'2 to 9% scales above the lateral line.
Steindachnerina gracilis, which has a spot
of dark pigmentation at the base of the mid-
dle rays of the dorsal fin 1s readily separable
from S. hypostoma which lacks such pig-
mentation. A variety of meristic and mor-
phometric features also distinguish S. graci-
lis from the three other species within the
complex (Table 1).

Description. — Body elongate, moderately
wide transversely. Dorsal profile of head
straight or very slightly concave. Dorsal
profile of body straight or very slightly con-
vex from rear of head to origin of dorsal fin;
Straight and very slightly posteroventrally
slanted at base of dorsal fin, straight or gently
convex from base of last dorsal-fin ray to
caudal peduncle. Dorsal surface of body with
distinct median keel anterior to dorsal fin,
keel more pronounced proximate to dorsal
fin; body surface smoothly rounded trans-
versely posterior to fin. Ventral profile of
body very slightly convex from tip of lower
jaw to caudal peduncle. Pre-pelvic region
wide, distinctly flattened transversely, with
obtuse lateral angles in body wall, five series
of scales transversely across flattened region.
Pelvic fins distinctly separated medially.
Post-pelvic region somewhat flattened
proximate to pelvic fin, gradually becoming
transversely rounded posteriorly.

Greatest body depth at origin of dorsal
fin, depth 0.25—0.27 [0.26]; snout tip to or-
igin of dorsal fin 0.46—-0.49 [0.46]; snout tip
to origin of anal fin 0.84—0.86 [0.83]; snout
tip to insertion of pelvic fin 0.53-0.55 [0.53];
snout tip to anus 0.73-0.75 [0.73]; origin of
dorsal fin to hypural joint 0.54—0.58 [0.56].
Dorsal-fin margin obtusely pointed, ante-
riormost rays slightly less than three times
length of ultimate ray. Pectoral-fin margin
pointed; length of pectoral fin 0.17—0.21
[0.19], extends about three-quarters dis-

VOLUME 102, NUMBER 2

ae e:

B®, _ aids. geen od

Fig. 5.
Araguaia, Aruana.

tance to vertical line through insertion of
pelvic fin. Pelvic-fin margin obtusely point-
ed, length of pelvic fin 0.19-0.21 [0.20],
reaches about three-fifths distance to origin
of anal fin. Caudal fin distinctly forked. Adi-
pose dorsal fin well developed. Border of
anal fin emarginate, anteriormost branched
rays about two and one-half times length of
ultimate ray. Caudal peduncle depth 0.11-
O20. 11].

Head distinctly pointed in profile, head
length 0.27—0.29 [0.27]; upper jaw distinctly
longer than lower, mouth inferior; snout
length 0.31-0.34 [0.32]; nostrils very close,
anterior circular, posterior crescent-shaped
with aperture closed by thin flap of skin
separating nares; orbital diameter 0.32-0.35
[0.32]; adipose eyelid poorly developed, with
rotund opening over center of eye; length of
postorbital portion of head 0.39-0.42 [0.39];
gape width 0.32-0.35 [0.33]; interorbital
width 0.39-—0.42 [0.41].

Pored lateral line scales from supraclei-
thrum to hypural joint 50 to 54 [53]; all
scales of lateral line pored, canals in scales
straight; 3 to 5 series of scales extend beyond
hypural joint onto caudal fin base; 10 to 10'4
[1014] scales in transverse series from origin
of dorsal fin to lateral line; 612 to 7!2 [74]
scales in transverse series from lateral line
to origin of anal fin.

Dorsal-fin rays 11,9 or 11,9 (when three
unbranched rays present, first very short)
[11,9]; anal-fin rays 11,7 or 111,7 (when three
unbranched rays present, first very short)
[11,7]; pectoral-fin rays 15 to 17 [16]; pelvic-
fin rays 1,8 [1,8].

475

Steindachnerina gracilis, new species, holotype, MZUSP 4857, 63.5 mm SL; Brazil, Goias, Rio

Total vertebrae 35 (8), 36 (5).

Color in alcohol. —Overall coloration of
specimens retaining guanine on scales sil-
very golden, darker on dorsal portions of
head and body. Overall coloration of spec-
imens fixed in formalin and lacking guanine
on scales tan to light brown, darker on dor-
sal portions of head and body. Pores of lat-
eral line scales outlined by dark pigmenta-
tion, spots forming irregular mid-lateral
stripe. Deeper lying dusky stripe extending
from supracleithrum to caudal peduncle,
stripe masked in individuals retaining gua-
nine on scales. Anterior margin and distal
portions of dorsal fin dusky. Distinct dusky
spot on middle rays of dorsal fin slightly
above their insertion, spot faint in some in-
dividuals. Rays of caudal fin outlined by
series of small chromatophores; lower lobe
of caudal fin dusky. Other fins hyaline.

Distribution. —Rio Tocantins system (Fig.
4).

Etymology. —The name gracilis, from the
Latin for slender, refers to the relatively
elongate body of the species.

Type material. — 16 specimens, 56.7—73.4
mm SL.

Holotype.—BRAZIL. Goids. Rio Ara-
guaia, Aruana (approx. 14°54’S, 51°05’W),
63.5 mm SL; Excursion de Departamento
de Zoologia (now MZUSP), MZUSP 4847,
15 to 19 Sep 1966.

Paratypes.—BRAZIL: Goids; Rio Ara-
guaia, Aruana, MZUSP 38591, 2 (56.7-63.5;
taken with holotype); USNM 293035, 1
(67.2; taken with holotype); USNM 191632,
1 (approx. 73.4), H. A. Axelrod, 1960. Para:

476

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Steindachnerina quasimodoi, new species, holotype, USNM 293041, 88.9 mm SL; Peru, Loreto, Rio

Yavari, near Petropolis.

Rio Tocantins, lagoons along margin of riv-
er near Tucurui (approx. 3°42’S, 49°27'W),
MZUSP 38592, 6 (61.5-64.9); USNM
293034, 6 (57.4-65.4, 2 specimens cleared
and counterstained for cartilage and bone),
Expedicao Permanente da Amazonas, 14
Sep 1970.

Steindachnerina quasimodoi, new species
Figs. 1, 6-10, Table 1

Diagnosis. — The 40 to 46 lateral line scales
to the hypural joint distinguish S. quasi-
modoi from S. gracilis which has 50 to 54
scales in that series and S. hypostoma which
has 46 to SO. The presence of a discrete spot
of dark pigmentation in the basal portions
of the middle rays of the dorsal fin further
separates S. guasimodoi from S. hypostoma
which lacks that pigmentation. Other dif-
ferences in meristic and morphometric fea-
tures further differentiate S. quasimodoi

Pigs:
opposite Colonia Angamos, Peru.

from S. hypostoma and S. gracilis (Table
1). Specimens of S. quasimodoi of greater
than 50 mm SL are distinguished from com-
parably sized individuals of S. planiventris
in their greatest body depths (0.30-0.34 of
SL versus 0.27—0.30, Fig. 8), by differences
in the degree of convexity of the dorsal sur-
face of the body (Figs. 6, 7, 11), length of
the pelvic fins (0.23-0.26 of SL versus 0.19-
0.23, Fig. 9), and degree of contact of fron-
tals anterior to cranial fontanel (Fig. 1B, C).
The relative orbital width and length of the
postorbital region of the head also discrim-
inate the species to a degree (Table 1).
Description. —Body elongate, deeper
bodied in larger specimens, somewhat com-
pressed laterally. Dorsal profile of head
slightly convex anterior to vertical through
nostrils, striaght from that line to rear of
head. Dorsal profile of body slightly convex
from rear of head to origin of dorsal fin,
slope more pronounced with increasing size;

Steindachnerina quasimodoi, new species, paratype, USNM 283040, 53.7 mm SL; Brazil, Rio Javari,

VOLUME 102, NUMBER 2

GBD

Fig. 8. Plot of greatest body depth (GBD) against
standard length (SL), both in millimeters, and regres-
sion lines for Steindachnerina quasimodoi (dots, Y =
—4.052 + 0.380X) and Steindachnerina planiventris
(stars, Y —4.089 + 0.351X) (some symbols represent
more than one data point).

straight and slightly posteroventrally slant-
ed at base of dorsal fin in juveniles, adults
with angle more pronounced; straight from
base of last dorsal-fin ray to caudal pedun-
cle. Dorsal surface of body transversely
rounded anteriorly, with indistinct median
keel immediately anterior to dorsal fin,
smoothly rounded transversely posterior to

PFL

SL

Fig. 9. Plot of length of pelvic fin (PFL) against
standard length (SL), both in millimeters and regres-
sion lines, for Steindachnerina planiventris (dots, Y =
0.851 + 0.185%) and Steindachnerina quasimodoi
(stars, Y = 0.561 + 0.241X), (some symbols represent
more than one data point).

477

) Ue. “J Z { { ; 1 -
at ee
ag ye

Fig. 10. Map of northern South America showing
geographic distribution of Steindachnerina quasimodoi
(stars) and Steindachnerina planiventris (dots). Type
localities: 1, Rio Machado; and 2, Rio Yavari, Petro-
polis. Some symbols represent more than one lot and/
or collecting locality.

fin. Ventral profile of body very gently
curved from tip of lower jaw to origin of
anal fin, sigmoid from that point to caudal
peduncle. Pre-pelvic region broadly flat-
tened, with three irregular series of scales
across flattened ventral surface. Scale series
of pre-pelvic region flanked on each side by
series of scales bent to conform to shape of
lateral angles of body. Obtuse median keel
posterior to pelvic fin insertion. Secondary
obtuse keel on each side of post-pelvic por-
tion of body two scales dorsal of ventral
midline.

Greatest body depth at origin of dorsal
fin, depth 0.27-0.34 (0.30-0.34 in speci-
mens above 50 mm SL) [0.33]; snout tip to
origin of dorsal fin 0.47—0.50 [0.49]; snout
tip to origin of anal fin 0.84—0.86 [0.86];
snout tip to insertion of pelvic fin 0.50—0.54
[0.50]; snout tip to anus 0.75-0.78 [0.76];
origin of dorsal fin to hypural joint 0.54—
0.59 [0.59]. Dorsal-fin margin poionted, less
SO with increasing age; anteriormost rays
about three to three and one-half times
length of ultimate ray. Pectoral-fin margin
pointed; length of pectoral fin 0.19-0.21
[0.21], extends posteriorly to point about
two scales anterior of vertical through in-
sertion of pelvic fin. Pelvic-fin margin

478

pointed, length of pelvic fin 0.23-0.26 [0.26],
reaches two-thirds distance to origin of anal
fin. Caudal fin forked. Adipose dorsal fin
well developed. Anal fin emarginate, ante-
riormost branched rays about three to three
and one-half times length of ultimate ray.
Caudal peduncle depth 0.11-—0.13 [0.13].

Head distinctly pointed, head length 0.27-—
0.33 [0.28]; upper jaw much longer than
lower, mouth inferior; snout length 0.31-
0.34 [0.33]; nostrils very close, anterior cir-
cular, posterior crescent shaped with aper-
ture closed by thin flap of skin separating
nares; orbital diameter 0.31-0.37 [0.32];
adipose eyelid present, more developed in
larger specimens, particularly anteriorly,
with broad vertically ovoid opening over
center of eye; length of postorbital portion
of head 0.36-0.40 [0.36]; gape width 0.30-
0.35 [0.33]; interorbital width 0.36-0.43
[0.42].

Pored lateral line scales from supraclei-
thrum to hypural joint 40 to 46 [44]; all
scales of lateral line pored, canals in scales
straight; 4 to 6 series of scales extend beyond
hypural joint onto caudal fin base; 72 to 9%
[84] scales in transverse series from origin
of dorsal fin to lateral line; 512 to 6% [614]
scales in transverse series from lateral line
to origin of anal fin.

Dorsal-fin rays 11,9 or 111,9 (when three
unbranched rays present, first very short)
[111,9]; anal-fin rays 11,7 or ili,7 (when three
unbranched rays present, first very short)
[11,7]; pectoral-fin rays 15 to 17 [16]; pelvic-
fin rays i,8 or rarely i,7,i [i,8].

Total vertebrae 35 (11), 36 (20).

Color in life. —(The following description
is based on a color transparency of a spec-
imen captured in the Rio Javari by S. O.
Kullander). Overall coloration bright silver,
somewhat darker on dorsal portions of head
and body. Iris silver. Fins yellowish with
light margins. Dorsal fin with a small dark
patch on basal portion of middle rays.

Color in alcohol. —Ground coloration of
specimens lacking guanine on scales tan,
darker on dorsal portion of body. Dorsal
portion of head dusky. Margins of scales on

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

lateral and dorsal portions of body outlined
by series of small chromatophores in ju-
veniles. Pigmentation field expanded in
larger specimens to cover exposed surface
of scales, particularly dorsally. Mid-dorsal
line somewhat darker, particularly imme-
diately anterior of dorsal fin. Pores in lateral
line scales outlined dorsally and ventrally
by chromatophores. Dorsal fin with diffuse
patch of dark pigmentation on basal portion
of membranes between third and fifth ray.
Dorsal and caudal fins dusky. Adipose dor-
sal fin margin dusky. Anal fin hyaline or
with scattered chromatophores. Paired fins
hyaline.

Distribution. — Rio Javari (=Rio Yavari)
system of Peru and Brazil, and proximate
portions of Peruvian Amazon (Fig. 10).

Etymology.—The specific name quasi-
modoi is taken from Quasimodo, the mis-
understood hunchback bell-ringer of the
Cathedral in Victor Hugo’s novel Notre
Dame de Paris, and refers to the pro-
nounced humped dorsal profile of the body
in larger specimens of the species.

Type material. —39 specimens, 34.3-—
122.6 mm SL.

Holotype.-PERU. Loreto. Rio Javari, near
Petropolis; Hérnan Ortega, 2 Oct 1982,
USNM 293041, 1 (88.9).

Paratypes.-PERU. Loreto. 13 specimens
taken with holotype, USNM 293042, 7
(49.7-122.6, two specimens cleared and
counterstained for cartilage and bone);
MHN-USM 1698, 4 (50.7—100.6); MZUSP
38593, 2 (58.4—-59.6).

BRAZIL: Amazonas; Rio Javari system,
immediately downstream of confluence of
Rio Jaquirana and Rio Galvez; S. Kullan-
der, et al., 31 July 1984. NRM SOK/
1984312.4093, 9 (33.3-121.8); MZUSP
38594, 3 (50.1-53.6). Rio Javari system,
opposite Colonia Angamos, Peru; S. Kul-
lander, et al., 5 Aug 1984, NRM SOK/
1984317.4091, 7 (40.0-69.2); MHN-USM
1699, 3 (49.8-57.0); USNM 293040, 3
(34.3-61.5).

The following non-type specimens were
also examined:

VOLUME 102, NUMBER 2

479

Pip? 11.
Rio Machado, near mouth.

PERU: Loreto; Rio Ampiyacu, near the
mouth, USNM 175868, 1 (48.4). Pebas,
CAS-SU 36888, 1 (81.7).

Steindachnerina planiventris, new species
Figs. 1, 8-11, Table 1

Curimatus bimaculatus, Eigenmann and Ei-
genmann, 1889:422 [in part, specimens
from Ica, Brazil].

Diagnosis. — The 40 to 46 lateral in scales
to the hypural joint distinguish S. planiven-
tris from S. gracilis which has 50 to 54 scales
in that series and S. hypostoma which has
46 to 50. The presence of a discrete spot of
dark pigmentation in the basal portions of
the middle rays of the dorsal fin further sep-
arates S. planiventris from S. hypostoma
which lacks that pigmentation. Other dif-
ferences in meristic and morphometric fea-
tures further differentiate S. planiventris
from S. hypostoma and S. gracilis (Table
1). Specimens of S. planiventris of greater
than 50 mm SL are distinguished from com-
parably sized individuals of S. guasimodoi
in their greatest body depths (0.27—0.30 of
SL versus 0.30-0.34, Fig. 8), differences in
the degree of convexity of the dorsal surface
of the body (Figs. 7, 11), pelvic-fin length
(0.19-0.23 of SL versus 0.23-0.26, Fig. 9),
and degree of contact of the frontals anterior
of the cranial fontanel (Fig. 1C, B). The rel-
ative orbital width and length of the post-
orbital region of the head also discriminate
the species to a degree (Table 1).

Description.—Body elongate, slightly com-
pressed. Dorsal profile of head distinctly

Steindachnerina planiventris, new species, holotype, MZUSP 35857, 65.7 mm SL; Brazil, Rondonia,

convex anterior to vertical through nostrils,
slightly convex or straight from that point
to rear of head. Dorsal profile of body slight-
ly convex from rear of head to origin of
dorsal fin; straight and slightly posteroven-
trally slanted at base of dorsal fin, straight
from base of last dorsal-fin ray to caudal
peduncle. Dorsal surface of body trans-
versely rounded anteriorly, with indistinct
median keel immediately anterior to dorsal
fin, smoothly rounded transversely poste-
rior to fin. Ventral profile of body gently
curved from tip of lower jaw to caudal pe-
duncle. Pre-pelvic region distinctly flat-
tened, with three longitudinal series of scales
on ventral flattened surface. Ventral scale
series flanked on each side by series of scales
bent to conform to shape of lateral angles
of body. Barely discernable median keel
posterior to pelvic-fin insertion. Secondary
obtuse keel on each side of postventral por-
tion of body about two scales dorsal of ven-
tral midline.

Greatest body depth at origin of dorsal
fin, depth 0.27—0.30 [0.29]; snout tip to or-
igin of dorsal fin 0.46—0.50 [0.49]; snout tip
to origin of anal fin 0.83—0.87 [0.87]; snout
tip to insertion of pelvic fin 0.52—0.56 [0.54];
snout tip to anus 0.72-0.76 [0.75]; origin of
dorsal fin to hypural joint 0.54—0.59 [0.57].
Dorsal-fin margin pointed, less so with in-
creasing age; anteriormost rays three to three
and one-half times length of ultimate ray.
Pectoral-fin margin pointed; length of pec-
toral fin 0.18-0.23 [0.20], extends about
three-quarters distance to vertical through
insertion of pelvic fin. Pelvic-fin margin

480 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

pointed, length of pelvic fin 0.19—0.23 [0.21],
reaches about two-thirds distance to origin
of anal fin. Caudal fin forked. Adipose dor-
sal fin well developed. Anal fin barely emar-
ginate, anteriormost branched rays three to
three and one-half times length of ultimate
ray. Caudal peduncle depth 0.11-0.12 [0.12].

Head distinctly pointed, head length 0.26—-
0.31 [0.30]; upper jaw distinctly longer than
lower, mouth inferior; snout length 0.33-
0.36 [0.33]; nostrils very close, anterior cir-
cular, posterior crescent-shaped with aper-
ture closed by thin flap of skin separating
nares; orbital diameter 0.29-0.33 [0.30];
adipose eyelid present, moderately devel-
oped, more so anteriorly, with broad ver-
tically ovoid opening over center of eye;
length of postorbital portion of head 0.36-
0.41 [0.38]; gape width 0.28-0.34 [0.32];
interorbital width 0.38-0.44 [0.40].

Pored lateral line scales from supraclei-
thrum to hypural joint 40 to 46 [41]; all
scales of lateral line pored, canals in scales
straight; 4 to 6 series of scales extend beyond
hypural joint onto caudal fin base; 612 to 812
[7'4] scales 1n transverse series from origin
of dorsal fin to lateral line; 52 to 6% [54]
scales in transverse series from lateral line
to origin of anal fin.

Dorsal-fin rays i1,9 or 11,9 (when three
unbranched rays present, first very short)
[11,9]; anal-fin rays 11,7 or i11,7 (when three
unbranched rays present, first very short)
[111,7]; pectoral-fin rays 15 to 17 [15]; pelvic-
fin rays 1,8 [1,8].

Total vertebrae 34 (19), 35 (109), 36 (7).

Color in alcohol.—Specimens retaining
guanine on scales with overall coloration
golden, somewhat purplish dorsally. Ground
pigmentation in specimens lacking guanine
on scales tan to light brown, darker on dor-
sal portions of head and body. Scales on
lateral and dorsal surfaces of body with
margins outlined by series of small chro-
matophores; chromatophore series most
developed dorsally, increasingly less so ven-
trally. Dorsal fin with patch of dark chro-
matophores on basal portion of membranes

between third and fifth rays. Median fins,
particularly lower lobe of caudal fin in some
individuals, dusky. Paired fins hyaline.

Distribution. —Rio Madeira and Rio Ne-
gro basins, main channel and tributaries of
middle Rio Solimoes (Fig. 10).

Etymology.—The specific name, plani-
ventris, from the Latin planum, flat, and
venter, belly, refers to the flattened ventral
surface of the body in this species.

Remarks. —In their revision of the then-
known species of curimatids, Eigenmann
and Eigenmann (1889:42) listed extensive
series of specimens of Curimatus bimacu-
latus Steindachner (=Steindachnerina bi-
maculata) from various central and western
Amazonian localities. At least some of the
specimens from “‘I¢a”’ (the Rio I¢a of west-
ern Amazonas State in Brazil) are Stein-
dachnerina planiventris (USNM 120250).

Type material. —80 specimens, 53.5—78.8
mm SL.

