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PROCEEDINGS

of the

Biological Society of
Washington

VOLUME 106
1993

Vol. 106(1) published 8 April 1993 Vol. 106(3) published 20 September 1993

Vol. 106(2) published 1 1 June 1993 Vol. 106(4) pubUshed ?? December 1993

WASHINGTON
PRINTED FOR THE SOCIETY

EDITOR
C. Brian Robbins

ASSOCIATE EDITORS

Classical Languages Invertebrates

George C. Steyskal Jon L. Norenburg

Frank D. Ferrari
Rafael Lemattre

Plants Vertebrates

David B. Lelunger Thomas A. Munroe

Insects
Wayne N. Mathis

All correspondence should be addressed to the

Biological Society of Washington, Smithsonian Institution

Washington, D.C. 20560

Allen Press Inc.
Lawrence, Kansas 66044

OFHCERS AND COUNCIL

of the

BIOLOGICAL SOCIETY OF WASHINGTON

FOR 1992-1993

OFFICERS

President
STORRS L. OLSON

President-Elect
JANET W. REID

Secretary
G. DAVID JOHNSON

Treasurer
T. CHAD WALTER

COUNCIL

Elected Members
STEPHEN D. CAIRNS JON L. NORENBURG

RICHARD C. FROESCHNER LYNNE R. PARENTI

ALFRED L. GARDNER F. CHRISTIAN THOMPSON

TABLE OF CONTENTS

Volume 106

Alonso de Pina, Gloria M. Linca pinita, a new phoxocephalid genus and species (Crus-
tacea: Amphipoda) from the Argentine continental shelf 497-507

Alvarez, Belinda and R. W. M. Van Soest. A new sponge species, Ceratopsion crustosum
(Demospongiae: Raspailiidae), from deep waters of the Gulf of Mexico 629-632

Baba, Keiji. Anomoeomunida, a new genus proposed for Phylladiorhynchus caribensis
Mayo, 1972 (Crustacea: Decapoda: Galatheidae) 102-105

Bamber, Roger N. A new species of Kalliapseudes (Crustacea: Tanaidacea: Kalliapseu-
didae) from Trinidad 1 22-1 30

Banford, Heidi M. and Bruce B. Collette. Hyporhamphus meeki, a new species of
halfbeak (Teleostei: Hemiramphidae) from the Atlantic and Gulf coasts of the United
States 369-384

Blake, James A. New genera and species of deep-sea polychaetes of the family Nau-
tiliniellidae from the Gulf of Mexico and the eastern Pacific 147-157

Brooks, Daniel R. and Barbara Holcman. Revised classification and phylogenetic hy-
pothesis for the Acanthostominae Looss, 1899 (Digenea: Opisthorchiformes: Cryp-
togonimidae) 207-220

Bruce, A. J. Potamalpheops darwiniensis (Crustacea: Decapoda: Alpheidae), the third
Indo-west Pacific species 698-704

Campos, Ernesto. Systematics and taxonomic remarks on Pinnotheres muliniarum
Rathbun, 1918 (Crustacea: Brachyura: Pinnotheridae) 92-101

Campos, Martha R. and Gilberto Rodriguez. Three new species of Strengeriana from
Colombia (Crustacea: Decapoda: Pseudothelphusidae) 508-5 1 3

Casanova, Jean-Paul. Spadellajaponica, a new coastal benthic chaetognath from Japan 359-365

Castro, Ricardo M. C. Prochilodus britskii, a new species of prochilodontid fish (Os-
tariophysi: Characiformes), from the rio Apiaca, rio Tapajos system, Mato Grosso,
Brazil 57-62

Couri, Marcia Souto and Carlos Jose Einicker Lamas. Revision of Ylasoia Speiser,

1920 (Insecta: Diptera: Bombyliidae: Lomatiinae) 450-454

Cumberlidge, Neil. Further remarks on the identity of Sudanonautes orthostylis Bott,
1955 (Crustacea: Decapoda: Potamoidea: Potamonautidae) with comparisons with
other species from Nigeria and Cameroon 514-522

Curino, Alejandro C. and Nestor J. Cazzaniga. A new species of freshwater planarian
from Chile (Platyhelminthes: Tricladida), with a nomenclatural note on Girardiafestae
(Borelli, 1898) 633-644

Desbruyeres, Daniel and Lucien Laubier. New species of Alvinellidae (Polychaeta) from
the north Fiji back-arc basin hydrothermal vents (southwestern Pacific) 225-236

Emmons, Louise H. On the identity oi Echimys didelphoides Desmarest, 1817 (Mam-
malia: Rodentia: Echimyidae) 1-4

Emmons, Louise H. A new genus and species of rat from Borneo (Rodentia: Muridae) 752-761

Erseus, Christer. A new marine species of Smithsonidrilus (Oligochaeta: Tubificidae)
from the Florida Keys _. 587-590

Erseus, Christer and Michael R. Milligan. A new species of Uniporodrilus (Oligochaeta:
Tubificidae) from the Gulf of Mexico coast of Florida, and a phylogenetic analysis of
the genus 243-250

Feduccia, Alan and A. Bradley McPherson. A petrel-like bird from the late Eocene of
Louisiana: earliest record for the order Procellariiformes 749-751

Ferrari, Frank D. and Deborah K. Steinberg. Scopalatum vorax (Easterly, 1911) and
Scolecithricella lobophora Park, 1970, calanoid copepods (Scolecithrichidae) associ-
ated with a pelagic tunicate in Monterey Bay 467-489

Fugate, Michael. Branchinecta sandiegonensis, a new species of fairy shrimp (Crustacea:
Anostraca) from western North America 296-304

Gardner, Alfred L. and Monica Romo R. A new Thomasomys (Mammalia: Rodentia)
from the Peruvian Andes 162-11 A

Ghosh, H. C. and Raymond B. Manning. A new deep-sea crab of the genus Chaceon
from India (Crustacea: Decapoda: Geryonidae 714-718

Goodman, Steven M. and Florent Ravoavy. Identification of bird subfossils from cave
surface deposits at Anhjohibe, Madagascar, with a description of a new giant Coua
(Cuculidae: Couinae) 24-33

Graves, Gary R. A new intergeneric wood warbler hybrid (Parula americana x Den-
droica coronata) (Aves: Fringillidae) 402-409

Graves, Gary R. A new hybrid manakin (Dixiphia pipra x Pipra fdicauda) (Aves:
Pipridae) from the Andean foothills of eastern Ecuador 436—441

Hardy, Laurence M. Activity and reproductive patterns of amphibians and reptiles
from the Engare Ondare River region of central Kenya, during the dry season 740-748

Harvey, Michael B. and Eric N. Smith. A new species of aquatic Bufo (Anura: Bufonidae)
from cloud forests in the Serrania de Siberia, Bolivia 442-449

Hershler, Robert and France Velkovrh. A new genus of hydrobiid snails (MoUusca:
Gastropoda: Prosobranchia: Rissooidea) from northern South America 182-189

Hobbs, H. H., III. Cambanis (Jugicambarus) subterraneus, a new cave crayfish (De-
capoda: Cambaridae) from northeastern Oklahoma, with a key to the troglobitic
members of the subgenus Jugicambarus 719-727

Hobbs, Horton H., Jr. and Daniel J. Peters. New records of entocytherid ostracods
infesting burrowing and cave-dwelling crayfishes, with descriptions of two new species 455-466

Holt, Perry C. and Brent D. Opell. A checklist of and illustrated key to the genera and
species of the Central and North American Cambarincolidae (Clitellata: Branchiob-
dellida) 251-295

Hotchkiss, Frederick H. C. A new Devonian ophiuroid (Echinodermata: Oegophiurida)
from New York state and its bearing on the origin of ophiuroid upper arm plates ._ 63-84

Humes, Arthur G. and Geoffrey A. BoxshaU. Pseudonicothoe branchialis (Crustacea:
Copepoda: Siphonostomatoida: Nicothoidae), living on the pandalid shrimp Hetero-
carpus sibogae off northwestern Australia 315-324

Jezerinac, Raymond F. A new subgenus and species of crayfish (Decapoda: Cambaridae)

of the genus Cambanis, with an amended description of the subgenus Lacunicambarus 5 3 2-5 44

Jezerinac, Raymond F. and G. Whitney Stocker. A new species of crayfish (Decapoda:
Cambaridae) belonging to the genus Cambanis, subgenus Hiaticambanis, from the
upper Elk River drainage of West Virginia 346-352

Komai, Tomoyuki. Two new records of the genus Heptacarpus (Crustacea: Decapoda:
Hippolytidae) from Japanese waters 545-553

Kudenov, Jerry D. A new species of Sphaerodoridae (Annelida: Polychaeta) from south-
em California 582-586

Leo L., Mariella and Alfred L. Gardner. A new species of a giant Thomasomys (Mam-
malia: Sigmodontinae) from the Andes of northcentral Peru 417-428

Liao, Yulin and David L. Pawson. Caudina intermedia, a new species of sea cucumber
from the South China Sea (Echinodermata: Holothuroidea: Molpadiida) 366-368

Lima, Idalina Maria Brasil and Cristiana Silveira Serejo. A new species of Benthana

Budde-Lund from Brazilian caves (Crustacea: Isopoda: Oniscoidea) 490-496

Loffler, Deborah L. and Michael Vecchione. An unusual squid paralarva (Cephalopoda)
with tentacular photophores _ 602-605

Lopez-Gonzalez, Pablo J., Mercedes Conradi, and J. Carlos Garcia-Gomez. Enterocola
africanus, a new species (Copepoda: Ascidicolidae) associated with a compound as-
cidian Synoicum species from North Africa (Strait of Gibralter) „ 131-136

Manning, Raymond B. Two new species of Neocallichirus from the Caribbean Sea
(Crustacea: Decapoda: Callianassidae) 106-1 14

Manning, Raymond B. Three genera removed from the synonym oi Pinnotheres Bosc,

1 802 (Brachyura: Pinnotheridae) 523-53 1

Manning, Raymond B. and David K. Camp. Erythrosquilloidea, a new superfamily,
and Tetrasquillidae, a new family of stomatopod crustaceans 85-91

McCranie, James R., Larry David Wilson, and Kenneth L. Williams. A new species
of Oedipina (Amphibia: Caudata: Plethodontidae) from northern Honduras 385-389

Morino, Hiroshi and Reuven Ortal. The identity of Talitroides alluaudi (Chevreux)

(Crustacea: Amphipoda: Talitridae) with notes on a new locality 332-338

Ng, Peter K. L. On a new genus and species of xantid crab (Crustacea: Decapoda:

Brachyura) from Chesterfield Island, Coral Sea 705-713

Norman, Mark D. Octopus ornatus Gould, 1852 (Cephalopoda: Octopodidae) in Aus-
tralian waters: morphology, distribution, and life history 645-660

Opresko, Dennis M. A new species of Sibopathes (Cnidaria: Anthozoa: Antipatharia:

Antipathidae) from the Gulf of Mexico 195-203

Pereira S., Guido A. A description of a new species of Macrobrachium from Peru, and

distributional records for Macrobrachium brasiliense (Heller) (Crustacea: Decapoda:

Palaemonidae) 339-345

Perez, Linnette Garcia and W. Ronald Heyer. Description of the advertisement call

and resolution of the systematic status of Leptodactylus gracilis delattini Miiller, 1968

(Amphibia: Leptodactylidae) 51-56

Petit, Richard E. and M. G. Harasewych. A new Mericella (MoUusca: Gastropoda:

Cancellariidae) from northeastern Africa 221-224

Pettibone, Marian H. Polynoid polychaetes associated with a whale skeleton in the

bathy al Santa Catalina basin 678-688

Pleijel, Fredrik. Taxonomy of European species of Amphiduros and Gyptis (Polychaeta:

Hesionidae) 158-181

Pleijel, Fredrik. Gyptis crypta, a new hesionid species from the U.S.A. east coast, with

a redescription of G. vittata Webster & Benedict, 1887 (Annelida: Polychaeta) 237-242

Pleijel, Fredrik and Kristian Fauchald. Scalispinigera oculata Hartman, 1967 (Scali-

bregmatidae: Polychaeta): senior synonym of Lacydonia antarctica (Lacydoniidae)

Hartmann-Schroder & Rosenfeldt, 1988 eilt-eil

Price, Roger D. and Robert M. Timm. Two new species of Gliricola (Phthiraptera:

Gyropidae) from the spiny tree rat, Mesomys hispidus, in Peru 353-358

Pybum, William F. A new species of dimorphic tree frog, genus Hyla (Amphibia: Anura:

Hylidae), from the Vaupes River of Colombia 46-50

Rausch, V. R. and R. L. Rausch. Karyotypic characteristics of Sorex tundrensis Merriam

(Mammalia: Soricidae), a Nearctic species of the S. araneus-group 4 1 0-4 1 6

Reid, Janet W. and Teruo Ishida. New species and new records of the genus Elaphoidella

(Crustacea: Copepoda: Harpacticoida) from the United States 137-146

Remsen, J. V., Jr. Zoogeography and geographic variation of Atlapetes rufinucha (Aves:

Emberizinae), including a distinctive new subspecies, in southern Peru and Bolivia 429-435
Rice, Mary E. Two new species of Phascolion (Sipuncula: Phascolionidae) from tropical

and subtropical waters of the central western Atlantic 591-601

Robinson, Harold. A review of the genus Critoniopsis in Central and South America

(Veronieae: Asteraceae) 606-627

Robinson, Harold. Three new genera of Vemonieae from South America, Dasyandan-

tha, Dasyanthina, and Quechualia (Asteraceae) 775-785

Rodrigues, Sergio de Almeida and Rosana Moreira da Rocha. Littoral compound

ascidians (Tunicata) from Sao Sebastiao, estado de Sao Paulo, Brazil 728-739

Roman-Contreras, Ramiro. Probopyrus pacificensis, a new parasite species (Isopoda:

Bopyhridae) of Macrobrachium tenellum (Smith, 1871) (Decapoda: Palaemonidae) of

the Pacific coast of Mexico 689-697

Rozbaczylo, N. and J. J. Canete. A new species of scale- worm, Harmothoe commensalis

(Poychaeta: Polynoidae), from mantle cavities of two Chilean clams 666-672

Rudjakov, J. A. The first finding of the male of Thaumatocypris echinata Miiller, 1906

(Crustacea: Ostracoda) 305-314

Saltzman, Jennifer and Thomas E. Bowman. Boreomysis oparva, a new possum shrimp

(Crustacea: Mysidacea) from an eastern tropical Pacific seamount 325-331

Schotte, Marilyn and Raymond B. Manning. Stomatopod Crustacea from Tobago, West

Indies 566-581

Smolen, Michael J., Richard M. Pitts, and John W. Bickham. A new subspecies of

pocket gopher (Geomys) from Texas (Mammalia: Rodentia: Geomyidae) 5-23

Solis- Weiss, Vivianne. Grassleia hy dr other malis, a new genus and species of Amphar-

etidae (Annelida: Polychaeta) from the hydrothermal vents off the Oregon coast (U.S.A.),

at Gorda Ridge 661-665

Takeuchi, Ichiro. Caprella arimotoi, a new species (Crustacea: Amphipoda: Caprellidea)

from the Seto Inland Sea, Japan 115-121

Thuesen, Erik V. Vampyrocrossota childressi, a new genus and species of black medusa

from the Bathypelagic zone off California (Cnidaria: Trachy medusae: Rhopalone-

matidae) 190-194

Weems, Robert E. and Peter G. Kimmel. Upper Triassic reptile footprints and a coel-

acanth fish scale from the Culpepper Basin, Virginia 390—401

Wynn, Addison H. and Alan E. Leviton. Two new species of blind snake, genus Typhlops

(Reptilia: Squamata: Typhlopidae), from the Philippine Archipelago 34-45

INDEX TO NEW TAXA

Volume 106

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

PORIFERA

Demospongea

Ceratopsion crustosum 629

CNIDARIA

Hydrozoa

Vampyrocrossota 190

childressi 191

Anthozoa

Sibopathes macrospina 195

PLATYHELMINTHES

Blairium, new subgenus 214

Gibsonium, new subgenus 215

Girardia canai 633

Maillardiella, new subgenus 212

Overstreetium, new subgenus 213

Timoniella ostrowskiae :. 208

MOLLUSCA

Gastropoda

Andesipyrgus 182

sketi 183

Mericella bozzettii 221

SIPUNCULOIDEA

Phascolion gerardi 591

psammophilum 594

ANNELIDA

Polychaeta

Amphiduros fuscescens n.c 176

Flascarpia 153

alvinae 153

Grassleia 662

hydrothermalis 662

Gyptis crypta 238

mackiei 165

mediterranea 168

Harmothoe commensalis 667

craigsmithi 683

Lacydonia oculata n.c 674

Laubierus 151

mucronatus ..^ 151

vui

Miralvinella, new subgenus..
Miura

spinosa

Nautalvinella, new subgenus.
Paral vinella fijiensis

unidentata

Peinaleopolynoe santacatalina.....

Santelma

miraseta n.c

Sphaerodoropsis sexantennella....

Smithsonidrilus exspectatus .
Uniporodrilus /7Mn^5

Oligochaeta

ARTHROPODA
Crustacea

Alachosquilla

digueti n.c

floridensis n.c

vicina n.c

A nomoeomunida

Benthana iporangensis^^
Bigelowina

biminiensis n.c.

235
148
148
235
232
226
680
149
150
582

587
244

572
572
573
572
102
490
574
574
325
296
346
719
535
536
115
714
705
707
459
141
138
132

Boreomysis oparva

Branchinecta sandiegonensis

Cambarus (Hiaticambarus) elkensis

Cambanis (Jugicambanis) subterraneus

Cambarus (Tubericambanis) acanthura n.c

Caprella arimotoi

Chaceon alcocki

Dactylocythere cryptoteresis

Elaphoidella amabilis

carterae

Enterocola africanus

ERYTHROSQUILLOIDEA 87

Gonodactylus caribbaeus 568

Juxtafabia 93

muliniarum n.c 96

Kalliapseudes (Mesokalliapseudes) soniadawnae 122

Linca „ 497

pinita „. „ 499

Macrobrachium depressimanum 339

Nannosquilla tobagoensis 575

Neocallichirus lemaitrei 107

nickellae 1 10

Phymocythere lophota 463

Potamalpheops danviniensis 698

Probopyrus /7(2c/^^«5/5 690

Pylopagurus par?/<3£' 554

Strengeriana cajaensis 508

flagellata 512

maniformis 51 1

TETRASQUILLIDAE 88

Tubericambarus, new subgenus 534

Xylopagurus tayrona 559

Insecta

Gliricola /2a/// 354

woodmani 353

CHAETOGNATHA

Spadella 7(2/7on/ca 359

ECHINODERMATA

Caudina intermedia 366

Strataster maciverorum 66

wrighti n.c 66

CHORDATA

Pisces

Hyporhamphus meeki 371

Prochilodus britskii 58

Amphibia

Bufo ambowensis 443

Hyla karenanneae 46

Otdvpindi gephyra 385

Reptilia

Typhlops castanotus 35

collaris 41

Aves

Atlapetes rufinucha terborghi 429

Coua berthae 26

Mammalia

Geomys texensis bakeri 19

Pithecheirops 752

otion 753

Thomasomys apeco 417

macrotis 762

PLANTAE

Critoniopsis cotopaxensis 610

dorrii 612

lewisii 615

palaciosii 617

quillonensis 620

steinbachii 622

uribei 625

Critoniopsis, 38 n.c.'s 609-626

Dasyandantha 778

cuatrecasasiana n.c 778

Dasyanthina 778

palustris n.c 780

serrata n.c 780

Quechnalia 780

cardenasii n.c 78 1

fiilta n.c 78 1

smithii 781

PROCEEDING^

OF THE

BIOLOGICAL SOCIETY

WASHINGTON

VOLUME 106 NUMBER 1
8 APRIL 1993

ISSN 0006-324X

THE BIOLOGICAL SOCIETY OF WASHINGTON

1992-1993
Officers

President: Storrs L. Olson
President-elect: Janet W. Reid

Stephen D. Cairns
Richard C. Froeschner
Alfred L. Gardner

Elected Council

Secretary: G. David Johnson
Treasurer: T. Chad Walter

Jon L. Norenburg

Lynne R. Parenti

F. Christian Thompson

Custodian of Publications: Austin B. Williams

PROCEEDINGS

Editor: C. Brian Robbins

Associate Editors

Classical Languages: George C. Steyskal

Plants: David B. Lellinger
Insects: Wayne N. Mathis

Invertebrates: Jon L. Norenburg
Frank D. Ferrari
Rafael Lemaitre

Vertebrates: Thomas A. Munroe

Membership in the Society is open to anyone who wishes to join. There are no prerequisites.
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PROC. BIOL. SOC. WASH.
106(1), 1993, pp. 1-4

ON THE IDENTITY OF ECHIMYS DIDELPHOIDES

SMAREST, 1817
(MAM]S^LIA:>|RODENTIA: ECHIMYIDAE)

.ouise H. Emmons

/xv

Abstrd^Tr^'-'^t^mr^rXy names for species of the genus Echimys {Makalatd),
E. didelphoides and Loncheres obscura, were erroneously assigned to the genus
Mesomys by Tate (1935). One of these, didelphoides, is the oldest name for the
red-nosed tree rats currently known as Makalata armata.

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

Taxonomic History

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

Geoffroy St.-Hilaire (1838) noted that
Lichtenstein's specimen was misidentified,
and, without seeing the specimen, renamed
it Nelomys armatus. In his complete revi-
sion of the echimyid rodents, Geoffroy St.-
Hilaire (1840) explained that armatus re-
sembled didelphoides except in details of
color, length of the haired part of the tail,
tail length, and width of the spines (p. 1 1),

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

Wagner (1843) commented that Echimys
didelphoides was closely related to Lonche-
res armata: Wagner, 1843, and perhaps
simply a young one. Waterhouse ( 1 848) said
the differences between the two did not seem
to warrant their separation.

Tate (1935) without seeing the specimens,
placed didelphoides in the genus Mesomys,
where it has remained since (e.g., Cabrera
1960, Honacki et al. 1982, Corbet & Hill
1991). At the same time, Tate (1935) also
placed Loncheres obscura Wagner, 1 840, in
the genus Mesomys.

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

Identity of the Holotype of
Echimys didelphoides

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

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

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

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

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

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

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

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

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

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

VOLUME 106, NUMBER 1

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

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

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

Nelomys armatus I. Geoffroy Saint-Hi-

laire, 1838
Loncheres guianae T\iov[i2iS, 1888
Echimys longirostris Anthony, 1921
Echimys castaneus Allen & Chapman,

1893

Identity of Loncheres obscura

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

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

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

Acknowledgments

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

Literature Cited

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

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

Cuvier, F. 1832. Description des characteres propres
aux genres Graphiureet Cercomys de I'ordre des
rongeurs. — Nouvelles Annates du Museum
d'Histoire Naturelle 1:449-452, pi. 18 fig. 2.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

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

GeofFroy Saint-Hilaire, I. 1838. Notice sur les ron-
geurs epineux designes par les auteurs sous les
noms d' Echimys, Loncheres, Heteromys et Ne-
/owy^.— Revue Zoologique 1:99-101.

. 1840. Notice sur les rongeurs epineux de-
signes par les auteurs sous les noms d^ Echimys,
Loncheres, Heteromys et A^6'/omy5. — Magazin
de Zoologie, Serie 2, 2:1-57, pis. 20-29.

Honaki, J. H., K. E. Kinman, & J. W. Koppl. 1982.
Mammal species of the world. Allen Press and
Association of Systematics Collections, Law-
rence, Kansas, 694 pp.

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

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

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

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

Wagner, J. A. 1 840. II. Stachelmalise. Abhandlungen
(Bayerische) Akademie Wissenschaften, Mu-
nich. Pp. 191-210, plate II.

. 1842. Diagnosen neuer Arten brasilischer

Saugethiere. — Archiv fiir Naturgeschichte 8:356-
362.

. 1843. Die Saugethiere in Abbilddungen nach

der Natur mit beschreibungen von Dr. Johann
Christian Daniel von Schreber. Leipzig. Sup-
plementband Erlangen, Expedition das Schre-
ber'schen Saugthier- und des Esper'sshen
Schmetterlingswerkes, und in Commission der
Voss'schen Buchhandlung in Leipzig, 3:xiv +
614 pp., pis. 85-165.

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

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

U.S.A.

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

A NEW SUBSPECIES OF POCKET GOPHER (GEOMYS) FROM
TEXAS (MAMMALIA: RODENTIA: GEOMYIDAE)

Michael J. Smolen, Richard M. Pitts, and John W. Bickham

Abstract.— Two isolated populations of pocket gophers were discovered far
outside the previously known distribution of geomyids in southern Texas. These
represent a new taxon, which is related to Geomys texensis in chromosome
morphology and biochemical properties. The two share identical diploid num-
ber (2N = 70), fundamental number (FN = 68), and the presence of a distinctive
large acrocentric X-chromosome. There are no fixed differences in any of the
1 8 loci analyzed with starch gel electrophoresis, when comparing the new taxon
with G. texensis. However, both of these taxa share alleles at two loci that are
present as fixed differences when compared to G. bursarius major. Analysis of
cranial morphology indicates that the new taxon resembles G. texensis, but
differs significantly in cranial dimensions of size and shape, related primarily
to measurements of basal and palatal lengths and mastoid breadth. Because of
the morphological distinctiveness and the extreme spatial separation (120 km)
from populations of G. texensis confined to the central basin of the Edwards
Plateau, we conclude that this new taxon is an isolated, relictual population of
G. texensis, and is a distinctive subspecies.

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

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

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

Materials and Methods

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

102

101

100

102

101

100

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

VOLUME 106, NUMBER 1

mosome morphology were described from
photographic prints of selected spreads.

Samples representing the unknown taxon
(n = 21), G. texensis (n = 13), and G. bur-
sarius {n= 10) were used to assay biochem-
ical variation. Heart and kidney tissues were
minced in a grinding solution (Tris/EDTA/
NADP) and homogenized using a mechan-
ical homogenizer (Tissue Tearor, Biospec
Products). Samples were loaded into 12%
starch gels (Starch Art). The techniques used
for visualizing the allozymes were those de-
scribed by Harris & Hopkinson (1976) and
Honeycutt & Williams (1982). Proteins were
examined on the following buffer systems:
Poulik, continuous Tris-citrate II (pH 7.0
and 8.0), Tris-malate EDTA (pH 7.4), and
Ridgway (pH 6.7). Eighteen presumptive
loci and their respective Enzyme Commis-
sion numbers (Murphy et al. 1990) are as
follows: glycerol- 3 -phosphate dehydroge-
nase (G3PDH; E.G. 1.1.1.8), malate dehy-
drogenase (MDH-1,2; E.G. 1.1.1.37), isoci-
trate dehydrogenase (IDH-1,2; E.G.
1.1.1.42), phosphogluconate dehydroge-
nase (6-PGDH; E.G. 1.1.1.44), superoxide
dismutase(SOD-l,2; E.G. 1.15.1.1), purine-
nucleoside phosphorylase (PNP; E.G.
2.4.2.1), aspartate aminotransferase (AAT-
1,2; E.G. 2.6.1.1), creatine kinase (GK; E.G.
2.7.3.2), phosphoglucomutase (PGM-1; E.G.
5.4.2.2), esterase (EST- 1,2; E.G. 3.1.1.-),
peptidase (PEP-1 leucyl glycyl glycine,
PEP-2 leucyl alanine; E.G. 3.4.-.-), aconitase
hydratase (AGOH; E.G. 4.2.1.3), glucose-
phosphate isomerase (GPI; E.G. 5.3.1.9).
The allozyme data were scored in a side by
side comparison of mobility where identical
mobilities were recorded as individual al-
leles. Data were analyzed using BIOSYS-1
(Swofford & Selander 1981), which pro-
duced Rogers' genetic similarity and genetic
distance values for the three taxa (Rogers
1972).

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

yses of morphological data. Geomys attwa-
teri was represented by two populations from
the western extent of its distribution and
thus in relatively close proximity to the new
taxon. Geomys personatus streckeri was se-
lected as a representative of the personatus
complex because its distribution also ap-
proaches that of the new gopher. Two pop-
ulations of G. texensis represent the previ-
ously recognized subspecies, G. bursarius
texensis and G. bursarius llanensis. They
were recently elevated to specific ranking
and synonymized as the monotypic G. tex-
ensis (Block & Zimmerman 199 1) based on
patterns of biochemical variation. Although
neither G. texensis nor G. bursarius are
closely distributed to the new taxon, they
are included because they are karyotypically
identical to it.

Geomyids show extensive variation in
secondary sexual characteristics (Baker &
Genoways 1975; Honeycutt & Schmidly
1979; Williams & Genoways 1977, 1978,
1980, 1981), which necessitated separate
analysis of males and females. Three exter-
nal measurements were recorded from spec-
imen labels, and 1 1 cranial characters were
measured using dial calipers. Skull mea-
surements were taken as described by Wil-
liams & Genoways (1977). Measurements
we evaluated are as follows: total length (TL),
length of tail (T), length of hind foot (HP),
condylobasal length (GBL), basal length
(BL), palatal length (PL), prefrontal depth
(PFD), length of nasals (LN), diastema
(DIAS), zygomatic breadth (ZB), mastoid
breadth (MB), squamosal breadth (SB), ros-
tral breadth (RB), interorbital constriction
(lOG), breadth across third molars (M3),
length of maxillary toothrow (Ml). Only
adult animals were used and they were iden-
tified by the degree of ossification of the
sutures of the skull, particularly the fusion
of the basisphenoid and basioccipital bones
(Williams & Genoways 1981). The aduhs
were classified into two categories based on
the degree of fusion and ossification. The

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

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

Phenetic relationships were assessed us-
ing correlation and distance matrices

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

Results

The diploid number of 1 6 specimens rep-
resenting the new populations in Medina,
Zavala, and Uvalde counties is 2N = 70,
and the fundamental number is 68. The
X-chromosome is a large acrocentric and
the Y-chromosome is medium-sized and
acrocentric. The morphology of the chro-
mosomes is indistinguishable from those
previously reported for both G. bursarius
and G. texensis (Baker et al. 1973, Hart
1978).

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

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

VOLUME 106, NUMBER 1

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

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

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

G. I. bakeri

G. texensis

G. bursarius

Pep-1

SOD-1

EST-1

A, B

EST-2

A,b

A, B

GPI

A, B

G3PDH

A, B

A,B

PGM-1

A, B

6-PGD

A, B

MDH-1

A, B

MDH-2

A, b

A, B

SOD-2

A, B

AAT-1

A,b

IDH-1

A,C

B, C

IDH-2

Pep-2

ACOH

PNP

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

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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VOLUME 106, NUMBER 1

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

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

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

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

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

Discussion

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

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

Males

G. t. baked
G. texensis
G. texensis
G. personatus
G. attwateri
G. attwateri

G. t. bakeri
G. texensis
G. texensis
G. attwateri
G. attwateri
G. personatus

G. t. bakeri
G. personatus
G. attwateri
G. attwateri
G. texensis
G. texensis

M3
G. t. bakeri
G. texensis
G. texensis
G. attwateri
G. personatus
G. attwateri

Ml
G. t. bakeri
G. texensis
G. texensis
G. attwateri
G. attwateri
G. personatus

IOC

G. texensis
G. texensis
G. t. bakeri
G. personatus
G. attwateri
G. attwateri

21.96

21.4

22.3

22.65

20.8

23.9

23.25

21.9

23.9

23.98

22.3

26.7

24.06

22.6

25.7

24.11

23.5

24.7

16.94

16.6

17.4

17.52

16.9

18.0

17.63

16.8

18.5

18.22

17.4

19.5

1S.63

18.1

19.5

18.69

17.5

19.8

9.19

8.9

9.5

9.37

8.9

10.7

9.43

9.1

9.9

9.74

8.9

10.4

9.95

9.8

10.2

10.17

9.3

10.9

6.90

6.8

7.1

7.48

7.2

7.8

7.61

7.3

8.0

7.74

7.1

8.1

7.80

7.3

8.8

7.92

7.6

8.5

8.49

8.2

8.8

8.63

8.1

9.0

8.91

8.6

9.6

9.36

9.1

9.8

9.36

9.1

9.6

9.43

8.4

10.6

5.87

5.5

6.3

5.88

5.6

6.2

5.96

5.7

6.4

6.18

5.9

6.6

6.40

6.0

6.7

6.53

6.1

6.9

served between G. texensis and the new tax-
on. Block & Zimmerman (1991) identified
fixed differences between G. bursarius and
G. texensis in a study involving species of

geomyids from central Texas. Genetic sim-
ilarities determined by them for G. bursari-
us and populations of G. texensis ranged
from 0.607 to 0.648, whereas genetic sim-

VOLUME 106, NUMBER 1

Table 3.— Continued.

Females

G. t. bakeri
G. texensis
G. texensis
G. personatus
G. att water i
G. attwateri

G. t. bakeri
G. texensis
G. texensis
G. attwateri
G. attwateri
G. personatus

G. t. bakeri
G. attwateri
G. personatus
G. texensis
G. texensis
G. attwateri

M3
G. t. bakeri
G. texensis
G. attwateri
G. attwateri
G. texensis
G. personatus

G. t. bakeri
G. texensis
G. texensis
G. attwateri
G. personatus
G. attwateri

IOC

G. texensis
G. t. bakeri
G. texensis
G. personatus
G. attwateri
G. attwateri

21.14

20.5

21.9

21.36

19.9

22.8

21.77

20.8

23.0

22.93

21.7

24.0

23.30

21.6

24.8

23.30

22.8

24.1

16.65

16.3

17.1

16.85

16.5

17.3

16.92

16.0

17.9

17.97

16.9

18.7

18.01

17.7

18.3

18.21

17.3

18.9

8.40

8.0

8.8

8.56

8.2

9.7

8.87

8.1

9.5

9.15

8.5

9.6

9.37

9.0

10.1

9.60

9.0

10.1

6.88

6.5

7.3

7.36

7.0

7.8

7.50

7.4

7.6

7.51

7.0

8.0

7.52

7.1

8.1

7.72

7.2

8.2

8.25

7.8

8.5

8.51

7.9

9.3

8.66

8.1

9.0

8.99

8.5

9.4

9.08

8.5

9.5

9.21

9.0

9.5

5.76

5.5

5.9

5.83

5.5

6.3

5.86

5.4

6.3

6.07

5.8

6.6

6.28

5.6

6.7

6.49

6.2

6.9

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

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

2.0 -■

< 0.0 4

-2.0 -■

-4.0 -■

t. bakeri

♦

p. streckeri

attwoteri

texensis

-6.0 -4.0

-2.0

0.0
CAN

2.0

4.0

6.0

-6.0

-4.0

-2.0 0.0
CAN

2.0

4.0

6.0

4.0 --

2.0-

0.0 --

-2.0. .

-4.0 -

Females

0.0 --

-2.0 --

-4.0 -■

Males

CP ■

-6.0 -4.0 -2.0 0.0 2.0 4.0 6.0

CAN I

-6.0 -4.0 -2.0 0.0 2.0 4.0 6.0

CAN I

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

ity values, seen when comparing the new
taxon to G. bursarius, are within the range
associated with interspecific comparisons
(Block & Zimmerman 1991, Dowler 1982).
Relationship between G. texensis and the
new taxon is supported further by the mor-
phometric analysis. In an analysis of the
cranial characters that appear to separate G.
personatus and G. attwateri from G. tex-
ensis, the new taxon was associated with G.
texensis in mastoid breadth, squamosal
breadth, interorbital constriction, and length

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

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

VOLUME 106, NUMBER 1

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

Canonical
Variate I

Canonical
Variate II

Canonical
Variate III

Character

Variable
coeff.

Males

CBL

-1.8433

28.51

-0.3363

3.36

-4.3026

41.24

0.6980

10.14

-4.5368

42.57

1.8989

17.09

-0.6005

6.09

2.1302

13.95

2.7239

17.11

PFD

-0.9800

5.87

-0.6656

2.58

0.9685

3.60

DIAS

1.1349

6.15

2.1888

7.67

-0.5400

1.81

-1.1519

11.26

0.6130

3.87

0.5727

3.47

2.0305

17.87

2.2454

12.77

-2.0193

11.01

0.8427

5.67

-1.8478

8.03

0.3642

1.52

-0.5925

2.10

0.0785

0.18

-0.1217

0.27

IOC

-0.6612

1.52

0.8264

1.23

0.6400

0.91

0.8382

2.38

-1.2829

2.35

0.6570

1.16

0.7154

2.44

0.6592
Females

1.45

0.3881

0.82

CBL

-1.9191

35.85

0.7594

15.53

-0.0083

0.28

0.4529

7.94

-1.7802

34.17

-0.2815

8.76

0.5315

6.44

0.9833

13.05

1.3095

28.18

PFD

-0.4949

3.63

0.1148

0.92

-0.3503

4.56

DIAS

0.6719

4.25

-1.1497

7.96

-0.7237

8.13

0.1029

1.18

0.3581

4.49

0.5166

10.49

2.4876

26.45

-0.5207

6.06

0.5198

9.81

-0.3687

3.06

0.9749

8.87

-1.3019

19.20

-1.0596

4.56

0.7393

3.48

0.6176

4.71

IOC

0.7276

2.09

-0.2932

0.92

0.7603

3.87

-0.4172

1.50

0.9869

3.88

-0.0849

0.54

0.7203

3.05

0.1459

0.68

-0.1930

1.45

ulations, isolated and 1 20 km distant from
the previously known distribution of G. tex-
ensis, constitute a new subspecies? An anal-
ysis of cranial morphology identifies the new
populations as being quite distinct from G.
texensis. They are smaller in 22 of 28 com-
parisons involving both sexes. Canonical
discriminant analysis identifies measure-
ments reflecting the length of the skull (BL,
PL) and mastoid breadth accounting for
much of the variation separating the taxa.
The centroids produced from plotting these
taxa along the first two canonical variates
are widely separated, although the ellipses
are overlapping. Separation is more clearly
seen in analysis of females, where the two
populations of G. texensis are closely as-
sociated with each other, and each is well

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

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Females

-- 2.0

-- 0.0

---2.0

C_3

Males

-- 2.0

-- 0.0

-2.0

Females
0.99

Correlation Matrix

1.00

G. p. streckeri
G. attwaterl
G. attwateri
G. texensis
G. texensis
G. t. bakeri

Males
0.99

Correlation Matrix

1.00
—I

G. p. streckeri

G. attwateri

G. texensis

G. t. bakeri

Females

1.12

-I

Males

0.64

—I

Distance

0.16

— I

G. p. streckeri
G. texensis
G. texensis
G. t. bakeri
G. attwateri
G. attwateri

1.75

-I

1.00

Distance

0.25

—I

G. p. streckeri
G. texensis
G. texensis
G. attwateri
G. attwateri
G. t. bakeri

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

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

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

VOLUME 106, NUMBER 1

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

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

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

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

Geomys texensis bakeri, new subspecies

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

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

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

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

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

Measurements.— MQasuTQmQnts (in mil-
limeters), as described in Williams & Geno-
ways (1977), from 1 2 adult individuals from
the two populations are listed in Table 4.
Measurements of the holotype (TCWC
52310) are: total length, 227; length of tail,
66; length of hind foot, 27; length of ear, 6;
greatest length of skull, 40.8; condylobasal
length, 40.4; basal length, 38.3; palatal
length, 26.3; prefrontal depth, 15.5; length
of nasals, 13.3; length of diastema, 14.3;
zygomatic breadth, 25.6; mastoid breadth,
22.4; squamosal breadth, 16.9; rostral
breadth, 9.4; interorbital constriction, 6.3;
breadth across third molars, 7.1; length of
maxillary toothrow, 8.5.

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

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

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

VOLUME 106, NUMBER 1

Acknowledgments

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

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gie Museum 50:435-473.

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

Appendix

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

Geomys attwateri.—(S5). Texas: Atascosa Co.: 6 mi
W Campbellton, l(TNHC); 2 mi NW Campbellton,
3(TCWC); 1 mi E Lytle, 16(TCWC); 7 mi E Lytle,
16(TNHC); 2.4 mi SE Lytle, 3(TCWC); 7 mi SE Lytle,
8(TCWC). Bexar Co.: 1 5 mi SE San Antonio, 1 (TSU).
Frio Co.: 1 mi N Moore, 3(TCWC); Pearsall city limits,
2(TCWC); 2.25 mi S, 1 mi E Pearsall, l(TCWC); Mc-
Coy, l(TNHC); 0.3 mi E McCoy, l(TNHC); 2 mi N
Pleasanton, 7(TNHC). Medina Co.: 5 mi W Devine,
4(TCWC); 7.2 miE Yancy, l(TCWC), l(TSU). Wilson
Co.: 11 mi NW Horesville on HWY 181, 6(TCWC);
4 mi W Floresville, l(TNHC); 1 mi W Floresville,
2(TCWC); % mi S, 2V2 mi E Floresville, 4(TCWC); 3.2
mi NW Poth, l(TNHC); 3.6 mi SSE Poth, l(TNHC);
5.4 mi W San Antonio River, between Floresville and
Pleasanton, l(TNHC).

Geomys personatus streckeri.— (68). Texas: Dimmit
Co.: 13 mi N Carrizo Springs on HWY 277, 3(TTU);
1 3 mi NE Carrizo Springs, 3(TTU); 1 3 mi NE Carrizo
Springs on US HWY 277, 5(TTU); 4 mi N, 9 mi W
Carrizo Springs, l(TCWC); 1 mi S Carrizo Springs,
700 ft, 2(TCWC); 15 mi S, 1 1 mi W Carrizo Springs,

VOLUME 106, NUMBER 1

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

Geomys texensis bakeri.— (35). Texas: Medina Co.:
1 mi E D'Hanis, 5(TCWC); D'Hanis, 5(TCWC); SVi
mi W Hondo, 6(TCWC); 6.2 mi W Hondo, 4(TCWC).
Uvalde Co.: 16 mi S Sabinal on FM 187, 6(TCWC);

17 mi S Sabinal on FM 187, 4(TCWC). Zavala Co.:

1 8 mi S Sabinal on FM 1 87, 1 (TCWC); y,o mi S Uvalde
County line, 4(TCWC).

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

2(TTU); 2.9 mi NW Llano on HWY 71, 2(TTU); 0.2
mi N, 8.7 mi W Llano, 3(TTU); Drier Cr at Lone
Grove, 7 mi W Llano, lO(TCWC); 4 mi W Llano
l(TCWC), Llano, 1(TCWQ; 0.2 mi E Llano, l(TCWC);
1 mi E Llano, 2(TCWC); 2 mi E Llano, 4(TCWC); 7.2
mi E Llano, l(TCWC); Oatman Cr, 3 mi S Llano,
6(TCWC); 51.6 mi W Austin on HWY 71, 2(TTU); 3
mi S Jet FM 268 and HWY 29 on 29, 2(TTU); 9.3 mi
N Jet Texas 29, Texas 1 6 on Texas 16,1 (TTU). Mason
Co.: 3. 1 mi E Art, l(TCWC); Art city limits, 3(TCWC);
1 2 mi N Mason, 6(MWU); 3.6 mi N, 1 .5 mi W Mason,
l(TTU); 1 mi N, 1.1 mi W Mason, 4(TTU); 12 mi W
Mason, 2(MWU); 9.4 mi W Mason on US 377, 3(TTU);
Mason, l(TCWC); 1 mi E Mason, 6(TCWC); 6.5 mi
E Mason on Texas 29, l(TTU); 2.0 mi S, 2.7 mi W
Castell, l(TTU); 5 mi S Mason, l(MWU). in Mason
Co.: 3'/2 mi W Castell, 3(TCWC); 0.3 mi S, 1.5 mi W
Castell, 3(TTU); 0.3 mi S, 0.9 mi W Castell, l(TTU);
0.3 mi S, 0.8 mi W Castell, l(TTU); 2.6 mi S, 3.0 mi
W Castell, l(TTU); 0.7 mi S, 2. 1 mi W Castell, 2(TTU);
1 .0 mi S, 2.3 mi W Castell, 2(TTU); 1 1 mi NE London,
HWY 377, 2(TCWC); 13 mi NE London, HWY 377,
2(TCWC).

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

IDENTIFICATION OF BIRD SUBFOSSILS FROM CAVE

SURFACE DEPOSITS AT ANJOHIBE, MADAGASCAR,

WITH A DESCRIPTION OF A NEW GIANT

COUA (CUCULIDAE: COUINAE)

Steven M. Goodman and Florent Ravoavy

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

Resume.— VnQ collection d'ossements subfossiles d'oiseau provenant des
gisements cavemicoles de surface du Nord-Ouest de Madagascar a ete decrite.
La plupart des specimens representant des taxons qui'existent encore dans la
region. La seule exception est constituee par un bassin incomplet pouvant se
rapporter au genre Coua (Famille Cuculidae: sous-famille Couinae), mais qui
est nettement plus grand que toute espece connue. Ce specimen est decrit
comme nouvelle espece.

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

The subfossil fauna ofMadagascar is well son & James 1991). While there are nu-
known for its remarkable array of lemurs merous archaeological sites on Madagascar
(e.g., Vuillaume-Randriamanantena 1982, that have yielded bird bones, this material
Simons et al. 1 990) and elephant birds (Ae- with the exception of elephant birds has been
pyomithiformes) (Andrews 1894, Lamber- rarely studied and thus not synthesized into
ton 1934, Battistini 1965). The study of the current working knowledge of the Ho-
hundreds of animal bones recovered at var- locene environment of the island,
ious sites on Madagascar has provided in- In 1983 and 1986 excavations were car-
sight into the Holocene faunas of the island, ried out in northwestern Madagascar near
inference about ecological change, as well Mahajanga in the Grottes d'Anjohibe (An-
as the reasons that a portion of these taxa dranoboka), by the Laboratoire de Prima-
have gone extinct in the past few thousand tologie et de Paleontologie des Vertebres,
years (Dewar 1984, MacPhee 1986). In par- Service de Paleontologie, Universite d'An-
allel situations on other islands, subfossil tananarivo (formerly Universite de Mada-
bird bones have provided important infor- gascar), and Duke University Primate Cen-
mation about paleoenvironments and the ter. The focus of these studies was primates
effects of anthropogenic perturbations (Vuillaume-Randriamanantena et al. 1985,
(Steadman 1989, James & Olson 1991, Ol- Simons et al. 1990), but a wide array of

VOLUME 106, NUMBER 1

Other animal remains were recovered. Dur-
ing the 1986 field season approximately
1 1 00 whole or fragmented subfossil bones
of non-primate vertebrates were excavated,
and, excluding the bird material, these have
been described by Ravoavy (1991). In this
paper we present information on 94 speci-
mens of avian bones recovered during the
1983 and 1986 field seasons at Anjohibe.

Description of Site

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

The cave is over 1 200 m long from north
to south, and with floor to ceiling heights
in some places exceeding 12 m. There are
numerous entrances and side passages to the
cave. It still has "active" calcite formations
(de Saint-Ours & Paulian 1953), and has
been used in recent times by local people
for a variety of activities (de Saint-Ours
1953).

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

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

Minimum number
of individuals

Taxa

1983

1984

Buteo brachypterus

Falco newtoni

Coturnix sp.

Numida meleagris

Turnix nigricollis

Coracopsis vasa

Tyto alba

Otus rutilus

Merops superciliosus

Hypsipetes madagascariensis

Foudia madagascariensis

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

Birds Recovered from the
Surface Deposits

A total of 1 2 and 82 bird bones recovered
during the 1983 and 1986 field seasons (re-
spectively) was used in this study. Most of
the species identified (Table 1) are taxa that
still occur in the area (Langrand 1990). The
present natural vegetation of the region is
dry deciduous forest; considerable portions
of this habitat have been degraded in the
past few centuries. The majority of birds
identified from the surface deposits are spe-
cies associated with open habitats and/or
the forest edge. The hawk {Buteo brachyp-
terus), falcon (Falco newtoni), parrot {Cor-
acopsis vasa), and owl {Tyto alba) presently
occur in a variety of habitats from grassland
savanna to wooded environments. A par-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

The most common bird species recovered
from these surface deposits was Numida.
Adults of this species weigh over 1 1 00 g
(Urban et al. 1986) and are too heavy to
have been carried into the cave by any rap-
tor known to occur on the island. Numida
may have been introduced to Madagascar,
and it is now extensively hunted on the is-
land by people (Langrand 1990). There was
no clear sign of carnivore gnawing, butch-
ering marks nor charring on the Numida
bones, nor on any of the other bird material
recovered from the cave. It is not clear what
agent(s) was (were) responsible for the de-
position of these bones in the cave. On a
few occasions during the excavations Numi-
da were observed flying in and out of the
cave in an area with a collapsed ceiling and
extensive vegetation. Thus, the Numida
bones recovered from the surface deposits
may be of individuals that naturally died
within the cave.

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

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

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

Coua berthae, new species
Figs. 1, 2

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

VOLUME 106, NUMBER 1

Fig. 1 . Pelvis of Coua berthae, new species, holotype UM 6264, (left) dorsal and (right) ventral views.

leontologie des Vertebres, Service de Pa-
leontologie, Universite d'Antananarivo,
UM 6264 (Figs. 1, 2). Collected during the
1983 field season.

Locality. —From surface deposits, Grotte
d'Anjohibe (Andranoboka), Grotte Prin-
cipaleno. 1 (de Saint-Ours & Paulian 1953),
Fivondronana (subprefecture) of Mahajan-
ga. Province of Mahajanga, about 80 km
NE Mahajanga, Madagascar (coordinates:

1) Laborde system— X = 1 172, y = 448 and

2) 15°32'S, 46°53'E).

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

Measurements of holotype. —hQngth —
from cranial border of the ilia to the Spinae
iliocaudales, 68.2 mm; length along the ver-
tebrae—from most cranial vertebra fused
with the Os lumbosacral to the most caudal
vertebra fused with the Os lumbosacral,

58.1 mm; smallest breadth across the Partes
glutaeae of the ilia, 18.3 mm; and greatest
breadth across the Partes glutaeae of the ilia,

29.2 mm. (See von den Driesch, 1976, fig.
59a, c for illustrations and descriptions of
these measurements.)

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

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

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

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

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

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

A.>.

2cm

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

length of the tarsometatarsus (Table 2). C surements in this species than the other

cristata shows some geographic variation in three. Within these four species there is no

size (Milon 1950), which accounts for the discemable sexual dimorphism in the skel-

greater variability in the range of these mea- etal measurements. A strong linear rela-

VOLUME 106, NUMBER 1

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

Pelvis

Tarsometatarsus

Species

Length'

Length along
the vertebrae

Cranial
breadth

Smallest
breadth of ilia

Greatest
length

berthae, sp. nov.^

68.2

58.1

-29.2

18.3

92.9

primavea

[62.5]

—

83.23

gigas{n = 1)

49.3

42.3

25.8

12.5

68.7

caerulea

44.28

38.70

24.20

13.75

55.20

n = 5

n = 6

n = 5

n = 6

A2= 5

(42.0^5.8)

(37.3-40.1)

(23.6-24.7)

(13.3-14.0)

(53.1-58.0)

cristata

33.30

29.10

11.44

10.29

42.96

n = l

n = 5

n = l

« = 5

(30.2-36.5)

(26.8-29.8)

(15.6-19.0)

(9.4-11.6)

(39.3-47.8)

reynaudii

35.98

31.33

17.78

9.80

47.60

« = 4

n = 4

« = 4

« = 3

(35.3-36.8)

(30.1-32.3)

(17.2-18.6)

(9.7-11.2)

(46.4-48.4)

ruficeps

40.15

32.28

21.05

10.97

57.68

« = 6

n = 6

« = 6

« = 5

(39.3-40.8)

(30.6-34.3)

(20.4-21.6)

(9.8-11.8)

(55.2-60.2)

serriana (« = 1)

45.9

37.6

22.6

13.3

61.0

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

- C berthae pelvic measurements from type specimen, Service de Paleontologie, Universite d' Antananarivo,
UM 6264; and tarsometatarsus from Museum National d'Histoire Naturelle, Service de Paleontologie, Paris,
MAD 5490.

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

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

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

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

Coua delalandei, a species that has gone
extinct in the past 1 50 years, has the longest
tarsometatarsus of any known recent Coua.
It is represented by less than 1 5 skin spec-
imens in museums, and no skeletal material
is available. The tarsometatarsus length of
this species, as measured from museum
skins, is 70 mm (Milne-Edwards & Gran-
didier 1895), considerably smaller than the
tarsometatarsus measurement oi primavea
or berthae.

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

CO
CO

on
+-•

+-•

E
o

CO
05

pelvis length

Fig. 3. Plot of tarsometatarsus length versus pelvis length in six species of modem Coua. The linear regression
equation is y = 2. 1 3 + 1 .29x (r^ = 0.80). Dotted lines are extrapolations of measurements based on the regression
analysis.

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

extremely large and swift-running species of
Coua.

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

VOLUME 106, NUMBER 1

berthae

C
Q)

• - gigas
■ - caerulea
A - reynaudii

o - cristata

100 200 300 400 500 600 700 800

weight (g)

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

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

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

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

Acknowledgments

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

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

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de Saint-Ours, J. 1953. Etude des grottes d'Andra-
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, & R. Paulian. 1953. Les grottes d'Andra-
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, L. R. Godfrey, & M. R. Sutherland. 1985.

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PROC. BIOL. SOC. WASH.

106(1), 1993, pp. 34^5

TWO NEW SPECIES OF BLIND SNAKE, GENUS

TYPHLOPS (REPTILIA: SQUAMATA: TYPHLOPIDAE),

FROM THE PHILIPPINE ARCHIPELAGO

Addison H. Wynn and Alan E. Leviton

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

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

Largely due to paucity of material, little
work has been done on Philippine typhlop-
ids. Until the mid- 1 950s fewer than 50 blind
snakes had been collected throughout the
whole of the Philippines. Most were ob-
tained between 1915 and 1921 by Edward
Harrison Taylor, who also described eight
of the 1 8 nominal species attributed to those
islands (Taylor 1917, 1918, 1919, 1922).
Moreover, most specimens collected before
Taylor were poorly documented as to prov-
enance, although all seem legitimately at-
tributable to the Philippines, except Typh-
lops dichromatus Jan (listed by Taylor [1 922]
in his synonymy of T. ruficaudus).

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

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

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

VOLUME 106, NUMBER 1

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

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

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

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

Typhlops castanotus, new species
Figs. 1, 2

Holotype.-CAS-SU 27940, an adult male
from 8 km west of Pulupandan, Inampu-
lugan Island, Negros Occidental Province,
Philippines, collected by Angel Alcala and
party, 23 May 1967.

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

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

Diagnosis.— A moderate-sized member
of the Typhlops ruficaudus group (McDow-
ell 1974) with 28 scale rows around the an-
terior body; dark dorsal stripe, nine or 1 1
scale rows wide, the lateral-most scale row
continuously pigmented for the entire body
length, sharply set off from the cream-col-
ored lateral and ventral scale rows; tail uni-
formly dark dorsally and laterally.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Description of holotype.— Total length
(TL), 224 mm; tail length, 7 mm; midbody
diameter (MBD), 7 mm; body diameter at
vent, 6 mm; TL/MBD, 32; 318 middorsal
scale rows; 307 midventral scale rows; 13
mid-subcaudal scales; 13 mid-dorsocaudal
scales; 28 scale rows around body anteriorly
reducing to 24 rows posteriorly; head
(viewed from above) tapered; in profile,
snout rounded, projecting anteriorly above
mouth, without transverse rostral keel; nos-
trils lateral, near tip of snout; eyes dorso-
lateral, visible beneath ocular shield im-
mediately behind posterior edge of
preocular, diameter of eye 1 6% of distance
from anterior edge to tip of snout on right,
1 7% on left; rostral oval, extends from up-
per lip ventrally to a level just anterior to
eyes on dorsum, about Vi width of head,
widest just posterior to level of nostrils, bor-
dered laterally by the nasals and by the pre-
frontal posteriorly; nasals incompletely di-
vided into pre- and postnasals, suture
dividing each nasal originating ventrally at
second upper labial, then extending dorsally
and anteriorly to nostril, beyond nostril
ending in a minute dimple-like depression
near edge of rostral; prenasal overlaps first
upper labial and anterior edge of second up-
per labial; postnasals separated from con-
tact behind rostral by prefrontal, posterior
edge overlaps supraocular, in broad contact
with preocular, ventral edge overlaps sec-
ond upper labial and anterior edge of third
upper labial; preocular inserts dorsally be-
tween postnasal and supraocular, borders
ocular posteriorly, inserts between postna-
sal and third upper labial ventrally (exclud-
ed from contact with the second upper labial
by postnasal); ocular slightly larger than
preocular, inserts dorsally between supra-
ocular and parietal, posteriorly contacts two
postoculars, inserts ventrally between third
and fourth upper labials, the ventroanterior
edge overlapped by third upper labial, the
ventral edge overlapping fourth upper la-
bial; four upper labials, first and second
smallest (second slightly larger than the first),

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

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

The number of scale rows around the body
decreases in two pairs of reductions from
28 rows anteriorly to 24 rows posteriorly.
Immediately following the fourth enlarged
middorsal head scale there are 28 scale rows
around the body (at 6 mm body length,
however, the midventral row splits produc-
ing 29 scales before fusing again four rows
posteriorly), reducing to 26 rows by fusion
of first and second para-midventral scale
rows on left at 109 mm body length and on
right at 1 1 9 mm body length, followed by
the second pair of reductions to 24 scale
rows at 207 mm body length.

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

VOLUME 106, NUMBER 1

Fig. 1 . Holotype (CAS-SU 27940) of Typhlops castanotus: left lateral view of head and ventral view of
tongue. The bar represents 5 mm.

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

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

is 3 mm in length. It is a single, soft organ.
The medial base is essentially smooth and
expands into a pair of smooth sacs, the larg-
est on the anterior side, a smaller sac on the
posterior side. At about half the length of
the hemipene, the lateral side expands into
a flat papillose disk covering the apicolateral
surface. The distal tip of the disk is enfolded
to produce a deep groove that extends onto
the medial shaft. The sulcus spermaticus is
a deep groove with smooth lips arising on
the posterior base and extending along the
posterior surface to the proximal lip of the
disk, where it ends in an area contiguous
with a series of grooves on the surface of
the disk. The remainder of the hemipene is
flounced with smooth parallel ridges (Fig.
2). The left hemipene is similar, but the
apicolateral disk appears to be flaccid.

Variation.— ThQ sex ratio in our sample
is seven males, six females. Two juveniles
could not be sexed. Total length ranges from
109-253 mm, with no apparent difference
between the sexes (Table 1). In contrast, al-
though our samples are small, both mid-
dorsal scale number and tail length appear

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

to be sexually dimorphic. In females, the
number of middorsal scale rows ranges from
324-339 {X= 331), in males 300-327 (X =
314), with only one male (CAS-SU 27942)
in the range of females. Although the total
number of middorsal scale rows is less in
most males, males have more mid-dorso-
caudal scales (12-14 [X = 13] vs. 11-12 [X
= 1 1.5] for females) and a tail that is pro-
portionately longer (as percent total length:
0.026-0.036 [X= 0.031] in males, 0.020-
0.027 [X = 0.025] in females).

In all specimens the number of scale rows
around the body reduces in two pairs of
reductions from 28 rows posterior to the
head to 24 rows anterior to the vent, al-
though reduction patterns vary (Table 1).
When reducing from an even to odd number
of scale rows, reductions involve the two
para-midventral scale rows, or the midven-
tral and one of the adjacent rows to either
side; reductions from an odd to even num-
ber occur by fusion of the first and second
rows to either the right or left of the midline,
or fusion of the first scale row to the right
with the first row to the left of the midline.
Only one specimen (CAS 127973) has a re-
duction (from 25 to 24 scale rows) that in-
volves other scale rows, rows seven and eight
to the right of the middorsal. The first pair
of reductions is offset in 14 of 1 5 specimens
(CAS-SU 27939 being the one exception).
The first reduction occurs on the left side
of the body in 1 3 of these specimens and
on the right side in one specimen (CAS-SU

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

The second pair of reductions, from 26
to 24, occurs together (or nearly so) at 89-
97% TL in 12 specimens. In three speci-
mens the second pair of reductions is offset:
the reduction from 26 to 25 occurs at 66%
or 70% TL (followed by a region of splitting
and refusing up to 1 6% TL), and the reduc-
tion to 24 at 84% or 89% TL.

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

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

VOLUME 106, NUMBER 1

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

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

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

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

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

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

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

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

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

VOLUME 106, NUMBER 1

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

Typhlops collaris, new species
Fig. 3

Holotype. — UF 55 1 23, an adult male from
Anuling Mt. (150 m elev.), Caramoan Mu-
nicipality, Camarines Sur, Luzon Island,
Philippines, collected by Walter Auffenberg
on 29 Jul 1982 (see Auffenberg 1988, for
additional information on this and the fol-
lowing localities).

Paratypes{\0).-V¥ 52866, 23 Jul 1982,
UF 54186 and USNM 319549 (formerly
UF 54187), 1 1 Oct 1982, base camp (Barrio
Terogo, about 2 km north of Caramoan;
Auffenberg 1988); UF 54188, 8 Aug 1982,
Kasini Mt. (250 m elev.); UF 54189, 22 Jul
1982, Ilawod; UF 54192, 26 Jul 1982, UF
55644 and USNM 319550 (formerly UF
55645), 27 Jul 1982, UF 55646, 7 Jul 1982,
Anuling Mt. (150 m elev.); UF 55648, 29
Jul 1982, Anuling Mt. (200 m elev.). All
collected by Walter Auffenberg in Cara-
moan Municipality, Camarines Sur, Luzon
Island, Philippines, except UF 52866, col-
lected by Walter Auffenberg et al.

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

Description of holotype.— Total length
(TL), 226 mm; tail length, 4 mm; mid-body
diameter (MBD), 5.5 mm; body diameter
at vent, 4.5 mm; TL/MBD, 41; 427 mid-
dorsal scale rows; 4 1 2 midventral scale rows;
1 2 mid-subcaudal scales; 1 1 mid-dorsocau-
dal scales; 28 scale rows around the body
anteriorly reducing to 26 scale rows poste-
riorly; head (viewed from above) tapered;
in profile, snout rounded, projecting ante-

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

riorly above mouth, without transverse ros-
tral keel; nostrils lateral, near tip of snout;
eyes dorsolateral, visible beneath ocular
shield, posterior edge of preocular covering
anterior third of eye on left and anterior
quarter of eye on right, diameter of both
eyes 1 5% of distance from anterior edge to
tip of snout; rostral oval, extending from
upper lip ventrally to a level just anterior
to eyes on dorsum, about Vi head width,
widest just posterior to level of nostrils, bor-
dered laterally by the nasals and by the pre-
frontal posteriorly; nasals incompletely di-
vided into pre- and postnasals, suture
dividing each nasal originating ventrally at
second upper labial, then extending dorsally
and anteriorly to nostril, beyond nostril

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

All head scales except the rostral have
basal glands forming a glandular line which
is overlapped by the posteriorly projecting
free edge of the preceding scale; glands lying
along intemasal suture between the nostril
and second upper labial expand into a stri-
ated organ; glands lying under posterior edge
of postnasal and preocular fuse at base of
third upper labial (under imbricate poste-
rior edge of postnasal); basal glands of scales
on body occupy anterior V3 of the scale (ex-
cluding posterior free edge of scale).

Posterior to the head shields, there are 26
scale rows around the body, increasing to
27 rows at 7 mm body length, and 28 rows
at 1 1 mm body length; the number of scale
rows then reduces in one pair of reductions
from 28 to 26 rows as follows: at 13 mm
body length the first and second para-mid-
ventral scale rows on the right fuse (followed
by a variable region in which this scale row
divides and refuses up to 42 mm body
length), and at 1 4 mm body length the first
and second para-midventral scale rows on
the left fuse; there are 26 scale rows for the
remainder of the body length, although
about 3 mm anterior to the vent the two
para-midventral rows on the right fuse, then
after three transverse scale rows divide again.

Light pigmentation, consisting of a fine
network of chromatophore reticulations,
covers non-glandular portions of dorsal head
scales (the three middorsal scales posterior
to the rostral; supraoculars; parietals; and
dorsal portions of the rostral, postnasals,
preoculars, and oculars); posterior to these
head scales, the scale inserted between the
interparietal and parietal, and the scale be-
tween the parietal and ocular are pigmented
on both sides of the head; posterior to these
pigmented scales is an unpigmented band
four scale rows wide middorsally and three
to four rows wide laterally; there are 15
lightly pigmented longitudinal rows of dor-
sal scales posterior to the collar reducing to
1 1 pigmented rows before the vent due to
gradual loss of pigmented scales in the lat-
eral-most row (row 7) over the body length,
and reduction in pigmentation density in
row 6 just before the vent; posterior to the
vent nine dorsal longitudinal scale rows are
pigmented on the anterior two-thirds of tail,
but unpigmented scales in the lateral-most
row and a band of four unpigmented mid-
dorsal scales produce a mottled appearance;
pigmented scales end about four transverse
scale rows anterior to the terminal spine;
terminal spine pigmented.

Scales in the dorsal stripe have a basal

VOLUME 106, NUMBER 1

Table 2.— Scale row reduction patterns in Typhlops collaris. See Table 1 for discussion.

Sex

MDC

MSC

Tail/TL

# of scale rows

Museum #

28-27

27-26

26-25

25-24

UF 54186

244

457

447

0.016

0.47 (0.52) L

0.52 (0.55) R

0.96 R

0.96 L

UF 54188

255

457

444

0.020

0.91 (0.93) R

0.92 L

—

UF 54189

255

460

448

0.014

0.55 (0.75) L

0.88 R

—

UF 54192

233

441

429

0.017

0.93 R

0.93 L

—

UF 55644

227

458

448

0.015

0.59 (0.65) L

0.74 (0.78) R

0.97

USNM 319550

255

434

427

0.016

0.58 (0.89) L

0.93 R

0.97 R

—

UF 55648

232

434

426

0.017

0.57 (0.63) L

0.66 (0.74) R

—

UF 52866

203

422

408

0.020

—

0.04 R

0.91

USNM 319549

207

412

396

0.024

0.62 (0.85) R

0.64 (0.67) L

—

UF 55123

226

427

412

0.018

0.06 (0.19) R

0.06 L

—

UF 55646

210

461

449

0.019

0.54 (0.68) L

0.56 (0.72) R

0.97 R

—

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

Variation. —T\iQ sex ratio in our sample
is four males, seven females. Total length
ranges from 203-226 mm {X = 212 mm)
in males and 227-255 mm {X = 243 mm)
in females (Table 2). The number of mid-
dorsal scale rows varies from 412-461 {X
= 430) in males and 434-460(1^ = 449) in
females; males have 11-13 {X = 12) mid-
dorsocaudal scales and females 10-12 (X =
1 1), and tail length (as percent total length)
is 0.018-0.024_(X = 0.020) in males and
0.014-0.020 {X = 0.016) in females, sug-
gesting tail length is sexually dimorphic.

Ten of 1 1 specimens have 28 scale rows
posterior to the head (although UF 54192
has an irregular ventral scale pattern for 1 1
mm posterior to the head with up to 30 scale
rows, and UF 55648 has an irregular pattern
in which both ventral and lateral scale rows

fuse and split before becoming regular at 35
mm total length). One specimen (UF 52866)
has only 27 scale rows posterior to the head.
Reduction patterns are variable but al-
ways involve the two para-midventral scale
rows, or the midventral and one of the two
adjacent scale rows, when the reduction is
from an even to odd number of rows. The
first and second scale rows to either the left
or right of the midline fuse when the re-
duction is from an odd to even number of
rows. The number of scale rows around the
body reduces in two pairs of reductions from
28 scale rows behind the head to 24 rows
in front of the tail in only two specimens
(Table 2). Six specimens have only one pair
of reductions (from 28 to 26) and two spec-
imens have one pair (28 to 26) followed by
a single reduction (26 to 25). In UF 52866
the single anterior reduction (27 to 26) is
followed by a pair of reductions to 24. In
the holotype (UF 55 123) and UF 52866 the
first reduction occurs anteriorly (6% and 4%
TL, respectively); in all other specimens the
first pair of reductions occurs near midbody
or on the posterior half of the body (47-
93% TL). The first pair of reductions is off-
set in ten specimens, the first reduction oc-
curring to the left of the middorsal in six of
these and to the right in four. All reductions
from 26 to 25, or 26 to 24 scale rows occur

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

The tongue of UF 54 1 86 is 4 mm in length,
forked at 1 .5 mm, and lacks lateral papillae.
All specimens have an unpigmented collar
immediately behind the head, beginning
middorsally with the fourth, fifth, or sixth
middorsal scale row and the first or second
scale behind the parietal and ocular shields.
Middorsally, the collar can be from one to
four scale rows wide; laterally, it varies in
width within individuals from one to five
scale rows, generally widest at the lateral-
most extent of the dorsal stripe, and bridged
in two specimens by pigmented scales. Pos-
terior to the collar, the dorsal stripe is either
17 or 15 scale rows wide. Contact between
the postnasals posterior to the rostral varies.
In seven specimens there is no overlap, al-
though the postnasals touch or come close
to touching behind the posterior edge of the
rostral in three of these specimens; in four
the postnasals overlap, the left postnasal
overlapping the right in two individuals and
the right overlapping the left in two. In all
specimens examined, the third upper labial
contacts the postnasal.

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

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

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

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

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

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

Preliminary Artificial Key to the

Species of Blind Snakes of the

Philippine Islands

The following key to Philippine scoleco-
phidians should serve as a useful prelimi-
nary guide to the currently recognized spe-
cies of Ramphotyphlops and Typhlops
known from the archipelago. We hesitate to
comment at this time on the status of sev-
eral problematic populations since our in-
vestigations of the T. ruficaudus group are
incomplete. For this key, we follow Mc-
Dowell (1 974) in including T. luzonensis and
T. hypogius in T. ruber, and T. jagorii in T.
ruficaudus. We tentatively synonymize T.
canlaonensis with T. ruficaudus. Although
McDowell (1974) stated that Typhlops hed-
raeus might be conspecific with Typhlops
ater, we recognize T. hedraeus here as a val-
id species.

1. Rostral without a sharp horizontal
ridge 2

- Rostral with a thickened horizontal
ridge; tail at least twice as long as
broad 7

2. Tail short, about as long as broad.
Glands on head confined to base of
scales 3

- Tail about two to three times as long
as broad. Head profusely covered
with glands, glands not confined to
base of scales; scales in 18 rows
around body Typhlops hedraeus

3. Scales in 20 rows around body; in-
temasal suture arising from preocu-
lar Ramphotyphlops braminus

- Scales in 26-30 rows around ante-

VOLUME 106, NUMBER 1

nor third of body; intemasal suture
arising from second upper labial ... 4

4. Preocular contacts second upper la-
bial forming a horizontal suture;
scales in 26 or 28 rows behind head;
15-21 dorsal rows of darkly pig-
mented scales, occasionally only the
centers of the scales are heavily pig-
mented giving rise to a lineate pat-
tern T. ruber

- Preocular separated from second
upper labial by postnasal, not form-
ing horizontal suture; scales in 28 or
30 rows behind head; 9-17 pig-
mented dorsal scale rows 5

5. Light nuchal collar present; trans-
verse scale rows >390; 11-17 light-
ly pigmented dorsal scale rows . . .

Typhlops coUaris

- Light nuchal collar absent; trans-
verse scale rows <390; 9-15 darkly
pigmented dorsal scale rows 6

6. Scale rows behind head 30-32. 11-
15 darkly pigmented dorsal scale
rows; scales on tail without dark pig-
mentation except for narrow mid-
dorsal stripe Typhlops ruficaudus

- Scale rows behind head 28. Nine or
1 1 darkly pigmented (usually black)
dorsal scale rows; tail darkly pig-
mented above and on sides

Typhlops castanotus

7. Scales in 20-22 rows around body . .

Ramphotyphlops olivaceus

- Scales in 24—28 rows around body . .

Ramphotyphlops cumingii

Acknowledgments

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

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

Literature Cited

Auffenberg, W. 1988. Gray's monitor lizard. Uni-
versity of Rorida Press, Gainesville, 4 1 9 pp.

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

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

McDowell, S. B. 1974. A catalogue of the snakes of
New Guinea and the Solomons, with special
reference to those in the Bemice P. Bishop Mu-
seum, Part I. Scolecophidia.— Journal of Her-
petology 8(1): 1-5 7.

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

Taylor, E. H. 1917. Snakes and lizards known from
Negros, with descriptions of new species and
new subspecies.— The Philippine Journal of Sci-
ence 12D(6):353-381.

. 1918. Reptiles of Sulu Archipelago.- The

Philippine Journal of Science 13D(5):233-267.

. 1919. New or rare Philippine reptiles. — The

Philippine Journal of Science 14(1): 105-1 25.

. 1922. The snakes of the Philippine Islands.—

Manila, Department of Agriculture and Natural
Resources, Bureau of Science Publication 16:1-
312.

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

(AHW) Department of Vertebrate Zool-
ogy, Division of Amphibians and Reptiles,
National Museum of Natural History,
Washington, D.C. 20560, U.S.A.; (AEL)
Department of Herpetology, California
Academy of Sciences, San Francisco, Cali-
fornia 941 18, U.S.A.

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

A NEW SPECIES OF DIMORPHIC TREE FROG,

GENUS HYLA (AMPHIBIA: ANURA: HYLIDAE),

FROM THE VAUPES RIVER OF COLOMBIA

William F. Pybum

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

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

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

Hyla karenanneae, new species
Fig. 1

Holotype. — The University of Texas at
Arlington Collection of Vertebrates (UTA)
A- 3 7 70, an adult female collected by J. K.
Salser, Jr. and the author on 7 June 1973
near Timbo, Department of Vaupes, Co-
lombia (01°06'N, 70°0rw, elev. 170 m).

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

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

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

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

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

VOLUME 106, NUMBER 1

Fig. 1. Hyla karenanneae, holotype, UTA A-3770.

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

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

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

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

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

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

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

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table L— Measurements in mm of the type series of Hy la karenanneae.

Catalog
no.

Sex

SVL

Head
length

Head
width

Tibia
length

Eye
length

Tym-
panum
diam-
eter

Eye to
nostril

Nostril
to tym-
panum

Width
third
finger
disc

Width
fourth
toe disc

UTA A-3768

26.6

8.2

8.8

13.6

2.7

1.5

3.0

8.3

1.4

1.2

A-3769

28.9

8.6

9.3

14.3

3.1

1.5

3.2

8.6

1.4

1.3

A-3770

30.5

10.7

10.3

15.8

2.6

1.7

3.3

9.2

1.6

1.4

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

Iris in both sexes bright golden bronze.

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

Discussion

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

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

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

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

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

Acknowledgments

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

VOLUME 106, NUMBER 1

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

for permission to examine his unpublished
manuscript dealing with hylid systematics.
Officials of INDERENA gave permission
to collect specimens in Colombia.