Holotype. —BRAZIL: Rondonia; Rio
Machado, near mouth (approx. 8°03’S,
62°53'W); M. Goulding, 4 Sep 1980,
MZUSP 38587, 1 (65.7).

Paratypes. —BRAZIL: Rondonia; 35
specimens taken with holotype: MZUSP
38588, 17 (55.5-74.1); USNM 267986, 18
(56.3-78.8). Rio Machado, Jauari, River
Channel Beach; M. Goulding, 5 Sep 1980,
USNM 267989, 18 (46.2-67.8), MZUSP
38590, 18 (53.5-65.2). Rio Machado, Santo
Antonio; M. Goulding, 3 Sep 1980, USNM
267987, 6 (56.5-65.1, two specimens cleared
and counterstained for cartilage and bone),
MZUSP 38589, 5 (59.0-67.9).

The 157 following non-type specimens
were also examined:

BRAZIL: Rio Jurua, BMNH 1897.12.1:
88-100, 13. Rondonia; Rio Machado, Cu-
ruru, USNM 267988, 8. Roraima: Rio
Branco, Boa Vista, NMW 68887, 9; 1 NMW
68888, 10; NMW 68889, 4; NMW 68890,
8; NMW 68892, 5; NMW 68897, 1. Rio
Branco, 20 km below Boiagu, MZUSP
21164, 4 (49.3-53.7). Conceicao (? =Con-
ceicao do Mau), NMW 68898, 6. Rio Bran-

VOLUME 102, NUMBER 2

co, Serra Grande, 15 miles from Boa Vista,
NMW 68885, 6 (53.1-87.0); NMW 68893,
3; NMW 68896, 2; NMW 697355, 4. Ama-
zonas; Rio Negro, just below mouth of Rio
Branco, USNM 293097, 22 (11, 52.5—80.6).
Rio Negro, Anvilhanas, Municipio de Ay-
rao, USNM 293098, 7. Rio Solimoes, Ilha
Sorubim, above Coari, MZUSP uncat., 24
(62.3—71.3). Rio Solimoes, Coari, USNM
293099, 12. Rio Solimoes, near Ilha Ba-
rurua, above mouth of Rio Jutai, MZUSP
20987, 3 (2, 63.2—71.7). Rio Solimoes, Ilha
Xibeco, above mouth of Rio Jutai, MZUSP
uncat., 1 (73.8). Iga, USNM 120250, 5 (for-
merly MCZ 19571).

BOLIVIA: Beni; Rio Mamoré, Puerto
Siles, AMNH uncat., 2 (65.7—73.5).

Acknowledgments

We are indebted to the following individ-
uals and institutions for the loan and ex-
change of specimens, information, hospi-
tality during visits, and other assistance: Dr.
Gareth Nelson and Dr. C. Lavett Smith
(AMNH); Mr. Gordon J. Howes, Dr. P.
Humphry Greenwood and Mr. Alwyne
Wheeler (BMNH); Dr. William Eschmeyer,
Dr. Tomio Iwamoto and Mr. David Catan-
ia (CAS); Dr. Jacques Geéry (GC); Dr. Camm
Swift and Mr. Jeffrey Seigel (LACM); Dr.
Naércio A. Menezes, Dr. Heraldo A. Brit-
ski, and Dr. José Lima de Figueiredo
(MZUSP); Prof. Hernan Ortega (MHN-
USM); Dr. Sven O. Kullander and Mr. E.
Ahlander (NRM); Dr. Barbara Herzig, Mr.
Harald Ahnelt, and the late Dr. Rainer
Hacker (NMW), and Dr. Gordon M. Reid
(Horniman Museum, London).

The type material of Steindachnerina
planiventris was collected by Dr. Michael
Goulding of the Museu Paraense “‘Emilio
Goeldi,’” Belem, and that of Steindachner-
ina quasimodoi by Prof. Hernan Ortega
(MHN-USM) and Dr. Sven O. Kullander
(NRM). Numerous specimens, including the
majority of the paratypes of Steindachner-
ina gracilis were made available through the

481

Expedicao Permanente de Amazonia (EPA),
directed by Dr. Paulo E. Vanzolini
(MZUSP). Collecting efforts in Peru were
carried out as a joint project with the In-
stituto Veterinario de Investigaciones Trop-
icales y de Altura, and the Museo de His-
toria Natural ““Javier Prado,’ Lima. The
Consejo Nacional de Ciencia y Tecnologia
of Peru, and in particular its director Dr.
M. Vegas Velez greatly facilitated studies in
that country. Dr. Naércio A. Menezes
(MZUSP) provided the Portuguese trans-
lation of the ““Abstract.’? Technical assis-
tance at the Smithsonian was provided by
Mr. Andrew G. Gerberich and Mr. Jeffrey
Clayton. Photographs were prepared by Mr.
Theophilus Britt Griswold.

Research associated with this study in
Peru and Brazil was partially supported by
the Neotropical Lowland Research Pro-
gram of the International Environmental
Sciences Program of the Smithsonian In-
stitution. This paper benefitted from the
comments and suggestions of Dr. Wayne C.
Starnes, Dr. Darrell Siebert, and Dr. Wil-
liam L. Fink.

Literature Cited

Boulenger, G. A. 1887. Description of South Amer-
ican characinoid fishes.— Annals and Magazine
of Natural History 9:172-174.

Eigenmann, C.H. 1910. Catalogue of the fresh-water
fishes of tropical and south temperate Ameri-
ca.—Report of the Princeton University Expe-
dition to Patagonia, 1896-1899, 3 (Zoology, 4):
375-511.

—,& W.R. Allen. 1942. Fishes of western South
America. I. The intercordilleran and amazonian
lowlands of Peru. II. The high pampas, Bolivia,
and Northern Chile. With a revision of the pe-
ruvian Gymnotidae, and of the genus Orestias.
494 p. Lexington: University of Kentucky.

—, & R.S. Eigenmann. 1889. A revision of the
edentulous genera of the Curimatinae. — Annals
of the New York Academy of Sciences 4:409-—
440.

——., & 1891. A catalogue of the fresh-
water fishes of South America.— Proceedings of
the United States National Museum 14:1-81.

Fernandez-Yeépez, A. 1948. Los curimatidos (peces
fluviales de Sur America), catalogo descriptivo

482

con nuevas adiciones genericas y especificas. —
Boletin Taxonomico del Laboratorio de Pes-
queria de Caiquire 1:1—86.
Fowler, H. W. 1906. Further knowledge of some het-
erognathus fishes, Part I.—Proceedings of the
Academy of Natural Sciences of Philadelphia,
58:293-351.

1942. Los peces del Peru.—Boletin Museo
de Historia Natural “Javier Prado.” Lima 6:
206-222.

1945. Los peces del Peru.—Boletin Museo
de Historia Natural “Javier Prado.” Lima 7:1-
298.

1950. Os peixes de aqua doce do Brasil (2a
entrega).— Arquivos de Zoologia do Estado de
Sao Paulo 6:205-404.

1975. Acatalogue of world fishes (XXIII). —
Quarterly Journal of the Taiwan Museum 28(3):
277-402.

Géry, J. 1977. Characoids ofthe world. Neptune City,
New Jersey, TFH Publications. 672 pp.
Ortega, H., & R. P. Vari. 1986. Annotated checklist

of the freshwater fishes of Peru.—Smithsonian
Contributions to Zoology 437:1-25.
Vari, R. P. 1982. Systematics of the neotropical char-
acoid genus Curimatopsis (Pisces: Characoid-
ei). —Smithsonian Contributions to Zoology 373:
1-28.

1984. Systematics of the neotropical chara-
ciform genus Potamorhina (Pisces: Characi-
formes).—Smithsonian Contributions to Zool-
ogy 400:1-36.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

. 1989. A phylogenetic study of the neotropical
characiform family Curimatidae (Pisces: Ostar-
iophysi).—Smithsonian Contributions to Zool-
ogy 471:1-71.

in press a. Systematics of the neotropical
characiform genus Curimata Bosc (Pisces:
Characiformes).— Smithsonian Contributions to
Zoology 474.

in press b. Systematics of the neotropical
characiform genus Psectrogaster Eigenmann and
Eigenmann (Pisces: Characiformes).—Smith-
sonian Contributions to Zoology 481.

in press c. Systematics of the neotropical
characiform genus Pseudocurimata Fernandez-
Yeépez (Pisces, Characiformes).—Smithsonian
Contributions to Zoology 490.
, & R. M.C. Castro. 1988. Prochilodus stig-
maturus Fowler, reassigned to the Curimatidae
from the Prochilodontidae, with comments on
other nominal curimatid and prochilodontid
species treated by Fowler.—Copeia 1988(3):777—
780.

(RPV) Department of Vertebrate Zoology
(Fishes), National Museum of Natural His-
tory, Smithsonian Institution, Washington,
D.C. 20560; and (AWV) ADP, National
Museum of Natural History, Smithsonian
Institution, Washington, D.C. 20560.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 483-490

DESCRIPTION OF TWO NEW SPECIES OF THE
ELEUTHERODACTYLUS MILESI GROUP
(AMPHIBIA: ANURA: LEPTODACTYLIDAE)
FROM NORTHERN HONDURAS

James. R. McCranie, Jay M. Savage, and Larry David Wilson

Abstract. —Two new species of the Eleutherodactylus milesi group (E. chry-
sozetetes and E. cruzi) are described from the Cordillera de Nombre de Dios
of northern Honduras. Eleutherodactylus milesi is shown to have a wide range
in Honduras on scattered montane ridges and is reported from Guatemala for

the first time.

The systematics of Eleutherodactylus ma-
tudai and E. milesi were reviewed by Savage
(1975), who included them in the E. rugu-
losus group. These species subsequently were
removed from the rugulosus group (Miya-
moto 1983, Savage 1987) and placed as a
separate infrageneric lineage, the E. milesi
group (Savage 1987). Members of this stock
differ in karyology (2N = 22; NF = 44) and
jaw musculature (DFSQAT + e) from those
of the rugulosus group (2N = 20, 22; NF =
36; jaw muscle formula: dfsq; dfsqat + e).
Externally, frogs of the mi/esi group are dis-
tinct (features for the rugu/osus group in pa-
rentheses) in: lacking tarsal folds (a well-
developed inner tarsal fold), having a
strongly tuberculate dorsum (dorsum
smooth to rugose), and having a pair of para-
anal light bars (never with such markings).
In addition E. matudai has an indistinct
tympanum in females and E. milesi has the
tympanum indistinct or hidden (distinct,
nearly transparent tympanum).

Recent fieldwork in Guatemala and Hon-
duras substantially increases the known
range for Eleutherodactylus milesi and this
information and data on variation are pre-
sented here. Among other newly collected
material from the Cordillera de Nombre de
Dios of northern Honduras are two forms,
clearly allied to E. matudai and E. milesi,
but trenchantly different from them and

from one another. These distinctive frogs
are regarded as representing previously un-
known species and are described below fol-
lowing the style and terminology of Savage
(1975); a few features not mentioned in that
paper follow the definitions of Savage (1987).

Eleutherodactylus chrysozetetes,
new species
Fig. 1

Holotype.—KU 209035, adult male, from
Quebrada de Oro (15°38'N, 86°47'W), ele-
vation 880 m, tributary of Rio Viejo, south
slope of Cerro Bufalo, Cordillera de Nombre
de Dios, Departamento de Atlantida, Hon-
duras, 17 Aug 1984, James R. McCranie,
Kenneth L. Williams, and Larry David Wil-
son. Original number LDW 6529.

Diagnosis. —Eleutherodactylus chrysoze-
tetes is a moderately large frog and is distin-
guished from the two previously described
members of the milesi group, E. matudai
and &. milesi (characteristics for these
species in parentheses), in having extensive
toe webbing with about three phalanges free
of the web on toe IV (basal to moderate
webbing, with four to four and one-half pha-
langes free of web); strongly expanded finger
and toe disks, about twice width of digit
(finger disks definite, less than twice width
of digit; toe disks weak, barely expanded);

ey mh “=
e~.

Fig. 1.

tympanum hidden in adult males and pre-
sumably females (tympanum evident in
adult males). Eleutherodactylus chrysoze-
tetes is further distinguished from E. ma-
tudai (states for matudai in parentheses) by
having: paired vocal slits in adult males (no
vocal slits); groin with a light spot on the
dark uniform flank color (groin mottled). In
addition, E. chrysozetetes can be distin-
guished from E. milesi (states of milesi in
parentheses) by having: a well-developed
fleshy fringe along margins of toes (a very
narrow ridge); adult males to 41.3 mm in
standard length (20-34 mm). E. chrysoze-
tetes 1s compared to the second new species
described in this paper in the diagnosis for
the latter form.

Summary of characteristics. —Dorsum
tuberculate; canthus rounded; tympanum
hidden in adult males; first finger equal to
or shorter than second; finger disks strongly

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Holotype, KU 209035. Eleutherodactylus chrysozetetes.

expanded, at least twice width of digit just
proximal to disks on fingers IIJ-IV; inner
tarsal fold absent; toe disks strongly devel-
oped, about twice width of digit; a well-
developed fleshy fringe along margins of
toes; toes extensively webbed, webbing for-
mula: I 1+—2, II 1+-2 1/2, III 2-3 1/4, IV
3+—2 V; paired vocal slits in adult males;
nuptial pads in adult males; throat and ven-
ter heavily suffused with pale purple; dor-
sum dark olive brown, blotched with pale
Olive brown; groin light spotted on a uni-
form dark flank; posterior surface of thigh
uniform purplish-brown with a light para-
anal bar; underside of tibial segment suf-
fused with pale purple.

Coloration. —In life the holotype was dark
olive brown, blotched with pale olive brown;
upper surfaces of limbs same as dorsum;
posterior surface of thigh uniform purplish-
brown with a light para-anal bar; venter and

VOLUME 102, NUMBER 2

throat pale purple; soles of hands and feet
purple; underside of tibial segment suffused
with pale purple; iris steel gray with rust-
red horizontal band.

Measurements. —Standard length in mil-
limeters, other measurements as percent-
ages of standard length: standard length 41.3;
head length 41.9; head width 42.1; orbit
13.3; snout length 17.2; loreal length 9.2;
hindleg length 166.1; tibia length 53.0.

Etymology. —The specific epithet is from
the Greek chrysos (=gold) and zététés
(=searcher), with reference to the occur-
rence of this form along the Quebrada de
Oro (stream of gold).

Natural history notes. —The holotype and
a juvenile specimen were collected during
the morning in the forest alongside the Que-
brada de Oro. The vegetation at the type
locality is in the Subtropical Wet Forest for-
mation (Holdridge 1967) and will be de-
scribed in more detail in a subsequent pa-
per. The known elevational range is 880-
1110 meters.

Referred specimen.—KU 209036, a ju-
venile from Quebrada de Oro, taken be-
tween 920-1110 m.

Eleutherodactylus cruzi, new species
Fis. 2

Holotype.—KU 209037, adult male, from
south slope of Cerro Bufalo (15°38'N,
86°47'W), elevation 1520 m, Cordillera de
Nombre de Dios, Departamento de Atlan-
tida, Honduras, 4 Jun 1980, Gustavo A.
Cruz Diaz, James R. McCranie, and Larry
David Wilson. Original number LDW 5606.

Paratopotype.—KU 209038, adult male.

Diagnosis.—A medium sized Eleuthero-
dactylus (males to 32.6 mm in standard
length, females presumably 8 to 10 mm
longer) differing from all other members of
the milesi group in having the following
combination of characteristics: moderate toe
webbing, with about four phalanges free of
web on toe IV; definite finger disks, almost
twice width of digit; toe disks weak, barely

485

expanded; a distinct ridge along toes, ba-
sally; tympanum hidden in adult males (and
presumably females); and a unique color-
ation with a narrow, median dorsal light
stripe, the venter cream and lightly punctate
and the throat punctate. Eleutherodactylus
cruzi most closely resembles the larger E.
matudai (adult males to 50 mm in standard
length) in disk size, toe webbing and fring-
ing. The latter form has the tympanum dis-
tinct in males and indistinct in females (hid-
den in male and almost certainly in female
E. cruzi), the venter uniform gray with sil-
ver or white spots, throat heavily pigment-
ed, dorsum without a median dorsal light
stripe and groin mottled (venter cream and
lightly punctate, throat punctate, a median
dorsal light stripe and a distinct white spot
in groin in cruzi). E. cruzi has more toe
webbing, somewhat broader finger disks and
somewhat better developed toe ridges than
E. milesi but is most obviously distinct in
having the tympanum hidden in males and
in coloration (the latter species has the tym-
panum evident but indistinct in males, the
venter and throat heavily marked and/or
suffused with dark pigment and the dorsum
never with a narrow median light stripe).
The newly described E. chrysozetetes can-
not be confused with E. cruzi (features for
the latter in parentheses) since the former
has strongly expanded finger and toe disks,
about twice as wide as digits (definite finger
disks, less than twice as wide as digits; bare-
ly expanded toe disks), extensive toe web-
bing with only about three phalanges on toe
IV free of web (about four phalanges free of
web), distinct toe fringes (toes ridged) and
a larger size to 41.3 mm in adult males (to
32.6 mm). Coloration also will separate these
forms as E. chrysozetetes resembles E. ma-
tudai and E. milesi in having the throat and
venter heavily marked with dark pigment
and lacks a mid-dorsal light stripe.
Summary of characteristics. —Dorsum
tuberculate; canthus rounded; tympanum
hidden in adult males; first finger equal to
or shorter than second; finger disks definite,

486

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Holotype, KU 209037. Eleutherodactylus cruzi.

almost twice width of digit just proximal to
disks on fingers IIJ-IV; inner tarsal fold ab-
sent; toe disks weakly expanded; ridge along
margins of toes basally; toes moderately
webbed, modal webbing formula: I 2—2 1/2,
II 2-3 1/2, II 3-4, IV 4-2 1/2 V; paired
vocal slits in adult males; nuptial pads in
adult males; venter cream, lightly punctate;
throat punctate; dorsum uniform brown ex-
cept for a narrow mid-dorsal light stripe;
groin with a light spot; posterior surface of
thigh uniform brown with a distinct light
para-anal bar; underside of tibial segment
banded; adult males 26.9—32.6 mm in stan-
dard length.

Coloration.—In preservative the holo-
type (KU 209037) is uniform brown with a
narrow mid-dorsal light stripe; dorsal sur-
faces of limbs brown with dark brown bands;
venter cream, lightly punctated with brown;

throat cream, punctated with brown; groin
brown with a cream colored light spot; pos-
terior surface of thigh brown with light
brown para-anal bar; underside of tibial sur-
face cream, banded with brown. The para-
topotype (KU 209038) is similar to the
holotype except that the overall appearance
of the dorsal surfaces is a slightly lighter
shade of brown.

Measurements. —Standard lengths in
millimeters, other measurements as per-
centages of standard length (holotype fol-
lowed by paratopotype). Standard length in
adult males 32.6, 26.9; head length 42.6,
45.0; head width 42.6, 44.2; orbit 16.3, 14.9;
snout length 17.5, 18.6; loreal length 10.4,
11.2; hind leg length 184.0, 201.4; tibia
length 58.0, 64.7.

Etymology.—The species is named in
honor of Gustavo A. Cruz Diaz, in recog-

VOLUME 102, NUMBER 2

nition of the many years of friendship
McCranie and Wilson have enjoyed with
this exemplary Honduran biologist and the
invaluable assistance he has proferred to
them.

Natural history notes.—Both specimens
of this species were collected during the
morning at 1520 m alongside a trickle at
the beginning of a small stream in the Lower
Montane Wet Forest formation (Holdridge
1967).

The Eleutherodactylus milesi
Population System

Recently collected material of frogs re-
sembling Eleutherodactylus milesi from the
Caribbean versant of Guatemala and Hon-
duras provide significant extensions in the
known geographic and elevational ranges for
the species group and exhibit substantial
variation in features previously used (Sav-
age 1975) to diagnose the species. E. milesi
was described from the mountains (Sierra
de Omoa) west of San Pedro Sula, Depar-
tamento de Cortés, Honduras. Additional
localities for the species were the Montanas
de Yoro, Departamento de Yoro and the
Cordillera de Nombre de Dios, Departa-
mento de Atlantida of north-central Hon-
duras.

These samples shared the following dis-
tinctive features: 1) dorsum tuberculate; 2)
canthus rounded; 3) tympanum indistinct
in males, hidden in females; 4) finger disks
definite; 5) no tarsal fold; 6) toe disks barely
expanded, hardly wider than digit just prox-
imal to disk; 7) a very narrow ridge along
margins of toe; 8) toes with no or at most
basal webbing; 9) paired vocal slits in adult
males; 10) nuptial pads in adult males; 11)
venter heavily suffused with light brown to
chestnut pigment; 12) throat heavily pig-
mented to almost uniform dark brown; 13)
dorsum brown, spotted to occasionally
blotched, rarely with a broad mid-dorsal
light stripe; 14) groin with a single light spot

487

on dark uniform flank color; 15) posterior
surface of thigh uniform reddish brown with
a distinct light para-anal bar; 16) underside
of tibial segment boldly marked with brown
spotting or mottling, rarely suffused with
chestnut pigment; 17) adult males 20-27 in
standard length, females to 34 mm. The
maximum size of adult males recorded as
22 mm in Savage (1975) was a typograph-
ical error. The newly acquired specimens
from east and west of previously known lo-
calities for the species all agree with the
above description in characteristics 1, 2, 4-6,
8-16. Variation in other features is briefly
described below by general locality from east
to west (populations numbered as on Fig. 3).

Samples 1-6 are very similar to one
another and to previously collected material
except that adult males attain lengths (in
mm) of 28 (sample 3) and 34 (sample 6) and
a female reaches 39 (sample 3).

Samples 7-10 have the tympanum indis-
tinct in males and most females (no males
present in sample 8, one female in sample
9 with the tympanum hidden on the right
side); adult males reach to 28 (in mm) (sam-
ple 7) and 30 (sample 9) and females 41
(sample 9).

Samples 7 and 8 from the sierra along the
border between Guatemala and Honduras
and sample 9 from the Sierra de las Minas,
Departamento de Baja Verapaz, Guatemala
are slightly different from other series of the
milesi complex in the condition of the tym-
panum in females (usually indistinct, not
hidden). However, we are reluctant to place
undue emphasis on this feature or the slight
variations in webbing, toe disk width and
fringing and adult size among samples 1-9
and conclude that they represent a single
morphospecies, Eleutherodactylus milesi.
This species sseems to be comprised of iso-
lated populations generally occurring on
widely separated ridges between 500-1720
m elevation in Caribbean versant Guate-
mala and Honduras. The single known fe-
male (BM 1985/1452) from extreme eastern
Honduras (sample 1) probably was taken

Figs3:

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ELEUTHERODACTYLUS
MILESI

KILOMETERS

Distribution of the Eleutherodactylus milesi group in Honduras and Guatemala. Numbers refer to

samples discussed in text. E. milesi and both new species, E. chrysozetetes and E. cruzi described here occur at

locality 5.

below 500 m elevation and this population
needs further study to establish its status
conclusively.

Sample 10, a single female (MVZ 159934),
is from the somewhat isolated Sierra Xuc-
aneb, Departamento de Alta Verapaz, Gua-
temala. It differs from all other examples of
the Eleutherodactylus milesi population
system in having well-developed toe fringes,
in this one feature it resembles E/leuthero-
dactylus matudai of northwestern Pacific
slope Guatemala and Eleutherodactylus
chrysozetetes (described in this paper from
northcentral Honduras). This specimen may
represent a distinct species within the com-
plex, but we are reluctant to recognize it as
new without seeing adult males.

Specimens examined.—Honduras: Co-
lon; Sample 1, Rio Guaraska, north slope
Montanas de Rio Platano, (BMNH 1985/
1452). Olancho; Sample 2, Montana de Ma-

lacate, 760 m (KU 209059). Sample 3, Sier-
ra de Agalta, 1450 m (ROM 18099-107),
1060 m (ROM 18113), 1050 m (ROM
18109-111), 1000 m (ROM 18108, 18112).
Yoro; Sample 4, Montanas de Yoro, Por-
tillo Grande, 1460 m (MCZ 21290; FMNH
21862-64), Santa Marta, 1460 m (MCZ
21291). Atlantida; Sample 5, Cordillera de
Nombre de Dios, above La Ceiba (MCZ
21293-294), Cerro Bufalo, Quebrada de
Oro, 1110 m (KU 209058), 940 m (KU
209083-091), 880 m (KU 209080-082,
209096, 209098-104, 209142-143, LACM
137306-311), 810 m (KU 209105-—106), 780
m (KU 209092-095). Cortés; Sample 6,
Sierra de Omoa, west of San Pedro Sula
(FMNH 4699-4711, 142147, 21820,
21829), MCZ 21295-99, UMMZ 120388,
USNM 118202), Cerro Cusuco, 1650-1720
m (LACM 137298-305), 1690 m (KU
209067-—070, 209107), 1580 m (KU

VOLUME 102, NUMBER 2

209040-051), 1570 m (KU 209060), 1560-
1570 m (KU 209065-066), 1550 m (KU
209052-055, 209061-064), 1540-1550 m
(KU 209071-075), 1530 m (KU 209056-
057, 209141). Copan; Sample 7, Sierra Es-
piritu Santo, Montana Cerro Azul, 1050 m
(KU 209077), 1100 m (KU 209076), 1300
m (KU 209078-079), 1350 m (KU 209097).
Guatemala: Zacapa; Sample 8, Sierra Es-
piritu Santo, La Union, 850 m (CM 57743-
44), Baja Verapaz; Sample 9, Sierra de las
Minas, Aldea Vista Hermosa, 500-650 m
(KU 189797-99, 189801-04). Alta Vera-
paz; Sample 10, Sierra de Xucaneb, Finca
Volcan, 875 m (MVZ 159934).
Distribution. —Premontane and lower
montane forests on the Atlantic versant of
northern Honduras and adjacent eastern
Guatemala (500-1720 m) (Fig. 3).

Co-occurrence and Relationships

The species Eleutherodactylus matudai
(northwestern Guatemala), the various
populations of Eleutherodactylus milesi
(Caribbean slope Guatemala and Hondu-
ras) and the montane form Eleutherodac-
tylus cruzi from above 1500 m in north-
central Honduras are allopatric, although
the latter ultimately may prove to be sym-
patric with E. milesi. Eleutherodactylus
chrysozetetes is sympatric with E. milesi be-
tween 880-1110 m elevation on Cerro Bu-
falo. That three distinctive species of this
group occur on Cerro Bufalo, and possibly
elsewhere in the Cordillera Nombre de Dios,
is almost inexplicible but perhaps reflects
the stochastic influence of the orogenic fac-
tor in tropical frog evolution.

In terms of relationships the Caribbean
versant species seem to form a derived clus-
ter, as compared to Eleutherodactylus ma-
tudai, and have the characteristic white groin
spot and reduction in distinctiveness of the
tympanum. E. matudai lacks vocal slits
which are found in adult males of Caribbean
versant populations. Other features (toe
webbing, fringing, and finger disk devel-
Opment) combine in a mosaic in the three

489

recognizable Caribbean slope forms. Of the
three, Eleutherodactylus chrysozetetes is
most similar to E. matudai in hand and foot
structure but most different from it in the
modification of the tympanum. Thus, while
E. matudai seems to be the primitive sister-
group (in which vocal slits have been lost)
to the eastern forms, it is not possible on
the basis of current data to explicate the
relationships of the Caribbean versant
species among themselves.

Acknowledgments

Gustavo A. Cruz Diaz, Mario Espinal,
Kelly M. Hogan, and Kenneth L. Williams
provided field assistance and companion-
ship. Don Tomas Meraz and his family as-
sisted McCranie and Wilson on all visits to
the Cerro Bufalo-Quebrada de Oro area.
Wilberto Aguilar N. and Gustavo A. Cruz
Diaz were instrumental in the acquisition
of collecting permits for Honduras. Brian I.
Crother of the University of Miami drafted
the map. We thank all of these individuals
for their aid and the authorities at the fol-
lowing institutions (abbreviations in paren-
theses used to identify these collections
throughout this paper) who arranged for us
to examine pertinent materials: British Mu-
seum (Natural History) (BMNH); Carnegie
Museum (CM); Field Museum of Natural
History (FMNH); Museum of Natural His-
tory, University of Kansas (KU); Museum
of Comparative Zoology, Harvard Univer-
sity (MCZ); Museum of Vertebrate Zoology,
University of California, Berkeley (MVZ);
University of Michigan, Museum of Zool-
ogy (UMMZ); National Museum of Natural
History, Smithsonian Institution (USNM).
The following abbreviations indicate col-
lections where material reported here has
been deposited: Natural History Museum
of Los Angeles County (LACM) and Royal
Ontario Museum (ROM).

Literature Cited

Holdridge, L. R. 1967. Life zone ecology. Tropical
Science Center, San José, Costa Rica. 206 pp.

490 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Miyamoto, M. M. 1983. Frogs of the Eleutherodac-
tylus rugulosus group: A cladistic study of al-
lozyme, morphological, and karyological data. —
Systematic Zoology 32(2):109-124.

Savage, J. M. 1975. Systematics and distribution of
the Mexican and Central American stream frogs
related to Eleutherodactylus rugulosus. —Copeia
1975(2):254-306.

—. 1987. Systematics and distribution of the
Mexican and Central American rainfrogs of the
Eleutherodactylus gollmeri group (Amphibia:

Leptodactylidae). — Fieldiana: Zoology, N.S. 33:
iv + 1-57.

(JRM) 10770 S.W. 164th Street, Miami,
Florida 33157; (JMS) Department of Biol-
ogy, University of Miami, Coral Gables,
Florida 33124; (LDW) Department of Bi-
ology, Miami-Dade Community College,
South Campus, Miami, Florida 33176.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 491-499

A NEW SPECIES OF DIMINUTIVE
ELEUTHERODACTYLUS (LEPTODACTYLIDAE)
FROM OAXACA, MEXICO

Jonathan A. Campbell, William W. Lamar, and David M. Hillis

Abstract. —Eleutherodactylus polymniae 1s described from the northern slopes
of the Sierra Juarez, Oaxaca, México. It is a small species, with distinctively
marked flanks and thighs, and has a vocal repertoire of at least four separate,
distinct calls. This species is apparently a member of the a/fredi group, and
may exist sympatrically with E. spatulatus.

Resumen.-—Se describe una nueva especie de rana, Eleutherodactylus polym-
niae, conocido del vertiente norteno de la Sierra Juarez, Oaxaca, México. Es
una especie diminutiva, con manchas distintivas sobre la parte lateral del
abdomen y sobre los muslos, y dispone de un repertorio vocal que consiste de
por lo menos cuatro cantos separados y distintos. Aparentemente dicha expecie
es miembro del grupo al/fredi, y posiblemente existe en simpatria con E. spat-

ulatus.

The vegetation covering the northern
slopes of the Sierra Juarez in Oaxaca com-
prises one of the most extensive tracts of
cloud forest remaining in México. Within
the last decade, however, agricultural clear-
ings have steadily advanced up the hillsides
so that most of the original forest below the
1200 m contour has been severely altered.
Also the leeward (southern) side of the range
is drier than the northern slopes, and is sub-
ject to periodic man-induced forest fires that
have damaged the higher portions of the
cloud forest. Previously, owing to the cold,
wet environment and poor soils, the portion
of the forest between about 1500 and 3000
m was almost uninhabited by humans.
However, with the recent advent of hard-
wood logging, the entire forest is in danger
of soon being irreversibly damaged with an
associated loss of many species of animals.

The cloud forest of the Sierra Juarez har-
bors a remarkably diverse herpetofaunal as-
semblage containing many endemic species.
Only one road traverses the region (Mexican
Highway 175), and it has been primarily

along this road that biologists have made
collections. Probably no other region of
similar size in México has yielded more new
species of amphibians and reptiles in the
last few decades, nor remains so superfi-
cially known. A few of the species, by no
means a complete listing, of recently de-
scribed species that, so far as is known, are
endemic to the region include Pseudoeury-
cea juarezi (Regal, 1966), Hyla cyvanomma
and H. sabrina (Caldwell, 1974), H. echi-
nata (Duellman, 1962), H. mixe(Duellman,
1965), Norops polyrhachis (Smith, 1968),
Abronia mitchelli (Campbell, 1982), A. ka-
laina (Good & Schwenk, 1985), Barisia
Juarezi (Karges & Wright, 1987), Exiliboa
placata (Bogert, 1968), Cryophis hallbergi
(Bogert & Duellman, 1963), and Geophis
duellmani (Smith & Holland, 1969).

In the summer of 1983, we discovered
yet another unknown anuran species in the
Sierra Juarez. This frog inhabits steep slopes
and is extremely difficult to collect. The re-
markable voice of this species, in combi-
nation with its habitat, where a misstep by

492

anyone pursuing it might have disastrous
consequences, moves us to name this frog

Eleutherodactylus polymniae, new species
Figs. 1-3

Holotype.— University of Texas at Ar-
lington (UTA) A-12976 (original number,
JAC 9007), an adult male, collected 0.8 km
N Vista Hermosa, northern slopes of the
Sierra Juarez, Oaxaca, México, at an ele-
vation of 1420 m, 25 Jul 1983, J. A. Camp-
bell, W. W. Lamar, and D. M. Hillis.

Paratype.—UTA A-23511, an adult to-
potypic male, taken by the same collectors,
17 Aug 1983.

Diagnosis. —A small species of Eleu-
therodactylus (SVL of largest male, 19.7
mm); possessing expanded digital pads, es-
pecially on the outer two fingers, which are
between three and four times wider than the
digits at their narrowest point; a relatively
small tympanum, the horizontal distance
across which is less than a third of the dis-
tance across the eye; and distinctive white
markings extending anterodorsally across
the flanks from the inguinal region. The
species differs from all Mexican congeners
in having four separate, distinct calls.

Description. —Dorsal outline of snout
broadly rounded; snout profile in lateral view
truncate; canthus rounded; choanae round
except along the anterior edge which is
straight owing to the shape of the lateral
wings of the prevomers; vomerine teeth
present, three or four teeth per fascicle;
tongue ovoid, notched posteriorly; glottal
disc large; vocal slits elongate, extending
from a point about equal to midlength of
tongue to the corner of the mouth; eyelids,
dorsum, forearms, and shanks, with a few
enlarged tubercles, otherwise skin of dor-
sum smooth, that of venter corrugate; tym-
panum small, dorsal margin of tympanic
ring indistinct; no glands apparent in the
axillary, lumbo-inguinal, or inguinal areas;
second finger longer than first; finger and

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

toe discs expanded and truncate (emargin-
ate in dorsal view), pads on outer fingers
between three and four times wider than
digit; a whitish nuptial thumb pad present;
subarticular tubercles rounded and raised;
supernumerary tubercles absent on digits;
three rounded or oblong palmar tubercles;
small calcar tubercles present; inner and
outer tarsal folds absent, but several small
tubercles along outer surface of tarsus, and
a single tubercle along inner surface of tar-
sus; inner and outer metatarsal tubercles
present, inner larger; several small tubercles
on distal plantar surface, not extending pos-
teriorly to level between metatarsal tuber-
cles; webbing on hands and feet vestigial.

The dorsum is mostly greenish brown with
some rust colored markings. Dark brown
markings are present along the posterior
margin of the nostrils and along the upper
lip, with large spots below the eye and above
the corner of the mouth. Dark brown mark-
ings also extend along the upper border of
the tympanum and the scapular region of
the dorsum. The inguinal region is bright
white with two irregular white bars extend-
ing forward anterodorsally to about the
midlength of the body (Fig. 1). The anterior
and posterior surfaces of the thighs are pat-
terned with sharply contrasting white and
brown bars; the forearms, tibia, and dorsal
surfaces of the thighs have moderately con-
trasting pale brown and darker brown bars.
The throat and venter are pale with scat-
tered black melanophores; the ventral sur-
faces of the thighs and the plantar surfaces
are darker, with numerous melanophores.

The snout-vent length (SVL) is 19.7 mm;
the head length, 8.2 mm; the head width,
8.1 mm; the horizontal distance across the
eye, 2.8 mm; the distance from the eye to
the tip of the snout, 3.6 mm; the distance
from the eye to the nostril, 2.5 mm; the
horizontal distance across the tympanum,
0.9 mm; the tibia length, 11.3 mm; and the
distance from the proximal edge of the inner
metatarsal tubercle to the distal tip of the
fourth toe, 9.7 mm.

VOLUME 102, NUMBER 2

; i
aa

Bip. 1.

Variation.—The paratype (UTA
A-23511) is similar to the holotype in most
aspects of morphology, color, and pattern.
Measurements for the paratype are as fol-
lows: SVL, 19.0 mm; head length, 7.9 mm;
head width, 7.3 mm: horizontal distance
across eye, 2.5 mm; eye to tip of snout, 3.4
mm; eye to nostril, 2.3 mm; horizontal dis-
tance across tympanum, 0.8 mm; tibia, 11.6
mm; and distance from proximal edge of
inner metatarsal tubercle to distal tip of
fourth toe, 9.5 mm.

Etymology. —The name polymniae is de-
rived from Polymnia, also known as “‘she
of the many hymns,” one of the Greek si-
rens that was believed to lure mariners to
destruction by her singing.

Remarks. —In many respects (size, color,
morphology) FE. polymniae resembles cer-
tain members of the genera Tomodactylus
and Syrrhophus (for reviews of these genera,

493

Eleutherodactylus polymniae, new species, holotype, SVL 19.7 mm, UTA A-12976.

see Dixon 1957, and Lynch 1970a, respec-
tively). It may be distinguished from species
contained in these genera by the absence of
lumbo-inguinal or inguinal glands, and by
the failure of the plantar supernumerary tu-
bercles to extend posteriorly to a level be-
tween the metatarsal tubercles. Most species
of Syrrhophus and Tomodactylus also have
supernumerary tubercles on the digits, and
especially on the toes; in E. polymniae the
digits lack supernumerary tubercles.
Eleutherodactylus polymniae conforms to
the diagnosis of the genus Eleutherodactylus
provided by Lynch (1968).

Mexican and Guatemalan members of
the genus

Eleutherodactylus having widely expand-
ed digital pads have been placed in the a/-
fredi group (Lynch 1965, 1966, 1967a, b, c,

494

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Pattern of posterior surface of thighs in Eleutherodactylus polymniae, paratype, UTA A-23511.

1970b). Other than sharing distinctive dig-
ital pads, few characters have been reported
to suggest that this group is monophyletic.
Geographically, the group is relatively com-
pact, with 11 species along the Atlantic
coastal plain and versant from Tamaulipas,
México, southward through much of the
Yucatan Peninsula to the mountains of
northern Guatemala. One species, E. guer-
reroensis, 1S restricted to the Pacific versant,
and another, FE. stuarti, occurs on both the
Atlantic and Pacific side of the Continental
Divide (Fig. 4). Most species have ex-
tremely restricted montane distributions,
and many are known from fewer than half
a dozen specimens. Eleutherodactylus alfredi
has a wide distribution, ranging from east-
central Veracruz (Smith & Taylor 1948) to
Tabasco (Lee 1980) and Piedras Negras,
Guatemala (Duellman 1960, Stuart 1963).
Eleutherodactylus decoratus occurs along the
Atlantic slopes of the Sierra Madre Oriental
from southern Tamaulipas to central Ve-

racruz (Martin 1958, Lynch 1967c). Eleu-
therodactylus stuarti ranges from western
Chiapas (Johnson 1974, Johnson et al. 1976)
into western Guatemala. Eleutherodactylus
xucanebi occurs in the northern cloud for-
ests of Guatemala and is the southernmost
member of the group (Stuart 1941). With
the possible exception of E. polymniae and
E. spatulatus, no two members of the group
are known to be sympatric with one another.

Males in this group of frogs range in size
from E. polymniae (19.7 mm) to E. guer-
reroensis (39.6 mm). Considerable sexual
dimorphism exists and females usually at-
tain from 1.3 to 1.4 times the SVL of males.
The relative size of the tympanum is also
usually sex dependent, being larger in the
males of most species. The tympanum to
eye ratio of all species in which the males
are known is 0.62 or greater, except for E.
polymniae and E. spatulatus which have ra-
tios of 0.25-0.30 and 0.32, respectively.
Eleutherodactylus megalotympanum was

VOLUME 102, NUMBER 2

495

Fig. 3. Ventral aspects of hand and foot of Eleutherodactylus polymniae, holotype, UTA A-12976.

named in allusion to what was perceived to
be the relatively large tympanum in fe-
males. However, the tympanum is not larg-
er than that known in females of several
other species. Eleutherodactylus spatulatus
differs from other species in the a/fredi group
in that males have a slightly smaller tym-
panum than females. A large series of E.
decoratus was reported to have little sexual
dimorphism in the size of the tympanum
(Martin 1958). The tarsal fold in this group

may be poorly to moderately developed or
absent. The color pattern of the flank in
most species is rather drab, being a pale
yellow, cream, gray, or tan with irregular
stippling or mottling. The specimens of E.
decoratus that we have seen in life had a
contrasting pattern of brown markings on a
pale background, often with a narrow pale
vertebral stripe. Eleutherodactylus polym-
niae has the most distinctive markings in
the al/fredi group, with a large irregular white

496

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

. decoratus

. spatulatus

- polymnius

. megalotympanum
. guerreroensts
. stlvtcola

. stuartt

. taylort

. glaucus

. xuecanebt

. yuecatanensts

@ ez
O fF
¥ EF
VeE
@ F£
AE
OB
ae
E
BeE
AE

Fig.4. Distributions of Mexican and Guatemalan species of Eleutherodactylus with greatly expanded, truncate
digital pads on outer two fingers. In some instances, for multiple localities that are in close proximity, a single
symbol is used. Localities for E. alfredi, ranging from east-central Veracruz to northwestern Guatemala, are not

indicated.

blotch in the inguinal region. The color pat-
tern of the anterior, dorsal, and posterior
surfaces of the thighs is unremarkable in
most species. Generally, there are about four
dark transverse bands on a somewhat paler
background, but the bands are often ill-de-
fined and, at best, only moderately contrast-
ing with the adjacent color. Eleutherodac-
tylus decoratus has relatively sharply
contrasting thigh bands, and, in E. polym-
niae, the bands are even more distinct (Fig.
2).