Literature Cited

Bokermann, W. C. A. 1962. Cuatro nuevos hylidos
del Brasil.-Neotropica 8:181-192.

. 1964. Dos nuevas especies de //y/a de Ron-

donia, Brasil.-Neotropica 10:2-6.

. 1966. Notas sobre Hylidae do Espirito Santo

(Amphibia, Salientia).— Revista Brasileira de
Biologia 26:29-37.

, & Ivan Sazima. 1973. Anfibios da serra do

cipo, Minas Gerais, Brasil. 2: duas especies no-
vas de Hyla (Anura, Hylidae).— Revista Brasi-
leira de Biologia 33:457-472.

Cochran, D. M., & C. J. Goin. 1970. Frogs of Co-
lombia.— United States National Museum Bul-
letin 288:1-655.

Delahoussaye, A. J. 1966. The comparative sperm
morphology of the Louisiana Hylidae (Am-
phibia: Anura).— Proceedings of the Louisiana
Academy of Sciences 29:140-152.

Duellman, W. E. 1 972. The systematic status and life
history of Hyla rhodopepla Giinther.— Herpe-
tologica 28:369-375.

. 1982. A new species of small yellow Hyla

from Peru (Anura: Hylidae).— Amphibia Rep-
tilia 3:153-160.

, & M. L. Crump. 1 974. Speciation in frogs of

the Hyla parviceps group in the upper Amazon
Basin.— University of Kansas Museum of Nat-
ural History, Occasional Papers 23:1-40.

, & M. J. Fouquette, Jr. 1968. Middle Amer-
ican frogs of the Hyla microcephalia group.—
University of Kansas Museum of Natural His-
tory Publications 17:517-557.

, & L. Trueb. 1983. Frogs of the Hyla Co-
lumbiana group: taxonomy and phylogenetic re-
lationships. Pp. 33-51 in A. G. J. Rhodin & K.
Miyata, eds.. Advances in herpetology and evo-
lutionary biology. Museum of Comparative Zo-
ology Harvard University, Cambridge.

, & . 1989. Two new treefrogs of the

Hyla parviceps group from the Amazon basin
in southern Peru.— Herpetologica 45:1-10.

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

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

Fouquette, M. J., Jr., & A. J. Delahoussaye. 1977.
Sperm morphology in the Hyla rubra group
(Amphibia, Anura, Hylidae) and its bearing on
generic status.— Journal of Herpetology 1 1 :387-
396.

Goin, C. J. 1957. Description of two new frogs from
Colombia.— Journal of the Washington Acad-
emy of Science 47:60-63.

. 1966. A new frog of the genus Hyla from

British Guiana.— Quarterly Journal of the Ror-
ida Academy of Science 29:39-42.

Haddid, C. F. B., & J. P. Pombal- Junior. 1987. Hyla
hiemalis, nova especie do grupo rizibilis do es-
tado de Sao Paulo (Amphibia, Anura, Hyli-
dae).— Revista Brasileira de Biologia 47:127-
132.

Heyer, W. R. 1977. Taxonomic notes on frogs from
the Madeira and Purus Rivers, Brasil.— Papeis
Avulsos de Zoologia 31:141-162.

. 1980. The calls and taxonomic positions of

Hyla qiesleri and Ololygon opalina (Amphibia:
Anura: Hylidae).— Proceedings of the Biological
Society of Washington 93:655-661.

Lutz, B. 1 973. Brazilian species of Hyla. —University
of Texas Press, Austin, 260 pp.

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

Rivero, J. A. 1961. Salientia of Venezuela. —Bulletin
of the Museum of Comparative Zoology 1 26: 1-
207.

. 1968. A new species of //y/a (Amphibia, Sali-
entia) from Venezuelan Guayana.— Breviora
307:1-5.

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

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

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

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

DESCRIPTION OF THE ADVERTISEMENT CALL

AND RESOLUTION OF THE SYSTEMATIC

STATUS OF LEPTODACTYLUS GRACILIS DELATTINI

MULLER, 1968 (AMPHIBIA: LEPTODACTYLIDAE)

Linnette Garcia Perez and W. Ronald Heyer

Abstract.— The advertisement call oi Leptodactylus gracilis delattini Muller
is described from Campeche, Santa Catarina, Brazil and compared with calls
from five populations of L. gracilis from mainland Brazil and calls of the closely
related L. furnarius and L. plaumanni. Based on comparison of call data and
re-examination of the holotype, Leptodactylus gracilis delattini Miiller, 1968
is considered to be a strict junior synonym of Leptodactylus gracilis (Dumeril
andBibron, 1841).

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

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

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

Materials and Methods

Recordings analyzed for this paper are:

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

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

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

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

— 20^

Fig. 1. Map of Brazilian States of Sao Paulo (SP), Parana (PR), Santa Catarina (SC), and Rio Grande do Sul
(RS), showing recording sites analyzed. 1 = Leptodactylus gracilis delattini, 2-6 L. gracilis, triangle = L.furnarius,
dot = L. plaumanni (see methods and materials section for further locality information).

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

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

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

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

ASN Tape AJC 73, cut 5, Leptodactylus

VOLUME 106, NUMBER 1

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

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

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

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

Results

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

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

Call of Leptodactylus gracilis delattini

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

Comparison With Other Calls

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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rsi

« -" -2 <N .^2 CO .^ -^ .22 in

"ob

05 so T- -5

. O

1-i

WD O
O

>^jk4jk4jk^j^j^

"Is

. o

VOLUME 106, NUMBER 1

^'^^^'^lft^^J^I

''ttelil(liiiiil!ltii

I-

(

' 1 1 1 1 1

I 2

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

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

Conclusion

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

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

Acknowledgments

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

Literature Cited

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

Miiller, P. 1968. Beitrag zur Herpetofauna der Insel

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Campeche (27°42'S/48°28'W).-Salamandra
4:47-55.
Straughan, I. R. 1973. Evolution of anuran mating
calls: bioacoustical aspects. Pp. 321-327 in J. L.
Vial, ed., Evolutionary biology of the anurans:
contemporary research on major problems.
University of Missouri Press, Columbia.

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

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

PROC. BIOL. SOC. WASH.

106(1), 1993, pp. 57-62

PROCHILODUS BRITSKII, A NEW SPECIES OF

PROCHILODONTID FISH

(OSTARIOPHYSI: CHARACIFORMES), FROM THE

RIO APIACA, RIO TAPAJOS SYSTEM,

MATO GROSSO, BRAZIL

Ricardo M. C. Castro

Abstract. —Prochilodus britskii is described from a single locality in the no
Apiaca, a tributary of the rio Arinos, of the rio Tapajos system, Brazil. The
species is distinguished from all other Prochilodus species by its slender caudal
peduncle (its depth 8.8 to 9.4% of SL versus 9.7% or higher in the remaining
species). Prochilodus britskii is a member of the assemblage of Prochilodus
species lacking dark marks on the caudal-fin. The 6 or 7 teeth in the inner
tooth row of each side of the lower jaw of P. britskii distinguish it from all
other species with plain caudal fins which have 8 to 1 8 teeth with the exception
of P. vimboides which has 6 to 1 3 teeth in that series. Prochilodus britskii differs
from P. vimboides in having 41 to 44 pored scales in the lateral line instead of
34 to 39.

Resumo.— Prochilodus britskii, uma nova especie da familia Prochilodon-
tidae e descrita de uma unica localidade no rio Apiaca, afluente do rio Arinos,
tributario do rio Juruena, pertencente a bacia do rio Tapajos, no Estado de
Mato Grosso, Brasil. A especie difere de todas as outras especies do genero
Prochilodus pelo fato da menor altura do seu pedunculo caudal variar de 8,8
a 9,4% do comprimento padrao contra valores iguais ou superiores a 9,7% nas
outras especies. Prochilodus britskii e parte do grupo de especies do genero sem
manchas negras na nadadeira caudal. Dentro de tal grupo pode ser distinguido
das outras especies por possuir 6 a 7 dentes na fileira interna da metade da
maxila inferior contra 8 a 18 nas especies restantes, com excegao de P. vim-
boides, que possui 6 a 1 3 dentes na mesma fileira de dentes. Prochilodus britskii
difere de P. vimboides por possuir 4 1 a 44 escamas perfuradas na linha lateral
contra 34 a 39.

The prochilodontid genus Prochilodus Posada Arango and Semaprochilodus Fow-
Agassiz is composed of medium to large ler, the two other recognized genera of the
sized (up to ~45 cm SL) fish species, widely family, is very confused, with the only pub-
distributed through South American wa- lished revisionary study dealing with the
ters. They are, wherever they occur, among Prochilodontidae (Mago-Leccia 1972) re-
the most important species in inland com- stricted to the Venezuelan species of the
mercial and subsistence fisheries (see Mago- family.

Leccia 1972; Roberts 1973; Vari 1983; This paper is part of ongoing phylogenetic

Lowe-McConnell 1975, 1987). Despite their and revisionary studies of the Prochilodon-

economic importance and widespread dis- tidae aiming to, among other things, solve

tribution, the state of the Prochilodus sys- the numerous taxonomic problems in the

tematics, as well as that of Ichthyoelephas family (Castro 1990). This work is based on

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

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

Prochilodus britskii, new species
Fig. 1, Table 1

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

^^^^^^^5.-6, MZUSP 38856-61, 195.5-
238.5 mm SL, same collection data as ho-
lotype.

Diagnosis. —Distinguished from all other
species of the genus Prochilodus by having
a more slender caudal peduncle, its depth
equal to 8.8 to 9.4% of standard length (SL)
instead of values equal to or higher than
9.7% in all other species. Prochilodus brit-
skii is a member of the assemblage of Pro-
chilodus species lacking dark marks on the
caudal-fin (Castro 1990). The 6 or 7 teeth
in the inner tooth row of each side of the
lower jaw of P. britskii distinguishes it from
all other species with plain caudal fins which
have 8 to 18 teeth in that series, with the
exception of P. vimboides which has 6 to 13
teeth in that series. Prochilodus britskii dif-
fers from P. vimboides in having 41 to 44
pored scales in the lateral line instead of 34
to 39.

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

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

VOLUME 106, NUMBER 1

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

Character

Holotype

Standard length

1 . Greatest body depth

2. Snout to dorsal-fin origin

3. Snout to p)elvic-fin origin

4. Snout to anus

5. Snout to anal-fin origin

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

7. Posterior termination of dorsal-fin
base to end of caudal peduncle

8. Dorsal-fin base length

9. Dorsal-fin length

10. Anal-fin base length

1 1 . Pectoral-fin length

12. Pelvic-fin length

13. Caudal peduncle length

14. Caudal peduncle depth

15. Bony head length

16. Snout length

17. Horizontal eye diameter

18. Postorbital length

19. Least interorbital width

20. Gape width

Lateral line scales

Scale rows between dorsal-fin origin

and lateral-line
Scale rows between anal-fin origin

and lateral-line
Scale rows between pelvic-fin origin

and lateral-line
Scale rows around caudal peduncle
Median predorsal scales
Median scales between posterior

termination dorsal-fin base and

adipose-fin origin
Vertebrae
Teeth in interior "V"-shaped tooth row

of upper jaw, left side
Teeth in interior "V"-shaped tooth row

of lower jaw, left side

)metrics
221.5

195.5-238.5

29.1

28.9-30.3

47.2

45.6-47.4

54.4

53.9-55.1

75.5

76.5-79.0

77.9

78.2-81.3

29.4

27.5-30.2

42.5

41.7-43.7

14.6

14.3-16.2

24.2

23.3-25.3

10.5

8.4-10.4

20.5

18.3-21.3

16.7

15.1-16.5

13.9

12.5-13.8

9.3

8.8-9.4

26.1

25.7-26.2

38.4

36.4-42.6

18.9

17.4-20.6

44.1

41.2-45.7

47.9

46.2-49.2

39.8

38.7-39.9

Meristics

41-43

6-7

5-6

195.5-238.5
28.9-30.3
45.6-47.4
53.9-55.1
75.5-79.0
77.9-81.3

27.5-30.2

41.7-43.7
14.3-16.2
23.3-25.3

8.4-10.5
18.3-21.3
15.1-16.7
12.5-13.9

8.8-9.4
25.7-26.2
36.4-42.6
17.4-20.6
41.2^5.7
46.2^9.2
38.7-39.9

41^4
6-7
5-6

6-7

13-14

13-15

40-41

10-13

6-7

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

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1 . Prochilodus britskii, new species, holotype, MZUSP 4 1 5 1 9, 22 L 5 mm SL, Brazil, Mato Grosso State,
no Apiaca, N of city of Juara, upriver from a fall (~10°36'S, 58°04'W), 15-24 Feb 1988.

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

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

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

Dorsal fin preceded by small anteriorly
bifurcated spine (see Gery 1977:367) con-
sidered herein as an unbranched ray in fin-
rays counts. Dorsal-fin rays iii,9 or iii,10
(iii,9 rare)[iii,10]; anal-fin rays iii,8 [iii,8];
pectoral-fin rays i, 1 3 or i, 1 4 (i, 1 4 most com-
mon) [i, 1 3]; pelvic-fin rays i,8 [i,8]; principal
caudal-fin rays 10/9 [10/9].

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

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

VOLUME 106, NUMBER 1

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

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

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

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

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

Acknowledgments

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

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

Literature Cited

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

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

Gery, J. 1977. Characoids of the world. Neptune City,
New Jersey, TFH Publications, 672 pp.

Lowe-McConnell, R. H. 1975. Fish communities in
the tropical fresh waters. New York, Longman,
337 pp.

. 1987. Ecological studies in tropical fish com-
munities. Cambridge, Cambridge University
Press, 382 pp.

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

Roberts, T. 1973. Osteology and relationships of the
Prochilodontidae, a South American family of
characoid fishes. — Bulletin of the Museum of
Comparative Zoology 145(4):21 3-235.

62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Vari, R. P. 1983. Phylogenetic relationships of the DepartamentO de Biologia, FFCLRP-

families Curimatidae, Prochilodontidae, An- Universidade de Sao Paulo, Avenida dos

ostomidae and Chilodontidae (Pisces: Chara- t> j • * ^c\r\r\ /^t-t-» i Ar\Ac\ t» -i- • ~

, „ ,. r- . V . . -7 Bandeirantes 3900, CEP 14049, Ribeirao

ciformes).— Smithsonian Contnbutions to Zo-

oiogy 378:1-60. Preto, SP, Brazil.

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

A NEW DEVONIAN OPHIUROID

(ECHINODERMATA: OEGOPHIURIDA) FROM

NEW YORK STATE AND ITS BEARING ON THE

ORIGIN OF OPHIUROID UPPER ARM PLATES

Frederick H. C. Hotchkiss

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

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

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

Systematic Paleontology

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

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

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

Class Ophiuroidea Gray, 1840

Order Oegophiurida Matsumoto, 1915

Suborder Lysophiurina Gregory, 1897

Family Protasteridae S. A. Miller, 1889

Strat aster Kesling & Le Vasseur, 1971

Strat aster Kesling & Le Vasseur, 1971:305,
317. -Kesling, 1972:10 in part {S.
ohioensis but not S. devonicus).—H2im-
mann & Schmincke, 1986:61.

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

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

"new genus". —Mclver & Mclver, 1955:159.

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

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

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

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

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

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

VOLUME 106, NUMBER 1

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

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

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

Strataster ohioensis
Kesling & Le Vasseur, 1971

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

Eugasterella ohioensis. — Harper, 1985:369,
371.

Diagnosis. — Carinal spines begin near the
periphery of the disc; there are approxi-
mately 1 3 spines in the space of four am-
bulacrals. The carinal spines are not pre-
served in a rigid erect position; instead they
are recumbent, usually all leaning in the
same direction, and certainly not fused to
the upper arm plates. The splay of the up-
permost vertical spines nearly doubles the
overall arm width so that it nearly equals
the disc radius. The disc tends to have con-
vex interradii, lacks spines, and is covered
above and below by minute granules which
conceal underlying plates. [Partly based on
Kesling & Le Vasseur, pi. 7, fig. 2.]

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

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

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

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

A A B A ?
A A B A B
A A B A B

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

I II III * IV V

Paratype UMMP

58332a

A A B * A

Paratype L-25e

A ? B * A

Paratype L-25i

? A ? * ?

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

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

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

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

Strataster wrighti (Kesling, 1970),
new combination

Drepanaster wrighti Kesling, 1970:75, pis.
l-2.-Hotchkiss, 1 978:542. -Hotchkiss,
1980:93.

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

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

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

Holotype UMMP

57497

A A B A B

Remarks.— ThQ disc radius of about 2.7
mm is in the middle of the size range for S.
ohioensis. The count of about five carinal
spines in the space of four ambulacrals is
distinctly different from the count of 1 3 in
four in 5. ohioensis. It agrees with S. ohioen-
sis in the obviously loose attachment of its
carinal spines. In fact, the spines are not
found on the holotype, but their former
presence is shown by a line of vacant sockets
which Kesling (p. 75) described as "pustular
bases for attachment of spines," and which
he noted extend from the aboral surface of
the disc [Note: typographical error in his
text says "oral surface"] onto the aboral
proximal part of the arms. The spine sockets
confirm the presence of upper arm plates.
The count of five carinal spines in the space
of four ambulacrals is probably not reliably
different from the count in the new species
to be described next. The disc outline of the
holotype has generally concave interradii.
The oral interradial spines are 0.3 to 0.4
mm long. The shape of the aboral surface
of the proximal ambulacrals is hidden by
upper arm plates and granules (presumably
the shape is trapezoidal, as in S. ohioensis).

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

Strataster maciverorum, new species
Figs. 1-5

"new genus and species," Mclver & Mclver,
1955:159. Strataster, n. sp. Hotchkiss,
1976:12.

Prot aster logani. — Mclver & Mclver, 1955:
159. [Not Protaster logani (Hall, 1868)]

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

VOLUME 106, NUMBER 1

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Strataster maciverorum, new species. 2.1, NYSM 13225, Paratype. Aboral view. Carinal spines
number approximately 1 3 in the space of eight laterals. Uppermost vertical spine short, splayed at a high angle
to the arm axis. Latex pull. Bar represents 2 mm. For less enlargement see Fig. 3.1. For counterpart see Fig.
3.2. 2.2, NYSM 13223, Paratype. Aboral view. Carinal spines diminish in height distally. Latex pull. Bar
represents 4 mm.

VOLUME 106, NUMBER 1

Fig. 3. Strataster macivewrum, new species. NYSM 13225, Paratype. Counterpart halves. Latex pulls. 3.1,
Aboral view. Alternating ambulacrals visible through integument near arm tip at left. Madreporic interradius
at lower right. Bar represents 4 mm. For greater enlargement see Fig. 2. 1 . 3.2, Oral view. Madreporic interradius
at lower right. Bar represents 2 mm.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

VOLUME 106, NUMBER 1

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

NYSM

R:r

w.r

Remarks

13222

5.5 mm

34 mm

2.5 mm

6:1

1:2.2

holotype; DS17

13223

26 +

4.5

—

1:1.8

paratype; DS9

13224

7.5

29+

—

1:1.9

paratype; DS37

13225

2.5

5:1

1:2.0

paratype; DSll, DS24

13226

4.5

15 +

—

1:2.3

paratype; DSll, DS24

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

Types. —There are six type-bearing pieces
of rock, numbered DS9, DSll, DS17,
DS17a, DS24, and DS37, collected by
Mclver and Mclver and given to the New
York State Museum (NYSM). DSll and
DS24 are counterpart halves of a single piece
of rock, as are DS17 and DS17a. The single
brittlestar (Fig. 1.1, 1.2) contained in rock
specimen DS17 and counterpart DS17a is
now catalogued as NYSM 13222; it was il-
lustrated by the Mclvers and is designated
the holotype of Strataster maciverorum.
There are four paratypes: one each in DS9
(Fig. 2.2) and DS37 (Fig. 5.1, 5.2), now cat-
alogued as NYSM 13223 and NYSM 13224,
respectively, and two in DS 1 1 and coun-
terpart DS24. Of the latter two, one is oral
side up and one is oral side down: NYSM
13225 refers to the individual (Figs. 2.1,3.1,
3.2) preserved as an aboral impression in
rock specimen DS 1 1 , and NYSM 1 3226 re-
fers to the individual (Fig. 4.1, 4.2) pre-
served as an aboral impression in rock spec-
imen DS24.

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

University, who pointed out that the spec-
imens may belong in a new genus and spe-
cies. Labels show that the ''Prot aster lo-
ganV of the Mclvers' article are contained
in rock specimens DSl , DS2 (figured by the
Mclvers), DS3, DSl 2, DSl 8, DS25, DS30,
DS70, and DS75. These rocks contain im-
pressions of the oral surface of a protasterid
brittlestar that resembles Eugasterella lo-
gani (Hall), the single type specimen of which
is itself an impression of the oral surface
without counterpart. These specimens are
most likely oral impressions of S. maciv-
erorum, an inference based on the obser-
vations (1) that the specimens which Prof.
Wells distinguished as a new generic type
are in every case impressions of the aboral
surface, and (2) that counterpart impres-
sions of these last are indistinguishable from
the oral impressions that do not have coun-
terparts. Hence these specimens are iden-
tified here as topotypes of S. maciverorum.
Remains of Devonaster are present in rock
specimens DS3, DS4, DSl 9 and DS24; re-
mains of Encrinaster sp. are present in DS23.

Etymology.— The species is named in
memory of Dr. Monroe A. Mclver and Eliz-
abeth P. Mclver of Cooperstown, New York,
who donated these prize fossils to science.
[According to ICZN Code Recommenda-
tion 21a (1985), the prefixes "Mac," "Mc"
or "M" should be spelled "mac," hence
''maciverorum^'; I thank Dr. David L.
Pawson for pointing this out to me.]

Measurements. — See Table 1 .

Loven's Law.— Two impressions of the

VOLUME 106, NUMBER 1

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

I II III * IV V

Topotype DS12

A A ? * A B

Topotype DS70

? ? B * ? ?

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

Orientation and behavior.— Tht carinal
spines and plates of S. maciverorum are very
asteroid-like, and suggest that the orienta-
tion in life was with the oral side toward the
substrate. Paratypes NYSM 13225 and
13226 that are preserved on the same piece
of rock (DS-1 1) but in opposite orientation
to each other probably indicate some sort
of turbulence at the time of burial.

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

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

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

Remarks. —All five specimens oiS. mac-
iverorum are larger than the largest of the
1 00 S. ohioensis that were measured by Kes-
ling & Le Vasseur (1971). The largest S.
maciverorum (NYSM 13223, Fig. 2.2) is
twice the size of the largest S. ohioensis. As
in S. wrighti, the shape of the aboral surface
of the proximal ambulacrals of S. maciv-
erorum is hidden by the upper arm plates
and granules.

The considerable differences between 5.
maciverorum and S. ohioensis are partly
bridged by S. wrighti. S. ohioensis has loose-
ly articulated crowded carinal spines and no
disc spines. S. maciverorum has rigid un-
crowded carinal spines and many such spines
on the disc. S. wrighti bridges the gap by
having loosely articulated uncrowded spines
and a few such spines on the disc (inferred
from spine sockets on the disc and arms).

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

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

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

Discussion of Ophiuroid Upper arm Plates

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

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

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

Ophiohelus (moved from the Ophiacanthi-
dae to the Ophiomyxidae by Spencer &
Wright 1966:U89), (9p/zzVzmZ?/x (Ophiacan-
thidae), and Astrophyton (Gorgonocephali-
dae) could be "due to degradation, or the
genera have not progressed through embry-
onic stages in which dorsals appear, if, as is
probably the case, dorsals have never ap-
peared." Mortensen (1913) showed that the
upper arm plates in the alleged primitive
ophiuroid Ophioteresis elegans Bell [shown
by Mortensen to be Ophiothela tigris Ly-
man] are being formed at the arm tip but
are being resorbed while still of microscopic
size. These possibilities offer a plausible ex-
planation for the lack of upper arm plates
in most of the previously described Oego-
phiurida and Stenurida.

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

Relation to the axial arm skeleton: Upper
arm plates of extant ophiuroids are perfectly
correlated with the arm segmentation of the
free portion of the arm, there being one up-
per arm plate to each segment. In Strataster
the halves of ambulacral vertebrae are not
in pairs simulating vertebrae, and so there
is no arm segmentation comparable to that
of extant ophiuroids. Even so, the upper
arm plates of the species oi Strataster might
be expected to show serial correlation with
the ambulacrals, but such is not the case. In
S. ohioensis there are approximately 1 3 up-
per arm plates in the space of four ambu-
lacrals, and in S. maciverorum approxi-
mately 13 upper arm plates occur in the
space of eight ambulacrals. Thus there is no
one-to-one serial correlation between upper
arm plates and ambulacrals in these lyso-
phiurine oegophiurids. Nor is there one-to-
one correlation between asteroid carinals
and the ambulacrals of the asteroid arm.

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

VOLUME 106, NUMBER 1

viewer, "they are not articulated and there-
fore not homologous with protasterid carinal
spines"; for examples see Clark 1911:figs.
38, 43-49, 100, 101, 106, 126, 131, 134.]
In contrast, carinal spines are a highly
conspicuous feature in S. ohioensis and S.
maciverorum. The carinal spines of 5. mac-
iverorum, particularly, are very "asteroid-
like." These spines almost certainly indicate
that these animals lived with their oral side
down as in modem sea stars.

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

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

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

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

Homologies of radial shields and upper
arm plates: Fewkes (1 887: 1 30) believed that
ophiuroid radial shields are homologous
with the first paired brachials of crinoids.
This view was discredited by Carpenter
(1887:308-309) based on study of crinoids.

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

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

Asteroids

Ophiuroids

Crinoids

Primary circlet:
centrale

primary interradials [cf. Zoro-
aster]

primary radials

carinal plates [= serial homo-
logues of primary radials]

centrale

primary interradials

primary radials

upper arm plates [= serial
homologues of primary ra-
dials]

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

? topmost stem penta-
meres [cf. Aethocrinus]

? infrabasals

—(absent)

Extraxial skeleton:

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

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

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

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

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

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

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

VOLUME 106, NUMBER 1

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

Asteroids

Ophiuroids

Crinoids

Axial skeleton:

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

terminal (ocular)
plates

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

terminal (ocular) plates

brachial plates [radials = 1st
plates of brachial series]
[pinnules = heterotomous
arm branching]

—(absent)

Adaxial skeleton:

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

adaxial virgalia
[evolved from stro-
matocystitid cover
plate series] [=aster-
opseid growth gradi-
ent metapinnules]

lateral arm plates [mouth angle
plates of jaw]

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

—(absent)

Admarginal skeleton:

admarginal virgalia
[=poraniid growth
gradient metapin-
nules] [=develop-
mental homologues
of inferomarginals]

—(unknown)

—(absent)

Marginal skeleton:

inferomarginal ossicles
[homologous with
the stromatocystitid
marginal frame]

[cf. marginalia in Rhopalo-
coma]

—(absent)

posed homology between radial shields and
upper arm plates is not supported by either
embryology or the new fossil evidence.

Homologies of the calycinal plates of
ophiuroids and crinoids: Carpenter (1884),
Sladen (1884), Fell (1963), and Hotchkiss
(1974, 1980) believed that the primary ra-
dial plates of ophiuroids and asteroids are
homologous with the radial plates of cri-
noids. This turns out to be a major source
of misdirection in inferring correct echi-
noderm plate homologies. If in crinoids the
brachials are serially homologous with the
radial plate, then where are the serial ho-
mologues of the primary radial plate in the
Asterozoa? For Carpenter (1887:309, foot-

note) the answer lay with the ophiuroid up-
per arm plates. For Fell (p. 4 1 9) it lay with
"asterozoan" ambulacral plates. To relocate
"asterozoan" ambulacral plates from the
aboral surface (where the calycinal system
develops) to the oral surface (where am-
bulacrals are part of the jaw apparatus), Fell
postulated a "dislocation of the main radial
growth gradient at the junction of the radial
calycinal plate and the first brachial ossicle"
(p. 382). This is Fell's mechanism for the
evolution of the Asterozoa from a crinoid-
like ancestor. Although Fell (p. 414) states
that evidence of a dislocation is directly ob-
servable in the ontogeny of Recent Astero-
zoa, this is contradicted by the fact that pri-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

mary radials develop in tissues derived from
the right side of the bilateral larva whereas
the ambulacrals develop in tissues derived
from the left side (Sladen 1884:37, Murak-
ami 1937:137, figs. 3-5).

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

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

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

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

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

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

VOLUME 106, NUMBER 1

per surface of the arms and that of the disc.
Important to the present context, it shows
that it is proper to compare upper arm plates
with disc plates. Judging from S. maciver-
orum it is very likely that the plates that
bear the first carinal spines are in fact the
primary radial plates of the rosette (cf Lud-
wig 1905:pl. 32, fig. 190). Hamling's ophiu-
roid and Rhopalacoma pyrotechnica (Salter)
seem to answer unequivocally which disc
plates hold the long sought homology. In
both there is a direct serial relation and a
virtually identical morphology between the
upper arm plates and the primary radials.
Upper arm plates are evidently fashioned
after the plan prescribed by the genetic in-
structions for fashioning of primary radials,
and therefore in every sense are serial ho-
mologues of the latter (cf Hubbs 1 944:293).

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

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

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

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

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

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

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

Oegophiurida Rediagnosed

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

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

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

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

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

VOLUME 106, NUMBER 1

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

Acknowledgments

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

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

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26 Sherry Road, Harvard, Massachusetts

01451, U.S.A.

PROC. BIOL. SOC. WASH.

106(1), 1993, pp. 85-91

ERYTHROSQUILLOIDEA, A NEW SUPERFAMILY,

AND TETRASQUILLIDAE, A NEW FAMILY OF

STOMATOPOD CRUSTACEANS

Raymond B. Manning and David K. Camp

Abstract.— The superfamily Erythrosquilloidea is erected for the family
Erythrosquillidae, which contains one genus and species, Erythrosquilla me-
galops Manning & Bruce, 1984. This new superfamily differs from the Bath-
ysquilloidea, Gonodactyloidea, and Squilloidea by its member having broad
and ventrally beaded propodi of the third and fourth maxillipeds; it differs
from the Lysiosquilloidea by having a distinct dorsal median carina on the
telson. The superfamily may represent a relict lineage within the Stomatopoda.
The family Tetrasquillidae is erected in the superfamily Lysiosquilloidea for
three genera, the monotypic Tetrasquilla Manning & Chace, 1990; Tectasquilla
Adkison & Hopkins, 1984; and Heterosquillopsis Moosa, 1991, which contains
three species. This new family can be distinguished from the three families
now remaining in the Lysiosquilloidea by its members having ovate rather
than styliform (as in the Lysiosquillidae) distal segments of the endopods of
the first two walking legs, by lacking an enlarged basal lobe on the dactylus of
the claw (Coronididae), and by lacking a strong proximal fold on the outer
margin of the uropodal endopod (Nannosquillidae). The only known pantrop-
ical stomatopod, Tetrasquilla mccullochae (Schmitt, 1 940), is included in this
family.

Manning & Bruce (1984:332) tentatively
placed their newly erected family Erythro-
squillidae in the superfamily Lysiosquilloi-
dea based on the presence of broad, ven-
trally beaded propodi of the third and fourth
maxillipeds of the only member of the fam-
ily, Erythrosquilla megalops Manning &
Bruce, 1 984 (Fig. 1). They pointed out, how-
ever, that the Erythrosquillidae differ from
other lysiosquilloids by having a sharp, dor-
sal median carina on the telson, by lacking
a ventrolateral projection on the sixth ab-
dominal somite overhanging the articula-
tion of the uropods, and by having a smaller
ventral papilla of the antennal protopod.
The first of those three characters is cur-
rently considered important in distinguish-
ing superfamilies of the stomatopods, and
we use it here to help differentiate the new
superfamily defined below. Our removal of

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

Camp & Kuck (1 990:852) pointed out that
a new family might have to be erected for
Meter osquilloides mccullochae (Schmitt,
1940), a species placed in the recently erect-
ed, monotypic genus Tetrasquilla by Man-
ning & Chace (1990) and assigned to the
Lysiosquillidae. Camp & Kuck (1990) not-
ed that characters of the species fit none of
the lysiosquilloid families then known
(Manning 1980), and that it also could not
be accommodated in the Erythrosquillidae
Manning & Bruce. A new family is diag-
nosed here for T. mccullochae and for the
related Tectasquilla lutzae Adkison & Hop-
kins, 1984 (Fig. 2). Keys to the superfami-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

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

(Lamarck, 1818), male, tl 227, Fort Pierce,
Florida, USNM 152469 (walking legs) and
female, tl 44, St. Lucie Inlet, Florida, USNM
256888 (uropod and claw); Nannosquilla
grayi (Chace, 1958), female holotype, tl 40,
Bass River, Massachusetts, USNM 100931
(claw) and female paratype, tl 4 1 , same lo-
cality, USNM 100932 (walking legs and
uropod); Tetrasquilla mccullochae, female,
tl 32, Alligator Reef, Florida, USNM
1 1 1028; Tectasquilla lutzae, male holotype,
tl 73, Gulf of Mexico, off northwestern Flor-
ida, USNM 204717.

VOLUME 106, NUMBER 1

Fig. 2. a, Tetrasquilla mccullochae (from Manning & Chace 1 990:fig. 46); b, Tectasquilla lutzae (from Adkison
& Hopkins 1984:fig. la).

Erythrosquilloidea, new superfamily movable apices. No more than 2 interme-

diate denticles present on telson.
Diagnosis. —Vropodi of third and fourth Type genus.— Erythrosquilla Manning &
maxillipeds broad, ventrally beaded. Telson Bruce, 1984, herein designated,
with distinct dorsal median carina. At most, Included families. — Erythrosquillidae
submedian marginal teeth of telson with Manning & Bruce, 1984.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

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

Superfamily Lysiosquilloidea

Giesbrecht, 1910
Tetrasquillidae, new family

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

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

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

1 970), H. philippinensis (Moosa, 1986), and
H. danielae Moosa, 1991, the type species.
Remarks. —This new family can be readi-
ly distinguished from the three other fam-
ilies now remaining in the Lysiosquilloidea.
The Tetrasquillidae differ from the Lysio-
squillidae in that the distal segment of the
endopods of the first two walking legs are
ovate (Fig. 3j, k, m) rather than slender and
styliform (Fig. 3d, e). The Tetrasquillidae
differ from the Nannosquillidae by lacking
a strong proximal fold on the outer margin
of the uropodal endopod (compare Fig. 4c
and 4d, e). The Tetrasquillidae can be dis-
tinguished from the Coronididae by the claw
(Fig. 5d, e) having the propodus pectinate
for all its length and by lacking the basal
inflation of the dactylus (Fig. 5 a).

Key to Superfamilies of

Recent Stomatopoda

(Modified from Manning 1980)

1 . Propodi of third and fourth maxil-
lipeds slender, not beaded or ribbed
ventrally 2

- Propodi of third and fourth maxil-
lipeds broad, usually beaded or
ribbed ventrally 4

2. All marginal teeth of telson with
movable apices Bathysquilloidea

- At most, submedian marginal teeth

of telson with movable apices .... 3

3 . Four or more intermediate denticles
present on telson Squilloidea

- Two or fewer intermediate denticles
present on telson . . . Gonodactyloidea

4. Telson lacking sharp dorsal median
carina Lysiosquilloidea

- Telson with sharp dorsal median ca-
rina Erythrosquilloidea

Key to Families of Lysiosquilloidea

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

- Dactylus of claw not inflated basal-
ly. Propodus of claw completely
lined with pectinations 2

VOLUME 106, NUMBER 1

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

2. Proximal portion ofouter margin of
uropodal endopod with strong fold

Nannosquillidae

- Proximal portion ofouter margin of
uropodal endopod lacking strong
fold 3

3. Distal segment of endopod of an-
terior two walking legs slender, sty-
liform Lysiosquillidae

- Distal segment of endopod of an-
terior two walking legs broadly ovate
Tetrasquillidae

Acknowledgments

The figures were prepared by Lilly King
Manning. Studies on stomatopod system-
atics have been supported by the Smithson-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

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

VOLUME 106, NUMBER 1

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

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

PROC. BIOL. SOC. WASH.

106(1), 1993, pp. 92-101

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

Ernesto Campos

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

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

The Pinnotheridae is a group of symbiotic termined that P. jamesi Rathbun, 1923, de-

and sexually dimorphic crabs that undergo scribed from a hard stage male (sensu Chris-

a series of morphological changes during tensen & McDermott 1958) is a junior

their postplanktonic development. This has synonym of P. reticulatus ^dUhbun, 1918,

resulted in some mistaken identifications a species which was described on the basis

representing developmental stages of the of a post-hard stage female. Postplanktonic

same species. For example, Williams (1965) stages of P. reticulatus collected in the upper

found that Pinnotheres depressus Rathbun, Gulf of California in 1986-1987 confirmed

1918 was the pre-hard stage male of P. os- Green's conclusions, and a comparison of

treum Say, 1817, and Campos (1989b) con- the pre-hard stage male with the holotype

eluded that P. pwZ7e'5ce«5 (Holmes, 1894) is of P. muliniarum Rathbun, 1918 revealed

a young female of Tumidotheres margarita no difference. In addition, a morphological

(Smith, 1869). Recently, Green (1985) de- analysis of these specimens has led to the

VOLUME 106, NUMBER 1

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

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

Systematics and Taxonomic Account
Juxtafabia, new genus

Diagnosis.— ¥Qvci2i\Q carapace subglobu-
lar, dorsal regions weakly defined. Third
maxilliped with ischium and merus indis-
tinguishably fused, ventral margin angular,
dorsal margin gently convex; palp slightly
shorter than ischium-merus, carpus and
propodus subequal in length, stout, latter
article suboblong with distal end obliquely
rounded, dactylus lunate, broad, ventrally
inserted near proximal end of propodus.
Abdomen with 7 free somites, laterally
reaching to coxae of walking legs, distally
covering buccal cavity.

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

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

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

i/05?5. — Mollusca, Rival via: species of
Chione, Polymesoda, Protothaca and Ta-
gelus.

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

Remarks.— Other genera in the Pinno-
theridae which also have the ischium and
merus of the third maxilliped indistinguish-
ably fused differ from Juxtafabia as follows:
Dissodactylus Smith, 1870, Clypeasterophi-
lus Campos & Griffith, 1990, Parapinnixa
Holmes, 1894 and Sakaina Serene, 1964
have an exopod without flagellum (Fig. 1 A);
Calyptraeotheres Campos, 1990, Ostraco-
theres H. Milne-Edwards, 1853 and Xan-
thasia White, 1846 have a palp with two
articles (Fig. IB); Limotheres Holthuis, 1975
and Orthotheres Sakai, 1969 have the dac-
tylus inserted distally on the propodus (Fig.
IC); Pinnixa White, 1846 and Sderoplax
Rathbun, 1893 have a lobe on the external
margin of the exopod (Fig. ID); Fabia Dana,
1851, Tumidotheres Campos, 1989b, and
Durckheimia De Man, 1889, have a small
palp with the dactylus inserted on the mid-
dle of the propodus (Fig. IE); and Pinnothe-
res Bosc, 1 802 s. str. has a linear and slender
dactylus inserted proximally on the inner
margin of the spatulate propodus (Fig. IF),
and seven free abdominal somites. Addi-
tional differences between Juxtafabia and
the above genera can be found in Biirger
(1895), Rathbun (1918), Tesch (1918), Se-
rene (1964, 1967), Holthuis (1975), Cam-
pos (1989a, 1989b, 1990) and Campos &
Griffith (1990).

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

VOLUME 106, NUMBER 1

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

Juxtafabia muliniarum (Rathbun, 1918),
new combination

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

Pinnotheres reticulatus Rathbun, 1918:93-
94, pi. 21, figs. 1, 2.-Glassell 1934:301;
1935:105. -Schmitt et al. 1973:5, 83.-
Green 1985: 611-614, figs. 1, 2.-
Campos-Gonzalez 1988:385.— Campos
1989b:672.

Pinnotheres jamesi Rathbun, 1 923:625-626,
pi. 29, figs. 1, 2, text-figs. 1, 2.— Glassell
1934:301. -Silas & Alagarswami 1967:
1200, 1223.-Schmittetal. 1973:50.

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

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

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

Material examined.— ^oXoXy^Q of Pin-
notheres muliniarum, Lower California,
male, cw 3.6 mm (fide Rathbun 1918),
USNM 23443.— Holotype of P. reticulatus,
off^ San Jose Island, Baja California Sur,
25°02'15"N, 110°43'30"W, 17 fms (31 m),
S.Sh., 17 Mar 1889, Station 3002 Albatross,
female, cw 9.2 mm (fide Rathbun 1918),
USNM 18217.-1 female, cw 2.9 mm, Gulf
of Santa Clara, Baja California, 25 Apr 1990,
in Chione fructifaga, UABC— 12 ovigerous
females, cw 5 to 10 mm, 4 young males, cw
1 to 2.2 mm, Campo El Pescador, on coast
4.5 km North of San Felipe, Baja California,
1988, in Protothaca grata, Chione califor-
niensis and C fructifaga, UABC. — 7 young
males, cw 1 to 2.3 mm, 1 1 adult males, cw

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

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

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

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

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

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

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

Chelipeds no more stout than walking legs;

VOLUME 106, NUMBER 1

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

mm.

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

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

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

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

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

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

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

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

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

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

VOLUME 106, NUMBER 1

Fig. 6. Juxtafabia muliniarum (Rathbun, 1 9 1 8); dorsal view of adult female. Carapace length
are dorsally visible when abdomen is folded towards carapace sternum).

7 mm. (Eyes

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

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

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

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

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

100

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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

Acknowledgments

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

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

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

ANOMOEOMUNIDA, A NEW GENUS PROPOSED FOR

PHYLLADIORHYNCHUS CARIBENSIS MAYO, 1972

(CRUSTACEA: DECAPODA: GALATHEIDAE)

Keiji Baba

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

In a previous paper, Baba (1 99 1 :484) sug-
gested that Phylladiorhynchus caribensis
Mayo, 1972 from the Caribbean Sea be re-
moved from that genus. It was suggested
that the species was close to Pleuroncodes
Stimpson, 1860, from the eastern Pacific
but that a new genus was needed. A more
thorough comparison now allows a new ge-
nus, Anomoeomunida, to be proposed to
include this species, and be discussed.

Anomoeomunida, new genus

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

Gender. —Feminine.

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

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

Remarks.— Mayo (1972:526) noted that
Phylladiorhynchus caribensis from 1 1-38 m
in the Caribbean Sea was intermediate be-
tween Galathea Fabricius, 1793, and Muni-
da Leach, 1820, and placed it in Phylla-
diorhynchus Baba, 1969. She stressed the
rostral shape in species of the genus and
amended the generic diagnosis to allow for
the lack of spination on the walking legs,
and for three rather than four or five ter-
minal spines (counting the double mesial
terminal spines as two) on the basal anten-
nular segment. Lewinsohn (1982:298) sug-
gested that this western Atlantic form should
eventually be placed in a new genus. Fol-
lowing the examination of the type material
deposited at the National Museum of Nat-
ural History, Smithsonian Institution,
Washington, D.C. (USNM 140187-8) of P.
caribensis, I am inclined to believe that
Lewinsohn was correct.

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

VOLUME 106, NUMBER 1

103

a o

Fig. 1 . Anterior part of carapace m Anomoeomunida caribensis [= Phylladiorhynchus caribensis Mayo, 1972]:
a, male holotype (USNM 140187), dorsal view, right supraocular spine partly broken; b, same, left lateral view.
Scales = 1 mm.

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

The unique morphology of the walking
legs also separates A. caribensis from species
of Phylladiorhynchus. The rostrum in spe-
cies of Phylladiorhynchus is broadly trian-
gular, dorsally excavated, and lacks the dor-
sal ridge seen in the Caribbean species. Mayo
(1 972:523) noted that the lateral limit of the
orbit in P. caribensis bears a small spine.
Examination of the type material discloses
that this spine can be discernible in dorsal
view (Fig. 1 a), but barely so in lateral view
(Fig. lb); in Phylladiorhynchus, the lateral
orbital angle is sharply produced (Baba 1991:
fig. 4). The distomesial margin of the an-
tennular basal segment in Phylladiorhyn-
chus bears double spines (see Miyake & Baba
1965:fig. 6A for P. ikedai\ Miyake & Baba
1 967: fig. 6b for P. pusillus; Tirmizi & Javed
1980:fig. 2D for P. bengalensis; Lewinsohn
1982:fig. le for P. integrirostris), while that
of P. caribensis bears a single thin small
spine (Mayo 1972:525, fig. le). The disto-
ventral margin of the first (basal) segment
of the antennal peduncle is strongly pro-
duced into an anterior prolongation in Phyl-
ladiorhynchus (see Miyake & Baba 1 965:fig.
4B for P. ikedai; Miyake & Baba 1967:fig.

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

Anomoeomunida is also related to the
eastern Pacific Pleuroncodes Stimpson,
1860, which contains two species: P. mon-
odon (H. Milne Edwards, 1837) and P. plan-
ipes Stimpson, 1860. Comparative material
from the Smithsonian Institution of these
two species was examined: P. planipes—4 S
(12.3-1 3.3 mm in carapace length excluding
rostral spine), 1 ovig. 9 (14.0 mm), USNM
81336, pelagic at 20-40 miles off" shore,
southern Lower California (Magdalena Bay),
Mar 1940, coll. & id. E. F. Ricketts; P.
monodon — 2 S (13.7, 16.3 mm), 2 9 (14.9,
17.3 mm), USNM 65710, "Albatross" Sta.
3396, off' Panama, 07°32'N, 78°36'W, 474

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

iilMli Ilk

a be

Fig. 2. Anterior part of carapace: a, Pleuroncodes planipes Stimpson, male (USNM 81336), dorsal view; b,
same, ventral view; c, Pleuroncodes monodon (H. Milne Edwards), female (USNM 65710), dorsal view. Scales
= 1 mm.

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

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

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

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

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

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

Acknowledgments

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

VOLUME 106, NUMBER 1

105

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

Literature Cited

Baba, K. 1969. Four new genera with their represen-
tatives and six new species of the Galatheidae
in the collection of the Zoological Laboratory,
Kyushu University, with redefinition of the ge-
nus Galathea.—OHyiU, Occasional Papers of
the Zoological Laboratory, Faculty of Agricul-
ture, Kyushu University 2(1): 1-32.

. 1991. Crustacea Decapoda: Alainius gen. nov.,

Leiogalathea Baba, 1969, and Phylladiorhyn-
chus Baba, 1969 (Galatheidae) from New Cal-
edonia. In Resultats des Campagnes MUSOR-
STOM, Volume 9, A. Crosnier ed.— Memoires
du Museum national d'Histoire naturelle, Zool-
ogie 152:479^91.

Fabricius, J. C. 1793. Entomologia systematica
emendata et aucta, secundum classes, ordines,
genera, species, adjectis synonimis, locis, ob-
servationibus, descriptionibus 2:viii + 519 pp.

Faxon, W. 1895. Reports on an exploration off the
west coasts of Mexico, Central and South Amer-
ica, and off the Galapagos Islands, etc. XV. The
stalk eyed Crustacea.— Memoirs of the Museum
of Comparative Zoology at Harvard College 18:
1-292, pis. A-K, 1-56.

Haig, J. 1955. 20. The Crustacea Anomura of Chile.
In Reports of the Lund University Chile Ex-
pedition 1948-49. — Kungliga Fysiografiska
Sallskapets Handlingar N.F. 66(1 2): 1-68.

Leach, W. E. 1820. Galateadees.— Dictionnaire des
Sciences Naturelles, Paris 18:48-56.

Lewinsohn, C. 1982. Phylladiorhynchus integrirostris
(Dana) und Lauriea gardineri (Laurie) (Decap-
oda, Anomura) aus dem nordlichen Roten
Meer.— Crustaceana 42:295-301.

Mayo, B. S. 1972. Three new species of the family
Galatheidae (Crustacea, Anomura) from the
western Atlantic — Bulletin of Marine Science
22:522-535.

Milne Edwards, H. 1837. Histoire naturelle des Crus-
taces, comprenant I'anatomie, la physiologic et
la classification de ces animaux. Libraire En-
cyclopedique de Roret, Paris, Volume 2, 532
pp.

Miyake, S., & K. Baba. 1965. Some galatheids ob-
tained from the Bonin Islands (Crustacea, An-
omura).— Journal of the Faculty of Agriculture,
Kyushu University 13:585-593.

,& . 1967. Galatheids of the East China

Sea (Chirostylidae and Galatheidae, Decapoda,
Crustacea).— Journal of the Faculty of Agricul-
ture, Kyushu University 14:225-246.

Stimpson, W. 1 860. Notes on North American Crus-
tacea, in the Museum of the Smithsonian Insti-
tution, No. II.— Annals of the Lyceum of Nat-
ural History of New York 7:177-246, pis. 2, 5.

Tirmizi, N. M., & W. Javed. 1980. Two new species
and one new record oi Phylladiorhynchus Baba
from the Indian Ocean (Decapoda, Galathei-
dae).— Crustaceana 39:255-262.

Von Martens, E. 1878. Einige Crustaceen und Mol-
lusken, welche das zoologische Museum in letzter
Zeit erhalten. — Sitzungsberichte der Gesell-
schaft naturforschender Freunde zu Berlin, 18
Juni 1878:131-135.

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

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

TWO NEW SPECIES OF NEOCALLICHIRUS

FROM THE CARIBBEAN SEA

(CRUSTACEA: DECAPODA: CALLIANASSIDAE)

Raymond B. Manning

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

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

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

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

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

In some of the species placed in Neocal-
lichirus by Manning & Felder (1991) the
propodus of Mxp3 is distally emarginate,
i.e., there is an indentation on the opposable
margin (see Fig. Id). This is shown by Ro-
drigues (1971:fig. 67) for A^. guara and ap-
pears to be characteristic of both species
described here. The indentation is present
in A^. trilobatus but is less well marked or
even absent in A^. grandimanus (see Biffar
1971:fig. 5f, and Manning 1987:fig. 2e), A^.
rathbunae (see Manning & Heard 1986:fig.
lb), or A^. mirim (see Rodrigues 1971:fig.
84). It is shown by Rodrigues (1971:fig. 48)
but not by Biffar (1971:fig. 9f) for A^. guas-
sutinga, although both figures suggest that
the opposable margin is irregular in each of
those species.

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

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

VOLUME 106, NUMBER 1

107

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

ters), Man (mandible), Max 1-2 (first and
second maxillae), mm (millimeters), Mxpl-
3 (first to third maxillipeds), PI -5 (first to
fifth pereopods), Pip 1-5 (first to fifth pleo-
pods), tl (total length, measured on midline,
in mm).

The measurement following the number
of specimens is carapace length; in some
cases total length also is given. Segments of
appendages are measured dorsally.

Family Callianassidae Dana, 1852

Subfamily Callichirinae

Manning & Felder, 1991

Gqwus Neocallichirus Sdikdii, 1988

Neocallichirus lemaitrei, new species

Figs. 1-3

Material— CoXombidi: Islas del Rosario
(10°10'N, 75°46'W), Isla del Rosario, beach
on south side, yabby pump, 17 Jul 1988,

108

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Neocallichirus lemaitrei, new species, female paratype, cl 18.5 mm, USNM 256875, Isla de Baru. a,
Man; b, Maxl; c, Max2; d, Mxpl; e, Mxp2; f, P2; g, P3; h, P4; i, P5; j, Plp3.

leg. R. Lemaitre: 1 (5 (1 1 .6) (holotype, USNM
256876).

Isla de Baru [10°10'N, 75°36'W], Aug
1986, yabby pump, leg. R. Lemaitre: 1 $
(11.7), 1 $ (18.5; tl 76 mm) (paratypes,
USNM 256875). -Isla de Barn, Cienaga de
Cholon, 0.5 m, yabby pump, 7 Aug 1988,
leg. R. Lemaitre: 2 (5 (8.9, 10.5), 5 non-ovig-
erous 9 (12.4, 15.7, 17.6, 18.1, 18.4), 2 ovi-
gerous 9 (12.3, 15.4) (paratypes, USNM
256877).

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

Description.— CsiTSLpsiCQ lengths of adults
to more than 1 8 mm, total lengths to at least
76 mm.

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

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

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

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

VOLUME 106, NUMBER 1

109

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

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

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

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

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

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

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Neocallichirus nickellae, new species, male holotype, USNM 256879, cl 1 5.2 mm, Tobago, a, Carapace
and frontal appendages, lateral view; b. Anterior part of carapace and frontal appendages, dorsal view; c, Mxp3,
inner surface; d, Major PI, outer surface; e. Minor PI, outer surface; f, Sixth abdominal somite, telson, and
uropods, dorsal view.

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

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

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

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

Neocallichirus nickellae, new species
Figs. 4-6

Material. —Republic of Trinidad and To-
bago: Coral Garden, Buccoo Reef (1 ri I'N,
60°49'W), Tobago, 28 Jul 1989, leg. Lois
Nickell: 2 d (14.1, 15.2) (smaller .5, tl 56 mm
is paratype, USNM 256878; larger 6, tl 58
mm, is holotype, USNM 256879).

Diagnosis.— C2ir2cp2iCQ with 3 unarmed
anterior projections, all low, obtuse. Mxp3
with length and height of propodus sub-

VOLUME 106, NUMBER 1

111

Fig. 5. Neocallichirus nickellae, new species, male holotype, USNM 256879, cl 1 5.2 mm, Tobago, appendages.
a, Man; b, Maxl; c, Max2; d, Mxpl; e, Mxp2; f, P2; g, P3; h, P4; i, P5.

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

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

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

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

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

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

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

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

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

112

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Fig. 6. Neocallichirus nickellae, new species, male holotype, USNM 256879, cl 15.2 mm, Tobago, a, Plpl;
b, Plp2; c, Plp3; d, Appendix interna of Plp3.

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

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

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

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

Remarks.— Neocallichirus nickellae re-
sembles A^. lemaitrei and differs fi*om A^.
grandimanus in having the opposable mar-
gin of the propodus of Mxp3 indented or

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

VOLUME 106, NUMBER 1

113

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

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

Key to Western Atlantic Species of
Neocallichirus

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

A^. rathbunae i^chmiXX, 1935); southern

Florida and Caribbean (Biffar

1971, Manning & Heard 1986)

- Frontal projections lacking anterior
spinule. Upper plate of uropodal ex-
opod shorter than lower plate. Me-
rus of major PI lacking erect spines

on ventral margin 2

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

A^. grandimanus (Gibbes, 1850);

Bermuda, southern Florida and

Caribbean to Brazil (Biffar 1971).

- Carpus of major PI more than half
as long as propodus. Opposable
margin of Mxp3 propodus with
notch or indentation 3

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

.... A^. nickellae, new species; Tobago

- Dactylus of major cheliped with
large, rectangular tooth basally on
cutting edge

. . A^. lemaitrei, new species; Caribbean

coast of Colombia

Acknowledgments

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

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

Literature Cited

Biffar, T. A. 1970. Three new species of callianassid
shrimp (Decapoda, Thalassinidea) from the
western Atlantic. —Proceedings of the Biological
Society of Washington 83(3):35-49.

. 1971. The genus Callianassa (Crustacea, De-
capoda, Thalassinidea) in south Florida, with
keys to the western Atlantic species.— Bulletin
of Marine Science 21(3):637-715.

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

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

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

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

Manning, R. B. 1 987. Notes on western Atlantic Cal-
lianassidae (Crustacea: Decapoda: Thalassinid-
ea).— Proceedings of the Biological Society of
Washington 100:386^01.

. 1988. T\\Q ^X2iXviS of Callianassa hart meyeri

Schmitt, 1935, with the description of Coral I i-
anassa xutha from the west coast of America
(Crustacea, Decapoda. Thalassinidae). — Pro-
ceedings of the Biological Society of Washington
101:883-889.

. 1992. A new genus for Corallianassa xutha

Manning (Crustacea: Decapoda: Callianassi-
dae).— Proceedings of the Biological Society of
Washington 105:571-574.

. & D. L. Felder. 1986. The status of the cal-
lianassid genus Callichirus Stimpson, 1866

114

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

, & . 1991. Revision of the American

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

, & . 1 992 [ 1 99 1 ]. Gilvossius, a new ge-
nus of callianassid shrimp from the eastern
United States (Crustacea: Decapoda: Thalassi-
nidea). — Bulletin of Marine Science 49(1-2): 558-
561.

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

Callianassa rathbunae Schmitt, 1935, from
Rorida and the Bahamas (Crustacea: Decapoda:
Callianassidae).— Proceedings of the Biological
Society of Washington 99:347-349.

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

ghost shrimp from the Americas (Crustacea: De-
capoda: Callianassidae).— Nauplius (Brazil) (in
press).

Rathbun, M. J. 1900. The decapod and stomatopod
Crustacea. Results of the Branner-Agassiz Ex-
pedition to Brazil, 1 . — Proceedings of the Wash-
ington Academy of Sciences 2:135-155, pi. 8.

Rodrigues, S. de A. 1 97 1 . Mud shrimps of the genus
Callianassa Leach from the Brazilian coast
(Crustacea, Decapoda).— Arquivos de Zoologia,
Sao Paulo 20(3): 19 1-223.

, & R. B. Manning. 1992a. Two new calli-
anassid shrimps from Brazil (Crustacea: Decap-
oda: Thalassinidea).— Proceedings of the Bio-
logical Society of Washington 105:324-330.

, & . i992b. Poti gaucho, a new genus

and species of ghost shrimp from southern Bra-
zil (Crustacea: Decapoda: Callianassidae).—
Bulletin of Marine Science 50:9-13.

Sakai, K. 1988. A new genus and five new species of
Callianassidae (Crustacea: Decapoda: Thalas-
sinidea) from northern Australia.— The Beagle,
Records of the Northern Territory Museum of
Arts and Sciences 5(1): 5 1-69.

Schmitt, W. L. 1935. Mud shrimps of the Atlantic
coast of North America.— Smithsonian Miscel-
laneous Collections 93(2): 1-21, pis. 1-4.

Werding, B. 1982. Porcellanid crabs of the Islas del
Rosario, Caribbean coast of Colombia, with a
description of Petwlisthes rosariensis new spe-
cies (Crustacea: Anomura).— Bulletin of Marine
Science 32:439-447.

Department of Invertebrate Zoology, Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington, D.C. 20560,

U.S.A.

PROC. BIOL. SOC. WASH.
106(1), 1993, pp. 115-121

CAPRELLA ARIMOTOI, A NEW SPECIES

(CRUSTACEA: AMPHIPODA: CAPRELLIDEA)

FROM THE SETO INLAND SEA, JAPAN

Ichiro Takeuchi

Abstract. — Caprella arimotoi is described based on the material collected
from the red alga Pterocladia capillacea (Gmelin) on the Mukaishima Island
in the Seto Inland Sea. The new species is close to C. verrucosa Boeck, 1872,
but differs in having few plumose setae on antenna II, forwardly vented pro-
jection on head and elongate gills. Caprella (Spinicephala) pseudoverrucosa
(nomen nudum) mentioned in Arimoto's essay of 1978 is synonymous with
the present species.

Caprella is the largest genus of the sub-
order Caprellidea (Crustacea: Amphipoda),
widely distributed from temperate to boreal
regions occurring primarily on seaweeds,
seagrasses, and hydroids. So far about 130
species of this genus have been reported (e.g.,
McCain 1968; Laubitz 1970, 1972; McCain
& Steinberg 1970; Vassilenko 1974; Ari-
moto 1976; Takeuchi 1989). During my
short visit to the Mukaishima Marine Bi-
ological Station of Hiroshima University in
June 1989, the author found numerous ma-
ture individuals of Caprella on the red alga
Pterocladia capillacea (Gmelin) in the sub-
tidal zone. A close examination of those ma-
terials revealed that some of them are iden-
tical with what has been called "young male"
of C {Spinicephala) verrucosa Boeck, 1872
in Arimoto (1976) and with C (5.) pseu-
doverrucosa in Arimoto (1978). In this pa-
per, the specimens are described as a new
species.

The type specimens have been deposited
in the National Science Museum in Tokyo
(NSMT), National Museum of Natural His-
tory in Washington, D.C. (USNM), and Ca-
nadian Museum of Nature in Ottawa
(NMCC). The definition of mature stages in
females was referred to Takeuchi & Hirano
(1991).

Caprella arimotoi, new species
Figs. 1-3

Caprella {Spinicephala) verrucosa, Arimo-
to, 1976, 122-129 (in part), fig. 67-D. (non
Caprella verrucosa Boeck, 1872)

Caprella {Spincephala [sic.]) pseudoverru-
cosa Arimoto, 1978, 14, fig. 7C. (nomen
nudum)

Material examined.— HoXoXypt (NSMT
11191), male from Pterocladia capillacea
(Gmelin) Bomet & Thuret found at the
highest level of subtidal zone, Mukaishima
Island, Seto Island Sea (34°22'N, 1 33°1 3'E),
June 6, 1989, coll. I. Takeuchi. Allotype
(NSMT 11192), female collected together
with holotype. Paratypes: NSMT 1 1 193 (14
males and 4 premature females), USNM
251762 (10 males and 1 mature and 2 pre-
mature females), and NMCC 1992-0603 (10
males and 1 mature and 3 premature fe-
males), all collected together with holotype.
Arimoto's private collection No. 877-8, 1
male from Sargassum sp., Tsushima Is-
lands, December 1946 (?).

Diagnosis.— Head with triangular for-
wardly pointing projection above eye. Pere-
onites II to VI each with 1 or 2 rounded
dorsal projections. Antenna II of large male
with 4-5 pairs of plumose setae on each of

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

peduncular segments II and III. Basis of
gnathopod II shorter than half of pereonite
II; propodus oval, palm with pointed pro-
jection near proximal end and shallow tri-
angular projection near distal end. Gills
elongate, small. Propodus of pereopods V
to VII each with a pair of proximal grasping
spines.

Description. —Holoiype (Male; Figs. lA,
2A-F', 3A-H). Body length 6.97 mm; length
of head 0.45 mm; length of pereonites I to
VII 0.34, 1.42, 1.37, 1.23, 1.05, 0.66, and
0.47 mm, respectively. Head anteriorly
round; dorsal projection curved and point-
ed forward from posterior end. Pereonite I
with small posterodorsal projection; pere-
onites II to IV each bearing a mid-dorsal
and a posterodorsal blunt rounded projec-
tion. Pereonite V with a minute anterolat-
eral and a mid-dorsal blunt rounded pro-
jection. Pereonite VI with a mid-dorsal
triangular projection.

Antenna I about Vi of body length. Pe-
duncular segments longer in the order of II,
I, and III; flagellum composed of 11 seg-
ments and somewhat longer than peduncle.

Antenna II about % length of antenna I.
Peduncular segments III and IV fringed with
5 pairs of plumose setae and with 5 pairs
and a plumose setae, respectively; flagellar
segment I with 4 pairs of short plumose
setae; flagellar segment II with a short plu-
mose setae followed by 3 setae on distal end.

Gnathopod I with propodus twice as long
as width; palm serriform, setose with a pair
of grasping spines near proximal end.

Gnathopod II with vestigial coxa insert-
ing V3 from anterior margin of pereonite II.
Basis about % of pereonite II. Propodus
oval, V3 of pereonite II. Palm with 2 pro-
jections; proximal one V^ from proximal end,
pointing distally, and carrying 4 setae on
basal part; distal one low triangular and
sparsely setose. Dactylus stout, distal % of
inner margin serrated.

Gills small, elongate, 3 times longer than
width.

Pereopods V to VII with vestigial coxae.

Basis of pereopod V with laterally expanded
projection carrying granulations; merus ex-
panded on outer side; palm of propodus
convex, coarsely setose, with a pair of grasp-
ing spines on a small knob near proximal
end. Features of pereopods VI and VII close
to that of pereopod V. Pereopods V to VII
longer progressively; pereopod VII especial-
ly large, twice as long as pereopod VI.

Abdomen. Distal segment of appendage
oval, shallowly divided into 3 apical teeth;
basal segment with 5 or 6 setae surrounding
distal segment. Lobes bearing several long
setae. Penes medial.

Mouthparts. Inner plate of maxilliped
round and distally expanded with 2 spini-
form setae on distal margin and a facial row
of several plumose setae; outer plate, sub-
equal to inner plate, round and bearing 2
curved spiniform setae and 3 long setae on
inner margin; segment II of palp with
scarcely setose inner margin; segment III
expanded distally, scarcely setose on lateral
face; segment IV, longer than III, with sharp
claw. Outer plate of maxilla I rectangular
and slightly curved, with 7 spiniform teeth;
distal segment of palp rectangular with 4
spiniform teeth on distal margin, 3 stout
setae on distal part of inner margin, and a
row of 5 long setae followed by a short seta
on lateral face. Maxilla II with oval inner
and rectangular outer plate; both with
densely setose margins. Incisor of left man-
dible divided into 6 teeth; lacinia mobilis
separated into 5 teeth followed by 3 setae;
molar with a long seta near outer edge. Right
mandible with 5 -toothed incisor, lacinia
mobilis carrying minute teeth on middle
margin followed by 2 setae; molar large, with
a long seta and bushy bundle of setae. Upper
lip finely setose. Inner lobe of lower lip
round, finely setose on distal part.

Female (allotype, figs. IB, 2G-I, 31). Body
length 5.91 mm. Head 0.43 mm. Pereonites
I to VII 0.22, 1.12, 0.97, 0.86, 0.87, 0.52,
0.49, and 0.44 mm, respectively.

Gnathopod II situated near anterior of
pereonite II. Basis slightly shorter than V2

VOLUME 106, NUMBER 1

117

Fig. 1. Caprella arimotoi, n. sp. from Mukaishima Island in the Seto Inland Sea. A, holotype (male), 6.97
mm; B, allotype (female), 5.91 mm.

of pereonite 11. Propodus oval, subequal to
basis; palm smooth, convex and setose, with
a grasping spine near proximal end.

Oostegite III setose on margin; oostegite
IV minutely setose on anterior margin and
moderately on posterior margin. Gills on
pereonite III oval and those on pereonite
IV smaller, elliptical. Abdomen with a pair
of lobes without setae.

Etymology.— The specific name, arimo-
toi, is in honor of the late Dr. Ishitaro Ari-
moto, who made contributions to the tax-
onomy of the Japanese caprellidean
amphipods and was the first person to find
the present species.

Localities.— Type locality: Mukaishima
Island, Seto Inland Sea (34°22'N, 1 33°1 3'E).
Others: Tsushima Islands, and Tateyama

(Arimoto 1976) and Amatsu-Kominato
(Takeuchi 1989; as Caprella sp. C), Chiba.

Remarks.— The present new species is
close to Caprella verrucosa Boeck, 1872 in
having blunt dorsal projections on pereon-
ites I to VI, short basis and oval-shaped
propodus in gnathopod II, and grasping
spines on pereopods V to VII.

Caprella verrucosa was first reported from
somewhere near San Francisco, California
(Boeck 1872), and later recorded from both
sides of the North Pacific; from the Queen
Charlotte Islands, British Columbia to Santa
Catalina Island, California (Dougherty &
Steinberg 1953, Laubitz 1970, McCain &
Steinberg 1970, Martin 1977, MarelH 1981),
in the east, and from the Tsugaru Straight
to the Kyusyu Islands along both Japanese

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

A: 0.20mm

B: 0.10mm
C-G: 0.20mm H, I: 0.20mm

Fig. 2. Caprella arimotoi, n. sp. from the Mukaishima Island in the Seto Inland Sea. Holotype (male), 6.97
.„m. A, antenna II; B, gnathopod I; C, gnathopod II; D, pereopod V; D', coxa of pereopod V; E, pereopod VI;
E', coxa of pereopod VI; F, pereopod VII; F', coxa of pereopod VII. Allotype (female), 5.91 mm. G, gnathopod
II; H, gill and oostegite on pereonite III; I, gill and oostegite on pereonite IV.

VOLUME 106, NUMBER 1

119

Fig. 3. Caprella arimotoi, n. sp. from the Mukaishima Island in the Seto Inland Sea. Holotype (male), 6.97
mm. A, maxilliped; B, maxilla I; C, maxilla II; D, right mandible; E, left mandible; F, upper lip; G, lower lip;
H, abdomen. Allotype (female), 5.91 mm. I, abdomen.

coasts (Utinomi 1943, 1947, 1964; Arimoto
1976; Takeuchi 1989) and the south coast
of the Korean Peninsula (Kim & Lee 1975,
Lee 1988) in the west.

Mature males of C arimotoi can be sep-
arated from those of C. verrucosa from the
Japanese coast (Utinomi 1943, 1947; Tak-

euchi 1989) and British Columbia (Laubitz
1970) with the following characters: 1) An-
tenna I equal to Vi of the body length {% in
C. verrucosa), 2) pedunclar segments of an-
tenna I slender, each about 4 times longer
than width (2 to 3 times in C verrucosa),
3) antenna II carrying 4-5 pairs of plumose

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

setae on peduncular segments II and III
(more than 8 pairs of longer plumose setae
in C. verrucosa), 4) mid-dorsal projection
on the head curved forward, (straight for-
ward in C verrucosa), 5) lacking ventrolat-
eral projections on pereonites III to IV (C.
verrucosa with distinct ventrolateral projec-
tions on pereonites III and IV), 6) in C
verrucosa, additional paired projections
on pereonites IV to V, and 7) gills 3 times
longer than width (1.5 times longer in C
verrucosa).

The author examined the specimens of C
verrucosa collected from Vancouver Island,
British Columbia, and deposited at the Ca-
nadian Museum of Nature (NMC 10867).
The characters given above were found ap-
plicable also to these Canadian specimens.

Arimoto (1976) described two types of C.
(Spinicephala) verrucosa from the Japanese
coast. His robust type is identical with C.
verrucosa as described by Utinomi (1943,
1947) and Takeuchi (1989), while the slen-
der type belongs to the new species. Ari-
moto (1976) stated that the slender type rep-
resented the young stage of C verrucosa,
although its body was 6.5 mm long. Later,
in his essay (Arimoto 1978), he showed the
lateral view of slender type labelled "Ca-
prella {Spincephala [sic.]) pseudoverrucosa.^^
During the reexamination of Arimoto' s col-
lection, a specimen labelled "C. pseudover-
rucosa'' from the Tsushima Islands was
found. The specimen, representing a large
mature male of 8.79 mm long, is identified
with C. arimotoi. This indicates that Ari-
moto had come to the same conclusion as
the present author that the slender type was
not a juvenile stage of C verrucosa, but rep-
resented another species. His essay (Ari-
moto 1978), however, was written in Jap-
anese and without any taxonomic account.
Thus, it does not constitute a valid publi-
cation as specified by the International
Commission on Zoological Nomenclature
(1985; Article 8-(a)-(l) on pp. 12-13). A
taxonomic account of C (S.) pseudoverru-
cosa was never prepared by Arimoto. Ca-

prella sp. C in Takeuchi (1989) is also syn-
onymous with C. arimotoi.

In conclusion, Caprella arimotoi, n. sp.
clearly differs from C verrucosa in several
characters, and the slender "young male"
of C. {Spinicephala) verrucosa in Arimoto
(1976) and C. {S.) pseudoverrucosa in Ari-
moto (1978) are junior synonyms of C ari-
motoi.

Acknowledgments

The author thanks Prof. Ju-shey Ho and
Dr. M. Takeda for reviewing the manu-
script. Dr. H. Katayama for use of the lab-
oratory facility at the Mukaishima Marine
Biological Station of Hiroshima University,
Dr. S. Yamato for assistance in the field,
and the family of the late Dr. I. Arimoto
for the loan of the specimens. Mrs. D. R.
Laubitz kindly arranged for my short visit
to the Canadian Museum of Nature (for-
merly National Museum of Natural Sci-
ences) in Ottawa. This study was partially
supported by the Research Aid of Inoue
Foundation for Science, Tokyo.

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VOLUME 106, NUMBER 1

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PROC. BIOL. SOC. WASH.
106(1), 1993, pp. 122-130

A NEW SPECIES OF KALLIAPSEUDES
(CRUSTACEA: TANAIDACEA: KALLIAPSEUDIDAE)

FROM TRINIDAD

Roger N. Bamber

Abstract. —A new species of kalliapseudid tanaidacean, Kalliapseudes sonia-
dawnae, is described from three specimens collected from the silty-clay benthos
in 8-10 m of water in the Gulf of Paria, off the northwest coast of Trinidad.
The new species is similar to congeners from the Bahamas and the Pacific coast
of Mexico, but can be distinguished by its stouter limb and antennal articles,
and elongate dactyli of pereopods 2 and 3. It is unique in bearing a stout
pectinate spine on the basis of pereopod 1 .

Kalliapseudid tanaidaceans have been
described previously from the Atlantic
coasts of Central and South America. Lang
(1956) established the family and described
several species from Brazil, while most re-
cently Sieg (1982) described Kalliapseudes
(Mesokalliapseudes) bahamensis from the
Caribbean. In Trinidad waters, Bacescu &
Gutu (1975) described Discapseudes suri-
namensis from the Caroni Swamp on the
northern west coast. The present material
was collected in 1982 during a study of the
benthos off the Port-of-Spain coastal area,
Trinidad, in the Gulf of Paria (Agard 1984).
Three specimens of a hitherto undescribed
kalliapseudid were taken from two separate
sites.

With this limitation on the quantity of
material, only one specimen was dissected
and mounted for microscopical examina-
tion; the holotype (which was missing its
antennae) and one male paratype were ex-
amined whole; consequently features of the
detailed anatomy (e.g., the mouthparts) re-
late only to the success of the single dissec-
tion and details of, for example, the epig-
nath must await further material. The
conspicuous distinction of the new species
does not, however, depend on any such sub-
tleties.

The specimens had little sclerotization and

long setae were commonly not straight. All
figures were drawn with the aid of the cam-
era lucida and represent the anatomy as it
exists, rather than any stylized symmetrical
interpretation.

The type material is lodged at the Na-
tional Museum of Wales, Cardiff, UK
(NMW).

Description

Order Tanaidacea Hansen, 1895

Suborder Apseudomorpha Sieg, 1980

Family Kalliapseudidae Lang, 1956

Kalliapseudes {Mesokalliapseudes)

soniadawnae, new species

Material.— One 9, 6 mm long, Holotype
(NMW.Z. 199 1.099.1); one S, 3.4 mm long,
Paratype (NMW.Z. 1991.099.2), both from
Station F4, 10 m depth, 28.5°C, salinity
20%o, pH 7.35. One S, 6 mm long, mounted
in polyvinyl lactophenol, Paratype (retained
in the collection of the author). Station D4,
8.5 m depth, 26°C, 29%o, pH 6.79. These
sampling stations were situated 2 to 3 km
off the Diego Martin shore, Trinidad, ap-
proximately 10°40'N 6r35'W, 9 Aug 1982,
in silty-clay, collected by John Agard.