Both the holotype and paratype of E. po-
lymniae were taken from the side of an ex-
tremely steep slope adjacent to a roadcut in
virgin cloud forest. They were found be-
tween 23:00 h and 03:00 h. They were call-
ing from vegetation; the holotype was about
3 m up in a tree, and the paratype was about
1 m above the ground. In both instances, a
low fog reduced visibility to only a few me-
ters and rain had fallen within the preceding
24 hr. Had we not fortuitously selected the
site to make camp for the night, we most
probably would not have discovered these
frogs. We became aware of their presence

by their remarkably voluminous and varied
calls. In fact, these tiny frogs are so loud
that we at first mistakenly thought that their
calls belonged to some large, night-calling
bird, unknown to us. Field notes taken im-
mediately after the capture of UTA A-23511
indicate calls include ‘“‘a low growl, a low
chuckle, single clicks, and a multinote
‘laugh.’ First two are given [relatively infre-
quently]; clicks (several) precede ‘laugh’.”
Four other species of Mexican Eleuthero-
dactylus with expanded toe pads are known
to have vocal slits (Table 1), including E.
spatulatus, with which E. polymniae may
be sympatric; specimens of the former have
been taken from the same hillside about 200
m above the type locality for E. polymniae.
Some of the structures associated with vo-
calization are relatively large in E. polym-
niae. The vocal slits extend forward to a
point about level with the midlength of the
tongue in E. polymniae; in E. spatulatus,
the slits extend only to about the posterior
edge of the tongue. Perhaps more remark-
able is the extent of development of the cir-
cular disc surrounding the glottis, which is

497

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498

supported by the arytenoid cartilages. In E.
polymniae, this disc is equal to 45% of the
head width, whereas in male E. spatulatus
it is only about 22% of the head width.

A number of species of the genus Eleu-
therodactylus occur on the mesic northern
slopes of the Sierra Juarez, including several
species apparently in the a/fredi group. With
E. polymniae, the number of species in this
group known to inhabit the Sierra Juarez
now stands at three. Eleutherodactylus al-
frediis known from the lower portion of the
range to elevations a little over 600 m, E.
polymniae is known only from the type lo-
cality at 1420 m, and E. spatulatus has been
collected at elevations from 1621 to 2235
m. Bogert (1969) found the latter species at
approximately 1900 m in the Sierra Juarez.
Although E. spatulatus has been collected
to the north, as well as to the south, of the
type locality of E. polymniae (Fig. 4), the
northern record was from a hillside at 1621
m, the lowest elevational record for the
species in the Sierra Juarez.

Eleutherodactylus bufonoides was de-
scribed as belonging to the a/fredi group from
‘““above Vista Hermosa,” in the Sierra Juar-
ez (Lynch 1965). This name was subse-
quently synonymized with E. spatulatus
(Lynch 1970b), an action with which we
concur. The holotype of E. bufonoides is a
male with a SVL of 29.7 mm and has none
of the distinguishing pattern on the groin
and thigh that characterizes E. polymniae.
The specimen was collected at 1700 m, an
elevation where we have encountered E.
spatulatus.

Acknowledgments

We thank the officials from the Direccion
General de Flora y Fauna Silvestre for is-
suing collecting permits for México. Greg-
ory S. Willis competently rendered the il-
lustrations of Fig. 3.

Literature Cited

Bogert,C. M. 1968. Anew genus and species of dwarf
boa from southern Mexico.—American Mu-
seum Novitates 2354:1-38.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

. 1969. The eggs and hatchlings of the Mexican
leptodactylid frog Eleutherodactylus decoratus
Taylor.—American Museum Novitates 2376:
1-9.

, & W.E. Duellman. 1963. A new genus and

species of colubrid snake from the Mexican state

of Oaxaca.— American Museum Novitates 2162:

1-15.

Caldwell, J. 1974. A re-evaluation of the Hyla bi-
stincta species group, with descriptions of three
new species (Anura: Hylidae).— Occasional Pa-
pers of the Museum of Natural History, The
University of Kansas 28:1-37.

Campbell, J. A. 1982. Anew species of Abronia (Sau-
ria, Anguidae) from the Sierra Juarez, Oaxaca,
México. — Herpetologica 38(3):355-361.

Dixon, J. R. 1957. Geographic variation and distri-
bution of the genus Tomodactylus in Mexico. —
The Texas Journal of Science 9(4):379-409.

Duellman, W.E. 1960. A distributional study of am-

phibians of the Isthmus of Tehuantepec, Méx-

ico.— University of Kansas Publications, Mu-
seum of Natural History 13(2):19-72.

1962. A new species of fringe-limbed tree
frog from México. Studies of American hylid
frogs VIII.—Transactions of the Kansas Acad-
emy of Science 64(4):349-352.

. 1965. Anew species of tree frog from Oaxaca,

México. — Herpetologica 21(1):32-34.

Good, D. A., & K. Schwenk. 1985. A new species of
Abronia (Lacertilia: Anguidae) from Oaxaca,
Mexico.—Copeia 1985(1):135-141.

Johnson, J. D. 1974 [1973]. New records of reptiles

and amphibians from Chiapas, México.—

Transactions of the Kansas Academy of Science

76(3):223-225.

,C. A. Ely, & R.G. Webb. 1976. Biogeograph-

ical and taxonomic notes on some herpetozoa

from the northern highlands of Chiapas, Mex-
ico.— Transactions of the Kansas Academy of

Science 79(3-4):131-139.

Karges, J. P., & J. W. Wright. 1987. A new species
of Barisia (Sauria, Anguidae) from Oaxaca,
Mexico.—Contributions to Science, Natural
History Museum of Los Angeles County 381:
1-11.

Lee, J. C. 1980. An ecogeographic analysis of the
herpetofauna of the Yucatan Peninsula.— Mis-
cellaneous Publication. Museum of Natural
History, University of Kansas 67:1-75.

Lynch, J. D. 1965. Two new species of Eleuthero-

dactylus from Mexico (Amphibia: Leptodactyl-

idae). — Herpetologica 20(4):246-252.

1966. A new species of Eleutherodactylus
from Chiapas, Mexico (Amphibia: Leptodac-
tylidae).— Transactions of the Kansas Academy
of Science 69(1):76-78.

. 1967a. Two new species of Eleutherodactylus

from Guatemala and Mexico (Amphibia: Lep-

VOLUME 102, NUMBER 2

todactylidae).— Transactions of the Kansas
Academy of Science 70(2):177—183.

1967b. Two new Eleutherodactylus from
western Mexico (Amphibia; Leptodactyli-
dae).— Proceedings of the Biological Society of

Washington 80:211-217.

. 1967c. Synonymy, distribution and variation

in Eleutherodactylus decoratus of Mexico (Am-

phibia: Leptodactylidae).— Transactions, IIli-

nois State Academy of Science 60(3):299-304.

. 1968. Genera of Leptodactylid frogs in Méx-

ico.— University of Kansas Publications, Mu-

seum of Natural History 17(11):503-515.
1970a. A taxonomic revision of the lepto-

dactylid frog genus Syrrhophus Cope.—Univer-

sity of Kansas Publications, Museum of Natural

History 20(1):1-45.

1970b. Taxonomic notes on some Mexican
frogs (Eleutherodactylus: Leptodactylidae). —

Herpetologica 26(2):172—180.

Martin, P. S. 1958. A biogeography of reptiles and
amphibians in the Gomez Farias region, Ta-
maulipas, Mexico.— Miscellaneous Publica-
tions, Museum of Zoology, University of Mich-
igan 101:1-102.

Regal, P. J. 1966. A new plethodontid salamander
from Oaxaca, Mexico.—American Museum
Novitates 2266:1-8.

Shannon, F. A., & J. E. Werler.

1955. Notes on am-

499

phibians of the Los Tuxtlas Range of Veracruz,
Mexico. — Transactions of the Kansas Academy
of Science 58(3):360-386.

Smith, H. M. 1968. Another new lizard from Mexico
of the schiedi group of Anolis.—The South-
western Naturalist 13(3):353-372.

—., & R. L. Holland. 1969. Two new snakes of
the genus Geophis from Mexico. — Transactions
of the Kansas Academy of Science 72(1):47-53.

— , & E.H. Taylor. 1948. An annotated checklist
and key to the Amphibia of Mexico. — Bulletin
of the United States National Museum 194:1-
118.

Stuart, L. C. 1941. Two new species of Eleuthero-

dactylus from Guatemala.— Proceedings of the

Biological Society of Washington 54:197—200.

. 1963. Achecklist of the herpetofauna of Gua-

temala.— Miscellaneous Publications, Museum

of Zoology, University of Michigan 122:1—150.

(JAC) Department of Biology, The Uni-
versity of Texas at Arlington, Arlington,
Texas 76019; (WWL) Department of Biol-
ogy, The University of Texas at Tyler, Tex-
as 75701; (DMH) Department of Zoology,
The University of Texas at Austin, Austin,
Texas 78712.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 500-506

PHYSALAEMUS CROMBIEI
(AMPHIBIA: LEPTODACTYLIDAE), A NEW
FROG SPECIES FROM ESPIRITO SANTO, BRAZIL WITH
COMMENTS ON THE P. SIGNIFER GROUP

W. Ronald Heyer and Alan J. Wolf

Abstract. — Physalaemus crombiei, a new species of the P. signifer group, is
described from the State of Espirito Santo, Brazil. Members of this group are
most easily distinguished from each other by advertisement call, although
morphological differences also exist. The relationships and distributions of
members of this species group are not well understood at present.

One species group of frogs of the genus
Physalaemus breeds in forest puddles and/
or swamps within the Atlantic Forest sys-
tem of Brazil. Members of this group are
more easily distinguished from each other
by advertisement calls than by their mor-
phologies. Perhaps because of this, several
species have been described since field re-
cording equipment has become more avail-
able. We describe another new species of
this group, to be known as:

Physalaemus crombiel, new species
Fig. 1

Holotype.—MZUSP 66252, adult male
from Brazil: Espirito Santo; adjacent to Re-
serva Biologica Nova Lombardia, near San-
ta Teresa, approximately 19°55’S, 40°36’W.
Collected on 31 Dec 1977 by one of collec-
tors of paratopotypes.

Paratopotypes.—MZUSP 66253-66284,
USNM 285038—285070, taken by Ronald
I. Crombie, Christina Duchéne, W. Ronald
Heyer, and Francisca C. do Val from 28 Dec
1977 toptayan 197.8.

Diagnosis.—Physalaemus crombiei is a
member of the P. signifer group. The sig-
nifer group differs from other Physalaemus
species groups (Lynch 1970) by the follow-
ing characteristics: absence of an inner tar-
sal tubercle, first finger shorter than second,

lack of parotoid glands, small to large in-
guinal glands, slender build and a size range
of 15-35 mm SVL.

Physalaemus crombiei differs from P. dei-
maticus Sazima and Caramaschi 1986
(which apparently does not belong to any
of Lynch’s (1970) species groups) by the fol-
lowing characteristics: relatively smooth
skin (granular in deimaticus), slender build
(stocky), and inverse V-shaped marks on
back (absent).

Physalaemus crombiei can be distin-
guished from other members of the P. sig-
nifer group [P. signifer (Girard, 1853), P.
olfersi (Lichtenstein & Martens, 1856), P.
nanus (Boulenger, 1888), P. maculiventris
(A. Lutz, 1925) (Frost 1985:253, is in error
in placing maculiventris in the cuvieri group),
P. obtectus Bokermann, 1966, P. boker-
manni Cardoso & Haddad, 1985, and P.
franciscae Heyer, 1985] as follows. Physa-
laemus crombiei differs from bokermanni,
obtectus, olfersi, and signifer by size (P.
crombiei 19-22 mm SVL, bokermanni 15—
17 mm, obtectus 22—27 mm, olfersi 30-35
mm, signifer 24-31 mm). Physalaemus
crombiei also differs from olfersi by the ab-
sence ofa light outline on the ventral border
of the lateral stripe from the eye to the arm
(present in o/fersi). It is distinguished from
bokermanni, maculiventris and nanus by the
presence of tubercles on the sole of the foot

VOLUME 102, NUMBER 2

(absent). Physalaemus crombiei differs from
obtectus by the lack of numerous small round
markings closely associated with the inverse
V-shaped marks on the back, by its sharp
canthus rostralis, and pointed snout (nu-
merous small round marks associated with
the dorsal pattern, rounded canthus ros-
tralis, less pointed snout in obtectus). Phys-
alaemus crombiei has a relatively light col-
ored posterior belly differing from the boldly
dark and light mottled posterior belly of
franciscae and maculiventris.

Description of holotype. —Snout subellip-
tically pointed from above, rounded in pro-
file; canthus rostralis rather sharp in cross
section; tympanum barely discernible ex-
ternally, large, about two-thirds eye diam-
eter; maxillary teeth not visible, but dis-
cernible by probe; vomerine teeth absent;
vocal slits present, elongate; vocal sac sin-
gle, noticeably but not greatly expanded ex-
ternally, extending to level in front of arms;
finger tips slightly expanded, not disked; fin-
gers lacking fringe and web; subarticular tu-
bercles large, globose, pungent; pair of tan
nuptial pads on inner surface of each thumb,
rounded-oblong medial pad associated with
inner metacarpal tubercle, narrowly sepa-
rated from elongate distal pad; no prepollex;
no ulnar ridge; forearm slightly hypertro-
phied; extremely weak supratympanic fold;
no dorsolateral fold: large ovate lumbar
gland in groin; dorsal skin texture smooth
in appearance but with scattered small fleshy
tubercles, especially on upper eyelid; pos-
terior part of vocal sac wrinkled, rest of
throat and belly smooth, under surfaces of
thighs areolate; toe tips slightly expanded,
toes II] and IV most expanded: toes without
webs, sides of toes with or without weak
ridges; subarticular tubercles globose,
rounded, pungent; well-developed rounded
metatarsal tubercle about half the size of
well-developed oblong inner metatarsal tu-
bercle; short, weakly developed inner tarsal
fold on distal one-fourth of tarsus, narrowly
separated from inner metatarsal tubercle,
remainder of tarsus smooth: heel smooth;

501

foot with four series of small but distinct
fleshy tubercles.

Color in preservative: Dorsum creamy tan
with indistinct brown markings including
an inversed V-shaped mark in shoulder area,
an indistinct interorbital mark, a pair of
acute marks in sacral area, and a pair of
very distinct black, white-outlined spots, one
on each upper portion of lumbar glands;
sides of head mottled brown, distinctly sep-
arated along canthus rostralis from lighter
dorsal color and leaving a light stripe on
very tip of snout; tympanum brown, distin-
guished from surrounding areas by color
pattern; distinct dark (almost black) supra-
tympanic stripe extending along flank to-
wards belly, terminating below mid-lumbar
gland level, distinct dorsal dark stripe with
white outline, brief white outline from pos-
terior tympanum midway to shoulder ven-
trally, rest of stripe blending into ventral
pattern and not distinct ventrally; upper limb
surfaces indistinctly cross-banded, fore-
arms with darker brown blotch medially,
each outer shank with series of three black-
ish spots; throat dark brown with small, light
irregular spots; anterior half of belly mottled
brown and cream, posterior belly cream;
ventral forearm region dark brown laterally,
irregularly but distinctly separated from
lighter dorsal color pattern; ventral surface
of thighs mottled brown and cream, lighter
medially, posterior portions with small white
dots; outer tarsus brown medially with ir-
regular dark brown elongate markings cen-
trally with lighter dorsal color laterally; sole
of foot brown, subarticular and metatarsal
tubercles cream; light dorsal pattern dipping
irregularly but with distinct border onto
posterior surface of each thigh, lowest on
medial half of thigh, remainder of lower
portions of thigh almost uniform brown,
with a few ventral white dots.

Snout—vent length 20.3 mm, greatest
tympanum diameter 1.7 mm, head length
7.0 mm, head width 7.7 mm, thigh length
9.3 mm, shank length 9.3 mm, foot length
9.7 mm.

502 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Pig 1:
views.

Variation.—The dorsal patterns range
from almost uniform brown to distinct dark
interorbital bands and two dark, white-out-
lined confluent arrows, one on the mid-
scapular area, the other on the mid-sacral
area. In a few specimens the interorbital and
arrow markings join. In some individuals a
distinct small mid-scapular light spot is
present. One individual (USNM 285048)
has a faint but distinct light mid-dorsal pin
stripe from the tip of the snout to just past
the sacrum. Most individuals lack a light
border posterior to the tympanum defining
a portion of the ventral border of the dark
lateral stripe. The stripe is distinct (black)
in most individuals, and fainter but distinct
in a few. The dark spots on the lumbar glands
are distinct in all individuals. The ventral
pattern described for the holotype is char-
acteristic of nearly all individuals, although
a couple have lighter bellies than does the
holotype. In some individuals the dorsal and
posterior thigh patterns are distinct only
medially, near the vent; most have the same
pattern as described for the holotype.

Holotype of Physalaemus crombiei (MZUSP 66252, SVL = 20.3 mm, male), dorsal and ventral

The following measurements are based
on 7 adult male (listed first) and 14 adult
female (listed second) paratopotypes: SVL
18.9-20.0 mm, 19.2—21.8 mm; head length
33-38% SVL (mean 35%), 32-36% SVL
(mean 34%); head width 35-39% SVL (mean
37%), 33-36% SVL (mean 35%); thigh length
43-48% SVL (mean 46%), 42-46% SVL
(mean 44%); shank length 45-48% SVL
(mean 47%), 43-48% SVL (mean 45%); foot
length 44-49% SVL (mean 47%), 40-46%
SVL (mean 44%).

Color in life. —Brief color notes are avail-
able for two individuals. In both, the iris
was bronze. In USNM 285051 a slight red
wash occurred in the groin and front and
back of the thighs. In USNM 285052 the
groin and belly had a pinkish cast and the
dorsum was red-brown and brown.

Advertisement call.—Data from a series
of calls presumably given by a single un-
captured individual; another captured in-
dividual (USNM 285070) was heard to give
a similar call. Call rate approximately one/
sec; average call duration 0.376—-0.418 sec

VOLUME 102, NUMBER 2

503

eed coed ae ede ee
| = SSS SS SS

= a
— _ == _ SY SY

=e ee

KILOHERTZ

SECONDS

Fig. 2. Advertisement call of Physalaemus crombiei. Wave form of second call shown on audiospectrogram,
total signal length 0.428 sec. Recorded (no voucher) by W. R. Heyer, 30 December 1977, air temperature 21.5°C

(USNM Tape 7, Cut 5).

(mean 0.399 sec); interval between calls
0.521-0.651 sec (mean 0.574 sec); 8-10
notes per call (mean 8.6); note length (other
than final note) 0.029-0.043 sec (mean 0.039
sec), final note length 0.043-0.088 sec (mean
0.061); notes weakly to markedly pulsatile,
for pulsed notes, 6-16 pulses/note (mean
13.7), pulse rate 344-454 pulses/sec (mean
394), first notes often not pulsatile, final note
pulsatile initially and unpulsed at end; fre-
quency range of call about 690-1840 Hz,
dominant frequency about 1060 Hz with a
second strong frequency band at about 1450
Hz; harmonic structure not apparent (Fig.
ya):

Etymology.—Named for Ronald I.
Crombie, who not only helped collect the
type series of this new species, but has added
to our knowledge of the Brazilian herpeto-
fauna in general through his determined col-
lecting efforts.

Distribution. —Known only from the type
locality.

Discussion

Our knowledge of members of the Phys-
alaemus signifer group is characterized
overwhelmingly by what we do not know
rather than any real understanding of the
relationships among members of this group

(and other species within the genus) or their
distributions. Of critical importance is the
number of species contained in the signifer
group. At first glance, the group seems to
represent a modest radiation. However,
three of the currently recognized species were
described prior to 1900, one was described
in 1925, another in 1966, and including P.
crombiei, three have been described since
1985. No doubt several more species in this
group are unnamed. Accordingly, we think
it premature to attempt to resolve the re-
lationships not only within the signifer
group, but also within the genus and concur
with Cannatella & Duellman (1984) on this
point.

Certain points resulting from our studies
deserve comment.

Definition of the Physalaemus signifer
group definition was unsatisfactory to Lynch
(1970:493—494) who reluctantly placed ol-
fersi in the signifer group. All signifer group
species described since Lynch’s 1970 defi-
nition are more similar externally to other.
signifer group members than they are to ol-
fersi. One character, presence (o/fersi) or ab-
sence (remaining species) of teeth, used by
Lynch no longer separates o/fersi from the
other species. Teeth are visible (under a dis-
secting microscope) in P. o/fersi. In all other
species of the signifer group, teeth are not

504

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Comparison of call characteristics for members of the Physalaemus signifer group. Call unavail-

able for P. nanus.

Calls Calls
with with Rate of smallest discrete Broadcast
Call duration notes pulses sound structure frequency range Source
bokermanni 0.25 =F uy 7 notes/s 2500-5200 Hz Cardoso & Haddad 1985
crombiei 0.45 se =F 390 pulses/s 690-1840 Hz - This study
franciscae 0.5-0.7 s af =F 250 pulses/s 600-1600 Hz Heyer et al. 1989
maculiventris-

Boracéia 0.2-0.8 s = + 130-170 pulses/s 700-3700 Hz Heyer et al. 1989
maculiventris-

Teresopolis Os = =F 120 pulses/s 860-2340 Hz This study
obtectus 0.1 s =F =F 250-300 pulses/s 1100-1500 Hz Bokermann 1966
olfersi 3.4-4.0 s = si 200-225 pulses/s 1000-2700 Hz Heyer et al. 1989
signifer 0.3 s = ae 300-350 pulses/s 700-2500 Hz Bokermann 1966

visible and are either very small or absent.
Teeth can be felt by probe in all individuals
of bokermanni, franciscae, and obtectus. No
teeth can be felt by probe in maculiventris
and nanus individuals. The condition is
variable within crombiei and conflicting lit-
erature reports and our observations suggest
the same might be true for signifer. As teeth
may be present and still not felt by probe,
final resolution of the toothed condition will
require extensive clearing and staining
preparations. It is clear with the evidence
at hand that the difference in tooth condi-
tion between o/fersi and the other group
members is qualitative, not discrete, how-
ever.

Each of the character states used by Lynch
(1970) to define the signifer group could be
primitive, as pointed out by Cannatella &
Duellman (1984); thus there is no assurance
that the signifer group is monophyletic. With
the exception of o/fersi, it is our intuitive
impression that the remaining species com-
prise a monophyletic group. The extreme
morphological similarity and available hab-
itat and reproductive data indicating inti-
mate association with Atlantic Forests and
small forest breeding pools support this
view. Without additional evidence, it is in-
appropriate to remove o/fersi from the sig-
nifer group, however.

As considerable confusion exists con-

cerning the identity of Physalaemus signifer,
an explanation of our use and understand-
ing of this species is in order. Rhinoderma
signifera Girard was described from Rio de
Janeiro (Girard 1853). We have no reason
to contradict Bokermann’s (1962) interpre-
tation that the locality represented the city
of Rio de Janeiro rather than being a general
locality for the region where collections were
made while the expeditionary ship was in
the port of Rio de Janeiro. With Boker-
mann’s (1962) removal of maculiventris
from the synonymy of signifer and the de-
scription of franciscae (Heyer, 1985), our
understanding is that signifer is the rela-
tively large species of the group presently
known only from around the city of Rio de
Janeiro and the Organ Mountains. B. Lutz
(e.g. 1954) clearly thought that a single
species inhabited both the city of Rio de
Janeiro and the Organ Mountains, although
she used the name bress/aui, described by
Muller (1924) from Teresopolis, Organ
Mountains for the taxon involved. We fol-
low Bokermann (1962) in considering bres-
slaui a synonym of signifer. The anuran fau-
na from Rio de Janeiro and the nearby coastal
lowland differs from the Organ Mountain
anuran fauna (examples in Heyer 1983,
1984). The only published call information
for signifer is from Itaguai, a coastal locality
near Rio de Janeiro (Bokermann 1966). If

VOLUME 102, NUMBER 2

505

N
Ee
oe
WJ
=x
ro)
=
3
SECONDS
Wh tlt hed bs ,
WW ayn \ Aedeagal + ¥ ATA Ma as Aen ee ae et
N
E
x
lJ
a
re)
a
x
SECONDS
Fig. 3. Advertisement calls of Physalaemus maculiventris. Upper set of figures recorded from Boracéia, Sao

Paulo, Brazil; wave form of call on left in audiospectrogram, total signal length 0.428 sec; recorded from USNM
209278 by W. R. Heyer, 13 December 1976, air temperature 17.6°C (USNM Tape 2, Cut 7). Lower set of figures
recorded from TeresOpolis, Rio de Janeiro, Brazil; wave form of call on left in audiospectrogram, total signal
length 0.428 sec; recorded from series of specimens, USNM 285076-78 and field numbers 6609-16 to be
catalogued in MZUSP collections, by W. R. Heyer, 20 December 1977, air temperature 18—20°C (USNM Tape

#5 Eut'3).

the Organ Mountains population proves to
be distinct at the species level, the name
bresslaui is available.

The distributions of signifer group mem-
bers are poorly known. Physalaemus olfersi
stands out in having a rather broad distri-
bution within the Atlantic Forest region
from the State of Santa Catarina to at least
the State of Espirito Santo (based on ex-
amination of museum specimens by WRH).
In contrast, P. bokermanni, franciscae, ob-
tectus, and crombiei currently are known
only from their type localities or nearby
areas. Physalaemus maculiventris and nan-
us apparently have moderate distributions,
but the situation with maculiventris requires
comment.

In the introduction, we stated that the

known advertisement calls of members of
the signifer group were distinctive (see Ta-
ble 1). We have two samples of calls of ma-
culiventris-like animals, one from Boracéia,
Sao Paulo, and one from Teresopolis, Rio
de Janeiro. The available recordings show
differences between calls from the two sites
(Table 1, Fig. 3), but not of the magnitude
found among other species in the group (Ta-
ble 1). We also found difficult to verbalize
subtle differences in the adult color patterns
of the two populations, involving the rela-
tive clarity of a light ventral outline of the
dark lateral stripe in the eye-shoulder region
and the appearance of the dark lateral stripe
itself. We have no geographically interme-
diate samples at hand. Two specimens of
maculiventris were collected with the type

506

series of crombiei; with only two specimens,
we are unable to determine any population-
al differences between these and the speci-
mens from the Teresopolis or Boracéia pop-
ulations. Our present conclusion is that the
variation observed in calls and morphology
is best considered intraspecific. Thus, it
would appear that some signifer group
species have broad distributions without
notable differentiation, others have mod-
erate distributions with some geographic dif-
ferentiation, but most apparently have very
localized distributions. Clearly, in order to
understand the systematics and distribu-
tions of species that have been included in
this group, much more field work, especially
with voice recordings, is necessary.

Acknowledgments

Stanley H. Weitzman, Smithsonian In-
stitution (USNM), kindly prepared the pho-
tographs used in Fig. 1. Ronald I. Crombie,
Smithsonian Institution, critically read the
manuscript. Research leading to this paper
was supported, in part, by the Museu de
Zoologia da Universidade de Sao Paulo
(MZUSP) through the kind offices of P. E.
Vanzolini and the I. E. S. P. Neotropical
Lowland Research Program, Smithsonian
Institution.

Literature Cited

Bokermann, W.C. A. 1962. Notas sobre trés espécies
de Physalaemus (Amphibia, Salientia, Lepto-
dactylidae).— Anais da Academia Brasileira de
Ciéncias 34:563-568.

1966. Dos nuevos especies de Physalaemus
de Espiritu Santo, Brasil (Amphibia, Leptodac-
tylidae).— Physis 26:193-202.
Cannatella, D. C., & W. E. Duellman.

1984. Lepto-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

dactylid frogs of the Physalaemus pustulosus
group.—Copeia 1984:902-921.

Cardoso, A. J., & C. F. B. Haddad. 1985. Nova es-
pecie de Physalaemus do grupo signiferus (Am-
phibia, Anura, Leptodactylidae).— Revista
Brasileira de Biologia 45:33-37.

Frost, D. R. (ed.). 1985. Amphibian species of the
world. Allen Press, Inc., and The Association of
Systematics Collections, 732 pp.

Girard, C. 1853. Descriptions of new species of Rep-
tiles, collected by the U.S. Exploring Expedition,
under the command of Capt. Charles Wilkes,
U.S. N. Second Part.—Including the species of
Batrachians, exotic to North America.—Pro-
ceedings of the Academy of Natural Sciences of
Philadelphia 6:420—424.

Heyer, W.R. 1983. Variation and systematics of frogs

of the genus Cycloramphus (Amphibia, Lepto-

dactylidae).— Arquivos de Zoologia 30:235-339.

1984. Variation, systematics, and zoogeog-
raphy of Eleutherodactylus guentheri and closely
related species (Amphibia: Anura: Leptodactyl-
idae).—Smithsonian Contributions to Zoology
402:1-42.

1985. New species of frogs from Boracéia,
Sao Paulo, Brazil.— Proceedings of the Biolog-
ical Society of Washington 98:657-671.

, A. S. Rand, C. A. G. da Cruz, O. L. Peixoto,

& C. E. Nelson. 1989. The frogs of Boracéia.

Arquivos de Zoologia (in press).

Lutz, B. 1954. Anfibios anuros do Distrito Federal. —
Memorias do Instituto Oswaldo Cruz 52:155-
238.

Lynch, J.D. 1970. Systematic status of the American
leptodactylid frog genera Engystomops, Eupem-
phix, and Physalaemus.—Copeia 1970:488-496.

Miller, L. 1924. Neue Batrachier aus Ost-Brasi-
lien. —Senckenbergiana 6:169-177.

(WRH) Amphibians & Reptiles, NHB
stop 162, Department of Vertebrate Zool-
ogy, Smithsonian Institution, Washington,
D.C. 20560; (AJW) Biology Department,
University of Wisconsin—Stevens Point,
Stevens Point, Wisconsin 54481.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 507-514

REDESCRIPTION OF THE GARTER SNAKE
THAMNOPHIS EXSUL ROSSMAN, 1969
(SERPENTES: COLUBRIDAE)

Douglas A. Rossman, Ernest A. Liner, Carlos H. Trevino, and
Alan H. Chaney

Abstract.— Thamnophis exsul, previously known only from the holotype from
Coahuila, is redescribed on the basis of nine recently collected specimens from
Nuevo Leon. An ontogenetic change in dorsal color pattern is apparent in the
Nuevo Leon sample. Notes on the habitat and associated herpetofauna are also

presented.

Thamnophis exsul was described by
Rossman (1969) from a single specimen col-
lected 17.7 km E, 5.6 km S San Antonio de
las Alazanas, Coahuila, Mexico, by M. D.
Sabath on 6 Jul 1961. Subsequent attempts
to find additional specimens at or near the
type locality by field crews from Louisiana
State University and Texas A&M Univer-
sity, aS well as by E. A. Liner, were unsuc-
cessful. In the summer of 1979, Trevino
sent Rossman two puzzling garter snakes
(Universidad Autonoma de Nuevo Leon,
UANL 2250-51) that had been collected in
southern Nuevo Leon the previous fall and
tentatively identified as 7. chrysocephalus.
Rossman soon concluded that, despite
striking differences in color pattern from the
holotype of T. exsul (Fig. 1), the Nuevo Leon
snakes were more closely related to exsul
than to any other known species of Tham-
nophis. Subsequent collecting in southern
Nuevo Leon by E. A. Liner, A. H. Chaney,
and the late R. M. Johnson has yielded sev-
en additional specimens; these animals shed
new light on the previously noted pattern
differences, and it now seems almost certain
that the Coahuila and Nuevo Leon snakes
are conspecific. The purpose of this article
is to redescribe what remains the rarest
species of garter snake.

Thamnophis exsul Rossman, 1969
Figs. 1-2

Specimens examined. —Coahuila: 17.7
km E, 5.6 km S San Antonio de las Alazanas
(holotype, National Museum of Natural
History, USNM 166423). Nuevo Leon:
Zaragoza, Rancho La Encantada (UANL
2250-51); 19.6 km NE San Antonio de Pena
Nevada (Ernest A. Liner, EAL 4820, 4837;
Louisiana State University Museum of Zo-
ology 43836); 17.9 km NE San Antonio de
Pena Nevada (EAL 4971, 4 specimens).

Definition. —The smallest species of
Thamnophis (maximum recorded snout-—
vent length = 360 mm), 7. exsul is char-
acterized by having: a maximum of 17 dor-
sal scale rows (in 90% of specimens), usually
with no posterior reduction to 15 rows
(80%); 7 supralabials on at least one side of
the head (90%); 8 infralabials on at least one
side (90%); 142—150 ventrals in 92, 152-156
in 66; 52-56 subcaudals in 2°, 63-65 in 3é;
tail length 18.9-20.0% of total length in 99,
21.8—22.2% in 36; 19-21 maxillary teeth,
the last 2 or 3 slightly enlarged; the dorsum
almost uniformly putty brown with 3 in-
distinct light stripes, or having 3 or 4 alter-
nating rows of spots or blotches with the
light stripes partially or completely sup-
pressed.

508

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Selected head scale proportions in adult Thamnophis exsul. All ratios expressed as a percent.

Character

Internasal-rostral contact/nasal-rostral contact
Prefrontal suture length/internasal suture length
Muzzle width'/muzzle length?

Muzzle length/frontal length

Eye diameter/frontal length

Frontal length/parietal length

Frontal width/frontal length

Frontal width posterior*/frontal width anterior

Loreal length dorsal*/loreal length ventral

Sex x Range n
3d 144.8 141.7-147.9 2
29 150.0 142.6-164.3 5
3d 101.6 100.0-103.1 2
29 143.3 139.6-147.2 3
36 114.4 111.3-117.4 2
29 1205 115.1-128.2 3
3d I39 72.3-75.4 2
29 65.0 60.2-70.2 3
ye) 67.5 66.0-68.9 Z
29 58.8 57.5-60.6 5
36 ONT, 74.2-77.1 2
29 78.0 74.2-81.0 3
33 50.6 41.4-59.7 2
29 62.4 60.6-64.1 B)
ie) 9775 91.1-103.8 2
29 88.0 85.0-92.2 3
le) 63.0 60.6-65.4 2
29 60.9 58.8-62.2 3

' Combined width of internasals at posterolateral corners.
> Combined length of internasal and prefrontal medial sutures.

> Where parietals meet supraoculars and frontal.
+ Measured to prefrontal-preocular junction.

Description of holotype.—See Rossman
(1969).

Variation. — Dorsal scale rows are 17-15-
17 (1 specimen), 17-17-15 (2), 17-17-17 (6),
and 19-17-17 (1). Where posterior reduc-
tion occurs, it results from the loss of row
four at about 60% of the distance from the
first to the last ventral scute.

Ventrals average 145.0 (142-150) in 8 99,
154.0 (152-156) in 2 46; subcaudals 53.4
(52-56) in 7 22, 64.0 (63-65) in 2 46; tail as
a % of total length 19.4 (18.9-20.0) in 7 99,
22.0 (21.8—22.2) in 2 66. Total supralabials
on both sides number 12 (1 specimen), 14
(7), and 15 (2); total infralabials 16 (8), 17
Ch): and? b9:(1):

In the five snakes that lack a well-defined
umbilical scar, head length as a percent of
snout-vent length is 5.3 (237 mm SVL), 5.2
(274), 4.7 (330), 4.8 (347), and 4.6 (360),
respectively. There appears to be some on-
togenetic change in this character, the large
specimens having a proportionately shorter

head. Head scale measurements were made
on the preceding five snakes and a series of
ratios generated (Table 1). Noticeable sex-
ual dimorphism appears to be present in
several characters (prefrontal suture length/
internasal suture length, eye diameter/fron-
tal length), but the sample size is so small
that these distinctions may disappear when
data from additional specimens become
available.

In the Nuevo Leon sample, there appears
to be a marked ontogenetic change in dorsal
color pattern. The juveniles (Fig. 2, lower)
are prominently spotted in three or four al-
ternating rows, the former condition re-
sulting from the fusion of spots in the upper
two rows. The anteriormost 0-7 spots may
be vertically enlarged to extend to, or fuse
across, the vertebral scale row. The verte-
bral stripe is absent or faint and interrupted
at intervals by the dorsal spotting. There is
a prominent dark nuchal blotch that may
or may not be very narrowly divided by the

VOLUME 102, NUMBER 2 509

Fig. 1. Upper: Holotype of Thamnophis exsul from 17.7 km E, 5.6 km S San Antonio de las Alazanas,
Coahuila, Mexico. Reprinted with permission from Rossman (1969). Lower: Adult Thamnophis exsul (EAL
4837) from 19.6 km NE San Antonio de Pea Nevada, Nuevo Leon, Mexico.

510

ye ee Se ee.
A

i |

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Upper: Subadult Thamnophis exsul (LSUMZ 43836) from 19.6 km NE San Antonio de Pena Nevada,
Nuevo Leon, Mexico. Lower: Three juvenile Thamnophis exsul (EAL 4971) from 17.9 km NE San Antonio de

Pena Nevada, Nuevo Leon, Mexico.

vertebral stripe. The nuchal blotch is either
continuous with the dark dorsal coloration
of the head or narrowly separated from it
by a lighter zone (less than one scale long)
immediately posterior to the parietals. The
spotted pattern of the subadult (Fig. 2, up-

per) is less distinct than that of the juveniles,
and is barely, if at all, discernible in the
larger adults (Fig. 1, lower).

The lone example from Coahuila, an
adult, differs from the Nuevo Leon animals
in retaining the juvenile pattern, in having

VOLUME 102, NUMBER 2

larger spots, and in having total suppression
of the vertebral stripe (Fig. 1, upper).

The everted hemipenis (Fig. 3) of EAL
4837 is single with the distal half greatly
expanded. The sulcus spermaticus is simple
and terminates apically between raised lips.
The apical surface of the expanded portion
is nude. The remainder of the organ is spi-
nose, the spines being very small on the
expanded portion, but becoming larger
proximally. One enlarged basal hook les
somewhat removed from the sulcus.

Teeth average 20.3 (19-21) on six max-
illae, 12.0 (12) on two palatines, 22.0 (21-
23) on two pterygoids, and 22.5 (22-23) on
two dentaries.

Relationships. —None of the data provid-
ed by the new material contradicts Ross-
man’s 1969 conclusion that Thamnophis
exsul more nearly resembles J. sca/aris than
it does any other garter snake. A detailed
comparison of the two species awaits com-
pletion of a variational study of the 7. sca-
laris complex now in progress by Rossman.
The lack of posterior dorsal scale row re-
duction in most (8 to 10) 7. exsul is an
unusual, but not unique, condition for a
Thamnophis. Most of the 7. godmani ex-
amined thus far from Oaxaca and Puebla
have 17-17-17 rows, as do 20% of the T.
scalaris from Veracruz.

Ecological notes. — Both the Coahuila and
Nuevo Leon localities are situated in mon-
tane areas at elevations between 2650-2800
meters (Fig. 4). The adult specimens from
Puerto de Pena Nevada were found in
meadows with mixed oak, pine, madrone,
and scrub brush, the meadows being used
for grazing (Fig. 5, upper). The snakes were
found in mid-morning after the cloud cover
had lifted, but while the ground cover was
still moist. When discovered in the open,
the snakes crawled rapidly toward the scrub
cover that was never far away. The young
of the year were collected from a rotten log
on an oak-madrone hillock with scattered
logs, agave, and rocks (Fig. 5, lower). No

Fig. 3. Drawing of everted hemipenis of adult male
Thamnophis exsul (EAL 4837). All spines save the
larger ones are diagrammatically approximated.

standing water or streams were found in the
area. The habitat at Rancho La Encantada
was generally similar to that described for
the Puerto de Pena Nevada area; but the
holotype from Coahuila was found in a
grassy “sump” above the head of a steep
canyon (Rossman 1969).

During Liner’s visits to the Puerto de Pena
Nevada area, all in July, clouds would start
to roll in around 1600 h; instead of the usual
mist, a light drizzle (along with thunder and
lightning) often would accompany the clouds
and the temperature would then turn de-
cidedly colder. In the morning, the sky would
clear around 0900 h; during mid-day it was
sunny and hot.

The associated herpetofauna in the Puer-
to de Pena Nevada area was: Chiropterotri-
ton priscus, Pseudoeurycea galeanae, Cro-
talus pricei, Storeria occipitomaculata,
Thamnophis cyrtopsis, Barisia imbricata,
Eumeces brevirostris, Sceloporus grammi-
GUS. US. sealaris, S. torquatus (kimer, pers.
obs.); at Rancho La Encantada, Crotalus
lepidus, Storeria occipitomaculata, Barisia
species, Sceloporus grammicus, and S. tor-
quatus (Trevino, pers. obs.); and at the type

512 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

hes jig looe 98° 96°
eS
Ln maw
ve \
/ \
ii *S
“>
| a
: im yh
—\ ve Be
N 5 L e
a ees G a 3
Spies Ee we |
se ea a
Var TROPIC OF CANCER
---—-—~— fe Mee
( -y |
\A ) Oe << (
28 be i — 22°
BX \
5 sige ‘SS
Pic coe ON
2
. sl 20°
?
Hn Se
ile eg
c ¢ wae
1ge Statute 250 25 50 75 100 Se ae as ic ; niger
dieses ee f A ae yy
| Dy a
104° 102° 100° 98° 96° 94

Fig. 4. Map of northwestern Mexico showing known localities for Thamnophis exsul. Triangle represents

type locality in Coahuila; circles represent the Nuevo Leon localities.

VOLUME 102, NUMBER 2 oye)

Fig. 5. Upper: Habitat 19.6 km NE San Antonio de Pefia Nevada, Nuevo Leon, Mexico, where adult
specimens of Thamnophis exsul were collected. Lower: Habitat 17.9 km NE San Antonio de Pena Nevada,
Nuevo Leon, Mexico, where juvenile specimens of Thamnophis exsul were collected.

514 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

locality in Coahuila, Crotalus pricei, Barisia
imbricata, Eumeces dicei (=brevirostris),
Sceloporus grammicus, and S. jarrovi (Ax-
tell and Sabath, 1963; Liner, Rossman, and
Johnson, 1974).