Body. —(Fig. 1 A) elongate (6 -times as long
as wide), unpigmented, with little scleroti-
zation.

Fig. 1. Kalliapseudes soniadawnae. A. Holotype female, body, dorsal. B. Paratype male, antenna 1. C.
Paratype male antenna 2. D. Uropod, entire and E. Basal articles, holotype female. Scale line 1 mm for A and
D, 0.2 mm for B, C and E.

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Cephalon. —Wider than long, longer than
first two pereonites together, with a distinct
semicircular rostrum, no eyes, a single an-
terior dorsolateral seta and a pair of pos-
terolateral setae on each side. Pereonites 1
to 5 increasing in length, pereonite 6 the
shortest; each pereonite with 1 anterolateral
and 1 or 2 posterolateral setae, the former
on a triangular extension on pereonites 2 to
6. Five free pleonites, of similar length, with
8 (pleonite 1) to 11 (pleonite 5) lateral setae;
each pleonite bearing pleopods. Pleotelson
(Figs. lA, 4 A) wider than long, with a pair
of terminal setae, 3 posterior setae on each
side of these, and 6 lateral setae on each side
anterior to uropod insertion; 1 pair of an-
terior dorsolateral setae.

Male antennule. —(Antenna. 1) (Fig. IB)
peduncle 4-articled; first article less than 3
times as long as wide; second as long as
wide; third and fourth wider than long, with
setation as figured. Accessory flagellum aris-
ing on fourth peduncle article and consisting
of 3 articles; third article bearing 2 distal
setae. Main flagellum of 10 articles; first as
long as wide, succeeding articles progres-
sively longer in relation to their width; tenth
article 5 times as long as wide; eighth and
tenth articles with 2 and 3 distal setae, re-
spectively; flagellum articles 1 to 4 with
dense rows of 6 to 10 aesthetascs distally;
articles 5 and 6 bearing paired aesthetascs
and article 9 with a single aesthetasc. Fe-
male antennule basal peduncle article of
similar proportions to that of the male; oth-
er articles not available.

Antenna. —{Antenna. 2) (Fig. IC) second
article with a lateral extension bearing 4 se-
tae, third article with an articulated "squa-
ma" with 5 setae; fourth article naked; fifth
setose as figured. Flagellum of 8 articles;
second to sixth articles with single tergal and
1 or 2 sternal distal setae; eighth article with
4 long and 1 short distal setae. No conspic-
uous sexual dimorphism.

Mouthparts.—Lahnim (Fig. 2C) simple,
setose; labium with very setose distal lobe
(Fig. 2E) wider than long. Mandibular palps

uniarticulate; left mandible (Fig. 2A) incisor
process with a large distal tooth and a row
of 5 smaller teeth appearing to connect to
the lacinia mobilis, itself with paired distal
teeth, lamina with 5 distal setae; right man-
dible (Fig. 2B) with a simple incisor process
with 2 or 3 teeth, lacinia mobilis reduced
or absent (not seen), lamina with 5 distal
setae. Maxilla 1 (maxillule) (Fig. 2D) inner
endite with 4 distal setae, outer endite setose
with a crown of 9 distal spines. Maxilla 2
(Fig. 2F) inner lobe with 4 plumose setae
on its outer lobe and 7 fine and 2 stout setae
on its inner lobe; endite distally with 4 wide
pectinate setae and 8 finer setae in 2 rows;
outer lobe with a wide plumose seta within
a row of 20 finer distal setae, and 3 short
lateral spines. Maxilliped (Fig. 2G) endite
typical for the subgenus, with a pair of cou-
pling hooks, all outer setae plumose; inner
edge with 3 simple setae and 6 comb-rows;
remaining articles furnished on their inner
margins with 2 parallel rows of plumose fil-
tering setae; 2 distal setae on the distal ar-
ticle. Epignath not seen in preparation.

Cheliped.—(Fig. 4C) long, slender filter-
ing structure typical of subgenus; basis 2.5
times as long as wide with a single distal
sternal seta; merus with 3 distal sternal se-
tae, carpus with 2 sternal rows of 42 and 32
filtering setae. Propodus slender (6 times as
long as wide) with a row of 1 6 filtering setae,
proximal ones as long as carpal filtering se-
tae; distally 3 rostral, 1 tergal and 8 caudal
simple setae a little longer than half length
of dactylus; distal finger extending only
slightly in sternal direction, half as long as
dactylus and with a serrated terminal spine
and 4 inner teeth. Dactylus with serrated
distal spine, a group of 3 mid-tergal setae
and a row of 7 sternal setae.

Pereopods.—FcTQopod 1 (PI) (Fig. 3 A)
proximal articles with few setae, but with a
conspicuous distal tergal pectinate spine on
the basis; distal articles with complex se-
tation and spination (Fig. 3G); merus, car-
pus and propodus armed distally with 1 , 2
and 3 stout spines tergally and 0, 1 and 2

VOLUME 106, NUMBER 1

125

Fig. 2. Kalliapseudes soniadawnae, mouthparts of paratype male. A. Left mandible. B. Right mandible. C.
Labrum. D. 1st maxilla. E. Distal lobe of labium. F. 2nd maxilla. G. Maxilliped. Scale line is 0.2 mm.

126

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Kalliapseudes soniadawnae, pereopods of paratype male. A to F. Pereopods 1 to 6, respectively. G
to K. Distal articles of pereopods 1, 2, 5 and 6, respectively. Scale line 0.4 mm for A to F, 0.2 mm for G to K.

VOLUME 106, NUMBER 1

127

Fig. 4. Kalliapseudes soniadawnae. A. Pleotelson of holotype female, dorsal. B. Pleopod (twisted) of paratype
male. C. Cheliped of paratype male. Scale line 0.26 mm for A, 0.2 mm for B and C.

128

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Spines stemally, respectively; dactylus blunt
and distally heavily setose (ca. 20 setae). P2
(Fig. 3B) with menis and carpus similarly
proportioned, with parallel sides and nearly
twice as long as wide; dactylus very long,
1.4 times length of carpus and propodus
together, with sensory "brush" at its base
and an adjacent toothed spine; setation and
spination of distal articles as in Fig. 3H. P3
(Fig. 3C) similar to P2; merus subtriangular;
carpus only 1 . 5 times as long as wide; dac-
tylus 1 .4 times length of carpus and prop-
odus together. P4 (Fig. 3D) and P5 (Figs.
3E, 3 J) similar to each other; carpus distally
with a row of 3 small spines on both caudal
and rostral edges; propodus with 2 parallel
combs of 7 spines of progressively increas-
ing length from proximal (tergal) to distal
(sternal) edges; dactylus a blunt, setose sen-
sory organ. P6 (Figs. 3F, 3K) carpus with a
tergal row of 5 long setae and a long distal
seta 1.8 times length of propodus; propodus
with 2 tergal "sole-" spines and an adjacent
comblike row of 1 4 smaller spines; dactylus
1.75 times length of propodus.

Pleopod. —(Fig. 4B) basis with 3 setae; en-
dopod first article with a single seta; second
article and exopod less than twice as long
as broad, with 1 1 and 12 setae, respectively.
Uropod (Figs. ID, IE) biramous, almost as
long as pleon; endopod with 3 articles, exo-
pod with numerous articles (1 5 to 18), some
irregular.

Male specimens with conspicuous genital
cone on sternum of pereonite six, showing
no obvious (without dismemberment) sig-
nificant differences in cheliped or antennal
morphology (one would not expect the dense
tufts of aesthetascs on the proximal articles
of the female antennule main flagellum). All
three type specimens had a cone-like ventral
process on the sternum of pleonite 1 .

Etymology.— This, most attractive spe-
cies is named after the marine biologist So-
ma Dawn Batten.

Remarks

Kalliapseiides soniadawnae is clearly close
to both K. viridis Menzies, 1953, from the

Pacific coast of Mexico, and K. bahamensis
Sieg, 1982 from the Bahamas. With the for-
mer it shares the rectangular pereonites with
anterolateral "triangular" spine-bearing
processes (a distinction between these de-
scribed species emphasized by Sieg 1982)
and a single seta on the first pleopodal en-
dopod article; with the latter it shares few
caudolateral setae on the telson and long
dactyli on pereopods two and three (al-
though these are even longer in proportion
to the combined length of carpus and prop-
odus in the present species— 1 .4 times— than
in K. bahamensis— 1.2 times). These long
dactyli and the distinct rounded rostrum,
better developed than in either of the other
two species, are distinguishing features of
K. soniadawnae evident from whole-animal
observation. Equally, the new species has
stouter articles on its limbs and antennae,
is generally less setose (although the filtering
setae of the propodus of the cheliped are as
long as those of the carpus in K. sonia-
dawnae, proximally shorter K. viridis and
in K. bahamensis), and, uniquely, a heavy
pectinate spine on the distal tergal comer
of the basis of pereopod 1 (only a long sim-
ple seta in the other two species).

Examination of 76 paratypes of K. ba-
hamensis, kindly loaned by the National
Museum of Natural History, Washington
(USNM 181901) revealed that, despite the
description in Sieg (1982), nearly half of
these specimens were male with genital
cones. This allowed the examination of sex-
ual dimorphism and variability in this spe-
cies.

The antennae of a male paratype of K.
bahamensis are shown in Fig. 5. Antenna 1
has stouter articles than those of the female,
and bears dense tufts of aesthetascs on the
proximal 5 articles of the main flagellum.
In addition, the distal setae on the propodus
of the male cheliped are elaborately pecti-
nate, being largely simple in the female (and
simple in the male of AT. soniadawnae). There
is no other conspicuous sexual dimorphism.
The proportions of the articles of antenna
2 are similar in both sexes (as in K. sonia-

VOLUME 106, NUMBER 1

129

Fig. 5.
mm.

Kalliapseudes bahamensis. A. Antenna 1, male paratype. B. Antenna 2, male paratype. Scale line 0.3

dawnae)\ it is therefore reasonable to as-
sume that the males of K. viridis will have
slender antenna 2 articles as found in the
female, and distinct from the more robust
morphology of K. soniadawnae.

Variability in the lengths of the dactyli of
pereopods 2 and 3 was analyzed by mea-
suring them in comparison to the length of
the adjacent, shorter toothed spine (see Fig
3H). This proportion showed no significant
difference between the two limbs. In K. ba-

hamensis the dactyl was 3 times the length
of the adjacent spine (mean from 10 spec-
imens 2.95, range 2.6 to 3.4) while in K.
soniadawnae XhQ dactyl was 4 times the spine
length (mean from all three specimens 4. 12,
range 3.75 to 4.45). In K. viridis this pro-
portion is about 2 (e.g., Sieg 1982:fig. 7).

Thus, of these three closely related spe-
cies, K. bahamensis is immediately distin-
guishable by its characteristic trapezoidal
pereonite morphology, in having more than

130

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1 Sternal setae on the cheliped dactylus and
its uropod basis exceeding the posterior tip
of the telson by about one-third of its length.
The remaining two zoogeographically iso-
lated species have rectangular pereonites,
less than 10 sternal setae on the cheliped
dactylus and uropod basal articles not or
only just exceeding the telson. They are dis-
tinguishable by the heavy spine on the basis
of pereonite 1 , the elongate dactyli of pere-
onites 2 and 3 and the stouter antennal ar-
ticles shown only by K. soniadawnae.

Acknowledgments

I am most grateful to J. Gobin for bring-
ing the specimens to my attention and for
supplying the data on the sample sites, to
Janice Clark of the National Museum of
Natural History, Washington, for the loan
of paratypes of Kalliapseudes bahamensis,
and to Dr. D. Holdich for access to relevant
literature.

Technical Report, Institute of Marine Affairs,
Trinidad, 79 pp.

Bacescu, M., & Gutu, M. 1975. A new genus {Dis-
capseudes n.g.) and three new species of Apseu-
didae (Crustacea, Tanaidacea) from the north-
eastern coast of South America.— Zoologische
Mededelingen 49:95-1 13.

Hansen, H. J. 1895. Isopoden, Cumacean und Sto-
matopoden der Plankton-Expedition. Ergeb-
nisse der Atlantischer Ozean Plankton Expedi-
tion Humboldt-Stiftung 2 (G), Lipsius & Tischer,
Kiel, 105 pp. + pis 1-8.

Lang, K. 1956. Tanaidacea aus Brasilien, gesammelt
von Professor Dr. A. Remane und Dr. S. Ger-
laeh.— Kieler Meeresforschungen 12:249-260.

Menzies, R. J. 1953. The apseudid Chelifera of the
eastern tropical and north temperate Pacific
Ocean.— Bulletin of the Museum of Compara-
tive Zoology 107(9):443-496.

Sieg, J. 1980. Sind die Dikonophora eine polyphy-
letische Gruppe?— Zoologischer Anzeiger 205:
401-416.

. 1982. Anmerkungen zum Genus ^a///a/75^w-

des Stebbing, 1910 mit Beschreibungen einer
neuen Art Kalliapseudes bahamensis n.sp.
(Crustacea, Tanaidacea).— Mitteilungen aus dem
Zoologischen Museum der Universitat Kiel 1(9):
3-17.

Literature Cited

Agard, J. B. R. 1984. A baseline study of the effects
of pollution on the benthos of the nearshore
Diego Martin to Port of Spain coastal area. —

FAWLEY Aquatic Research Laborato-
ries, Marine & Freshwater Biology Unit,
Fawley, Southampton S04 ITW, Hants.,
United Kingdom.

PROC. BIOL. SOC. WASH.

106(1), 1993, pp. 131-136

ENTEROCOLA AFRICANUS, A NEW SPECIES

(COPEPODA: ASCIDICOLIDAE) ASSOCIATED WITH A

COMPOUND ASCIDIAN SYNOICUM SPECIES FROM

NORTH AFRICA (STRAIT OF GIBRALTAR)

Pablo J. Lopez-Gonzalez, Mercedes Conradi, and
J. Carlos Garcia-Gomez

Abstract. —The genus Enterocola van Beneden, 1860 is principally recorded
from European waters. In this paper Enterocola africanus, new species, from
the Strait of Gibraltar (North African side) is described from the compound
ascidian Synoicum sp. Enterocola fulgens, Enterocola clavelinae, Enterocola
hessei, Enterocola precaria and Enterocola ianthina are morphologically similar
to the new species. All are discussed and compared with Enterocola africanus.
Resumen. —1.3. mayor parte de las especies descritas dentro del genero En-
terocola van Beneden, 1860 proceden de las costas Europeas. Se describe una
nueva especie, Enterocola africanus del Estrecho de Gibraltar (vertiente Norte
Africana). Enterocola fulgens, Enterocola clavelinae, Enterocola hessei, Enter-
ocola precaria y Enterocola ianthina son morfologicamente proximas a la nueva
ijp especie. Todas ellas son comparadas y discutidas con Enterocola africanus.

Recently the Laboratorio de Biologia Ma-
rina of the University of Sevilla and Cadiz
(Spain) initiated a program to study the co-
pepods associated with marine inverte-
brates from the coasts of the Strait of Gib-
raltar and nearby areas. So far, three marine
biological expeditions "Bahia 90" and "Ba-
hia 91" in Algeciras Bay (Southern Iberian
Peninsula) and "Ceuta 91" in Ceuta (North
Africa) were carried out within a more com-
prehensive program of marine benthos. The
studies were centered mainly on the cope-
pod fauna associated with molluscs, ascid-
ians, anthozoans, and echinoderms. The first
results have already been reported (Lopez-
Gonzalez et al. 1992).

Four female parasitic copepods belonging
to the genus Enterocola were found in the
colonies of the compound ascidian, Synoi-
cum sp. collected during the "Ceuta 91"
Expedition. They were later determined to
be new to science.

Studies of the genus Enterocola have been
concentrated in European waters with lim-

ited references to the North American coast
and the Philippine Islands (Illg & Dudley
1980). Recently, Ooishi (1987) recorded an
undetermined species of this genus from
Okinawa. Shellenberg (1922) reported Bo-
tryllophilus sp. associated to Polycitor ren-
ieri from Plattenberg Bucht (South Africa),
and not Enterocola sp. as Illg & Dudley
(1980) referred to in their monograph on
the Ascidicolidae.

In this work, Enterocola africanus from
the North African coast (Ceuta) is de-
scribed. It represents the only member of
its genus described from this continent. Al-
though it is not the only species reported,
Barnard (1955) quotes Enterocola fulgens
from South Africa.

Material and Methods

The compound ascidians were collected
on stones from the infralittoral zone (6-12
m deep). They were maintained in separate
glass bottles. The copepods were removed

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

through dissection of hosts and preserved
in formahn (4% in sea water). The speci-
mens were stained with cotton blue, dis-
sected under a stereomicroscope, and semi-
permanent mounts were made using
lactophenol. All figures were drawn with the
aid of a camera lucida. The letter after the
explanation of each figure refers to the scale
at which it was drawn.

Family Ascidicolidae Thorell, 1859

Subfamily Enterocolinae Delia Valle, 1883

Genus Enter ocola van Beneden, 1860

Enterocola africanus, new species

Figs. 1-2

Type material— A 22 from 3 colonies of
Synoicum sp. at Ceuta (Spain, North Africa)
(35°53'430''N; 15°17'W), 18 Aug 1991. The
holotype has been deposited in the Museo
Nacional de Ciencias Naturales de Madrid
(Spain) (MNCNM 20.04/3504). The three
paratypes (two dissected) in the collection
of the authors.

Female.— ^ody (Fig. la-c) of relaxed
specimens 0.97 mm total length (based on
4 specimens). Proportions of cephalosome :
metasome : urosome, 1:4.2:1.48. Uncon-
tracted specimen with dorsal sutures and
urosome folded without segmentation. Body
covered ventrally with discontinuous rows
of spinules.

Antennule (Fig. le) unimerous, apically
narrowing suggesting 2 possible segments.
Basal article covered with discontinuous
rows of spinules. Distal component with
minute conical apical protuberance. Junc-
tion of first and second components with
about 5 setules. Antenna (Fig. If) obscurely
bimerous. Basal segment unarmed, but with
several rows of minute spinules. Distal seg-
ment with 1 subterminal and 5 terminal
setae and several rows of minute spinules.
Labrum (Fig. Ig) with 2 lateral spinose lobes;
ventral surface with rows of spinules. Max-
illule (Fig. 2a, b) bilobed. Basal portion la-
melliform, somewhat trilobed distally; dis-
tal third of anterior margin with

unomamented lobe (Fig. 2a, b). A promi-
nence near articulation of palp bearing
barbed seta and short setule. Palp with 5
spinulose setae and 1 unomamented seta on
its distal truncate margin. Maxilla (Fig. 2c)
with a massive basal segment bearing an
articulated digitiform, spinulose endite at
its distal medial comer. Apical segment bi-
fid distally, with one process somewhat
shorter than the other, narrower than basal
segment, but also heavily sclerotized. Junc-
tion of basal and apical segments with 1
short seta.

Intercoxal area of leg 2-4 with pro-
nounced mammilliform processes (Fig. la,
b). Legs 1-4 (Fig. 2d-g) biramous. Anterior
surface of legs bearing rows of spinules. Pro-
topodites with small seta at distal lateral
comer. All legs with bimerous protopodites
and unimerous rami. Exopodites terminat-
ing in a pointed process and slightly curved
laterally. Exopodite of third leg (Fig. 2f) with
characteristic styliform outline in most spe-
cies of the genus. Endopodites approxi-
mately equal to exopodites; lateral margin
more convex than medial margin; 2 apical
setae of each endopodite set close together
and longer than ramus. Outer apical seta
about 1.7 times the length inner apical seta
on all legs. Pediform projection (probably
leg 5) (Fig. lb) a plate with subcircular mar-
gin, bearing 2 separate minute setules. Cau-
dal rami (Fig. la-c) conical and apparently
forming a definitive articulation with uro-
some.

Male unknown.

Etymology.— ThQ specific name africa-
nus was chosen because this is the first spe-
cies of this genus described from Africa.

Discussion

There are five species of Enterocola van
Beneden, 1 860 with mammiliform process-
es present between at least one pair but not
all pairs of legs: Enterocola fulgens van Be-
neden 1860, Enterocola clavelinae Chatton
& Harant 1924, Enterocola hessei Chatton

VOLUME 106, NUMBER 1

133

Fig. 1 . Enterocola africanus, new species, female: a, habitus, ventral (A); b, habitus, lateral-oblique (A); c,
habitus, dorsal (A); d, oral region (B); e, antennule (C); f, antenna (C); g, labrum (C). Scale bars, A: 600 fim\ B:
50 MHi; C: 50 ^ni.

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Enterocola africanus, new species, female: a and b, maxillule (A), ul: unomamented lobe of the distal
third anterior margin; c, maxilla (A); d, first leg (B); e, second leg (B); f, third leg (B); g, fourth leg (B). Scale
bars, A: 25 tivrv; B: 50 /um.

VOLUME 106, NUMBER 1

135

Table 1.— Comparison of selected features between Enterocola clavelinae, Enterocola precaria, Enterocola
ianthina and Enterocola africanus, new species.

E. clavelinae

E. precaria

E. ianthina

E. africanus

Antennule

Unsegmented lobe

Unimerous with

Obscurely bimerous

Unimerous with 5

with 2 anterior setae

small apical setules

with about 7 setules

setules

and 3 terminal setae

Antenna

Strongly bimerous

Unimerous

Bimerous

Obscurely bimerous

Ratio

Endopodite longer

In the first, second

Endopodite approxi-

Endopodite:

than exopodite

and fourth legs exo-

mately as long as

exopodite

podite are shorter
than endopodite but
in the third leg exo-
podite is longer
than endopodite

exopodite

Length of 2 apical

Equal

Outer apical seta 1.7

setae

times as long as
inner

pediform projection

With single small

With single setule

Not element of

With 2 separated

setule on the margin

in the middle

armature although
there are 4 shallow
emarginations

minute setules

Caudal rami

Are not delimited

Without apparent

With apparent

from urosome

articulations

& Harant 1924, Enterocola precaria Illg &
Dudley 1980, and Enterocola ianthina Illg
& Dudley 1980. None of these species have
a body covered ventrally with discontinu-
ous rows of spinules like the new species.

Enterocola fulgens has setae of antenna
short, hooked, while E. africanus has these
setae very long and flexible.

The most important difference between
E. hessei and the other species is that the
two apical setae of each endopodite are set
on the lateral rather than on the apical sur-
face, and so are diverging from the axis of
the ramus.

Differences between the new species and
the other three species are summarized in
Table 1.

The diagnostic feature of Enterocola af-
ricanus are: presence of mammiliform pro-
cesses at the bases of legs 2-4, antennule
being unimerous with five setules, antenna
obscurely bimerous with six long flexible
setae, length ratio of endopodite : exopo-
dite, length of two apical setae of the endo-

podite of the leg, armature of pediform pro-
jection, and caudal rami articulated.

Acknowledgments

We thank Dr. P. L. Illg, Dr. P. L. Dudley
and Dr. V. Gotto for their generous help
with information and literature in the course
of this work. Rocio Juan provided valuable
assistance during the field work. We are
grateful to Dr. L. Cervera for help during
this project.

Appreciation is extended to CEPSA, Se-
villana de Electricidad, Excmo. Ayunta-
miento de los Barrios, Mancomunidad de
Municipios del Campo de Gibraltar and
Agencia de Medio Ambiente (Junta de An-
dalucia) for financial support of this work.

Literature Cited

Barnard, K. H. 1955. South African parasitic Co-
pepoda.— Annals of the South African Museum
41:223-312.

136

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Beneden, van P. J. 1860. Sur un nouveau genre de
crustace Lemeen. — Bulletin de I'Academie de
Belgique, serie 2, 9:151-160.

Chatton, E., & H. Harant. 1924. Notes sur les Co-
pepodes ascidicoles. XV. Sur trois formes nou-
velles du genre Enterocola P. J. van Beneden.
Etat actuel de la systematique des Enterocolinae
n. subf.— Bulletin de la Societe Zoologique de
France 49:354-364.

Delia Valle, A. 1883. Sui copepodi che vivono nelle
Ascidie composite del Golfo di Napoli.— Atti
della R. Accademia dei Licei, serie 3, Memoire
della Clase de Scienze Fisiche, Matematiche e
Naturali 15:242-253.

Illg, P. L., & P. L. Dudley. 1980. The family Asci-
dicolidae and its subfamilies (Copepoda, Cyclo-
poida) with descriptions of new species.— Me-
moires du Museum National d'Histoire
Naturelle, ser. A, T. 117:1-192.

Lopez-Gonzalez, P. J., M. Conradi, S. Naranjo, & J.
C. Garcia-Gomez. 1 992. A new species of ^«-
thessius (Copepoda: Poecilostomatoida) asso-
ciated with Berthella stellata Risso, 1826 (Gas-
tropoda: Opisthobranchia). — Proceedings of the
Biological Society of Washington 105:240-248.

Ooishi, S. 1987. A preliminary list of copepods as-
sociated with ascidians collected around Sesoko
Island, Okinawa.— Galaxea 6:95-98.

Schellenberg, A. 1922. Neue Notodelphyiden des
Berliner und Hamburger Museums mit ein
Ubersicht der ascidienbewohnenden Gattungen
und Arten.— Mitteilungen aus dem Zoologi-
schen Museum in Berlin 10:277-362.

Thorell, T. 1859. Till Kannedomen om vissa par-
asitiskt lefvande Entomostraceer.— Ofversigt at
Kongl. Vetenskaps-Academiens Forhandlingar
16, 8:335-362.

(PJLG) (JCGG) Laboratorio de Biologia
Marina, Departamento de Fisiologia y Bio-
logia Animal, Facultad de Biologia. Univer-
sidad de Sevilla, Apdo. 1095, 41080 Sevilla,
Spain; (MC) Laboratorio de Biologia, Fa-
cultad de Ciencias del Mar. Universidad de
Cadiz. Apdo. 40, 1 1510, Puerto Real (Cad-
iz), Spain.

PROC. BIOL. SOC. WASH.
106(1), 1993, pp. 137-146

NEW SPECIES AND NEW RECORDS OF THE GENUS
ELAPHOIDELLA (CRUSTACEA: COPEPODA: HARPACTICOIDA)

FROM THE UNITED STATES

Janet W. Reid and Teruo Ishida

Abstract. —We list the first record ofElaphoidella wilsonae from New Mexico
and new records of Elaphoidella bidens from Maryland, Tennessee, Virginia
and the District of Columbia. Two new species of harpacticoid copepods from
the eastern United States, Elaphoidella carterae from Virginia and Elaphoidella
amabilis from Maryland differ from congeners in the shapes of the caudal ramus
and caudal setae and the spine formulas of the swimming legs. We provide
keys to the known species of Elaphoidella from North America.

Collections of harpacticoid copepods
from springs and streams in the District of
Columbia, Maryland, New Mexico and Vir-
ginia included several species, two previ-
ously undescribed, belonging to the harpac-
ticoid copepod genus Elaphoidella. We list
the new records of the previously known
species and describe the new ones. The de-
scription of each species was authored by
its collector. We furnish identification keys
and a table of distinguishing characters of
both sexes of the known North American
species. For taxonomic examination, spec-
imens were drawn with the aid of drawing
tubes before dissection in lactic acid and
after dissection in polyvinyl lactophenol
with a little chlorazol black E added, or in
gum-chloral medium. Lengths were mea-
sured from the anterior tip of the rostrum
to the end of the caudal ramus. Specimens
were deposited in the National Museum of
Natural History, Smithsonian Institution
(USNM).

Hamond (1987) returned several generic
and subgeneric taxa including Elaphoidella
to the synonymy of the genus Canthocamp-
tus Westwood, 1836 s. 1. pending eventual
revision of the family Canthocamptidae.
However, we have employed the more fa-
miliar genus name without wishing to imply

recognition at the generic level of this poorly
defined group of species.

Order Harpacticoida G. O. Sars, 1903

Family Canthocamptidae G. O. Sars, 1906;

Monard, 1928; Lang, 1948

Genus Elaphoidella Chappuis, 1929

Elaphoidella wilsonae YiunX, 1979

Elaphoidella wilsonae \^vin\, 1979:248-253,
figs. 1-21.

Material examined. — 1 9, in 70% ethanol,
Guadalupe River, Jemez National Forest,
about 40 km NE of San Ysidro, New Mex-
ico, about 35°45'N 106°50'W, elevation
about 2100 m, damp moss by streamside,
26 May 1991, col. E. Warner (USNM
251152).

Remarks. —The specimen from New
Mexico agrees in all respects with the de-
scription of females from the type popula-
tion (Hunt 1979).

Distribution and habitat.— The Guada-
lupe River is a third-order stream in the
drainage basin of the Rio Grande. This spe-
cies was formerly known only from alkaline
spring-fed ponds in Garfield and Rio Blanco
Counties, Colorado, in the basins of the Col-
orado and Green Rivers respectively. The
find reported herein extends its known dis-

138

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

tribution some 400 km to the south and
newly includes the Rio Grande drainage ba-

sm.

Elaphoidella bidens (Schmeil, 1893)

Synonymy.— GivQn by Lang (1948) and
Apostolov(1985).

Material examined. — $, ethanol-pre-
served, Lonaconing Creek, south of Raw-
lins, Allegany County, Maryland, about
39°31'N 78°54'38"W, 2 Feb 1988, col. M.
C. Swift (USNM 242082). 2 $, ethanol-pre-
served, Piney Creek, Garrett County, Mary-
land, about 39°42'21"N 78°57'45"W, 6 Apr
1988, col. M. C. Swift (USNM 242084). 3
2, Mountain Lake, Giles County, Virginia,
37°2r22"N80°32'll"W, elevation 1181 m,
sandy bottom at small swimming beach on
south shore, 25 May 1990, together with E.
carterae, col. J. W. Reid (USNM 250448).
3 9 on 2 slides, and 4 9, 1 copepodid, eth-
anol-preserved, near-bank sediments of
Rock Creek, District of Columbia, just east
of Maryland border, about 38°59'00"N
77°03'10"W, 5 Oct 1990, col. T. Ishida
(USNM 251796).

Remarks. —Elaphoidella bidens is a usu-
ally parthenogenetic species recorded from
nearly every continent. The taxonomy of
the two subspecies, E. bidens s. s. and E.
bidens coronata (G. O. Sars, 1 904), recorded
from North America was discussed by Wil-
son (1956, 1975) and Wilson & Yeatman
(1959). More recently Apostolov (1985),
citing morphological variations including
the coronata-iorvci present in topotypic pop-
ulations of £". bidens, returned several sub-
species to the nominate species taxon.

North American records of £". bidens co-
ronata reviewed by Wilson (1975) include
Florida, Georgia, Louisiana, Minnesota,
North Carolina, Ohio, Pennsylvania, and
Virginia. These records include a report by
Carter (1944) from the region of Mountain
Lake. Subsequent records of E. bidens s. 1.
from North America include Coahuila,
Mexico by Reid (1988) and New York by
Strayer [1988 (1989)]. H. C. Yeatman col-

lected an ovigerous female E. bidens from
a small lake at Sewanee, Tennessee on 8
April 1973; this is a new record for that state
(H. C. Yeatman, pers. comm.). The new
records herein, including the first reports
from the District of Columbia, Maryland
and Tennessee, are well within the known
range of this species in North America.

Elaphoidella carterae Reid, new species
Figs. 1, 2

Material examined.— ^oloXypt 9, dis-
sected and mounted on slide in polyvinyl
lactophenol (USNM 25 1 767), and paratype
9, in 70% ethanol (USNM 251768), from
Mountain Lake, Giles County, Virginia,
37°21'22"N80°32'11"W, elevation 1181 m,
sandy bottom at small swimming beach on
south shore, 25 May 1990, col. J. W. Reid.

Female.— YidihiXus (Fig. la) cylindrical.
Length of holotype 0.76 mm, of paratype
0.62 mm. Cephalosome (Fig. la, b) with
elongate ovoid nuchal organ. Hyaline fring-
es of posterior margins of all somites smooth.
All somites with scattered long hairs and all
somites except cephalosome with trans-
verse rows of tiny hairs, surface of all so-
mites also finely punctate as in area within
dotted line (indicated by arrow) on lateral
surface of pediger 2 (Fig. la). Genital seg-
ment (Fig. la, c) with remnant of division
visible laterally beneath integument (indi-
cated by dotted line in Fig. la); ornamented
with short transverse row of small spines
lateral to genital field; genital field reaching
midlength of segment. Two urosomites pos-
terior to genital segment (Fig. la, c) each
with one row of small spines on ventral and
lateral margin. Anal somite (Fig. la, c-e)
with two spines near posteroventral margin
above each caudal ramus; anal operculum
smooth, slightly convex. Caudal ramus (Fig.
la, c-e) about 1.2 times longer than broad,
ovate, with dorsal, terminally hooked lon-
gitudinal keel extending about % length of
ramus, and small subdistal medial lobe. Ra-
mus with basally biarticulate dorsal seta in-
serted lateral to end of keel, two lateral se-

VOLUME 106, NUMBER 1

139

Fig. 1. Elaphoidella carteme Reid, new species, female, holotype (USNM 251767): a. Habitus, left lateral
(arrow indicates detail of somitic punctations); b, Cephalosome, dorsal; c, Urosome, ventral; d. Anal somite
and caudal ramus, left lateral; e. Part of anal somite and right caudal ramus, dorsal (somewhat compressed in
permanent mount); f, Antennule; g, Antenna; h. Mandible; i, Maxillule (part); j. Maxilla; k, Maxilliped. Scales
= 50 ^m.

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Elaphoidella carterae Reid, new species, female, holotype (USNM 25 1767): a, Left leg 1 and coupler;
b, Right leg 2 and coupler; c, Left leg 3 and coupler; d, Left leg 4 and coupler; e, Right leg 5 and coupler. Scale
= 50 Mm.

tae, longitudinal row of five spines slightly
ventral to insertion of distal lateral seta,
group of fine hairs distal to medial lobe, and
three terminal setae. Median terminal setae
broken in both specimens, lacking proximal
breaking plane and ornamented with few
spiniform setules; lateralmost terminal seta
with bulbous base and slender tip; medi-
almost terminal seta stout, tapering, slightly
shorter than lateralmost terminal seta; both
medialmost and lateralmost terminal setae
ornamented with fine hairs. Holotype bear-
ing long ovoid spermatophore (Fig. Ic).
Rostrum (Fig. 1 a, b) short, subtriangular.

with two sensillae. Antennule (Fig. If) of
eight articles, article 4 with long broad es-
thetasc reaching past end of antennule, ar-
ticle 8 with shorter slender esthetasc. An-
tenna (Fig. Ig) biarticulate, exopodite
uniarticulate with four setae. Exopodite of
mandible (Fig. Ih) biarticulate, proximal
and distal articles with one and four setae
respectively. Maxillule (Fig. li) partly ob-
scured in mount, basis with three visible
setae and long terminal claw. Maxilla (Fig.
Ij) also with three setae on basis. Maxilliped
(Fig. Ih) prehensile, with basis broken but
lacking seta present on some congeners.

VOLUME 106, NUMBER 1

141

Legs 1-4 (Fig. 2a-d) each with triarticu-
late exopodite; endopodite of leg 1 triarticu-
late, longer than exopodite; endopodites of
legs 2-4 each biarticulate. Formula for ma-
jor armament as follows:

Legl

basis 1-1

exp 0- 1 :

1-1; 0,2,2

enp 1-0

, 1-0; 1,2,0

Leg 2

basis 0-1

exp 0- 1 •

1-1; 1,2,2

enp 1-0

, 1,2,1

Leg 3

basis 0- 1

exp 0-1

0-1; 2,2,2

enp 0-0

, 1,2,1

Leg 4

basis 0- 1

exp 0-1

1-1; 2,2,2

enp 1-0

,2,1,1

Major lateral spines of leg 3 exopodite ar-
ticles 1 and 2 very large, curved posteriorly.
More distal setae of exopodites of legs 2-4
and endopodites of legs 3 and 4 unusually
stout, almost spiniform. Couplers of all legs
without ornament.

Leg 5 (Fig. 2e), medial expansion of ba-
soendopodite reaching less than Vi length of
exopodite. Basoendopodite and exopodite
each with four setae of which lateral and
medialmost setae are very short and two
medial setae longer, all setae stout, spini-
form.

No variation was observed between the
two specimens.

Male. —Unknown.

Etymology. —Dr. Marjorie Estelle Carter
collected copepods from Mountain Lake and
its environs for nearly two decades, but pub-
lished only two articles, one posthumously,
from those studies (Carter 1944, Carter &
Bradford 1972). Her collection, which ap-
parently included numerous undescribed
species, no longer survives (H. H. Hobbs,
Jr., pers. comm. to JWR). It is a pleasure
to pay tribute to Dr. Carter's contributions
to knowledge of American harpacticoid co-
pepods by naming this species for her.

Comparisons.— The form of the caudal
ramus, especially the medial protrusion, and
the terminal caudal setae resemble those of
no known member of the Elaphoidella-
group. The major setation of legs 1-4 also

Table 1.— Number of major setae and spines on
proximal: distal articles of endopodites of legs 2-4 (fe-
males) and legs 2 and 4 (males) and on basoendopod-
ites: exopodites of leg 5 of species of Elaphoidella re-
corded from North America. (Males of £". californica,
E. carterae, and E. kodiakensis are unknown.)

Species

Leg 2

Leg 3

Leg 4

Legs

Females

amabilis

1:3

0:3

4:3

shawangunkensis

1:3

0:5

0:3

4:3

carterae

1:4

0:4

1:4

4:4

subgracilis

1:4

1:5

1:4

4:4

californica

1:5

1:4

4:5

bidens, kodiakensis.

reedi, wilsonae

1:5

1:6

1:4

4:5

Males

shawangunkensis

1:3

—

0:3

subgracilis

1:3

—

0:4

amabilis

1:3

—

0:3

0:4

reedi, wilsonae

1:4

—

0:3

0:4

bidens

1:4

—

0:4

^ The leg 4 endopodite of the male of E. bidens is
uniarticulate.

differs from known North American species
(Table 1).

Elaphoidella amabilis Ishida, new species

Material examined. —Holotype 2, dis-
sected and mounted on slide (USNM
25 1 799). Allotype 6, dissected and mounted
on slide (USNM 251800). Paratypes: 2 $
and 1 6, mounted whole together on slide
(USNM 251798), and 2 9, in 70% ethanol
(USNM 25 1797). All from perennial spring,
southeast comer of Maryland Maintenance
Facility, Rock Creek Stream Valley Park,
Montgomery County, Maryland, approxi-
mately 100 m west of boundary with Wash-
ington, D.C., 38°59'16"N 77°03'18"W, 5 Oct
1990, col. T. Ishida. Mounted specimens in
gum-chloral medium.

Female. —Habitus (Fig. 3a) cylindrical.
Length of holotype 0.57 mm, of mounted
paratypes 0.55 and 0.56 mm. Hyaline fring-
es of posterior margins of all somites smooth.
Surface of all somites rather smooth, faintly

142

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

HC,d,e,g

Fig. 3. Elaphoidella amabilis Ishida, new species, female, b-f, holotype (USNM 251799), a, g, paratype
(USMM 25 1 798): a, Habitus (somewhat compressed in permanent mount), left lateral; b, Antennule; c. Antenna;
d, Genital field; e, Penultimate urosomite, anal somite, and caudal rami, ventral; f, Anal somite and caudal
rami, dorsal; g. Anal somite and caudal ramus, left lateral. Scales =100 jum.

punctate. Posterior end of seminal recep-
tacle (Fig. 3d) reaching % length of genital
segment. Penultimate urosomite (Fig. 3a, e)
with rows of small spines on lateral and
ventral margin. Anal somite (Fig. 3e-g) with
transverse row of lateral spines on each side,
and with two spines near posteroventral
margin above each caudal ramus; anal oper-
culum convex, with marginal comb. Caudal
ramus (Fig. 3e-g) about 1.5 times longer

than broad, rectangular, with dorsal, ter-
minally hooked longitudinal keel extending
about % length of ramus. Caudal ramus with
basally biarticulate dorsal seta inserted lat-
eral to end of keel, two lateral setae, each
with transverse row of four spines at base,
and three terminal setae. Median terminal
seta lacking proximal breaking plane, ba-
sally expanded with ventral knob at inser-
tion, remaining part of seta slender, about

VOLUME 106, NUMBER 1

143

Fig. 4. Elaphoidella amabilis Ishida, new species, female, holotype (USNM 251799): a, Right leg 1 and
coupler: b, Left leg 2 and coupler; c. Right leg 3 and coupler; d, Left leg 4 and coupler; e, Left leg 5 and coupler.
Male, allot>T)e (USNM 25 1800): f. Anal somite and caudal rami, dorsolateral; g. Left leg 3 and coupler; h, Right
leg 5. Scale = 100 jum.

1.3 times longer than urosome. Lateralmost
terminal seta slender, ventrally curved
proximally, base bulbous with acute dorsal
process. Medialmost terminal seta stout
proximally, tapering distally, about V2 length
of lateralmost terminal seta. Medialmost and
lateralmost terminal setae ornamented with
fine hairlike setules, median terminal seta
ornamented with short stiff setules.

Rostrum short. Antennule (Fig. 3b) of
eight articles, article 4 with long esthetasc
reaching past end of antennule, article 8 with
shorter esthetasc. Antenna (Fig. 3c) biartic-
ulate, exopodite uniarticulate with four se-
tae.

Legs 1-4 (Fig. 4a-d) each with triarticu-
late exopodite; endopodite of leg 1 triarticu-
late, longer than exopodite; endopodites of
legs 2—4 each biarticulate. Formula for ma-
jor armament as follows:

Leg I

basis 1-1

exp 0-1

1-1; 0,2,2

enp 1-0

, 1-0; 1,2,0

Leg 2

basis 0-1

exp 0-1

1-1; 1,2,2

enp 1-0

, 1,1,1

Leg 3

basis 0-1

exp 0-1

0-1; 2,2,2

enp 0-0

, 1,1,1

Leg 4

basis 0-1

exp 0-1-

1-1; 2,2,2

enp 0-0

, 1,1,1

Major lateral spines of leg 3 exopodite ar-

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

tides 1 and 2 large, curved posteriorly. Dis-
tal medial setae of endopodites of legs 3 and
4 short. Couplers of all legs without orna-
ment.

Leg 5 (Fig. 4e), medial expansion of ba-
soendopodite reaching less than V2 length of
exopodite. Basoendopodite with four setae,
lateralmost seta very short and two medial
setae longest. Exopodite with three setae,
medialmost seta very short.

Male. —l^engih of allotype 0.52 mm, of
paratype 0.51 mm. Urosomite 3 with one
row of small spines on ventral and lateral
margin. Anal somite (Fig. 4f) similar to fe-
male, but with one spine near posteroven-
tral margin above each caudal ramus. Cau-
dal ramus (Fig. 4f) subrectangular, with
dorsal keel extending about Vi length of ra-
mus, dorsal seta biarticulate at base, two
lateral setae each with transverse row of four
spines at base. Median and lateralmost ter-
minal setae without bulbous bases.

Legs 1 , 2, and 4 similar to those of female.
Leg 3 (Fig. 4g) exopodite, major lateral
spines of articles 1 and 2 very large, curved
posteriorly; major setae and spines of article
3 shorter than those of female. Leg 3 en-
dopodite triarticulate, modified, spiniform
process of article 2 reaching only midlength
of exopodite article 3, article 3 with two
short apical plumose setae.

Leg 5 (Fig. 4h) basoendopodite reduced,
lacking armament; exopodite slightly longer
than broad and bearing four spines, next
innermost spine longest.

No variation was observed between spec-
imens of either sex.

Etymology.— NsLTned for the lovely as-
pect of the body, especially the female cau-
dal rami.

Remarks. —The type locality is the same
as that of Attheyella (Mrazekiella) spinipes
Reid, 1987.

Comparisons. —Elaphoidella amabilis,
like E. carterae is highly distinctive in the
structure of the caudal rami and caudal se-
tae. The major setation of the swimming
legs of the female is the most reduced of
known North American species (Table 1).

The male resembles E. subgracilis in seta-
tion of the swimming legs, as far as the latter
species has been described, but differs in
lacking a papilla on the lateral surface of the
caudal ramus.

Keys to Continental North American
Species of Elaphoidella

The following key to females of conti-
nental North American species oi Elaphoi-
della represents a considerable departure
from the previous key of Hunt (1979) in
that it is based primarily on the setation of
legs 2-4. These meristic characters are eas-
ier to interpret than are descriptions of form
in a key without illustrations. However there
is always the possibility of variation in num-
ber of setae and users should consult the
original species descriptions. As an addi-
tional aid, a more complete description of
setation is given in Table 1 .

Hunt (1979) did not furnish a key to males
of continental North American Elaphoidel-
la. The males of E. californica, E. carterae
and E. kodiakensis are undescribed. The
male of E. subgracilis is incompletely de-
scribed.

Key to females:

1 . Leg 2 endopodite article 2 with total

of three setae 2

- Leg 2 endopodite article 2 with total

of four or five setae 3

2. Leg 3 endopodite article 2 with three
setae .... amabilis Ishida, new species

- Leg 3 endopodite article 2 with five
setae

shawangunkensis Strayer, 1988 (1989)

3. Leg 3 endopodite article 1 with one
medial seta, article 2 with five or six
setae 4

- Leg 3 endopodite article 1 naked,
article 2 with four setae

carterae Reid, new species

4. Caudal ramus with small or no dor-
sal keel, not ending in hook; hyaline
membranes of somites smooth ... 5

- Caudal ramus with pronounced

VOLUME 106, NUMBER 1

145

dorsal keel, ending in large hook;
hyaline membranes of somites

coarsely toothed

bidens (Schmeil, 1893) s. 1.

5. Leg 3 endopodite article 2 with
six setae, leg 5 exopodite with five
setae 6

- Leg 3 endopodite article 2 with five
setae, leg 5 exopodite with four setae

subgracilis (Willey, 1934)

- Leg 3 endopodite article 2 with five
setae, leg 5 exopodite with five setae

californica Wilson, 1975

6. Caudal ramus 2-3 times longer than
broad, anal somite not expanded
dorsally over caudal ramus, most of
ramus visible in dorsal view 7

- Caudal ramus slightly longer than
broad, anal somite with postero-
dorsal expansions, most of caudal
ramus not visible in dorsal view . . .

wilsonae Hunt, 1979

7. Apex of ramus with three processes,
two of these digitiform and flexible

kodiakensis Wilson, 1975

- Apex of ramus with normal apical
setae only reedi Wilson, 1975

Key to males:

1 . Leg 5 exopodite with four setae; leg
2 endopodite article 2 with three or
four setae 2

- Leg 5 exopodite with three setae;
leg 2 endopodite article 2 with three
setae shawangunkensis

2. Leg 2 endopodite 2 with three
setae 3

- Leg 2 endopodite 2 with four
setae 4

3. Caudal ramus with small distally
directed papilla at posterior Va of
lateral surface subgracilis

- Caudal ramus lacking ornament on
distal Vi of lateral surface . . . amabilis

4. Leg 4 biarticulate, article 1 with
none, article 2 with three setae . . 5

- Leg 4 uniarticulate, with two setae
bidens

5. Caudal ramus tapering distally,

medial and lateral surfaces slightly
expanded in dorsal view .... wilsonae
- Caudal ramus bottle-shaped, me-
dial and lateral surfaces incurv ed in
dorsal view reedi

Acknowledgments

The cordial assistance of many people
contributed to this report. Members of the
Department of Biology, Virginia Pohtech-
nic Institute & State University, Blacks-
burg, conducted an excursion to Mountain
Lake subsequent to the 1990 meeting of the
North American Benthological Society, in
which JWR participated. Mr. William B.
Yeaman of the U.S. National Park Serv'ice
facilitated collections by both of us in Rock
Creek Park, District of Columbia and Mar>-
land, in 1990. Mr. and Mrs. William C.
Warner and Miss Elizabeth Warner guided
and assisted JWR with collections in New
Mexico in 1991. Dr. Horton H. Hobbs, Jr.
provided information on the career of Dr.
Marjorie Estelle Carter. Dr. Michael C. Swift
collected the specimens of Elaphoidella bi-
dens from Maryland and donated them to
the National Museum of Natural History-.
Dr. Harry C. Yeatman contributed a pre-
viously unpublished record of E. bidens.

Literature Cited

Apostolov. A. 1985. Etude sur quelques copepodcs
harpacticoides du genre Elaphoidella Chappuis,
1929 de Bulgarie avec une revision du genre.—
Acta Musei Macedonici Scientianim Natural-
ium 17(7/145):133-163.

Carter. M. E. 1944. Harpacticoid copepods of the
region of Mountain Lake. Virginia (With de-
scription of.\foraria virginiana n. sp.).— Journal
of the Elisha Mitchell Scientific Society 60: 1 58-
166. plates 65-67.

. & J. M. Bradford. 1972. Postembnonic de-
velopment of three species of freshwater har-
pacticoid Copepoda. — Smithsonian Contribu-
tions to Zoology 1 19:1-26.

Chappuis, P. A. 1929. Revision du genre Cantho-
camptus Westwood (Note preliminaire). — Bu-
letinul Societatii de §tiinte din Cluj 4:41-50.

Hamond. R. 1987. Non-marine harpacticoid cope-
pods of Australia. I. Canthocamptidae of the
genus Canthocamptus Westwood s. lat. and Fi-

146

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

bulacamptus, gen. nov., and including the de-
scription of a related new species of Cantho-
camptus from New Caledonia.— Invertebrate
Taxonomy 1:1023-1247.

Hunt, G. W. 1979. Description of Elaphoidella wil-
sonae n. sp. (Canthocamptidae: Copepoda) from
Colorado.— Transactions of the American Mi-
croscopical Society 98:248-253.

Lang, K. 1 948. Monographic der Harpacticiden. Vols.
I, II. Nordiska Bokhandeln, Stockholm, 1648
pp.

Monard, A. 1928. Synopsis universalis generum har-
pacticoidarum.— Zoologische Jahrbiicher, Ab-
teilung fur Systematik, Okologie und Geogra-
phic der Tiere 54:139-176.

Reid, J. W. 1987. Attheyella (Mrazekiella) spinipes,
a new harpacticoid copepod (Crustacea) from
Rock Creek Regional Park, Maryland.- Pro-
ceedings of the Biological Society of Washington
100:694-699.

. 1988. Cyclopoid and harpacticoid copepods

(Crustacea) from Mexico, Guatemala, and Co-
lombia.— Transactions of the American Micro-
scopical Society 107:190-202.

Sars, G. O. 1903-191 1. An account of the Crustacea
of Norway. V. Copepoda Harpacticoida. Bergen
Museum, Bergen, 449 pp., 284 pis.

. 1904. Pacifische Plankton-Crustaceen. (Er-

gebnisse einer Reise nach dem Pacific.
Schauinsland 1896/97.) — Zoologische Jahr-
biicher, Abtheilung fiir Systematik, Geographic
und Biologic der Thiere 19:629-646 + Tafeln
33-38.

Schmeil, O. 1893. Deutschlands freilebende Siis-
swasser-Copepoden. II. Teil: Harpacticidae. Er-

win Nagele Verlag, Stuttgart, 1 00 pp. + Tafeln
I-VIII.

Strayer, D. 1988 (1989). Crustaceans and mites (Ac-
ari) from hyporheic and other underground wa-
ters in southeastern New York.— Stygologia
4:192-207.

Westwood, J. O. 1836. Canthocamptus. —Faning-
ton: British Cyclopaedia of Natural History,
London 2:221.

Willey, A. 1934. Some Laurentian copepods and their
variations.— Transactions of the Royal Cana-
dian Institute 20:77-98 + Plates XIII-XV.

Wilson, M. S. 1956. North American harpacticoid
copepods. 1. Comments on the known fresh-
water species of the Canthocamptidae. 2. Can-
thocamptus oregonensis n. sp. from Oregon and
California.— Transactions of the American Mi-
croscopical Society 75:290-307.

. 1975. North American harpacticoid cope-
pods II. New records and new species of Ela-
phoidella from the United States and Canada. —
Crustaceana 28: 125-1 38. [The article is actually
part 1 1 of a series.]

, & H. C. Yeatman. 1959. Harpacticoida. Pp.

815-861 in W. T. Edmondson, ed.. Ward &
Whipple's Fresh- Water Biology. John Wiley &
Sons, New York.

(JWR) Research Associate, Department
of Invertebrate Zoology, NHB 163, Nation-
al Museum of Natural History, Smithson-
ian Institution, Washington, D.C. 20560,
U.S.A.; (TI) 372 Irifunecho, Yoichimachi,
Hokkaido, 046 Japan.

PROC. BIOL. SOC. WASH.
106(1), 1993, pp. 147-157

NEW GENERA AND SPECIES OF DEEP-SEA

POLYCHAETES OF THE FAMILY NAUTILINIELLIDAE

FROM THE GULF OF MEXICO AND THE

EASTERN PACIFIC

James A. Blake

Abstract.— Three new genera and species of deep-sea polychaetes are de-
scribed. Pilar gis mirasetis Fauchald, 1972 is redescribed and referred to a new
genus Santelma. These species are tentatively referred to the family Nautili-
niellidae that heretofore has included commensals of bivalve molluscs. One
species, Laubierus mucronatus, was dissected from a mussel during its collec-
tion, but the other three species were not associated with bivalves during the
sample sorting process. Two genera and species, Miura spinosa and Santelma
miraseta come from the eastern Pacific, while Laubierus mucronatus and Flas-
carpia alvinae come from cold-seep communities on the Florida Escarpment.
The genera presently assigned to the Nautiliniellidae are heterogeneous and
diverse. These genera are compared with previously described nautiliniellids
and with the closely related family Antonbrunnidae.

The nautiliniellids are a small group of
polychaetes that live as commensals or par-
asites of deep-sea bivalve molluscs. Al-
though only first reported in 1989, six spe-
cies have already been described (Miura &
Laubier 1989, 1990; Blake 1990; Miura &
Ohta 1991), and there are probably many
additional species awaiting discovery. In-
dividual species tend to be small and have
probably been overlooked by biologists ow-
ing to their association with molluscs. The
morphology of the prostomium and the
types of setae have proven to be quite vari-
able and the status of this new group is not
fully understood.

Four additional genera and species of this
family have been discovered from North
America. One species comes from mussels
at cold-seep sites on the Florida Escarpment
and is assigned to a new genus Laubierus.
Another Florida Escarpment species is as-
signed to a new genus Flascarpia. A third
species has been identified from a low oxy-
gen site in the Santa Maria Basin off central
California, and is assigned to a new genus

Miura. A fourth species from deep-water off
Western Mexico and previously described
as Pilar gis mirasetis by Fauchald (1972) is
redescribed and assigned to a new genus
Santelma.

The types of the new species are deposited
in the collections of the National Museum
of Natural History (USNM), Smithsonian
Institution, Washington, D.C.

Systematic Account

Family Nautiliniellidae Miura & Laubier

Diagnosis.— bodies elongate, cylindrical,
and smooth. Prostomium variable, with 1-
2 pairs of antennae, medial antenna or pa-
pilla present or absent, or antennae entirely
absent; eyes absent; palps lacking. Foregut
expanded into a muscular pharynx, termed
a proventriculus (Miura & Laubier 1989,
1 990); this pharynx partially eversible (Blake
1990, this paper). Peristomial segment well
developed with distinct tentacular cirri
present in two genera (Santelma and Flas-
carpia), reduced in three genera {Petrecca,

148

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Laubierus, and Miura), and lacking in three
other genera {Nautiliniella, Shinkai, and
Natsushima). Parapodia subbiramous to ses-
quiramous, with reduced notopodia, usu-
ally with acicula; neuropodia with internal
aciculae and one to several simple hooked
spines. Pygidium simple, lacking append-
ages.

Miura, new genus

Type species.— Miura spinosa, new spe-
cies. Gender feminine.

Diagnosis.— Body, widest anteriorly, ta-
pering posteriorly. Prostomium rounded;
antennae and eyes lacking. Pharynx weakly
muscularized, prominent in first 3 seg-
ments. Peristomial segment achaetous,
smaller than following setigers; with 2 pair
of short lobes present. Parapodia sesquira-
mous, with thick notopodial lobe, acicula
not observed; neuropodium with fascicle of
5-6 simple hooks, with single acicula, some-
times with tip protruding; ventral cirrus
present. Pygidium unknown.

Etymology. —This genus is named for Dr.
Tomoyuki Miura, who was one of the first
describers of the nautiliniellid polychaetes.

Remarks.— T\\\s genus differs from pre-
viously described nautiliniellids by lacking
antennae altogether and in having sesquira-
mous parapodia where the notopodial lobes
are large and thick, but apparently lacking
aciculae. Like Petrecca, Miura has a reduced
and achaetous tentacular segment; like
Shinkai it has several neuropodial spines
instead of one. A single species, Miura spi-
nosa, is known.

Miura spinosa, new species
Fig. 1

Material examined.— Califomia: Santa
Maria Basin, Sta. R-7, 34°52.90'N,
121°10.30'W, May 1989, 565 m, holotype
(USNM 148677).

Description.— A small species, holotype
an anterior fragment measuring 1.5 mm

long, 0.7 mm wide for 10 setigers. Color in
alcohol: light tan with scattered orange pig-
ment spots on dorsal and ventral surfaces.
Body subcylindrical to oval in cross section,
widest through first 6-7 setigers, then ta-
pering posteriorly.

Prostomium twice as wide as long, broad-
ly rounded anteriorly (Fig. lA); antennae
and eyes absent. Mouth with broad ventral
lip (Fig. IB); pharynx weakly muscularized,
prominent in first 3 segments (Fig. lA).
Peristomial segment achaetous, smaller than
following setigers, with 2 pairs of small stub-
by lateral lobes. Parapodia sesquiramous,
with thick notopodial lobe lacking acicula
(Fig. IC); neuropodium with fascicle of 4-
6 simple hooks; each hook sharply pointed,
with subapical notch and boss (Fig. ID);
acicula present, distal end slightly protrud-
ing on some setigers; ventral cirrus thick,
rounded lobe. Pygidium unknown.

Biology. —Miura spinosa was collected at
a depth of 565 m in a sea valley in the Santa
Maria Basin that has very fine silty sedi-
ments with high clay content. This depth is
also characterized by low dissolved oxygen
concentrations (X = 0.898 ml/1, SD =
±0.152; n = 4) in the near-bottom water.
It is not known if hydrogen sulphide or
methane is also present at this site. This
specimen was not associated with a bivalve
during the sorting process, but may have
been washed from a host animal during pro-
cessing.

Etymology. —The specific name refers to
the neuropodial spines.

Remarks.— The shape of the setae of
Miura spinosa is similar to Pilargis mira-
setis, with which this species was identified
in the MMS monitoring program where it
was collected. Pilargis mirasetis has also
been found to be a species of Nautilinielli-
dae and has been referred to a new genus,
Santelma (see below). Santelma miraseta
differs from Miura spinosa in having short
notopodial lobes instead of ones that are
large and thick, in having long protruding

VOLUME 106, NUMBER 1

149

300nm

lO^m

lOOjim

Fig. 1. Miura spinosa (USNM 148677). A, anterior end, dorsal view; B, anterior end ventral view; C, tenth
parapodium; D, neurosetae.

aciculae instead of lacking these spines, and
in having two distinct tentacular cirri in-
stead of two pair of short, stubby lobes.

Distribution.— CdMiomidi, upper conti-
nental slope, 565 m.

Genus Santelma, new genus

Type species.— Santelma miraseta (Fau-
chald, 1972). Gender feminine.

Diagnosis. —Body dorsoventrally flat-
tened. Prostomium rounded anteriorly,
bearing 2 lateral and 1 medial antenna or

papilla (scar only present on S. miraseta);
eyes and palps absent. Pharynx enlarged,
muscular. Peristomial segment achaetous,
bearing 2 pair of tentacular cirri. Parapodia
sesquiramous, with prominent notopodial
lobe bearing large internal acicula; neuro-
podium with single large acicular protrud-
ing acicula and fascicle of numerous, small,
simple setae; dorsal and ventral cirri absent;
branchiae absent. Pygidium a simple
rounded lobe.

Etymology.— The name for this genus is
coined from the town of San Telmo, Mex-

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Santelma miraseta (LACM-AHF 097). A, anterior end, dorsal view; B, eighth parapodium; C, middle
parapodium; D, neurosetae from an anterior setiger.

ico, which is the closest point of land near
the collecting site of Santelma miraseta.

Remarks. —The presence of two antennae
and a third median antenna or papilla dis-
tinguishes Santelma from other genera in
the nautiliniellid complex. The four small
tentacular cirri and large, protruding neu-
ropodial acicula are also unusual.

Santelma miraseta (Fauchald, 1972),

new combination

Fig. 2

Pilargis mirasetis Fauchald, 1 972:59-60, pi.
8, figs. a-c.-Salazar-Vallejo, 1986:200,
pi. 2, figs. 9-10.

Material examined. —Off San Telmo,

VOLUME 106, NUMBER 1

151

Mexico, Velero IV Sta. 13744-70, 15 Jan
1970, 18°12'00"N, 104°00'00"W, 2340 m,
holotype (LACM-AHF 097).

Description. —Hololype complete, only
known specimen, 25 mm long, 2 mm wide,
with about 1 10 segments. Color in alcohol:
tan. Body generally robust, dorsoventrally
compressed, elongate.

Prostomium wider than long, broadly
rounded on anterior margin; with 2 lateral
antennae (bases only) and single median an-
tenna or papilla (scar only), all located near
border of prostomium and peristomial seg-
ment (Fig. 2A); palps and eyes absent. Phar-
ynx muscular, not everted. Peristomial seg-
ment slightly larger than following segment,
bearing 2 pair of short, tapering tentacular
cirri (Fig. 2A).

Parapodia sesquiramous, with notopo-
dium formed into thickened lobe, distally
prolonged in middle and posterior seg-
ments; notopodia bearing large internal
acicula, typically bending sharply near tip
(Fig. 2B); dorsal cirrus absent. Neuropo-
dium with large protruding acicula and fas-
cicle of 18-25 simple bidentate setae (Fig.
2B); ventral cirrus absent. Setae bidentate,
tapering apically to fine, mucronate tip; sub-
apical tooth blunt (Fig. 2D); bidentate setae
mostly lost in middle and posterior para-
podia, leaving only protruding spine (Fig.
2C). Pygidium simple, rounded lobe; with-
out cirri.

Remarks. — Santelma miraseta is here re-
moved from the Pilargidae because palps
are absent. However, the referral of this ge-
nus and species to the Nautiliniellidae is
preliminary pending further review of the
family. The presence of a medial antenna
or papilla is new to species of the nautili-
niellid-complex of genera. The very large
and conspicuous protruding neuropodial
acicula was not mentioned by Fauchald
(1972). This acicula accompanies a cluster
of very minute bidentate setae in anterior
setigers. In posterior setigers, the bidentate
setae are absent and the protruding acicula
is the only visible seta and provides this
species with a distinct posterior armature.

Two types of emergent neurosetae have also
been reported for Natsushima and Laubier-
us, but these genera are very distinct from
Santelma (see Discussion for overall sum-
mary).

Laubierus, new genus

Type species.— Laubierus mucronatus,
new species. Gender masculine.

Diagnosis. —Body dorsoventrally flat-
tened, ribbonlike. Prostomium rounded an-
teriorly, bearing a single pair of antennae;
eyes lacking. Pharynx muscularized, form-
ing distinctive proventriculus. Achaetous
peristomial segment present, bearing pair of
short cirri homologous to normal ventral
cirrus. Parapodia sesquiramous, with acic-
ulae in both noto- and neuropodia; setal
fascicles limited to neuropodia; including
large and small types of simple setae. Ven-
tral cirrus present. Pygidium a simple lobe.

Etymology. —This genus is named for Dr.
Lucien Laubier, polychaete systematist, in
recognition of his first descriptions of nau-
tiliniellids.

Remarks.— The appearance of the phar-
ynx resembles the proventriculus that has
been reported for the Japanese genera and
species. The presence of large and small
simple spines in the neuropodia of Laubier-
us is similar to that of the genus Natsushima
Miura & Laubier. In comparing these gen-
era, Laubierus has an achaetous peristomial
segment, large notopodial acicula, and small
simple neurosetae, whereas Natsushima has
no peristomial segment, thin notopodial
acicula, and small bifid neurosetae.

Laubierus mucronatus, new species
Fig. 3

Material examined. —¥\or\d.di Escarp-
ment, Alvin Dive 1756, 26°01'N, 84°55'W,
3243 m, 17 Oct 1986, R. Lutz and G. Tien,
observers, dissected from mussel, 1 frag-
ment (JAB); Alvin Dive 1758, 26°01.8'N,
84°54.9'W, 3266 m, Oct 1986, C Wirsen
and B. Tilbrook, observers, holotype
(USNM 148678).

152

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

300jim

W/ V-f I

1 10^

. ,.,,^. iv..v. .! .; .l., >i !? T g?r7?''??S:^X:

T" .- ■•'^y.\\■^i'^r\\'■v^iiy^y}|fi^ ^^■'\'■■: ■■"■ • •• ■.11-'---'" .'--:^.

Fig. 3. Laubierus mucronatus (USNM 148678). A, anterior end, dorsal view; B, middle parapodium; C,
small neurosetae; D, large neuroseta.

VOLUME 106, NUMBER 1

153

Description. — A small species, holotype
complete, 5 mm long and 0.4 mm wide for
34 setigerous segments; fragment larger, but
anteriorly incomplete. Color in alcohol:
opaque white. Body elongate, dorsoven-
trally flattened, tapering gradually anteri-
orly and posteriorly.

Prostomium wider than long, rounded
along anterior margin, bearing single pair of
sublateral antennae (Fig. 3A); tentacular
segment poorly developed, only vaguely
separated from prostomium, bearing pair of
short cirri. Anterior part of digestive tract
modified into proventriculus, with distinct
musculature apparent through body wall
(Fig. 3A).

Parapodia sesquiramous, with notopodia
bearing only a single, large acicula, with tip
sometimes protruding; notopodium elon-
gate, tapering, extended apically into nar-
row lobe (Fig. 3B). Neuropodia longer,
broader than notopodium, with fingerlike
ventral cirrus; bearing large internal acicula
and 2 types of simple neurosetae; 1 type
small, numbering 1 5-20 in 2 rows; each with
fringed tip from which tapering mucronate
tip emerges (Fig. 3C); second type 1-2 large,
falcate hooks, with each bearing delicate
subapical fringe of fine bristles (Fig. 3D).
Pygidium a simple lobe, lacking cirri.

Etymology.— Tht specific name comes
from the latin, mucros, referring to the fine,
tapering point of the smaller neurosetae that
characterize this species.

Genus Flascarpia, new genus

Type species.— Flascarpia alvinae, new
species. Gender feminine.

Diagnosis.— Body elongate, compressed,
ventrum flattened, dorsum rounded. Pro-
stomium with broadly rounded anterior
margin, lacking eyes, with 2 lateral anten-
nae. Distinct peristomial segment, bearing
a pair of tentacular cirri. Pharynx soft, ever-
sible. Parapodia sesquiramous, with noto-
podial lobe containing blood loop and ap-
parently functioning as gill; acicula absent.

Neuropodium well-developed, with large
acicula and fascicle of large simple neuro-
setae of 1 type; ventral cirrus present. Py-
gidium a simple lobe, lacking cirri.

Etymology. —The generic name is a com-
posite of Florida and Escarpment, denoting
the Florida Escarpment where the type spe-
cies was collected.

Remarks. —The notopodium lacks all se-
tae, including the acicula, and by having a
blood loop, apparently functions as a gill.
Flascarpia and Miura are the only nautili-
niellid-like genera known to lack notoacicu-
lae and to have the notopodium modified
into a soft, fleshy lobe. In Flascarpia, a dis-
tinct blood loop is present, suggesting that
it functions as a gill. In Miura, no blood
loop was observed. The relationships of
Flascarpia with all of the nautiliniellid-like
genera are compared in the Discussion (see
below).

Flascarpia alvinae, new species
Fig. 4

Material examined. —Florida Escarp-
ment, Alvin Dive 1754, 26°02.4'N,
84°55.3'W, 3303 m, 15 Oct 1986, R. Carney
and B. Hecker, observers, holotype (USNM
148679).

Description.— A moderate-sized species,
holotype complete with 66 segments, mea-
suring 2 1 mm long; 1 mm wide anteriorly,
2 mm wide in middle. Body widest in mid-
dle of body, tapering anteriorly and poste-
riorly; body flattened ventrally, rounded
dorsally. Pygidium a simple lobe lacking
cirri. Color in alcohol: tan.

Prostomium wider than long, broadly
rounded on anterior margin; with 2 short
antennae on lateral anterior margins of pro-
stomium (Fig. 4A). Peristomial segment
present, not distinctly separated from pro-
stomium, bearing a single pair of long, fin-
gerlike tentacular cirri (Fig. 4A). Parapodia
sesquiramous, with notopodium reduced to
soft lobe bearing internal blood vessel (Fig.
4B); acicula absent. Neuropodium elongate,

154

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

lOO^m

Fig. 4. Flascarpia alvinae (USNM 148679). A, anterior end, dorsal view (broken line indicates outline of
everted pharynx); B, middle parapodium; C, D, neurosetae.

thickened, triangular, tapering to point;
ventral cirrus short, cirriform. A single large
internal acicula present and fascicle of 7-8
large protruding simple spines; each with
curved tip and 2 thick subterminal protu-
berances (Fig. 4C-D). Pygidum a simple
lobe.

Etymology.— TYiis species is named for
the DSRV Alvin, which was the vehicle used
to collect specimens.

Remarks.— The habitat of Flascarpia al-
vinae is not known. It is likely, however,
that this species is a commensal of bivalves
and was washed from its host during the
sorting process.

Discussion

The first published description of a nau-
tiliniellid was by Miura & Laubier (1989),
who described Nautilina calyptogenicola
from a deep-sea vesicomyid clam collected
from the Japan Trench at a depth of 5960
m. This species was characterized by having
two pairs of small antennae, short dorsal
and ventral cirri, and a single, large pro-
truding neuropodial hooked spine. The au-
thors assigned this new genus to a new fam-
ily, the Nautilinidae. In a subsequent paper,
the same authors renamed the genus and
family Nautiliniella and Nautiliniellidae,

VOLUME 106, NUMBER 1

155

because the earlier names were preoccupied
in the Cephalopoda (Miura & Laubier 1 990).
In this same paper, the authors described
two additional new genera and species:
Shinkai sagamiensis Miura & Laubier from
the bivalve, Calyptogena soyoae; and Na-
tsushima bifurcata Miura & Laubier from
Solemya sp. Both of these latter species were
collected at the Hatsushima cold-seep site
off Japan in depths of 1130-1170 m. The
genus Shinkai was superficially similar to
Nautiliniella, but differed in having up to
eight hooks per neuropodium instead of one
and in having only a single pair of antennae
instead of two. Natsushima differed from
the other two genera in having two types of
neuropodial spines instead of one. Blake
(1 990) described another genus and species,
Petrecca thyasira, from the mantle cavity of
a thyasirid clam, Thyasira insignis collected
from seep-like communities off Newfound-
land at a depth of 3700 m. Petrecca differed
from the previously described genera in
having greatly elongated notopodia and an
achaetous peristomial segment. Miura &
Ohta (1991) described Shinkai longipeda
from the mantle cavity of Calyptogena sp.
collected at active hydrothermal vents in
the Okinawa Trough in 1400 m. This spe-
cies appears to be transitional between the
genera Shinkai and Petrecca in the devel-
opment of the notopodia.

The new taxa described in the present pa-
per do not agree with any of the previously
described genera. All have an achaetous
peristomial segment with either small or
well-developed tentacular cirri. Miura lacks
antennae, has a thickened achaetous noto-
podial lobe, and a fascicle of small neuro-
podial spines. Laubierus has two small an-
tennae, a long notopodial lobe with a large
acicula, and both large and small types of
neuropodial spines. Flascarpia has two an-
tennae, two well-developed tentacular cirri,
a reduced notopodium that may function as
a gill, and a fascicle of heavy neuropodial
spines. Santelma has three antennae, two

pairs of tentacular cirri, a single large pro-
truding acicula and a fascicle of very fine
bidentate setae.

The Nautiliniellidae now include eight
genera, the characters of which are com-
pared and contrasted in Table 1 . The group
is very heterogeneous and it is likely that
the genera will need to be redefined and
reorganized after additional species are dis-
covered. In general, two distinct groups are
apparent. One group lacks a peristomial seg-
ment and includes Nautiliniella, Shinkai,
and Natsushima. The second group has a
peristomial or achaetous segment and in-
cludes Petrecca, Miura, Laubierus, Flascar-
pia, and Santelma. Among the second group,
the "peristomial segment" oi Petrecca is ac-
tually a reduced setigerous segment that has
a ventral cirrus and neuroacicula, but en-
tirely lacks a notopodium. In Laubierus, the
"peristomial segment" is also a reduced seg-
ment that bears a ventral cirrus, but no acic-
ula. Thus, the presence or absence of a peri-
stomial segment appears to depend upon
the degree to which the first setiger is re-
duced and this undoubtedly contributes to
the variability in this character that is ex-
hibited by the different genera that have been
described. All genera are characterized by
having simple neuropodial spines, and ex-
cept for Santelma, these setae are usually
heavier and of a different form than those
of the closely related family Pilargidae.

All of these new species are tentatively
assigned to the Family Nautiliniellidae based
on absence of notosetae, presence of simple
spinous neurosetae, absence of anal cirri,
and lack of palps. None of these species
agrees with any genus of the Pilargiidae al-
though they appear to be closely related.
The nautiliniellids are also related to An-
tonbruunia viridis Hartman & Boss (1965)
from a bivalve dredged in the Mozambique
Channel off Madagascar in 80-90 m. This
genus and species was assigned to a separate
family, the Antonbruunidae by Fauchald
(1977) and to the Pilargidae by Salazar-Va-

156

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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VOLUME 106, NUMBER 1

157

llejo (1986). The characteristics of Anton-
bruunia are compared with those of the var-
ious nautilinieUids in Table 1. The setal
characteristics of A. viridis are more similar
to those of the Nautiliniellidae than to the
Pilargidae (See also Miura & Laubier 1990)
and I prefer to follow Fauchald (1977) in
placing Antonbrunnia in its own family.

All members of the Nautiliniellidae are
believed to be associated with the mantle
cavities of deep-sea bivalve molluscs. There
is insufficient data to determine if the worms
are commensals or parasites with the clams.
Four nautiliniellid species are associated
with bivalves at cold-seep communities
(Miura & Laubier 1989, 1990; Blake 1990,
this paper), while a fifth comes from a bi-
valve at an active hydrothermal vent (Miura
& Ohta 1 99 1). The Santa Maria Basin spec-
imen was found in an upper slope sea valley
at a depth of 565 m in an area of low dis-
solved oxygen. The specimen was not as-
sociated with a bivalve when it was sorted
from the mud, but may have been washed
from a bivalve in the same sample. The
exact habitats of Santelma miraseta and
Flascarpia alvinae are not known. Infesta-
tion rates of nautilinieUids in bivalves may
be fairly high when the worms are present.
For example, out of ten specimens of Thy-
asira insignis collected from off Newfound-
land, five were found with a specimen of
Petrecca thyasira (Blake 1990). These clams
were each relatively small, ranging from 29
X 30 cm to 37 x 43 cm in width x length.
The worms themselves were up to 16 mm
long and thus occupied a considerable space
among the gill filaments of the clams. In-
festation data is not available for other spe-
cies.

Acknowledgments

This study was funded in part by the Pa-
cific Outer Continental Shelf Region of the
Minerals Management Service (MMS), U.S.
Department of the Interior, Washington,
D.C. under Contract No. 14-35-0001-
30484. The specimen of Miura spinosa was

collected as part of the MMS Phase II Mon-
itoring Program in the Santa Maria Basin.
The specimens of Laubierus mucronatus and
Flascarpia alvinae were provided by Dr.
Barbara Hecker, Lamont-Doherty Geolog-
ical Observatory. The holotype of Pilar gis
mirasetis was provided by Dr. Kirk Fitz-
hugh of the Los Angeles County Museum
of Natural History (LACM-AHF). The
manuscript was greatly improved by in-
sightful comments provided by Dr. Chris
Glasby and an anonymous reviewer.

Literature Cited

Blake, J. A. 1990. A new genus and species of Poly-
chaeta commensal with a deep-sea thyasirid
clam. — Proceedings of the Biological Society of
Washington 103:681-686.

Fauchald, K. 1972. Benthic polychaetous annelids
from deep water off western Mexico and adja-
cent areas in the eastern Pacific Ocean.— Allan
Hancock Monographs in Marine Biology No.
7:1-575, 69 pis.

. 1977. The polychaete worms. Definitions and

keys to orders, families and genera.— Natural
History Museum of Los Angeles County, Sci-
ence Series 28:1-188.

Hartman, O., & K. J. Boss. 1965. Antonbrunnia viri-
dis, a new inquiline annelid with dwarf males,
inhabiting a new species of pelecypod, Lucina
fosteri, in the Mozambique Channel.— Annals
& Magazine of Natural History, series 13,8:177-
186.

Miura, T., & L. Laubier. 1989. Nautilina calypto-
genicola, a new genus and species of parasitic
polychaete on a vesicomyid bivalve from the
Japan Trench, representing a new family Nau-
tilinidae.— Zoological Science 6:387-390.

, & . 1990. Nautiliniellid polychaetes

collected from the Hatsushima cold-seep site in
Sagami Bay, with descriptions of new genera
and species.— Zoological Science 7:319-325.

, & S. Ohta. 1991. Two polychaete species

from the deep-sea hydrothermal vent in the
Middle Okinawa Trough.— Zoological Science
8:383-387.

Salazar-Vallejo, S. I. 1986. Pilargidae (Annelida:
Polychaeta) de Mexico: listade especies, nueva
especie y biografia. — Cahiers de Biologic Marine
27:193-209.

Science Applications International Cor-
poration, 89 Water Street, Woods Hole,
Massachusetts 02543, U.S.A.

PROC. BIOL. SOC. WASH.
106(1), 1993, pp. 158-181

TAXONOMY OF EUROPEAN SPECIES OF

AMPHIDUROS AND GYPTIS

(POLYCHAETA: HESIONIDAE)

Fredrik Pleijel

Abstract.— ThQ taxonomy of the European species of Amphiduros Hartman,
1959, and Gyptis Marion & Bobretzky, 1875, is reviewed. Amphiduros is re-
corded for the first time from European waters. Amphiduros fuscescens (Maren-
zeller, 1875), new combination (originally described in Oxydromus), G. pro-
pinqua Marion & Bobretzky, 1875, and G. rosea (Malm, 1874) are redescribed
based on available types and newly collected specimens from the vicinity of
the type localities, and A. fuscescens is removed from synonymy with G. pro-
pinqua. Two new species are described: G. mackiei from Sweden and the Faroes,
unique within the genus in having setae from segment 4 rather than segment
5, and G. mediterranea from southern France and Sicily, a species close to G.
rosea. A key to the European species is included, and a checklist for species
and subspecies described in or later referred to the two genera is appended.

The genus Amphiduros Hartman, 1959,
is only known for four species, all described
from the Pacific. Examination of type and
newly collected Mediterranean specimens
of Oxydromus fuscescens Marenzeller, 1875,
indicates that this species also belongs to
Amphiduros, and constitutes the first record
of the genus from Europe. Previously O.
fuscescens has been treated as a junior syn-
onym of G. propinqua Marion & Bobretzky,
1875 (Fauvel 1923, Hartman 1959).

Two European species of Gyptis Marion
& Bobretzky, 1875 sensu stricto (excluding
Podarkeopsis Laubier, 1961; see below) are
recorded in the literature from this century:
the type species G. propinqua Marion &
Bobretzky, 1875, and G. rosea (Malm,
1874). The status of these two species is
currently uncertain. For example, Eliason
(1962) treated G. propinqua as a doubtful
junior synonym of G. rosea, and Hartmann-
Schroder (1971) referred Fauvel's (1923)
description of G. propinqua to G. rosea.
Haaland & Schram (1982) also treated G.
propinqua as a possible junior synonym, but
remarked that the larvae from the Oslofjord

differed from those described as G. propin-
qua by Bhaud (1971).

I believe that these synonymies are in-
correct, and that the number of species oc-
curring in the area is underestimated. This
study aims to describe all the European spe-
cies of Gyptis (including two new species)
and the closely related Amphiduros, to re-
solve their synonymies and to provide a key
facilitating their correct identification. Apart
from the interpretation of tentacular cirri
(see below), the generic delineations follow
traditional ones (e.g., Fauchald 1977) and
no phylogenetic considerations are made at
this point; these will have to await future
studies treating the relationships within
family (Pleijel, in prep.). The study is based
on museum specimens as well as newly col-
lected material during trips to the northern
part of the Swedish west coast, northwestern
Iceland, eastern Sicily, and Banyuls-sur-Mer
in southern France. A checklist for species
originally or later referred to the two genera
is appended. The list also serves as a record
of other congeneric species examined in the
course of this study.

VOLUME 106, NUMBER 1

159

Species belonging to Podarkeopsis are ex-
cluded. These are distinguished from Gyptis
and Amphiduros by the presence of an an-
teriorly inserted median antenna (rather than
medially on the dorsal surface of the pro-
stomium), by ten terminal proboscideal pa-
pillae (rather than absence or a larger num-
ber), and by the presence of furcate notosetae
(rather than absence). Species recorded from
European waters which I consider belong to
the Podarkeopsis -group are: P. arenicolus
(La Greca, 1946), described from the Gulf
of Naples, Italy; P. galangaui Laubier, 1961,
described from Banyuls-sur-Mer, southern
France; P. capensis (Day, 1963), described
from South Africa but recorded from south-
em England (Gibbs & Probert 1973) and
the Tyrrhenian Sea, Italy (Gravina and
Giangrande 1988); and Gyptis helgolandica
Hilbig & Dittmer, 1979, described from
Helgoland in the North Sea.

In hesionids the number of tentacular cir-
ri has generally been considered diagnostic
at the generic level (e.g., Fauchald 1977).
Gyptis and Amphiduros are described as
having eight pairs of tentacular cirri, but the
character "number of tentacular cirri"
probably conceals a mixture of several char-
acters. In the literature the tentacular cirri
appear to be defined by absence of para-
podia, i.e. if parapodial lobes are absent they
are named tentacular cirri, otherwise dorsal
and ventral cirri. However, the tentacular
cirri also differ from cirri in "normal" seg-
ments in the usually more pronounced cir-
rophores, and in being stouter and longer.
Further, many adult hesionids have dorsal
cirri on the first setigerous segment that are
of the same shape as the preceding dorsal
tentacular cirri. In considering these am-
biguities I prefer to use the absence of no-
topodia on segment 4 to define Gyptis and
Amphiduros and presently avoid the char-
acter "number of tentacular cirri." Gyptis
mackiei (which has setigerous neuropodia
on segment 4 and thus may be interpreted
as having six pairs of tentacular cirri) will
then also be included in Gyptis. Possible

reassignement will have to await future phy-
logenetic analyses.

Materials and Methods

The collected specimens were relaxed, ei-
ther with menthol or magnesium chloride
(7% in distilled water), studied alive, pre-
served in formalin (5-10% in seawater) for
a few days, rinsed in fresh water and trans-
ferred to 80% alcohol. For SEM specimens
were similarly relaxed, preserved for a few
hours in osmium tetraoxide (1% in artificial
seawater), rinsed in distilled water, trans-
ferred to 80% alcohol in a graded series, and
subsequently critical-point dried. All draw-
ings were made with a camera lucida; those
of Figs. lA, B, K, 4A-C, and 9A, B from
live, relaxed specimens, and remaining ones
from preserved specimens.

All measurements were carried out either
on live, relaxed specimens or specimens re-
laxed prior to preservation. Width mea-
surements were taken from median seg-
ments and include parapodia but exclude
cirri and setae.

Institutions and museums are indicated
by the following abbreviations: BIOFAR
(Marine benthic fauna of the Faroe Islands,
Kaldbak), BMNH (The Natural History
Museum, London), LACM (Los Angeles
County Museum of Natural History),
NHMG (G5teborg Natural History Muse-
um), NHMR (Natural History^ Museum,
Reykjavik), NHMW (Naturhistorisches
Museum Wien), NMCA (National Museum
of Canada, Ottawa), NMW (National Mu-
seum of Wales, Cardiff), SMNH (Swedish
Museum of Natural History, Stockholm),
USNM (National Museum of Natural His-
tory, Smithsonian Institution, Washington,
D.C.), ZMB (Universitat Humboldt, Mu-
seum fur Naturkunde, Berlin). ZMH (Uni-
versitat Hamburg, Zoologisches Institut und
Museum), ZMUU (Uppsala Universitet,
Zoologiska Museet). All material is depos-
ited at SMNH unless otherwise indicated.

160

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1 . Gyptis propinqua. A. Anterior end, dorsal view. Setae omitted. B. Anterior end, ventral view. Setae
omitted. C. Parapodium segment five, posterior view. D. Parapodium segment six, anterior view, ca. half number
of setae shown. E. Median parapodium, anterior view, ca. half number of setae shown. F. Acicular notoseta. G.
Spiked capillary notoseta. H. Serrated capillary notoseta. I. Median neuroseta. J. Ventral neuroseta. Specimens
from Koster area, Sweden. Scales A-E, K, 0.25 mm; F-J, 50 nm.

VOLUME 106, NUMBER 1

161

Gyptis Marion & Bobretzky, 1875
Gyptis Marion & Bobretzky, 1875:50-51.

Type species. — Gyptis propinqua Marion
& Bobretzky, 1875, either by monotypy or
subsequent designation (see Remarks).

Diagnosis (provisional).— }:!iQsiomds with
two palps and two frontal antennae. Median
antenna present, inserted dorsally on pro-
stomium. Proboscis with terminal ring of
papillae. Jaws absent. Segment 4 without
notopodia or notosetae, with or without
neuropodia with neurosetae. Segment 5 with
or without notopodia or notosetae, with
neuropodia and neurosetae. Following
parapodia with well-developed noto- and
neuropodia. Notosetae include one to sev-
eral acicular and large number of capillary
setae; furcate setae absent. Neurosetae nu-
merous, all usually compound.

Remarks.— \X is not obvious from Mari-
on & Bobretzky' s original description that
G. propinqua should constitute the type spe-
cies of the genus, but the matter is probably
without practical significance because it has
been designated subsequently, if not earlier
then at least by Hartman (1965).

The name Oxydromus Grube, 1855 has
variously been treated as confused (Marion
& Bobretzky 1875), as a senior synonym to
Gyptis (e.g., Fauvel 1923, Hartman 1959)
or, more recently (Hartman 1965), as pre-
occupied (in Aves by Oxydromus Schlegel,
1854). However, examination of the type
material for the type species for the genus,
O.fasciatusGrubQ, 1855 (ZMB 3825), shows
it to be a junior synonym to Ophiodromus
flexuosus (delle Chiaje, 1827). Apart from
possibly being a junior homonym the name
is thus also a synonym oi Ophiodromus Sars,
1862, and of no relevance to Gyptis.

Pending further investigations (see In-
troduction) the generic diagnosis is provi-
sionally emended to include G. mackiei, a
new species with neurosetae present on seg-
ment 4.

Gyptis propinqua Marion & Bobretzky, 1875
Figs. 1-3, 12

Gyptis propinqua Marion & Bobretzky,

1875:51-54, pis. 5-6, fig. 15.
Oxydromus propinquus. — Fauvel, 1923:

241-242, fig. 90. [Not Oxydromus pro-

pinquus sensu Ushakov, 1955:196-197,

fig. 58]
Gyptis r(95^a. — Hartmann-Schroder, 1971:

132-134, fig. 42.-Helgason et al., 1990:

205.

Material examined. -IcQland: ca. 200
specimens (NHMR), Breidafiodur, 65°05'N,
23°17'W, 43 m, van Veen grab and Agassiz
trawl, shell gravel, 9 Aug 1979; 3 specimens
(NHMR), Breidafjordur, 65°05'N, 23°16'W,
53 m, van Veen grab, shell gravel, 9 Aug
1979; 15 specimens, Breidaflordur, Selsker,
65°05.03'N, 23°16.91'W, 43 m, dredge,
coarse shell gravel, 16 Jul 1991. Sweden: 7
specimens, Vaderoarna, SE Norra Ragstu-
ten, 58°32.2'N, 1 1°5.0'E, 12 m, dredge, shell
sand and gravel, 7 Oct 1984; 1 specimen,
Koster area, SW Yttre Vattenholmen,
58°52.5'N, 11°06.3'E, 50-100 m, dredge,
mixed sediments, 7 January 1985; 1 spec-
imen, Koster area, W Yttre Vattenholmen,
58°52.4'N, 1 1°06.5'E, 20-30 m, dredge, shell
sand, 29 Aug 1985; 4 specimens, Koster
area, SW Yttre Vattenholmen, 58°52.5'N,
11°06.3'E, 30 m, dredge, shell sand and
gravel, 30 May 1989; 7 specimens, Koster
area, E Yttre Vattenholmen, 58°52.6'N,
11°06.7'E, 20-30 m, dredge, coarse gravel,
29 Aug 1989; 16 specimens, Koster area, W
Yttre Vattenholmen, 58°52.4'N, 11°06.5'E,
20-40 m, dredge, shell sand and gravel, 1
Sep 1989; 1 specimen, Koster area, Koster-
grund, 58°52.5'N, ll°05.rE, 20-40 m,
dredge, mixed sediments, 11 Apr 1990; 4
specimens (all mounted for SEM), Koster
area, W Yttre Vattenholmen, 58°52.4'N,
1 r06.5'E, 30-60 m, dredge, shell sand and
gravel, 13 Apr 1990; 2 specimens, Koster
area, SW Yttre Vattenholmen, 58°52.5'N,
1 1°06.3'E, 40 m, dredge, mixed sediments,
28 Jun 1 990; 2 specimens, Koster area, Kos-
tergrund, 58°52.5'N, ir05.rE, 15 m,
dredge, sand and gravel, 18 Oct 1990; 3
specimens, Koster area, SW Yttre Vatten-

162

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

holmen, 58°52.5'N, 11°06.3'E, 30-50 m,
dredge, 6 Jun 1991; 4 specimens, Koster
area, E Krugglo, 58°53.4'N, ir05.9'E, 10-
20 m, dredge, coarse shell gravel, 13 Jun
1991; 7 specimens, Koster area, E Yttre
Vattenholmen, 58°52.6'N, 1 r06.7'E, 30 m,
dredge, shell gravel, 18 Jun 1991. Faroes:
1 specimen (BIOFAR), 61°54.78'N,
06°28.84'W, 77 m, epibenthic sledge, 20 Jul
1987; 4 specimens, Torshavn, Hognabodi,
62°05'N, 06°33'W, 20-40 m, dredge, 21 Nov
1988. Italy: 1 specimen, Sicily, Acitrezza,
37°34.4'N, 15°11.7'E, 41 m, SCUBA, 17
May 1990; Sicily, Brucoli, 37°16.8'N,
15°11.7'E, 10 m, SCUBA, 21 May 1990.
France: ca. 75 specimens (5 specimens
mounted for SEM), Banyuls-sur-Mer, Cap
d'Osne, 42°29.48'N, 03°08.29'E, 24 m,
dredge, silty coarse sand with shell gravel,

9 Oct 1991; 4 specimens, Banyuls-sur-Mer,
42°29.48'N, 03°08.29'E, 18 m, dredge, sand
and shell gravel, 13 Oct 1991; 3 specimens,
Banyuls-sur-Mer, lie Grosse, 42°29.0'N,
03°08.1'E, 10 m, SCUBA, gravel, 13 Oct
1991; 1 specimen, Banyuls-sur-Mer, Cap
Oullestrell, 42°30.13'N, 03°08.18'E, 18 m,
dredge, 14 Oct 1991; 1 specimen, Banyuls-
sur-Mer, He Grosse, 42°29.0'N, 03°08.rE,

10 m, SCUBA, gravel, 18 Oct 1991.
Description. —Body, excluding parapo-

dia, cylindrical, posteriorly tapered; venter
flattened, without distinct median longitu-
dinal furrow. Median parapodia only slight-
ly longer than anterior, posterior ones suc-
cessively shorter, yielding outline of fairly
equal width with tapering posterior end.

Prostomium about as wide as long, an-
teriorly straight, laterally and posteriorly
with rounded lobes separated by deep pos-
terior incision (Figs. lA, 2 A, B). Proximal
parts of palps cylindrical; distal parts widest
medially, anteriorly rounded (Fig. IB).
Proximal and distal parts of equal length.
Paired antennae situated on small cerato-
phores, longer and thinner than palps, with
pointed tips. Median antenna club-shaped,
widest subdistally, inserted half-way be-
tween anterior pair of eyes and anterior

margin of prostomium on small, rather in-
distinct ceratophore. Anterior pair of eyes
rounded to reniform, twice as large as pos-
terior pair and situated further apart; pos-
terior pair rounded, both pairs with lenses.
Nuchal organs well-developed, lateral to
prostomium but extending and almost co-
alescing mid-dorsally (Fig. 2 A).

Large, distinct lip glands present (Figs.
IB, 2C, E). Proboscis smooth, cylindrical,
sometimes with enlarged opening in pre-
served, unrelaxed specimens, divided in
proximal and distal parts (not obvious on
specimens with strongly extended probos-
cides), distal part smaller. Number of ter-
minal papillae size-dependent (Fig. 12),
usually numbering 35-50. Papillae long (ca.
125 nm) and thin with ciliated tips, arranged
in single ring.

Tentacular cirri distinctly annulated, rings
basally about two to three times as long as
wide, distally usually shorter, tips rounded.
Dorsal tentacular cirri of segment 2 longest,
reaching to about segment 10-12; ventral
tentacular cirri of segment 3 shortest, reach-
ing to about segment 6. Cirri of segments 3
and 4 more ventrally displaced than ante-
rior ones (Fig. 2D). Aciculae present in cir-
rophores of all tentacular cirri, two to three
in dorsal ones and one in ventral ones. An-
terior dorsal segmental delineations not dis-
tinct, several segments fused or reduced.
Segment 4 with first dorsally fully distin-
guishable segment, often forming elevated
ridge anteriorly (Fig. 2A).

Segment 5 (setiger 1) with from one to
three notoaciculae within cirrophores of
dorsal cirri, without setigerous lobes or se-
tae (Fig. IC). Dorsal cirri similar to those
of segment 4. Neuropodia similar to follow-
ing ones but slightly smaller. Ventral cirri
similar to following ones. Segment 6 similar
to median ones but slightly smaller (Fig.
ID).

Dorsal cirri of median segments distinctly
annulated with about 10-15 rings; rings from
one to three times as long as wide (Fig. IE).
Dorsal cirri longer than setae, slightly dif-

VOLUME 106, NUMBER 1

163

prjFj

IbRl'' ^^^HI^^^S

-»^a

Fig. 2. SEM micrographs of Gyptis propinqua. A. Anterior end, dorsal view. B. Prostomium, dorsal view.
C. Anterior end, ventral view. D. Anterior end, right side. E. Lip glands. F. Median parapodium, right side,
antero-ventral view. A & F specimens from Koster area, Sweden, B-D specimens from Banyuls, southern France.
Scale lines A-D, F, 0.1 mm; E, 50 mhi.

164

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

QOO

I I I I — I I I I I I — I — r

10 15

-I— I — I I I I I I
25

No. of segments
Fig. 3. Gyptis propinqua. Relationship between
number of segments and length. Specimens from Ban-
yuls, southern France (squares), Koster area, Sweden
(circles), and Breidafjordur, northwestern Iceland (tri-
angles).

ferentiated in length and orientation, those
of segment 5, 8, 10, 12, 15, 17, 19, 21 and
23 slightly longer and oriented more dor-
sally than other ones (best observed on live
specimens). Notopodial lobes conical, usu-
ally with two internal aciculae (one small
and difficult to detect), and one emerging,
anteriorly situated, dorsally bent smooth
acicular seta (Fig. IF). About 10-15 capil-
lary setae inserted behind lobe, with smooth
proximal parts and two alternating rows of
teeth medially and subdistally (Fig. IG). A
few ventrally situated serrated capillaries
usually present (Fig. IH).

Neuropodia of median segments conical,
slightly asymmetrical, usually with two
internal aciculae (one larger than other)
and 20-30 compound setae. Distal part of
setal shafts internally reticulated (Fig. II, J).
Blades of median and dorsal setae long, ven-
tral ones very short. Ventral cirri annulated
with about 5-10 rings, longer than neuro-
podial lobe, on distinct cirrophores situated
far back on neuropodia (Fig. 2F).

Pygidium a rounded cone. Pygidial cirri
long and annulated, median papilla absent
(Fig. IK).

Color: Live specimens transparent with
dark brown pigmentation forming segmen-
tally arranged transverse bands across dor-
sum that also connect laterally. First dor-
sally visible segment with distinct dark

brown band dorsally. Eyes orange-red.
Specimens vary from dark to pale, depend-
ing on amount of pigmentation. Venter al-
ways unpigmented. Eggs uncolored. Mature
males whitish. Pigmentation usually fades
in alcohol.

Measurements: Up to 7.5 mm long for 32
segments (see Fig. 3).

Habitat.— CosiTSQ and fine shell gravel
from 10-100 m.

Distribution. —IcQland, Faroes, Swedish
west-coast, southern France, Sicily.

Reproduction. —Mature specimens found
in May and June at Koster, Sweden, in July
at Iceland. Eggs large, about 175 ixm in di-
ameter. Bhaud (1971) described the larvae
of G. propinqua.

Remarks.— Eliason (1962), Haaland &
Schram (1982) and Helgason et al. (1990),
treated G. propinqua as a possible junior
synonym of G. rosea. The two most recent
studies, however, commented on inconsis-
tencies between different descriptions. Fur-
ther, Hartmann-Schroder (1971) and Hel-
gason et al. stated that Fauvel's (1923)
description of G. propinqua referred to G.
rosea. There is no remaining type material
of G. propinqua, either at the museum in
Paris, or at Station Marine d'Endoume in
Marseille (J.-C. Dauvin and G. Bellan, in
litt.); it is presumed lost. Nevertheless, new-
ly collected material from Banyuls-sur-Mer
was found to correspond well with the ex-
cellent original description based on speci-
mens from Marseille. Comparison of the
Banyuls specimens with both the type and
newly collected material of G. rosea leaves
no doubt that G. propinqua is a distinct and
very different species (see key for the best
diagnostic characters). Hartmann-Schro-
der' s (1971) description and examination of
the specimens of Helgason et al. (1990)
shows both accounts of G. rosea actually
refer to G. propinqua. By contrast, the ac-
counts of G. rosea by Eliason (1962) and
Haaland & Schram (1982) are considered
correct and refer to G. rosea as described
here. Fauvel's (1923) description of G. pro-

VOLUME 106, NUMBER 1

165

pinqua (as Oxydromus propinquus) is in good
agreement with Marion & Bobretzky's spe-
cies.

Ushakov (1955) reported G. propinqua
(as Oxydromus propinquus) from the Sea of
Okhotsk, but judging from his description
this presumably represents a different spe-
cies.

Gyptis mackiei, new species
Figs. 4-5

Material examined. —Sv/eden: 1 speci-
men, GuUmarsfjord, Skar, 110 m, mud, 25
May 1963, (NHMG 12805c); 4 paratypes
(SMNH 4395), Koster area, SW Yttre Vat-
tenholmen, 58°52.1'N, 11°06.8'E, 100-140
m, detritus sledge, mud, 8 Aug 1987; 2 para-
types (SMNH 4396), Singlefiord, 59°04'N,
1 riO'E, 80 m, detritus sledge, mud, 1 1 Apr
1990; holotype (SMNH 4397) and 2 addi-
tional specimens (used for dissection and
SEM), Koster area, S Yttre Vattenholmen,
58°52.0'N, 11°06.6'E, 110-130 m, detritus
sledge, mud, 1 Jan 1991; 1 specimen
(mounted for SEM), Koster area, S Yttre
Vattenholmen, 58°52.25'N, 11°06.30'E, 90-
140 m, detritus sledge, mud, 19 Aug 1991;
3 paratypes (NMW.Z. 1992.007.1-2) and
one additional specimen (mounted for
SEM), Single^ord, 59°04.5'N, 1 1°10.6'E, 78-
84 m, detritus sledge, mud, 26 Aug 1991;
1 specimen (mounted for SEM), Singleflord,
59°04.8'N, lin0.8'E, 82-83 m, detritus
sledge, mud, 15 Sep 1991. Skagerrak: 3
specimens, 58°08'N, 10°07'E, 295 m, grab,
27 Jun 1933; 1 specimen (ZMUU), 58°N,
09°33'E, 271 m, Agassiz trawl, 30 Jun 1933;
7 specimens (ZMUU), 58°02.5'N, 09°29.5'E,
478 m, Agassiz trawl, 30 Jun 1933; 1 spec-
imen (ZMUU), 57°50'N, 08°51'E, 358 m,
grab, 5 Jul 1933; 1 specimen (ZMUU),
57°41'N, 08°35'E, 191m, grab, 6 Jul 1933;
6 specimens (ZMUU), 58°22'N, 10°34'E,
270 m, Agassiz trawl, 14 Jul 1933; 2 spec-
imens (ZMUU), 58°30'N, 10°26'E, 300 m,
Agassiz trawl, 15 Jul 1933. Faroes: 4 spec-
imens (BIOFAR), 62°31.40'N, 05°02.30'W,

430 m, epibenthic sledge, 17 Jul 1987;
1 specimen (BIOFAR), 6r41.75'N,
05°47.71'W, 354 m, epibenthic sledge,
18 Jul 1987; 1 specimen (BIOFAR),
61°13.30'N, 04°46.50'W, 780 m, epibenthic
sledge, 19 Jul 1987; 4 specimens (BIOFAR),
60°31.34'N, 08°25.07'W, 732 m, epibenthic
sledge, 22 Jul 1987; 11 specimens (BIO-
FAR), 62°05.32'N, 10°06.71'W, 859 m, epi-
benthic sledge, 15 May 1988; 1 specimen
(BIOFAR), 62°12.30'N, 03°59.54'W, 402
m, detritus sledge, 27 May 1989; 2 speci-
mens (BIOFAR), 62°41.3rN, 10°03.90'W,
500 m, epibenthic sledge, 1 Jun 1989.

Description. —Body, excluding parapo-
dia, cylindrical; venter flattened, without
distinct longitudinal furrow. Median para-
podia only slightly longer than anterior,
posterior ones successively shorter, result-
ing body-outline of fairly constant width,
tapering slowly posteriorly.

Prostomium rounded, as wide as long,
anteriorly straight, posterior incision often
indistinct (Fig. 4A), often more conspicuous
on specimens with everted proboscis. Prox-
imal parts of palps cylindrical, distal parts
widest medially, anteriorly rounded (Fig.
4B); distal parts longer than proximal. Paired
antennae without distinct ceratophores, as
long as palps but thinner, with pointed tips.
Median antenna widest medially, without
distinctly pointed tip, inserted in front of
anterior pair of eyes. Anterior pair of eyes
rounded to reniform, slightly larger than
posterior pair and situated farther apart;
posterior pair rounded. Nuchal organs lat-
eral to prostomium, not coalescing dorsally.

Large, distinct lip glands present (Fig. 4B).
Proboscis short, smooth, divided into prox-
imal and distal parts; distal part smaller (Fig.
4C). Terminal papillae 25-26 (observed in
two specimens only; one 5.75 mm long for
26 segments, and one with posterior end
missing), short (about 60-70 fxtn) and blunt,
distally ciliated, arranged in single ring.

Tentacular cirri distinctly annulated, rings
from one to four times as long as wide, tips
rounded. Dorsal ones of segment 2 longest

166

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Gyptis mackiei, new species. A. Anterior end, dorsal view. Setae omitted. B. Anterior end, ventral
view. Setae omitted. C. Head end with protruded proboscis, dorsal view. D. Parapodium segment four, anterior
view, ca. half number of setae shown. E. Parapodium segment five, anterior view, ca. half number of setae

VOLUME 106, NUMBER 1

167

and stoutest, reaching to about segment 1 2-
16; ventral ones of segment 3 shortest,
reaching to about segment 5, similar to fol-
lowing ventral cirri but larger. Single acicula
present in all cirrophores of all dorsal ten-
tacular cirri; not observed in ventral ten-
tacular cirri (but may be present). Segment
1 reduced dorsally (possibly fused to seg-
ment 2), segment 2 fully developed.

Notopodia of segment 4 (setiger 1) with
one notoacicula situated in cirrophores of
dorsal cirri, without setigerous lobes or se-
tae (Fig. 4D). Dorsal cirri similar to those
of segment 3. Neuropodia similar to follow-
ing ones but slightly smaller, with about 1 5
compound setae. Ventral cirri similar to fol-
lowing ones. Segment 5 similar to segment
4 but slightly larger (Fig. 4E).

Segment 6 similar to median ones but
slightly smaller (Fig. 4F).

Notopodia of median segments with an-
nulated dorsal cirri with from three to seven
rings, shorter than notosetae, with pointed
tips (Fig. 4G). Dorsal cirri without obvious
differentiation in length and orientation.
Notopodial lobes conical, with one or two
(usually one) internal aciculae (smaller one
difficult to detect), and from one to three
emerging, anteriorly situated dorsally bent
acicular setae (Fig. 41). About 20-30 cap-
illary setae inserted behind lobe, with
smooth proximal parts and two alternating
rows of teeth medially and subdistally (Fig.
4H). Serrated notosetae not observed.

Neuropodia of median segments conical,
with single internal acicula and about 30-
50 compound setae. Distal part of setal shafts
internally reticulated (Fig. 4J, K). Blades of
median and dorsal setae long, ventral ones
short. A few dorsally situated serrated cap-
illary setae occasionally present. Ventral cir-
ri smooth or indistinctly annulated, as long

No. of segments

Fig. 5. Gyptis mackiei, new species. Relationship
between number of segments and length. Specimens
from Koster area, Sweden.

as or slightly longer than neuropodial lobe,
on distinct cirrophores situated far back on
the neuropodium.

Pygidium rounded. Pygidial cirri long,
annulated, median papilla absent.

Color: Live specimens transparent with
dark brown pigmentation forming trans-
verse stripes dorsally on anterior and pos-
terior sides of each segment, especially con-
spicuous on first dorsally visible segment.
Eyes red. Eggs rose-colored, mature males
whitish. Brown pigmentation fades in al-
cohol.

Measurements: Up to 5.75 mm long for
29 segments (see Fig. 5).

Habitat. — Found on mud bottoms from
78-859 m.

Distribution. —^orXYitm part of Swedish
west coast, Skagerrak, Faroes.

Reproduction. —Mature specimens found
in January in Sweden; not fully mature spec-
imens found in August. Eggs about 100 ^m
in diameter.

Remarks.— In many hesionids the ante-
rior parapodia are successively reduced dur-
ing ontogeny (e.g., Blake 1975; Haaland and
Shram 1983; Schram and Haaland 1984),
and the first segment carrying setae and

shown. F. Parapodium segment six, anterior view, ca. half number of setae shown. G. Parapodium segment 13,
anterior view, ca. one third of setae shown. H. Acicular notoseta. I. Spiked capillary notoseta. J. Median neuroseta.
K. Ventral neuroseta. A-C paratypes (NMW.Z. 1 992.007. 1-2), D-K specimen from Koster area, Sweden. Scales
A-C, 0.25 mm; D-G, 0.1 mm; H-K, 50 ^im.

168

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

number of tentacular cirri should be com-
pared between corresponding semapho-
ronts only. That the description above is
based on adults is evidenced by the presence
of sexual products in many of the speci-
mens.

Gyptis mackiei is unique among hesio-
nids in the character combination of "me-
dian antenna with dorsal insertion," and
"neurosetae present but notosetae absent on
segment 4 and 5." It is united with other
species of Gyptis by the place of insertion
of median antenna, by the presence of a
large number of proboscideal papillae, by
the absence of furcate setae, and by the ab-
sence of noto- but presence of neurosetae
on segment 5.

Gyptis mackiei seems closely related to
G. propinqua and the two species are rather
similar in general appearance. In addition
to the fourth setigerous segment G. mackiei
differs in having a median antenna which
is widest in its midregion rather than sub-
distally, in its prostomium being less deeply
incised posteriorly, in having much shorter
and more pointed dorsal cirri, and in having
rose-colored rather than colorless eggs.

Eliason's specimens from Gullmarsfjord
and of the Skagerrak-Expedition 1933 con-
stitute a mixture of G. rosea and G. mackiei;
they have now been relabelled.

Etymology. —This species is named for
Andrew Mackie, friend and collaborator.

Gyptis mediterranea, new species
Figs. 6-8, 12

Material examined. —France: 3 para-
types (NMW.Z. 1992.007.3), Banyuls-sur-
Mer, 42°29.92'N, 03°09.22'E, 35 m, dredge,
sandy mud with detritus, 3 Oct 1991; ho-
lotype (SMNH 4398), 7 paratypes (SMNH
4399, 4400), and 4 additional specimens
mounted for SEM, Banyuls-sur-Mer, Cap
Oullestrell, 42°30.17'N, 03°09.48'E, 40 m,
dredge, mud, 7 Oct 1991; 13 paratypes
(SMNH 4401), Banyuls-sur-Mer, 42°30.00'N,
03°11.75'E, 80 m, dredge, mud, 13 Oct 1991.

Italy: 1 specimen, eastern Sicily, BrucoU,
37°17'N, 15°13'E, 60 m, dredge, mud, 21 May
1990.

Description.— Body, excluding parapo-
dia, cylindrical, tapering posteriorly; venter
flattened without distinct median longitu-
dinal furrow. Parapodia long, often directed
anteriorly, median ones longest, yielding el-
liptical and flattened outline of animal.

Prostomium trapezoidal with rounded
comers (Figs. 6A, 7A), with small posterior
incision (often visible only in SEM). Prox-
imal parts of palps cylindrical, distal parts
thinner, narrowing to rounded ends. Prox-
imal parts slightly longer than distal. Paired
antennae slightly shorter and narrower than
palps, with thin tips, situated on small cera-
tophores. Median antenna cylindrical, end-
ing without well defined, prolonged tip, in-
serted on line between anterior pair of eyes.
Anterior pair of eyes rounded, larger than
posterior pair and situated farther apart,
posterior pair rounded, both pairs with lens-
es. Nuchal organs lateral to prostomium,
not dorsally coalescing (Fig. 7B).

Lip glands absent (Fig. 6B). Proboscis di-
vided in proximal and distal parts. Proxi-
mal part larger and longer, with small, poor-
ly defined papillae (Fig. 7A); distal part short
and smooth. Number of terminal papillae
size-dependent (Fig. 12), about 20-32. Pa-
pillae long (ca. 60-70 /iim) and thin with
ciliated tips, arranged in single ring (Fig.
7C).

Tentacular cirri thin, annulated (not al-
ways obvious proximally) with rings (one
to three times as long as wide), tips distinctly
pointed. Dorsal tentacular cirri of segment

2 longest, reaching to about segment 10-12,
ventral tentacular cirri of segment 3 short-
est, reaching to about segment 5-6. Cirri of
segment 3 and 4 more ventrally displaced
than anterior ones. Aciculae present in all
cirrophores of tentacular cirri (small acces-
sory ones not observed but may be present).
Segment 1 dorsally reduced, segments 2 and

3 dorsally fused.

Notopodia of segment 5 (setiger 1) with

VOLUME 106, NUMBER 1

169

^^^ ^/i/i

p\x^'7///,/7^^

f\(\AA/7

'%u,^^''^\

Fig. 6. Gyptis mediterranea, new species. A. Anterior end. dorsal view. Setae omitted. B. Anterior end,
ventral view. Setae omitted. C. Parapodium segment five, anterior view. D. Parapodium segment six, anterior
view, ca. half number of setae shown. E. Parapodium segment 14, anterior view, ca. half number of setae shown.
F. Acicular notoseta. G. Spiked capillar>' notoseta. H. Serrated capillary notoseta. I. Median neuroseta. H.
Serrated capillar>' notoseta. I. Median neuroseta. J. Ventral neuroseta. A-B hololype, C paratype (SMNH 4401),
D-J paratype (SMNH 4399). Scales A-B, 0.5 mm; C-E, 0.25 mm; F-J, 50 urn.

170

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

IBI^ ff^^jL^: . ^

r * -^

%^i^'sfe^^

c • ^^^^*f 1

Fig. 7. SEM micrographs of Gyptis mediterranea, new species. A. Anterior end, dorsal view. B. Anterior
end, right side. C. Terminal ring of proboscis, anterior view. D. Anterior-dorsal view of dorsum, median segments.
E. Median parapodium, left side, postero-dorsal view. F. Median parapodium, right side, antero- ventral view.
Specimens from Banyuls, southern France. Scale lines, 0.1 mm.

VOLUME 106, NUMBER 1

171

one notoacicula in cirrophores of dorsal cir-
ri, without setigerous lobes or setae. Dorsal
cirri similar to those of segment 4. Neuro-
podia similar to following ones but slightly
smaller and with fewer setae (Fig. 6C). Ven-
tral cirri similar to following ones. Segment
6 similar to median ones but slightly smaller
(Fig. 6D).

Elevated dorsal ridges present across pos-
terior side of each segment (Fig. 7D, E), less
distinct on anterior segments and more pro-
nounced on median and posterior ones. In
one specimen the ridges are provided with
distinct cylindrical papillae (length ca. 15
Mm).

Notopodia of median segments with more
or less distinctly annulated dorsal cirri
(smooth proximally), with about five rings,
about three times as long as wide. Dorsal
cirri shorter than setae, inserted posteriorly
to those (Fig. 6E). All dorsal cirri thin, of
approximately similar length, but those of
segment 5, 8, 10, 12, 15, 17, 19, 21, and 23
oriented slightly more dorsally than other
ones (best observed on live specimens). No-
topodial lobes conical, with one or two in-
ternal aciculae (small one difficult to detect)
and from zero to four emerging, dorsally
bent acicular setae, situated anteriorly to
other setae (Fig. 6F); tapering but termi-
nated bluntly, often with fine spines distally.
Large number of long capillary setae in-
serted behind lobe, median ones twice as
long as dorsal and ventral ones, with smooth
proximal parts and two alternating rows of
medial and subdistal teeth (Fig. 6G). A few
serrated notosetae usually present, situated
ventrally (Fig. 6H).

Neuropodia of median segments conical,
usually with two internal aciculae, one large
and one small, about 20-40 compound se-
tae, and, occasionally, one or two dorsally
serrated capillaries. Distal part of setal shafts
with transverse striation internally. Blades
thin, dorsal side from distinctly serrated to
almost smooth, median and dorsal ones long
(Fig. 61), ventral ones shorter (Fig. 6J). A
few additional serrated capillary setae often

o o°

^-^6-

£5-

.C 4-

ts-

_J 2-

1 1 1 1 1 1 1
5

1 1 1 1 1

1 1 1 1 1
15

1 1 1 1

1 1 1 1 1
25

1 1 1 I 1 1 1
30 3J

No. of segments

Fig. 8. Gyptis meditenanea, new species. Relation-
ship between number of segments and length. Speci-
mens from Banyuls, southern France.

present, situated dorsally. Ventral cirri
smooth with fine tapering tips (Fig. 7F),
without cirrophores, situated distally on
neuropodium.

Pygidium rounded. Pygidial cirri annu-
lated, longer than dorsal cirri, with pointed
tips, median papilla absent.

Color: Live specimens transparent. Eyes
red. Small brown pigment spots may be
present ventrally on posterior side of para-
podia. Eggs colorless. Preserved specimens
white, brown spots retained.

Measurements: Up to 7 mm long for 32
segments (see Fig. 8).

Habitat. —Mud and sandy mud from 35-
80 m.

Distribution. —Presently known only from
southern France and eastern Sicily.

Reproduction. —Several of the specimens
collected in Banyuls in October were mature
females with an egg size of 50-60 yum in
diameter. No males observed.

Remarks.— ^iXhirv Gyptis G. rosea and
G. mediterranea are unique in having dis-
tally inserted ventral cirri. They share with
G. hians Fauchald & Hancock, 1981, the
elliptical and flattened body-shape, but the
latter differs in having ventral cirri inserted
subdistally, and in having the distal part of
setal shafts internally reticulated rather than
striated.

Gyptis mediterranea differs from G. rosea
in being smaller, having red rather than black
eyes, having a median antenna without ex-

172

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

tended tip, having a smaller number of ter-
minal proboscis papillae (even when ad-
justed for size; see Fig. 12), having shorter
dorsal cirri, having much more pronounced
dorsal ridges, and in having colorless rather
than pink eggs.

Etymology.— ISiamed for the Mediterra-
nean Sea.

Gyptis rosea (Malm, 1874)
Figs. 9-12

Ophiodromus roseus Malm, 1874. (p. 82)
Gyptis rosea. — Haaland & Schram, 1982.—

Eliason, 1962 (p. 238-240, fig. 9), in part.

[Not Gyptis rosea sensu Hartmann-

Schroder, 1971:132-134, fig. 42.-Hel-

gasonetal., 1990:205]

Material examined.— ^"^Qd^n: Holotype
(NHMG 901), Gullmarsjgord, 45 fathoms,
mud; 2 specimens (NHMG 12805a & b),
Gullmarsfjord, Skar, 110 m, mud, 25 May
1963; 1 specimen, Single^ord, 59°04.9'N,
11°10.8'E, 80 m, detritus sledge, mud, 27
Dec 1988; 2 specimens, SingleQord,
59°04.9'N, 1 1°10.8'E, 80 m, detritus sledge,
mud, 14 Jun 1989; 2 specimens, Koster area,
WSvartskar, 58°54.5'N, 11°05.0'E, 100-150
m, dredge, mud, 22 Sep 1989; 6 specimens
(2 mounted for SEM), Singleflord,
59°04.9'N, 1 1°10.8'E, 80 m, detritus sledge,
mud, 11 Apr 1990; 1 specimen, Koster
area, S Yttre Vattenholmen, 58°52.1'N,
11°06.9'E, 50-100 m, dredge, mud, 1 Oct
1990; 3 specimens (mounted for SEM), Sin-
glefjord, 59°04.9'N, 11°10.8'E, 83 m, detri-
tus sledge, mud, 26 Feb 1991; 1 specimen,
Koster area, S Yttre Vattenholmen,
58°52.1'N, 11°06.9'E, 80-140 m, dredge,
mud, 17 Aug 1991; 5 specimens, Sin-
glefjord, 59°04.5'N, 11°10.6'E, 82-84 m,
detritus sledge, mud, 2 Sep 1991; 2 speci-
mens (1 mounted for SEM), Koster area, S
Yttre Vattenholmen, 58°52.0'N, 11°06.6'E,
100-110 m, detritus sledge, mud, 7 Sep
1991; 5 specimens, Singlefiord, 59°04.8'N,
11°10.8'E, 82-83 m, detritus sledge, mud,
15 Sep 1991. Skagerrak: 3 specimens

(ZMUU), 58°08'N, 10°07'E, 295 m, grab,
27 Jun 1933; 2 specimens (ZMUU),
58°2.5'N, 09°29.5'E, 478 m, grab, 30 Jun
1933; 1 specimen (ZMUU), 58°02.5'N,
09°29.5'E, 427 m, grab, 1 Jul 1933; 1 spec-
imen (ZMUU), 58°21'N, 08°56'E, 225 m,
grab, 2 Jul 1933; 1 specimen (ZMUU),
58°02.7'N, 08°13.5'E, 241 m, grab, 4 Jul
1933; 4 specimens (ZMUU), 57°50'N,
08°51'E, 358 m, grab, 5 Jul 1933; 1 speci-
men (ZMUU), 57°45'N, 08°07'E, 421 m,
grab, 6 Jul 1933; 3 specimens (ZMUU),
57°52'N, 08°01'E, 510 m, Agassiz trawl, 6
Jul 1933; 2 specimens (ZMUU), 57°58'N,
06°44'E, 384 m, grab, 7 Jul 1933; 1 speci-
men (ZMUU), 58°59.5'N, 06°27'E, 290 m,
dredge, 12 Jul 1933; 6 specimens (ZMUU),
58°22'N, 10°34'E, 270 m, Agassiz trawl, 14
Jul 1933; 5 specimens (ZMUU), 58°30'N,
10°26'E, 300 m, Agassiz trawl, 15 Jul 1933;
1 specimen (ZMUU), 58°30'N, 10°32.5'E,
175 m, Agassiz trawl, 15 Jul 1933; 1 spec-
imen, 57°59'N, 08°40'E, 500 m, RP-sledge,
15 Mar 1990; 1 specimen, 57°49.5'N,
08°12.5'E, 500 m, RP-sledge, 15 Mar 1990.

Description. —Body, excluding parapo-
dia, cylindrical, tapered posteriorly; venter
flattened, with distinct median longitudinal
furrow. Parapodia long, median ones lon-
gest, resulting in elliptical and flattened out-
line of animal.

Prostomium rounded rectangular to trap-
ezoidal, almost twice as wide as long (Fig.
9 A), with small posterior incision (often not
visible except in SEM; Fig. lOA). Proximal
parts of palps cylindrical, distal parts thin-
ner, anteriorly rounded; proximal and distal
parts of equal length. Paired antennae with-
out ceratophores, as long as palps but thin-
ner, with prolonged tips. Median antenna
similar in shape to frontal ones but smaller,
inserted just in front of anterior pair of eyes.
Anterior and posterior pairs of eyes small,
rounded; anterior pair slightly larger. Nu-
chal organs lateral to prostomium, not co-
alescing dorsally.

Lip glands absent. Proboscis divided into
proximal and distal parts (not obvious on

Fig. 9. Gyptis rosea. A. Anterior end, dorsal view. Proboscis partly everted. Setae omitted. B. Anterior end
with proboscis everted (some papillae lacking), ventral view. Setae omitted. C. Right parapodium, segment 5,
posterior view. Full number of setae (ca. 30) not shown. D. Right parapodium, segment 6, anterior view. Full
number of notosetae (ca. 25) or neurosetae (ca. 40) not shown. E. Right parapodium, segment 21. anterior view.
Full number of notosetae (ca. 40) or neurosetae (ca. 40) not shown. F. Acicular notoseta. G. Spiked capillary
notoseta. H. Median neuroseta. I. Ventral neuroseta. Specimens from Singlefjord and Koster area. Scales A-B,
0.5 mm; C-E, 0.25 mm; F-I, 50 ^m.

174

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

^-^^H

^^feb ' j^^sSMMHt^

• \&- § jM*'^ ^^^^1

^'^g

Ls^^^^^^^^^^^^^^^^^^^^^^^^l

ig^

Fig. 10. SEM micrographs of Gyptis rosea. A. Anterior end, dorsal view. B. Distal part of proboscis, ventral
view. C. Insertion of tentacular cirri, right side. D. Parapodia from segment 25-26, right side, antero-dorsal
view. E. Parapodium from segment 1 1 , left side, antero-ventral view. F. Spiked capillary notoseta from median
segment. G. Articulation between shaft and blade of median neuroseta from segment 18. Specimens from
Singlefjord, Sweden. Scales A-E, 0.1 mm; F-G, 10 )um.

VOLUME 106, NUMBER 1

175

Specimens with strongly extended probos-
cides) (Fig. 9B). Proximal part larger and
longer, covered with poorly defined, dif-
fusely distributed papillae, about 30-40 ijim
in diameter (Fig. 10 A) (surface appearing
slightly rugose in lower magnifications); dis-
tal part short and smooth. Number of ter-
minal papillae size-dependent (Fig. 12),
about 35-80. Papillae long and thin with
ciliated tips (Fig. lOB), arranged in single
ring (may appear as several rings in speci-
mens with proboscis incompletely protrud-
ed).

Tentacular cirri thin, annulated (not al-
ways obvious proximally) with median rings
about three times as long as wide, tips dis-
tinctly pointed. Dorsal tentacular cirri of
segment 2 longest, reaching to about seg-
ment 12-18; ventral tentacular cirri of seg-
ment 3 shortest, reaching to about segment
5-7. Cirri of segment 3 and 4 more ventrally
displaced than anterior ones (Fig. IOC). Sin-
gle acicula present in all cirrophores of ten-
tacular cirri except ventral ones of segment
3 and 4, which have double ones (smaller
ones difficult to detect).

Anterior segmental delineations not very
distinct. Segment 1 reduced dorsally except
for small middorsal part (Fig. lOA), seg-
ments 2 and 3 appearing fused dorsally, fol-
lowing segments fully developed.

Notopodia of segment 5 (setiger 1) with
one notoacicula situated in cirrophore of
dorsal cirrus, but without setigerous lobes
or setae. Dorsal cirri similar to those of seg-
ment 4. Neuropodia similar to following
ones but shorter (Fig. 9C). Ventral cirri sim-
ilar to following ones. Segment 6 similar to
median ones but smaller (Fig. 9D).

Notopodia of median segments with more
or less distinctly annulated dorsal cirri
(smooth proximally), with about eight to ten
rings; median rings about three times as long
as wide (Fig. 9E). Dorsal cirri thin, longer
than notosetae, inserted posteriorly to those
(Fig. lOD). All dorsal cirri of similar length,
those of segment 5, 8, 10, 12, 15, 17, 19,
21, 23 and 26 oriented slightly more dor-

No. of segments

Fig. 1 1 . Gyptis rosea. Relationship between num-
ber of segments and length. Specimens from Koster
area and Singlefjord, Sweden.

sally than other ones (best observed on live
specimens). Notopodial lobes conical, usu-
ally with two internal aciculae (one small
and difficult to detect) and from one to six
(usually from three to five) emerging, dor-
sally bent acicular setae, situated anteriorly
to other setae (Fig. 9F), tapering but ter-
minated bluntly, occasionally with fine
spines distally. Large number of long cap-
illary setae inserted behind lobe, median
ones twice as long as dorsal and ventral ones,
with smooth proximal parts and two alter-
nating rows of teeth medially and subdis-
tally (Figs. 9G, lOF). A few serrated noto-
setae situated ventrally occasionally present
(Fig. 9H).

Neuropodia of median segments conical
to rounded, usually with two internal acic-
ulae, one large and one small, and about
40-50 compound setae (Fig. lOG). Distal
part of setal shafts with internal transverse
striation. Blades thin, dorsal side varying
from distinctly serrated to almost smooth;
median and dorsal ones long (Fig. 91), ven-
tral ones short (Fig. 9J). A few additional
serrated capillary setae may occur dorsally
(difficult to detect). Ventral cirri smooth with
fine, evenly tapering tips (Fig. lOE), situated
distally on neuropodium, without cirro-
phores.

Pygidium rounded. Pygidial cirri very
long, annulated, median papilla absent.

Color: Live specimens transparent, stom-
ach yellowish to orange. Mature females

176

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

20 25 30

No. of segments

Fig. 1 2. Relationship between number of segments
and number of terminal proboscideal papillae in Gyptis
propinqua (triangles), G. mediterranea, new species
(squares), and G. rosea (circles).

pink-orange. Eyes dark brownish-black.
Dark brown pigment spots may be present
dorsally and ventrally on posterior side of
parapodia. Preserved specimens yellowish-
white. Brown pigment usually retained.

Measurements: Up to 1 1 mm long for 36
segments (see Fig. 11).

Habitat. —Mud bottoms from 50-5 10 m.

Distribution. —Presently known only from
Skagerrak, OsloQord and northern part of
Swedish west coast.

Reproduction. —Females with eggs found
in Sweden in April and August, the former
immature. Mature eggs bright pink-orange,
small, about 50-60 ixva in diameter. The
holotype is full of eggs which may explain
Malm's choice of specific name; "rosea" in-
dicating the color of the live mature female.

Remarks. — Eliason's specimens from
Gullmarsflord and the Skagerrak-Expedi-
tion of 1 933 constitute a mixture of (z. rosea
and G. mackiei, and have not been re-la-
belled. His description (Eliason 1962), how-
ever, clearly is based on specimens of G.
rosea. Haaland & Schram (1982) provided
descriptions of both adults and juvenile
stages from Oslofjord. For previous syn-
onymy with G. propinqua, see Remarks for
that species.

Amphiduros Hartman, 1959

Amphiduros Hartman, 1959:182 (replace-
ment name for Amphidromus Hessle,
1925).

Type species. —Amphidromus setosus
Hessle, 1925, by subsequent designation
(Hartman 1959:182).

Diagnosis (provisional) . —l:^QS\owids with
two palps and two frontal antennae. Median
antenna present, inserted dorsally on pro-
stomium. Proboscis without terminal pa-
pillae. Jaws absent. Segment 5 without no-
topodia or notosetae, with neuropodia and
neurosetae. Following parapodia with well-
developed noto- and neuropodia and large
number of noto- and neurosetae. Furcate
notosetae absent.

Remarks. —Amphiduros is presently sep-
arated from Gyptis solely on the absence of
terminal ring of proboscideal papillae. One
of the states (presence or absence of papil-
lae) is presumably ancestral, and the genus
defined on that state may be paraphyletic
unless supported by other characters. The
matter warrants further investigation.

A mphiduros fuscescens (Marenzeller, 1875),

new combination

Fig. 13

Oxydromus fuscescens Marenzeller, 1875:
143-146, pl.2, fig. 1.

Material examined. — Italy: 1 syntype,
(NHMW 2446), Trieste, St. Servola, 3-4 m,
stones with bore-holes; 1 specimen, eastern
Sicily, Acitrezza, 37°33.5'N, 15°11.1'E, 42-
45 m, dredge, muddy sand, 5 Apr 1990; 1
specimen, eastern Sicily, Brucoli, 37°17.rN,
15°12.6'E, 24 m, SCUBA, mixed bottom
with gravel, boulders and algae, 23 Apr 1 990.
France: 1 specimen (NMW), Banyuls-sur-
Mer, lie Grosse, 42°29.0'N, 03°08.1'E, 10
m, SCUBA, shell gravel, 13 Oct 1991; 1
specimen, Banyuls-sur-Mer, He Grosse,
42°29.0'N, 03°08.rE, 10 m, SCUBA, shell
gravel, 13 Oct 1991; 2 specimens, Banyuls-
sur-Mer, Cap Oullestrel, 42°30.22'N,
03°08.30'E, 18 m, dredge, shell gravel, 15
Oct 1991. Israel: 4 specimens. Flat, 28°30'N,
34°34'E, 4 m, SCUBA, 8 Mar 1986.

Description. —Body, excluding parapo-
dia, cylindrical, venter flattened, with dis-
tinct median longitudinal furrow (in larger

VOLUME 106, NUMBER 1

177

Fig. 13. Amphiduros fuscescens. A. Prostomium, dorsal view. B. Parapodium segment 5, anterior view, ca.
half number of setae shown. C. Parapodium segment 6, anterior view, ca. half number of setae shown. D.
Parapodium segment 14, anterior view, ca. one-third of number of setae shown. E. Spiked capillary notoseta.
F. Serrated capillary notoseta. G. Median neuroseta. H. Ventral neuroseta. A & E-H specimens from eastern
Sicily, B-D specimens from Banyuls, southern France. Scales A, 0.5 mm; B-D, 0.25 mm; E-H, 50 /im.

Specimens only). Median parapodia only
slightly longer than anterior ones, posterior
ones successively shorter, resulting body-
outline of fairly equal width with slowly ta-
pered posterior end.

Prostomium wider than long, anteriorly
straight, laterally and posteriorly with
rounded lobes separated by deep posterior
incision (Fig. 13A). Palps long and thin,

proximal parts cylindrical, distal parts ta-
pering to a point. Proximal and distal parts
of equal length. Paired antennae situated on
small ceratophores, as long as palps, slightly
more slender, with drawn-out tips tapering
to a point. Median antenna pointed, much
shorter than paired ones, inserted on line
between anterior side of anterior pair of eyes.
Eyes very large with lenses; anterior eyes

178

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

twice as large as posterior. Eye pigment
poorly delineated, spreading across anterior
part of prostomium. Nuchal organs lateral
to prostomium, coalescing dorsally, large
and distinct.

Lip glands absent. Proboscis smooth, di-
vided in proximal and distal parts; proximal
part slightly larger. Terminal ring of papillae
lacking, dense ciliation present. One small
specimen (anterior end, 3.0 mm for 1 1 seg-
ments) with (probably) 10 papillae in ter-
minal ring.

No specimen observed with complete
tentacular cirri, but dorsal ones of segments
3 and 4 much stouter than others. Ventral
tentacular cirri shorter and thinner than
dorsal. Cirri of segments 3 and 4 more ven-
trally displaced than anterior ones. Tentac-
ular cirri without obvious annulation, all
situated on long and large cirrophores with
several (2-6) internal aciculae.

Anterior segmental delineations uncer-
tain, segment 1 probably reduced dorsally.
Notopodia of segment 5 (setiger 1) with two
notoaciculae situated in cirrophores of dor-
sal cirri, but without setigerous lobes or se-
tae (Fig. 13B). Dorsal cirri as stout as those
of segment 4. Neuropodia and ventral cirri
similar to following ones. Segment 6 similar
to median ones (Fig. 1 3C).

Notopodia of median segments with short
dorsal cirri, lacking apparent annulation;
subdistally widened and tapering to a point
(Fig. 13D). Notopodial lobes conical, with
one or two internal aciculae, without emerg-
ing acicular setae. Large number of capillary
setae (ca. 20-30) inserted behind lobe, with
smooth proximal parts and double rows of
small teeth medially and subdistally (Fig.
13E). A few, ventrally situated serrated no-
tosetae present (Fig. 13F).

Neuropodia of median segments conical,
with one or two internal aciculae and a large
number of stout compound neurosetae.
Distal part of setal shafts internally reticu-
lated. Blades of median (Fig. 13G) and dor-
sal setae much longer than ventral (Fig.
13H). Ventral cirri smooth, tapering to a
point; inserted distally without cirrophores.

Pygidium rounded. Pygidial cirri not ob-
served, median papilla absent.

Color: Live specimens with white pig-
ment spots distally on dorsal cirri. Eyes or-
ange. Eggs dark red. Preserved specimens
yellowish, white pigment spots and red col-
or of eggs disappear.

Measurements: No entire, non-regener-
ating Mediterranean specimens observed.
Measurements for most complete speci-
mens: length 15 mm, width 3.3 mm for 28
segments (posterior end lacking); length 6.2
mm, width 1.5 mm for 27 segments (regen-
erating). Two of the Gulf of Aqaba speci-
mens (see Remarks) are complete: length 14
mm, width 3.2 mm for 40 segments; length
10 mm, width 2.4 mm for 37 segments.

Habitat. —StonQs and coarse shell gravel
from 2-45 m. One specimen from Sicily
found in muddy sand.

Distribution. —Southern France, eastern
Sicily, northern Adriatic, Gulf of Aqaba (see
Remarks).

Reproduction. —Females with eggs found
in May at Sicily and March in Gulf of Aqa-
ba, about 100-1 10 fxm in diameter.

Remarks.— The newly collected speci-
mens are in good agreement with Maren-
zeller's type. According to his original de-
scription this species should have black
rather than orange eyes. What is remaining
of eye pigment on his type, however, sug-
gests reddish (or orange) rather than black
eyes.

The Gulf of Aqaba specimens are in good
condition and could not be distinguished
from Mediterranean ones. Common Med-
iterranean and Red Sea distribution may,
however, not be very common, and until
further material is available these speci-
mens are labelled Amphiduros cf. fusces-
cens.

Fauvel (1923) and Hartman (1959) syn-
onyvcnzed A. fuscescens (as Oxydromus) with
Gyptis propinqua. This is obviously incor-
rect, and the species differ in many respects,
most notably size, proboscideal papillation,
and annulation, shape and length of dorsal
and ventral cirri.

VOLUME 106, NUMBER 1

179

Key to European Species of
Amphiduros and Gyptis

1. Prostomium wider than long, lip
glands absent, ventral cirri inserted
distally 2

- Prostomium as wide as long, lip
glands present, ventral cirri inserted
subdistally 4

2. Eyes small with well delineated pig-
ment, adults (> ca. 20 segments)
with terminal papillae on proboscis,
dorsal cirri distinctly annulated and
tapering evenly to a point, acicular
notosetae present 3

- Eyes large with poorly delineated
pigment, adults without terminal
papillae on proboscis, dorsal cirri
subdistally widened, not annulated,
acicular notosetae absent A. fuscescens

3. Median antenna with distinct, well
delineated tip, eyes brownish-black,
proboscis of adults (> ca. 25 seg-
ments) with more than 35 papillae
in terminal ring, dorsal cirri reach-
ing farther than setae, distinct ele-
vated dorsal ridges absent . . . G. rosea

- Median antenna without delineated
tip, eyes red, proboscis of adults with
less than 35 papillae in terminal ring,
dorsal cirri not reaching farther than
setae, distinct elevated dorsal ridges
present G. mediterranea

4. Median antenna widest subdistally,
adults (> ca. 20 segments) with neu-
rosetae from segment five, dorsal
cirri much longer than setae

G. propinqua

- Median antenna widest medially,
adults with neurosetae from seg-
ment four, dorsal cirri much shorter
than setae G. mackiei

Acknowledgments

I wish to thank K. Fauchald and L. Ward
(USNM), K. Fitzhugh and L. Harris
(LACM), G. Hartwich (ZMB), G. V. Hel-
gason, E. Kritscher (NHMW), A. Mackie
(NMW), A. Muir (BMNH), A. Norrevang

(BIOFAR), L. Orrhage (NHMG), and L.
Wallin (ZMUU) for access to material and/
or working facilities, the crews of R/V Mi-
mir and R/V Nereus for field assistance, and
the staffs of Laboratoire Arago and Tjamo
Marine Biological Laboratory where part of
the work was conducted. Special thanks to
P. Bouchet for admitting a polychaete work-
er to join the "Fifth European Malacological
Workshop" at Sicily, to G. V. Helgason for
hostship on Iceland, and to A. Mackie for
field collaboration, discussions as well as
comments on the manuscript. Financial
support was provided by Helge Ax: son
Johnsons Stiftelse and the Swedish Natural
Science Research Council (contracts 9555-
302 & -307).

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Appendix

Checklist of species and subspecies of Amp hiduros-
and Gyptis, with notes on material examined.

Amphiduros axialensis Blake & Hilbig, 1990. Hydro-
thermal vents of northeast Pacific, Juan de Fuca
Ridge. Type material NMCA. No material ex-
amined.

A. fuscescens (Marenzeller, 1875), new combination.
As Oxydromus fuscescens. Muggia, Trieste,
Adriatic. Syntype (NHMW) and non-types
(SMNH, NMW) examined.

A. izukai (Hessle, 1925). As Amphidromus izukai. Ja-
pan. Syntype (ZMUU) examined. Synonymous
to A. setosus (Hessle, 1925).

A. pacificus Hartman, 1961. California. Holotype
(LACM) and non-types (LACM) examined.

A. setosus (Hessle, 1925). As Amphidromus setous. Ja-
pan. Syntype (ZMUU) examined. Synonymous
\o A. zzw/ca/ (Hessle, 1925).

Gyptis brunnea (Hartman, 1961). As Oxydromus brun-
nea. California. Holotype (LACM) examined.

G. helgolandica Hilbig & Dittmer, 1979. North Sea.
Non-types (SMNH) examined. Belongs to Po-
darkeopsis.

G. hians Fauchald & Hancock, 1981. Off Oregon. Ho-
lotype (LACM), paratypes (LACM) and non-
types (LACM) examined.

G. incompta Ehlers, 1912 (complementary description
based on other specimens in Ehlers 1913). Kai-
ser Wilhelm II Land or Victoria Land, Antarc-
tic. Type material lost? Non-types (ZMH,
SMNH) examined.

G. lobata (Hessle, 1925). As Oxydromus lobatus. Ja-
pan. Syntype (ZMUU) examined. Synonymous
to G. pacifica.

G. maraunibi nae Gibbs, 1971. Solomon Islands. Ho-
lotype (BMNH) examined. Belongs to Podar-
keopsis.

G. ophiocomae Storch & Niggemann, 1967. Red Sea.
Type material in author's collection (Storch).
Non-type (SMNH) examined.

G. pacifica (Hessle, 1925). As Oxydromus pacificus.
Japan. Syntype (ZMUU) examined. Synony-
mous to G. lobata.

G. propinqua Marion & Bobretzky, 1875. Marseille,
France. No type material. Non-types (SMNH)
examined.

G. raluanensis (Augener, 1927). As Oxydromus ralu-
anensis. New Guinea. Syntype (ZMH) exam-
ined.

G. rosea (Malm, 1874), As Ophiodromus roseus. Gull-
marsfjorden, Sweden. Holotype (NHMG) and
non-types examined (NHMG, NHMR).

G. vittata Webster & Benedict, 1887. Maine, U.S. Syn-
types (USNM) examined.

Ophiodromus roseus Malm, 1874. See Gyptis rosea.

Oxydromus arenicolus La Greca, 1946. No type ma-
terial? No material examined. Belongs to Po-
darkeopsis.

O. arenicolus glabrus Hartman, 1961. Holotype
(LACM) examined. Belongs to Podarkeopsis.

O. aucklandicus Willey, 1902. Auckland Islands. Type
material lost. No material examined. Genus un-
certain (not Gyptis or Amphiduros). Nomen du-
bium.

O. brevipalpa Hartmann-Schroder, 1959. Type mate-
rial ZMH. No material examined. Belongs to
Podarkeopsis.

O. brunnea Hartman, 1961. See Gyptis brunnea.

O. capensis Day, 1963. Holotype (BMNH) examined.
Belongs to Podarkeopsis.

O. fasciatus Grube, 1855. Trieste, Italy, or Ville-
franche, southern France. Syntype (ZMB) ex-
amined. Junior synonym to Ophiodromus flex-
uosus (Delle Chiaje, 1827).

O.flaccidus Grube, 1857. St. Croix, West Indies. Type
material probably lost. No material examined.
Nomen dubium.

O. fuscescens yidivenztWer, 1875. 'S>tQ Amphiduros fus-
cescens, new combination.

O. /z^?ero<:M/a?w5 Hartmann-Schroder, 1965. Valdivia,
Chile. Type material ZMH. No material ex-
amined. Genus uncertain (not Gyptis or Am-
phiduros).

O. lobatus Hessle, 1925. See Gyptis lobatus.

O. longisetisGnibe, 1857. St. Croix, West Indies. Type
material probably lost. No material examined.
Nomen dubium.

O. pacificus Hessle, 1925. See Gyptis pacifica.

O. pallidus Qaparede, 1 864. No type material. Non-types
(SMNH) examined. Belongs to Podarke.

O. raluanensis Augener, 1927. See Gyptis raluanensis.

PROC. BIOL. SOC. WASH.

106(1), 1993, pp. 182-189

A NEW GENUS OF HYDROBIID SNAILS

(MOLLUSCA: GASTROPODA: PROSOBRANCHIA: RISSOOIDEA)

FROM NORTHERN SOUTH AMERICA

Robert Hershler and France Velkovrh

Abstract.— An aquatic hydrobiid cavesnail from Andes Mountains of Co-
lombia and Ecuador is described (Andesipyrgus sketi, new genus, new species).
The genus is placed in the Cochliopinae based on females having a sperm tube
separated from the glandular gonoduct. Diagnostic features oi Andesipyrgus
include a minute, near-pupiform shell with thickened aperture; unpigmented
animal (except for eyespots); reduced ctenidium; female genitalia featuring
complex histology of glandular gonoduct, oviduct coiled onto right side of
albumen gland, and absence of seminal receptacle; and male with simple,
slender penis. Andesipyrgus does not closely resemble other cochliopinids hav-
ing a simple penis, and appears remotely related to other South American
hydrobiid fauna.

The aquatic prosobranch snails of the
family Hydrobiidae comprise a large cos-
mopolitan group of several hundred genera
and a few thousands of Recent species. While
the group is highly diverse at lower taxo-
nomic levels in most of the major physio-
graphic regions where it occurs, one of the
few exceptions is South America, where only
seven genera (and about 1 20 Recent species)
are found. (North America has about 40
genera and over 200 species by compari-
son.) Of these, each of five are represented
by less than 10 species on the continent,
while the remaining two {Heleobia Stimp-
son, \%65\ Potamolithus ViXshry, 1896) have
radiated extensively. The apparent paucity
of hydrobiid fauna in South America surely
relates in part to hydrographic/geologic and
historical features of the continent, but also
probably reflects inadequate sampling of the
brackish coastal and inland waters of this
huge land mass.

One habitat that has been especially ne-
glected in the search for these animals in
South America is cave streams, whose hy-
drobiid fauna often is diverse and strongly
differentiated from local epigean (surface-
dwelling) taxa. While caves are not extreme-

ly numerous in the continent, there are large
areas of karstic terrain to the north (es-
pecially in the Andes Mountains and in the
Amazon basin of Brazil) which are poten-
tially fertile areas for discovery of cavesnail
fauna. A recent compendium of subterra-
nean aquatic mollusks (Bole & Velkovrh
1986) listed only a single species of hydro-
biid cavesnail from South America, which
was collected from the Andes Mountains of
Ecuador by a Yugoslavian expedition dur-
ing 1978 (Sket 1985). Restudy of this ma-
terial and other collections made by a Yu-
goslavian expedition to the Colombian
Andes in 1984 (Sket 1988) confirmed the
uniqueness of this animal, which we de-
scribe as a new genus and species below.

Material studied is from the personal col-
lection of the second author (FV). Types
have been deposited in the National Mu-
seum of Natural History, Smithsonian In-
stitution (USNM).

Family Hydrobiidae Troschel, 1857

Subfamily Cochliopinae Try on, 1866

Andesipyrgus, new genus

Type species.— Andesipyrgus sketi, new
species.

VOLUME 106, NUMBER 1

183

Diagnosis. —Shell minute-small, smooth,
narrow, with thickened aperture. Opercu-
lum corneous, thin, with eccentric nucleus
and slightly thickened ventral attachment
scar. Radula taenioglossate; central teeth
with two pairs of basal cusps, marginal and
lateral teeth with relatively numerous, fine
cusps. Rectum with bend or loop in pos-
terior pallial cavity. Stomach without pos-
terior caecum. Animal unpigmented except
for eyespots. Ctenidium reduced in length,
and with relatively few filaments. Females
oviparous; glandular gonoduct of complex
histology. Oviduct coiled behind (onto right
side) of albumen gland, bursa copulatrix of
posterior position, seminal receptacle ab-
sent. Bursal duct opening to sperm tube,
which has an anterior connection to the cap-
sule gland. Male prostate gland with prom-
inent pallial section. Penis slender, simple,
without lobes or enlarged glands.

Remarks. —The pupiform shell with
thickened aperture of Andesipyrgus (similar
to that of unrelated Bythinella Moquin-
Tandon, 1855, from Europe) does not re-
semble that of any other South American
hydrobiid. A remote affinity with other fau-
na of the region is further suggested by the
configuration of female genitalia (notably
the coiled oviduct), which is unique in the
Cochliopinae (and substantially different
from the usual pattern of a single coil of
oviduct on the left side of the albumen
gland). The simple penis (without terminal
constriction or large-sized glands) of An-
desipyrgus is shared by 10 other cochliopi-
nid genera, including several other subter-
ranean forms. This is considered a
phylogenetically diverse group (almost cer-
tainly representing a grade of organization),
whose genera are of uncertain relationship
both to one another (in most cases) and to
other cochliopinid groups (Hershler &
Thompson 1992). Andesipyrgus does not
closely resemble any of these genera and
further evaluation of its affinities cannot be
made at this time.

Several morphological features oi Ande-

sipyrgus, including pigmented eyespots and
brown periostracum, suggest that this ani-
mal only recently invaded cave waters.

Etymology.— From Andes, referring to
distribution along the Andes Mountains, and
Classical Greek pyrgos, meaning tower and
referring to the moderately elongate shell.
Gender masculine.

Andesipyrgus sketi, new species
Figs. 1-5, Table 1

Littoridinai?) jumandi Bole & Velkovrh

1986:196. [Nomen nudem.]
"Hydrobioidea-Gastropoda."-Sket 1985:

84.
"Hydrobioidea."-Sket 1988:55, 58.

Material examined. —Colombia (Santan-
der Department): La Paz area (6°irN,
73°35'W)— Cueva de los Indios, ca. 6 km
SW of La Paz, 1995 m elevation, VI- 1984,
FV 43575, FV 43689, USNM 860574 (ho-
lotype), USNM 860575 (paratypes); Hoyo
(de) Colombia, ca. 6 km SSE of La Paz, 1775
m (1 broken shell), VI-1984, FV 43691;
Hoyo del Aire, ca. 6 km SSW of La Paz,
1800 m, VI-1984, FV 43690. San Gil area
(6°33'N, 73°08'W)-Cueva del Indio, ca. 7.5
km S of San Gil, 1270 m, VI-1984, FV
43685. Ecuador: Cavemas de Jumandi, near
Archidona (0°55'S, 77°48'W), 140 km SW
of Quito, Napo Province, about 500 m, XII-
1978, FV 38129, FV 38130.

Description. —Shell (Figs. 1, 2a) narrowly
conic to pupiform, 1.6-2.5 mm tall, with
4.0-4.5 whorls. Protoconch (Fig. 2b) blunt,
smooth except for very faint pattern of low
wrinkles. Teleoconch whorls near flat to
slightly convex, rarely with slight adapical
shoulders and/or weak adapical angulation;
sutures very shallow. Teleoconch sculpture
of moderately strong growth lines. Aperture
ovate, less than 50% of shell height, broadly
adnate or very slightly separated from body
whorl, usually chalk-white (possibly due to
lengthy preservation in alcohol). Inner lip
complete, moderately reflected, thickened,
sometimes markedly so in parietal region.