Acknowledgments

We wish to express our gratitude to the
late Richard M. Johnson, who helped to
collect some of the Nuevo Leon specimens;
to Richard M. Blaney for serving as a cour-
ier to bring the UANL specimens to Ross-
man; and to the officials of the Direccion
General de la Fauna Silvestre in Mexico
City for issuing the necessary scientific col-
lecting permits to Liner over the years.

Literature Cited

Axtell, R. W., & M. D. Sabath. 1963. Crotalus pricei
miquihuanus from the Sierra Madre of Coa-
huila, Mexico.—Copeia 1963(1):161-164.

Liner, E. A., D. A. Rossman, & R. M. Johnson.
1973[1974]. Life history: Gerrhonotus (Bari-
sia) imbricatus ciliaris. —HISS News-Journal
1(6):185.

Rossman, D. A. 1969. A new natricine snake of the
genus Thamnophis from northern Mexico.—
Occasional Papers of the Museum of Zoology,
Louisiana State University (39): 1-4.

(DAR) Museum of Zoology, Louisiana
State University, Baton Rouge, Louisiana
70803; (EAL) 310 Malibou Boulevard,
Houma, Louisiana 70364; (CHT) Facultad
de Ciencias Biologicas, Universidad Auton-
oma de Nuevo Leon, San Nicolas de los
Garza, Nuevo Leon, Mexico; (AHC) De-
partment of Biology, Texas A&I University,
Kingsville, Texas 78363.

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 515-519

THE TYPE SPECIMENS OF HEKSTRA’S OWLS

M. Ralph Browning

Abstract.—The erroneous holotype designations and errors in the type lo-
calities in Hekstra’s descriptions of 24 new subspecies of New World owls are
corrected. Several scientific names are emended and taxonomic comments are
appended for a small number of Hekstra’s proposed taxa.

Twenty-four new subspecies of New
World owls of the genus Otus were named
in a paper by Hekstra (1982b) issued 17
December 1982. These same new names
also appeared in Hekstra’s (1982a) unpub-
lished thesis. The holotypes of the new taxa
are in 10 different collections, most of which
are in the United States. Incorrect infor-
mation was published with regard to mu-
seum designation, museum number, and
collecting locality of many of the holotypes.
I here list the holotypes with their correct
specimen label data.

The species names are here presented in
the sequence proposed by Marshall & King
(1988). The subspecies are generally ar-
ranged from north to south. The type lo-
calities given are standardized, and the
spellings are corrected where required. Ab-
breviations for museum designations are
given under acknowledgments. The num-
bers following the names refer to the pages
where the descriptions were given (Hekstra
1982b). Several of the scientific names pro-
posed were spelled incorrectly and these
have been emended in accord with Article
31c and Appendix D of the International
Code (International Commission on Zoo-
logical Nomenclature 1985). Taxonomic
comments are appended for a small number
of Hekstra’s proposed taxa. The remaining
forms he named require further study.

Otus flammeolus borealis, p. 56

Holotype. —MVZ 101700, adult female,
Penticton, Okanagan Valley, British Co-

lumbia, shore of Okanagan Lake, 22 Oct
1901, collected by Allan Brooks (original
number 1818).

Remarks. — The catalogue number of the
holotype was given as “10,700.” The type
locality, as published, did not include Oka-
nagan Lake. Additional specimen label data
are the collector’s original number and that
the holotype was found dead, with the spec-
imen label notation “‘probably killed two
weeks before” (N. K. Johnson, in litt.).

Otus flammeolus frontalis, p. 56

Holotype.—USNM 119650, female, Estes
Park, Colorado, 20 Jun 1890, collected by
W. G. Smith.

Remarks. —Hekstra (1982b) stated that
the holotype was a breeding individual but
the specimen label does not so indicate. This
form is a recognizable subspecies (J. T. Mar-
shall, pers. comm.).

Otus flammeolus meridionalis, p. 55

Holotype. —MVZ 109708, adult male, 25
Aug 1938, Cuapongo, Guerrero, Mexico,
collected by W. W. Brown.

Remarks.—The catalogue number was
published as “10978.” The original label of
the holotype indicates that the testes were
minute (N. K. Johnson, in litt.).

Otus seductus colimensis, p. 61

Holotype.—LSUMZ 39853, adult male,
7 miles south of Colima, Colima, [Mexico],

516

24 Dec 1958, collected by W. J. Schaldach,
Jt

Remarks.—This specimen was number
7001 in the Sheffler Collection, and on an
additional label is recorded as “‘S. no. 4808,”
which presumably is Schaldach’s personal
catalogue number (J. V. Remsen, in litt.).

Otus trichopsis inexpectus, p. 58

Holotype. —USNM 152828, adult fe-
male, Porto Jimenz, Costa Rica, 30 May
1892, collected by Verrill.

Remarks. —Hekstra (1982b) gave the
museum designation as Chicago (=Field)
Museum of Natural History and the cata-
logue number as “52828.” The primaries of
the holotype are clipped and no doubt the
specimen was a Captive bird. The specimen
is an example of Otus guatemalae in the red
phase (J. T. Marshall, pers. comm.).

Otus choliba caucae, p. 60

Holotype. —RMNHL 8091, [adult] male,
El Tambo, 5100 ft., Rio Cauca, Colombia,
25 June 1938 [collected by Kjell von
Schneidern].

Remarks. — There is no indication of the
age or collector on the specimen label of the
holotype but the published information (as
above) is correct (G. F. Mees, in litt.).

Otus choliba montanus, p. 61

Holotype. —BM(NH) 1914.11.22.219,
adult male, Valle, Montana Sierra, 2000 m,
Mérida, [Venezuela], 15 Apr 1909, collect-
ed by S. Briceno Gabaldon.

Otus choliba kelsoi, p. 61

Holotype. —AMNH 59483, adult male,
Princes Town, Trinidad, 10 Mar 1893, col-
lected by Frank M. Chapman.

Remarks. —In naming this form Hekstra
(1982b) stated that kelsoi “‘replaces Otus
choliba portoricensis Kelso ... preoccu-
pied.””’ However, Kelso (1942) clearly did
not introduce a new name but only referred
the population from Trinidad to the name

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Scops portoricensis Lesson, 1831. Hekstra
(1982b) proposed ke/soi as the name of a
new taxon (Greenway 1987). Hekstra
(1982a) spelled the specific locality as
“Princetown” which is the same as on the
specimen label. Modern gazetteers spell the
locality as Princes Town.

Otus choliba guyanensis, p. 60

Holotype. —BM(NH) 1888.7.20.63, adult
male, Mt. Roraima, 3500 ft., Guyana; 9 Jan
1884, collected by H. Whitely, Jr. (P. R.
Colston, in litt.).

Remarks.—The catalogue number was
published as “*88.7.20.63.”? The type local-
ity of guyanensis is the same as that for
Scops roraimae Salvin, 1897 (=Otus gua-
temalae roraimae, cf. Peters 1940), a name
based on topotype 1888.7.20.62 that was
collected by H. Whitley, Jr., on 1 Oct. 1883
(Warren 1966).

Otus choliba caatingensis p. 59

Holotype. —FM 191645, adult male, Jan-
auba, northern Minas Gerais, [Brazil], -- Jul
1949, collected by Ricardo Medeiros Berla.

Otus choliba chapadensis, p. 59

Holotype. —AMNH 34595, adult female,
Chapada, Mato Grosso, [Brazil], 9 Feb 1883,
collected by H. H. Smith.

Remarks.—The holotype of O. atricap-
illus fulvescens Hekstra, AMNH 34597, was
collected by Smith from the same locality
as the holotype of chapadensis (see beyond).

Otus choliba uruguaiensis, p. 59

Holotype. -AMNH 769736, adult fe-
male, Arroyo, Urugua-i, km 10, Misiones,
Argentina, 29 May 1958, collected by W.
H. Partridge.

Remarks. — Hekstra (1982b) proposed the
trinomial as “urugaii” and under the head-
ing of the derivation he spelled the name as
“uruguail.’’> The AMNH catalogue number
was given by Hekstra (1982b) as “76736”
(see Greenway 1987). Hekstra (1982b) gave

VOLUME 102, NUMBER 2

the collecting locality as “Argentina, Mi-
siones, Arroyo, River Uruguai, km 10.”

Otus choliba alilicuco, p. 59

Holotype. —BM(NH)_ 1899.1.27.236,
adult male, Rosario, Province Salta, Argen-
tina, 15 Sep 1896, collected by Gesling, for-
merly R. P. Moreno collection (P. R. Col-
ston, in litt.)

Remarks. — Hekstra (1982b) published the
catalogue number as “99.1.27.235, and
year the of collection as “1895.”

Otus choliba koepckeae, p. 60

Holotype. —-AMNH 802425, adult fe-
male, near Yungai, 4000 m, Quebrada Yun-
gay, Cordillera Blanca, Departamento An-
cash, Peru, 3 Aug 1960, collected by Maria
Koepcke.

Remarks. —Hekstra (1982b) referred to
the holotype as no. 1699 from the Koepcke
collection “‘on loan at AMNH.”’ The name
was originally spelled by Hekstra as
“koepckei’’ and was properly emended to
to koepckeae by Greenway (1987). This form
is considered a distinct species by Marshall
& King (1988).

Otus atricapillus moreliensis, p. 62

Holotype. —AMNH 115738, male, Mo-
relia (Rio Bodoquera), Caqueta, 600 ft., Co-
lombia, 22 Jul 1912, collected by L. E. Mil-
ler.

Remarks. —The trinomial was originally
spelled ‘‘morelius’’ and the locality as ‘“‘La
Morelia” by Hekstra (1982b), who referred
to the locality Caqueta as ““Rio Caqueta.”’
Otus atricapillus is known only from Par-
aguay and adjacent Brazil, and the holotype
may represent either O. sanctaecaternae or
O. choliba (J. T. Marshall, pers. comm.).

Otus atricapillus inambariensis, p. 62

Holotype. —FM 222284, female, Hua-
jyumbe, near Quincemil on the Rio Inam-
bari, 630 m Cuzco, Peru, 28 Jun 1953, col-
lected by C. Kalinowski.

517

Remarks. —Hekstra (1982b) spelled the
trinomial as “inambarii,” from the locality
Rio Inambari. He gave the type locality as
“Quince Mil. Huajyumba, at side of river
of the Inambari, Cuzco, 630 m, Peru.”
However, the Rio Inambari is 45 km SE of
Quincemil (see Stephens & Traylor 1983).
The holotype, an adult (D. Willard, in litt.),
resembles examples of O. watsonii in the
brown phase (D. Willard & J. Fitzpatrick,
pers. comm.).

Otus atricapillus ater, p. 61

Holotype.—USNM 513892, adult male,
Belem, Para, [Brazil], 22 Jul 1964, collected
by Phillip S. Humphrey.

Remarks.—The ‘“SUSNM field nr.
196899,” given for the holotype by Hekstra
(1982b), is the collector’s catalogue number.
Additional information on the specimen la-
bel include reproductive data and soft part
colors. The holotype is an example of O.
watsonil (pers. observ., J. T. Marshall, pers.
comm.).

Otus atricapillus fulvescens, p. 62

Holotype. —-AMNH 34597, adult female,
Chapada, Mato Grosso, [Brazil], 20 Jul
1883, collected by H. H. Smith.

Remarks.—The holotype of O. choliba
chapadensis Hekstra, AMNH 34595, was
collected by Smith from the same locality
as the holotype of fulvescens.

Otus guatemalae pettingilli, p. 56

Holotype. —DMNH 52892 (ex Cornell
Univ. 10445), female, above Rio Savinas
near Gomez Faria, Tamaulipas, Mexico, 11
Apr 1941, collected by O. S. Pettingill, Jr.
(original no. 1046).

Remarks. — Hekstra (1982b) did not name
the repository or give a catalogue number
for this holotype. Under the heading ““Type”’
and following the new name, Hekstra
(1982b) wrote, in quotes: “G. M. Sutton,
1949 (manuscript name).’’ Under the head-
ing “Derivation of name,” Hekstra stated

518

that the name pettingilli was “from a manu-
script name on a label (by Griscom?).”” The
holotype of pettingilli is the specimen of Otus
guatemalae referred to by Sutton & Pettin-
gill (1942) as being similar in size but grayer
above than the holotype of O. g. cassini.

Otus guatemalae petenensis, p. 57

Holotype. -UMMZ 137395, adult fe-
male, Laguna Perdida, Peten, Guatemala,
18 Apr 1920, collected by P. W. Shufeldt
(R. W. Storer, in litt.).

Remarks. —Hekstra (1982b) spelled the
trinominal as “‘peteni”’ from the locality Pe-
ten. The name of the lake was incorrectly
spelled as “‘Perida.”’

Otus guatemalae centralis, p. 57

Holotype. —USNM 484980, adult fe-
male, Cerro Mali, 4100 feet, Darien, Pan-
ama, 14 Feb 1964, collected by C. O. Han-
dley, Jr.

Remarks. — Additional information on the
specimen label: “skull oss; ovary 15 x 10
[mm]; largest ova 3.”

Otus guatemalae pallidus, p. 57

Holotype. —AMNH 476699, adult, sex
unknown, Andes de Cumana, northern
Venezuela, -- Mar 1897, collected by Eng.
André, formerly Museo Delmas (no.
B.11.3.C.-81) and Rothchild Collection.

Remarks. —Hekstra (1982b) omitted the
name of the collector and data on the mu-
seums formerly holding the specimen (see
Greenway 1987).

Otus guatemalae rufus, p. 58

Holotype.—BM(NM) 1888.7.20.64, adult,
sex unknown, Balzar Mtns., western Ecua-
dor, -- Mar 1880, collected by Illingsworth.

Remarks. —The museum designation was
given by Hekstra (1982b) as “USNM”’ al-
though the catalogue number was given as
above. Derek Goodwin (in litt.) confirmed

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

that this specimen is in the British Museum
(Natural History).

Otus guatemalae pacificus, p. 58

Holotype. —BM(NH) 1902.3.13.1564,
adult female, Morropon, 140 m, Piura,
northwestern Peru, 27 Aug 1899, collected
by P. O. Simons.

Remarks. —The catalogue number of the
holotype was given by Hekstra (1982b) as
“BM 023.13.1564.’

Acknowledgments

I thank the following who verified the in-
formation on the specimen labels and the
museum catalogue numbers of the holo-
types: M. LeCroy, American Museum of
Natural History (AMNH); D. Goodwin and
P. R. Colston, British Museum (Natural
History) (BM(NH)); D. Niles, Delaware
Museum of Natural History (DMNH); D.
Willard and S. Lanyon, Field Museum of
Natural History (FM); J. V. Remsen, Jr.,
Louisiana State University Museum
(LSUMZ); N. K. Johnson, Museum of Ver-
tebrate Zoology (MVZ); G. F. Mees, Rijks-
museum van Natuurlike Histoire (RMNH);
and R. W. Storer, University of Michigan
Museum of Zoology (UMMZ). The other
holotypes are in the collection of the U.S.
National Museum of Natural History
(USNM).

R. C. Banks, N. K. Johnson, M. LeCroy,
and J. T. Marshall read the manuscript and
offered helpful suggestions.

Literature Cited

Greenway, J. C., Jr. 1987. Type specimens of birds
in the American Museum of Natural History.
Pt. 4.—American Museum Novitates No. 2879,
63 pp.

Hekstra, G. 1982a. “I don’t give a hoot...” A re-
vision of the American Screech owls (Otus;
Strigidae). Unpublished thesis, Universiteit te
Amsterdam. The Netherlands.

Hekstra,G. P. 1982b. Description of twenty-four new
subspecies of American owls (Aves: Strigi-

VOLUME 102, NUMBER 2

dae).— Bulletin Zoologisch Museum Universi-
teit Amsterdam 9(7):49-63.

International Commission on Zoological Nomencla-
ture. 1985. International code of zoological
nomenclature. 3rd ed. International Trust No-
menclature, British Museum (Natural History),
London, 338 pp.

Kelso, L. 1942. The ear of Otus asio. Biological Leaf-
let Number 14, 2 pp.

Marshall, J. T., & B. F. King. 1988. Subfamily Strigi-
nae typical owls genus Otus. Pp. 331-336 in D.
Amadon and J. Bull, ed., Hawks and owls of
the World: A distributional and taxonomic list. —
Proceedings of the Western Foundation of Ver-
tebrate Zoology 3(4):294-357.

Peters, J. L. 1940. Check-list of birds of the world.
Volume 4. Museum of Comparative Zoology,
Harvard University, Cambridge. 291 pp.

519

Stephens, L., & M. A. Traylor, Jr. 1983. Ornitholog-
ical gazetteer of Peru. Museum of Comparative
Zoology, Harvard University, Cambridge. 271
pp.

Sutton, G. M., & O. S. Pettingill. 1942. Birds of the
Gomez Farias region, southwestern Tamauli-
pas.— Auk 59(1):1-34.

Warren, R. L. M. 1966. Type-specimens of birds in
the British Museum (Natural History). Vol. 1.
Non-passerines. Trustees British Museum (Nat-
ural History), London. 320 pp.

U.S. Fish and Wildlife Service, National
Ecology Research Center, U.S. National
Museum of Natural History, Washington,
DG .20560:

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 520-531

CHROMOSOMES OF TEN SPECIES OF PHILIPPINE
FRUIT BATS (CHIROPTERA: PTEROPODIDAE)

Eric A. Rickart, Lawrence R. Heaney, and Mark. J. Rosenfeld

Abstract.—Standard karyotypes and silver-stained nucleolar organizer re-
gions (Ag-NORs) of ten species of Philippine pteropodids are described. Results
are discussed in the context of an updated version of Andersen’s (1912) phy-
logeny. Data for Cynopterus brachyotis (2N = 34), Eonycteris spelaea (2N =
36) and Macroglossus minimus (2N = 34) agree with previous reports. Pteropus
hypomelanus (2N = 38) and Rousettus amplexicaudatus (2N = 36) have stan-
dard karyotypes identical to those of congeners. The Philippine endemic species
Haplonycteris fischeri (2N = 58), Ptenochirus jagori (2N = 44) and Ptenochirus
minor (2N = 46) have distinctive karyotypes consisting primarily of acrocentric
elements. Haplonycteris has the highest diploid number for the suborder. The
karyotypes of Harpyionycteris whiteheadi (2N = 36) and Nyctimene rabori (2N
= 38) are distinctive, but share some apparenntly derived features with cy-
nopterine genera. In all taxa examined, the Ag-NORs corresponded to the
secondary constrictions on the pair of ““marker’’ chromosomes. These results
demonstrate that the cynopterine section is the most chromosomally variable

clade in the Pteropodidae.

The family Pteropodidae is a large and
diverse assemblage of Old World bats that
appears to constitute a natural group dis-
tinct from other chiropterans (Smith 1980,
Koopman 1984). In the only monographic
study of the entire family, Andersen (1912)
used morphological criteria to construct a
detailed phylogeny. Andersen’s work re-
mains the most complete statement of re-
lationships within the family. However, it
is important to test and, if necessary, modify
his kinship hypotheses using independent
evidence.

Prior analysis of standard karyotypes of
pteropodids had led to the early assumption
that chromosomal variation within the
family is relatively limited (Haiduk et al.
1980). However, differential staining tech-
niques have shown that several genera with
similar standard karyotypes substantially
differ in banding patterns (Haiduk et al.
1981). Nevertheless, fewer than half of the
recognized pteropodid genera have been
karyotyped to date, and fewer have been

examined for banding patterns (Haiduk et
al. 1981).

In this paper, we present standard karyo-
types of ten species of Philippine pteropod-
ids representing nine genera and two
subfamilies. Chromosomal data for the gen-
era Haplonycteris, Harpyionycteris, Nycti-
mene, and Ptenochirus, and for eight of the
species are reported for the first time. We
also present results on silver-stained nu-
cleolar organizer regions (Ag-NORs) for each
species and discuss variation in pteropodid
““marker’” chromosomes. We have inter-
preted our results in the context of Ander-
sen’s (1912) phylogeny, as updated to in-
clude genera described since 1912.

Materials and methods.—Our animals
were all freshly collected from wild popu-
lations and killed with sodium pentabar-
bitol (Nembutal) or with chloroform within
24 h of capture. Chromosome terminology
and preparation methods followed those of
Patton (1967) with the exception that 0.4%
potassium chloride was used for the hypo-

VOLUME 102, NUMBER 2

tonic treatment. Cells were processed and
fixed in the field, and suspensions were
stored at 0O-10°C within two weeks of fixa-
tion. After three to seven months, air-dried
slides were made at the University of Utah.
Standard karyotypes were prepared for each
specimen from photographs of slides stained
with Giemsa. Silver-stained nucleolar or-
ganizer regions (Ag-NORs) were examined
using a procedure modified from that de-
scribed by Howell & Black (1980). Deter-
minations of diploid number were based on
minimal counts of 10 mitotic spreads per
individual. Fundamental numbers (FN) re-
fer to numbers of autosomal arms. Due to
variable specimen quality and the presence
of minute chromosomes in some species,
we consider some FN values to be provi-
sional, as indicated by question marks.
Specimens examined were prepared as skins
with partial skeletons or preserved in fluid
and are deposited in the National Museum
of Natural History (USNM), Washington,
DDG:

Specimens Examined

Cynopterus brachyotis (Muller, 1838).—
Leyte Island, Leyte Province, 7 km N Bay-
bay, cley. 10 m, 10°45'’N, 124°47’E (1 9);
Negros Island, Negros Oriental Province,
Mumacnete City, clev. 5 m,. 09°18’N,
Pooks E (13, 2 29).