184

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. I. Andesipyrgns sketi, holotype, USNM 860574, standard and side views. Shell height, 2.0 mm.

Outer lip thick, slightly expanded, weakly
sinuate, near orthocline. Umbilicus absent
or very narrowly rimate. Periostracum
brown.

Measurements (mm) of the shells of the
holotype and nine paratypes are in Table 1 .

Operculum (Fig. 2c-e) light brown, trans-
parent, ovate, paucispiral. Attachment scar
with weakly developed central callus and
slight thickening along inner margin.

Radular ribbon about 470 ^m long and
70 Mm wide, of about 66 rows of teeth,

Table 1.— Shell measurements (mm) for Andesipyrgus sketi. SH = standard height, HBW = height of body
whorl, SW = standard width, AH = aperture height, AW = aperture width, WH = number of whorls.

HBW

SW/SH

Holotype

2.0

1.5

1.2

0.87

0.79

4.0

0.58

Paratypes

2.1

1.4

1.2

0.83

0.81

4.25

0.58

2.2

1.5

1.2

0.91

0.87

4.5

0.55

2.0

1.3

1.1

0.75

0.73

4.5

0.55

2.2

1.5

1.2

0.81

0.73

4.5

0.54

2.1

1.4

1.2

0.87

0.81

4.25

0.58

2.2

1.5

1.3

0.89

0.85

4.5

0.58

2.1

1.4

1.2

0.83

0.79

4.5

0.57

2.2

1.5

1.2

0.85

0.89

4.25

0.54

2.1

1.5

1.2

0.81

0.83

4.25

0.56

2.0

1.4

1.2

0.85

0.75

4.0

0.61

VOLUME 106, NUMBER 1

185

Fig. 2. Scanning electron micrographs of shell and opercula of ^. sketi, USNM 860575. a. Shell (height, 2. 1
mm), b. shell apex (bar = 100 Mm), c. Operculum, dorsal surface (bar = 176 Mm), d. e. Operculum, ventral
surface (bar = 200 Mm).

scarcely extending beyond edge of well-de-
veloped buccal mass and without posterior
coil. Central teeth (Fig. 3a, b) trapezoidal,
with well indented dorsal edge; lateral an-
gles narrow, slightly thickened, well ex-
panded and sometimes broadened distally.
Lateral cusps of central teeth narrow, 4-5;
central cusp slightly longer than laterals;
basal cusps moderate to long, arising from

face of tooth near origin of lateral angles,
inner cusp much broader and slightly longer
than outer. Basal process of central teeth
narrow, well excavated. Lateral tooth (Fig.
3d) formula, 2(inner)/l/3-4(outer): cusps
narrow, curved, with central cusp enlarged.
Marginal teeth (Fig. 3c, d) with numerous
(about 1 7-20) cusps. Rectum broadly over-
lapping glandular gonoducts, with bend (fe-

186

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Radula of A. sketi, USNM 860575. a. Central teeth (bar = 6 ^im). b. Central teeth (bar = 10 ^m). c.
Outer marginal teeth (bar =10 /urn), d. Lateral and inner marginal teeth (bar = 13.6 ^m).

males) or pronounced U-shaped loop
(males) in posterior portion of pallial cavity;
anus near mantle edge.

Animal white (except for black eyespots),
without melanin pigment. Cephalic tenta-

cles with narrow central band of elongate
cilia on dorsal surface.

Ctenidium filling about 66% of pallial
cavity length, extending from well anterior
to pericardium almost to mantle edge. Cte-

VOLUME 106, NUMBER 1

187

nidial filaments about 12, short and narrow.
Osphradium narrowly ovate, about 33% of
ctenidium length, positioned along poste-
rior half of ctenidial axis. Kidney with slight
bulge into pallial cavity; kidney opening
thickened, white. Hypobranchial gland
weakly developed.

Ovary a small unlobed mass filling about
0.25 whorls and slightly overlapping pos-
terior stomach chamber. Albumen gland
(Fig. 4a, Ag) with large pallial section (> 50%
of length). Capsule gland (Cg) about equal
to albumen gland in length, of two tissue
sections (posterior, orange; anterior, white).
Capsule gland composed of folded glandu-
lar cells, thick-walled, with central lumen.
Coiled oviduct (Co) bends posteriorly be-
hind pallial wall, loops behind albumen
gland to right side, then twists back to left
side of gland, where it coils once before
looping to right side of bursal duct (Dbu)
before joining the duct at the pallial wall.
Coiled oviduct with thin muscular coat,
strongly ciliated. Bursa copulatrix (Bu) small
(about 33% of albumen gland length), ovate,
scarcely extending posterior to albumen
gland. Bursa of tall glandular cells having
basal nucleii, lined with thick muscular coat.
Bursal duct emerging from anterior tip of
bursal copulatrix; distal section of duct
broadened, with an internal division into
two sections suggesting an enclosed seminal
receptacle. (Study of thin sections could not
confirm this possibility.) Duct to albumen
gland (Dag) opening posteriorly from point
where bursal duct and coiled oviduct join.
Sperm tube (St) broad, thick, muscular,
strongly ciliated; positioned ventro-laterally
to capsule gland, joining capsule gland dis-
tally. Genital aperture (Ga) a short terminal
slit.

Testis an unlobed mass, orange-colored,
filling about 0.5 whorl posterior to stomach.
Prostate gland narrow, bean-shaped; walls
thick, lumen slit-like; pallial section prom-
inent (about 60% of gland length). Posterior
vas deferens opening to prostate gland just
behind pallial wall; anterior vas deferens

Fig. 4. Genitalia of A. sketi, USNM 860575. a. Left
side of female glandular oviduct (bar = 0.5 mm). The
thick curving line represents the posterior wall of pallial
cavity. In the drawing to the right (slightly enlarged),
the coiled oviduct has been cut and rotated to expose
the bursa copulatrix and associated ducts. Abbrevia-
tions: Ag, albumen gland; Bu, bursa copulatrix; Cg,
capsule gland; Co, coiled oviduct; Dag, duct to albu-
men gland; Dbu, bursal duct; Ga, genital aperture; St,
sperm tube. b. Male penis, dorsal surface (bar = 0.25
mm).

Opening subterminally. Pallial section of vas
deferens with proximal coil. Penis (Fig. 4b)
small, tightly coiled, positioned well behind
cephalic tentacles slightly to right of center
of "neck." Penis vermiform, unciliated,
slightly broadened distally, with small folds
on inner curvature near base. Distal tip of
penis strongly tapered, vas deferens opening

188

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

TROPICAL AMERICA

Rora Neotropjca base map no 1

O 1 979 by ths University of Utracht

Published by the State University of Utracht, the Netherlands Department of Systematic B^otany

Fig. 5. Map showing distribution of A. sketi. 1, Colombian localities near La Paz and San Gil, Santander
Department; 2, Ecuador locality near Archidona, Napo Province.

through short terminal papilla. Proximal
course of vas deferens not discemable.

Type locality. —Stream flowing into Cueva
de los Indios, vicinity of La Paz, Santander
Department, Colombia.

Distribution. — K trans-Andean species,
occurring in cave streams of Rio Magdalena
basin (draining to Caribbean), Colombia;
and Rio Napo basin (Amazon basin), Ec-
uador.

Habitat.— Andesipyrgus sketi occurs in
small Andean cave streams at elevations be-
tween 500-1990 m. Water temperatures
among sites varied from 15-23°C. Some of
the cave streams represent sinking rivers

whereas others probably originate from per-
colating waters. All of the streams contained
some surface fauna, particularly insect lar-
vae (Chironomidae, Elminthidae, etc.) and
cyclopoid Copepoda, and most also were
inhabited by more or less troglomorphic an-
imals: variable catfish populations (Tricho-
myoterus sp. in Colombia, Astroblepus pho-
leter CoUette, in Ecuador) and probable
troglobiont, amphibious crabs {Neostren-
geria sketi Rodriguez, in Colombia) or am-
phipods (Bogidiella gammariformis Sket, in
Ecuador). Snails were most common in the
only cave lacking troglomorphic fauna
(Cueva de los Indios) and in which Tricho-

VOLUME 106, NUMBER 1

189

myoterus was normally pigmented and
shaped. For additional details, the reader is
referred to Sket (1979, 1988).

Remarks.— Avdii\2ih\Q material was un-
relaxed, and details of shape of head and
cephalic tentacles could not be discerned.

Populations varied in several shell fea-
tures (width, angulation of aperture, whorl
outline) and while available material does
not permit meaningful analysis, further
study may reveal that a species complex is
involved.

Etymology.— ^2iTs\Q& in honor of Dr.
Boris Sket, for his discovery of this species
and for his encouragement of this collabo-
rative study.

Acknowledgments

The authors thank Dr. Boris Sket for
making collections of the Yugoslavian cav-
ing expeditions available for study and for
providing useful comments on the manu-
script. Scanning electron micrographs were
prepared by Susann Braden of the Scanning
Electron Microscopy Laboratory, National
Museum of Natural History. Shells were
drawn by Molly Ryan, Dept. of Invertebrate
Zoology, National Museum of Natural His-

tory. Anatomical illustrations were inked by
Susan Escher.

Literature Cited

Bole, J., & F. Velkovrh. 1986. Mollusca from con-
tinental subterranean aquatic habitats. Pp. 1 77-
208 in Stygofauna mundi, a faunistic, distri-
butional, and ecological synthesis of the worid
faunas inhabiting subterranean waters (includ-
ing the marine interstitial). E. J. Brill/Dr. W.
Backhuys, Leiden, 740 pp.

Hershler, R., & F. G. Thompson. 1992. A review of
the genera of the aquatic gastropod subfamily
cochliopinae (Prosobranchia: Hydrobiidae). —
Malacological Review Supplement 5:1-140.

Sket, B. 1979. Fauna in the Cavema de Jumandi.—
Nase Jame 20:85-91.

. 1985. Bogidiella (s.l.) gammariformis sp. n.

(Amphipoda) from Ecuador.— Bioloski Vestnik
33:81-88.

. 1988. Speleobiological investigations in the

Colombian Andes 1984.— Bioloski Vestnik 36:
53-62.

(RH) Department of Invertebrate Zool-
ogy, National Museum of Natural History,
Smithsonian Institution, Washington, D.C.
20560, U.S.A.; (FV) Department of Biolo-
gy, Biotechnical Faculty, University of Lju-
bljana, Askerceva 12, 61001 Ljubljana, Slo-
venia.

PROC. BIOL. SOC. WASH.
106(1), 1993, pp. 190-194

VAMPYROCROSSOTA CHILDRESSI, A NEW GENUS

AND SPECIES OF BLACK MEDUSA FROM THE

BATHYPELAGIC ZONE OFF CALIFORNIA

(CNIDARIA: TRACHYMEDUSAE: RHOPALONEMATIDAE)

Erik V. Thuesen

Abstract.— A. new genus and species of deep-sea medusa, Vampyrocrossota
childressi, is described from the eastern North Pacific. It has been found in San
Clemente Basin off Baja CaHfomia, Mexico, and from the waters off Point
Conception, CaUfomia, U.S.A., at depths between 600 and 1475 meters. This
genus is alHed to the cosmopoHtan rhopalonematid genus Crossota, but differs
notably in the shape and position of the gonads. It is the only described species
of hydromedusa with black pigmentation.

During the course of an ongoing project
on the physiology and biochemistry of mid-
water gelatinous organisms off California, a
distinctive black medusa was commonly re-
covered in trawls taken deeper than 600 m.
The animals were collected by a 10 m^
Mother Tucker trawl using a specially de-
signed 30 1 insulated cod end to protect the
animals from heat and light as they are
brought to the surface (Childress et al. 1978).
Medusae were captured in very good con-
dition, and fragile hydromedusae of the
families Halicreatidae and Rhopalonema-
tidae were often brought aboard ship with
tentacles several body heights in length. The
black medusa is not included in reports of
Pacific Ocean hydromedusae (Alvarifio
1967, Kramp 1968, Segura-Puertas 1984),
and it is not one of the several new species
of mesopelagic rhopalonematid medusae
currently being described by Mills & Larson
(C. E. Mills, pers. comm.).

Vampyrocrossota, new genus
Figs. 1, 2

Diagnosis. — Rhopalonematidae without
gastric peduncle; stomach with four oral lips,
extending to or just past the velum when
empty; with eight tubular gonads attached

longitudinally to the eight radial canals in
all the specimens observed; exumbrellar
furrows present; with tentacles all of one
kind.

Type species. — Vampyrocrossota chil-
dressi, new species.

Etymology.— From. Serbian vampira, a
nocturnal demon supposed to eat the heart,
blood and soul of its victim, with reference
to Vampyroteuthis infernalis, the black me-
sopelagic squid often captured in the same
trawls as this animal, and Crossota the
closely allied rhopalonematid genus.

Relationships. —Both Bigelow (1913) and
Kramp (1947) discuss the genus Crossota
Vanhoffen, 1902 in some detail and are ex-
plicit that the pendant nature of the gonads
is an important characteristic distinguishing
this genus from other related genera. Given
the importance that this characteristic has
had in conserving the genus Crossota, I have
erected the genus Vampyrocrossota in this
paper. This genus is closely related to Cros-
sota in general appearance, lack of a pedun-
cle, the large number of tentacles and rib-
bon-like nature of the radial canals.

Recently, Larson & Harbison (1990) es-
tablished the new rhopalonematid genus
Benthocodon which also differs from Cros-
sota by having gonads attached to the radial

VOLUME 106, NUMBER 1

191

canals. They reported that the gonads in B.
hyalinus are ribbon-like and run along most
of the length of the eight gastric canals with
the most distal portions hanging free. Vam-
pyrocrossota is also different from Bentho-
codon in that it lacks a gastric peduncle and
has numerous exumbrellar furrows.

Vampyrocrossota childressi, new species
Figs. 1, 2

7>'/7€'5. — Holotype: a 12 mm tall speci-
men (USNM 91883) taken from 777 m
depth on 3 1 July 199 1 off Point Conception
from the RV Point Sur. Paratypes: two spec-
imens, 1 1 mm (Paratype A: USNM 91884)
and 6 mm (Paratype B: USNM 91885) from
984 m depth, captured on 30 July 1991 off
Point Conception, California. All types are
deposited in the National Museum of Nat-
ural History, Smithsonian Institution.

Description.— This description is based
upon observations of ~20 living animals
ranging in size from 6 to 14 mm in height
and up to 475 mg wet weight. Up to 14 mm
tall; up to 10 mm in diameter; velum up to
3 mm; jelly fairly thin, especially at apex;
mesoglea colorless; numerous exumbrellar
furrows; inner surface of bell is black with
pigment fading out posteriorly towards the
velum (particularly in immature animals);
velum is black in some specimens; up to
400 tentacles, all the same kind; tentacles
and radial canals reddish-orange; eight
cream-colored tubular gonads attached one-
eighth from the top to five-eighths the length
of the radial canal; stomach without pedun-
cle, reaching past the velum when extended;
stomach cream colored with a wide hori-
zontal black pigmented band located half
way to the four oral lips. The immature
paratype specimen had a completely orange
manubrium when it was collected before
preservation in formalin. Upon first in-
spection with the naked eye, this species
looks remarkably like Cross ot a rufobrunnea
with black rather than burgundy pigmen-
tation. Vampyrocrossota childressi is much

less active after capture and has a lower met-
abolic rate than many other rhopalonema-
tids which have been captured in the same
trawls. These other species include Crossota
alba, C. rufobrunnea, Pantachogon sp.,
Sminthea eurygaster, and Colobonema seri-
ceum (Thuesen & Childress, unpublished).

Etymology.— ^diVCiQd. in honor of James
J. Childress of the Marine Science Institute,
University of California at Santa Barbara
who has devoted a considerable part of his
life to the study of midwater organisms off
the California coast and is in part respon-
sible for the discovery of this medusa.

Distribution. —This animal has been re-
covered in trawls from San Clemente Basin,
off Baja California, Mexico where the bot-
tom depth can be greater than 2000 m to
northwest of Point Conception, California,
U.S.A. where the bottom depth is over 4000
m. The shallowest discrete depth tow in
which it has been taken was 600 m and it
has been taken in discrete depth trawls
reaching to 1475 m. We have not routinely
fished at depths greater than this and it is
not known how deep V. childressi occurs.
Although not abundant (never more than
three specimens in a trawl), it is routinely
taken in the above region all four seasons
of the year.

Coloration. —The pigmentation of V.
childressi is neither a dark blue nor deep
burgundy but is truly black. The color does
not fade in specimens preserved in 1 0% for-
malin in filtered seawater after storage in
the dark for two years. Other bathypelagic
organisms including fish, crustaceans and
molluscs are known to have black pigmen-
tation (Wimpenny 1 966, color frontispiece),
however no other species of hydromedusa
with black pigmentation is recorded in the
literature. Kramp (1961) describes the gut
of the coronate scyphozoan Nausithoe glo-
bifera as being black, however the guts of
other coronates, such as A^. rubra, Periphylla
periphylla, Atolla wyvillei and A. vanhoef-
feni, are not black but rather a densely-pig-
mented deep burgundy in living specimens.

192

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. L Side view of Vampyrocrossota childressi, new genus, new species, holotype, collected from 777 m
depth off Point Conception, California. Inner-bell pigmentation and velum pigmentation is black. Bell is 12
mm in height.

Anecdotal accounts reporting large num-
bers of a black medusa (Semaeostomeae;
Chrysaora sp.?) in the Los Angeles, Cali-
fornia, area were prevalent during July-Au-
gust of 1989, however the organism has yet
to be described in the scientific literature.
Other Cnidaria with black pigmentation are
known. The anemone Metridium senile has
a black endodermal melanin (Fox & Pantin
1941), and the siphonophore Erenna ri-

chardi has black endodermal pigmentation
which is thought to be acquired by feeding
on black midwater fishes (Totton 1965). The
small size of V. childressi suggests that me-
sopelagic fishes are not the source of its black
pigmentation. Some of the burgundy-col-
ored deep-sea medusae, including Crossota
rufobrunnea, contain porphyrin pigments
(Herring 1972, Bonnett et al. 1979). The
exact nature of the pigment in V. childressi

VOLUME 106, NUMBER 1

193

Fig, 2. Side view of Vampyrocrossota childressi, new genus, new species, drawn with inner-bell pigmentation
"removed" to reveal the gonads and manubrium. The manubrium can reach past the velum when extended in
living individuals.

is not known, although spectrophotometric
analysis of pigment extracted in ethanol re-
veals an absorption peak at 479 nm indi-
cating that it has blue-light absorbing com-
ponent (Thuesen, unpublished data).

Acknowledgments

I am grateful to J. J. Childress, K. L. Smith
and the captains, crews and scientists aboard

the Research Vessels Point Sur and New
Horizon for their assistance at sea. Research
cruises were supported through National
Science Foundation grants OCE 85-00237
to J. J. Childress and OCE 89-22620 to K.
L. Smith. I thank C. E. Mills for informative
discussion of mesopelagic medusae and for
comments on the manuscript. The figures
were drawn by P. Schalk of Stichting ter
Bevordering van de Nederlandse Oceano-

194

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

grafie. I was supported in part by a Califor-
nia Sea Grant Fellowship through NOAA,
National Sea Grant College Program, De-
partment of Commerce, under grant num-
ber NA89AA-D-SG138, project number
USDC EG-10-8B, through the California
Sea Grant College, and in part by the Cal-
ifornia State Resources Agency. The U.S.
Government is authorized to reproduce and
distribute for governmental purposes.

Literature Cited

Alvarino, A. 1967. Bathymetric distribution of Chae-
tognatha, Siphonophorae, Medusae, and Cte-
nophorae off San Diego, California.— Pacific
Science 21:474-485.

Bigelow, H. B. 1913. Medusae and Siphonophorae
collected by the U.S. Fisheries steamer "Alba-
tross" in the northwestern Pacific, 1906.— Pro-
ceedings of the United States National Museum
44:1-119, 6 pis.

Bonnett, R., E. J. Head, & P. J. Herring. 1979. Por-
phyrin pigments of some deep-sea medusae. —
Journal of the Marine Biological Association of
the United Kingdom 59:565-573.

Childress, J. J., A. T. Barnes, L. B. Quetin, & B. H.
Robison. 1978. Thermally protecting cod ends
for recovery of living deep-sea animals. —Deep-
Sea Research 25:419^22.

Fox, D. L., & C. F. A. Pantin. 1941. The colours of
the plumose anemone Metridium senile (L.).—

Philosophical Transactions of the Royal Soci-
ety, Series B 230:415-450.

Herring, P. J. 1972. Porphyrin pigmentation in deep-
sea medusae.— Nature 238:276-277.

Kramp, P. L. 1947. Medusae. Part III. Trachylina
and scyphozoa.— Danish Ingolf-Expedition Re-
ports 5:1-66, 6 pis.

. 1961. Synopsis of the medusae of the world.—

Journal of the Marine Biological Association of
the United Kingdom 40: 1-469.

. 1968. The hydromedusae of the Pacific and

Indian Oceans.— "Dana"-Reports 72:1-200.

Larson, R. J., & G.R.Harbison. 1990. Medusae from
McMurdo Sound, Ross Sea including the de-
scriptions of two new species, Leuckartiara
brownei and Benthocodon hyalinus. —Folar Bi-
ology 11:19-25.

Segura-Puertas, L. 1984. Morphology and zoogeog-
raphy of medusae (Cnidaria: Hydrozoa and Scy-
phozoa) from the eastern tropical Pacific— In-
stituto de Ciencias del Mar y Limnologia
Universidad Nacional Autonoma de Mexico
Publicaciones Especiales 8:1-320.

Totton, A. K. 1965. A synopsis of the Siphonophora.
British Museum (Natural History), London 230
pp., 40 pis.

Wimpenny, R. S. 1966. The plankton of the sea.
American Elsevier, New York, 426 pp.

Marine Science Institute, University of
Cahfomia, Santa Barbara, California 93106,
U.S.A.

PROC. BIOL. SOC. WASH.
106(1), 1993, pp. 195-203

A NEW SPECIES OF SIBOPATHES
(CNIDARIA: ANTHOZOA: ANTIPATHARIA: ANTIPATHIDAE)

FROM THE GULF OF MEXICO

Dennis M. Opresko

Abstract. —A new species of antipatharian (Cnidaria: Anthozoa: Antipathar-
ia), Sibopathes macrospina, is described from the Gulf of Mexico. The species
differs from the only other species in the genus, S. gephura van Pesch, by having
longer spines and anterior pinnules positioned slightly above, rather than below,
the adjacent lateral pinnules. The taxonomic relationships of Sibopathes with
Cladopathes, Hexapathes, Taxipathes, Bathypathes, Schizopathes, and Par-
antipathes are discussed.

An unusual antipatharian coral was re-
cently collected in the northern Gulf of
Mexico during submersible investigations
conducted by Dauphin Island Sea Lab un-
der the direction of T. Hopkins. This coral
proved to be related to a species in the genus
Sibopathes previously known only from the
Indo-Pacific. Comparisons with the type
material of the Pacific species have revealed
that the Gulf specimen represents a new
species. The holotype has been deposited in
the National Museum of Natural History,
Smithsonian Institution, Washington, D.C.
(USNM).

Family Antipathidae

Subfamily Cladopathinae

Genus Sibopathes van Pesch, 1914

Type species.— Sibopathes gephura van
Pesch, 1914:203-205, pi. VI, figs. 3, 5-6,
15; pi. VII, fig. 3. (Type locality: Indo-Pa-
cific, east of Timor, Indonesia, 8° 17. 4'S,
127°30.7'E, 1224 m, Siboga Stn. 280).

Diagnosis. —VoXyps transversely elongat-
ed with six reduced primary mesenteries,
no secondary mesenteries, and no actino-
pharynx.

Sibopathes macrospina, new species
Figs. 1, 2, 3, 4A-C, E

Holotype. - USNM 91417. Gulf of Mex-
ico, off Alabama, 29°09'30"N, 88°01'10"W,

UNCW 9119, Johnson Sea Link Stn. JSL
3097, 26 Aug 1991, 489-559 m, Coll. W.
W. Schroeder.

Diagnosis. —CovdiWum. branched and pin-
nulate (Fig. 1); pinnules simple, 1-2 cm long,
arranged in four longitudinal rows and
grouped in alternating pairs with each pair
consisting of one lateral and one anterolat-
eral pinnule (Fig. 2A, B). Spines simple, tri-
angular, smooth, 0.07-0.12 mm in height
(Fig. 3); arranged in longitudinal rows; from
four to six spines per millimeter in each row.
Polyps elongated, about 2 mm in transverse
diameter (Fig. 4A); in a single row with four
polyps per centimeter.

Description. —Holotype about 36 cm high
and 16 cm wide (Fig. 1). Basal holdfast ab-
sent; lowermost part of the stem about 2
mm in diameter. Corallum branched irreg-
ularly to the fourth order with branches 0.5-
2.0 cm apart. Branches and branchlets
straight or curved, and directed upward
(branch angle 60-75°). Overall branching of
corallum planar with branchlets arising pri-
marily from sides of lower order branches,
occasionally from front (corresponding to
polyp-bearing side of stem) and rarely from
back.

Stem and branches pinnulated; branch-
lets developing from elongated pinnules
which become pinnulated in turn. Pinnules
simple (without subpinnules), relatively
straight and stiff; not strictly uniform in size,

196

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. I. Sibopathes macrospina. Holotype, USNM 91417; height about 36 cm.

number, or arrangement on branchlets but erally and laterals more posterolaterally.

usually in four longitudinal rows with two Anterolaterals absent near base of some

nearly opposite lateral or posterolateral rows branchlets. Pinnules 1.8-3.5 mm apart

and two anterolateral rows (Fig. 2 A). An- (mean 2.75 mm, n = 20) with four or five

terolaterals occasionally located more lat- per centimeter in each longitudinal row.

VOLUME 106, NUMBER 1

197

Fig. 2. A-B. Sibopathes macrospina, holotype. USNM 91417. A, Cross sectional view of branchlet sho\\ing
arrangement of pinnules around axis: scale equals 0.2 mm. B. Frontal (polypar) view of branchlet sho\\ing
arrangement of pinnules; scale equals 0.4 mm.

Pinnules in lateral rows arranged alternate-
ly; anterolateral pinnules placed 0.4-0.7 mm
above (distal to) adjacent lateral pinnule.
Overall, pinnules form alternating pairs,
each consisting of one lateral and one slight-
ly higher anterolateral member (Fig. 2B).
Lateral pinnules 0.9-1.9 cm long (mean 1.36
cm. « = 14) and 0.20-0.32 mm in diameter
(near base). Axial canal of pinnules 0.06-
0.08 mm in diameter. Pinnules project up-

ward slightly forming angle of 75° or more
with branchlet. Anterolateral pinnules usu-
ally shorter, but occasionally as long as or
longer than adjacent lateral pinnules. Ad-
jacent pinnules from different branches
anastomose near base of corallum.

Spines usually simple, but occasionally
bifid: triangular, acute, and smooth-sur-
faced (Fig. 3); small and relatively unde-
veloped at tip of pinnules (Fig. 4B) but up

198

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3.
0.1 mm.

Sibopathes macrospina, holotype, USNM 91417. Stereo SEM of pinnule with axial spines; scale equals

to 0.12 mm high (from midpoint of base to
apex) on lower portion of pinnules (Fig. 4C).
Spines unequally developed around circum-
ference of pinnules; largest spines usually
on front or polyp side of axis but, in places
nearly equal in size on three sides, leaving
abpolypar side with smallest spines. Poly-
par spines usually extending out perpendic-
ular to pinnular axis, but occasionally di-
rected distally or proximally. Abpolypar
spines 0.02-0.05 mm high. Spines on stem
and larger branches rarely more than 0.04
mm high. Spines on pinnules arranged in
longitudinal rows; three to five rows visible
in lateral view (excluding rows in which
spines only partially visible). Distance be-
tween adjacent spines in each row 0.20-0.36
mm; generally from four to six spines per
millimeter in each row.

Polyps elongated along transverse axis;

distance from distal edge of distal lateral
tentacles to proximal edge of proximal lat-
eral tentacles 1.8-2.3 mm (Fig. 4A). Polyps
arranged in a single row with about four
polyps per centimeter; interpolypar space
0.5-0.8 mm. Peristomal folds absent. Oral
cone usually elongated transversely 0.28-
0.44 mm; sagittal diameter 0.14-0.28 mm.
Tentacles 0.12-0.25 mm in length. Ova
present in lateral sections of coelenteron (Fig.
4E). Polyps with only six rudimentary and
incomplete mesenteries, no actinopharynx,
and no evidence of mesogloeal partitions
separating coelenteron into central and lat-
eral chambers.

Etymology.— The; specific name is de-
rived from the Latin "macros" and "spina"
in reference to the relatively large spines on
the pinnules.

Comparisons.— T\As species is very sim-

VOLUME 106, NUMBER 1

199

Fig. 4. A-C. Sibopathes macrospina, holotype, USNM 91417. A, Stereo SEM of polyp. B, Distal end of
pinnule. C, Center of pinnule. D. S. gephura van Pesch, holotype, Siboga Stn. 280, center of pinnule. E. S.
macrospina, holotype, USNM 91417, polyp with ova. Scale in A and E equals 0.5 mm; scale in B (also for C
and D) equals 0.1 mm.

200

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table L— Morphometries for Sibopathes gephura and S. macrospina.

Character

S. gephura

S. macrospina

Corallum

Height (cm)

Basal stem diameter (mm)
Highest order of branching
Rows of pinnules

Pinnules

Length of lateral pinnules (cm)
Basal diameter (mm)
Diameter of central canal (mm)
Distance apart in one row (mm)
Angle between lateral and

anterior pinnules
Distal angle with stem ^

Spines
Height, polypar (mm)
Height, abpolypar (mm)
Distance apart in one row (mm)
Average number per millimeter
Number of rows on pinnules (one view)

Polyps

Length (mm)

Interpolypar space (mm)

Number per centimeter

Length of tentacles (mm)

Oral cone, transverse diameter (mm)

Oral cone, sagittal diameter (mm)

-0.85

-2

4^-5

2-4"

0.7-2.2

0.9-1.9

0.22-0.28

0.20-0.32

0.10-0.16

0.06-0.08

2.2-3.0

1.8-3.5

45-90°

30-45°

-60°

>75°

0.03-0.04

0.07-0.12

<0.03-0.04

0.02-0.05

0.12-0.22

0.20-0.36

5-7

4-6

3-4

3-5

2.0-2.5

1.8-2.3

0.5-1.1=

0.5-0.8

3-3.5

3.5-4

<0.5

0.12-0.25

0.6-0.87^^

0.28-0.44

0.2

0.14-0.28

^ specimen may only be a branch from a larger colony.

*' Most common condition.

'^ Maximum value reported by van Pesch (1914).

ilar to Sibopathes gephura van Pesch (1914).
Both species have a branched pinnulated
corallum with simple pinnules arranged for
the most part in four longitudinal rows. The
major differences between the two species
involve the orientation and arrangement of
the pinnules and the size of the spines (Table
1). Pinnules on S. gephura appear somewhat
less regular in length than those on S. mac-
rospina, although in both cases they reach
about the same maximum size of about 2
cm before becoming pinnulated branchlets.
In S. gephura the pinnules tend to be more
curved and more distally directed (relative
to the branch), whereas in S. macrospina
they are straight, stiff, and extend out more
horizontally. The central axial canal in the

pinnules of S. macrospina is smaller in di-
ameter than that in S. gephura (Table 1).

Van Pesch (1914) described the arrange-
ment of the pinnules in S. gephura as being
in four longitudinal rows, two lateral, one
anterior, and one posterior; however, in re-
examining the type specimen it was found
that in places the rows are also arranged
biserially, that is, with two rows on the right
and two on the left. In such cases the pin-
nules are placed in alternating pairs with the
front or anterolateral pinnule of each pair
located slightly below the adjacent lateral
(as viewed from the front or polyp-side of
the corallum). This arrangement differs from
that found in S. macrospina where the an-
terolaterals are placed slightly above the ad-

VOLUME 106, NUMBER 1

201

jacent laterals. Furthermore, on the holo-
type of 5. gephura a few additional pinnules
were found representing a fifth longitudinal
row. These occurred in front of and slightly
below the adjacent anterolateral pinnules
such that each group of three pinnules
formed a descending series going from the
side to the front of the branch (as viewed
from the polyp-side of the corallum). In
contrast, in S. macrospina the pattern is re-
versed, with the pinnules in each group
forming an ascending series (Fig. 2B).

In both species the spines are simple, tri-
angular and acute; however, in S. gephura
they measure not more than 0.04 mm in
height (Fig. 4D), whereas in S. macrospina
they attain a maximum size of 0. 12 mm. In
addition, the spines in S. macrospina are
spaced farther apart than those in S. ge-
phura (Table. 1). In both species the spines
are largest on the side of the axis bearing
the polyps; however, this is not easily seen
in S. gephura because the abpolypar spines
are only slightly smaller than the polypar
spines.

There are no major differences in the ex-
ternal morphology of the polyps of the two
species except that the polyps in S. macro-
spina are slightly smaller and closer together
and have shorter tentacles than those in S.
gephura (Table 1). These differences are not
very great and may be due, in part, to state
of preservation. In both species the polyps
are arranged in a single row; on the upper
side of the pinnules in S. gephura, and on
the upper and occasionally front and lower
sides of the pinnules in S. macrospina. In
neither species is there any sign of peristo-
mal folds dividing the polyps into central
and lateral sections.

Although the specimen of S. macrospina
was not originally fixed for histological ex-
amination, several polyps were removed
from the corallum, sectioned, stained, and
examined microscopically. Internal features
of the polyp were difficult to distinguish be-
cause of the poor condition of the tissue;
however, as in the case of 5. gephura, there

was no sign of an actinopharynx, and the
mesenteries were rudimentary and ap-
peared to be no more than six in number.
In addition, there was no indication of me-
sogoeal septa separating the coelenteron into
central and lateral chambers. Van Pesch re-
ported the same condition in S. gephura.
Although nematocysts could not be seen in
the polyps of S. macrospina, they were re-
ported as occurring in batteries on the ten-
tacles of 5. gephura polyps (van Pesch 1914).

Discussion

The two species of Sibopathes differ from
all other antipatharians in that the polyps
lack an actinopharynx and, consequently,
the six mesenteries are incomplete. Van
Pesch (1914) created the subfamily Homoe-
otaeniales to contain Sibopathes and a sec-
ond genus Cladopathes Brook, whose pol-
yps also have only six mesenteries; however,
in Cladopathes the mesenteries are attached
to a well-developed actinopharynx. Clado-
pathes was originally included with Schi-
zopathes, Bathypathes, and Taxipathes, in
the subfamily Schizopathinae by Brook
(1889). Polyps in the latter three genera have
ten mesenteries, six primary and four sec-
ondary. In establishing the Schizopathinae,
Brook considered the number of mesenter-
ies to be of secondary importance when
compared to the transverse elongation of
the polyps into what he interpreted as "di-
morphic" structures consisting of "gastro-
zooids" and "gonozooids." In describing the
polyps ofSchizopathes, Brook reported that
the "zooids" were isolated from one anoth-
er externally by peristomal folds and inter-
nally by mesogloeal partitions extending
down from the upper, interior surface of the
coelenteron. These features were not spe-
cifically described for other genera in the
subfamily, and Brook's own illustrations in-
dicate that peristomal folds are not typical
of Bathypathes or Taxipathes polyps. Con-
sequently, the only remaining diagnostic
character of the Schizopathinae is the ex-

202

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

treme transverse elongation of the polyp. In
1896, Schultze proposed a new classifica-
tion of the Antipathidae based on the num-
ber of mesenteries, rather than on the pre-
sumed dimorphism of the polyps. He
therefore established a separate subfamily,
the Hexamerota for Cladopathes. The sub-
family was renamed the Cladopathinae by
Kinoshita in 1910, who added to it a new
monotypic genus Hexapathes. The polyps
of//, heterosticha reportedly contain an ac-
tinopharynx, six complete primary mesen-
teries and no secondaries; however, the
skeletal morphology is quite different from
that of Cladopathes (and Sibopathes); in-
stead, it is similar to that of species of Bath-
ypathes. Thus, van Pesch's Homoeotaeni-
ales is equivalent to the Cladopathinae and
contains those genera having polyps with
six mesenteries, but in which an actino-
pharynx may be present (Cladopathes and
Hexapathes) or absent {Sibopathes).

In describing Sibopathes gephura, van
Pesch (1914) noted that many of the ana-
tomical features of the polyps were sugges-
tive of a very primitive condition. These
features included: absence of an actino-
pharynx and secondary mesenteries; small
size of the sagittal mesenteries and their oc-
currence only near the base of sagittal ten-
tacles; thinness of the mesogloea; presence
of epidermally derived cells within the me-
sogloea; very wide connection between the
axis epithelium and the body wall; occur-
rence of deeply staining gland cells through-
out the gastrodermis; and absence of mes-
enteric filaments (defined by van Pesch as
extensions of the mesenteries at the aboral
edge of the actinopharynx). Furthermore,
van Pesch noted that the transverse mes-
enteries disappeared at the junction of the
central and lateral parts of the polyp, but
reappeared, with a club-shaped edge, in the
lateral chambers. Van Pesch did not con-
sider the club-shaped edge of the transverse
mesenteries to represent a mesenteric fila-
ment.

If Schultze' s classification is followed, it
could be argued that the absence of an ac-
tinopharynx and the presence of other prim-
itive characters would warrant the taxo-
nomic recognition of Sibopathes at least at
the same level as the Cladopathinae, and
perhaps even at the family level (with the
recent removal of the Dendrobrachiidae, the
order currently contains only the single fam-
ily Antipathidae, see Opresko & Bayer
1991). However, if one considers the pos-
sibility that the characters oi Sibopathes are
secondarily derived as a result of the ex-
treme transverse elongation of the polyp (a
character which in itself would be difficult
to view as a primitive state), and if one eval-
uates overall similarities in external mor-
phology, then it might be argued that Si-
bopathes is indeed related to Cladopathes
but not to Hexapathes. Like Sibopathes,
Cladopathes is quasi-sympodial (i.e., with-
out a single continuous stem), multi-
branched, and pinnulated. The pinnules are
simple or forked and arranged in three or
four longitudinal rows "showing a subspiral
arrangement." In contrast, Hexapathes het-
erosticha Kinoshita is clearly monopodial
with two rows of lateral pinnules and one
row of anterior pinnules, a pattern identical
to that occurring in Bathypathes lyra Brook.

Sibopathes and Cladopathes also show
external similarities to the genera Taxi-
pathes and Parantipathes. All four genera
(Sibopathes, Taxipathes, Parantipathes, and
Cladopathes) have transversely elongated
polyps of relatively small size (generally <3
mm). In contrast, the polyps of Hexapathes
heterosticha were described as being 5-9 mm
long, and this is also the case for some spe-
cies of Bathypathes. Taxipathes and Par-
antipathes also have pinnules arranged in
longitudinal rows and alternating semispiral
groupings, but they differ from Sibopathes
and Cladopathes in having secondary mes-
enteries in the polyps. However, this differ-
ence is certainly not as significant as that
separating Sibopathes from Cladopathes

VOLUME 106, NUMBER 1

203

(i.e., the absence of an actinopharynx). Fur-
ther analysis may eventually show that these
four genera form a natural assemblage.

Acknowledgments

The author wishes to thank S. Cairns for
his helpful suggestions and for taking the
scanning electron micrographs; T. Hopkins
and W. W. Schroeder of the University of
Alabama, Marine Environmental Sciences
Consortium, Dauphin Island Sea Lab for
providing the specimen of S. macrospina
which was collected during field studies con-
ducted under NSF EPSCOR Grant No. R 1 1 -
8996 1 52 and NOAA/NURC/UNCW Grant
No. NA88AA-D-UR004; R. W. M. van
Soest of the Riksmuseum van Natuurlijke
Historic in Amsterdam for the loan of the
type specimen of S. gephura; T. Bayer, C.
Bast, and M. Bogle for reviewing the manu-
script; S. Braden of the Smithsonian Insti-
tution for preparing the samples for the
scanning electron microscope (SEM); and J.
Wesley of Oak Ridge National Laboratory

for preparing the histological sections. This
work was supported in part by the Smith-
sonian Institution and by Oak Ridge Na-
tional Laboratory, Oak Ridge, Tennessee.

Literature Cited

Brook, G. 1889. Report on the Antipatharia. — Re-
ports of the Scientific Results of the Voyage of
the Challenger. —Zoology 32:5-222.

Kinoshita, K. 1910. On a new antipatharian //e-xa-
pathes heterosticha n. gen. and n. sp.— Anno-
tationes Zoologicae Japonenses 7:231-234.

Opresko, D. M., & F. M. Bayer. 1991. Rediscovery
of the enigmatic coelenterate Dendwbrachia,
(Octocorallia: Gorgonacea) with descriptions of
two new species.— Transactions of the Royal
Society of South Australia 115:1-19.

Schultze, L. S. 1896. Beitrag zur Systematik der An-
tipatharien.— Abhandlungen der Senckenber-
gischen naturforschenden Gesellschaft 23: 1^0.

Van Pesch, A. J. 1914. The Antipatharia of the Siboga
Expedition.— ^z^c^a-Expeditie Monographe 17:
1-258.

Health and Safety Research Division, Oak
Ridge National Laboratory, P.O. Box 2008,
Oak Ridge, Tennessee 37831-6050, U.S.A.

PROC. BIOL. SOC. WASH.
106(1), 1993, pp. 204-205

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London, SW7 5BD, U.K.
Tel. 071-938 9387

Applications published in the Bulletin of Zoological Nomenclature

The following Applications were published on 30 September 1992 in Vol. 49,
Part 3 of the Bulletin of Zoological Nomenclature. Comment or advice on these
Applications is invited for publication in the Bulletin, and should be sent to the
Executive Secretary, I. C.Z.N. , % The Natural History Museum, Cromwell Road,
London SW7 5BD, U.K.

Case No.

2806 Zanclea costata Gegenbaur, 1856 (Cnidaria, Hydrozoa): proposed conser-
vation of both generic and specific names.

2827 Gebia major capensis Krauss, 1843 (currently Upogebia capensis; Crustacea,

Decapoda): proposed replacement of neotype, so conserving the
usage of capensis and also that of G. africana Ortmann, 1894 (cur-
rently Upogebia africana).

2828 Podisus Herrich-Schaeffer, 1851 (Insecta, Heteroptera): proposed conserva-

tion of P. vittipennis Herrich-Schaeffer, 1851 as the type species.
2795 ANTHRiBiDAE Billbcrg, 1820 (Insecta, Coleoptera): proposed precedence over
CHORAGIDAE Kirby, 1819.

2811 Catocala connubialis Guenee, 1852 (Insecta, Lepidoptera): proposed con-

servation of the specific name.
2793 METOPiiNAE Foerster, 1868 (Insecta, Hymenoptera), metopiini Raffray, 1904
(Insecta, Coleoptera), and metopiini Townsend, 1908 (Insecta, Dip-
tera): proposed removal of homonymy.

2812 Acamptopoeum Cockerell, 1905 (Insecta, Hymenoptera): proposed desig-

nation of Camptopoeum submetallicum Spinola, 1851 as the type

species.
2792 Cynolebias opalescens Myers, 1942 and Cynolebias spendens Myers, 1942

(Osteichthyes, Cyprinodontiformes): proposed conservation of the

specific names.
2601 Filimanus Myers, 1936 (Osteichthyes, Perciformes): proposed designation

of Filimanus perplexa Feltes, 1991 as the type species.
2821 Rana megapoda Taylor, 1942 (Amphibia, Anura): proposed conservation of

the specific name.
2382 Megophrys montana Kuhl & van Hasselt, 1 822 (Amphibia, Anura): proposed

placement of both the generic and specific names on Official Lists,

and Leptobrachium parvum Boulenger, 1893 (currently Megophrys

parva): propposed conservation of the specific name.
2802 Anisolepis grilli Boulenger, 1891 (Reptilia, Squamata): proposed conservation

of the specific name.

VOLUME 106, NUMBER 1 205

Opinions published in the Bulletin of Zoological Nomenclature

The following Opinions were published on 30 September 1992 in Vol. 49, Part
3 of the Bulletin of Zoological Nomenclature.

Opinion No.

1689. Epizoanthus Gray, 1867 (Cnidaria, Anthozoa): conserved.

1 690. Helix (Helicigona) barbata Ferussac, 1832 (currently Lindholmiola barbata;

Mollusca, Gastropoda): lectotype designation confirmed.

1691. Polygyra Say, 1818 (Mollusca, Gastropoda): Polygyra septemvolva Say, 1818

designated as the type species, and polygyridae Pilsbry, 1895 given
precedence over mesodontidae Tryon, 1866.

1692. Phyllodoce Lamarck, 1818 and Polyodontes de Blainville, 1828 (Annelida,

Polychaeta): conserved.

1693. Coccinella undecimnotata Schneider, [1792] (currently Hippodamia (Sem-

iadalid) undecimnotata; Insecta, Coleoptera): specific name con-
served.

1694. Rhinapion Beguin-Billecocq, 1905 (Insecta, Coleoptera): conserved.

1695. Acanthophthalmus van Hasselt in Temminck, 1824 (Osteichthyes, Cyprin-

iformes): not conserved.

1 696. HYDROBATiDAE Mathews, 1912 (1865) (Aves, Procellariiformes): conserved.

PROC. BIOL. SOC. WASH.

106(1), 1993, p. 206

REVIEWERS

The following people reviewed manuscripts for the Proceedings in 1 992. P. Alderslade, F. Alvarez,
A. Alvariiio, W. D. Anderson, Jr., M. Aoki, A. Asakura, K. Baba, I. Ball, S. Bandoni, A. M. Bauer,
F. M. Bayer, J. A. Blake, T. E. Bowman, R. Brinkhurst, J. Burch, S. D. Cairns, E. Campos, M. D.
Carleton, R. Carney, K. Carpenter, J. Chess, K. Coates, C. O. Coleman, B. B. Collette, M. R.
Cooper, J. S. Costlow, Jr., N. Cumberlidge, H.-E. Dahms, D. Davis, M. Dillon, M. Dojiri, R. C.
Dowler, L. H. Emmons, K. Fauchald, D. L. Felder, R. M. Feldmann, G. Fenton, K. Fitzhugh, J.

F. Fitzpatrick, O. S. Flint, Jr., J. Friend, R. J. Gagne, A. L. Gardner, S. Gelder, A. C. Gill, C. J.
Glasby, W. Goldberg, I. Goodbody, J. Goy, M. Grasshoff, G. R. Graves, D. E. Hahn, R. Hanley,
E. Harada, W. Hartman, P. C. Heemstra, G. Hendler, D. Hendrickson, B. Hilbig, S. Hiruta, J.-s.
Ho, H. H. Hobbs, Jr., D. Holdich, J. Holsinger, R. W. Holzenthal, T. S. Hopkins, G. D. Johnson,
N. K. Johnson, J. K. Jones, Z. Kabata, Y. Kikuchi, G. L. Kirkland, Jr., G. Klassen, L. S. Komicker,
R. Kropp, J. Kudenov, S. Lanyon, R. Larson, D. Laubitz, R. Lemaitre, W. G. Lyons, E. Macpherson,
L. Madin, C. Magalhaes, R. B. Manning, J. W. Martin, W. N. Mathis, J. Mauchline, D. McKinnon,

G. A. de Melo, H. Michel, M. Milligan, C. Mills, W. L. Minckley, R. F. Modlin, R. D. Mooi, J.
C. Morse, A. Muir, M. Murano, T. Newberry, P. K. L. Ng, J. L. Norenburg, R. D. Owen, L. R.
Parenti, K. Parkes, C. Patterson, J. L. Patton, D. L. Pawson, T. Perkins, M. H. Pettibone, F. Pleijel,
G. Pohle, W. F. Presch, G. Pretzmann, J. Pruski, A. Rea, J. W. Reid, C. B. Robbins, S. de A.
Rodrigues, G. Rodriguez, C. F. E. Roper, F. Rowe, K. Ruetzler, C. W. Sabrosky, J. Sieg, T. Simpson,
R. Sluys, A. E. Smalley, J. Smith III, G. S. Steyskal, J. H. Stock, F. C. Thompson, F. G. Thompson,
M. Thurston, R. Toll, S. Tyler, J. L. Villalobos, R. Vonk, N. Voss, E. Walker, V. Wallach, L. A.
Ward, M. J. Wetzel, M. K. Wicksten, A. B. Williams, J. T. Williams, K. Wittmann, K. Wouters,
H. Yeatman, R. Young, W. Zeider, R. Zottoli, G. R. Zug.

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CONTENTS

On the identity of Echimys didelphoides Desmarest, 1817 (Mammalia: Rodentia: Echimyidae)

Louise H. Emmons 1

A new subspecies of pocket gopher (Geomys) from Texas (Mammalia: Rodentia: Geomyidae)

Michael J. Smolen, Richard M. Pitts, and John W. Bickham 5

Identification of bird subfossils from cave surface deposits at Anjohibe, Madagascar, with a

description of a new giant Coua (Cuculidae: Couinae)

Steven M. Goodman and Florent Ravoavy 24
Two new species of blind snake, genus Typhlops (Reptilia: Squamata: Typhlopidae), from the

Philippine Archipelago Addison H. Wynn and Alan E. Leviton 34

A new species of dimorphic tree frog, genus Hyla (Amphibia: Anura: Hylidae), from the Vaupes

River of Colombia William F. Pybum 46

Description of the advertisement call and resolution of the systematic status of Leptodactylus

gracilis delattini Muller, 1968 (Amphibia: Leptodactylidae)

Linnette Garcia Perez and W. Ronald Heyer 5 1
Prochilodus britskii, a new species of prochilodontid fish (Ostariophysi: Characiformes), from

the rio Apiaca, rio Tapajos system, Mato Grosso, Brazil Ricardo M. C. Castro 57

A new Devonian ophiuroid (Echinodermata: Oegophiurida) from New York state and its

bearing on the origin of ophiuroid upper arm plates Frederick H. C. Hotchkiss 63

Erythrosquilloidea, a new superfamily, and Tetrasquillidae, a new family of stomatopod crus-
taceans Raymond B. Manning and David K. Camp 85
Systematics and taxonomic remarks on Pinnotheres muliniarum Rathbun, 1918 (Crustacea:

Brachyura: Pinnotheridae) Ernesto Campos 92

Anomoeomunida, a new genus proposed for Phylladiorhynchus caribensis Mayo, 1972 (Crus-
tacea: Decapoda: Galatheidae) Keiji Baba 102
Two new species of NeocalUchirus from the Caribbean Sea (Crustacea: Decapoda: Callianas-

sidae) Raymond B. Manning 106

Caprella arimotoi, a new species (Crustacea: Amphipoda: Caprellidea) from the Seto Inland

Sea, Japan Ichiro Takeuchi 1 1 5

A new species of Kalliapseudes (Crustacea: Tanaidacea: Kalliapseudidae) from Trinidad

Roger N. Bamber 1 22
Enterocola africanus, a new species (Copepoda: Ascidicolidae) associated with a compound

ascidian Synoicum species from North Africa (Strait of Gibraltar)

Pablo J. Lopez-Gonzalez, Mercedes Conradi, and J. Carlos Garcia-Gomez 131
New species and new records of the genus Elaphoidella (Crustacea: Copepoda: Harpacticoida)

from the United States Janet W. Reid and Teruo Ishida 137

New genera and species of deep-sea polychaetes of the family Nautiliniellidae from the Gulf

of Mexico and the eastern Pacific James A. Blake 147

Taxonomy of European species of Amphiduros and Gyptis (Polychaeta: Hesionidae)

Fredrik Pleijel 158
A new genus of hydrobiid snails (Mollusca: Gastropoda: Prosobranchia: Rissooidea) from

northern South America Robert Hershler and France Velkovrh 182

Vampyrocrossota childressi, a new genus and species of black medusa from the Bathypelagic

zone off California (Cnidaria: Trachy medusae: Rhopalonematidae) Erik V. Thuesen 1 90

A new species of Sibopathes (Cnidaria: Anthozoa: Antipatharia: Antipathidae) from the Gulf

of Mexico Dennis M. Opresko 195

International Commission on Zoological Nomenclature: Applications and Opinions 204

Reviewers -1992 - 206

PROCEEDINGS

OF THE

BIOLOGICAL SOCIETY

WASHINGTON

VOLUME 106
1 1 JUNE 1993

NUMBER 2

ISSN 0006-324X

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PROC. BIOL. SOC. WASH.

106(2), 1993, pp. 207-220

REVISED CLASSIFICATION AND PHYLOGENETIC
HYPOTHESIS FOR THE
ACANTHOSTOMINAE LOOSS, 1899
(DIGENEA: OPISTHORCHIFORMES: CRYPTOGOJ^^^JiSD^fe^

Daniel R. Brooks and Barbara Holcman

JUN

2 1W911

Abstract.— SpecimQns of an acanthostome digenean originally identified a^o
Acanthostomum scyphocephalum and later transferred to Timonietitt-M^ ^e^
scribed and named as a distinct species of Timoniella. Acanthostomum scy-.
phocephalum sensu strictu is included in an updated phylogenetic analysis of
the acanthostome digeneans. The new analysis differs from an earher one by
Brooks (1980) by allowing reversals (Wagner criterion vs. Camin-Sokal crite-
rion), producing a more parsimonious representation of character data; no
transformation series needed re-polarization. Acanthostomum scyphocephalum
is a member of the clade containing all other species of Acanthostomum oc-
curring in North, Central and South America. Acanthostomum is paraphyletic
if Atrophecaecum is excluded from it; accordingly, the two genera are synon-
ymized. No other changes from the hypothesis of Brooks (1980) resulted. The
resulting annotated phylogenetic classification, with synapomorphic diagnoses,
includes Acanthostomum as the sister-group of Caimanicola, Proctocaecum as
their sister-group, the monotypic Gymnatrema as their sister-group, and Ti-
moniella as the basal sister-group. Four new subgenera are proposed, one in
Timoniella, one in Proctocaecum, and two in Acanthostomum.

The acanthostome digeneans (Opisthor-
chiformes: Cryptogonimidae: Acanthosto-
minae) inhabit a variety of piscivorous poi-
kilotherm amniote vertebrates throughout
the tropical and subtropical regions of the
world. Brooks (1980) provided the first phy-
logenetic systematic analysis of the acan-
thostomes. He recognized six genera, Ti-
moniella Rebecq, 1960, Proctocaecum
Baugh, 1951 , Gymnatrema MorozoY, 1955,
Caimanicola Teixeira de Freitas & Lent,
1938, Acanthostomum Looss, 1899 and
Atrophecaecum Bhalerao, 1940, although
Acanthostomum had no synapomorphy to
support its recognition as a monophyletic
group.

Since that time, publications have de-
scribed two additional species and dis-
cussed their phylogenetic relationships
(Brooks & Caira 1982, Blair et al. 1988),

and have added data about host and geo-
graphic distributions, as well as valuable
taxonomic information (Ostrowski de Nu-
iiez 1984a, 1984b, 1986, 1987). This study
provides a description of a new species, and
incorporates the new data provided by Os-
trowski de Nunez into the phylogenetic data
base for the acanthostomes, resulting in an
updated phylogenetic hypothesis and an an-
notated classification with cladistic diag-
noses for all taxa.

Methods

In addition to the material specified in
Brooks (1980), Brooks & Caira (1982) and
Blair et al. (1988), we have examined the
following material loaned from Dr. Mar-
garita Ostrowski de Nunez: Acanthosto-
mum gnerii (19 specimens from Rhamdia

208

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

sapo, Laguna Chis-Chil, Prov. Buenos Ai-
res, Argentina); Acanthostomum megace-
tabulum (1 specimen from Drymarchon
corais melanurus, Villahermosa, Mexico);
Caimanicola marajoarum (14 specimens
from Crocodylus intermedius, Caracas,
Venezuela; 3 specimens from Paleosuchus
sp., Antioquia, Colombia); Caimanicola
brauni (13 specimens from Phrynops hilarii,
Buenos Aires, Argentina); Timoniella loossi
(3 specimens from Crocodylus intermedius,
Caracas, Venezuela); Acanthostomum sp. VI
of Ostrowksi de Nunez (1984b) (2 speci-
mens from Caiman fuscus, Rio Chagras,
Panama). All measurements are in nm un-
less otherwise noted. TBL = total body
length. Figures were drawn with the aid of
a drawing tube.

Results and Discussion

Timoniella ostrowskiae, new species
Figs. 1-3

Synonyms. —Acanthostomum scypho-
cephalum of Mane-Garzon & Gil, 1961; Ti-
moniella scyphocephala of Brooks, 1980.

Description.— (yydiSQd on holotype and 2
paratypes) Body 2.98-3. 1 5 mm long by 420-
540 wide at midbody. Tegument covered
with spines of uniform size. Oral sucker ter-
minal, bell-shaped, 410-517 long by 443-
517 wide, armed with single row of 23 spines
68-78 long by 20-25 wide. Pharynx 197-
205 long by 187-230 wide. Prepharynx 377-
426 long. Ratio of oral sucker width to pha-
ryngeal width 1:0.42-0.45. Ceca opening
separately at posterior end of body. Ace-
tabulum 1.22-1.47 mm from anterior end;
forebody 41-47% TBL. Acetabulum 140-
156 long by 131-147 wide; ratio of oral
sucker width to acetabulum width 1:0.27-
0.32. Testes tandem, intercaecal, near pos-
terior end of body; posttesticular space 5.2-
5.5% TBL. Anterior testis 187-205 long by
1 23-1 39 wide, posterior testis 1 80-22 1 long
by 1 15-123 wide. Male genitalia consisting
of coiled external seminal vesicle lying pos-
terodorsal to acetabulum and musculo-

glandular pars prostatica and ejaculatory
duct extending anteriorly dorsal to acetab-
ulum, opening into genital atrium. Gonotyl
lacking. Genital pore ventral, medial, im-
mediately preacetabular. Ovary 90-139 an-
terior to anterior margin of anterior testis,
123-139 long by 115-123 wide. Seminal
receptacle posterodorsal to ovary, between
ovary and anterior testis. Mehlis gland pres-
ent, Laurer's canal short. Uterine loops ex-
tending posteriorly to lateral margin of an-
terior testis, anteriorly to posterior margin
of seminal vesicle, occupying 38-43% TBL;
terminal portion of uterus opening into gen-
ital atrium. Vitellaria follicular, in two lon-
gitudinal extracecal fields extending from
level of posterior margin of seminal vesicle
to slightly posterior to anterior margin of
anterior testis. Eggs 25-27 long by 10-12
wide.

Type host.— Phrynops hilarii (Dumeril
and Bibron).

Type locality.— ^10 Negro, Paso de los
Toros, Departamento de Tacuarembo,
Uruguay.

Holotype. -U'^YC Helm. Coll. No.
11038. Paratypes: URFC Helm. Coll. No.
11039-11040.

Etymology.— ThQ species is named in
honor of Dr. Margarita Ostrowski de Nu-
nez, who first recognized its distinct iden-
tity.

Brooks (1980) was unable to locate the
holotype of ^. scyphocephalum Braun, 1 899
which had been collected in ^'Testudo ma-
tamata^' from southern Brazil. Based on ex-
amination of specimens collected in Phry-
nops hilarii from Uruguay (reported by
Maiie-Garzon & Gil 1961), Brooks (1980)
transferred the species to Timoniella be-
cause it possessed preovarian rather than
postovarian seminal receptacles, a unique
and unreversed synapomorphy that diag-
noses Timoniella among the acantho-
stomes. Ostrowski de Nunez (1986) found
the holotype of Acanthostomum scypho-
cephalum Braun, 1899, and showed that it
was a member of Acanthostomum. The

VOLUME 106, NUMBER 2

209

Figs. 1-3. Timoniella (Maillardiella) ostrowskiae. 1 . Ventral view of holotype. 2. Ootype region. 3. Terminal
genitalia.

210

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.— Homoplasious changes for 17 characters
of acanthostomes. Character consistency index = num-
ber of apomorphic states for each character divided by
total number of changes postulated on the phylogenetic
trees. For character identities, see Appendix 1 .

Char-

Character

acter

Figure #-Character

consistency

state

number on figure

index

1-0

9-36

50% (1/2)

2-0

9-38

50% (1/2)

6-0

5-3

50% (1/2)

7-0

4-17,5-12

50% (1/2)

9-0

8-10

9-1

4-10,5-1,6-3,7-5,8-7,9-32

33% (3/9)

10-0

6-1

10-1

5-2,7-4,8-2,8-17,9-29,9-35

17% (1/7)

11-1

5-4,9-21

50% (1/2)

12-1

5-5,6-11,8-14

33% (1/3)

13-1

5-6,6-6,8-13,9-22,9-33

33% (2/6)

14-2

5-13,6-7,8-1

14-3

5-14,9-19

14-5

4-12,6-4,8-6,9-26

57% (8/14)

15-1

5-8,8-18,9-24

50% (2/4)

16-0

7-3

16-1

4-20, 5-9, 6-8, 9-20, 9-28, 9-34

17% (1/7)

17-1

5-10,8-3

67% (2/3)

18-0

8-30

18-1

5-11,7-1,8-9

40% (2/5)

22-0

7-2,8-11

50% (2/4)

23-1

6-5,6-13,9-25

33% (1/3)

29-1

5-15,6-12,8-15

50% (2/4)

Specimens reported by Mafie-Garzon & Gil
(1961) therefore appear to represent a pre-
viously unnamed species of Timoniella,
which we formally described and named
above. Timoniella ostrowskiae is most
closely related to T. incognita Brooks, 1980,
T. loossi (Perez Vigueras, 1957) Brooks,
1980, T. absita Blair et al., 1988, and T.
w«am/ (Pelaez & Cruz, 1957) Brooks, 1980,
all of which lack gonotyls. Of those species,
T. ostrowskiae is the only one exhibiting
ceca opening separately at the posterior end
of the body; T. incognita has blindly-ending
ceca, and the remaining three species have
ceca opening into the excretory vesicle near
the posterior end of the body.

As indicated in the introduction, new in-
formation concerning acanthostomes has
accumulated since Brooks (1980) produced

the first phylogenetic systematic hypothesis
for the group. Brooks (1980) used the Ca-
min-Sokal (also known as the Weighted In-
variant Step Strategy [WISS]— Wiley et al.
1991) criterion, allowing no evolutionary
reversals, in producing a phylogenetic tree
for the acanthostomes. He found no syn-
apomorphy to support the monophyly of
Acanthostomum, nor any character that
could link Acanthostomum with either Cai-
manicola or Atrophecaecum, two genera
having synapomorphic support that were
shown forming an unresolved trichotomy
with Acanthostomum. In this study we re-
analyzed the data using the less restrictive
Wagner criterion (Wiley et al. 1991) after
checking character polarizations using es-
timates of higher-level digenean phyloge-
netic relationships (Brooks et al. 1985,
Brooks et al. 1989, Brooks & McLennan
1993) not available to Brooks (1980). We
found support for the original character po-
larizations of Brooks (1980, see also Brooks
& Caira 1982, Blair et al. 1988) but, as
would be expected, we also found more par-
simonious optimizations for some homo-
plasious characters when reversals were al-
lowed (for a summary of transformation
series, see Appendix 1; for a summary of
homoplasious changes, including reversals,
see Table 1). Based on the information pre-
sented by Ostrowski de Nunez (1986), we
included Acanthostomum scyphocephalum
in the phylogenetic framework; it appears
to be a member of the clade containing all
the other species of Acanthostomum occur-
ring in North, Central and South America
(Fig. 8). Most importantly, the new analysis
showed that Acanthostomum is paraphylet-
ic (Fig. 8), and should be combined with
Atrophecaecum (Figs. 8, 9).

Systematic theorists have begun investi-
gating the problems of providing robust
means for assessing the results of phyloge-
netic analyses (e.g., Archie 1989, Farris
1989, Sanderson & Donoghue 1989, Klas-
sen et al. 1991, Meier et al. 1991). These
studies have produced some interesting, and

VOLUME 106, NUMBER 2

211

in some cases initially counter-intuitive,
findings. For example, the minimum sig-
nificant value for the most commonly used
indicator, the consistency index (CI— Wiley
et al. 1991), drops as one adds taxa and
characters to a study; for example, a study
using 50 characters for 20 taxa and reporting
a CI of 65% may actually be more robust
than using 10 characters for 7 taxa and re-
porting a CI of 80%. This happens because
there are often apomorphic character
changes occurring once within a given taxon
that also occur once in another taxon. If the
scope of a study were expanded to include
both taxa, the estimate of homoplasy would
increase (and the CI would drop) even if the
hypothesized phylogenetic relationships of
the (now) subgroups did not change. Or, to
use current terminology, we would say that
a global phylogenetic analysis had discov-
ered homoplasy that the two local analyses
failed to recognize. In some cases such glob-
al homoplasy could affect the hypotheses of
relationships, so recognizing global homo-
plasy may play an important role in deter-
mining robust character polarities during
outgroup comparisons at the inception of a
phylogenetic analysis (e.g., Maddison et al.
1984, Wiley etal. 1991).

The acanthostomes provide an excellent
illustration of the relationship between local
and global parsimony considerations in
phylogenetic analysis. If we treat all the
acanthostomes as a single taxon, the con-
sistency index for the characters reported
herein is 50.5% (49 apomorphic character
states and 97 character changes). This value
is low for digeneans in general, the consis-
tency index based on global parsimony con-
siderations for all digeneans being approx-
imately 72% (Brooks & McLennan 1993).
By contrast, if we treat each of the clades
denoted as a genus separately, the estimated
homoplasy is generally much lower (100%
for the characters used at the generic level
by themselves, including the monotypic
Gymnatrema, for the species of Timoniella,
and for the species of Caimanicola, and

84.6% for the species of Proctocaecum), the
exception being Acanthostomum, for which
the consistency index based on local par-
simony considerations is 51%. This means
that although there is much homoplasy
among the acanthostomes as a whole, most
of it is dispersed among clades rather than
concentrated within clades. Thus, deter-
mination of plesiomorphic states by out-
group comparisons is not problematical, and
there is presently a single most parsimoni-
ous tree, for the group.

Mensural, or continuous variable, char-
acters are problematical for phylogenetic re-
construction. The determination of discrete
character states is often problematical for
such traits; options seem to range from con-
sidering almost every species autapomorph-
ic to recognizing very few states and there
is no consensus approach among phyloge-
neticists. Brooks (1980) adopted a conser-
vative approach to recognizing character
states based on his examination of available
specimens. Taking a conservative approach
to such traits often results in considerable
homoplasy. In the present study, the ho-
moplasy is distributed among half (1 7 of 34)
of the transformation series (1, 2, 6, 7, 9-
1 8, 22-23, and 29 in Appendix 1), 9 of which
(10, 12-13, 15-18, 22-23) are mensural in
nature. Of the 48 homoplasious character
transformations, 28 (58.3%) stem from the
mensural characters and 20 (41.7%) from
the qualitative traits; moreover, the men-
sural traits that show homoplasy have a
combined character consistency index of
28.6% (14/42), while the qualitative traits
that show homoplasy have a combined
character consistency index of 48.7% (19/
39) (Table 1). While suggestive, these data
are actually moot with respect to the ques-
tion of whether or not such characters pro-
vide adequate phylogenetic information,
because they exhibit high levels of homo-
plasy but do not support relationships that
are contradicted by non-mensural charac-
ters. A strong test of these characters re-
quires a search for intrinsically qualitative

212

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

characters whose apomorphic states sup-
port phylogenetic relationships that conflict
with the ones supported by the present data
base.

Conclusions

The changes discussed above are reflected
in the phylogenetic trees, cladistic diagno-
ses, and classification below (the monotypic
Gymnatrema has no separate cladogram).
In the following, italicized numbers refer to
synapomorphies listed by number on Fig.
4; other numbers refer to synapomorphies
listed by number on Figs. 5-8. Four new
subgenera are proposed. Each is named for
a digenean systematist who has advanced
our understanding of acanthostome rela-
tionships: Dr. Claude Maillard, University
of Montpellier, France; Dr. David Gibson,
British Museum (Natural History), London,
England; Dr. David Blair, James Cook Uni-
versity, Townsville, Australia; and Dr.
Robin Overstreet, Gulf Coast Marine Re-
search Laboratory, Ocean Springs, Missis-
sippi, USA.

Subfamily Acanthostominae Poche, 1926
(Fig. 4)

Diagnosis.— CvypXogomrmddiQ with ter-
minal oral sucker (7); armed with single row
of spines (2); preacetabular pit (5); genital
pore not in preacetabular pit {4)\ seminal
vesicle coiled posteriorly (5); suckerlike
gonotyl present (6).

Genus Timoniella Rebecq, 1960
(Fig. 5)

Diagnosis. —Acanthostominae preovari-
an seminal receptacle (7).

Subgenus Timoniella Rebecq, 1960

Diagnosis. — Timoniella having vitelline
follicles not extending anteriorly to poste-
rior margin of seminal vesicle* (1); length
of body occupied by uterine loops more than
50% TBL* (2); seminal vesicle not coiled

posteriorly* (3); prepharynx shorter than
pharynx* (4); ratio of oral sucker width to
pharyngeal width 1:0.25-0.40* (5); ratio of
body length to width averaging 7.5-15:1*
(6).

T. praeterita (Looss, 1901) Maillard, 1974

Diagnosis. -CycXoco^X (7); forebody 10-
20% TBL* (8); maximum body length 7-16
mm* (9).

T. imbutiformis (Molin, 1859) Brooks, 1980

Diagnosis. —Ratio of oral sucker : acetab-
ular width 1:0.8-1.3* (10); oral spines av-
eraging 25-30 in number* (11).

Subgenus Maillardiella, new subgenus

Diagnosis. — Timoniella lacking gono-
tyls* (12).

T. incognita Brooks, 1980

Diagnosis.— With characters of the sub-
genus.

Remarks: This species was originally re-
ported by Nasir (1974) as Acanthostomum
scyphocephalum inhabiting Caiman croco-
dilus crocodilus. According to Ostrowski de
Nunez (1984b), the acanthostomes reported
as Acanthostomum. scyphocephalum from
Drymarchon corais by Nasir (1974) may be
a still undescribed species.

T. ostrowskiae Brooks & Holcman, 1993

Diagnosis.— CQca opening separately at
posterior end of body* (13).

T. unami (Pelaez & Cruz, 1957) Brooks,
1980

Diagnosis. — Ceca opening into excretory
vesicle* (14); vitelline follicles extending
posteriorly to middle of posterior testis* (15).

T loossi (Perez Vigueras, 1957) Brooks,
1980

Diagnosis. —Ceca opening into excretory
vesicle* (14); vitelline follicles extending

VOLUME 106, NUMBER 2

213

posteriorly to middle of posterior testis* (1 5);
vitelline follicles confluent dorsally ( 1 6).

T. absita Blair et al., 1988

Diagnosis. — Ceca opening into excretory
vesicle* (14); vitelline follicles extending
posteriorly to middle of posterior testis* (1 5);
constriction in seminal vesicle (17).

Genus Gymnatrema Morozov, 1955

Diagnosis. — Acanthostominae having
some uterine loops lateral to testes but none
posttesticular (5); vitelline follicles not ex-
tending anteriorly to posterior margin of
seminal vesicle* {10)\ vitelline follicles con-
fluent posttesticularly (77); one cecum at-
rophied* (12); one cecum opening laterally
and one cecum ending blindly {13).

G. gymnarchi (Dollfus, 1950) Morozov,
1955

Diagnosis. —With characters of the genus.

Genus Proctocaecum Baugh, 1957
(Fig. 6)

Diagnosis. —Acanthostominae having
some uterine loops lateral to testes but none
posttesticular {8)\ ceca opening separately
and laterally at even levels (9); excretory
vesicle Y-shaped with short stem and con-
striction of arms in middle {14)\ eggs av-
eraging more than 30 txm. long (75); gonotyl
large, solid-muscular (76).

Subgenus Proctocaecum Baugh, 1957

Diagnosis.— Proctocaecum having rela-
tive length of uterine loops less than 45%
TBL* (1).

P. gonotyl {"DoWius, 1950) Brooks, 1980

Diagnosis.— V^iXh characters of the sub-
genus.

P. vicinum (Odhner, 1902) Brooks, 1980

Diagnosis. —Ceca opening separately and
laterally at uneven levels (2).

P. coronarium (Cobbold, 1861) Brooks,
1980

Diagnosis. — Ceca opening separately and
laterally at uneven levels (2); vitelline fol-
licles not extending anteriorly to posterior
margin of seminal vesicle* (3); one cecum
atrophied* (4); maximum oral spine length
more than 100 )um* (5).

Subgenus Overstreetium, new subgenus

Diagnosis.— Proctocaecum having ratio
of body length to width averaging 7.5-15:
1*(6).

P. productum (Odhner, 1902) Brooks, 1980

Diagnosis.— With characters of the sub-
genus.

P. elongatum (Tubangui & Masilungen,
1936) Brooks, 1980

Diagnosis.— Ceca. opening separately at
posterior end of body* (7); maximum body
length 7-16 mm* (8); ratio of body length
to width averaging more than 20:1 (9).

P. crocodili (Ysiimiguxi, 1954) Brooks, 1980

Diagnosis.— Ceca. opening separately at
posterior end of body* (7); maximum body
length 7-16 mm* (8); forebody less than
10% TBL (10).

P. atae (Tubangui & Masiluiigen, 1936)
Brooks, 1980

Diagnosis.— Ceca opening separately at
posterior end of body* (7); maximum body
length 7-16 mm* (8); ratio of oral sucker
width to pharyngeal width 1 :0.25-0.40* (1 1).

P. nicolli Brooks, 1980

Diagnosis.— Ceca opening separately at
posterior end of body* (7); maximum body
length 7-16 mm* (8); ratio of oral sucker
width to pharyngeal width 1:0.25-0.40* (1 1);
vitelline follicles extending anteriorly to
posterior margin of acetabulum* (12); max-
imum oral spine length more than 100 iim*
(13).

214

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Genus Caimanicola

Teixeira de Freitas & Lent, 1938

(Fig. 7)

Diagnosis. — Acanthostominae having
some uterine loops lateral to testes but none
posttesticular (5); ceca opening separately
and laterally at even levels (9); excretory
vesicle Y-shaped with short stem and con-
striction of arms in middle {14)\ eggs av-
eraging more than 30 iim long (75); gonotyl
lacking* (77); esophagus longer than phar-
ynx (18)\ tegumental spines unusually ro-
bust in mid-forebody (79); maximum body
length 7-16 mm* {20).

C. pavidus (Brooks & Overstreet, 1977)
Brooks, 1980

Diagnosis. —Oral spines averaging 25-30
in number* (1).

C. caballeroi (Pelaez & Cruz, 1953) Brooks,
1980

Diagnosis. —Eggs averaging less than 30
jLim long (2); maximum body length 2-6 mm
(3).

C. marajoarus Teixeira de Freitas & Lent,
1938

Diagnosis. —Eggs averaging less than 30
lim long (2); length of body occupied by
uterine loops more than 50% TBL* (4); vi-
telline follicles not extending anteriorly to
posterior margin of seminal vesicle* (5).

C. brauni (Mane-Garzon & Gil, 1961)
Brooks, 1980

Diagnosis. —Eggs averaging less than 30
ixm long (2); length of body occupied by
uterine loops more than 50% TBL* (4); ratio
of oral sucker width to acetabular width
1:0.3-0.7 (6).

Remarks: According to Ostrowksi de Nu-
iiez (1984b), Acanthostomum brauni of
Caballero (1 955) is an undetermined species
of acanthostome, possibly undescribed.

Genus Acanthostomum hooss, 1899
(Figs. 8, 9)

Diagnosis. —Acanthostominae having
some uterine loops lateral to testes but none
posttesticular (5); ceca opening separately
and laterally at even levels (9); eggs aver-
aging more than 30 ^ni long (75); gonotyl
lacking* (77); excretory vesicle with long
stem and short arms (27).

Subgenus Blairium, new subgenus

Diagnosis. —Acanthostomum having ceca
opening separately at posterior end of body*
(1).

A. scyphocephalum (Braun, 1899) Hughes
etal., 1941

Diagnosis.— V^iXh characters of the sub-
genus.

Remarks: Ostrowski de Nunez (1986) re-
described this species from the type mate-
rial, which had been missing and presumed
lost (see Brooks 1980).

A. americanum(FQTQz yigueras, 1957) Her-
ber, 1961

Diagnosis. —Eength of body occupied by
uterine loops more than 50% TBL* (2).

A. megacetabulum Thatcher, 1963

Diagnosis. —EQnglh of body occupied by
uterine loops more than 50% TBL* (2); ratio
of oral sucker : acetabular width 1:0.8-1.3*
(3).

A. gnerii Szidat, 1954

Diagnosis.— TqsXqs oblique (4); vitelline
follicles sparse (5).

A. minimum Stunkard, 1938

Diagnosis.— TqsXqs oblique (4); vitelline
follicles sparse (5); one cecum atrophied*
(6).

A. astorquii Watson, 1976

Diagnosis.— Testes oblique (4); vitelline

VOLUME 106, NUMBER 2

215

Timoniella Gymnatrema Proctocaecum Caimanicola Acanthostomum \ Pwaocaecum 1 ( Oversireetium - - I

coronarium vicinum gonotyl productum elongatum crocodili aloe nicolli

14- 15

Fig. 4. Phylogenetic relationships among the gen-
era of the subfamily Acanthostominae. Numbers refer
to apomorphic traits listed in diagnoses in text. Each
asterisk (*) indicates the presence of a homoplasious
character; the particular homoplasious characters are
denoted by an asterisk in the diagnoses in the text.

follicles sparse (5); one cecum atrophied*
(6).

Subgenus Gibsonium, new subgenus

Diagnosis.— Acanthostomum having vi-
telline follicles not extending anteriorly to
posterior margin of seminal vesicle* (7); oral
spines averaging less than 20 in number*
(8).

16 I ngure 4 )

Fig. 6. Phylogenetic relationships among species of
Proctocaecum. Numbers refer to apomorphic traits list-
ed in diagnoses in text. Each asterisk (*) indicates the
presence of a homoplasious character; the particular
homoplasious characters are denoted by an asterisk in
the diagnoses in the text.

Subgenus Acanthostomum Looss, 1899

Diagnosis. —Acanthostomum having oral
spines averaging 25-30 in number* (9); vi-
telline follicles extending anteriorly to pos-
terior margin of seminal vesicle* (10); eggs
averaging less than 30 fiva long (11); cecal
bifurcation approximately 1 0% TBL preac-
etabular (12).

A. absconditum (Looss, 1901) Poche, 1926 A. knobus Issa, 1962

Diagnosis.— V\fi\h. characters of the sub-
genus.

[ — Timoniella — ] [ - — Maillardiella —

praeterita imbutiforme incognita osirowskiae unami loossi

absita

Diagnosis.— Ratio of body length to width
averaging 7.5-15:1* (13).

A. spiniceps (Looss, 1896) Looss, 1899

Diagnosis. —Ratio of oral sucker width to
pharyngeal width 1:0.25-0.40* (14).

C. pavida C. caballeroi C. marajoara C. brauni

l*\r 3X- 5*\- -7^6

1-6 »*«*«*

7 |Dgure41

Fig. 5. Phylogenetic relationships among species of
Timoniella. Numbers refer to apomorphic traits listed
in diagnoses in text. Each asterisk (*) indicates the pres-
ence of a homoplasious character; the particular homo-
plasious characters are denoted by an asterisk in the
diagnoses in the text.

Fig. 7. Phylogenetic relationships among species of
Caimanicola. Numbers refer to apomorphic traits list-
ed in diagnoses in text. Each asterisk (*) indicates the
presence of a homoplasious character; the particular
homoplasious characters are denoted by an asterisk in
the diagnoses in the text.

216

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

[ Blairium ] [ Gibsonium] [ - Acanthostomum - )

astorquii minimum gnerii megaceiahulum americanum sc ahscondiium niloiicum spiniceps knobus

Fig. 8. Phylogenetic relationships among species of Acanthostomum excluding Acanthostomum (Atrophe-
caecum). Numbers refer to apomorphic traits listed in diagnoses in text. Each asterisk (*) indicates the presence
of a homoplasious character; the particular homoplasious characters are denoted by an asterisk in the diagnoses
in the text.

A. niloticum Issa, 1962

Diagnosis. —^diXio of oral sucker width to
pharyngeal width 1:0.25-0.40* (14); vitel-
line follicles extending anteriorly to poste-
rior margin of acetabulum* (15).

Subgenus Atrophecaecum Bhalerao, 1940

Diagnosis. —Acanthostomum having oral
spines averaging 25-30 in number* (9); vi-
telline follicles extending anteriorly to pos-

terior margin of seminal vesicle* ( 1 0); vi-
telline follicles terminating preovarially (16);
length of body occupied by uterine loops
more than 50% TBL* (17); forebody 10-
20%TBL*(18).

A. indicum Sinha, 1942

Diagnosis.— ^ixh characters of the sub-
genus.

A. 5/w5<2r5/:zz Kalyankar, 1977

[ Atrophecaecum

proctophorum
indicum slusankii pakistanensis asymmeiricum simhai

burminis lobaceiabulare c^rberi marini

Fig. 9. Phylogenetic relationships among species of Acanthostomum {Atrophecaecum). Numbers refer to
apomorphic traits listed in diagnoses in text. Each asterisk (*) indicates the presence of a homoplasious character;
the particular homoplasious characters are denoted by an asterisk in the diagnoses in the text.

VOLUME 106, NUMBER 2

217

Diagnosis. — Ceca opening into excretory
vesicle* (19); maximum body length 7-16
mm* (20).

A. pakistanense Coil & Kuntz, 1960

Diagnosis. — Prepharynx shorter than
pharynx* (21); forebody 10-20% TBL* (22);
ratio of body length to width averaging 7.5-
15:1* (23); oral spines averaging 20-24 in
number* (24); maximum oral spine length
more than 100 fxm* (25).

A. asymmetricum (Simha, 1958) Khalil,
1963

Diagnosis. —Prepharynx shorter than
pharynx* (21); one cecum lost (27); maxi-
mum body length 7-16 mm* (28).

A. proctophorum (Dwivedi, 1966) Yama-
guti, 1971

Diagnosis. —Prepharynx shorter than
pharynx* (21); one cecum lost (27); length
of body occupied by uterine loops more than
50% TBL* (29); oral spines averaging 20-
24 in number* (30); vitelline follicles con-
fluent preovarially (31).

A. simhai Khalil, 1963

Diagnosis. —Prepharynx shorter than
pharynx* (21); one cecum atrophied* (26);
vitelline follicles not extending anteriorly to
posterior margin of seminal vesicle* (32);
ratio of body length to width averaging 7.5-
15:1* (33); maximum body length 7-16
mm* (34).

A.burminis(&h2i\QV2iO, 1926)Bhalerao, 1936

Diagnosis. —VrQ^hdiVyny. shorter than
pharynx* (21); one cecum atrophied* (26);
vitelline follicles not extending anteriorly to
posterior margin of seminal vesicle* (32);
length of body occupied by uterine loops
more than 50% TBL* (35).

A. lobacetabulare Brooks & Caira, 1982

Diagnosis. —VrQx>^2ivynx shorter than
pharynx* (21); one cecum atrophied* (26);
vitelline follicles not extending anteriorly to

posterior margin of seminal vesicle* (32);
subterminal mouth* (36); lobate acetabu-
lum (37).

A. cerberi (Fischthal & Kuntz, 1965) Brooks,
& Caira 1982

Diagnosis. —Prepharynx shorter than
pharynx* (21); one cecum atrophied* (26);
vitelline follicles not extending anteriorly to
posterior margin of seminal vesicle* (32);
subterminal mouth* (36); no oral spines*
(38); no esophagus (39).

A. marinum (Coil & Kuntz, 1960) Brooks
& Caira, 1982

Acknowledgments

We gratefully acknowledge the loan of
specimens from Doctora Margarita Ostrow-
ski de Nunez, Buenos Aires, Argentina. This
study was supported by funds from oper-
ating grant A7696 from the Natural Sci-
ences and Engineering Research Council of
Canada (NSERC) to DRB.

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(DRB) Department of Zoology, Univer-
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Canada; (BH) Laboratorio de Zoologia In-
vertebrados, Facultad de Ciencias, Uni-

220

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

versidad de la Republica, Tristan Narvaja
1674, 11200 Montevideo, Uruguay.

Appendix 1

Transformation series for characters used to for-
mulate phylogenetic hypotheses for acanthostome di-
geneans (for outgroup argumentation, see Brooks 1980,
Brooks &Caira 1982, Brooks et al. 1985, Brooks et al.
1989). = plesiomorphic; 1 or higher indicates apo-
morphic states. U = nonlinear transformation series,
run unordered in computer-assisted analyses (numbers
assigned to each apomorphic state are arbitrary).

1. Oral sucker subterminal (0); terminal (1).

2. Oral sucker lacking spines (0): armed with single
row of spines (1).

3. Preacetabular pit lacking (0); present (1).

4. Ventrogenital pit present (0); lacking (1).

5. Genital pore in preacetabular pit (0); not in pre-
acetabular pit (1).

6. Seminal vesicle not coiled posteriorly (0); coiled
posteriorly (1).

7. Gonotyl lacking (0); suckerlike (1); large, solid-
muscular (2). U

8 . Seminal receptacle posto varian (0); preo varian ( 1 ) .

9. Vitelline follicles extending anteriorly to posterior
margin of seminal vesicle (0); not extending an-
teriorly to posterior margin of seminal vesicle (1);
confluent dorsally (2); confluent posttesticularly
(3). C/

10. Space occupied by uterine loops less than 45%
TBL (0); more than 50% TBL (1).

1 1 . Prepharynx longer than pharynx (0); shorter than
pharynx (1).

12. Ratio of oral sucker width to pharyngeal width
averaging 1:0.5 (0); 1:0.25-0.40 (1).

1 3. Ratio of body length to width averaging less than
7.5:1 (0); 7.5-15:1 (1); more than 20:1 (2). U

1 4. Ceca ending blindly near posterior end of body (0);
cyclocoel (1); opening separately at posterior end
of body (2); opening into excretory vesicle (3);
opening separately and laterally at even levels (4);
one cecum atrophied (5); one cecum opening lat-

erally and one cecum ending blindly (6); opening
separately and laterally at uneven levels (7); one
cecum lost (8). U

15. Forebody more than 20% TBL (0); 10-20% TBL
(1); less than 10% TBL (2).

16. Maximum body length less than 7 mm (0); 7-16
mm (1).

17. Ratio of oral sucker : acetabular width 1:0.6-0.9
(0); 1:0.8-1.3(1); 1:0.3-0.7(2).

18. Oral spines averaging 20-24 in number (0); 25-
30 (1); less than 20 (2). U

19. No constriction in seminal vesicle (0); constriction
present (1).

20. Posttesticular loops present (0); some uterine loops
lateral to testes but none posttesticular (1).

2 1 . Excretory vesicle Y-shaped with long stem (0); with
short stem and constriction of arms in middle ( 1 );
long stem and short arms (2).

22. Eggs averaging less than 30 ^m long (0); more than
30 ^ni long (1).

23. Maximum oral spine length less than 100 nm. long
(0); more than 100 Mm (1).

24. Esophagus shorter than pharynx (0); longer than
pharynx (1).

25. Tegumental spines not unusually robust in mid-
forebody (0); unusually robust in mid-forebody
(1).

26. Testes tandem (0); oblique (1).

27. Vitelline follicles numerous (0); sparse (1).

28 . Cecal bifurcation averages 20% TBL preacetabular
(0); 10% TBL preacetabular (1).

29. Vitelline follicles terminating at least at ovarian
level and no further posterior than posterior mar-
gin of the ovary (0); extending posteriorly to mid-
dle of posterior testis (1); terminating preovarially
(2). U

30. Vitelline follicles not confluent preovarially (0);
confluent preovarially (1).

31. Acetabulum not lobate (0); lobate (1).

32. Esophagus present (0); lacking (1).

33. Prepharynx present (0); lacking (1).

34. Secondary group of vitelline follicles surrounding
testes lacking (0); present (1).

PROC. BIOL. SOC. WASH.

106(2), 1993, pp. 221-224

A NEW MERICELLA

(MOLLUSCA: GASTROPODA: CANCELLARIIDAE)

FROM NORTHEASTERN AFRICA

Richard E, Petit and M. G. Harasewych

Abstract. —Mericella bozzettii, new species is described from nine shells trawled
off Cape Ras Hafun, Somalia in shrimp and lobster nets at a depth of 200-250
m. This new species may be distinguished from its western Indian Ocean
congeners by its large size and coarsely cancellate sculpture.

Thiele (1929) originally erected Mericel-
la, as a subgenus of Cancellaria, to contain
a single Recent species from bathyal depths
off Tanzania that he had previously attrib-
uted (Thiele 1925) to Cancellaria (Merica).
Mericella paschalis (Thiele, 1925), a closely
related species from the Zanzibar Channel
that was described in the same publication
as the type species, has not previously been
ascribed to this genus. Olsson & Bayer (1972)
proposed the generic name Gerdiella to ac-
commodate three newly discovered Recent
species taken in bathyal depths (5 1 6-897 m)
of the northern Caribbean Sea and the Straits
of Florida. They recognized that Gerdiella
was closely related to, and possibly conge-
neric with, Mericella, but distinguished these
taxa on the basis of size and geographical
distribution. Subsequent authors have as-
signed Cancellaria {Merica) corbicula Dall,
1908, which occurs in bathyal to abyssal
depths off southern California, to Mericella
(Abbott 1974:247) or Gerdiella (Kaicher
1978). Due to the rarity of material, the
genus Mericella has received little subse-
quent attention. An additional species of
Mericella, known from shells of nine spec-
imens collected by shrimp trawlers off the
coast of Somalia, is described in this report.

Abbreviations used in the text: AMNH,
American Museum of Natural History, New
York; MNHN, Museum national d'Histoire
naturelle, Paris; USNM, National Museum

of Natural History, Smithsonian Institu-
tion, Washington, D.C.

Mericella bozzettii, new species
Figs. 1-3, Table 1

Diagnosis.— A large species with thick,
high-spired, coarsely cancellate shell with
irregularly spaced varices. Aperture exceeds
half the shell length, with weak denticles
along flaring, strongly sinuate outer lip.

Description. —Shell (Fig. 1, Table 1) large
for genus, reaching 37 mm, heavy, with tall,
conical spire, rounded anterior. Protoconch
(Figs. 2-3) conical, of 2^4 smooth, rounded
whoris, aligned with coiling axis, increasing
in diameter from 368 ^m to 1.87 mm. Tran-
sition to teleoconch (Figs. 2-3, arrow)
abrupt, demarcated by weak varix, followed
immediately by four faint spiral cords, and
within Va whorl by axial costae. Teleoconch
with up to five strongly convex whorls. Su-
ture impressed. Shoulder indistinct or ab-
sent. Spiral sculpture of 23-25 strong cords
on body whorl, 8-9 on penultimate whorl,
lacking intervening threads. Axial sculpture
of 25-28 rounded, regularly-spaced, sinu-
ate, opisthocline, axial costae on body whorl
(16-18 on first teleoconch whorl) producing
beaded, cancellate appearance at intersec-
tions with spiral cords, with four to six fine
axial lamellae in intervening concave spac-
es. Varices up to seven in number, initially

222

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-3. Mericella bozzettii, new species. 1, Apertural, lateral and dorsal views of holotype. Scale bar = 1
cm. 2, Lateral and 3, apical views of the protoconch. Scale bars = 500 /um.

weak and irregularly placed (90-270° apart),
more prominent and regularly spaced
(^220°) on third and subsequent teleoconch
whorls. Aperture elongate, elliptical, de-
flected from coiling axis by 13-16°. Outer
lip thickened, forming flaring, strongly sin-

uate varix, with 12-18 weak denticles lim-
ited to the base of the varix. Inner lip ad-
pressed posteriorly, with an angle of 153-
157° between parietal region and columella.
Columella with two weak columellar folds
and siphonal fold, posteriormost fold most

VOLUME 106, NUMBER 2

223

Table l.—Mericella bozzettii, new species. Measurements of shell characters. Linear measurements in mm {n
9).

Character

Mean

Range

Shell length (SL)

Aperture length (AL)

AL/SL

No. of whorls, protoconch^

No. of whorls, teleoconch

No. of varices

No. of axial ribs between

varices 5-6
No. of axial ribs between

varices 6-7
No. spiral cords on

penultimate whorl
No. of spiral cords on

whorl 5-6
No. of teeth on outer lip

30.6

3.9

22.1-37.0

16.9

2.3

12.2-20.3

0.553

0.018

0.512-0.578

1.92

0.12

1.75-2.0

4.72

0.27

4.0-5.0

7.0

0.0

7.0

14.1

2.0

11-18

17.4

3.1

13-24

7.8

0.6

7-9

7.0

0.8

5-8

13.3

2.0

10-16

'' « = 3 for this character.

pronounced. Anterior slope of shell round-
ed, lacking clear distinction between body
whorl and siphonal canal. Shell color uni-
formly white. Periostracum, soft parts un-
known.

Material examined.— YloXoXy^Q, USNM
860315, 28.7 mm; Paratype 1, AMNH
226453; Paratype 2, MNHN; Paratype 3;
Petit collection; Paratypes 4-8, Bozzetti col-
lection; all from the type locality.

Type locality. —Off CsLpQ Ras Hafun, ap-
proximately 150 km S of Cape Guardafni,
Somalia. Trawled in shrimp and lobster nets
at 200-250 m.

Etymology.— This species is named in
honor of Mr. Luigi Bozzetti, who first
brought it to our attention and kindly pro-
vided the type material.

Remarks.— Although this new species
corresponds more closely in size to species
of the western Atlantic genus Gerdiella, it
is placed in Mericella because of its large
aperture (>y2 shell length), smooth proto-
conch lacking axial sculpture, as well as be-
cause of its geographic proximity to other
species of Mericella. It is readily distin-
guished from Mericella jucunda and M. pa-
schalis on the basis of its large size, much
broader axial costae, thick shell and white

color. Mericella corbicula more closely ap-
proaches M. bozzettii in size, but differs in
having a more finely reticulate surface
sculpture, a lower spire, and a chalky shell
surface.

Although Olsson & Bayer (1972:880)
confirmed the inclusion of the closely re-
lated genus Gerdiella in Cancellariidae on
the basis of the morphology of the radula
of the type species, nothing is known of the
anatomy or radular morphology of any spe-
cies of Mericella. Mericella bozzettii occurs
in somewhat shallower depths (200-250 m)
than other species of Mericella (404-20 1 2
m).

Literature Cited

Abbott, R. T. 1974. American seashells, second edi-
tion. Van Nostrand Reinhold, New York, 663
pp., 24 pis.

Dall, W. H. 1908. [Reports on the dredging opera-
tions off the west coast of Central America to
the Galapagos, to the west coast of Mexico, and
in the Gulf of California, in charge of Alexander
Agassiz, carried on by the U.S. Fish Commis-
sion Steamer "Albatross." during 1891, Lieut.
Commander Z. L. Tanner, U.S.N.. command-
ing. XXXVIl. Reports on the scientific results
of the expedition to the eastern tropical Pacific,
in charge of Alexander Agassiz, by the U.S. Fish

224

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Commission Steamer "Albatross," from Octo-
ber 1904, to March, 1905, Lieut. Commander
L. M. Garrett, U.S.N., commanding. XIV] The
Mollusca and Brachiopoda. — Bulletin of the
Museum of Comparative Zoology 43(6):205-
487, pis. 1-22.

Kaicher, S. D. 1978. Card catalogue of world-wide
shells. Pack no. 19— Cancellariidae. Privately
published, St. Petersburg, Rorida. Card nos.
1859-1964.

Olsson, A. A., & F. M. Bayer. 1972. Gerdiella, a. new
genus of deep-water cancellariids. — Bulletin of
Marine Science 22:875-880.

Thiele, J. 1925. Gastropoda der Deutschen Tiefsee-
Expedition. II. — Deutsche Tiefsee-Expedition
17:35-382 + pis. 13-46.

. 1929. Handbuch der systematischen Weich-

tierkunde.— Gustav Fischer, Jenna 1:1-376.

Department of Invertebrate Zoology,
NHB stop 118, National Museum of Nat-
ural History, Smithsonian Institution,
Washington, D.C. 20560, U.S.A.

PROC. BIOL. SOC. WASH.

106(2), 1993, pp. 225-236

NEW SPECIES OF ALVINELLIDAE (POLYCHAETA)

FROM THE NORTH FIJI BACK- ARC BASIN

HYDROTHERMAL VENTS (SOUTHWESTERN PACinC)

Daniel Desbruyeres and Lucien Laubier

Abstract. —The polychaete family Alvinellidae Desbruyeres & Laubier, 1986
comprises two genera, Alvinella and Paralvinella, and ten species or subspecies.
All species are strictly associated with hydrothermal vents in the Pacific Ocean.
The genus Paralvinella includes eight species or sub-species, plus one additional
new species presently being described from North East Pacific hydrothermal
fields. In 1989, the French research submersible Nautile had 12 successful dives
in the North Fiji back-arc Basin and explored two active hydrothermal vents.
Numerous specimens of two additional new species of the genus Paralvinella
were collected using the manipulator of the submersible. Paralvinella uniden-
tata, new species, exhibits several features that lead us to erect three different
subgenera, Paralvinella s. s., Miralvinella, new subgenus and Nautalvinella, new
subgenus, within the genus Paralvinella. This species lives within the anhydrite
mass, very close to hot fluid openings. The second new species, P. fijiensis,
found in the anhydrite mass, but also on basaltic rocks, is closely related to P.
grasslei, the type species, and to P. palmiformis.

While the first discovery of hydrothermal
phenomena occurring at the axes of oceanic
ridges goes back to 1976 (Lonsdale 1977),
the exploration of hydrothermal systems in
back-arc basins is rather recent (Both et al.
1986, in Manus Basin, Hessler et al. 1988,
in Marianas back-arc Basin, Fouquet et al.
1990, 1991, in Lau Basin, Ohta 1990, in
Okinawa back-arc Basin) (Auzende et al.
1989, Jollivet et al. 1989). During the early
summer of 1989, a French- Japanese bio-
logical cruise, STARMER 2 (30 June 1989
to 19 July 1989) was devoted to the study
of biological communities associated with
deep-sea hydrothermal vents in the South-
western Pacific, in the North Fiji back-arc
Basin (Desbruyeres et al. 1991). The French
research submersible Nautile, operated from
R/V Le Nadir, had 12 successful dives in
these areas. Two different active sites were
explored. White Lady vent has one large
diffuser plus several small cylindrical chim-
neys; the edifice is a few meters high and is

built up by anhydrite, with hydrothermal
fluid temperature up to 285°C; the fluid is
translucent, relatively depleted of metal due
to subsurface phase separation and shows a
low content of hydrogen sulfide. The site is
located at a depth of 2000 m at 16°59'S and
173°55'E. Another hydrothermal field
named Mussel Valley consists of Bathy-
modiolus spp. beds developing on the ba-
salt; there is no chimney and the hydro-
thermal fluid diffuses from cracks between
basaltic rocks; the ffuid temperature does
not exceed 8.5°C. This site is located at a
depth of 2700 m at 18°49'S and 173°29'E.
A large collection of polychaetes including
representatives of the family Alvinellidae
were collected from the submersible.

Since our last publication on Alvinellidae
(Desbruyeres & Laubier 1991), the total
number of known Paralvinella species in-
cludes eight species or subspecies (Detinova
1988). At least one additional new species
from North East Pacific hydrothermal fields

226

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

is presently being described and additional
material tentatively identified as Paralvi-
nella hessleri has recently been collected
from Okinawa vent fields (Miura & Ohta
1991).

During the dives in the North Fiji Basin,
numerous specimens of two different spe-
cies of the genus Paralvinella were collected
using the manipulator of the submersible.
Morphological study of these animals
showed that they represent two new species,
Paralvinella unidentata and P.fijiensis. Par-
alvinella unidentata exhibits several fea-
tures that lead us to erect three different sub-
genera within the genus Paralvinella. This
species lives in the anhydrite mass, very close
to hot fluid openings. The second new spe-
cies, Paralvinella fijiensis, found in the an-
hydrite mass and on basaltic rocks, is closely
related to P. grasslei, the type species, and
P. palmiformis.

Paralvinella unidentata, new species
Figs. 1, 3, 4

Type locality, material examined. —Sev-
enty-eight specimens collected during Nau-
tile dives PL 1 (3 specimens. White Lady
site), PL 1 1 (5 specimens. White Lady site),
PL 16 (37 specimens. White Lady site), PL
20 (32 specimens. White Lady site), PL 21
(1 specimen, small hydro thermal vent lo-
cated 150 m from White Lady site in the
south-west). Extra specimens from dive PL
10 and PL 20 deep frozen for biochemical
analyses. Most specimens come from White
Lady vent site (depth 2000 m, 16°59'50"S
and 173°55'47"E). Holotype (dive PL 16, 1 1
July 1989, on White Lady site) deposited in
the collections of the Museum national
d'Histoire naturelle, Laboratoire de Biolo-
gic des Invertebres marins et Malacologie
(n°UC 350). Paratypes from same dive de-
posited in the collections of the National
Museum of Natural History, Smithsonian
Institution, Washington, D.C. (USNM
157044).

Etymology. —The specific name refers to

the peculiar unidentate uncini. The lack of
a secondary tooth on the uncini is unique
within the family Alvinellidae.

Description. —Holotype 1 1 mm long and
1 . 1 mm wide with 8 1 setigerous segments.
Paratypes (14 specimens from PL 16) range
from 77 to 88 setigerous segments, with the
majority (1 1 specimens) having from 79 to
83. Length of paratypes ranges from 4.8 mm
to 9.2 mm, with an average of 5.2 mm.
Color pale grey-pinkish in ethanol, with
capillary setae and acicular notopodial hooks
yellow; integuments iridescent. Body grad-
ually tapering from about setigerous seg-
ment 50 to the end of the body. A medio-
ventral row of small shields present.

Prostomium well developed, with ovi-
form shield shape, clearly separated from
buccal segment by deep grooves, with an-
terior median incision on two thirds of
length; with small glandular notch visible
(with SEM) at base of incision. Buccal ap-
paratus, from dorsum to ventrum, with
many grooved tentacles in several rows on
buccal membrane arising from dorsal side
of buccal cavity; length of tentacles highly
variable from one individual to the next
one, possibly due to preservation. Paired
large ventral tentacles and ventral organ ab-
sent.

Buccal segment laterally and ventrally
visible, well separated from prostomium and
from segment IL First visible segment (II)
achaetous, laterally and ventrally discern-
ible, clearly separated from peristomium and
from branchial region. First 25 to 30 setig-
erous segments with notopodia only.

Branchial region with four segments, pre-
ceded by one reduced asetigerous segment,
visible laterally and ventrally (segment II).
First branchial segment (segment III) ase-
tigerous, totally fused with two first setig-
erous segments. Third setigerous segment
well separated from others. Following three
segments setigerous, with notopodia dor-
sally elevated in laterodorsal row, with no-
topodia of same size as those of following
segments.

VOLUME 106, NUMBER 2

227

Branchiae four pairs, all similar, arranged
as funnel-like structure, with strong basal
stem bearing small secondary filaments and
thin terminal tip devoid of secondary fila-
ments as long as basal stem. Branchial stem
bearing one, sometimes two, bean-shaped
vesicles on internal side near base. Second-
ary filaments inserted along stem on two
opposite areas; each leaf-shaped, strongly
flattened, with median ciliated area, and
pointed tip. These secondary filaments rem-
iniscent of species of Alvinella.

Notopodia, from setiger 1 to end of body
(7th setiger excepted) each cylindrical, bear-
ing two groups of capillary setae. Notopodia
without digitiform lobes. Setiger 7 strongly
modified, with slightly reduced cylindrical
notopodia, bearing two to three straight short
acicular notopodial setae on each side. Se-
tiger 8 not modified.

Cylindrical notopodia and uncinigerous
neuropodial tori on each segment from se-
tigerous segment 26 to 29 (with a majority
from 28 to 29). Uncini numerous (20 to 50
per torus), in single rows, with teeth directed
anteriorly (retrogressive arrangement). Un-
cini with only a single main tooth, lacking
a secondary tooth.

Pygidium rounded, with five conspicuous
rounded papillae, two ventral paired papil-
lae and three smaller, dorsal ones.

Tubes unknown.

Ecology.— From observations made on
the White Lady hydrothermal site, the al-
vinellid worms colonize the whitish mass
of anhydrite, living close to high tempera-
ture (285°C) translucent desalinated hydro-
thermal fluid openings. On videotapes ob-
tained by the submersible 3 CCD TV
camera, the branchial fans of Paralvinella
spp. are visible at the surface of the mass
of anhydrite. The worms belong to P. uni-
dentata and to P. fijiensis, another species
(see below), that are not distinguishable dur-
ing the sampling procedure.

Discussion. — Since the discovery of the
first species oi Paralvinella, P. grass lei Des-
bruyeres & Laubier, 1982, six species or

subspecies have been described within the
genus: P. palmiformis Desbruyeres & Lau-
bier, 1986, P. pandorae pandorae Desbru-
yeres & Laubier, 1986, P. pandorae irlandei
Desbruyeres & Laubier, 1986 (Desbruyeres
& Laubier 1986), P. dela Detinova, 1988,
P. hessleri Desbruyeres & Laubier, 1989
(Desbruyeres & Laubier 1989) and P. bac-
tericola Desbruyeres & Laubier 1991 (Des-
bruyeres & Laubier 1991).

Among these species, P. pandorae and its
two subspecies are clearly distinguished by
the presence of uncini from setigerous seg-
ments 5 or 6 (depending on the subspecies),
a unique situation within alvinellids con-
sidered as a plesiomorphous character.

The following morphological features can
be used to assess the relationship between
species and groups of related species: The
total number of segments and its range of
variation (primitive situation: 100 to 150
segments, with a large range of individual
variation; apomorphous situation, 60 to 80
segments, with a reduced range of individ-
ual variation); the rank of occurrence of the
anteriormost neuropodial uncinigerous to-
rus (first uncinigerous torus anterior to the
modified setigerous segment in plesiomor-
phous situation, first uncinigerous torus
from segment 13 to more than segment 60
in apomorphous situation). The range of
variation increases with the rank of occur-
rence; the buccal apparatus, with two dif-
ferent basic types: the first one bears two
large paired lateral tentacles in addition to
the small ciliated dorsal tentacles, and a re-
versible ventral globular organ; the second
one lacks the large paired tentacles and has
an unpaired pointed organ with a longitu-
dinal slit and a reversible ventral globular
organ. The development of the large paired
tentacles adapted for different trophic be-
haviors is considered as an apomorphic fea-
ture. P. unidentata exhibits a unique buccal
structure, reduced to the numerous grooved
tentacles inserted on a well developed buc-
cal membrane: this structure basically dif-
fers from all previously described species of

228

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1 . Paralvinella unidentata, new species. Entire animal in ventro-lateral view.

Paralvinella; the shape and position of the posite or adjacent longitudinal rows. There

secondary filaments of the branchiae. These is no obvious reason to consider either of

can be cylindrical and slender or flattened these structures or situations more primi-

and leaf-shaped, and inserted on two op- tive than the other one. Nevertheless, con-

VOLUME 106, NUMBER 2

229

•mm

*%..«»

Ss-SWfigi

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Fig. 2. Parahinella fijiensis, new species. Entire animal in ventro-lateral view.

230

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

sidering the facts that the genus Alvinella
possesses flattened leaf-shaped secondary
filaments and is clearly apomorphic to the
genus Paralvinella (Desbruyeres & Laubier
1986), the question of the systematic rank
of P. unidentata, with identical flattened leaf-
shaped secondary filaments, must be con-
sidered carefully; and the shape of the un-
cini, in all alvinellids but P. unidentata, is
characterized by one main tooth and one
secondary smaller tooth, while in P. uni-
dentata there is only one main tooth. Still,
the general shape of the uncinus is identical
in both groups and the presence or absence
of a secondary tooth is not considered an
important phylogenetic significance. How-
ever, the absence of a secondary tooth can
be considered plesiomorphic to the biden-
tate semiavicular uncini previously known
for all alvinellids.

Within the genus Paralvinella, P. uniden-
tata can be best compared with P. pandorae
and its two subspecies in the structure of
the buccal apparatus and the funnel-like ar-
rangement of the branchiae. However, P.
unidentata can be easily distinguished by a
series of characters including the rank of
occurrence and shape of neuropodial uncini
and the leaf-shaped secondary filaments of
the branchiae.

Paralvinella fijiensis, new species
Figs. 2, 5

Type locality, material examined. —
Ninety-five specimens collected and pre-
served during Nautile dives PL 1 1 (3 spec-
imens, White Lady site), PL 12 (1 specimen,
White Lady site), PL 14 (47 specimens;
White Lady site), PL 16 (10 specimens.
White Lady site), PL 20 (34 specimens,
White Lady site). All specimens come from
White Lady vent site (depth 2000 m,
16°59'50"S and 173°55'47"E). Holotype
(dive PL 14) deposited in the collections of
the Museum national d'Histoire naturelle,
Laboratoire de Biologic des Invertebres
marins et Malacologie (n°UC 439). Para-

types from same dive deposited in the col-
lections of the National Museum of Natural
History, Smithsonian Institution (USNM
157043).

Etymology. —The species is named for its
geographic origin.

Description. —Holotype 24 mm long and
5 mm wide with 63 setigerous segments.
Paratypes ( 1 5 specimens from PL 1 4) range
from 50 to 68 setigerous segments. Color
pale grey-pinkish in ethanol, with capillary
setae and acicular notopodial hooks yellow;
integument iridescent. Body maggot-shaped
in large specimens; small animals with body
gradually tapering posteriorly; medioven-
tral area slightly depressed. When viewed
under scanning microscope, integument of
intersegmental areas densely covered with
small circular glandular areas and secretions
from same.

Prostomium reduced medially with two
anterior lobes and two nucal grooves
obliquely situated near base. Prostomium
laterally separated from lateral parts of buc-
cal segment. Buccal segment laterally and
ventrally visible, well separated from pro-
stomium and segment IL Buccal apparatus
comprising many grooved tentacles inserted
dorsally and two large paired grooved ten-
tacles inserted ventrally, ending with three
unequally developed rounded lobes. Edge
of main lobe provided with several rows of
small rounded internal papillae.

Branchial region comprised of four seg-
ments, preceded by one reduced asetigerous
segment still visible laterally and ventrally
(segment II). First branchial segment ase-
tigerous, ventrally visible. Next three bran-
chial segments (setigerous segments 1 to 3)
totally fused. Notopodia of first setigerous
segment very reduced; notopodia of 2nd se-
tigerous segment reduced; notopodia of 3rd
setigerous segment similar to the following
notopodia. Notopodia of setigers 1 to 3 ad-
jacent to stem of external pair of branchiae.
Fourth setigerous segment fused ventrally
to fourth branchial segment.

Branchiae four pairs, all similar, with a

VOLUME 106, NUMBER 2

233

1mm

200um

20um

500um

50um

Fig. 5. Paralvinella fijiensis, new species. A, anterior part in lateral view. B, buccal apparatus, showing
grooved tentacles (right) and the terminal lobes of one of the ventral large tentacles (left). C, left parapod from
anterior setiger, showing dorsal lobe. D, capillary seta covered with small spines and two rows of larger ones
on the edge. E, modified notopodium of setiger 7, with large acicular hooks. F, uncinigerous torus, right side of
body.

234

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Strong basal stem. Secondary filaments very
abundant, inserted on two opposite areas of
stem to end of branchia, reminiscent of Par-
alvinella grasslei and P. palmiformis.

Notopodia, from setiger 4 to the end of
body (7th setiger excepted) cylindrical,
bearing two groups of capillary setae heavily
coated with filamentous bacteria. Notopo-
dia from about setiger 9 to setiger 30 bearing
dorsal and ventral rounded lobes. Setiger 7
strongly modified, with two reduced dorsal
and ventral lobes surrounding a very re-
duced notopodium bearing three to four
curved acicular notopodial setae on each
side. Surface of acicular setae covered with
minute spinelets; these are longer on convex
side of acicular hooks when viewed under
SEM. This structure similar to ornamen-
tation of notopodial setae in other segments.
Notopodia of setiger 8 with anterior part
enlarged.

Uncinigerous neuropodial tori present on
each segment from setiger 12 to 19 (13 on
the holotype). Occurrence of first uncini-
gerous torus appears to be inversely pro-
portional to total number of setigerous seg-
ments: on a total of 23 individuals, with
total number of setigerous segments from
46 to 68, first uncinigerous segment occurs
from 19th to 12th setigerous segment. Re-
gression curve:

Ro = (96 - N)/2.54

where Rq = rank of occurrence of the first
uncinigerous torus and N = total number
of setigerous segments of the body, indi-
cating that the anteriormost uncinigerous
tori appear latest.

Uncini numerous (20 to 50 per row) in
single rows, with teeth facing anteriorly (ret-
rogressive arrangement). Uncini increasing
in number posteriorly. Each uncinus with
one main tooth surmounted by smaller sec-
ondary tooth, as in other species of Paral-
vinella (except P. unidentata, see above).

Pygidium blunt, without conspicuous pa-
pillae.

Ecology.— yiosX specimens were collect-
ed in anhydrite samples together with P.
unidentata; on one occasion, a tube inhab-
ited by P. fijiensis was found on a piece of
basalt.

Discussion.— VsfiXhm the genus Paralvi-
nella, P. fijiensis exhibits clear relationships
with the stem species P. grasslei and its close
relative P. palmiformis. These three species
have in common: the structure of the buccal
apparatus, with two large trilobate paired
ventral tentacles; the general shape of the
branchiae; the rank of occurrence of the un-
cinigerous tori, starting between setiger 12
(P. fijiensis, new species) and setiger 3 1 {P.
palmiformis)', and the first setigerous seg-
ment is very reduced, and the second setig-
erous segment reduced, emphasizing the
importance of the cephalization processes.

Within this group of three species, P. fi-
jiensis is characterized by the number of
setigerous segments (maximum 68 in P. fi-
jiensis, compared to 1 10 in P. grasslei and
118 in P. palmiformis), the small papillae
of the main end of the trilobate paired buc-
cal tentacles and the shape and relative size
of the prostomium. Additional minor diff-
ences can be found in the presence of spine-
lets on the acicular hooks and the enlarged
base of the 7th notopodium in P. fijiensis.

Conclusion

All Paralvinella species except P. hessleri
and the two new species described herein
are known from the East Pacific Rise and
related ridge systems in the Eastern Pacific.
Within this general framework, two differ-
ent species groups can be distinguished in
the Northern and Southern parts of the East
Pacific ridge system. These parts have been
separated by the subduction of the Ameri-
can plate over the oceanic crust off Oregon
starting 35 MY (TunnicHffe 1988). P. pal-
miformis and P. pandorae pandorae live in
the Northern part, while P. grasslei and P.
pandorae irlandei live in the Southern part.

VOLUME 106, NUMBER 2

235

These two pairs of species (P. grasslei and
P. palmiformis) or subspecies (P. p. irlandei
and P. p. pandorae) have been regarded as
examples of sibling species (Desbruyeres &
Laubier 1986). Two additional species, P.
bactericola in the Southern area and P. dela
in the Northern one, have been discovered,
providing a third example of a closely re-
lated paired species. More recently, P. hess-
leri, collected in the Mariana back-arc Basin
in the Western Pacific, was found to exhibit
a close relationship with P. bactericola and
P. dela. Paralvinella fijiensis and P. uniden-
tata herein described from the North Fiji
Basin provide additional information about
the striking similarities which can be found
between the different groups of species liv-
ing in these three major areas:

Evolutionary groups:
Group 1 Group 2

Group 3

East Pacific Rise:
P. grasslei P. p. irlandei P. bactericola

Juan de Fuca- Explorer:
P. palmifor- P. p. pandorae P. dela
mis

Western Pacific:
P. fijiensis P. unidentata P. hessleri

It must be emphasized that each evolu-
tionary group within the genus Paralvinella
is present in each of the three geographic
areas. Moreover, morphological differenti-
ation is not related to the distance between
the hydrothermal areas: the three species
from the Western Pacific are much more
different from one another than they are
from related species in other biogeograph-
ical areas.

We hypothesize that these three different
groups of Paralvinella species evolved in-
dependently from three ancestors adapted
to hydrothermal environmental conditions.
As a consequence of this hypothesis, we pro-
pose the establishment of three subgenera,
that can be diagnosed as follows:

Subgenus Paralvinella

Type species. —Paralvinella {Paralvinella)
grasslei Desbruyeres & Laubier, 1982

Other species. —P. (P.) palmiformis Des-
bruyeres & Laubier, 1986 and P. (P.) fijien-
sis

Diagnosis.— Buccal apparatus, compris-
ing, from dorsum to ventrum, numerous
buccal grooved retractile tentacles inserted
on a semi-circular buccal membrane, two
large paired trilobate appendages and a
globular eversible ventral organ. Branchiae
with cylindrical secondary filaments, in-
serted on two opposite areas on the stem up
to its end. Digitiform or rounded notopo-
dial lobes present on some anterior setig-
erous segments.

Subgenus Miralvinella, new subgenus

Type species.— Paralvinella {Miralvinel-
la) dela Detinova, 1988

Other species.— P. {M.) hessleri Desbru-
yeres & Laubier, 1989 and P. {M.) bacter-
icola Desbruyeres & Laubier, 1991.

Etymology. —Miralvinella, from Alvinel-
la and MIR, the name of the two deep-sea
Russian submersibles recently built in a
Finnish shipyard for the Shirshov Institute
of the former Academy of Sciences of the
Soviet Union.

Diagnosis.— BwccdiX apparatus complex,
comprising from dorsum to ventrum nu-
merous grooved tentacles inserted on a buc-
cal membrane, two large deeply grooved ta-
pering paired tentacles and a globular
eversible ventral organ. Branchiae with cy-
lindrical secondary filaments, inserted on
two opposite areas on the stem up to its end.
Digitiform notopodial lobes present on some
anterior segments.

Subgenus Nautalvinella, new subgenus

Type species. —Paralvinella {Nautalvinel-
la) pandorae Desbruyeres & Laubier, 1986
Other species.— P. {N.) unidentata

236

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Etymology. —Nautalvinella, from Alvi-
nella and Nautile, the French deep-sea sub-
mersible of IFREMER which was used dur-
ing the STARMER cruise and previous
expeditions on hydrothermal vents areas.

Diagnosis.— ^uccdl apparatus comprising
numerous grooved tentacles inserted dor-
sally on a buccal membrane. A median hol-
lowed pointed lobe can be present in some
species. No large paired ventral tentacles.
No ventral organ. Branchiae with numerous
leaf-shaped secondary filaments inserted on
two more or less adjacent lines on the stem.
Terminal part of the stem devoid of sec-
ondary filaments. No digitiform notopodial
lobes.

Literature Cited

Auzende, J. M., et al. 1989. Le cadre geologique d'un
site hydrothermal actif: la campagne STAR-
MER 1 du submersible Nautile dans le Bassin
Nord-Fidjien. — Comptes Rendus de I'Acade-
mie des Sciences, Paris, Serie II 309: 1 787-1 795.

Both, R., et al. 1986. Hydrothermal chimneys and
associated fauna in the Manus Back-Arc Basin,
Papua New Guinea.— Eos 67:489-490.

Desbruyeres,D.,A. M.Alayse,&S. Ohta. 1991. Deep-
sea hydrothermal communities in two back-arc
basins of the south west Pacific (the North Fiji
and Lau basins): composition, microdistribu-
tion and food- web. — 6th Deep-Sea Biology
Symposium, Copenhagen, July 1991, 20 pp.

, & L. Laubier. 1982. Paralvinella grasslei, new

genus, new species of Alvinellinae (Polychaeta:
Ampharetidae) from the Galapagos rift geo-
thermal vents.— Proceedings of the Biological
Society of Washington 95:484-494.

, & . 1986. Les Alvinellidae, une famille

nouvelle d'annelides polychetes infeodees aux
sources hydrothermales sous-marines: syste-
matique, biologic et ecologie.— Journal Cana-
dien de Zoologie 64:2227-2245.

, & . 1989. Paralvinella hessleri, new

species of Alvinellidae (Polychaeta) from the

Mariana back-arc basin hydrothermal vents. —
Proceedings of the Biological Society of Wash-
ington 102:761-767.

, & . 1991. Systematics, phylogeny,

ecology and distribution of the Alvinellidae
(Polychaeta) from deep-sea hydrothermal
vents.— Ophelia Supplement 5:31-45.

Detinova, N. N. 1988. New species of polychaetous
annelids from hydrothermal vents of the Juan
de Fuca ridge (Pacific Ocean).— Zoologichesky
Zhumal 57:858-864.

Fouquet, Y., et al. 1990. Hydrothermal activity in
the Lau Basin.— Eos, May 1:678-679.

, et al. 1991. Hydrothermal activity and me-

tallogenesis in the Lau back-arc basin.— Nature
349:778-780.

Hessler, R. R., P. Lonsdale, & J. Hawkins. 1988. Pat-
terns on the ocean floor.- New Scientist 1605:
47-48.

Jollivet, D., et al. 1989. Premieres observations de
communautes animales associees a I'hydrother-
malisme arriere-arc du bassin Nord Fidjien.—
Comptes Rendus de I'Academie des Sciences,
Paris, Serie III 309:301-308.

Lonsdale, P. 1977. Clustering of suspension-feeding
macrobenthos near abyssal hydrothermal vents
at oceanic spreading centers.— Deep Sea Re-
search 24: 857-863.

Miura, T., & S. Ohta. 1991. Two polychaete species
from the deep-sea hydrothermal vent in the
middle Okinawa Trough.- Zoological Science
8:383-387.

Ohta, S. 1990. Deep-sea submersible survey of the
hydrothermal vent community on the north-
eastern slope of the Iheya Ridge, the Okinawa
trough. —Jamstec Deepsea Research 2: 145-1 56.

Tunnicliffe, V. 1988. Biogeography and evolution of
hydrothermal vent-fauna in the eastern Pacific
Ocean. — Proceedings of the Royal Society of
London B 223:347-366.

(DD) Ifremer, Centre de Brest, B.P. 70,
29263 Plouzane, France; (LL) Institut
Oceanographique, Laboratoire de Physio-
logie des Etres marins, 195, rue Saint-
Jacques, France.

PROC. BIOL. SOC. WASH.
106(2), 1993, pp. 237-242

GYPTIS CRYPT A, A NEW HESIONID SPECIES FROM

THE U.S.A. EAST COAST, WITH A REDESCRIPTION OF

G. VITTATA WEBSTER & BENEDICT, 1887

(ANNELIDA: POLYCHAETA)

Fredrik Pleijel

Abstract. — Gyptis crypta, new species, is described from North Carolina,
Florida, and off Mississippi, and several previous records of G. vittata Webster
& Benedict, 1887 are referred to this species. The new species is unique within
the genus in having only ten terminal papillae on the proboscis, and in having
segment 5 equipped with both noto- and neurosetae, rather than neurosetae
only. The European species G. rosea (Malm, 1874) and G. mediterranea Pleijel,
1993 are considered the most closely related. Gyptis vittata is redescribed from
type material.

Subsequent to the original account, Gyp-
tis vittata Webster & Benedict, 1887 has
been reported on several occasions from the
U.S.A. east coast (e.g., Pettibone 1963, Day
1974, Gardiner 1975, Uebelacker 1984).
Reexamination of the currently available
specimens on which these records were
based shows them to represent, in part (Pet-
tibone 1963), a species of Podarkeopsis,
possibly P. levifuscina Perkins, 1984, and,
in part (Gardiner 1975, Uebelacker 1984),
an undescribed species of Gyptis, herein de-
scribed as G. crypta. Day's specimens were
not available for examination, and are con-
sidered of uncertain status. Except for the
original description from Maine, no valid
literature records of G. vittata appear to ex-
ist. To avoid further confusion, G. vittata is
redescribed from the syntypes.

A provisional diagnosis and a brief dis-
cussion of the genus Gyptis Marion & Bob-
retzky, 1875 is provided in Pleijel (1993).

All drawings were made with a camera
lucida. Width measurements are from me-
dian segments and include parapodia but
exclude cirri and setae. Apart from material
of congeneric species used for comparison
(detailed in Pleijel 1993), the study is based
on specimens from the National Museum

of Natural History, Smithsonian Institu-
tion, Washington, D.C. (USNM).

Gyptis vittata Webster & Benedict, 1887
Fig. 1

Gyptis vittata Webster & Benedict, 1887:
715-716, pi. 1, figs. 21-22, pi. 2, fig. 23.
[Not descriptions and figures of Gyptis
vittata sensu Pettibone, 1963: 106-107, fig.
28c-d.- Gardiner, 1975:119, fig. 81-p.-
Uebelacker, 1984:28-29 to 28-31, fig. 28-
28a^g]

Material examined. —MainQ: 3 syntypes
(USNM 452, including slides 503-506),
Eastport, low water, rocks; 25-30 fm, shells.
(Specimens apparently from two different
but fused samples.)

Description. — Prostomium rounded,
about as wide as long, posteriorly with
rounded lobes separated by posterior inci-
sion (Fig. 1 A). Palpophores cylindrical; pal-
postyles widest medially, anteriorly round-
ed (Fig. 1 B), longer than palpophores. Paired
antennae thinner and possibly longer (see
Remarks) than palps, cylindrical with
pointed tips. Median antenna short, club-
shaped, widest subdistally (but see Re-
marks); inserted half-way between anterior

238

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

pair eyes and anterior margin of prosto-
mium. Anterior pair eyes rounded to reni-
form, twice as large as posterior pair and
situated further apart; posterior pair round-
ed; both pairs with lenses. Nuchal organs
not discernible.

Large, distinct lip glands present laterally
on the ventral lips (Fig. IB). Proboscis prob-
ably divided in proximal and distal parts
(visible on a mounted specimen only where
it is partly everted and difficult to observe).
Number of terminal proboscideal papillae
difficult to discern, possibly 1 5-20. Papillae
in single ring, conical to cylindrical.

Proximal parts of tentacular cirri indis-
tinctly annulated, medial and distal parts
distinctly annulated; rings about as long as
wide; tips not observed (no complete ten-
tacular cirri present). At least one acicula
present in all cirrophores of dorsal tentac-
ular cirri, not discernible in ventral ones.
Anterior dorsal segmental delineations not
distinct; segment 1 probably reduced and
following segments fused.

Notopodia of segment 5 (setiger 1 ) with-
out setigerous lobes or setae. Dorsal cirri
incomplete, slightly stouter than those of
following segments. Neuropodia similar to
following ones. Ventral cirri similar to fol-
lowing ones but slightly smaller. Segment 6
similar to median ones.

Notopodia of median segments situated
on cirrophores, with distinctly annulated
dorsal cirri, longer than notosetae. Rings ca.
1.5 times as long as wide. Possible alter-
nation of length and orientation of dorsal
cirri not discernible. Notopodial lobes con-
ical, with one or two internal aciculae. All
notosetae simple; setal details not discern-
ible.

Neuropodia of median segments conical,
with two internal aciculae, one larger and
one smaller. All neurosetae compound; dis-
tal part of shafts internally reticulated. Ven-
tral cirri apparently smooth, shorter than
neuropodial lobe, on small cirrophores sit-
uated far back on the neuropodium (Fig.
IC).

Pygidium not observed.

Color: Eyes brown. No other pigmenta-
tion retained.

Habitat.— Currently known only from
rocks, intertidal, and shells, 25-30 fm.

Distribution. —Known only from the type
locality.

Remarks.— Webster & Benedict's mate-
rial consists of three syntypes: one in alco-
hol, two mounted on slides (one dorsally
and one ventrally), and two additional slides
with mounted parapodia. All three speci-
mens lack posterior ends. The unmounted
specimen consists of an anterior end plus a
median part, both in very poor condition.
The mounted syntypes are both females with
eggs, one approaching maturity with eggs
about 120-140 ixm in diameter. A median
antenna is present only on one of the
mounted specimens, and, being of unusual
shape for the group, it cannot be excluded
that it is distorted (e.g., flattened by prep-
aration).

As seen from the descriptions as well as
specimens deposited at USNM, Gardiner's
(1975) and Uebelacker's (1984) descrip-
tions of G. vittata both refer to G. crypta,
new species, whereas Pettibone's (1963) de-
scription refers to a species ofPodarkeopsis,
possibly P. levifuscina, from Hadley Har-
bor, Woods Hole, Massachusetts. Day
(1973) recorded G. vittata from off'Beaufort,
and stated that the specimens were depos-
ited at USNM and the Duke University Ma-
rine Laboratory. Since these specimens are
absent, however, the record could not be
verified and is considered uncertain.

Gyptis crypta, new species
Fig. 2

Gyptis vittata. —Gardiner, 1975: 1 19, fig. 81-
p.-Uebelacker, 1984:28-29 to 28-31, fig.
28-28a-g [not Webster & Benedict, 1887].

Gyptis sp. -Taylor, 1 97 1 : 1 67-1 7 1 , fig. 4a-f.

Material examined. —^orth. Carolina: 1
paratype (USNM 52892) Wrightsville
Beach, Banks Channel, intertidal in burrow

VOLUME 106, NUMBER 2

239

Fig. 1 . Gyptis vittata, syntypes. Setae omitted. A. Anterior end, dorsal view. B. Same as A but drawn with
focus moved to ventral side of specimen; arrows indicate lip glands. C. Parapodium. A & B slide 505. C slide
503. Scale lines 0.25 mm.

of Notomastus lobatus, 20 Jul 1974; holo-
type (USNM 52893) and 5 paratypes
(USNM 15761 5), Wrightsville Beach, Banks
Channel, intertidal in burrow of Notomas-
tus lobatus, 18 Aug 1974; 5 paratypes
(USNM 52894), Wrightsville Beach, Banks
Channel, intertidal in burrow of Notomas-
tus lobatus, 27 Feb 1975. Gulf of Mexico:
18 paratypes (USNM 45534), Florida,
Tampa Bay, 1963; 1 paratype (USNM
75478), off Mississippi, 30°16.27'N,
88°36.42'W, 3.4 m, 24 Oct 1980; 1 paratype
(USNM 75319), off Rorida, 24°47.5'N,
82°13.16'W, 24 m, Jul 1981.

Description.— ^ody, excluding parapo-
dia, cylindrical, tapered posteriorly. Venter

slightly flattened, without distinct median
longitudinal furrow.

Prostomium rectangular to trapezoidal
with rounded comers (Fig. 2A). with small
posterior incision (often difficult to discern).
Palpophores cylindrical, palpostyles thin-
ner, narrowing to rounded ends. Palpo-
phores as long as or slightly longer than pal-
postyles. Paired antennae probably situated
on small ceratophores, slightly longer than
or as long as palps, with fine tips. Median
antenna inserted on or just in front of line
between anterior pair of eyes, cylindrical or
evenly tapering to a point, without extended
tip. Anterior pair eyes rounded, larger than
posterior pair and situated further apart,

240

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

/v-^^^-A^

Fig. 2. Gyptis crypta. A. Anterior end, dorsal view. Setae omitted. B. Anterior end, ventral view. Setae
omitted. C. Left parapodium, segment 5, anterior view. D. Right parapodium, median segment, anterior view.
About half of full number of setae shown. E. Acicular notoseta. F. Spinose capillary notoseta. G. Serrated
capillary notoseta. H. Median neuroseta. I. Ventral neuroseta. A & C-I paratypes USNM 157615, B paratype
USNM 45534. Scale lines A-B 0.25 mm, C 0.2 mm, D 0.2 mm, E-G 50 mhi, H-I 50 Mm.

VOLUME 106, NUMBER 2

241

posterior pair rounded, both pairs with lens-
es. Nuchal organs lateral to prostomium,
not coalescing dorsally.

Lip glands absent (Fig. 2B). Proboscis
short, divided in proximal and distal parts,
without discernible papillae. Distal part
narrow and smooth. Ten short, conical ter-
minal papillae in single ring.

Tentacular cirri thin and smooth, indis-
tinctly annulated distally, tips distinctly
pointed. Dorsal tentacular cirri of segments
3 and 4 reaching to about segment 1 or 11,
ventral tentacular cirri of segment 3 short-
est, reaching to about segment 5. Aciculae
in cirrophores of tentacular cirri not dis-
cernible. Segment 1 dorsally reduced, seg-
ments 2 and 3 probably fused dorsally.

Notopodia of segment 5 (setiger 1) similar
to following notopodia but slightly smaller
and with fewer setae (Fig. 2C). Dorsal cirri
similar to and as long as those of segment
4. Neuropodia similar to following ones but
slightly smaller and with fewer setae. Ven-
tral cirri similar to following ones.

Elevated dorsal ridges present across pos-
terior side of each segment (Fig. 2D), most
distinct on median and posterior segments.

Notopodia of median segments with more
or less distinctly annulated dorsal cirri
(smooth proximally). Dorsal cirri about as
long as setae, slightly shorter on anterior
segments, with about five or six rings, each
about three times as long as wide (Fig. 2D).
All dorsal cirri slender, possible alternation
of length and orientation not discernible.
Notopodial lobes conical, usually with one,
occasionally two, internal aciculae and three
kinds of notosetae: 5-10 anteriorly situated,
slightly curved acicular setae (Fig. 2E), ta-
pering, distally with fine spines; slender cap-
illary setae smooth proximally with two al-
ternating rows of spines medially and
subdistally (Fig. 2F); and few, rather stout,
ventrally situated, serrated capillary noto-
setae (Fig. 2G).

Neuropodia of median segments conical,
with one internal acicula, and about 20-40
compound setae. Distal part of setal shafts

with transverse striation internally. Blades
thin, dorsal serrated, median and dorsal ones
long (Fig. 2H), ventral ones short (Fig. 21).
Dorsally situated serrated capillary setae
absent. Ventral cirri smooth with fine ta-
pering tips (Fig. 2D), without cirrophores,
situated distally on neuropodium.

Pygidium rounded, pygidial cirri long and
thin with pointed tips, longer than dorsal
cirri, median papilla not observed.

Color: Live specimens not observed. Pre-
served specimens yellow with fine brown
pigment spots, usually denser distally on
noto- and neuropodia and on all cirri. Eyes
reddish brown. Small dark spots may be
present ventrally at parapodial bases.

Measurements. —Only two complete
specimens observed; length 7 mm, width
1.4 mm for 39 segments; length 7.5 mm,
width 1.2 mm for 39 segments.

//<2Z?/^a^— Currently known intertidally
from burrows of Notomastus lobatus and
down to 24 m depth.

Distribution.— North Carolina, west coast
of Florida and off Mississippi.

Etymology. —The name crypta is trans-
literated from the Greek noun "/cpuTrrr?,"
meaning haunt and hiding-place and refers
to the habit of living in burrows of Noto-
mastus lobatus.

Remarks.— The holotype is a mature
male, and the paratypes include several ma-
ture females with eggs (50-60 ^m in di-
ameter). Gametes were observed in speci-
mens collected from August to February in
North Carolina, although those collected in
February did not appear fully mature.

Gyptis crypta differs from all other known
members of the genus in having a small and
fixed number of terminal papillae on the
proboscis, and in being provided with both
noto- and neurosetae on segment 5 (rather
than only neurosetae). Together with G. ro-
sea Malm, 1874 and G. mediterranea Plei-
jel, 1993 G. crypta shares the two putative
derived characters distally inserted ventral
cirri that taper evenly to a point (rather than
subdistally inserted ventral cirri with short

242

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

tips as in other Gyptis). Apart from the
smaller number of proboscis papillae and
the biramous parapodia of the fifth segment,
G. crypta is separated from these two species
in having lighter colored eyes (probably red
on live specimens) and a more elongated
body.

Although Gardiner (1975) examined the
syntypes of G. vittata, his description of G.
vittata and that of Uebelacker (1 984) clearly
characterize the new species, and specimens
from both authors now constitute part of
the type material of G. crypta.

Acknowledgments

I wish to thank H. G. Hansson and Ou-
rania Papakosta for nomenclatural advice,
K. Fauchald and L. Ward for loan of spec-
imens as well as working facilities, B. Hilbig
and T. Perkins for comments on the manu-
script, and the Swedish Natural Science Re-
search Council (contracts 9555-306 and
-307) for financial support.

Literature Cited

Day, J. H. 1973. New Polychaeta from Beaufort, with
a key to all species recorded from North Car-
olina. -NO AA Technical Report NMFS 375:
1-140.

Gardiner, S. L. 1975. Errant polychaete annelids from
North Carolina.— Journal of the Elisha Mitchell
Scientific Society 91:77-220.

Malm, A. W. 1874. Annulater i hafvet utmed Sveri-

ges vestkust och omkring Goteborg.— Gote-
borgs K. vetenskaps- och vitterhetssamhalles
handlingar 14:67-105.

Marion, A. P., & N. Bobretzky. 1875. Etude des An-
nelides du Golfe de Marseille. —Annales des sci-
ences naturelles 2:1-106.

Perkins, T. H. 1984. New species of Phyllodocidae
and Hesionidae (Polychaeta), principally from
Florida. — Proceedings of the Biological Society
of Washington 97:555-582.

Pettibone, M. H. 1963. Marine polychaete worms of
the New England region. — Bulletin of the Unit-
ed States National Museum 227:1-356.

Pleijel, F. 1993. Taxonomy of European species of
Amphiduros and Gyptis (Polychaeta: Hesioni-
dae).— Proceedings of the Biological Society of
Washington 106:158-181.

Taylor, J . L. 1971. Polychaetous annelids and benthic
environments in Tampa Bay, Florida. Unpub-
lished, Ph.D. Dissertation, University of Flor-
ida, Gainesville, 1332 pp.

Uebelacker, J. M. 1 984. Chapter 28. Family Hesioni-
dae Sars, 1862. Pp. 28:1-28:39 in J. M. Uebe-
lacker and P. G. Johnson, eds., Taxonomic guide
to the polychaetes of the northern Gulf of Mex-
ico. Final report to the Minerals Management
Service, contract 14-12-001-29091. Barry A.
Vittor & Associates, Inc., Mobile, Alabama. 7
vols.

Webster, H. E., & J. E. Benedict. 1 887. The Annelida
Chaetopoda from Eastport, Maine.— Report of
the United States Commissioner of Fisheries
1885:707-755.

Swedish Museum of Natural History,
Stockholm, and (postal address): Tjamo
Marine Biological Laboratory, PL 2781,
S-452 96 Stromstad, Sweden.

PROC. BIOL. SOC. WASH.
106(2), 1993, pp. 243-250

A NEW SPECIES OF UNIPORODRILUS

(OLIGOCHAETA: TUBIFICIDAE) FROM THE GULF OF

MEXICO COAST OF FLORIDA, AND A

PHYLOGENETIC ANALYSIS OF THE GENUS

Christer Erseus and Michael R. Milligan

Abstract. — Uniporodrilus purus, new species, from subtidal sand near Pen-
sacola in northwestern Florida, is described. It is distinguished from all con-
geners by lacking penial setae. A most parsimonious cladistic hypothesis of the
species of Uniporodrilus (subfamily Phallodrilinae) is presented. It suggests a
basal dichotomy of the genus, with U. purus and U. granulothecus, both subtidal
species, constituting one clade characterized by unpaired male and sperma-
thecal pores, and the four remaining species, all intertidal, representing another
clade characterized by a high number of somatic setae and enlarged pharyngeal
cavity.

The genus Uniporodrilus was established
for Uniporodrilus granulothecus Erseus,
1979, a subtidal marine tubificid from the
east coast of the United States (Erseus 1979).
Davis (1985) described a second species as-
signed to this genus, Uniporodrilus vestig-
ium, from Georges Bank off Massachusetts.
In a recent revision of the subfamily Phal-
lodrilinae (Erseus 1992a), however, the lat-
ter species was transferred to the genus At-
lantidrilus Erseus, 1982, and at the same
time, Uniporodrilus was enlarged to include
also three taxa previously placed in Phal-
lodrilus Pierantoni, 1902: Phallodrilus scir-
piculus Erseus, 1985 (from Saudi Arabia),
Phallodrilus nasutus Erseus, 1990(a), and
Phallodrilus bipartitus Erseus, 1 990(a) (both
from the Caribbean area). A fifth species
recently described from Hong Kong, Uni-
porodrilus furcatus Erseus, 1992(b) is also
regarded as a member of this genus, which
has been defined by its "varying" penial se-
tae (setae different in size and morphology
within bundles) and its granulated sper-
mathecal ducts (Erseus 1992a).

In oligochaete material collected in north-
western Rorida by Mr. Jerry McLelland
(Gulf Coast Research Laboratory, Ocean

Springs, Mississippi), an additional species
of Uniporodrilus was discovered. It is de-
scribed in the present paper, which also in-
cludes a cladistic analysis of the species
within this genus.

All specimens of Uniporodrilus purus, new
species, were stained in paracarmine and
mounted whole in Canada balsam. The type
series is deposited in the U.S. National Mu-
seum of Natural History (USNM), Smith-
sonian Institution, Washington, D.C. For
comparisons, the whole-mounted speci-
mens in the type material of U. granulothe-
cus (also in the USNM) were re-examined.

Cladograms were constructed under the
principle of maximum parsimony, using the
program PAUP (Phylogenetic Analysis Us-
ing Parsimony, version 3.0) by Swoffbrd
(1990).

Uniporodrilus Erseus, 1979

Uniporodrilus Erseus, 1979:414-415.—

1992a:22-23.
Uniporodrilus (partim): Davis 1985:169.
Phallodrilus Pierantoni (partim): Erseus

1990b:54.

Diagnosis (emended after Erseus

244

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1992a) (assumed autapomorphies under-
lined).— Small marine tubificids. Prosto-
mium distinctly longer than its width at base;
generally well set off from peristomium (see
Fig. 1). Somatic setae bifid with upper tooth
thinner and shorter than lower. [Posterior
dorsal setae sharply single-pointed in U. na-
sutus.] Penial setae, when present, in fan-
shaped bundles or in rings, with bifid or
single-pointed, hooked tips. Penial setae ab-
sent [in U. purus], or from about 6 to about
22 per bundle. Spermathecal pores un-
paired, mid- ventral in posterior part of IX
[in U. granulothecus and U. purus], or paired
in line with ventral setae in X [in all other
species]. Pharyngeal cavity large and much
ciliated in some species. Vasa deferentia ei-
ther entering apical end of, or somewhat
subapical on, atria. Atria small, oval, or
elongate, curved; when paired, each with
two large prostate glands [atrium unpaired
in U. granulothecus and U. purus, with four
prostates, at least in U. granulothecus]. Atria
opening into more or less developed cop-
ulatory sacs. Spermathecae [unpaired in U.
granulothecus and U. purus] elongate, with
thick-walled, granulated ducts , and small
ampullae.

Type species. — Uniporodrilus granulothe-
cus Erseus, 1979.

Remarks. —The characteristic shape of the
prostomium was referred to as "snout-like"
by Erseus (1992a:23). For differences be-
tween this diagnosis and the one by Erseus
(1992a), see Discussion below.

Uniporodrilus granulothecus Erseus, 1979
Fig. lA-B

Uniporodrilus granulothecus Erseus, 1979:
415-417, figs. 1-3 .-Erseus 1992a:22.

Material re-examined. —The holotype
(USNM 56307) and 3 paratypes (USNM
56308, 56309), all whole-mounted.

Remarks. — In the character matrix used
for the cladistic analysis of the Phallodrili-
nae (Erseus 1992a:table I), the prostomium
of U. granulothecus was coded as "not snout-

like." This was inferred from the original
description (Erseus 1979), which does not
include any particular note on the prosto-
mium, except that it is "rounded, about as
long as its width at peristomium." The re-
examination of the type specimens, how-
ever, showed that the prostomium of this
species (Fig. lA-B) is very similar to that
of any congener.

All the re-examined specimens have di-
atoms in their guts, which suggests that U.
granulothecus selectively feeds on these al-
gae.

Uniporodrilus purus, new species
Figs. IC-D, 2

Holotype. -U^IS^M 157046, whole-
mounted specimen.

Type locality.— V^rdido Key, near Pen-
sacola. Gulf of Mexico coast of Florida; 500
m from shore, about 6 m depth, sand (Oct
1989; J. McLelland).

Paratypes. -USNM 157047-157049, 3
specimens from type locality.

Other material.— Milligsin collection: 2
specimens from type locality.

Etymology.— The species epithet purus
(Latin for 'simple,' 'plain') refers to the lack
of penial setae, and to the single unpaired
atria and spermathecae, in this species. All
congeners have penial setae, and all but one
(U. granulothecus) have paired genitalia.

Description. -Fixed worms stout, 1.5-1.9
mm long, 0.16-0.23 mm wide at segment
XI; 25-35 segments. Prostomium (Fig. IC-
D) large, clearly longer than its width at
base. Clitellum extending over V2X-XII. Se-
tae (Fig. 2A-B) bifid with upper tooth re-
duced; reduction more pronounced in pos-
terior setae (Fig. 2B) than in anterior ones
(Fig. 2A); lower tooth is prolonged in pos-
terior setae (Fig. 2B). Setae 30-45 ixm long,
about 1-2 /Lim thick, three per bundle
throughout body; but absent ventrally from
XI (penial setae absent). Male pore un-
paired, mid-ventral, posterior to middle of
XL Spermathecal pore unpaired, mid- ven-
tral, in most posterior part of IX.

VOLUME 1 06, NUMBER 2

245

prm

200 pm

Fig. 1. A-B, Uniporodrilus granulothecus, anterior ends of two paratypes (USNM 56308, 56309); C-D, U.
purus, new species, anterior ends of holotype (C) and one paratype (D). Abbreviation: prm, prostomium. Dense
shading indicates epidermal lining, light shading (coelomic?) space within prostomium.

Pharyngeal cavity hollow, but not large.
Pharyngeal glands in (III)IV-V. Male gen-
italia (Fig. 2C) partly paired, partly un-
paired. Vasa deferentia paired, much longer
than atrium, conspicuously widened (up to
1 5-22 ixm wide) along most parts; cilia few
(or absent?), but scattered spermatozoa of-
ten present in vasa. Both vasa deferentia
entering unpaired atrium somewhat sub-

apically. Atrium oval, 30-40 txva long, 27-
30 jLini wide, with thin outer muscular lining
(1-2 fxvci thick), and ciliated and somewhat
granulated inner epithelium. Atrium open-
ing into unpaired copulatory sac (but details
not clear in available material). Lobed bod-
ies of prostate glands located posterio-lat-
eral to atrium (on both sides). They appear
broadly attached to atrium, but a division

50 jjm

Fig. 2. Uniporodrilus purus, new species. A, Free-hand drawing of anterior seta. B, Free-hand drawing of
posterior seta. C, Lateral view of spermatheca and male genitalia in segments X-XI. Abbreviations: a, atrium;
cs, copulatory sac; pr, prostate gland; pro, prostate gland of other side of worm; s, spermatheca; vd, vas deferens;
vdo, vas deferens of other side of worm.

246 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

granulothecus
purus
furcatus
nasutus

bipartitus

scirpiculus

furcatus

scirpiculus

nasutus

bipartitus

Fig. 3. The two most parsimonious trees (A-B) obtained with the PAUP program for the six ingroup taxa
of Uniporodrilus (Table 1) and their 13 characters (listed in text, coded in Table 1). Search method: branch-
and-bound. Length of trees 17 steps, consistency index 0.765. Rooting at outgroup (Nootkadrilus), which is
excluded from trees. The clade granulothecus /purus is identical in both trees, and therefore only shown in A.
The numbers refer to characters. Filled rectangle, autapomorphy; open rectangle, apomorphy that is later followed
by reversal; two parallel lines, convergence; cross, reversal.

into anterior and posterior glands (as is nor- part of spermatheca somewhat granulated,

mal for Phallodrilinae) not obvious; either longer and more thick- walled than inner part

anterior or posterior prostates possibly ab- (ampulla); lumen of duct, however, irregu-

sent. Spermatheca (Fig. 2C:s) unpaired, lar. Sperm scattered throughout duct as well

elongate pear-shaped, 85-100 iim long, as ampulla,

maximally 33-45 ij,m wide. Outer, duct-like Remarks. —This new species is closely re-

VOLUME 106, NUMBER 2

247

Table 1.— Data matrix for parsimony analysis of the species of Unipowdrilus (Fig. 3). For characters and
character states, see text.

Taxon

Character states, characters 1-13

Outgroup:

Nootkadrilus Baker, 1982

00700

00000

000

Ingroup:

Unipowdrilus granulothecus Erseus, 1979

10001

01011

oil

U. purus, n. sp.

10???

?1011

oil

U. furcatus Etsqus, 1992b

11110

01100

oil

U. scirpiculus Etsqus, 1985

11011

00100

001

U. nasutus Erseus, 1990a

11100

10100

101

U. bipartitus Erseus, 1 990a

11100

10100

101

lated to U. granulothecus (cf. Erseus 1979).
Both taxa have somewhat inflated vasa def-
erentia, as well as unpaired atrium and sper-
matheca. Moreover, in these two species,
the spermathecal pore is in segment IX, not
in X as is normal for Tubificidae. Unipo-
wdrilus purus is, however, unique within
the genus by its lack of penial setae. It is
further distinguished from U. granulothecus
by its stouter atrium, and its less clearly
bipartite spermatheca (in U. granulothecus
the spermathecal duct is much longer than
ampulla and has densely granulated walls).