Eonycteris spelaea (Dobson, 1871).—
Leyte Island, Leyte Province, Cathedral
Cave, 4 km S, 1 km E Inopacan, elev. 50
m, 10°28’N, 124°45’E (2 22); Negros Island,
Negros Oriental Province, Caves at 4 km N
Manjuyod, elev. 20 m, 09°43'N, 123°10’E
(4 38).

Haplonycteris fischeri Lawrence, 1939.—
Biliran Island, Leyte Province, 5 km N, 10
km E Naval, elev. 850 m, 11°36’N, 124°29’E
(ives 12); Leyte Island, Leyte Province,
Mount Pangasugan, 10.5 km N, 4 km E
Baybay, elev. 700 m, 10°47’'N, 124°50’E (1
6, 1 9).

Harpyionycteris whiteheadi Thomas,
1896.—Leyte Island, Leyte Province, Mount

521

Pangasugan, 10.5 km N, 4 km E Baybay,
elev. 700 m, 10°47’N, 124°50’E (3 3, 4 99);
Negros Island, Negros Oriental Province,
Mount Guinsayawan, 3 km N, 17 km W
Dumaguete City, elev. 1280 m, 09°22’N,
123°09’E (1 &).

Macroglossus minimus (E. Geoffroy,
1810).—Negros Island, Negros Oriental
Province, Dumaguete City, elev. 5 m,
09°18'N, 123°18’E @G 6é, 1 9).

Nyctimene rabori Heaney & Peterson,
1984.—Negros Island, Negros Oriental
Province, Mount Guinsayawan, 3 km N, 17
km W Dumaguete City, elev. 1280 m,
Q9°22'N, 123°O9EAGWS):

Ptenochirus jagori (Peters, 1861).—Leyte
Island, Leyte Province, 7 km N Baybay, elev.
10 m, 10°45'N, 124°47’E (1 6, 6 22), Mount
Pangasugan, 10.2 km N, 2.2 km E Baybay,
elev. 320 m, 10°46'N, 124°49’E (1 8).

Ptenochirus minor Yoshiyuki, 1979.—
Leyte Island, Leyte Province, Mount Pan-
gasugan, 10.2 km N, 2.2 km E Baybay, elev.
320 m, 10°46’N, 124°49’E (2 64, 5 29).

Pteropus hypomelanus Temminck,
1853.—Negros Island, Negros Oriental
Province, 9 km N, 14 km W Dumaguete
City, elev..600 m, 09223'N, 123°L17E (1 6,
2 99).

Rousettus amplexicaudatus (E. Geoffroy,
1810).—Leyte Island, Leyte Province, 7 km
N Baybay, elev. 10 m, 10°45'N, 124°47’'E
(2 46, 4 22); Negros Island, Negros Oriental
EFOvinee ~Oumacuete City, clev. 5 m,
09°18'N, 123°18’E (2 99).

Results

We present standard karyotypes and sil-
ver-stained marker chromosomes for male
specimens of eight pteropodid species in
Figs. 1—4 and briefly discuss them below.
Karyotypes for two additional species are
discussed, but not illustrated.

Subfamily Pteropodinae

Cynopterus brachyotis. 2N = 34, FN =
58, Fig. 1A.—The karyotype is indistin-

522 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Af SC AR RO BR RE BR
*

X Y

Fig. 1. Standard karyotypes and silver-stained marker chromosomes showing nucleolar organizer regions
(insets) of (A) Cynopterus brachyotis 6 (USNM 458083), 2N = 34, FN = 58; (B) Haplonycteris fischeri 6 (USNM
458196), 2N = 58, FN = 66(?).

VOLUME 102, NUMBER 2 523

A le
WX 6G AR ORR kK oa

i Ab GA AR GS GK
ne Ah AA 0H Of §b.
mY

B

AY AR AR WR OBR ke
RG Oe Ax
44 44 RA Bb ax

“a6 a - ~*~ * ~ -£ = a » *

Fig. 2. Standard karyotypes and Ag-NORs (insets) of (A) Harpyionycteris whiteheadi 6 (USNM 458213),
2N = 36, FN = 58; (B) Nyctimene rabori 6 (USNM 458906), 2N = 38; FN = 60.

‘teh = 3.
XY
Ak bp 4

*
ea 20 88 @0 46 “* AA

aoe se e8@ a@ 6 #*#* «ee

B
44 Bh aK tk

Aa a@ 80 @8 An ee a8

sz «?
ya ;

ae an esp 028 AH 47 2

oe « Ae
XY

Fig. 3. Standard karyotypes and Ag-NORs (insets) of (A) Ptenochirus jagori 6 (USNM 458322), 2N = 44,
FN = 56(?); (B) Ptenochirus minor 6 (USNM 458424), 2N = 46, FN = 56(?).

VOLUME 102, NUMBER 2

i Wo St UR xy ay ak

aR 4S te

ye La
+ "

XY

B
SU

AM ux KA XR OH B®

525

SA AA AA AA i -
XY

Fig.4. Standard karyotypes of (A) Pteropus hypomelanus 6(USNM 458447), 2N = 38, FN = 72; (B) Rousettus
amplexicaudatus 6 (USNM 458486), 2N = 36, FN = 68. Asterisks indicate marker chromosomes.

526

guishable from those reported by previous
workers (Yong et al. 1973, Ando et al. 1980,
Harada & Kobayashi 1980). The autosomal
complement consists of 11 pairs of meta-
centrics or submetacentrics, 2 pairs of sub-
telocentrics, and 3 pairs of acrocentric chro-
mosomes. A_ pair of medium-sized
metacentrics are marker chromosomes, that
have a secondary constriction on the short
arm near the centromere. In C. brachyotis,
and in other species (see below), the sec-
ondary constriction coincides with the Ag-
NOR site. The X chromosome is a medi-
um-sized submetacentric and the Y, a small
acrocentric.

Haplonycteris fischeri. 2N = 58, FN =
66(?), Fig. 1B.—The autosomal comple-
ment consists of 3 pairs of small to medium-
sized submetacentrics, 2 pairs of small to
medium-sized subtelocentrics, and 23 pairs
of small to medium-sized acrocentric (or
possibly subtelocentric) chromosomes. The
marker chromosome is a medium-sized ac-
rocentric. The X is a medium-sized sub-
metacentric and the Y, a small acrocentric.

Harpyionycteris whiteheadi. 2N = 36, FN
= 58, Fig. 2A.—The autosomal complement
consists of six pairs of medium to large
metacentric or submetacentric chromo-
somes, six pairs of medium-sized subtelo-
centrics, and five pairs of medium-sized ac-
rocentrics. The marker chromosome is a
large metacentric. The X chromosome is a
medium-sized subtelocentric and the Y, a
small acrocentric.

Nyctimene rabori. 2N = 38, FN = 60(?),
Fig. 2B.— The autosomal complement con-
sists of eight pairs of medium to large meta-
centrics and submetacentrics, four pairs of
medium-sized subtelocentrics, and six pairs
of small acrocentric (or subtelocentric)
chromosomes. The marker chromosome is
a medium-sized metacentric. The X chro-
mosome is a medium-sized metacentric and
the Y, a small submetacentric.

Ptenochirus jagori. 2N = 44, FN = 56(?),
Fig. 3A.—The autosomal complement has 4
pairs of small to large metacentric or sub-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

metacentric chromosomes, 3 pairs of me-
dium-sized subtelocentrics, and 14 pairs of
small to medium-sized acrocentrics (or sub-
telocentrics). The marker chromosome is a
medium-sized acrocentric. The X chro-
mosome is a medium-sized submetacentric
and the Y, a small acrocentric.

Ptenochirus minor. 2N = 46, FN = 56(?),
Fig. 3B.—The standard karyotype is similar
to that of the P. jagori. However the auto-
somal complement includes 2 subtelocen-
tric pairs and 16 acrocentric pairs (as op-
posed to 3 and 14, respectively, for P. jagori).

Pteropus hypomelanus. 2N = 38, FN =
72, Fig. 4A.—A diploid number of 38 for
this species was reported by Yong & Dhal-
iwal (1976), but the karyotype was not il-
lustrated. The autosomal complement con-
sists of 10 pairs of small to large metacentrics
or submetacentrics, and 8 pairs of small to
medium-sized subtelocentrics. The marker
chromosome is a medium-sized submeta-
centric. The X chromosome is a medium-
sized subtelocentric and the Y, a small ac-
rocentric.

Rousettus amplexicaudatus. 2N = 36, FN
= 68, Fig. 4B.—The autosomal group con-
sists of 13 pairs of small to large metacentric
and submetacentric chromosomes, and 4
pairs of medium-sized subtelocentrics. The
marker chromosome is a medium-sized
metacentric. The X is a medium-sized sub-
metacentric and the Y, a small acrocentric.

Subfamily Macroglossinae

Eonycteris spelaea. 2N = 36, FN = 66,
not figured. — Our specimens yielded karyo-
types similar to those reported previously
(Yong & Dhaliwal 1976, Ando et al. 1980,
Harada et al. 1982). The autosomes consist
of 14 pairs of small to large metacentrics or
submetacentrics, 2 pairs of medium-sized
subteocentrics, and 1 pairs of small acro-
centrics. The marker chromosome is a me-
dium-sized metacentric. The X is a medi-
um-sized metacentric and the Y, a small
submetacentric.

VOLUME 102, NUMBER 2

Macroglossus minimus. 2N = 34, FN =
62, not figured. — Our results agree with those
of Ando et al. (1980) and Yong & Dhaliwal
(1976). The autosomal complement in-
cludes 12 pairs of small to large metacentric
and submetacentric chromosomes, 3 pairs
of medium-sized subtelocentrics, and 1 pair
of minute acrocentrics. The marker chro-
mosome is a medium-sized metacentric. The
X chromosome is a medium-sized meta-
centric and the Y, a small acrocentric.

Discussion

Karyotypic data represent a source of in-
formation that may be used for studies of
phylogenetic relationships. Because our data
and those of previous reports are, with a
single exception (Haiduk et al. 1981), re-
stricted to unbanded standard karyotypes,
we believe that there is not yet an adequate
basis for an independent analysis of rela-
tionships. Instead, we use Andersen’s (1912)
morphologically-based phylogeny as a
framework in which to examine chromo-
somal data.

Andersen (1912) did not use current ter-
minology in his monograph of the Ptero-
podidae. However, his general methodol-
ogy is acceptable by today’s standards. He
differentiated character polarities, prefer-
entially used derived characters in con-
structing the phylogeny, and recognized
monophyly of groups. Andersen’s (1912: pp.
lu, 1x1) graphic depiction of relationships
within the portion of the subfamily Ptero-
podinae relevant to this study is shown in
Fig. 5, with several slight modifications.
First, he considered Cynopterus to be near
or part of the ancestral stock that gave rise
to other members of the Cynopterus group,
and so placed Cynopterus at the base of the
group rather than on a terminal branch. The
arrangement shown represents our inter-
pretation of his conclusions based on his
discussion of characters. Second, we have
included six genera that have been named
since 1912. The genera Aethalops, Latidens,

S27

and Paranyctimene were described within
the context of Andersen’s character system,
and their cladistic positions are unambig-
uous (Thomas 1932, Tate 1942, Thonglon-
gya 1972). Unfortunately, the Philippine
endemic genera Alionycteris, Haplonycteris,
and Otopteropus were not described within
Andersen’s framework, so their positions are
harder to determine. The three genera are
very similar and undoubtedly closely relat-
ed (Lawrence 1939; Kock 1969a, b). They
resemble several other small cynopterines
(e.g., Aethalops and Balionycteris), but most
similarities involve character losses. Pend-
ing a thorough analysis of characters, we
tenatively associate them with the Cynop-
terus group.

Several authors have differed with An-
dersen’s interpretation of pteropodid phy-
logeny: these differences indicate areas of
uncertainty. Miller (1907) and Simpson
(1945) listed Harpyionycteris as the sole
member of a separate subfamily Harpyio-
nycterinae. Anderson himself recognized the
subfamily in his formal classification. How-
ever, his discussion and diagrams indicate
that he did so because of the many unique
features of Harpyionycteris, and that he con-
sidered Dobsonia, a member of the rouset-
tine section, to be its sister-taxon (Fig. 5).
Similarly, Miller (1907), Simpson (1945),
and others have recognized the unique fea-
tures of Nyctimene and Paranyctimene, and
have placed them in a separate subfamily.
However, Andersen noted that Nyctimene
shared derived characters with members of
the cynopterine section, and he placed it in
that group. Andersen considered most char-
acters of Myonycteris to be primitive for the
family. However, on the basis of several
derived features he united this genus with
the cynopterines. Bergmans (1976) trans-
ferred Myonycteris to the rousettine section
based principally on characters that Ander-
sen believed to be primitive for the family,
but that Bergmans considered to be useful
for defining the rousettines.

Previous karyotypic work on megachi-

528

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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* = o = "” = .
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ee te Sn a oma Mr Fa x 4“ wm 2D <x
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7

?
CYNOPTERUS
group
CYNOPTERINE ROUSETTINE
section section

Fig. 5. Phylogeny of the cynopterine and rousettine sections of the subfamily Pteropodinae modified from

Andersen (1912). Asterisks indicate genera described since 1912 (see text).

ropterans suggested a rather narrow range
of variation in chromosome number and
shape, most taxa having diploid numbers
ranging from 34 to 38 and a majority of
biarmed elements (Summarized in Haiduk
et al. 1980; 1981). Exceptions include Bal-
ionycteris maculata (2N = 24) and Mega-
erops ecaudatus (2N = 24, 26) with lower
diploid numbers, and Penthetor lucasi (2N
= 48) with a higher count (Yong & Dhaliwal
1976, Harada et al. 1982, Yong 1984). Our
data on the Philippine endemic Haplonyc-
teris (2N = 58; Fig. 1B) now place it at the
upper extreme for the suborder Megachi-
roptera. In contrast to all other pteropodids
examined except Ptenochirus (see below),
the karyotype of Haplonycteris consists pri-
marily of acrocentric elements. In contrast
to Cynopterus (2N = 34; Fig. 1A), the FN
differences (66 vs. 58) indicates a series of
changes including both whole arm translo-
cations and non-Robertsonian events.

The two species of Ptenochirus, another
genus endemic to the Philippines, also have

high diploid numbers (2N = 44 and 46).
The difference in 2N supports the specific
status of the recently described P. minor
(Yoshiyuki 1979, Heaney & Rabor 1982).
The species share an FN of 56 and appear
to be separated by a single Robertsonian
whole-arm translocation. As is the case with
Haplonycteris, the karyotypes include a high
proportion of acrocentrics (Fig. 3A, B).
However, Ptenochirus also possess a num-
ber of large biarmed elements not present
in Haplonycteris.

Andersen (1912) considered Nyctimene a
specialized member of the cynopterine sec-
tion. With FN = 60 and six pairs of small
acrocentric elements, the standard karyo-
type of Nyctimene rabori (Fig. 2B) does re-
semble those of several cynopterines. How-
ever, the small metacentric marker
chromosome and the metacentric Y distin-
guish it from other genera in this section.

The relationship of Harpyionycteris to
other pteropodids is uncertain. Andersen
(1912) associated it with his rousettine sec-

VOLUME 102, NUMBER 2

tion as the sister-taxon to Dobsonia, where-
as Tate (1951) postulated a closer associa-
tion with Nyctimene on the basis of certain
shared characters (fusion of premaxillae and
contact of lower canines). Although the dip-
loid number of Harpyionycteris (2N = 38;
Fig. 2A) is near the median value for the
family (2N = 36), the standard karyotype
is unique. Several pairs of relatively large
acrocentric elements distinguish it from
known karyotypes of epomophorine, rou-
settine and macroglossine genera. This fea-
ture allies it with the cynopterines, although
the absence of small acrocentrics is distin-
guishing.

The rousettine section of the Pteropodi-
nae seems relatively conservative karyotyp-
ically. All members of the genus Pferopus
examined to date, including P. hypomelan-
us (Fig. 4A), exhibit similar standard karyo-
types with 2N = 38, FN = 72 (Haiduk et
al. 1980, Harada & Kobayashi 1980, Ka-
sahara & Dutrilaux 1983). However, there
are some interspecific differences in hetero-
chromatin content (Kasahara & Dutrilaux
1983). Rousettus karyotypes resemble those
of Pteropus in that they consist almost ex-
clusively of biarmed elements. The stan-
dard karyotype of Rousettus amplexicau-
datus (2N = 36, FN = 68; Fig. 4B) is
indistinguishable from that reported by
Harada et al. (1982) and Ray-Chaudhuri et
al. (1968) for R. leschenaulti. The only other
member of the genus that has been exam-
ined, Rousettus aegyptiacus (2N = 36, FN
= 66), differs from R. amplexicaudatus and
R. leschenaulti by a single rearrangement of
the smallest autosomal element, which is
acrocentric rather than biarmed (Dulic &
Mutere 1973, Haiduk et al. 1981).

Our data on the widespread macroglos-
sine species Eonycteris spelaea and Mac-
roglossus minimus agree with previous
findings (Yong and Dhaliwal 1976). Little
karyotypic variation has been detected
among macroglossines, but few taxa have
been examined (Haiduk et al. 1980).

Extreme diploid numbers (24 to 58) for

529

the pteropodids are confined to the cynop-
terine section of the subfamily Pteropodi-
nae, whereas in the rousettine section and
in the Macroglossinae, presumably out-
groups to cynopterines, diploid numbers
range only from 34 to 38 (Haiduk et al.
1980). This pattern suggests that both higher
and lower diploid numbers represent de-
rived states within the cynopterine section.
The morphological specializations of both
Haplonycteris and Megaerops relative to the
presumed primitive genus Cynopterus (An-
dersen 1912; Lawrence 1939) support this
contention. The broad range in FN (44-66)
indicates that karyotypic evolution within
the cynopterine section has involved more
than Robertsonian rearrangements.

With the possible exception of Scotonyc-
teris ophiodon, all pteropodids examined
possess a pair of marker chromosomes with
secondary constriction sites (Haiduk et al.
1980). Yong (1984) demonstrated that the
secondary constriction corresponds to the
silver-stained nucleolar organizer region in
Megaerops. This same relationship, which
also exists for several vespertilionids (Vol-
leth 1987), was observed for all taxa in this
study. In a few of our specimens, prepara-
tions consistently revealed only one mark-
er/NOR element (e.g., Fig. 1B). No speci-
men had more than one pair. In
Haplonycteris and Ptenochirus, the NOR
sites are located on medium-sized acrocen-
trics. In the other genera, they are on a pair
of medium-sized or large metacentrics or
submetacentrics. For all taxa, the NORs are
located interstitially near the centromere.
The general uniformity of the marker arms
suggests that they are homologous through-
out the family. However, variation in the
size of biarmed marker chromosomes (e.g.,
between Harpyionycteris and Nyctimene)
may indicate non-homologous Robertson-
ian events or differences in heterochromatin
content.

Data currently available allow us to draw
several general conclusions. First, the
subfamily Macroglossinae and the rouset-

530

tine section of the subfamily Pteropodinae
exhibit low variability in gross chromo-
somal morphology. Second, our data rein-
force earlier observations of the cynopterine
section of the Pteropodinae as the most
karyotypically variable clade in the family.
Third, the karyotype of Harpyionycteris fur-
ther highlights the uncertainty of its phy-
logenetic placement. Although these con-
clusions demonstrate the utility of
karyotypes in investigating pteropodid phy-
logenetic relationships, they also indicate the
need for more extensive studies involving
banding data.

Acknowledgments

We thank P. Heideman, J. Klompen, and
R. Utzurrum for their valuable field assis-
tance in the Philippines. M. Carleton and
L. Wilburn provided helpful comments on
the manuscript. We gratefully acknowledge
the cooperation and assistance of the Phil-
ippine Bureau of Forest Development, the
Philippine National Museum, Silliman
University, and the Institute of Philippine
Culture at Ateneo de Manila University.
This study was supported by National Sci-
ence Foundation grant number BSR-
8514223 (awarded to L. R. Heaney).

Literature Cited

Andersen, K. 1912. Catalogue of the Chiroptera in
the collection of the British Museum, Vol. 1:
Megachiroptera. 2nd ed. British Museum (Nat-
ural History), London. pp. ci + 854.

Ando, K., T. Tagawa, & T. A. Uchida. 1980. A karyo-
typic study on four species of the Indonesian
fruit-eating bats belonging to Cynopterus, Eonyc-
teris, and Macroglossus (Chiroptera: Pteropi-
dae). —Caryologia 33:41-53.

Bergmans, W. 1976. A revision of the African genus
Myonycteris.—Beaufortia 24:189-216.

Dulic, B. & F. A. Mutere. 1973. Les chromosomes
de trois especes des Mégachiropteres (Mam-
malia; Chiroptera) d’Afrique oriental.—
Caryologia 26:389-396.