The condition of the prostate glands (with
one pair possibly absent) needs to be con-
firmed when new, preferably sectioned, ma-
terial becomes available. If any of the pros-
tate glands is absent, this feature is an
additional autapomorphy for U. purus.

The guts appear empty in all examined
specimens. Thus the diet of U. purus is un-
known (cf. Remarks for U. granulothecus
above).

Distribution and habitat.— Kno'wn only
from the type locality in the northeastern
part of the Gulf of Mexico. Subtidal sand,
at 6 m depth.

Phylogenetic Analysis
Taxa.— The six species of Uniporodrilus
(Table 1) are the ingroup taxa in the anal-
ysis. Nootkadrilus Baker, 1982, was selected
as the outgroup; according to the cladistic
analysis of the whole subfamily Phallodri-

linae (Erseus 1992a), this genus possibly is
the sister group of Uniporodrilus. For the
coding of most of the outgroup character
states, the descriptions by Baker (1982) were
used. For character 9 in the list below, how-
ever, material of an unidentified Nootkad-
rilus species and of A^. longisetosus (Brink-
hurst & Baker, 1979) in the senior author's
collection was examined.

Characters.— T\\Q following characters
and character states were used. In the PAUP
run, reversals were allowed for all charac-
ters.

1. Prostomium not prolonged (0); dis-
tinctly longer than its width at base ("snout-
like"), and well set off* from peristomium

(1).

2. Somatic setae maximally three or four
(occasionally five) per bundle (0); maxi-
mally five to seven (occasionally even eight
or nine) per bundle (1).

3. Penial setae all single-pointed (0); at
least some penial setae bifid (1).

4. Penial setae of about the same size
within bundle (0); some penial setae dis-
tinctly larger than others within bundle (1).

5. Within each bundle, penial setae
forming a straight row that is parallel to long
axis of worm (0); penial setae forming ring
within each bundle (1).

6. Penial setae one continuous group
within each bundle (0); penial setae in two
groups within each bundle (1).

7. Male pores paired (0); unpaired (1).

248

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

8. Pharyngeal cavity normal, not en-
larged (0); enlarged (1).

9. Pharyngeal glands clearly extending
into segment VI (occasionally into VII) (0);
not extending further than into V (1).

10. Atria and spermathecae paired, sper-
mathecae in segment X (0); atrium and sper-
matheca unpaired, spermatheca in IX (1).

1 1 . Atria elongate, not small (0); pear-
shaped, small (1).

12. Prostate glands not broadly attached
to atria (0); at least one prostate gland
broadly attached to atrium/atria (1).

13. Spermathecal ducts not granulated
(0); granulated (1).

Results. —The branch-and-bound-algo-
rithm of PAUP 3.0 was used to find the two
most parsimonious trees (A and B), which
both are shown in Fig. 3. Both trees are fully
resolved, with U. granulothecus and U. pu-
rus forming one clade, and the four other
species as the sister group of this clade. The
topologies of the two trees differ with regard
to the branching pattern within the larger
sister group: U. furcatus is either grouped
with U. nasutus and U. hipartitus (tree A),
or with U. scirpiculus (tree B). Uniporodrilus
nasutus and U. bipartitus are always togeth-
er, as they have identical character codings
(see Table 1).

However, the character transformations
indicated in Fig. 3 A-B are not the only most
parsimonious optimizations of the present
data set. Although not affecting the branch-
ing pattern, in tree A, the apomorphic state
of character 4 can be interpreted as con-
vergent for U. furcatus and U. scirpiculus,
instead of assuming reversal in the clade
nasutus /bipartitus (as shown in Fig. 3 A). In
tree A also, character 5 could be regarded
as a basal apomorphy for the whole ingroup,
assuming a reversal in the clade furcatus/
nasutus /bipartitus, or it could be seen as
independent autapomorphies for U. gran-
ulothecus and U. scirpiculus (U. purus was
coded as "?" for this character as it lacks
penial setae altogether). In a similar way, in
tree B, character 3 could be regarded as con-

vergent apomorphies for U. furcatus and the
nasutus/ bipartitus clade. Still, none of these
alternative optimizations, or combinations
of them, will give other tree topologies than
those in Fig. 3.

Discussion

In the previous revision (Erseus 1992a),
the varying size and morphology of the pe-
nial setae within the bundles, and the gran-
ulation of the spermathecal ducts, were
assumed to be autapomorphies of Unipo-
rodrilus. The "snout-like" prostomium
(character 2 in Erseus 1992a:fig. 4) was then
interpreted as a synapomorphy of U. scir-
piculus, U. nasutus, U. bipartitus and U. fur-
catus (the last-mentioned called "Hong
Kong sp. r' by Erseus 1992a). The present
study has shown that U. granulothecus as
well as the new taxon U. purus have pro-
longed prostomia (Fig. 1) and thus, this fea-
ture (character 1 in Fig. 3) can be used to
define the genus as a whole.

The character state "penial setae of at least
two different kinds within bundle" (char-
acter 1 in Erseus 1 992a), on the other hand,
is treated in a more restricted sense here
(character 4 in present paper). In the present
analysis, only those two species {U. furcatus
and U. scirpiculus) with distinctly aniso-
morphic penial setae are coded as apomor-
phic for this character (character 4 in Fig.
3). The variation of setal morphology in the
penial bundles of U. granulothecus, U. na-
sutus and U. bipartitus is slight, and in U.
purus, penial setae are absent.

The spermathecal duct is not as long (in
relation to the ampulla) in U. purus as in
the other species, but it is granulated (al-
though not as heavily as in the congeners),
and therefore the granulation of the sper-
mathecal ducts (character 1 3 in Fig. 3) is an
apomorphy that still supports the mono-
phyly of Uniporodrilus.

The present parsimony analysis suggests
a basal dichotomy of Uniporodrilus, with U.
granulothecus and U. purus forming the sis-

VOLUME 106, NUMBER 2

249

ter group of the rest of the genus (Fig. 3).
Monophyly of the granulothecus/purus clade
is unequivocally supported by the unpaired
male pore (character 7), the restricted ex-
tension of the pharyngeal glands (character
9), the unpaired atrium and spermatheca,
with the latter in segment IX (character 10),
and the broad attachment of the anterior
prostates (character 12); characters 7 and
12, however, also (convergently) apomor-
phic in U. furcatus. Depending on which
character optimization one prefers, one may
also use the ring-shaped penial bundles
(character 5) to define this group, based on
the assumption that the penial setae were
forming rings in the most recent ancestor of
U. granulothecus and U. purus, but not in
the common ancestor of the whole genus
(see Remarks for the Phylogenetic analysis
above). Moreover, both U. granulothecus
and U. purus have distinctly widened parts
of the vasa deferentia. This feature appears
synapomorphic, but it is not conclusive
whether it is unique to these two species, as
the vasa deferentia have not been visible in
their full lengths in the other members of
the genus.

Monophyly of the four other species is
supported by the high number of somatic
setae (character 2 in Fig. 3) and the enlarged
pharyngeal cavity (character 8). Further, two
characters suggest that U. nasutus and U.
bipartitus are closely related, the disjunct
distribution of setae within the penial bun-
dles (character 6), and the small, pear-shaped
atria (character 1 1). With regard to the phy-
logenetic position of U. furcatus and U. scir-
piculus, the parsimony analysis is less de-
cisive. Depending on which one of characters
3 (the bifid penial setae) or 4 (the distinctly
anisomorphic penial setae) is considered as
homoplasious, U. furcatus may either be re-
garded as the sister taxon of U. scirpiculus
(Fig. 3B), or as the sister taxon of the na-
sutus /bipartitus group (Fig. 3A).

Habitat and geographical distribution
were not used as "characters" in the par-
simony analysis, but the monophyly of U.

granulothecus and U. purus seems to be sup-
ported by their subtidal habitat, as all the
other four species, as well as the outgroup
(Nootkadrilus), are intertidal. Furthermore,
the possibility that U. furcatus (from Hong
Kong) and U. scirpiculus (from Saudi Ara-
bia) are endemic to the Indo-West Pacific
region appears to favor their monophyletic
status, i.e., in accordance with tree B (Fig.
3); the congeners are all from the Northwest
Atlantic and Nootkadrilus is so far only
known from the Northeast Pacific.

Acknowledgments

We are indebted to Mr. Jerry McLelland
for providing the material of U. purus; to
Ms. Barbro Lofnertz (University of Gote-
borg), and Ms. Christine Hammar (Swedish
Museum of Natural History), for technical
assistance; and to the Swedish Natural Sci-
ence Research Council, for financial sup-
port.

Literature Cited

Baker, H. R. 1982. Two new phallodriline genera of
marine Oligochaeta (Annelida: Tubificidae) from
the Pacific Northeast. — Canadian Journal of Zo-
ology 60:2487-2500.

Brinkhurst, R. O., & H. R. Baker. 1979. A review of
the marine Tubificidae (Oligochaeta) of North
America. — Canadian Journal of Zoology 67:
1553-1569.

Davis, D. 1985. The Oligochaeta of Georges Bank
(NW Atlantic), with descriptions of four new
species. — Proceedings of Biological Society of
Washington 98:158-176.

Erseus, C. 1979. Uniporodrilus granulothecus n.g.,
n.sp., a marine tubificid (Oligochaeta) from east-
em United States.— Transactions of the Amer-
ican Microscopical Society 98:414-418.

. 1982. Atlantidrilus, a new genus of deep-sea

Tubificidae (Oligochaeta). — Sarsia 67:43-46.

. 1985. Annelida of Saudi Arabia. Marine Tu-
bificidae (Oligochaeta) of the Arabian Gulf coast
of Saudi Arabia. — Fauna of Saudi Arabia 6
(1984): 130-1 54.

. 1 990a. The marine Tubificidae (Oligochaeta)

of the barrier reef ecosystems at Carrie Bow Cay,
Belize, and other parts of the Caribbean Sea,
with descriptions of twenty-seven new species
and revision of Heterodrilus. Thalassodrilides

250

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

and Smithsonidrilus. — Zoologica Scripta 1 9:243-
303.

-. 1990b. The marine Tubificidae and Naididae
(Oligochaeta) of Southwestern Austraha. Pp. 43-
88 in F. E. Wells, D. I. Walker, H. Kirkman, &
R. Lethbridge, eds., Proceedings of the Third
International Marine Biological Workshop: the
marine flora and fauna of Albany, Western Aus-
tralia. Volume 1 . Western Australian Museum,
Perth, 437 pp.

-. 1992a. A generic revision of the Phallodri-
linae (Oligochaeta, Tubificidae. — Zoologica
Scripta 21:5-48.

-. 1992b. Hong Kong's marine Oligochaeta: a
supplement. Pp. 157-1 80 in B. Morton, ed., The
marine flora and fauna of Hong Kong and south-
em China III. Proceedings of the Fourth Inter-
national Marine Biological Workshop: the ma-
rine flora and fauna of Hong Kong and Southern
China, Hong Kong, 1 1-29 April 1989. Volume

1. Hong Kong University Press, Hong Kong,
526 pp.

Pierantoni, U. 1902. Due nuovi generi di Oligocheti
marini rinvenuti nel Golfo di Napoli.— Bollet-
tino della Societa di Naturalisti i Napoli 16:113-
117.

Swoffbrd, D. L. 1990. PAUP. Phylogenetic Analysis
Using Parsimony. Version 3.0. Computer pro-
gram distributed by the Illinois Natural History
Survey, Champaign.

(CE) Department of Invertebrate Zoolo-
gy, Swedish Museum of Natural History,
Box 50007, S-104 05 Stockholm, Sweden;
(MRM) Mote Marine Laboratory, 1600
Thompson Parkway, Sarasota, Florida
34236, U.S.A.

PROC. BIOL. SOC. WASH.

106(2), 1993, pp. 251-295

A CHECKLIST OF AND ILLUSTRATED KEY TO THE

GENERA AND SPECIES OF THE CENTRAL AND

NORTH AMERICAN CAMBARINCOLIDAE

(CLITELLATA: BRANCHIOBDELLIDA)

Perry C. Holt and Brent D. Opell

Abstract.— This paper treats the 90 species of Cambarincolidae known from
Central and North America. Each species is illustrated and its synonyms, tax-
onomic references, type specimen disposition, and distribution are given. Keys
are provided to the family's nine genera and to the 48 species of Cambarincola,
2 species of Ceratodrilus, 1 species of Ellisodrilus, 4 species of Oedipodrilus, 8
species of Pterodrilus, and 1 8 species of Sathodrilus, Magmatodrilus, Tetto-
drilus, and Triannulata are each represented by a single species.

The branchiobdellidans are obligate epi-
zoites on freshwater crustaceans throughout
the Holarctic region except, apparently, in
Central Asia between the Ural Mountains
and the Amur drainage (Holt 1968a). The
greatest diversity of families and species is
found in North America. This may be an
artifact of collecting, as we have less infor-
mation from other regions except Europe
where only a few species are found. In Asia
only Yamaguchi has done any significant
work on the Japanese and Korean faunas.
Despite the greater body of work on the
North American branchiobdellidans, it is
likely that only a small fraction of this fauna
has been accorded taxonomic treatment.
Most genera and species of North American
branchiobdellidans are assigned to the fam-
ily Cambarincolidae (Holt 1986). The phy-
logenetic position and classification of the
Branchiobdellida is discussed by Gelder &
Brinkhurst (1990), Holt (1986, 1989a), and
Sawyer (1986).

Methods

Types or topotypes of all taxa of the cam-
barincolids have been studied in Holt's lab-
oratory and detailed methods for preserving

and studying specimens are found in Holt
(1986). Additional methods include Gelder
& Hall's (1990) use of a mixture of clove
oil and methyl salicylate (oil of winter-
green), as branchiobdellidans, particularly
gill-inhabiting forms, are often twisted and
distorted when methyl salicylate alone is
used in the preparation of entire animals.
The use of Nomarski differential interfer-
ence contrast optics gives a much clearer
view of internal structures in whole animals
than does bright field illumination. When
time and the availability of specimens per-
mit, the worms should be relaxed with an
appropriate agent, such as chloral hydrate
and, after fixation in alcohol-formalin,
lightly stained. For field collection, alcohol-
formalin (Holt 1 963) still seems the best and
easiest preservative to use.

All of the branchiobdellidan material, in-
cluding types, studied by Holt and their ac-
companying field notes have been deposited
in the collections of the National Museum
of Natural History, Smithsonian Institu-
tion, Washington, D.C., U.S.A. These spec-
imens are identified by the accession num-
bers (USNM) of the National Museum, or
by Holt's accession numbers (PCH . . .). A
few paratypes are in the collection of the

Fig. 1. A, generalized branchiobdellidan in lateral view; Abbreviations as follows: I-XI, Trunk Segments;
B, Bursa; BR, Brain; C, Circulatory System; DR, Dorsal Ridge; DRP, Digital Projections of the Dorsal Ridge;
EBP, Ental Bulb of Prostate; ELS, Ental Lobe of the Seminiducal Gland; EPS, Ental Process of Spermatheca;
ED, Ejaculatory Duct; EP, Everted Penis; EVE, Funnel of Vas Deferens; HDP, Histologically Differentiated
Prostate; INT, Intestine; IPR, Incompletely Separated Prostate; J, Jaws; NP, Nephridiopore; OV, Ovary; PR,
Prostate; PS, Penial Sheath; PT, Peristomal Tentacles; PP, Prostatic Protuberance; PB, Prostate Bulb; RP,
Retracted Penis; SPG, Seminiducal Gland; SB, Bulb of Spermatheca; SD, Spermathecal Duct; SMR, Super-
numerary Muscles of Dorsal Ridge; VN, Ventral Nerve Cord; VD, Vas Deferens (from Holt 1969, 1986); B-C,
eversible penis of Pterodrilus alcicornis; B, withdrawn; C, everted; D-E, eversible penis ofCambarincolapamelae;
D, withdrawn; C, everted; F-G, protrusible penis of Oedipodrilus oedipus; F, withdrawn; G, protruded, H,
withdrawn protrusible penis of Oedipodrilus cuetzalanae; I, protruded protrusible penis of Oedipodrilus macbaini.
Unless otherwise noted, the figures of each species on the following plates are arranged, from left to right, as
follows: lateral view of whole specimen, lateral view of reproductive system, ventral view of upper jaw (top)
and dorsal view of lower jaw (bottom); and lateral views of upper jaw (top) and lower jaw (bottom). Abbreviations
are given above.

VOLUME 106, NUMBER 2

253

Institute de Biologia, Universidad Auto-
noma de Mexico, Ciudad Mexico, D.F.,
Mexico, and indicated by the initials IBUM.

The characters used in the taxonomy of
the branchiobdellidans (Fig. lA) have
changed over the years with the recognition
that structures once thought to be common
to all members of the class are not so and
the discovery of previously unrecognized
features. Summaries, in each case incom-
plete, may be found in Yamaguchi (1934),
Goodnight (1940), Holt (1953, 1960a,
1960b), Hoffman (1963), and Gelder &
Brinkhurst (1990). An example of this is the
use by Gelder & Hall (1990) of the number
of oral papillae to separate species. Previ-
ously these putative sensory structures were,
at best, simply noted as present or absent.
Further studies should be done to confirm
the taxonomic usefulness of these papillae.

In the preparation of this paper all figures
were either redrawn from the originals or
from enlarged illustrations in the literature
in order to achieve uniformity of style. The
correctly constructed term "seminiducal
gland" replaces "spermiducal gland" in the
taxonomic keys. This paper cites primarily
taxonomic papers and its literature cited is,
therefore, not intended to be a complete
bibliography of the branchiobdellidans. The
species names of all crayfish host records
have been updated according to Hobbs
(1989). As the literature citations for these
names are provided by Hobbs, they are not
repeated in this paper. Unless otherwise
noted, all hosts are crayfish.

Nomen inquirendum. — Cambarincola
okadai Yamaguchi, 1933. This nominal
species was described from a specimen in-
troduced with its American host into Lake
Chuzenji, Nikko, Japan. Unfortunately, the
location of the type(s) is unknown and the
description could apply to any one of sev-
eral American species.

The Checklist
Cambarincola ^Wis, 1912
Cambarincola Ellis, 1912:481.

Type species. — Cambarincola macrodon-
ta Ellis, 1912, by original designation.
G^/7(i^r. —Masculine.

Cambarincola acudentatus Holt, 1973
Figs. 2-5

Cambarincola acudentatus Holt, 1 973b: 1 1-
13.

7y/7e5. — Holotype and 1 paratype, USNM
45435, 6 paratypes, PCH 489, 4 paratypes,
IBUM, on the isopods Speocirolana bolivari
and S. pelaezi, from Grutas de Quintero. 1 1
km SW of Ciudad Monte, Tanaulipas, Me-
xico, 14 May 1950, by Alejandro Villalo-
bos.

Distribution. —Known only from the type
locality.

Cambarincola alienus Holt, 1963
Figs. 6-10

Cambarincola aliena Holt, 1963:97-100.
Cambarincola alienus.— Holt, 1973c: 10.

ry/7^5. -Holotype, USNM 30415, 7
paratypes, USNM 30416, 7 paratypes, PCH
1325, on the isopod Asellus alabamensis
(Stafford), from Wet Cave, Franklin Co.,
Tennessee, by T. C. Barr [date lost].

Distribution. —Known only from the type
locality.

Notes: Cambarincola steevesi Holt, 1 973c,
may be conspecific with or a local race of
C aliena.

Cambarincola barbarae Holt, 1981
Figs. 11-14

Cambarincola barbarae Holt, 1981a:677-
679.

ry/7^5. -Holotype, USNM 54639, 5
paratypes, PCH 1101, on Procambarus
(Scapulicambarus) clarkii (Girard), from ir-
rigation ditch 3.1 mi E of Salvang, Santa
Barbara Co., California, by Perry C. and
VirgieF. Hoh, 3 Jul 1960.

Distribution.— Santa. Barbara and Sono-
ma Counties, California.

Notes: ''Cambarincola barbarae occurs

254

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

pr ebp

C. acudentatus

C. alienus

C. bobbi

C. branchiophilus

eps

8
9

Figs. 2-22. 2-5, Cambarincola acudentatus Holt, 2, ventro-lateral view; 6-10, Cambarincola alienus Holt;
11-14, Cambarincola barbarae Holt; 15-18, Cambarincola bobbi Holt; 19-22, Cambarincola branchiophilus
Holt.

VOLUME 106, NUMBER 2

255

Upon an introduced species (Hobbs, 1972:
72) which also serves as a host for C. me-
sochoreus Hoffman, 1963, and C. fallax
Hoffman, 1963. The former species is wide-
spread in the Mississippi Valley and the lat-
ter in the Appalachians and were possibly
introduced with their host. This may be so
for C. barbarae'' (Holt 1981a:679).

Cambarincola bobbi Holt, 19SS
Figs. 15-18

Cambarincola bobbi Hoh, 1988b:794-808.

Types. -HoXoXy^Q, USNM 101496, 4
paratypes, USNM 101497-101499, on
Cambarus bartonii bartonii (Fabricius), from
a medium-sized stream in Tom's Brook (a
town) 5.7 mi S of Strasburg, Shenandoah
Co., Virginia, by Marvin L. Bobb and Perry
C. Holt, 22 Jul 1948.

Distribution. —Known only from the type
locality.

Cambarincola branchiophilus Holt, 1954
Figs. 19-22

Cambarincola branchiophila Holt, 1954:
168-172.-Hoffman, 1963:317-319.

Cambarincola branchiophilus. —Holt,
1973b:10.

Types. -HoXoXy^Q, USNM 25855, 6
paratypes, PCH 407 USNM, on Cambarus
bartonii bartonii (Fabricius), and C. sci-
otensis Rhoades, from Sinking Creek at
crossing of State Highway 700, Giles Co.,
Virginia, by Frank D. Kiser, Cornelia Tuten
and Perry C. Holt, 3 Jul 1950.

Distribution. — Known only from the type
locality.

Cambarincola carcinophilus Holt, 1973
Figs. 23-28

Cambarincola carcinophilus Holt, 1973b:
13-14.

Types. -WoXoXy^Q, USNM 45439, 1
paratype, IBUM, 1 paratype, PCH 698, on

the freshwater crab Pseudothelophusa ve-
racruzana, from Rio Tapalapa, Santiago,
Tuxtla, Mexico, by Alejandro Villalobos and
Horton H. Hobbs, Jr., 1957.

Distribution.— Two locations in the low-
lands of Southern Veracruz.

Notes: The specimens taken in 1957 by
Villalobos and Hobbs from the Arroyo de
Zapoapan de Cabana were associated with
not only the crab P. veracruzana, but also
with the crayfish Procambarus zapoapensis
Villalobos.

Cambarincola chirocephala Ellis, 1919
Figs. 29-34

Cambarincola chirocephala Ellis, 1 9 1 9:26 3-
264. -Goodnight, 1940:37-38. -Holt &
Hoffman, 1959:103. -Hoffman, 1963:
348-351.

Cambarincola chirocephalus. —Holt,
1973b:9 [unjustified emendation].

Types. -Holotype, USNM 1 77 1 3, on Or-
conectes virilis Hagen, from Rolla, Phelps
Co., Missouri, by J. Barley, date unknown.

Distribution. —¥r ova western Tennessee
north to Indiana, westward to Oklahoma
and Kansas.

Notes: Cambarincola chirocephala ap-
pears to intergrade with C philadelphicus
in western Tennessee (Hoffman 1963:345).

Cambarincola demissus Hoffman, 1963
Figs. 35-36

Cambarincola demissa Hoffman, 1963:365-

367.
Cambarincola demissus. —Holt, 1 973b: 10.

Types. —Holotype and 4 paratypes,
USNM 29948, on Orconectes erichsonianus
Faxon, and O. rusticus (Girard), from the
Powell River at Big Stone Gap, Wise Co.,
Virginia, by H. H. Hobbs, Jr. and C. W.
Hart, Jr., 17 Jun 1950.

Distribution. —Mountain regions of
southwestern Virginia.

256

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Cambarincola dubius Holt, 1973
Figs. 37-42

Cambarincola dubius Holt, 1 973c:234-236.

ry;7^5. -Holotype, USNM 49673, 2
paratypes, USNM 49674, 4 paratypes, PCH
2763, on Cambarus laevis Faxon, and Or-
conectes inermis testii (Hay), from May's
Cave, Monroe Co., Indiana, by Horton H.
Hobbs, III, 20 Sep 1969.

Distribution. —CavQS in Monroe Co., In-
diana.

Cambarincola ellisi Holt, 1973
Figs. 43-48

Cambarincola ellisi Holt, 1973b: 14-1 6.

r>;/7^5. -Holotype, USNM 45433, on
Procambarus regiomontanus Villalobos,
from Rio San Juan, San Juan, Nuevo Leoni,
Mexico, by Salvador Contreras, 18 Apr
1964. One paratype, PCH 1844, on P. re-
giomontanus, from Arroyo de la Cruz, km
245 de la carretera Ciudad Mexico-Mon-
terey, by A. Villalobos, 14 Feb 1964.

Distribution. —Known only from the type
locality.

Notes: Cambarincola ellisi is known from
a tributary on the Rio Grande (Rio Bravo)
and is geographically nearer the branchiob-
dellidans of the United States than of Mex-
ico.

Cambarincola fallax IrLoffrmin, 1963
Figs. 49-54

Cambarincola fallax Hoffman, 1963:256-
259.-Hobbs et al., 1967:54-55. -Holt,
1969:207; 1973c:238; 1981a:679-680.

Types. — Holotype and 4 paratypes,
USNM 29945, additional paratypes, PCH
904, on '^Cambarus longulus subsp." [either
C. longirostris Faxon, or C longulus Gi-
rard], from Maiden Spring Creek, Tazewell
Co., Virginia, by Richard L. Hoffman, 19
Jun 1959.

Distribution. —Appalachian Mountains
from Canada to Georgia; one specimen from

Sonoma Co., California. In the southern
parts of its range this species is confined to
higher elevations.

Notes: The specimen from California rep-
resents either an introduction, or less likely
a new, but closely related species.

Cambarincola floridanus Goodnight, 1941
Figs. 55-60

Cambarincola floridanus Goodnight, 1941:
73-74. -Hoffman, 1963:368. -Holt,
1973a:90-93.

r>;;7^5. -Holotype, USNM 20570, on
Procambarus fallax (Hagen), collected 6.8
mi S of Lamont, Taylor Co., Rorida, by
Horton H. Hobbs, Jr., 18 Mar 1939.

Distribution. —ThQ type locality and one
locality in Liberty Co., Florida.

Notes: In the paragraph beginning "The
spermiducal gland . . ." (Hoh 1973a:92) a
lapsus resulted in "ectally" being written
instead of the obviously correct "entally."

Cambarincola goodnighti Holt, 1973
Figs. 61-66

Cambarincola goodnighti Holt, 1973a:88-
90.

ry;?^^. -Holotype, USNM 49706, 1
paratype, USNM 48707, 1 paratype, PCH
2607, on Procambarus fallax (Hagen), and
P. paeninsulanus (Faxon), collected 5 mi N
of St. Augustine, St. Johns Co., Florida
(Collector and date unknown).

Distribution. —Known only from the type
locality.

Cambarincola gracilis Robinson, 1954
Figs. 67-70

Cambarincola gracilis Robinson, 1 954:467-
468.-Hoh& Hoffman, 1959:97-103.-
Hoffman, 1963:369. -Holt, 1981a:680-
685.-Gelder&Hall, 1990:2354.

ryp^5. —Holotype and 10 paratypes,
USNM 26110, on Pacifastacus leniusculus
kalamathensis (Stimpson), from creek on

VOLUME 106, NUMBER 2

257

C. carcinophilus
dr.

C. chirocephala

C. demissus

C. dubius

27
28

33
34

41
42

Figs. 23-48. 23-28, Cambarincola carcinophilus Holt; 29-34, Cainbarincola chirocephala Ellis; 35-36, Cam-
barincola demissus Hoffman; 37^2, Cambarincola dubius Holt; 43^8, Cambarincola ellisi Holt.

258

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

campus of Whitman College, Walla Walla,
Walla Walla Co., Washington, by A. G.
Rempel, 1 1 Aug 1952; 2 paratypes, USNM
26 1 1 1 , on P. /. klamathensis, from Klamath
River, 1 mi W of Ash Creek, Siskiyou Co.,
California, by Harold Wolf, 27 Oct 1952; 1
paratype, USNM 261 12, on P. I. trowbridgii
(Stimpson), from Burdette Creek, Bumaby,
British Columbia, by G. CHfford Carl, 21
May 1942.

Distribution.— ¥r ova southern California
to southern British Columbia in streams of
the Pacific versant.

Notes: Holt's statement (Hoh 198 la:682)
that the prostate is subequal in length and
diameter to the spermiducal gland is an in-
explicable error (see Holt 1 98 la:68 1 , fig. 2).
Cambarincola gracilis is the second most
common species of branchiobdellidan in
collections from the Pacific drainage in the
United States and Canada.

Cambarincola heterognathus Hoffman,

1963

Figs. 71-76

Cambarincola heterognatha Hoffman, 1963:

362-365.
Cambarincola heterognathus. —Holt, 1 973a:

95; 1973b:10.

Types. — Holotype and 1 paratype, USNM
29947, on Cambarus sp., from a tributary
to Big Wilson Creek, 4 mi N of Mouth of
Wilson, on State Highway 16, Grayson Co.,
Virginia, by Horton H. Hobbs, Jr., and C.
W. Hart, 14 Jun 1950.

Distribution. —From northwestern Vir-
ginia and adjacent West Virginia south and
west to the central eastern parts of Kentucky
and Tennessee; disjunct populations in Leon
and Calhoun Counties, Florida.

Cambarincola hoffmani Holt, 1973
Figs. 77-82

Cambarincola hoffmani Holt, 1 973b: 1 6-1 7.

ry;7^5. -Holotype, USNM 45447, on
Procambarus hoffmanni (Villalobos), from

Arroyo de Tlatentilojan, at Los Estajos, 6
km NE of Zihuateutla, Puebla, Mexico, by
Alejandro Villalobos, 1 1 Nov 1949; 2 para-
types, PCH 1622, on Procambarus cabal-
leroi Villalobos, from Villa Juarez, Puebla,
Mexico, by Alejandro Villalobos, May 1944.

Distribution. — Kjiown only from the type
locality.

Notes: Cambarincola hoffmani may be an
inhabitant of the gill chambers of its hosts
(Holt 1973b: 17). The spelling is correct, as
the species was named from R. L. Hoffman,
not for the host crayfish.

Cambarincola holostoma Hoffman, 1963
Figs. 83-87

Cambarincola holostoma Hoffman, 1963:

359-361.
Cambarincola holostomus. —Holt, 1 973b: 1

[unjustified emendation].

Types. —Holotype and 4 paratypes,
USNM 29946, 2 paratypes, PCH 599, on
"Cambarus bartoniV [probably C b. ca-
vatus Hay] and C longulus Girard, from
Crab Run, Highland Co., Virginia, by L. B.
Holthuis, 25 Oct 1952.

Distribution. —Western Virginia in the
headwaters of the Potomac and James Riv-
ers (Hobbs et al. 1967:57).

Cambarincola holti Hoffman, 1963
Figs. 88-89

Cambarincola holti Hoffman, 1963:314-
316.

Types. —Holotype and paratypes, USNM
29940, on Cambarus sp., from a stream in
the southern part of Somerset, Pulaski Co.,
Kentucky, by Perry C. and Virgie F. Holt,
28 Jul 1958.

Distribution. —Known only from the type
locality.

Cambarincola illinoisensis Holt, 1982
Figs. 90-93

Cambarincola illinoisensis Holt, 1982:25 1-

255.

VOLUME 106, NUMBER 2

259

C. fallax

C. floridanus

C. goodnighti

C. heterognathus 72

51 53

Figs. 49-76. 49-54, Cambarincola fallax Hoffman; 55-60, Cambarincola floridanus Goodnight; 61-66,
Cambarincola goodnighti Holt; 61-lQ, Cambarincola gracilis Robinson; 71-76, Cambarincola heterognathus
Hoffman.

260

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Types. -HolotypQ, USNM 65225, 3
paratypes, USNM 65226, 5 paratypes, PCH
840, on Orconectes virilis Hagen, from a
prairie stream (possibly one of the two Sugar
or two Mud Creeks that drain into the Ir-
oquois River) N of Stockland, Iroquois Co.,
lUinois, by Perry C. and Virgie F. Holt, 25
Jul 1958.

Distribution. —Known only from the type
locality.

Notes: The protruded penes of C illinois-
ensis are similar in some respects to those
of species of Sathodrilus, but there are not
enough data now to unite these genera (see
Holt 1982:254-255).

Cambarincola ingens Hoffman, 1963
Figs. 94-97

Cambarincola ingens Hoffman, 1963:333-
336.

Types. — Holotype and 2 paratypes,
USNM 29944, on Cambarus sciotensis
Rhoades, from Sinking Creek, Giles Co.,
Virginia, by Ben I. Johns, 27 Jun 1953.
Topotypes, taken by others, PCH 234, 407,
499.

Distribution. —The southern Appala-
chians in North Carolina, Tennessee, Vir-
ginia, and West Virginia.

Cambarincola jamapaensis Holt, 1973
Figs. 98-103

Cambarincola jamapaensis Holt, 1 973b: 1 7-
20.

7ype'5.— Holotype and 5 immature para-
types, USNM 45438, 1 paratype, PCH 1592,
on Procambarus mexicanus (Erichson), from
the Rio Jamapa, 7 km NE of Coscomatepec,
Veracruz, Mexico, by Perry C. and Virgie
F. Holt, 9 Jul 1962.

Distribution. —Known with certainty only
from the type locality; specimens from two
localities in Puebla, Mexico, have tenta-
tively been assigned to this species (Holt
1973b:20).

Cambarincola leoni Holt, 1973
Figs. 104-106

Cambarincola leoni Holt, 1973c:226-229.

r;;/7^5. -Holotype, USNM 49676, 1
paratype, USNM 40677, on Procambarus
orcinus Hobbs & Means, from Gopher Sink,
Leon Co., Florida, by D. Bruce Means and
Joseph Halusky, 3 Apr 1971; 5 paratypes,
PCH 2756, on P. orcinus, from Gopher Sink,
by D. Bruce Means and J. F. Berry, 26 Feb
1971.

Distribution.— Caves in Leon, Alachua
and Marion Counties, Florida.

Cambarincola leptadenus Holt, 1973
Figs. 107-111

Cambarincola leptadenus Holt, 1973c:231-
234.

Types. -HolotypQ, USNM 49678, 1
paratype, USNM 49679, 2 paratypes, PCH
2739, on Cambarus tenebrosus Hay, from
Bethel Cave, Perry Co., Tennessee, by Hor-
ton H. Hobbs, III, 6 Oct 1968.

Distribution. —Known only from the type
locality.

Cambarincola macrocephala Goodnight,

1943

Figs. 112-116

Cambarincola macrocephela (inadvertent

misspelling) Goodnight, 1943:100-101.
Cambarincola macrocephala. —Holt &

Hoffman, 1959:103. -Hoffman, 1963:

312-314.
Cambarincola macrocephelus. —Holt,

1973b: 10 [unjustified emendation]; Holt,

1981a:685, fig. 3.
Cambarincola macrocephalus. —Holt,

1981a:685-689.

Types. -Holotype, USNM 20598, on Pa-
cifastacus gambelii (Girard), from Polecat
Creek, Teuton Co., Wyoming, by Robert C.
Brown, 16 Aug 1941.

Distribution. —The, upper reaches of the

VOLUME 106, NUMBER 2

261

C. hoffmani

C. holostoma

C. hoiti

C. illinoisensis

eps

C. ingens

Figs. 11-97. 77-82, Cambarincola hoffmani Holt; 83-87, Cambarincola holostoma Hoffman; 88-89, Cam-
barincola holti Hoffman; 90-93, Cambarincola illinoisensis Holt; 94-97, Cambarincola ingens Hoffman.

262

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

C. jamapaensis

C. leptadenus

105

108

113

C. macrocephala

O C. macrodontus 118

100

101

114

<
115

119

120

102

103

106
109 110

111

116

121

122

Figs. 98-122. 98-103, Cambarincola jamapaensis Holt; 104-106, Cambarincola leoni Holt; 107-1 11, Cam-
barincola leptadenus Holt; 112-116, Cambarincola macrocephala Goodnight; 117-122, Cambarincola macro-
dontus Ellis.

VOLUME 106, NUMBER 2

263

Snake River in Idaho and Wyoming (Holt
1981a:688).

Cambarincola macrodontus Ellis, 1912
Figs. 117-112

Cambarincola macrodonta Ellis, 191 2:48 1-
484; 1919:257. -Hall, 1914:190.-
Goodnight, 1940: 31. -Holt & Hoffman,
1959:97._Hoffman, 1963:352.

Cambarincola macrodontus. —Holt,
1973b:9.

Types. — Holotype and 2 paratypes,
USNM 53794, on Cambarus diogenes Gi-
rard, from a stream in Boulder, Boulder Co.,
Colorado, by Max M. Ellis, date unknown.

Distribution. —The high plains of the cen-
tral United States, doubtfully from Las Ve-
gas, Nevada (see Hoffman 1963:353-354).

Cambarincola manni Holt, 1973
Figs. 123-128

Cambarincola manni Holt, 197 3a: 8 5-8 8.

ry/7e^5.— Holotype, 9 paratypes, USNM
48700, on Procambarusfallax(tiagQn), from
Lake Martha, at 6 1 2 E. Lake Martha Drive,
Winter Haven, Polk Co., Florida, by Ches-
ter A. Mann, 12 Jan 1964; 1 paratype,
USNM 48701, on P. fallax, from canal be-
tween Lake Buckeye and Lake Fanny, Win-
ter Haven, Polk Co., Rorida, by Chester A.
Mann and Perry C. Hoh, 20 Apr 1963; 1
paratype, PCH 1670, on P. fallax, from Lake
Martha, Winter Haven, Polk Co., Florida,
by Chester A. Mann and Perry C. Holt, 20
Apr 1963; 10 paratypes, PCH 1793, from
the type locality; 5 paratypes, PCH 1673,
on Procambarus paeninsulanus (Faxon),
from slough, 1 3 mi NW of Inglis, Levy Co.,
Florida, by Perry C. and Virgie F. Holt, 26
Apr 1963.

Distribution. —Florida peninsular.

Notes: Cambarincola manni appears to
be confined to peninsular Florida and is the
only branchiobdellidan from the southern
portion of the state, but overlaps C. osceolai

in the northern part of the state (see Holt
1973a:88, fig. 1).

Cambarincola marthae Holt, 1973
Figs. 129-134

Cambarincola marthae Holt, 1973c:221-

224.

Types. -Holoty^t, USNM 49509, 1
paratype, PCH 2161 , on the isopod Asellus
alabamensis (Stafford), from Carter's Cave,
Jackson Co., Tennessee, by John E. and
Martha R. Cooper, 21 Sep 1968.

Distribution. —Known only from the type
locality.

Cambarincola mesochoreus Hoffman, 1963
Figs. 135-137

Cambarincola vitrea (in part).— Ellis, 1919:

257-258.
Cambarincola macrodonta (in part).— Ellis,

1919:257.
Cambarincola mesochorea Hoffman, 1963:

307-311.
Cambarincola mesochoreus. —Holt, 1973b:

10; 1981a:689.

Types. —Holotype and 4 paratypes,
USNM 29934, additional paratypes, PCH
815, on Orconectes sp., from stream 1.5 mi
E of Adyville, Perry Co., Indiana, by Perry
C. and Virgie F. Holt, 28 Jul 1958.

Distribution. — Central Mississippi drain-
age with disjunct populations in Massachu-
setts and California (Hoffman 1963:308,
Hoh 1981a:689).

Notes: In Hoh, 1981a:689, the last entry
in the synonymy should be ^^Cambarincola
mesochoreus.— Holt, 1973b:10" instead of
"1963:10."

Cambarincola meyeri Goodnight, 1 942
Figs. 138-143

Cambarincola m^ym Goodnight, 1942:272-
273. -Hoffman, 1963:354.-Holt, 1973d:
678-682.

264

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

125 ^ 127

.««\i 128

C. manni

C. marthae

C. mesochoreus

C. meyeri

130

136

139

C. micradenus ''^^

131

132

146

147

133

134

148

149

Figs. 123-149. 123-128, Cambarincola manni Hoh; 129-134, Cambarincola marthaeHoW, 135-137, Cam-
barincola mesochoreus Hoffman, 137, dorsal view of reproductive system, showing recurved prostate; 138-143,
Cambarincola meyeri Goodnight; 144—149, Cambarincola micradenus Holt.

VOLUME 106, NUMBER 2

265

Types. -Hololype, USNM 20597, on
''Cambarus bartoniV [probably C. b. bar-
tonii (Fabricius)], from Raven's Creek, Fay-
ette Co., Kentucky, by Marvin C. Meyer
(date unknown).

Distribution. —Raven Run (="Raven's
Creek"), Fayette Co., Kentucky (Holt 1 973d:
681).

Cambarincola micradenus Holt, 1973
Figs. 144-149

Cambarincola micradenus Holt, 1973b:20-

22.

Types. — Holotype and 2 paratypes,
USNM 45448, 1 paratype, PCH 1615, on
Procambarus paradoxus (Ortmann), from
La Canada y Tetela de Ocampo, Puebla,
Mexico, by Alejandro Villalobos, May 1944.

Distribution. —Known only from the type
locality.

Cambarincola montanus (Goodnight), 1940
Figs. 150-153

Triannulata montana Goodnight 1940:57-

58.
Cambarincola montanus. —Holt, 1974:67-

70; 1981a:690-691.

Types. -Holotype, USNM 2056, on Pa-
cifastacus sp., from the Kalami River,
Washington (Collector and date unknown).
Paratypes were left at the University of Il-
linois in the collections of H. J. Van Cleave
(Goodnight 1940:58).

Distribution. —Streams of the Coastal and
Cascade Ranges of the Pacific drainage in
western North America from Santa Barbara
Co., California to northern Washington
(Holt 1981a:690).

Cambarincola nanognathus Holt, 1973
Figs. 154-159

Cambarincola nanognathus Holt, 1973b:

22-23.

ry/7^5. -Holotype, USNM 45444, 1
paratype, PCH 1830, 1 paratype, IBUM, on

the freshwater crab Potamocarcinus nica-
raguensis, from Lago de Nicaragua (Isleta
de Granada), G. Alvilez, 13 Jul 1964.

Distribution.— From southern Veracruz,
Mexico, to Nicaragua.

Cambarincola olmecus Holt, 1973
Figs. 160-163

Cambarincola olmecus Holt, 1973b:24-26.

r>^;7^5. -Holotype, USNM 45445, 1
paratype, USNM 45446, 1 paratype, PCH
201, 1 paratype, IBUM, on Procambarus
mexicanus (Erichson), from Tomatlan, Ve-
racruz, Mexico, by Alejandro Villalobos, 3
Nov 1948.

Distribution. -'Sidin Luis Potosi and Ve-
racruz, Mexico.

Cambarincola osceolai Hoffman, 1963
Figs. 164-165

Cambarincola osceola Hoffman, 1963:330-

333.
Cambarincola osceolai.— Holt, 1973a:93-

95; 1973b:10.

Types. —Holotype and 7 paratypes,
USNM 29943, on Procambarus paeninsu-
lanus (Faxon) and Fallicambarus uhleri
(Faxon) from Dry Creek, 3.1 mi N of Iron
City, Seminole Co., Georgia, by Horton H.
Hobbs, Jr. and C. W. Hart, Jr., 9 Sep 1955.

Distribution. -From southeastern Vir-
ginia to northwest peninsular Florida.

Notes: Attention may again be called to
the suspicions of both Hoffman (1963:331)
and Hoh (1973a:94) that C. osceolai and C
vitreus Ellis, 1918:51 are conspecific.

Cambarincola ouachita Hoffman, 1963
Figs. 166-167

Cambarincola ouachita Hoffman, 1963:
303-305.

Types. -Holotype, USNM 29937, on Or-
conectes sp., from a small stream 4.3 mi W
of the Montgomery Co. line in Chautaugua

266

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Co., Kansas, 8 Jul 1958, by Perry C. and
Virgie F. Holt.

Distribution. —Known only from the type
locality.

Notes: Holt (1973b:9-10) changed the
endings of several species to conform to the
masculine gender of the generic name Cam-
barincola. He did not emend the name C
Ouachita which may be considered a noun
in apposition, referring to the Ouachita
Mountains. Hoffman gives no derivation for
the name ouachita, but Holt knows he meant
it to refer to the mountains.

Cambarincola pamelae Holt, 1984
Figs. 168-171

Cambarincola pamelae Holt, 1984b:544-
549.

Types. -Yio\oXyx>Q, USNM 080687, 4
paratypes, USNM 080688-080691, 10
paratypes, PCH 4065, on Procambarus
clarkii (Girard), from an irrigation canal that
drains into the San Joaquin River in the
western part of Stanislaus Co., California,
by J. A. Meeuwse, 2 Dec 1982.

Distribution. —Stanislaus, Santa Barbara,
Merced and Sonoma Counties, California.

Notes: Cambarincola pamelae has been
found only on P. clarkii, a crayfish species
that has been widely introduced throughout
the United States (Hobbs 1989). Therefore,
it is possible that C. pamelae has also been
introduced into its present range and that it
may be a geographical variant of C meso-
choreus, to which it is similar (Holt 1984b).

Cambarincola philadelphicus (Leidy, 1851)
Figs. 172-177

Astacobdella philadelphica Leidy, 1851:209.
Branchiobdella philadelphica. —Moore,

1893:427-428.
Bdellodrilus philadelphicus. — Moore, 1895:

497; 1901:542.-Pierantoni, 1912:17.
Cambarincola philadelphica. — Ellis, 1912

484.-Hall, 19 14: 190. -Ellis, 1918:49

1919:260-263. -Goodnight, 1939:11

1940:38. -Holt, 1954:169. -Holt &
Hoffman, 1959:103. -Hoffman, 1963:
341-348.
Cambarincola philadelphicus. —Holt,
1973b:9.

Types. — Holotype, lost. Topotypes, PCH
695, from Wissahicon Creek in Philadel-
phia, Pennsylvania (Hoffman 1963:342).

Distribution. —From Minnesota and New
York south through the Appalachians and
Piedmont to South Carolina (Hoffman 1963:
344).

Notes: This species is reported in the lit-
erature as unusually variable. Indeed, in
southwestern Virginia it appears to be so,
but many references to it undoubtedly apply
to other related forms. The problem of vari-
ability in C philadelphicus should be re-
studied.

Cambarincola restans Hoffman, 1963
Figs. 178-179

Cambarincola re5^a«5 Hoffman, 1963:305-
307.

Types. —Holotype and 2 paratypes,
USNM 29938, on Orconectes sp., from Sug-
ar Creek, 2 mi N of Avoca, Benton Co.,
Arkansas, by Perry C. and Virgie F. Holt,
6 Jul 1958.

Distribution. —Known only from the type
locality.

Cambarincola serratus Holt, 1981
Figs. 180-183

Cambarincola serratus Holt, 1981a:691-
693.

r>;p^5. -Holotype, USNM 54638; 3
paratypes, PCH 795, on Pacifastacus con-
nectens (Faxon), from Idaho State Fish
Hatchery, Riley Creek, Gooding Co., Idaho,
by Perry C. and Virgie F. Hoh, 14 Jul 1958;
2 paratypes, PCH 784, on P. connectens,
from spring tributary to Snake River, Ha-
german, Gooding Co., Idaho, by Perry C.
and Virgie F. Holt, 14 Jul 1958.

VOLUME 106, NUMBER 2

267

152
153

C. montanus

154 -C)

C. nanognathus 155

156

.Ui

£>>

157

159

160
C. olmecus ''"''

162

163

C. osceolai

165

C. Ouachita

167

Figs. 150-167. 150-153, Cambarincola montanus Goodnight; 154-159, Cambarincola nanognathus Holt;
160-163, Cambarincola olmecus Holt; 164-165, Cambarincola osceolai Hoffman; 166-167, Cambarincola
Ouachita Hoffman.

268

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Distribution. —TribulanQS of the Snake
River, Gooding Co., Idaho.

Cambarincola sheltensis Holt, 1973
Figs. 184-189

Cambarincola sheltensis Holt, 1973c:229-
231.

Types. -\lo\oXyvQ, USNM 49683, 2
paratypes, PCH 1846, on Orconectes aus-
tralis australis (Rhoades), from Shelta Cave,
Hunts ville, Madison Co., Alabama, by John
E. and Martha Cooper, 24 Aug 1 963; 2 para-
types, USNM 49684, 1 paratype, PCH 1863,
on O. a. australis, from Shelta Cave, Mad-
ison Co., Alabama, by James E. Larimer,
1965.

Distribution. —Known only from the type
locality.

Cambarincola shoshone Hoffman, 1963
Figs. 190-191

Cambarincola shoshone Hoffman, 1963:
3 19-320. -Holt, 1981a:693.

Types. — Holotype and 3 paratypes,
USNM 29941, topotypes, PCH 785, on Pa-
cifastacus connectens (Faxon), from Riley
Creek (Idaho State Fish Hatchery), Hager-
man, Gooding Co., Idaho, by Perry C. and
VirgieF. Holt, 14 Jul 1958.

Distribution. —Known only from the type
locality.

Cambarincola smalleyi Holt, 1964
Figs. 192-195

Cambarincola smalleyi Holt, 1964:1-4;
1973b:26-27.

Typ^^. -Holotype, USNM 20940, 1
paratype, 30941, 1 paratype in the collec-
tions of Tulane University, 1 paratype, PCH
1702, on the freshwater crab Pseudothel-
phusa tuminenus, from Rio Hondura, 8 mi
N of San Jeronimo de Moravia, San Jose
Province, Costa Rica, by A. E. Smalley, 9
Jul 1962.

Distribution. — Known only from the type
locality.

Notes: Holt (1964:3) postulated that C
smalleyi was carried to Costa Rica by cam-
barines and qualified (Holt 1973b:27) this
conclusion by suggesting that the transfer to
crabs could have occurred much further
north in Mexico. In truth we know too little
to make any credible conjectures about the
ancient origin and subsequent wanderings
of branchiobdellidans. They may as easily
have originated as symbionts of freshwater
crabs and shrimps and later taken up a sym-
biotic relationship with crayfishes. Cam-
barincola smalleyi is the southernmost
known species of the family.

Cambarincola speocirolanae Holt, 1984
Figs. 196-199

Cambarincola speocirolanae Holt, 1984a:
36-38.

ry/7^5. -Holotype, USNM 80221, 1
paratype, PCH 4054, on the isopod Speo-
cirolana palaezi, from Sotao del Arroyo, San
Luis Potosi, Mexico, by Peter Sprouse, 22
Feb 1980.

Distribution. —Known only from the type
locality.

Cambarincola steevesi Holt, 1973
Figs. 200-203

Cambarincola steevesi Holt, 1 973c:224-226.

r>;p^5. -Holotype, USNM 40680, 2
paratypes, USNM 40681, 1 paratype, PCH
1883, on the isopod Asellus alabamensis
(Stafford), from Glover's Cave, Todd Co.,
Kentucky, by R. M. Norton, 17 Apr 1964;
1 paratype, USNM 49682, on Asellus ala-
bamensis, from Brown Cave, Barren Co.,
Kentucky, by R. M. Norton, 25 Sep 1965.

Distribution. —The type locality and Tay-
lor Cave, Trigg Co., Kentucky.

Cambarincola susanae Holt, 1973
Figs. 204-207

Cambarincola susanae Holt, 1973b:27-29.

VOLUME 106, NUMBER 2

269

168

C. pamelae

169

170

171

173

C. philadelphicus

174 176

175

177

C. restans

179 —^ /-^

C. serratus

C. sheltensis

185

182

186

183 O^

^J 188

187

189

Figs. 1 68-189. 1 68-17 1 , Cambarincola pamelae Holt; 1 72-177, Cambarincola philadelphicus (Leidy); 1 78-
179, Cambarincola restans Hoffman; 180-183, Cambarincola serratus Holt; 184-189, Cambarincola sheltensis
Holt.

270

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ry/?^^. — Holotype and 2 paratypes,
USNM 45441, 10 paratypes, PCH 1529, 3
paratypes, IBUM, on Procambarus acutus
cuevachicae (Hobbs), from Cueva Chica, El
Pujal, 3 km NE of Valles, San Luis Potosi,
Mexico, by Alejandro Villalobos, 9 May
1950.

Distribution.— In eastern Mexico from
Rio San Juan, Nuevo Leon to western Cam-
peche.

Notes: Cambarincola susanae may be an
ectoparasite in the gill chambers of the host
(Hoh 1973b:29).

Cambarincola toltecus Holt, 1973
Figs. 208-211

Cambarincola toltecus Holt, 1973b:29-31.

rj;^^^. -Holotype, USNM 45436, 2
paratypes, USNM 45437, 2 paratypes,
IBUM, 2 paratypes, PCH 697, on the fresh-
water crab Pseudothelphusa veracruzana,
from Rio Tapalapa, Santiago, Tuxtla, Ve-
racruz, Mexico, by Alejandro Villalobos and
Horton H. Hobbs, Jr., 17 Apr 1957.

Distribution. —Tropical lowlands of Ve-
racruz, Mexico.

Cambarincola virginicus Hoffman, 1963
Figs. 212-215

Cambarincola virginica Hoffman, 1963:

322-323.
Cambarincola virginicus. —Holt, 1 973b: 1 0.

Types.— Holotype and 1 paratype, USNM
29942, on Cambarus acuminatus Faxon,
from a small, slow stream 4.7 mi N of Pe-
tersburg, Chesterfield Co., Virginia, by Mar-
vin L. Bobb and Perry C. Holt, 31 May
1949.

Distribution.— Along the Fall Line in
eastern Virginia (Hoffman 1963:323) and
eastern North Carolina.

Cambarincola vitreus Ellis, 1918
Figs. 216-217

Cambarincola vitrea Ellis, 1918:49-51;
1919:257-258. -Goodnight, 1940:33-34;

1943:100. -Holt & Hoffman, 1959:
103. -Hoffman, 1963:324-329.
Cambarincola vitreus.— Holt, 1973b:9.

Types. -Holotype, USNM 17667, on Or-
conectes immunis (Hagen), from Douglas
Lake, Cheboygan Co., Michigan, by Max
M. Ellis, Jul 1915.

Distribution.— Western portions of the
upper Mississippi valley from north and west
of Arkansas.

Notes: See Cambarincola osceola.

Ceratodrilus Kail, 1914

Ceratodrilus Hall, 1914:190-191.
Ceratodrilus Stephenson, 1930:901.
Cirrodrilus Goodni^t, 1940:63-64 [in part].

Type species. — Ceratodrilus thysanoso-
mus Hall, 1914, by original designation.
Gender. —Masculine.

Ceratodrilus ophiorhysis Holt, 1960
Figs. 218-221

Ceratodrilus orphiorhysis Holt, 1960a:60

(incorrect spelling).

Ceratodrilus ophiorhysis. — Holt, 1 9 60a: 6 1 ;

1988a:308.

Types. -Kolotype, USNM 29910, 4
paratypes, USNM 29911,3 paratypes, PCH
786, on Pacifastacus gambelii (Girard), from
the Snake River between Buhl and Wendel,
Gooding Co., Idaho, by Perry C. and Virgie
F. Holt and Judson Ford, 14 Jul 1958.

Distribution.— The Snake River and its
tributaries in Idaho and Oregon.

Ceratodrilus thysanosomus Hall, 1914
Figs. 222-223

Ceratodrilus thysanosomus Hall, 1914:

191. -Stephenson, 1930:801. -Yama-

guchi, 1932:367.
Cirrodrilus thysanosomus. —Goodnight,

1940:64-65 [in part].
Ceratodrilus thysanosomus. —Holt, 1960a:

58-60.

Types. -Holotype, USNM 17708, from

VOLUME 106, NUMBER 2

271

190

C. Shoshone

C. smalleyi

193

194

196

C. speocirolanae '^^^

C. steevesi

205

C. susanae

195

198

199

It r-

202

203

fve 206

207

Figs. 190-207. 190-191, Cambarincola shoshone Hoffman; 192-195, Cambarincola smalleyi Holt; 196-
199, Cambarincola speocirolanae Holt; 200-203, Cambarincola steevesi Holt; 204-207, Cambarincola susanae
Holt.

272

Streams of Great Basin, Salt Lake City, Utah
(no date).

Distribution. —StTQams of the Great Sah
Lake Basin.

Notes: The two species of Ceratodrilus
differ only in the length of their dorsal ap-
pendages (Holt 1960a); their distinctiveness
should be reinvestigated.

Ellisodrilus Holt, 1960

EUisodrilus Holt, 1 960b: 1 70.

Type species. —Ellisodrilus clitellatus Holt,
1960, by original designation.
Gender. —Masculine.

Ellisodrilus carronamus Holt, 1988
Figs. 224-227

Ellisodrilus carronamus, 1988b:796-798.

r>;p^5. -Holotype, USNM 119539, 2
paratypes, USNM 119540-119541, on Or-
conectes sp., from Carr Creek, Overton Co.,
Tennessee, ca. 3 mi S of Livingston, on State
Highway 42, by Perry C. and Virgie F. Holt,
26 Jul 1961.

Distribution.— C^xr Creek and Roaring
River, Overton Co., Tennessee.

Notes: There appears to be a N-S gradient
in Ellisodrilus species with E. carronamus
the southern-most, E. durbini the northern-
most and E. clitellatus found in between.
This occurs in part in areas scoured by the
Pleistocene glaciation (Holt 1 960b: 171,174;
1988b:798).

Ellisodrilus clitellatus Holt, 1960
Figs. 228-231

Ellisodrilus clitellatus Holt, 1 960b: 1 69-1 76.

r>;;?^5. -Holotype, USNM 29935, 1
paratype, USNM 29936, 18 paratypes, PCH
827, on Cambarus distans Rhoades, from
stream 8.9 mi E of Columbia on Kentucky
Highway 80, Adair Co., Kentucky, by Perry
C. and Virgie F. Holt, 28 Jul 1958.

Distribution. —Kentucky.

Notes: The first paragraph on page 171

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(Holt 1 960b) should be transposed to follow
the diagnosis of the genus on page 170.

Ellisodrilus durbini (Ellis, 1918)
Figs. 232-234

Pterodrilus durbini Ellis, 1918:49.— Ellis

1919:254.-Goodnight 1940:61-62.
Ellisodrilus durbini. — Holt, 1 9 6 Ob: 1 7 3 .

Types. -Holotype, USNM 17655, on Or-
conectes barrenensis Rhoades, from White
River, at Irondale, Anderson Co., Indiana.

Distribution. -From. Anderson Co., In-
diana, into Michigan (Ellis 1918:50).

Magmatodrilus Holt, 1967

Stephanodrilus Goodnight, 1940:55 [in

part].
Magmatodrilus Holt, 1967b: 3-4.

Type species.— Magmatodrilus obscurus
(Goodnight, 1 940), by original designation.
Gender. —Masculine.

Magmatodrilus obscurus (Goodnight, 1 940)
Figs. 235-238

Stephanodrilus obscurus Goodnight, 1940:

55-56.
Magmatodrus obscurus. — Holt, 196 7b: 4-5 .

Types. -Holotype, USNM 20568, on Pa-
cifastacus nigrescens (Stimpson) (Good-
night, 1940:55), (collector and date un-
known), from Fall River, Shasta Co.,
California; 2 topotypes, USNM 45696, 11
topotypes, PCH 1818, on unknown host,
from the head of Fall River, Thousand
Springs Ranch, Shasta Co., California, by
Perry C. and Virgie F. Holt, Aug 1964.

Distribution. —Known only from the type
locality.

Notes: Some ambiguity may exist as to
the type locality. Holt was refused permis-
sion to trap for crayfish in Fall River by the
local game warden. The river is a deep, ditch-
like stream that does not lend itself to col-
lecting with a dipnet. After several futile
attempts to do so, the Holts obtained per-

VOLUME 106, NUMBER 2

273

210

C. toltecus

209

C. virginicus

213

C. vitreus

^^.

217

drp

219

C. ophiorhysis

211

214

215

C. thysanosomus

Figs. 208-223. 208-2 1 1 , Cambarincola toltecus Holt; 2 1 2-2 1 5, Cambarincola virginicus Hoffman; 2 1 6-2 1 7,
Cambarincola vitreus Ellis; 218-221, Ceratodrilus ophiorhysis Holt; 222-223, Ceratodrilus thysanosomus Hall.

274

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

mission to take a collection from the spring
from which Fall River arises. The holotype
and topotypes appear to be identical (Holt
1967b:4-5).

Oedipodrilus Holt, 1967

Oedipodrilus Holt, 1967a:58.

Type species. — Oedipodrilus oedipus Holt,
1967a, by original designation.
Gender. —Masculine.

Oedipodrilus anisognathus Holt, 1988
Figs. 239-242

Oedipodrilus anisognathus Holt, 1988b:
798-800.

Types. -YloXoXy^Q, USNM 119534, 1
paratype, USNM 1 19535, on Orconectes sp.,
from a small stream in Montgomery Bell
State Park, Dickson Co., Tennessee, by Per-
ry C. and Virgie F. Hoh, 4 Jul 1958; 3 para-
types, USNM 119536-119537, on Orco-
nectes sp., from Carr Creek, ca. 3.0 mi S of
Livingston, Overton Co., Tennessee, by
Perry C. and Virgie F. Holt, 26 Jul 1961.

Distribution. —Dickson and Overton
Counties, Tennessee, the Central (Nash-
ville) Basin and the eastern Highland Rim,
respectively (Hoh 1988b:800).

Oedipodrilus cuetzalanae Holt, 1984
Figs. 243-246

Oedipodrilus cuetzalanae Holt, 1984a:38-
41.

Type^. -Holotype, USNM 80223, 8
paratypes, PCH 4050, on Procambarus
cuetzalanae Hobbs, from Sima Zoquiapan,
1 . 1 km N of Cuetzalan, Puebla, Mexico, by
L. Wilk, W. Hooper and M. Minton, 2 Jan
1980.

Distribution. —Known only from the type
locality.

Oedipodrilus macbaini (Holt, 1955)
Figs. 247-248

Cambarincola macbaini Holt, 1955:27-31.

Oedipodrilus naacbaini.— Holt, 1969:205;
1984a:39; 1988b:800-804.

ryp^5. -Holotype, USNM 25952, 6
paratypes, PCH 1 34, on Orconectes sp., from
Charles Creek, 8 mi W of Ashland, on State
Highway 5, Boyd Co., Kentucky, by Rod-
ney Macbain, Jul 1948.

Distribution. —Known only from the type
locality.

Oedipodrilus oedipus Holt, 1967
Figs. 249-252

Oedipodrilus oedipus Holt, 1967a:58-60.

Types. -Holotype, USNM 34086, 7
paratypes, USNM 34087, 4 paratypes, PCH
756, on Orconectes compressus (Faxon),
collected 10.2 mi E of Waverly, Humphreys
Co., by Perry C. and Virgie F. Holt, 5 Jul
1958.

Distribution. —Known only from the type
locality.

Pterodrilus Moore, 1894
Pterodrilus Moore, 1894:449-450.

Type species. —Pterodrilus alcicornus
Moore, 1894, subsequent designation by
Goodnight (1940).

Gender. —Masculine.

Pterodrilus alcicornus Moore 1894
Figs. 253-254

Pterodrilus alcicornus Moore, 1894:450-
453.-Pierantoni, 1912:25. -Ellis, 1919:
245. -Goodnight, 1940:58-60. -Holt,
1968c:6-12.

7>pe'5. —Apparently lost (Holt 1968c:6),
but was on Cambarus acuminatus Faxon,
from Johns River at Blowing Rock, Watau-
ga Co., North Carolina, in the summer of
1893, presumably by Moore himself (Moore
1894:453).

Distribution. —The mountain streams of
the Southern Appalachians in Virginia,
North Carolina, Tennessee, and West Vir-

VOLUME 106, NUMBER 2

275

226

227

E. carronamus

229

230

231

E. clitellatus

233

234

spg

235

M. obscurus

237

238

O. anisognathus

240

242

Figs. 224-242. 224-227, Ellisodrilus carronamus Holt; 228-231, Ellisodrilus clitellatus Holt; 232-234, El-
lisodrilus durbini (Ellis), 234, longitudinal section through a dorsal ridge, showing supernumerary muscles; 235-
238, Magmatodrilus obscurus (Goodnight); 239-242, Oedipodrilus anisognathus Holt.

276

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

O. cuetzalanae 244

247 248

O. macbaini

O. oedipus

250

246

251

252

p. alcicornus

254

P. cedrus

256

257

258

Figs. 243-258. 243-246, Oedipodrilus cuetzalanae Holt; 247-248, Oedipodrilus macbaini (Holt); 249-252,
Oedipodrilus oedipus Holt; 253-254, Pterodrilus alcicornus Moore; 255-258, Pterodrilus cedrus Holt.

VOLUME 106, NUMBER 2

277

ginia (Holt 1968c: 12) in which the species
is abundant.

Pterodrilus cedrus Holt, 1968
Figs. 255-258

Pterodrilus cedrus Holt, 1968c:21-23.

Types. — Holotype and 5 paratypes,
USNM 36464, 3 paratypes, PCH 1396, on
Orconectes placidus (Hagen) and Cambarus
tenebrosus Hay, from a small stream at the
intersection of State Highways 5 1 and 53 in
Celina, Clay Co., Tennessee, by Perry C.
and Virgie F. Holt, 25 Jul 1961.

Distribution.— T\iQ eastern Highland Rim
and Nashville Basin regions of Tennessee.

Pterodrilus choritonamus Holt, 1968
Figs. 259-262

Pterodrilus choritonamus Holt, 1968c:26-
28.

r};/7^5. -Holotype, USNM 36471, 2
paratypes, USNM 36472, on Cambarus te-
nebrosus Hay, from a tributary (Holt Spring
Branch) ca. 4.5 mi N of Livingston, Overton
Co., Tennessee, by Perry C. and Virgie F.
Holt, 24 Jul 1961; 5 paratypes, PCH 1393,
on C. tenebrosus and Orconectes placidus
(Hagen), from Little Eagle Creek, Overton
Co., Tennessee, by Perry C. and Virgie F.
Holt, 24 Jul 1961.

Distribution. —TnbuX2ir\Qs of the Cum-
berland River in the eastern Highland Rim
region in Tennessee (see locality records for
Pterodrilus manuscript by Holt in the Li-
brary of Congress and the USNM). Some
conspecific material is deposited in the Na-
tional Museum of Natural History, USNM
36473-36476.

Pterodrilus distichus Moore, 1894
Figs. 263-266

Pterodrilus distichus Moore 1894:453-
454.-Pierantoni, 1912:25. -Hall, 1914:
190. -Ellis, 1919:254.-Goodnight,
1940:60; 1943:100.-Holt, 1968c:12.

Types.— Yrova western New York, loca-
tion unknown (Holt 1968c: 12).

Distribution.— Ohio, Mississippi, Great
Lakes, drainage systems in Kentucky, Ohio,
Indiana, Illinois, Michigan, and New York.

Pterodrilus hobbsi Holt, 1968
Figs. 267-270

Pterodrilus hobbsi HoU, 1968c: 18.

r>;p^5. -Holotype, USNM 36486, 5
paratypes, USNM 36487, on Cambarus
rusticiformis Rhoades, Orconectes rusticus
(Girard), and Orconectes placidus (Hagen),
from Spring Creek, 1.4 mi N of the Putnam
Co. line on State Highway 42, Overton Co.,
Tennessee, by Perry C. and Virgie F. Holt,
26 Jul 1961.

Distribution. —Cumberland River in
Tennessee and Kentucky, the upper Ten-
nessee drainage in Tennessee, the New Riv-
er drainage in Virginia and North Carolina,
the Big Sandy drainage in Virginia (Holt
1968c:20, 38)

Pterodrilus mexicanus Ellis, 1919
Figs. 271-272

Pterodrilus mexicanus Ellis, 1919:254.—
Goodnight, 1940:63. -Holt, 1968c:15;
1973b:32.

r>^/?^5. -Holotype, USNM 17654, on
Procambarus mexicanus (Erichson), from
Mirador, Veracruz, Mexico, by Nelson and
Goldman.

Distribution.— MounXsLins of Arkansas,
Oklahoma, and Missouri.

Notes: The apparent disjunct distribution
of this species is difficult to explain. Hobbs
(1989) lists the type locality of P. mexicanus
as "El Mirador de Zacuapan, 8 km NE of
Huatusco, Veracruz, Mexico." However,
Holt was unable to find P. mexicanus in
Mexico (Hoh 1973b:32).

Pterodrilus missouriensis Holt, 1968
Figs. 273-274

Pterodrilus missouriensis Holt, 1 968c:28-32.

278

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

P. choritonamus

260

P. distichus

264

265

266

268

P. mexicanus 272

273

274

P. missouriensis

269

270

Figs. 259-274. 259-262, Pterodrilus choritonamus Holt; 263-266, Ptewdrilus distichus Moore; 267-270,
Pterodrilus hobbsi Holt; 271-272, Pterodrilus mexicanus Ellis; 273-274, Pterodrilus missouriensis Holt.

VOLUME 106, NUMBER 2

279

Types. -HolotypQ, USNM 36469, 2
paratypes, USNM 36470, 2 paratypes, PCH
1476, on Orconectes luteus (Creaser), from
Whetstone Creek, 5 mi W of Mountain
Grove, Wright Co., Missouri, by Perry C.
Holt, 23 Aug 1961.

Distribution. —Known only from the type
locality.

Pterodrilus simondsi Holt, 1968
Figs. 275-276

Pterodrilus simondsi Holt, 1968c:23-26.

ryp^5. -Holotype, USNM 26477, 5
paratypes, USNM 36478, 4 paratypes, PCH
989, on Cambarus bartonii bartonii (Fabri-
cius), from a tributary to the Oconee River
S of Morganton, Fannin Co., Georgia, by
Kenneth W. Simonds, 6 Nov 1958.

Distribution. —TributSLTies to the Oconee
River in Fannin and Union Counties, Geor-
gia and Cherokee Co., North Carolina.

Sathodrilus Holt, 1968

Sathodrilus Holt, 1968b:294.

Type species.— Sathodrilus carolinensis
Holt, 1968, by original designation.
Gender. —Masculine.

Sathodrilus attenuatus Holt, 1981
Figs. 277-280

Sathodrilus attenuatus Holt, 1981b:849-
853.

r^i^e^. -Holotype, USNM 65227, 26
paratypes, PCH 1 1 13, on Pacifastacus leni-
usculus klamathensis (Stimpson), from Elk
Creek, ca. 12.6 mi S of Cottage Grove,
Douglas Co., Oregon, by Perry C. and Virgie
F. Holt, 11 Jul 1960.

Distribution. —Streams of the Cascade and
Coastal Ranges in Oregon and Washington
to the headwater streams of the Snake River
in Wyoming.

Sathodrilus carolinensis Holt, 1968
Figs. 281-284

Sathodrilus carolinensis Holt, 1968b:296-
299.

Types. -Holotype, USNM 37107, 1
paratype, USNM 37108, 1 paratype, PCH
1333, on Cambarus latimanus (Le Conte)
and Cambarus sp., from a small stream ca.
11.5 mi SW of Anderson, Anderson Co.,
South Carolina, on U.S. Highway 29, by
Perry C. and Virgie F. HoU, 21 Mar 1961.

Distribution. —Known only from the type
locality.

Sathodrilus chehalisae Holt, 1981
Figs. 285-286

Sathodrilus chehalisae Holt, 1981b:853-
855.

Types. — Holotype and 7 paratypes,
USNM 65228, 3 paratypes, PCH 1813, on
Pacifastacus leniusculus trowbridgii (Stimp-
son), from Chehalis River at Adna, Lewis
Co., Washington, by Perry C. and Virgie F.
Holt, 15 Aug 1964.

Distribution. —Known only from the type
locality.

Sathodrilus dorfus Holt, 1977
Figs. 287-290

Sathodrilus dorfus HoU, 1977:120-122.

Types. -Holotype, USNM 53643, 3
paratypes, PCH 1 1 20, on Pacifastacus leni-
usculus klamathensis (Stimpson), from a
small tributary to the Yaguina River, 13.4
km NE of Toledo, Lincoln Co., Oregon, by
Perry C. and Virgie F. HoU, 12 Jul 1960.

Distribution. —Known only from the type
locality.

Sathodrilus elevatus (Goodnight, 1940)
Figs. 291-294

Cambarincola elevata Goodnight, 1 940:34-

35.
Cambarincola ? elevata. —Hof[man, 1963:

275.
Sathodrilus elevatus. —Holt, 1978:473-48 1 .

280

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

7y/7£'5. — Holotype, on Orconectes virilis
Hagen, from Leaf River, Illinois; paratypes,
on O. virilis, from: Macoupin Creek near
Carlinville, Illinois; Buck Creek, near Pen-
field, Illinois; Leaf River, near Bryon, Illi-
nois; Seven Mile Creek, Rock River drain-
age, Illinois; and Lake Geneva, Wisconsin.

Distribution. —The upper Mississippi and
Red Rivers and Great Lakes drainages in
Illinois, Indiana, Iowa, Michigan, Minne-
sota, South Dakota, Wisconsin in the Unit-
ed States and Ontario, Canada (Holt 1978:
478-481).

Notes: The redescription of Sathodrilus
elevatus and the determination that it is con-
specific with Goodnight' s holotype were
based upon the comparison of the holotype
and numerous specimens from the localities
listed by Hoh (1 978:479-48 1). Goodnight's
material, including types, was lost (Good-
night, pers. comm.).

Sathodrilus hortoni Holt
Figs. 295-298

Sathodrilus hortoni Holt, 1973a:97-99.

r>;;7^5. -Holotype, USNM 48713, 2
paratypes, USNM 497 14, 2 paratypes, PCH
2716, on Cambarus diogenes Girard and
Cambarus sp., from Pond Creek, 2. 1 mi NE
of Laurel Hill, Okaloosa Co., Florida, by
Horton H. Hobbs, III, 10 Aug 1968.

Distribution. —Known only from the type
locality.

Sathodrilus inversus (Ellis, 1919)
Figs. 299-302

Cambarincola inversa Ellis, 1919:259-260.
Cambarincola ? zTiv^r^a.- Hoffman, 1963:

294.
Sathodrilus virgiliae. —Holt, 1977:128-131.
Sathodrilus inversus. —Holt, 1981b:855-

856.

ryp^5. -Holotype, USNM 16780, 5
paratypes, USNM 17680, Eugene, Oregon,
on Pacifastacus leniusculus klamathensis

(Stimpson), from Eugene, Oregon, by J. E.
Gutberlet.

Distribution. —Common in streams of the
Pacific versant in Oregon and Washington.

Sathodrilus lobatus Holt, 1977
Figs. 303-306

Sathodrilus labatus Holt, 1977:122-125.