Haiduk, M. W., R. J. Baker, L. W. Robbins, & D. A.
Schlitter. 1981. Chromosomal evolution in
African megachiroptera: G- and C-band assess-
ment of the magnitude of change in similar stan-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

dard karyotypes.—Cytogenetics and Cell Ge-

netics 29:221-—232.

, L. W. Robbins, R. L. Robbins, & D. A. Schlit-

ter. 1980. Karyotypic studies of seven species

of African megachiropterans (Mammalia:

Pteropodidae).—Annals of the Carnegie Mu-

seum of Natural History 49:181-191.

Harada, M., & T. Kobayashi. 1980. Studies on the
small mammals of Sabah, east Malaysia II.
Karyological analysis of some Sabahan mam-
mals (Primates, Rodentia, Chiroptera).—Con-
tributions from the Biological Laboratory, Kyo-

to University 26:83-95.

, M. Minezawa, S. Takada, S. Yenbutra, S. Nun-

pakdee, & S. Ohtani. 1982. Karyological anal-

ysis of 12 species of bats from Thailand. —Car-

yologia 35:269-278.

Heaney, L. R., & D. S. Rabor. 1982. Mammals of
Dinagat and Siargao islands, Philippines. —Oc-
casional Papers of the Museum of Zoology, Uni-
versity of Michigan 699:1-—30.

Howell, W. M., & D. A. Black. 1980. Controlled
silver staining of neucleolus organizer regions
with a protective colloidal developer: a 1-step
method.— Experientia 36:1014—1015.

Kasahara, S., & B. Dutrilaux. 1983. Chromosome
banding patterns of four species of bats, with
special reference to a case of X-autosome trans-
location.— Annales de Génétique 26:197-201.

Kock, D. 1969a. Eine neue Gattung und Art cynop-

teriner Flughunde von Mindanao, Philippinen

(Mammalia, Chiroptera).—Senckenbergiana

Biologica 50:319-—327.

1969b. Eine bemerkenswerte neue Gattung
und Art Flughunde von Luzon, Philippinen

(Mammalia, Chiroptera).—Senckenbergiana

Biologica 50:329-338.

Koopman, K. F. 1984. Bats. Pp. 145-186 in S. An-
derson and J. K. Jones, Jr., eds., Orders and
families of Recent mammals of the world. John
Wiley, New York, 686 pp.

Lawrence, B. 1939. Collections from the Philippine
Islands. Mammals. — Bulletin of the Museum of
Comparative Zoology, Harvard University 86:
28-73.

Miller, G. S., Jr. 1907. The families and genera of
bats.— Bulletin of the United States National
Museum 57:1-282.

Patton, J. L. 1967. Chromosome studies of certain
pocket mice, genus Perognathus (Rodentia: Het-
eromyidae).— Journal of Mammalogy 48:27-37.

Ray-Chaudhuri, S. P., S. Pathak, & T. Sharma. 1968.
Chromosomes and affinities of Pteropidae
(Megachiroptera) and Rhinopomatidae (Micro-
chiroptera).—The Nucleus (Supplement):96—
101.

Simpson, G. G. 1945. The principles of classification

VOLUME 102, NUMBER 2

and a classification of mammals.—Bulletin of
the American Museum of Natural History 85:
1-350.

Smith, J.D. 1980. Chiropteran phylogenetics: intro-
duction. Pp. 233-244, in D. E. Wilson and A.
L. Gardner, eds., Proceedings fifth international
bat research conference, Texas Tech Press, Lub-
bock, 434 pp.

Tate, G. H. H. 1942. Results of the Archbold expe-

ditions. No. 46. A new genus and species of fruit

bats, allied to Nyctimene.— American Museum

Novitates 1204:1-2.

1951. Harpyionycteris, a genus of rare fruit
bats.— American Museum Novitates 1522:1-9.
Thomas, O. 1932. On some small mammals, chiefly

bats, from the East Indian archipelago. — Annals
and Magazine of Natural History, series 9, 11:
250-255.

Thonglongya, K. 1972. A new genus and species of
fruit bat from south India (Chiroptera; Ptero-
podidae).—Journal, Bombay Natural History
Society 69:151-158.

Volleth, M. 1987. Differences in the location of nu-
cleolus organizer regions in European vesper-
tilionid bats.—Cytogenetics and Cell Genetics
44:180-197.

Yong, H. S. 1984. Robertsonian translocation, peri-
centric inversion and heterochromatin block in

531

the evolution of the tailless fruit bat.—Exper-

ientia 40:875-876.

, & §.S. Dhaliwal. 1976. Chromosomes of the

fruit-bat subfamily Macroglossinae from pen-

insular Malaysia. —Cytologia 41:85-89.

abe Lam, Kook. Teh, & A; N. Start:
1973. Uniformity in the karyotype of the fruit
bats Cynopterus (Mammalia: Chiroptera: Pter-
opodidae).— Malaysian Journal of Science 2:19-
2.

Yoshiyuki, M. 1979. Anewspecies of the genus Pren-
ochirus (Chiroptera, Pteropodidae) from the
Philippine Islands.—Bulletin of the National
Science Museum (Tokyo), series A (Zoology) 5:
75-81.

(EAR AND MJR) Utah Museum of Nat-
ural History, University of Utah, Salt Lake
City, Utah 84112; (LRH) Department of
Vertebrate Zoology, Division of Mammals,
National Museum of Natural History,
Smithsonian Institution, Washington, D.C.
20560; (Present address: Field Museum of
Natural History, Roosevelt Road at Lake
Shore Drive, Chicago, Illinois 60605).

PROC. BIOL. SOC. WASH.
102(2), 1989, pp. 532-534

TAXONOMIC STATUS OF THE
DELPHINID (MAMMALIA: CETACEA)
TURSIO? PANOPE PHILIPPI, 1895

Robert L. Brownell, Jr. and James G. Mead

Abstract.—Tursio? panope Philippi, 1895 is a junior synonym of Lageno-
rhynchus obscurus (Gray, 1828) and not Cephalorhynchus eutropia (Gray, 1846)

as noted by previous authors.

Rodolpho Amando Philippi (1895) men-
tioned a new species of dolphin as Tursio?
panope based on a skull in the Museo Na-
cional de Historia Natural, Santiago, Chile
(MNHN-S). He described his specimen in
more detail in 1896 (p. 14, pls. 4, 5, and 6).
No locality was given for the holotype but
Philippi (1896:14) implied that it was from
Chile. The generic and specific identity of
this delphinid have remained uncertain.

True (1903:141) was unable to determine
the generic affinity of 7.? panope with cer-
tainty, but suggested that it probably rep-
resented a new genus. He based this idea on
Philippi’s drawing, which showed a bowed
rostrum in lateral view (Fig. 1). Trouessart
(1904:766) referred to the specimen as Lis-
sodelphis? panope. Harmer (1922:631) be-
lieved that 7.? panope might belong to the
genus Cephalorhynchus. Miller (1928:171)
was also uncertain about the generic affinity
of the specimen but felt that it eventually
would be placed in the genus Cephalorhyn-
chus. He made this determination based on
photographs of the type specimen taken by
Waldo L. Schmitt in 1926. Unfortunately,
these photographs were not published until
now (Fig. 2). Cabrera (1961:615), Hersh-
kovitz (1966:75), Donoso-Barros (1975:33)
and Tamayo & Frassinetti (1980:364) listed
T. panope as a junior synonym of Cepha-
lorhynchus eutropia. Sielfeld (1980:277), in
his list of marine mammal specimens in
Chilean museums, included a _ skull
MNHN-S 584, under C. eutropia. This

specimen was later correctly identified as
the holotype of Tursio? panope by Goodall
et al. (1988:203). They noted that the known
range of condylobasal length (CBL) in C.
eutropia 1s 302 to 364 mm (mean 341.3
mm, n = 13), whereas the CBL of the type
of 7. panope was slightly greater than 379
mm (the tip of the rostrum is damaged).
They left 7. panope as incertae sedis.

All previous authors have cited Philippi,
1896, as the type description of Tursio? pan-
ope. We have used Philippi, 1895. Although
Philippi (1895) provided only a synopsis of
his 1896 account of Tursio? panope and sev-
eral other new small cetaceans, the descrip-
tion given is valid under the International
Code of Zoological Nomenclature. We be-
lieve that Philippi’s 1895 work was pub-
lished before Philippi (1896) because a re-
print of the 1895 paper in the Museo
Nacional de Historia Natural, Montevideo
has a separate cover that is also dated 1895.
The year of publication, therefore, should
be Philippi, 1895, not Philippi, 1896.

Much confusion has exited regarding the
specimens, numbers, and labels of the small
cetaceans in the collection at the MNHN-S.
In 1986 RLB examined the type specimen
of Tursio? panope and confirmed its identity
as the type. Cranial damage matches that
shown in the photographs taken by Waldo
L. Schmitt in 1926 (see Fig. 2) and the orig-
inal drawings of the type specimen (see Fig.
1). The specimen bears the (old) number
937. The catalog entry for the specimen

VOLUME 102, NUMBER 2

533

Pipe A:
5(2) and 6(2).

states: ““Tursio? panope Ph. craneo San Vi-
cente 1887 hallado en la playa.’ San Vicente
is near Concepcion, Chile (36°44’S). This
catalog number was probably assigned by
Philippi. The new number 584 was assigned
by the present Curator of Mammals, José
Yanez, who started a new catalog and num-

Original drawings of the holotype of Tursio? panope Philippi 1895, from Philippi 1896, Plates 4(2),

bering system. The cards and catalog for the
old system were retained (see discussion of
this in Goodall et al. 1988:213).

Very little has been published on the type
of Tursio? panope. With no other descrip-
tion available except for the drawings of the
skull, all subsequent authors were puzzled

Fig. 2. Dorsal, ventral and lateral views of the holotype of Tursio? panope Philippi, 1895. The condylobasal
length is 380+ mm. Photography by Waldo L. Schmitt, 1926.

534

by the extraordinary downward curvature
of the rostrum. The CBL (380+ mm as
measured by Brownell; the rostrum is dam-
aged) of 7.? panope exceeds that of the larg-
est known physically mature specimens of
Cephalorhynchus eutropia by more than 15
mm (see Goodall et al. 1988). The type of
Tursio? panope is very similar in appear-
ance and size to specimens of Lagenorhyn-
cus obscurus (Gray, 1828). The CBL range
for 16 specmens of L. obscurus is 333 to 421
mm with a mean of 373 mm (specimens in
USNM).

Although the age of the holotype is not
apparent in Philippi’s drawings, examina-
tion of the specimen revealed that it is im-
mature. The basioccipital has not yet fused
with the vomer, and the left zygomatic has
not yet fused with the parietal and exoccip-
ital. The upper tooth count of the holotype
is 28 on the left and 29 or 30 on the right.
This is a minimum count because of the
damage to the tip of the rostrum. The man-
dibular tooth count is 29 on each side.

The CBL, skull shape and ventral topog-
raphy of the type of Tursio? panope corre-
spond closely to all specimens of Lageno-
rhynchus obscurus examined by us. Based
on CBL and immaturity alone, the skull is
too large to be from a specimen of Cephalo-
rhynchus eutropia. Therefore, Tursio? pan-
ope Philippi, 1895 is reidentified here as a
junior synonym of Lagenorhynchus obscu-
rus (Gray, 1828).

We thank José Yanez V., Museo Nacional
de Historia Natural, Santiago, Chile, for al-
lowing us access to various small cetacean
specimens under his care, and Alvaro
Mones, Museo Nacional de Historia Nat-
ural, Montevideo, Uruguay, for his help with
the Philippi literature. Ricardo Praderi,
William F. Perrin and Clayton E. Ray pro-
vided useful comments on this manuscript.

Literature Cited

Cabrera, A. 1961. Catalogo de los mamiferos de
America del Sur.— Revista del Museo Argenti-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

no de Ciencias Naturales ““Bernardino Riva-
davia,”’ Buenos Aires, Ciencias Zoologicas 4(2):
309-732.

Donoso-Barros, R. 1975. Contribucion al conoci-
miento de los cetaceos vivientes y fosiles del
territorio de Chile.—Gayana, Universidad de
Concepcion, Zoologia 36:1—127.

Goodall, R. N. P., K. S. Norris, A. R. Galeazzi, J. A.
Oporto, & I.S. Cameron. 1988. On the Chil-
ean dolphin, Cephalorhynchus eutropia (Gray,
1846).—Reports of the International Whaling
Commission (Special Issue 9):197—257.

Harmer, S. F. 1922. On Commerson’s dolphin and
other species of Cephalorhynchus.—Proceed-
ings of the Zoological Society of London 43:
627-638.

Hershkovitz, P. 1966. Catalog of living whales.—
United States National Museum, Bulletin 246:
1-259.

Miller, G. S., Jr. 1928. The generic position of the
porpoise described by Philippi as Tursio? pan-
ope.— Proceedings of the Biological Society of
Washington 41:171.

Philippi, R. A. 1895. Los delfines Chilenos.—Anales

de la Universidad de Chile, Santiago, 1895:281-

285.

. 1896. Los craneos de los delfines Chilenos. —

Anales del Museo Nacional de Chile, Santiago

1 (Zoologia) 12:1-18 + 6 pl.

Sielfeld K., W. 1980. Mamiferos marinos en colec-
ciones y museos de Chile. —Anales del Instituto
de la Patagonia, Punta Arenas, Chile, 11:273-
280.

Tamayo H., M., & D. Frassinetti C. 1980. Catalogo
de los mamiferos fosiles y vivientes de Chile. —
Boletin del Museo Nacional de Historia Natu-
ral, Chile 37:323-405.

Trouessart, E.-L. 1904. Catalogus mammalium tam
viventium quam fossilium. R. Friedlande and
Sohn, Quinquennale supplementum, 929 pp.

True, F. W. 1903. On the species of the South Amer-
ican Delphinidae described by Dr. R. A. Phi-
lippi in 1893 and 1896.— Proceedings of the Bi-
ological Society of Washington 16:133-143.

(RLB) U.S. Fish and Wildlife Service, Post
Office Box 70, San Simeon, California
93452; (JGM) Department of Vertebrate
Zoology, National Museum of Natural His-
tory, Smithsonian Institution, Washington,
D.C. 20560.

INTERNATIONAL COMMISSION ON ZOOLOGICAL
NOMENCLATURE

% British Museum (Natural History)
Cromwell Road
London SW7 5BD

January 1989

The following applications were published on 16 December 1988 in Vol. 45, Part
4 of the Bulletin of Zoological Nomenclature. Comment or advice on these appli-
cations is invited for publication in the Bulletin and should be sent to the Executive
Secretary, ICZN, British Museum (Natural History), Cromwell Road, London SW7
SBD, U.K.

Case No.

2663 Orbitolina dOrbigny, 1850 (Foraminiferida): proposed confirmation of Or-
bulites concava Lamarck, 1816 as the type species.

2653 Hapalorhynchus beadlei Goodman, 1987 (Trematoda, Digenea): proposed
replacement of the holotype by a lectotype.

2633 Phyllodoce (Carobia) rubiginosa Saint-Joseph, 1888 (currently also Nerei-
phylla rubiginosa; Annelida, Polychaeta): proposed conservation of
the specific name.

2636 Fizesereneia Takeda & Tamura, 1980 (Crustacea, Decapoda): proposed con-
firmation of 7roglocarcinus heimi Fize & Seréne, 1956 as the type
species.

2645 Bodotria Goodsir, 1843 (Crustacea, Cumacea): proposed conservation.

2644 Leucon Kr@yer, 1846 (Crustacea, Cumacea): proposed conservation.

2643 Iphinoe Bate, 1856 (Crustacea, Cumacea): proposed conservation.

N.B. Would Editors of Molluscan journals note that this application
involves the suppression of the gastropod name Iphinoe H. & A.
Adams, 1854?

Aleuropteryx Low, 1885 (Insecta, Neuroptera): proposed designation of Aleu-
ropteryx loewii Klapalek, 1894 as the type species.

Sialis Latreille, 1802 (Insecta, Megaloptera): proposed conservation by the
confirmation of Phryganea phalaenoides Linnaeus, 1758 as the type
species of Semblis Fabricius, 1775 (Insecta, Trichoptera).

Ophonus Dejean, 1821 and Tachys Dejean, 1821 (Insecta, Coleoptera): pro-
posed designation of type species.

Papilio carthami Hubner, [1813] and Syrichthus serratulae major Staudinger,
1879 (currently both in Pyrgus; Insecta, Lepidoptera): proposed con-
servation of the names carthami and major.

Rapport sur les Myodaires du Docteur Robineau Desvoidy, (1826): proposed
nomenclatural suppression.

Physcus Howard, 1895 (Insecta, Hymenoptera): proposed conservation.

J DD Smith
Scientific Administrator

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CONTENTS

A revision of Arabella mutans (Chamberlin, 1919) and related species (Polychaeta: Arabellidae)
G. Kent Colbath
A new species of Benhamipolynoe (Polychaeta: Polynoidae: Lepidastheniinae) from Australia,
associated with the unattached stylasterid coral Conopora adeta Marian H. Pettibone
Two new species of Harmothoinae (Polychaeta: Polynoidae) from the East Pacific Rise, collected
by Alvin dives 2000 and 2003 Marian H. Pettibone
Varichaetadrilus angustipenis (Brinkhurst and Cook, 1966), new combination for Limnodrilus
angustipenis (Oligochaeta: Tubificidae) Ralph O. Brinkhurst
The adult male of the troglobitic ostracode Spelaeoecia bermudensis Angel and Iliffe, 1987,
from an anchialine cave in Bermuda (Crustacea: Ostracoda: Halocypridoidea)
Louis S. Kornicker
New records of entocytherid ostracods infesting burrowing crayfishes, with the description of

a new species, Ascetocythere stockeri Horton H. Hobbs, Jr. and Daniel J. Peters
Pseudocyclops lepidotus, a new species of demersal copepod (Calanoida: Pseudocyclopidae)
from the northwestern Pacific Douglas J. Barr and Susumu Ohtsuka

A revision of the genus Microprotus Richardson with descriptions of two new species, M.
acutispinatus and M. lobispinatus (Asellota, Isopoda, Crustacea)
George D. F. Wilson, Oleg G. Kussakin, and Galina S. Vasina
Four species of Synopiidae from the Caribbean region (Crustacea: Amphipoda)
J. L. Barnard and James Darwin Thomas
A new species, Ampelisca burkei, (Crustacea, Amphipoda) from Florida
J. L. Barnard and James Darwin Thomas
Aegla denticulata lacustris, new subspecies, from Lake Rupanco, Chile (Crustacea: Decapoda:
Anomura: Aeglidae) Carlos G. Jara
Freshwater crabs associated with caves in southern Mexico and Belize, with descriptions of
three new species (Crustacea: Decapoda) Gilberto Rodriguez and Horton H. Hobbs, Jr.
A new deep-sea crab of the genus Chaceon from Chile (Crustacea, Decapoda, Geryonidae)
Luis A. Chirino-Galvez and Raymond B. Manning
Upogebia corallifora, a new species of coral-boring shrimp from the West Indies (Decapoda:

Upogebiidae) Austin B. Williams and P. J. B. Scott
Additional records for an Atlantic reef lobster, Enoplometopus antillensis Liitken, 1865 (Crus-
tacea, Decapoda, Enoplometopidae) Raymond B. Manning and David K. Camp

Studies of neotropical caddisflies, XL: new species of Smicridea (Smicridea) from middle
America and the West Indies (Trichoptera: Hydropsychidae)

Oliver S. Flint, Jr. and D. G. Denning

A review of the shore-fly genus Polytrichophora Cresson from Asia (Diptera: Ephydridae)
Wayne N. Mathis and Jin Zuyin
Adelphydraena, new genus, and two new species from Venezuela, and remarks on phylogenetic
relationships within the subtribe Hydraenina (Coleoptera: Hydraenidae) Philip D. Perkins
Symphurus callopterus (Cynoglossidae, Pleuronectiformes), a new deepwater tonguefish from

the eastern Pacific Thomas A. Munroe and Madhu N. Mahadeva
Systematics of the Steindachnerina hypostoma complex (Pisces, Ostariophysi, Curimatidae),
with the description of three new species Richard P. Vari and Ann Williams Vari

Description of two new species of the Eleutherodactylus milesi group (Amphibia: Anura:
Lepucdeet eae) from northern Honduras
James R. McCranie,.Jay M. Savage, and Larry David Wilson
A new species of diminutive Eleutherodactylus (Leptodactylidae) from Oaxaca, Mexico
Jonathan A. Campbell, William W. Lamar, and David M. Hillis
Physalaemus crombiei (Amphibia: Leptodactylidae), a new frog species from Espirito Santo,
Brazil with comments on the P. signifer group W. Ronald Heyer and Alan J. Wolf
Redescription of the garter snake Thamnophis exsul Rossman, 1969 (Serpentes: Colubridae)
Douglas A. Rossman, Ernest A. Liner, Carlos H. Trevifio, and Alan H. Chaney
The type specimens of Hekstra’s owls M. Ralph Browning
Chromosomes of ten species of Philippine fruit bats (Chiroptera: Pteropodidae)
Eric A. Rickart, Lawrence R. Heaney, and Mark J. Rosenfeld
Taxonomic status of the delphinid (Mammalia: Cetacea) Tursio? panope Philippi, 1895
Robert L. Brownell, Jr. and James G. Mead
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