Types. —Holotype and 3 paratypes,
USNM 53644, 5 paratypes, PCH 1117, on
Pacifastacus leniusculus klamathensis
(Stimpson), from Mary's River, 7.7 km E
of Blodgett, Benton Co., Oregon, by Perry
C. and Virgie F. Holt, 12 Jul 1960.

Distribution. —Western Oregon and
Washington (Hoh 1977:125).

Sathodrilus megadenus Holt, 1968
Figs. 307-310

Sathodrilus megadenus Holt, 1968b:302-
305.

Types. -YioXoXy^Q, USNM 37109, 2
paratypes, USNM 37 1 1 0, 2 paratypes, PCH
1346, on Cambarus latimanus (Le Conte),
from a small stream, 3.1 mi N of Buchanan,
Haralson Co., Georgia, by Perry C. and Vir-
gie F. Holt, 25 Mar 1961.

Distribution. —Known only from the type
locality.

Sathodrilus nigrofluvius Holt, 1989
Figs. 311-314

Sathodrilus nigrofluvius Holt, 1989b:738-
741.

rvp^5. -Holotype, USNM 118199, 3
paratypes, USNM 118200-118202, on un-
known host, from a tributary of the Black
River, 2 mi NE of Lesterville, Reynolds Co.,
Missouri, on State Road 2 1 , by Perry C. and
Virgie F. Holt, 22 Aug 1961.

Distribution. —Known only from the type
locality.

VOLUME 106, NUMBER 2

2S1

P. simondsi

276

278

279

S. attenuatus

282

S. carolinensis

285

S. chehalisae

286

288

280

283

284

289 V>^
290

Figs. 275-290. 275-276, Pterodrilus simondsi Holt; 277-280, Sathodrilus attenualus Holt; 281-284, 5<2(A-
odrilus carolinensis Holt; 285-286, Sathodrilus chehalisae Holt; 287-290. Sathodrilus dorfus Holt.

282

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Sathodrilus norbyi Holt, 1977
Figs. 315-318

Sathodrilus norbyi Holt, 1977:125-128.

Types. — Holotype and 3 paratypes,
USNM 53642, 15 paratypes, PCH 920, on
Pacifastacus leniusculus klamathensis
(Stimpson), from Union Rat Creek, ca. 1 3
km W of Pullman, Whitman Co., Wash-
ington, by Darwin E. Norby, 1 1 Jul 1958.

Distribution. —Idaho and Washington
(Holt, 1977:127).

Sathodrilus okaloosae Holt, 1973
Figs. 319-325

Sathodrilus okaloosae Holt, 1973a:99-102.

ry;7^5. -Holotype, USNM 49715, 5
paratypes, USNM 49716, 2 paratypes, PCH
2720, on Procambarus evermanni (Faxon)
and P. versutus (Hagen), collected 1.0 mi E
of Santa Rosa Co. line on U.S. Highway 90,
Okaloosa Co., Florida, by Horton H. Hobbs,
III, 12 Aug 1968.

Distribution. —Known only from the type
locality.

Sathodrilus prostates Holt, 1973
Figs. 326-331

Sathodrilus prostates Holt, 1973b:33-36.

Types.— HolotypQ, USNM 4532 [sic]
(should be 45431), on Procambarus acutus
cuevachicae (Hobbs), from El Ajenjibre,
Mesa de San Diego, km 262 de la carretera
Mexico [Tuspan, Puebla, Mexico], by A.
Villalobos and H. H. Hobbs, Jr., 12 Apr
1957; 2 paratypes, USNM 45432, on the
freshwater crab Pseudothelphusa veracru-
zana, from Rio Tapalapa, Veracruz, Mexi-
co, by A. Villalobos and H. H. Hobbs, Jr.;
1 paratype in the IBUM from the latter lo-
cality; 7 paratypes, PCH 700, from the latter
locality.

Distribution. —From the lower slopes of
the Sierra Oriental in Puebla to the lowlands
of Veracruz, Mexico.

Sathodrilus rivigeae Holt, 1988
Figs. 332-335

Sathodrilus rivigeae Holt, 1988b:804-807.

ry/7^5. -Holotype, USNM 119545, 2
paratypes, USNM 119546 (PCH 1089), on
Orconectes palmeri longimanus (Faxon),
from cool pools in a medium sized stream
in Ouachita National Forest 3.2 mi E of
Joplin, Montgomery Co., Arkansas, at
crossing of U.S. Highway 270, by Perry C.
and Virgie F. Holt, 23 Jun 1960.

Distribution. —Known only from the type
locality.

Sathodrilus shastae Holt, 1981b
Figs. 336-338

Sathodrilus shastae Holt, 1981b:856-859.

Ty/j^s. -Holotype, USNM 65230, 4
paratypes, PCH 1 8 1 8, on Pacifastacus fortis
(Faxon), from the headwaters of Fall River,
Thousand Springs Ranch, Shasta Co., Cal-
ifornia, by Perry C. and Virgie F. Holt, 1 9
Aug 1964.

Distribution. —Known only from the type
locality.

Sathodrilus veracruzicus Holt, 1968
Figs. 339-344

Sathodrilus veracruzicus Holt, 1968b:305-
308.

rj;;?^^. -Holotype, USNM 37105, 3
paratypes, USNM 37 106, 3 paratypes, PCH
1623, on Procambarus hoffmanni (Villalo-
bos), from waters near Coyutla, Veracruz,
by Alejandro Villalobos, 16 Apr 1949.

Distribution. —Known only from the type
locality.

Notes: In the description of this species
Hoh (1968b:299, 307) correctly states that
oral papillae are absent. In the description
ofS. veracruzicus it is incorrectly stated that
oral papillae are present in both species.
They are present only in S. villalobosi.

VOLUME 106, NUMBER 2

283

S. hortoni

eps

301

302

305 iVt

306

307

S. megadenus

eps

309

310®

Figs. 291-310. 291-294, Sathodrilus elevatus (Goodnighl), 292, left lateral view of reproductive system, 293,
spermatheca, 294, right lateral view of reproductive system; 295-298, Sathodrilus hortoni Holt; 299-302,
Sathodrilus inversus (Ellis); 303-306, Sathodrilus lobatus Holt; 307-310, Sathodrilus megadenus Holt.

284

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

S. nigrofluvius

312

S. norbyi

316

S. okaloosae 320

313 /,-

314

318^^

324
325

S. prostates 327

328

329

330

3 ^

331

S. rivigeae 333

334

335

C=^

Figs. 311-335. 311-314, Sathodrilus nigrofluvius Holt; 315-318, Sathodrilus norbyi Holt; 319-325, Sath-
odrilus okaloosae Holt, 321, lateral view of everted penis; 326-331, Sathodrilus prostates Holt; 332-335, Sath-
odrilus rivigeae Holt.

VOLUME 106, NUMBER 2

285

Sathodrilus villalobosi Holt, 1968
Figs. 345-348

Sathodrilus villalobosi Holt, 1 968b:299-302,
1973b:36-38.

ryp^5.- Holotype, USNM 37101, 4
paratypes, USNM 37 102, 4 paratypes, PCH
208, on Procambarus paradoxus (Ort-
mann), from Tetela de Ocampo, Puebla,
Mexico, by Alejandro Villalobos, May 1941.

Distribution. —The type locality and Aqua
Fria, Puebla, Mexico.

Sathodrilus wardinus Holt, 1981
Figs. 349-352

Sathodrilus wardinus Holt, 1 9 8 1 b: 8 5 9-8 6 1 .

Types.— HolotypQ and 1 paratype, USNM
65229, 5 paratypes, PCH 921, on Pacifas-
tacus leniusculus klamathensis (Stimpson),
from Purdy Creek, 6 mi N of Gig Harbor,
Pierce Co., Washington, by Darwin E. Nor-
by, 26 Jun 1939.

Distribution. —FiercQ Co., Washington.

Tettodrilus Holl, 1968

Tettodrilus Holt, 1968b:312.

Type species. — Tettodrilus friaufi Holt,
1968, by original designation.
Gender. —Masculine.

Tettodrilus friaufi YioXx, 1968
Figs. 353-357

Tettodrilus friaufi Holt, 1968b:314-317.

r>;pe'5. -Holotype, USNM 37099, 1
paratype, USNM 37100, on Orconectes mi-
rus (Ortmann), O. rhoadesi Hobbs, Cam-
barus graysoni Faxon, and C tenebrosus
Hay, from a small stream ca. 8.5 mi S of
Lewisburg, Marshall Co., Tennessee, on U.S.
Highway 431, by Perry C. and Virgie F.
Holt, 18 Apr 1960; 1 paratype, PCH 1007,
on C. graysoni and C. tenebrosus, from a
stream tributary to the Harpeth River, 2.4
mi S of Franklin, Williamson Co., Tennes-

see, on U.S. Highway 432, by Perry C. and
Virgie F. Holt, 18 Apr 1960; 1 paratype,
PCH 1008, on C. graysoni, from a small
stream, 5.3 mi S of Franklin, Williamson
Co., Tennessee, by Perry C. and Virgie F.
Holt, 18 Apr 1960.

Distribution. —StvesiTns of the Nashville
Basin in middle Tennessee.

Triannulata Goodnight, 1940

Triannulata Goodnight, 1940:56.

Type species. —Triannulata magna
Goodnight, 1940, by original designation.
Gender. —Feminine.

Triannulata magna Goodnight, 1940
Figs. 358-361

Triannulata magna Goodnight, 1940:56-
57. -Holt, 1974:63-66.

Types. -Holotype, USNM 20567, on Pa-
cifastacus sp., from Naches, Washington
[collector and date unknown].

Distribution. —Yakima and Cowlitz
Counties, Washington (Holt 1974:66).

Key to the Genera of Cambarincolidae
(modified from Pennak, 1978)

1. Penis a protrusible muscular

cone (Fig. IB-E) 2

- Penis eversible (Fig. IF-I) . . 4
2(1). Spermatheca absent; bursa

asymmetrical and rounded
(Figs. 225, 229) Ellisodrilus

- Spermatheca present (Fig. 3) 3
3(2). Length 2.5 mm or greater; no

dorsal appendages present (Fig.

2) Cambarincola

- Length less than 2.0 mm; cy-
lindrical or fan-shaped dorsal
appendages usually present on
dorsal ridges (Figs. 253, 255)
Pterodrilus

4(1). Penis a cuticular tube, often

very long (Fig. 1 F-I)

Oedipodrilus

286

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

S. veracruzicus

349

S. wardinus

T. friaufi

340

345 346

S. villalobosi

350

354

355

341

338

343

/::>

342 344

347

348

351

(
352

£>

356

357

T!>>

Figs. 336-357. 336-338, Sathodrilus shastae Holt; 339-344, Sathodrilus veracruzicus Holt; 345-348, Sath-
odrilus villalobosi Holt; 349-352, Sathodrilus wardinus Holt; 353-357, Tettodrilus friaufi Holt, 355, lateral view
of bursa and penis.

VOLUME 106, NUMBER 2

287

spg

359

361

360

Figs. 358-361. Triannulata magna Goodnight, 361, ventral view of upper jaw.

- Penis with internal strands
(Fig. 321) or an epithelial or
muscular wall (Fig. 355) .... 5

5(4). Body segments II to VIII with
cylindrical projections on dor-
sal surface (Fig. 2 1 8), upper lip
with four tentacles . Ceratodrilus

- Without projections on body

or tentacles on lip 6

6(5). Penis and ejaculatory duct a
continuous, muscular eversi-
ble tube (Fig. 359) . . Triannulata

- Penis and ejaculatory duct dis-
tinct regions, ejaculatory duct
not eversible (Figs. 350, 354) 7

7(6). Bursal atrium much longer
than penial sheath; penial
sheath eversible (Fig. 236) . .
Magmatodrilus

- Bursal atrium shorter than
penial sheath; penial sheath not
eversible (Fig. 278) 8

8(7). Seminiducal gland small and
slender, length and diameter
about equal to that of ejacula-
tory duct (Fig. 354) ... Tettodrilus

- Seminiducal gland larger than
ejaculatory duct (Fig. 278) . .

Sathodrilus

Key to the Species of Cambarincola

1. Cephalic area conspicuously

enlarged (Figs. 1 12, 178) ... . 2

- Cephalic area of normal size
(Fig. 2) 3

2(1). Prostomial tentacles present
(Fig. 1 1 2); spermathecal bulb

cylindrical (Fig. 113)

C. macrocephala

- Prostomial tentacles absent
(Fig. 178); spermathecal bulb
oval (Fig. 179) C. restans

3(1). Prostomial tentacles present

(Figs. 49, 88) 4

- Prostomial tentacles absent
(Fig. 2) 8

4(3). Spermathecal bulb with cylin-
drical ental process; semini-
ducal gland with two lobes (Fig.
89) C holti

- Spermathecal bulb without en-
tal process; seminiducal gland
without lobes (Fig. 50) 5

5(4). Body without prominent dor-
sal ridges (Fig. 94); prostate
longer than seminiducal gland
(Figs. 95, 151) 6

- Body with prominent dorsal
ridges (Fig. 49); prostate short-
er than seminiducal gland
(Figs. 50, 173) 7

6(5). Prostomial tentacles small (Fig.
94); diameter of prostate less
than that of seminiducal gland
(Fig. 95) C ingens

- Prostomial tentacles large (Fig.
150); diameter of prostate
greater than that of semini-
ducal gland (Fig. 151)

C montanus

288

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

7(5). Prostomial tentacles large (Fig.
49); diameter of prostate three
times that of seminiducal gland
(Fig. 50) C fallax

- Prostomial tentacles small (Fig.
172); diameter of prostate two
times that of seminiducal gland
(Fig. 173) C philadelphicus

8(3). Length of prostate greater than
that of seminiducal gland (Figs.
78, 145) 9

- Length of prostate equal to or
less than that of seminiducal
gland (Figs. 7, 99) 11

9(8). Length of spermathecal bulb
three times that of duct (Fig.
169); lower jaw with four teeth
(Fig. 171) C. pamelae

- Length of spermathecal bulb
about 1.5 times that of duct
(Fig. 78); lower jaw with two
teeth (Fig. 80) 10

10(9). Spermathecal bulb oval, di-
ameter two times that of duct
(Fig. 78) C. hoffmani

- Spermathecal bulb lobed, di-
ameter 1.5 times that of duct
(Fig. 145) C micradenus

1 1 (8). Length of prostate no more than
one-third that of seminiducal
gland (Figs. 72, 191) 12

- Length of prostate at least half
that of seminiducal gland (Figs.

12, 16) 13

12(11). Seminiducal gland not bi-
lobed; spermathecal duct lon-
ger than bulb (Fig. 72)

C heterognathus

- Seminiducal gland bilobed;
spermathecal duct shorter than
bulb (Fig. 191) C. Shoshone

13(11). Length of prostate about half
that of seminiducal gland (Figs.
11, 16) 14

- Length of prostate at least two-
thirds that of seminiducal
gland (Figs. 7, 99) 25

14(13). Seminiducal gland abruptly
bent and U-shaped (Figs. 12,
14) 15

- Seminiducal gland not
U-shaped (Figs. 16, 20) 19

15(14). Diameter of spermathecal bulb

and duct similar (Figs. 44, 62) 16

- Diameter of spermathecal bulb
greater than that of duct (Fig.

12) 17

16(15). Spermatheca oval; length of
prostate about half body di-
ameter (Fig. 44) C ellisi

- Spermatheca spherical; length
of prostate one-fourth that of
body diameter (Fig. 62)

C goodnighti

17(15). Length of spermathecal duct
three times that of bulb (Fig.
12) C. barbarae

- Length of spermathecal duct
about 1.3 times that of bulb
(Fig. 205) 18

18(17). Spermathecal bulb nearly
spherical; length of prostate 1.5
times that of bursa (Fig. 205)
C susanae

- Spermathecal bulb elongate;
length of prostate twice that of
bursa (Fig. 217) C vitreus

19(14). Diameter of seminiducal gland
1.5 times that of prostate (Fig.
193) C smalleyi

- Diameter of seminiducal gland
at least twice that of prostate
(Figs. 16,20) 20

20(19). Seminiducal gland with two

lobes (Figs. 20, 213) 21

- Seminiducal gland not bilobed
(Fig. 16) 22

21(20). Length of spermatheca about
one-half body diameter (Fig.
20) C branchiophilus

- Length of spermatheca nearly
as great as body diameter (Fig.
213) C. virginicus

VOLUME 106, NUMBER 2

289

22(20). Length of bursa greater than
that of seminiducal gland (Fig.
36) C. demissus

- Length of bursa less than that
of seminiducal gland (Figs. 1 6,
24) 23

23(22). Spermathecal bulb spherical

(Fig. 16) C bobbi

- Spermathecal bulb elongate
(Figs. 24, 68) 24

24(23). Diameter of ejaculatory duct
one-fifth that of bursa: diam-
eter of prostate one-fourth that
of seminiducal gland (Fig. 24)
C. carcinophilus

- Diameter of ejaculatory duct
one-half that of bursa; diam-
eter of prostate one-half that
of seminiducal gland (Fig. 68)

C. gracilis

25(13). Seminiducal gland bilobed

(Figs. 3, 167) 26

- Seminiducal gland not bilobed
(Figs. 7, 30) 27

26(25). Prostate and bursa of similar

length (Fig. 3) . . . C acudentatus

- Length of prostate three times
that of bursa (Fig. 167) .....

C Ouachita

27(25). Diameter of seminiducal gland
equal to that of prostate (Fig.
105) 28

- Diameter of seminiducal gland
at least 1.5 times that of pros-
tate (Figs. 7, 30, 84) 32

28(27). Spermathecal bulb smaller

than bursa (Fig. 124) . . C. manni

- Spermathecal bulb equal to or
larger than bursa (Figs. 105,
108) 29

29(28). Spermathecal duct about twice
as long as spermathecal bulb
(Fig. 197) C. speocirolanae

- Spermathecal bulb longer than
spermathecal duct (Fig. 105) 30

30(29). Diameter of spermathecal bulb

four times that of duct (Fig.
105) C. leoni

- Diameter of spermathecal bulb

1 . 5 times that of duct (Fig. 108) 31

31(30). Length of spermatheca half

that of body diameter; length

of prostate two-thirds that of

seminiducal gland (Fig. 108)

C leptadenus

- Length of spermatheca nearly
equal to body diameter; length
of prostate equal to that of
seminiducal gland (Fig. 136)

C. mesochoreus

32(27). Diameter of spermathecal bulb
at least twice that of duct (Figs.
38, 56) 33

- Diameter of spermathecal bulb
no more than 1.5 times that of
duct (Figs. 7, 84) 39

33(32). Length of prostate equal to that
of seminiducal gland (Figs. 56,
91) 34

- Length of prostate two-thirds
that of seminiducal gland (Figs.

38, 130) 36

34(33). Lengths of prostate and bursa

equal (Fig. 91) ... C illinoisensis

- Prostate twice as long as bursa
(Figs. 56, 161) 35

35(34). Prostate straight; diameter of
spermathecal bulb four times
that of duct (Fig. 56); lower jaw
with four teeth (Fig. 58) ....
C. floridanus

- Prostate J-shaped; diameter of
spermathecal bulb 1.5 times
that of duct (Fig. 161); lower
jaw with two teeth (Fig. 163)

C olmecus

36(33). Spermathecal bulb with a small

ental lobe (Fig. 139) . . . C meyeri

- Spermathecal bulb without an
ental lobe (Fig. 38) 37

37(36). Seminiducal gland L-shaped

(Fig. 38) C. dubius

290

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

- Semini ducal gland oval or cy-
lindrical (Figs. 130. 210) ... . 38

38(37). Spermathecal bulb oval, di-
ameter 3 times that of duct;
prostate longer than bursa (Fig.
130) C marthae

- Spermathecal bulb spherical,
diameter four times that of
duct; lengths of prostate and
bursa similar (Fig. 201)

C. steevesi

39(32). Prominent dorsal ridges pres-
ent (Figs. 29, 180) 40

- Dorsal ridges absent (Figs. 6,
83) 41

40(39). Bursa longer than prostate;
spermathecal duct and bulb
equal in length (Fig. 30) ....
C chirocephala

- Bursa shorter than prostate;
spermathecal duct half as long

as bulb (Fig. 181) . . . . C serratus
41(39). Length of seminiducal gland
less than that of bursa; sper-
mathecal bulb with a large en-
tal lobe (Fig. 7) C. alienus

- Length of seminiducal gland
equal to or greater than bursa;
spermathecal bulb without
large ental lobe (Figs. 84, 99) 42

42(39). Seminiducal gland L- or

U-shaped (Figs. 165, 209) . . 43

- Seminiducal gland straight or
slightly curved (Figs. 84, 99) 45

43(42). Bursa cylindrical, length twice

its diameter (Fig. 118)

C macrodontus

- Bursa oval, length no greater
than 1.5 times its diameter
(Figs. 165, 209) 44

44(43). Seminiducal gland U-shaped;
diameter of spermathecal duct
less than that of prostate (Fig.
165) C osceolai

- Seminiducal gland L-shaped;
diameter of spermathecal duct
similar to that of prostate (Fig.
209) C toltecus

45(42). Upper and lower jaws with
three teeth (Figs. 85, 86); sper-
mathecal bulb cylindrical (Fig.
84) C holostoma

- Upper jaw with five teeth (Figs.
100, 156), lower with four or
five teeth (Fig. 157); sperma-
thecal bulb oval (Figs. 99, 155) 46

46(45). Lower jaw with one large and
four small teeth (Fig. 187);
length of prostate 1.5 times that
of bursa (Fig. 185); body length
2.5 mm C. sheltensis

- Lower jaw with four teeth of
equal size (Figs. 101, 157);
length of prostate and bursa
similar (Fig. 99); body length
less than 2.0 mm 47

47(46). Length of prostate 0.8 times
that of seminiducal gland;
length of ejaculatory duct twice
its diameter (Fig. 99); central
tooth of upper jaw larger than

lateral teeth (Fig. 100)

C. jamapaensis

- Length of prostate 0.6 times
that of seminiducal gland;
length and diameter of pros-
tate equal (Fig. 155); teeth of
upper jaw similar in size (Fig.
156) C. nanognathus

Key to the Species of Ceratodrilus

1 . Length of tentacles on head 200
)um (Fig. 218); projections on
segments II-VII 146 ^iva long
C. ophiorhysis

- Length of tentacles on head 90
IJLVCi (Fig. 222); projections on
segments II-VII 50 iiva long .

C thysanosomus

Key to the Species of EUisodrilus

\. Bursa large and bent, length
about two-thirds body diam-
eter (Fig. 229) E. clitellatus

- Bursa not bent, length about

VOLUME 106, NUMBER 2

291

half body diameter (Figs. 225,

233) 2

2(1). Bursa wider than long; pros-
tate diameter equal to that of
seminiducal gland (Fig. 225)
E. carronamus

- Width and length of bursa
about equal; diameter of pros-
tate half that of seminiducal
gland (Fig. 233) E. durbini

Key to the Species of Oedipodrilus

1. Prominent dorsal ridges pres-
ent O. oedipus

- No dorsal ridges present .... 2
2(1). Spermathecal bulb and duct

slender, diameters equal; sper-
mathecal bulb bent at right an-
gle to duct (Fig. 244)

O. cuetzalanae

- Spermathecal bulb and duct
robust (Figs. 240, 248) 3

3(2). Length of spermathecal duct
half that of bulb; bursa straight
(Fig. 240) O. anisognathus

- Length of spermathecal duct
1.5 times that of bulb; bursa
long and bent (Fig. 248) ....

O. macbaini

Key to the Species of Pterodrilus

1 . Four of five pairs of dorsal, fan-
like projections on body (Fig.
253) 2

- Body with one to seven pairs
of dorsal, finger-like projec-
tions (Figs. 255, 263) 3

2(1). Diameter of spermathecal bulb
1.5 times that of duct; diam-
eter of seminiducal gland equal
to that of bursa (Fig. 254) ...
P. alcicornus

- Diameter of spermathecal bulb
three times that of duct; di-
ameter of seminiducal gland
half that of bursa (Fig. 276) .

P. simondsi

3(1). Length of prostate equal to that
of seminiducal gland (Figs.
256, 260) 4

- Length of prostate 0.6 to 0.5
times that of seminiducal gland
(Figs. 264, 274) 7

4(3). Diameter of bursa twice that
of seminiducal gland; length of
bursa half that of body diam-
eter (Fig. 272) .... P. mexicanus

- Diameter of bursa one to 1.5
times that of seminiducal
gland; length of bursa one-third
or less that of body diameter
(Figs. 256, 260) 5

5(4). Length of spermathecal duct
twice that of bulb (Fig. 256) .
P. cedrus

- Length of spermathecal duct
less than that of bulb (Fig. 260)
6

6(5). Diameter of spermathecal bulb
and duct equal; length of sem-
iniducal gland and bursa equal
(Fig. 260) P. choritonamus

- Diameter of spermathecal bulb
three times that of duct; length
of seminiducal gland twice that

of bursa (Fig. 268) .... P. hobbsi
7(3). Finger-like projections on the
dorsal surface of seven body
segments (Fig. 263); length of
bursa about one-fourth that of
body diameter (Fig. 264) ....
P. disdchus

- Finger-like projections on the
dorsal surface of only one body
segment (Fig. 273); length of
bursa about half that of body
diameter (Fig. 274)

P. missouriensis

Key to the Species of Sathodrilus

1. Tentacles on dorsal lips (Fig.

303) S. lobatus

- Dorsal lips without tentacles
(Fig. 277) 2

2(1). Ejaculatory duct spherical, di-

292

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ameter about half that of bursa
(Fig. 292) S. elevatus

- Ejaculatory duct cylindrical,
diameter less than half that of
bursa (Fig. 278) 3

3(2). Spermathecal bulb spherical or

oval (Figs. 346, 350) 4

- Spermathecal bulb cylindrical
(Figs. 278, 288) 5

4(3). Length of spermathecal duct
four times the diameter of the
bulb; prostate absent (Fig. 346)
S. villalobosi

- Length of spermathecal duct
1.3 times the diameter of the
bulb; prostate present (Fig.
350) S. wardinus

5(3). Prostate present (Fig. 278) . . 6
Prostate absent (Fig. 282) ... 12

6(5). Length of spermathecal duct
three times that of bulb (Fig.
288) 5. dorfus

- Length of spermathecal bulb
equal to or greater than that of
duct (Figs. 278, 308) 7

7(6). Ejaculatory duct long, diame-
ter equal to that of bursa (Fig.
308) S. megadenus

- Ejaculatory duct short, diam-
eter less than half that of bursa
(Fig. 278) 8

8(7). Length of prostate about half
that of seminiducal gland (Fig.
278) 9

- Lengths of prostate and sem-
iniducal gland equal (Fig. 286)
10

9(8). Bursa spherical, length about
one-third body diameter (Fig.
278) S. attenuatus

- Bursa cylindrical, length about
half body diameter (Fig. 300)
S. inversus

10(8). Length and width of bursa

equal (Fig. 286) . . . . S. chehalisae

- Length of bursa two to three
times its width (Figs. 327, 333)
11

11(10). Length of spermathecal bulb
four times that of duct (Fig.
327) S. prostates

- Lengths of spermathecal bulb
and duct equal (Fig. 333) ...

S. rivigeae

1 2(5). Seminiducal gland slender and
looped, length greater than
body diameter (Fig. 337) ...

S. shastae

- Seminiducal gland short,
length less than half body di-
ameter (Figs. 282, 316) 13

13(12). Diameter of seminiducal gland
equal to that of bursa (Fig. 316)
S. norbyi

- Diameter of seminiducal gland
no more than 0.6 that of bursa
(Figs. 312, 320) 14

14(13). Length of seminiducal gland

greater than that of bursa;

length of spermatheca less than

half body diameter (Fig. 296)

S. hortoni

- Length of spermathecal gland
less than that of bursa; length
of spermatheca nearly equal to
or greater than body diameter
(Figs. 282, 312) 15

15(14). Spermatheca bent, length 1.5
times body diameter (Fig. 340)
S. veracruzicus

- Spermatheca not bent, length
equal to body diameter (Fig.
282) 16

16(15). Spermathecal bulb with a slen-
der ental lobe whose length is
equal to that of the sperma-
thecal duct (Fig. 320)

S. okaloosae

- Spermathecal bulb without a
long ental lobe (Figs. 282, 3 1 2)

17(16). Diameter of seminiducal gland
about one-third that of bursa;
spermathecal duct cylindrical;
spermathecal bulb without en-
tal lobe (Fig. 282) S. carolinensis

VOLUME 106, NUMBER 2

293

- Diameter of seminiducal gland

half that of bursa; spermathe-
cal duct constricted; sperma-
thecal bulb with short ental
lobe (Fig. 312) .... 5. nigrofluvius

Acknowledgments

We thank Dr. Richard L. Hoffman and
Dr. Mark Wetzel for detailed comments and
Dr. Ralph O. Brinkhurst for general sug-
gestions that helped improve this paper. Mr.
George C. Steyskal reviewed the paper's no-
menclatural citations. Support from Virgin-
ia Polytechnic Institute and State Univer-
sity made possible the publication of this
manuscript. Perry Holt thanks the Virginia
Academy of Science, the National Science
Foundation (grants G-4439, G-9828, and
GB-372), and Virginia Polytechnic Institute
and State University's Biology Department
and Agricultural Research Division for sup-
porting his collecting trips and research. He
thanks Dr. Horton H. Hobbs, Jr., for the
encouragement and advice provided
throughout his career. Virgie F. Holt and
Dr. Susan E. H. West provided invaluable
help on numerous collecting trips. Drs. Jo-
seph F. Fitzpatrick, John Holsinger, Denton
W. Crocker, and Harrison R. Steeves, III,
and others cited in the test also helped with
collecting or provided specimens.

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Department of Biology, Virginia Poly-
technic Institute and State University,
Blacksburg, Virginia 24061, U.S.A.

PROC. BIOL. SOC. WASH.
106(2), 1993, pp. 296-304

BRANCHINECTA SANDIEGONENSIS, A NEW

SPECIES OF FAIRY SHRIMP

(CRUSTACEA: ANOSTRACA) FROM

WESTERN NORTH AMERICA

Michael Fugate

Abstract.— Branchinecta sandiegonensis, a. new species of fairy shrimp is
described from vernal pools on Del Mar Mesa, San Diego County, California.
The species is found within 50 km of the Pacific Ocean from Santa Barbara,
California to Valle de las Palmas, Baja California Norte, Mexico. It can be
distinguished from the other six branchinectids inhabiting southern California
by the combination of thoracic spine pattern, ovary length, ovisac length and
shape, and egg morphology of females and the form of the second antenna of
males.

Vernal pools, so named for their colorful,
springtime floral displays as the pools dry
after winter rains, have long been known for
their endemic floras (Crampton 1976, Hol-
land & Jain 1977). Only one fairy shrimp
however was known to be endemic to the
extensive vernal pool habitat ranging from
southern Oregon through the Central Valley
of California and into northern Baja Cali-
fornia, Mexico (Dodds 1923, Brtek 1964)
before a recent monograph on the Anostra-
ca of California described four endemic,
vernal pool species (Eng et al. 1990). Here
I describe another species found primarily
in vernal pools atop mesas in San Diego
County, California.

Methods

Animals were obtained either from filled
pools or by hydrating soil samples from dry
pools. Freshly molted individuals were fixed
for 3 hours in 3% glutaraldehyde in 0.1 M
sodium cacodylate at pH 7.4, post-fixed with
2% osmium tetroxide in sodium cacodylate
for 2 hours, dehydrated in a graded series
of 1 0% glacial acetic acid in absolute etha-
nol, and transferred to absolute ethanol. The
eggs were air-dried from absolute ethanol,

the mandibles were air-dried after 10 min-
utes in tetramethylsilane (Dey et al. 1989),
and all other parts were critically point-
dried, coated with gold-palladium and ob-
served on a Philips 515 scanning electron
microscope.

Branchinecta sandiegonensis,

new species

Figs. 1-13

Type material. —One 6, holotype, USNM
256557, 1 9, allotype, USNM 256558, 2 6,
3 9, paratypes, USNM 256556, 3 <5, 5 9,
paratypes, Los Angeles County Museum of
Natural History (LACM), LACM 90-356. 1 ,
2 (5, 3 9, paratypes, Hungarian Museum of
Natural History, Del Mar Mesa, San Diego
County, California, USA (32°51'N,
1 17°15'W), 17 Mar 1990 (coll. M. Simovich
& M. Fugate); 4 6, 8 9, paratypes, LACM
89-357. 1 , 4 3, 5 9, paratypes Museo Ciencias
Naturales de La Plata, Kearney Mesa (Mir-
amar Naval Air Station), San Diego County,
California (32°50'N, 1 17°09'W), 8 Feb 1990
(coll. M. Simovich); 50 S, 50 9, paratypes,
USNM 294523, Ramona, San Diego Coun-
ty, California (33°02'N, 116°52'W), 4 Mar
1962 (coll. J. E. Lynch); 50 ^, 50 9, para-

VOLUME 106, NUMBER 2

297

types, USNM 305974, Poway, San Diego
County, California (32°55'N, 117°04'W), 3
Mar 1962 (coll. J. E. Lynch).

Type locality.— An extensive network of
vernal pools surrounded by chaparral on
Del Mar Mesa, San Diego County, Califor-
nia, USA, 32°51'N, 117°15'W, elev. 100
meters, south and east of junction of Inter-
state 5 and Carmel Valley Road (Green-
wood 1984).

Etymology. —Named for San Diego
County, California, USA.

Af<3/e. —Antenna 1 slender, cylindrical,
approximately 10 times as long as wide, with
3 long setae and 8 or more shorter aesthe-
tascs (type 1 and type 2 sensilla, respec-
tively—Tyson & Sullivan 1979; Fig. 1). An-
tenna 2 biarticulate, cylindrical, reaching to
thoracic segment 8 (Fig. 6). Basal and distal
segments of approximately equal length.
Basal segment with medial, oval pulvillus,
near proximal end, of short spines inter-
spersed with slightly longer, stouter, conical
spines (Fig. 10a), medial, elevated cluster of
6-10 short, stout spines, half distance from
end of basal segment (Fig. 6a), and single
row containing clusters of papillae, each with
sensory seta, on distal 0.67 of anterolateral
surface. Distal segment slightly arcuate, up-
per 0.25 cylindrical, remainder mediolater-
ally flattened (Fig. 2a-e). Breadth 0.2 length
at joint with basal segment expanding to
0.33 at tip. Mediolateral surfaces 6 times as
broad as anteroposterior surfaces. Medial
surface flat and lateral concave. Anterior 0.5
of distal portion inflated, turned in medially
at 45°, ovoid anteriorly, triangular laterally.
Posterior edge of distal portion with patch
of raised, ovoid papillae arranged in rows
(Fig. 10b).

Body of mandible with spinelike protu-
berance on posterolateral surface. Molar
surfaces of mandibles asymmetrical, broad-
ly oval with dorsal edge concave and ventral
convex, divided into 2 basic regions, an an-
tero ventral and posterodorsal (Tyson & Sul-
livan 1981, Fig. 11). Anteroventral region
of both molar surfaces similar, with ap-

proximately 45 ridges and furrows, some
bifurcating, running dorsoventrally. Ridges
of rectangular cuticular projections bearing
many conical protuberances, both projec-
tions and protuberances becoming taller near
edges of molar surface. Ridges extend to
dorsal edge on anterior half of right man-
dible, but only on anterior 0.15 of left. Pos-
terodorsal region with 1 long, thin spine at
posterior edge and row of spines, decreasing
in size anteriorly, along dorsal edge. On right
mandible, spines sickle-shaped, each bor-
dering short row of widely spaced cuticular
projections, and each row in turn bordering
small area of relatively unadorned cuticle
dorsal to ridges of anteroventral region. On
left mandible, spines squat, conical, and
more widely spaced, becoming longer and
thinner anteriorly and bordering large area
of unadorned cuticle extending almost en-
tire length of molar surface.

Maxilla 1 with short, stout spine on ven-
tral edge and 1 8 setae, approximately 3 times
as long as ventral spine, on medial edge (Fig.
3). Proximal 0.33 of seta with 4-7 stout spi-
nules, arranged in single row, along medial
surface, distal 0.67 with 2 rows of setules.
Maxilla 2 small, with 6-10 setae on crown
(Fig. 4). Setae pliant, with 2 rows of setules
on distal 0.8. Medial surface of maxilla 2
with 2-3 short, pliant setae covered with
setules, not arranged in rows, and 1 small,
stout spine nearer base. Ventral surface of
labrum broadly triangular with lateral flap
on each side, posterior edge of flaps spinose.
Oral surface with small, distal lobe covered
with fine setae.

Thoracic segments appendage-bearing,
with paired dorsolateral, cuticular papillae,
each papillus with sensory seta (Figs. 6, 9).
Papillae not on raised protuberances, lateral
on segments 1-3, 5-8, and 10-11, dorsal on
4 and 9 (compare Lynch 1960, fig. 4 and
Cohen 1983, fig. 22). All 11 pairs of ap-
pendages similar in form, but those of seg-
ment 1 1 reduced (Fig. 8). There is much
confusion as to which lobes of anostracan
phyllopodous limbs are homologous with

298

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-5. Branchinecta sandiegonensis, new species male. 1. antenna, 1, distal tip. 2a-e. antenna 2, distal
segment, a. lateral, b. anterolateral, c. anterior, d. medial, e. dorsomedial. 3. maxilla 1, medial view. 4. maxilla
2, posterior view. 5a, b. right penis, a. ventral view. b. distal tip, lateral view, (scale bars 2, 4, 5, 6: 0.1 mm, 3:
1 mm)

VOLUME 106, NUMBER 2

299

Fig. 6. Branchinecta sandiegonensis, new species male. a. ventral view. b. lateral view, (scale bar 1.0 mm)

stenopodous limbs. Here the nomenclature
of Linder (1941) is employed. Endite 1 with
3 centers of origin for its numerous poste-
rior setae. Middle series with 7-9 posterior
setae and long, slightly toothed anterior seta.
Distal series with 11-16 posterior setae and
2 anterior setae on basal edge, distal seta
cornblike, with single row of stout setules,
basal seta shorter, spiniform. Appendage 1 1
with 3-5 posterior setae and short, spini-
form anterior seta in middle series and 4-
6 posterior setae in distal series (Fig. 8c).
Endite 2 similar to distal series of endite 1 ,
with 13-18 posterior setae and 2 anterior
setae. Appendage 1 1 with posterior setae
reduced to 2-4. Endites 3, 4, and 5 with 3,
2, and 2 posterior setae, respectively (Figs.
8 & 12a). Endites 3 and 4 with 2 anterior
setae on appendages 2-1 1 and 4-7 anterior
setae on appendage 1. Endite 5 with 2-6
anterior setae on appendages 2-1 1 and 4-8
on appendage 1 . Endopodite large, elongate,
shaped like broad scimitar (Fig. 8). Setae

along medial edge with several rows of fine
setules surrounding distal 0.67 (Fig. 12b),
becoming comblike toward ventral edge,
along ventral edge, long and thin, with 2
rows of fine setules. Exopodite oval to tri-
angular, surrounded by setae resembling
those of ventral edge of endopodite. Epi-
podite smooth, inflated, without setae.
Preepipodite thin, semicircular, with
coarsely toothed edge, reduced on append-
age 1 1 . Early in development with 2 lobes,
in adults with slight notch along lateral edge.

Genital segments only slightly expanded,
paired papillae dorsal, paired penes arising
ventrolaterally. Non-rectractile portion of
penis with ventral, fleshy lobe and medially-
directed, sclerotized spur (Fig. 5a). Distal
portion of penis eversible with two small,
sclerotized lobes on lateral surface each with
six to ten pyramidal teeth (Fig. 5b).

Post-genital segments with paired papil-
lae in following positions: one, dorsal, two,
ventral, three, dorsal, four, lateral, five and

300

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 7-8. Branchinecta sandiegonensis, new species, 7a-c. thoracic appendages female, a. appendage 1. b.
appendage 5. c. appendage H. 8a-c. thoracic appendages male. a. appendage 1. b. appendage 5. c. appendage
1 1 . (scale bar 0. 1 mm)

VOLUME 106, NUMBER 2

301

^^=

Fig. 9. Branchinecta sandiegonensis, new species female, a. dorsal view. b. lateral view, (scale bar 1.0 mm)

six, dorsal. Cercopods on anal segment
bearing setae with 2 rows of fine setules (Co-
hen 1983).

Length of mature individuals, from front
of head to end of anal segment, excluding
cercopods, 9.0-16.0 mm.

Female.— Yitdid. similar to that of male,
except for pair of cuticular papillae on dor-
sal surface, posterior to mandibular crease,
and form of antenna 2 (Fig. 9). Antenna 2
cylindrical, approximately 3 times as long
as wide basally, gradually tapering to 5 times
as long as wide at 0.8 its total length, then
rapidly to sharp point (Fig. 9b). Anterior
surface with 2 patches of sensory setae, each
seta borne on a cuticular papillus.

Thorax with cuticular papillae in same
locations as male, but segments 3 and 5-8
with 2 pairs of dorsolateral spines arranged
above and below papillus, dorsal normally
smaller than lateral spines, segment 4 with
1 pair of large bilobed spines (Fig. 9). Tho-
racic appendages similar to male, but en-
dopodite lobelike (Fig. 7a-c) with stout,
comblike setae along entire medial edge (Fig.
12c).

Genital segments slightly inflated, with
paired papillae lateral on segment 1 and
dorsal on 2 (Fig. 9a). Ovisac fusiform, on
average ending under post-genital segment
4, occasionally under segment 5. Paired
ovaries t-shaped, extending from thoracic
segment 9 or 10 to post-genital segment 4,
and into ovisac at junction of two genital
segments, forming an oviductal pouch.
Resting eggs spherical, diameter A^ = 269
Atm, Si) = 18 MHi, range = 227-309 jum, n
= 100, with numerous shallow hemispher-
ical depressions, approximately 40 ^m in
diameter, covering surface (Fig. 13a, b).

Post-genital segments similar to male, but
arrangement of papillae as follows: one, lat-
eral, two, ventral, three, dorsal, four, ven-
tral, five, dorsal, six, lateral.

Length of mature individuals, from front
of head to end of anal segment, excluding
cercopods, 8.0-15.0 mm.

Remarks.— T\iQ form of the male second
antenna has served as the cardinal character
for species identification in the genus and
although all males are currently distinguish-
able on that basis, the use of that character

302

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 10-13. Branchinecta sandiegonensis, new species, 1 Oa, b. antenna 2 male. a. pulvillus on basal segment,
b. tip of distal segment, dorsolateral view. 11a, b. mandibles male. a. left mandible, molar surface, b. right
mandible, transition between posterodorsal and anteroventral regions. 1 2a-c. setae on medial surfaces of thoracic
appendages, a. appendage 1 male, endites 3-5. b, c. medial edge of endopodite. a. male. b. female. 13a, b. resting
egg. a. whole egg. b. surface detail, (scale bars, la, b, 3a, 4a: 0.1 mm, 2a, b, 3b, c, 4b: 0.01 mm)

VOLUME 106, NUMBER 2

303

alone can often lead to difficulties. Bran-
chinecta sandiegonensis has been reported
as Branchinecta lindahli Packard, 1883, a
common species throughout western North
America, by Ebert & Balko (1 987) in a study
of the vernal pools on Kearney Mesa, San
Diego County, California (D. Belk, pers.
comm.; vouchers in personal collection of
D. Belk) and by Dr. J. E. Lynch in two ear-
lier collections from San Diego County
(USNM 305974, 3 Mar 1962, Poway,
32°55'N, 1 17°04'W; USNM 294523, 4 Mar
1962, Ramona, 33°02'N, 116°52'W). Five
additional species in the genus are also
known from southern California, but of
those only Branchinecta lynchi Eng et al.,
1990 is likely to be confused with B. san-
diegonensis. Branchinecta mackini Dexter,
1956 and Branchinecta gigas Lynch, 1937
are found in large playa lakes in the Mojave,
while Branchinecta conservatio Eng et al.,
1 990 and Branchinecta longiantenna Eng et
al., 1990, although found in vernal pools,
are readily distinguished morphologically.
In general form, B. sandiegonensis females
resemble those of B. lindahli due to the fu-
siform shape of the ovisac, however that of
B. lindahli is slightly longer, typically ending
under post-genital segment 5. Branchinecta
lindahli differs also in having a longer ovary,
extending from thoracic segment 4-7 to
post-genital segment 4, an egg with hemi-
spherical surface depressions, approximate-
ly 20 jLim in diameter, and a single dorso-
lateral spine, always below the papillus, on
each side of thoracic segments 3-11. Bran-
chinecta lynchi females share a similar ova-
ry length, egg surface (Mura 1991), and dor-
solateral spine pattern with B.
sandiegonensis, but differ in having a short,
conical ovisac, typically ending under post-
genital segment 3.

Males ofB. sandiegonensis share the large,
oval pulvillus on the basal segment of the
second antenna with B. lindahli, but the me-
dial series of spines is less diffuse. The distal
segment is broadest in B. sandiegonensis jusi
proximal to the tip, while that ofB. lindahli

is at 0.75 the distance from base. The breadth
at the tip in B. lindahli is only slightly larger
than at the joint with the basal segment and
the entire tip is bent medially at a right angle
to the segment of the antenna just preceding
it (Shantz 1905, Lynch 1964). The second
antenna of B. lynchi is quite different; the
pulvillus is smaller, there is a small apoph-
ysis, slightly distal and posterior to the pul-
villus, the distal segment has a narrower
breadth, and the entire tip is bent medially
(Engetal. 1990).

Distribution and habitat— Branchinecta
sandiegonensis is found after winter rains in
vernal pools from northern Baja California
Norte, Mexico (Valle de las Palmas, Baja
California Norte, Mexico 32°28'N,
116°37'W, 15 Nov 1987, soil sample, coll.
M. Fugate & G. Pratt) to Santa Barbara,
California (Isla Vista, Santa Barbara Coun-
ty, California 34°24'N, 119°51'W, 1, 7 Apr
1991, coll. J. Kommeyer). Its current range
is centered in San Diego County, California
(Del Mar Mesa, Kearney Mesa, Ramona,
Otay Mesa, 32°34'N, 1 16°58'W). All known
localities are within 50 km of the Pacific
Ocean and at elevations less than 700 m.
There are no records from Los Angeles and
Orange counties and it is unknown if the
Santa Barbara population is either disjunct
or the northern end of a formally continu-
ous distribution.

The pools in San Diego County are shal-
low (<30 cm) and often on chaparral cov-
ered mesas (Greenwood 1984, Ebert & Bal-
ko 1987). Zedler (1984) found 46 plant
species common in pools at Kearney Mesa,
one of which is endangered (Pogogyne
abramsii) and Ebert & Balko (1987) found
B. sandiegonensis associated with six cla-
docerans, one copepod, nine ostracods and
at least 21 rotifer species at the same site.
Streptocephalus woottoni Eng et al., 1990
and B. lindahli have subsequently been
found at Kearney Mesa, the former in a pool
containing B. sandiegonensis and the latter
in a road rut, but not in pools containing B.
sandiegonensis (M. Simovich, pers. comm.).

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Branchinecta sandiegonensis has been found
in a disturbed pool with B. lindahli at Del
Mar Mesa and in two pools with S. woottoni
and a clam shrimp, Cyzicus sp. at Otay Mesa.

Acknowledgments

Marie Simovich and her students sup-
plied me with many field-collected speci-
mens and soil samples. Clay Sassaman pro-
vided financial support, lab space, criticism
and encouragement. Denton Belk provided
useful comments on the text and figure pre-
sentations and examined his collections of
material from San Diego County, Califor-
nia. Marcia Hartz and Chris Williams as-
sisted with and the Chancellor's Patent Fund
at the University of California, Riverside
provided funds for the electron microscopy.
Three anonymous reviewers substantially
improved the manuscript.

Literature Cited

Brtek, J. 1964. Ein neue Gattung und Familie der
Ordnung Anostraca. — Annotationes Zoologicae
et Botanicae (Bratislava) 7:1-7.

Cohen, R. G. 1983. Notas sobre anostracos neotrop-
icales (Crustacea). III. Branchinecta somuncu-
remis y Branchinecta prima spp. nov. — Physis
(Buenos Aires) 41B:69-80.

Crampton, B. 1976. A historical perspective on the
botany of the vernal pools in California. Pp. 5-
10 in S. Jain, ed., Vernal pools: their ecology
and conservation. Institute of Ecology Publi-
cation No. 9, University of California, Davis,
93 pp.

Dey, S., T. S. Basu Baul, B. Roy, & D. Dey. 1989. A
new rapid method of air-drying for scanning
electron microscopy using tetramethylsilane.—
Journal of Microscopy 156:259-261.

Dodds, G. S. 1923. A new species of a phyllopod. —
Occasional Papers of the Museum of Zoology,
University of Michigan 141:1-3.

Ebert, T. A., & M. L. Balko. 1987. Temporary pools
as islands in space and time: the biota of vernal
pools in San Diego, southern California, USA. —
Archiv fur Hydrobiologie 1 10:101-123.

Eng, L. L., D. Belk, & C. H. Eriksen. 1990. Califor-

nian Anostraca: distribution, habitat, and sta-
tus.— Journal of Crustacean Biology 10:247-277.

Greenwood, N. 1984. The physical environment of
series H, vernal pools in San Diego County, Cal-
ifornia. Pp. 30-36 in S. Jain & P. Moyle, eds.,
Vernal pools and intermittent streams. Institute
of Ecology Publication No. 28, University of
California, Davis, 280 pp.

Holland, R. P., & S. K. Jain. 1977. Vernal pools. Pp.
515-533 in M. Barbour & J. Major, eds., Ter-
restrial vegetation of California. John Wiley &
Sons, New York, 1002 pp.

Linder, F. 1941. Contributions to the morphology
and the taxonomy of the Branchiopoda Anos-
traca. — Zoologiska Bidrag fran Uppsala 20: 101-
302 + 1 plate.

Lynch, J. E. 1960. The fairy shrimp Branchinecta
campestris from northwestern United States
(Crustacea: Phyllopoda).— Proceedings of the
United States National Museum 1 12:549-561.

. 1964. Parkard's and Pearse's species of 5ra/7-

chinecta: analysis of nomenclatural involve-
ment.— American Midland Naturalist 71:466-
488.

Mura, G. 1991. SEM morphology of resting eggs in
the species of the genus Branchinecta from North
America.— Journal of Crustacean Biology 11:
432-436.

Shantz, H. L. 1905. Notes on North American species
of Branchinecta and their habitats.— Biological
Bulletin 9:249-263 + plates X-XII.

Tyson, G. E., & M. L. Sullivan. 1979. Antennular
sensilla of the brine shrimp, Artemia salina. —
Biological Bulletin 156:382-392.

. 1981. A scanning electron microscopic study

of the molar surface of the mandibles of the
brine shrimp (CI. Branchiopoda: Or. Anostra-
ca).— Journal of Morphology 170:239-251.

Zedler, P. H. 1984. Micro-distribution of vernal pool
plants at Kearney Mesa, San Diego County. Pp.
185-197 in S. Jain & P. Moyle, eds., Vernal
pools and intermittent streams. Institute of
Ecology Publication No. 28, University of Cal-
ifornia, Davis, 280 pp.

Department of Biology, University of
California, Riverside, California 92521,
U.S.A.; (Current Address) Department of
Biology, University of Oregon, Eugene, Or-
egon 97403, U.S.A.

PROC. BIOL. SOC. WASH.

106(2), 1993, pp. 305-314

THE FIRST FINDING OF THE MALE OF

THAUMATOCYPRIS ECHINATAMiJlA.YK, 1906

(CRUSTACEA: OSTRACODA)

J. A. Rudjakov

Abstract.— Two adult males and two juveniles of Thaumatocypris echinata
Miiller, 1906 were found in the near-bottom layer off the west coast of Mad-
agascar at the depth of 180-1220 m. The first description of male morphology
is given. The sexual dimorphism manifests itself in the structure of mandibular
endopodite instead of endopodite of second antenna as it was known for other
halocypridids previously. The contents of the gut suggest that the species is
either a carnivore or scavenger.

The myodocopid ostracod genus Thau-
matocypris with only species T. echinata was
established by Miiller (1906) on six females
derived from a depth of 1 1 00 m near In-
donesia. Since then, only one female of the
type species was found nearly in the same
region at the depth of about 2000 m (Poul-
sen 1969).

Two other living genera of the family
Thaumatocyprididae, all benthic— T/z^w-
matoconcha (eight species) and Danielopo-
lina (six species) were thoroughly investi-
gated by Komicker & Iliffe (1989a, 1989b)
and by Komicker & Sohn (1 976). The latter
authors also used the opportunity to rein-
vestigate the specimen of Thaumatocypris
echinata described by Poulsen (1969).

Plankton investigation in the near-bot-
tom layer of the Indian Ocean off the west
coast of Madagascar (17th trip of R/V Vi-
tyaz) revealed a few specimens of the genus
Thaumatocypris. The Madagascar speci-
mens (two males and two juveniles) were
preliminarily studied by Dr. Louis S. Kor-
nicker (National Museum of Natural His-
tory, Smithsonian Institution, Washington,
D.C.), who returned them to me with the
kind suggestion to prepare the first male de-
scription. Two males have been deposited
to the U.S. National Museum of Natural
History and given USNM numbers.

Materials and Methods

Ostracods were collected in the 1 7th cruise
of R/V Vityaz by the towing underwater
apparatus "SOUND" equipped with open-
ing/closing plankton sampler (Biryukov et
al. 1990). Observations are briefly sum-
marized below.

Station (St.) 2649; 3 Dec 1988; 0206-0306
h; 22°25'S, 43°00"E; 970-950 m; 30 m above
bottom. 1 male USNM 194110(1.16 mm),
1 juvenile specimen (0.65 mm).

St. 2655; 3 Dec 1988; 1917-2017 h;
22°22'S, 42°54'E; 1212-1220 m; 1.5 m
above bottom. 1 juvenile specimen (0.72
mm).

St. 2661; 4 Dec 1988; 1200-1230 h;
22°13'S, 43°07'E, 260-180 m; 30 m above
bottom. 1 male USNM 1941 16 (1.20 mm).

The specimens were preserved with form-
aldehyde and after about two-years storage
transferred into alcohol. The males were
dissected in water diluted glycerol. Ap-
pendages were placed on slides moistened
with Faures liquid diluted 3-4 times with
water. After hardening of the fluid with the
appendages mounted in a necessary order
and position, a drop of molten glycerol-gel-
atin was deposited on a membrane formed
on the fluid, and a cover glass added (Rud-
jakov 1968). Separated valves were mount-

306

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ed on the shallow-well slides with glycerol-
gelatin.

Results

The following description of the adult
male was mostly prepared using the speci-
men from St. 2661, supplemented with the
features of the specimen from St. 2649 when
necessary.

Thaumatocypris echinata Miiller, 1906
Figs, la-d, 2a-c, 3a-d, 4a-f, 5a, b

Thaumatocypris echinata Muller, 1906:42,
pi. 6, figs. 1-18; 1912:54 [listed]. -Poul-
sen, 1969:7, fig. 1.— Komicker & Sohn,
1976:34, figs. 14c, d, 15.

Description. —Sh.Q\\ (Fig. la). Surface
smooth, without discernible anterior ridges
and serration along posterior ridge. Each
valve with short upper and long lower tube-
like anteroventral horns, appearing broken.
The shell can firmly stand on its four horns,
anterior side down, with furcal lamellae just
above ventral pair of horns and with anten-
nal and mandibular bristles protruding be-
tween dorsal and ventral pairs of horns.
Postero-dorsal ridge of right valve with short
rounded tubercle (hardly discernible on the
male from St. 266 1). Antero-ventral margin
of each valve with hairs distally bifurcate
or split into several branches. Indistinct ad-
ductor muscle attachment scar (Fig. lb) does
not form radial pattern (patterns of left and
right valves seem to differ significantly). Scar
consists of about 9-18 muscle segments.
Length 1 . 1 6-1 .20 mm, height 0.90-1 .05 mm
(without horns).

First antenna (Fig. Id): 8 -jointed, third
and fourth joints being not clearly separat-
ed. First joint without spines and hairs, with
1 bare dorsal bristle about as long as ventral
side of second through fourth joints com-
bined and 1 proximally spinous lateral bris-
tle, as long as 6 distal joints together, di-
rected dorsally. Second joint with hairs
dorso-laterally and proximo-ventrally, with

1 ventral bristle reaching to eighth joint or
exceeding it and 1 dorsal bristle reaching to
fifth joint, both bristles without discernible
spines. Third joint with long hairs on dorsal
and ventral sides, its dorsal margin some-
what shorter than dorsal margin of fourth
joint. Fourth joint with ventral group of hairs
and 2 spinous disto-ventral bristles, short
bristle reaching or exceeding eighth joint and
long bristle as long as total length of dorsal
margin of 7 distal joints. Fifth joint with
ventral group of hairs and 3 bristles on dis-
to-ventral comer, 1 bristle (not the longest)
medially with short hairs in proximal part,
long bristle about 2.2-2.8 times total length
of 7 distal joints, others about 2 times their
total length. Sixth joint with dorsal group
of hairs, without bristles. Seventh joint with
1 short dorsal spinous bristle as long as com-
bined length of 3-4 distal joints and with 2
long disto-ventral bristles, the longest of
them being about 3.1-3.5 times total length
of 7 distal joints. Eighth joint with two bris-
tles, dorsal bristle covered with stout small
spines along dorsal margin being about 3.9-
4.3 times total length of 7 distal joints.

Second antenna: Protopodite with cluster
of hairs in proximo-ventral area, without
bristles. Exopodite 9 -jointed, first joint di-
vided into long proximal and short distal
parts. Joints 2-8 each with 1 long bristle
with natatory hairs. Ninth joint with 2 bris-
tles, short and long, both devoid of natatory
hairs. Endopodite 2-jointed (Fig. Ic), but
very short distal part of second joint may
be interpreted presumably as third joint.
First joint with long hairs, 1 ventral and 2
dorsal bristles, ventral bristle with marginal
spines. Second joint with 2 transverse rows
of hairs, 1 lateral bristle bearing spines along
ventral margin and 2 pre- terminal bristles.
Terminal part of second joint (or third joint)
with 2 bristles, ventral one being the longest.

Mandible (Fig. 2a-c): Coxale endite seems
to have the same character set as Thau-
matoconcha radiata Komicker et Sohn,
1976: proximal set of teeth with four broad
teeth, some of them bifurcate. Distal set with

VOLUME 106, NUMBER 2

307

two large flat teeth bearing several cusps and
distally flattened spinous bristle. Basale with
knife-like process and 1 1 bristles, which can
be subdivided into 4 groups: 3 posterior
bristles, 1 anterior bristle, 5 lateral bristles
and 2 medial bristles, the longest of the lat-
ter with long hairs. First endopodite joint
with 1 dorsal bristle, second joint with 4
bristles on ventral margin and 2 bristles on
dorsal margin. Third endopodite joint with
7 bristles, the longest bristle with ca. 10-12
long spines directed distally in the middle
part of its posterior side, with ca. 80-90
short spines along posterior side and sparse
short spines along anterior side more dis-
tally. Shorter of the 2 stouter bristles seems
wavy bent with long spine-like processes (ca.
10 along each side) in the middle and with
very short fine spines more distally. Distal
half of the longest bristle of the terminal
joint with fine short spines.

Maxilla (Fig. 3a-d): First endite with 1
proximal and 7 distal bristles, second endite
with 7-9 bristles, third endite with 6 bristles
distal of which being not partitioned off'the
endite body. Basale with 1 long dorsal (cov-
ered with long hairs along ventral side) and
1 shorter lateral bristles. There is 1 lateral
bristle between basale and first endopodite
joint. First endopodite joint with long hairs
bearing 5 dorsal bristles and 2-3 disto-lat-
eral bristles. Second endopodite joint with
5-6 bristles, of which 1 distal bristle being
claw-like and not clearly partitioned off the
joint.

Fifth limb (Fig. 4a, b): Epipodial append-
age with 1 4 bristles arranged in 3 groups of
5 (dorsal), 5 and 4 (ventral). Protopodite
and endopodite with total of 19 bristles in
3 indistinct groups each with 6 (proximal),
7 and 6 (distal) bristles. First exopodite joint
hirsute with 1 long disto-dorsal bristle and
9 ventral bristles. Second joint with hairs
and 2 mid ventral bristles. Third joint with
3 terminal bristles, length of the shortest of
them (ventral) being about 36% of others.

Sixth limb (Fig. 4c): Epipodial appendage
with 1 5 bristles arranged in groups of 6 (dor-

sal), 4 and 5 (ventral) bristles. Hirsute pre-
coxale, coxale and basale each with 2 ven-
tral bristles. Disto-dorsal process of basale
(endopodite?) with 1 long bristle and small
spine. First exopodite joint with 2 disto-
ventral bristles, second with 1 dorsal and 2
mid ventral bristles. Third joint with 3 ter-
minal bristles, length of the shortest of them
(ventral) being about 39% of others.

Seventh limb (Fig. 4d): With 2 finely spi-
nous bristles of nearly equal length.

Copulatory organ (Fig. 4e, f) consists of
elongate anterior part and tapered posterior
part with 3 hardly discernible transparent
hair-like appendices. Anterior part with a
single long tooth-like process not reaching
distal part of the organ.

Lip morphology (Fig. 5 a) is consistent with
family diagnosis given by Komicker & Sohn
(1976).

Rod-shaped organ not discernible.

Furca (Fig. 5b): Each lamella with 2 long
anterior claws separated from lamella, fol-
lowed by 6 short spinous claws joined to
lamella and by 1 bare triangular process ori-
ented like preceding claws (the latter is ab-
sent on right lamella of the specimen from
St. 2649).

Posterior of body with hook-like process
proximal to furcal lamellae oriented pos-
teriorly.

Discussion

A comparison of the described specimens
of T. echinata with descriptions and draw-
ings by Miiller (1906) and Poulsen (1969)
and with supplemental description and
summary of characters given by Komicker
& Sohn (1976) reveals a set of differences
(Table 1). Only one of them may be attrib-
uted to sexual dimorphism for certain: the
number of ventral bristles of fourth joint of
first antenna (as in the other Thaumatocy-
pridid genera). Some differences may be
sexually dimorphic: presence of the lateral
bristle of first antenna's first joint, the bristle
number of first antenna's seventh joint, the

308

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

500

Fig. L Thaumatocypris echinata Miiller, 1906, adult male: a, Outside view of right valve of the specimen
from St. 2649; b, Adductor muscle-scar of the right valve of the specimen from St. 2649; c, Endopodite of left
2nd antenna of the specimen from St. 2661, medial view; d, Left 1st antenna of the specimen from St. 2661,
lateral view. (Scale in micrometers.)

VOLUME 106, NUMBER 2

309

Fig. 2. Thaumatocypris echinata Muller, 1906, adult male: a, 3rd endopodite joint of right mandible of the
specimen from St. 2649, lateral view; b, Basale of right mandible of the specimen from St. 2661, lateral view;
c, Endopodite of right mandible of the specimen from St. 2661, 3rd joint missing, medial view. (Scale in
micrometers.)

310

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Thaumatocypris echinata Muller, 1 906, adult male, St. 266 1 : a, Basale and endopodite of left maxilla,
lateral view (R— bristle found on right limb only); b, 1st endite of right maxilla, medial view; c, 2nd endite of
right maxilla, medial view (L— bristle found on left limb only); d, 3rd endite of left maxilla, lateral view. (Scale
in micrometers.)

VOLUME 106, NUMBER 2

311

Fig. 4. Thaumatocypris echinata Miiller, 1906, adult male: a. Distal joint of 5th limb of the specimen from
St. 2649; b, 5th limb of the specimen from St. 2661, distal bristles missing; c, 6th limb of the specimen from
St. 2661 (dotted lines denote folds which can be erroneously interpreted as sutures between joints); d, 7th limb
of the specimen from St. 2661; e, Copulatory organ of the specimen from St. 2661; f. Tip of copulatory organ
of the specimen from St. 2661. (Scale in micrometers.)

armament of male mandibular endopodite
terminal bristles (on two longest bristles the
male has spines in place of hairs drawn for
females by Miiller and Poulsen) though these

suggestions are not supported by the sexual
dimorphism in other genera of Thaumato-
cypridids (Komicker & Sohn 1976, Kor-
nicker & Iliffe 1989). Some differences may

312

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Thaumatocypris echinata Miiller, 1906, adult male, St. 2661: a. Upper lip; b, Furca. (Scale in mi-
crometers.)

prove to be a result of individual damage
or variability (total number of bristles on
mandible basale, of second joint of man-
dible endopodite or of first joint of fifth limb
exopodite). Some bristle groups are difficult
to count: epipodite bristles, bristles of max-
illae coxale and of fifth limb. In many other
instances there is much more similarity of
the male with the female described by Poul-
sen and reinvestigated by Komicker & Sohn
than with Miiller's non-adult female, though
judging by Miiller's drawing and by his
specimen length (1.55 mm) it could not be
younger than the A- 1 developmental stage
of Komicker & Sohn (1976).

As it follows from the comparison above
there are no morphological differences of
Thaumatocypris specimens investigated so
far which could be treated as important on
the species level. The length of the males
found is smaller than one could predict on
the basis of Miiller's and Poulsen's female
measurements (1.55 and 1.4 respectively)
and male : female length ratio (1.06) deter-
mined by Komicker & Sohn for Thauma-
toconcha radiata. But the ratio may be quite
different for Thaumatocypris and geograph-
ical variability also may prove to be quite

significant. The latter may be the situation
for the taxonomical importance of the pos-
terodorsal process of the right valve in the
males studied. Dr. Louis S. Komicker
pointed out in his letter reviewing the
manuscript that "... there does exist some
difference in the endopodite of the male and
female 2nd antenna in that Poulsen and
Miiller show it to be truncate. ..." This ob-
servation is true but the difference in ques-
tion may be attributed to sexual dimor-
phism. These considerations form the basis
for referring the males described to Thau-
matocypris echinata, but the final species
identification will become possible after
finding and investigation of Thaumatocy-
pris females from the Madagascar area.

The diagnosis of the genus Thaumato-
cypris given by Komicker & Sohn (1 976:34)
should be changed in several respects to in-
clude adult male and an emended diagnosis
follows:

Each valve with upper and lower long an-
teroventral protuberances. First antenna:
first joint with 0-1 lateral bristles; seventh
joint with 2-3 bristles, 1-2 ventral, 1 dorsal;
eighth joint with 2 bristles. Second antenna:
first endopodial joint with 3 bristles, 1 ven-

VOLUME 106, NUMBER 2

313

Table 1.— Main morphological differences of Thau-
matocypris specimens known so far (number of bristles
if other not indicated).

Female

Poulsen

Miiller

Character

1969

1906

Male

First antenna

1st joint: lateral

4th joint: ventral

6th joint

7th joint: ventral

Mandible

Basale

12?

Endopodite

2nd joint: ventral

2nd joint: dorsal

3rd joint

Maxilla

Basale: dorsal

Basale: ventral

Endopodite

1st joint: anterior

1st joint: posterior

2-3

Fifth limb

Epipodite

Protopodite +

endopodite

Exopodite

1st joint: ventral

Sixth limb

Epipodite

ca. 12

Furca

(no. of short claw-

like processes)

6-7

tral, 2 dorsal; terminal endopodial joint of
male without hook-like process. Mandible:
one of male endopodite terminal bristles
wavy bent and with long spine-like pro-
cesses in the middle. Maxilla: second en-
dopodite joint with 6 bristles. Fifth limb:
second exopodial joint without terminal
bristle on ventral margin; third exopodial
joint with 3 bristles. Sixth limb: process on
dorsal comer of first exopodial joint with 1
bristle and 1 minute spine; third exopodite
joint with 3 bristles. Rod-shaped organ is
minute cone-shaped or not discernible.

The most striking peculiarity of the spe-
cies is the absence of the sexual dimorphism
in the endopodite structure of the second

antenna: there is no hook-like process typ-
ical for other halocypridinid males. Instead,
the sexual dimorphism manifests itself in
the mandibular endopodite terminal bristle
morphology, not known in the other genera
of the Halocyprida order. In other respects
Thaumatocypris males do not differ from
other Thaumatocyprididae genera. For ex-
ample, the copulatory organ of T. echinata
differs from those described by Komicker
& Sohn (1976:38, fig. 18) for Thaumato-
concha, but the extent of the difference is
within the range of variability within the
latter genus.

The contents of the gut (crustacean-like
claws) suggest that the species is either a
carnivore or scavenger. The species has not
been found in the vertical plankton catches
in the localities where the "SOUND" ap-
paratus tows were made. Therefore the spe-
cies may be attributed presumably to the
near-bottom fauna as it was described by
Heinrich & Rudjakov (1991).

Acknowledgments

I thank Dr. Louis S. Komicker and anon-
ymous reviewers who offered valuable sug-
gestions for improvement of the manu-
script. Thanks to the crew and scientific team
of the R/V Vityaz for help collecting. Special
thanks to A. K. Heinrich, my wife and son
who helped with the figures. Manuscript
preparation was supported by the Soros
Foundation.

Literature Cited

Bir\ukov. S. G., V. N. Mar\atkin. A. S. Matveev. V.
A. Popov. & J. A. Rudjakov. 1 990. Experience
of usage of towed underwater apparatus
''SOUND" for study of near-bottom plank-
ton.— Okeanologiya 30:152-156 (in Russian,
English summan,).

Heinrich, A. K.. & J. A. Rudjakov. 1991. Vertical
distribution of plankton animals in the near-
bottom layer and the biological structure of the
ocean. — Okeanologiya 31:146-150 (in Russian,
English summar>').

Komicker. L. S.. & T. M. Iliffe. 1989a. New Ostra-
coda (Thaumatocyprididae. Halocyprididae)
from anchialine caves in the Bahamas. Palau,

314

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

and the Yucatan peninsula, Mexico. — Smith-
sonian Contributions to Zoology 470:1-47.

, & . 1989b. Troglobitic Ostracoda (Cy-

pridinidae, Thaumatocyprididae) from anchia-
line pools on the Galapagos Islands. — Smith-
sonian Contributions to Zoology 483:1-38.

, & I. G. Sohn. 1976. Phylogeny, ontogeny,

and morphology of living and fossil Thauma-
tocypridacea (Myodocopa, Ostracoda).—
Smithsonian Contributions to Zoology 219:1-
124.

Miiller, G. W. 1906. Ostracoda.— Wissenschaftliche
Ergebnisse der Deutschen Tiefsee-Expedition auf
dem Dampfer "Valdivia" 1898-1899 8:1-128
+ pis. 1-31.

. 1912. Ostracoda. — Das Tierreich 3 1 :i-xxxiii

+ 1-434, figs. 1-92.

Poulsen, E. M. 1969. Ostracoda-Myodocopa, 3A:
Halocypriformes-Thaumatocypridae and Hal-
ocypridae.— Dana-Report 75:1-100.

Rudjakov, J. A. 1968. A method of making perma-
nent preparations of small Arthropoda.- Zoo-
logichesky Zhumal 47:453-454 (in Russian, En-
glish summary).

P. P. Shirshov Institute of Oceanology,
Russian Academy of Sciences, 23 Krasikov
St, Moscow 117218, Russia.

PROC. BIOL. SOC. WASH.

106(2), 1993, pp. 315-324

PSEUDONICOTHOE BRANCHIALIS (CRUSTACEA:

COPEPODA: SIPHONOSTOMATOIDA: NICOTHOIDAE),

LIVING ON THE PANDALID SHRIMP

HETEROCARPUS SIBOGAE
OFF NORTHWESTERN AUSTRALIA

Arthur G. Humes and Geoffrey A. Boxshall

Abstract. —BoXh. sexes of Pseudonicothoe branchialis (Siphonostomatoida: Ni-
cothoidae), living on the marine shrimp Heterocarpus sibogae off the coast of
northwestern AustraUa, are described, the male for the first time. Both sexes
have hyaline flaps (suckers?) on the exopods of legs 1-3. The innermost element
on the free segment of leg 5 is sexually dimorphic. Pseudonicot hoe is recognized
as a valid genus, to which Paranicothoe procircularis (Carton) is transferred as
a new combination.

The siphonostomatoid copepod family
Nicothoidae Dana, 1852, includes several
genera whose species live on decapod crus-
taceans. Except Choniostoma Hansen, 1897,
all genera living on the decapods belong to
the Nicothoe group, as recognized by Box-
shall & Lincoln (1983). These genera are
Nicothoe Audouin & Milne Edwards, 1 826,
Choniosphaera Connolly, 1929, Choniomy-
zon Pillai, 1 962, Paranicothoe Carton, 1 970a
(see also 1970b), Hadrothoe Humes, 1975,
and Pseudonicothoe Avdeev & Avdeev,
1978. In Nicothoe, Choniosphaera, Choni-
omyzon, and Hadrothoe, the prosome of the
female is swollen. Only Paranicothoe and
Pseudonicothoe have a flattened prosome in
the female.

All species of Paranicothoe live on the
gills of penaeid and pandalid shrimps in
warm regions of the Indo-Pacific. As men-
tioned by Boxshall & Lincoln (1983), two
species of Paranicothoe, P. procircularis
(Carton, 1967) (see also Carton, 1970b) and
P. cladocera Carton, 1970a, may parasitize
epicaridean isopods which are in turn par-
asitic on shrimps. Paranicothoe procircu-
laris parasitizes Pseudione affinis (Sars)
(Isopoda) which lives in the branchial cavity
of the pandalid shrimp Plesionika ensis (A.

Milne Edwards) in the Java Sea. Parani-
cothoe cladocera inhabits the brood cavity
of Orbione natalensis Bourdon (Isopoda)
found in the gill cavity of the penaeid shrimp
Hymenopenaeus triarthrus Stebbing, in the
Mozambique Channel, off' Natal, South Af-
rica.

Pseudonicothoe branchialis Avdeev &
Avdeev, 1978, lives on the gills of the pan-
dalid shrimp Heterocarpus laevigatus Bate
in the Marshall Islands.

The purpose of this paper is to describe
the male of Pseudonicothoe branchialis for
the first time, and to redescribe certain fea-
tures of the female.

Siphonostomatoida Thorell, 1859

Nicothoidae Dana, 1852

Pseudonicothoe Avdeev & Avdeev, 1978

Redescription of the genus Pseudonico-
thoe, based on both sexes: Body cyclopi-
form, flattened, relatively unmodified. So-
mite bearing leg 1 fused with cephalosome.
Urosome in female 5 -segmented, in male
6-segmented. Caudal ramus with very long
inner terminal seta.

Rostrum weakly developed. Antennule
1 1 -segmented. Antenna 4-segmented with
small exopod bearing 1 seta. Siphon short

316

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

with terminal sucking disk. Mandible slen-
der blade. Maxillule with 2 lobes. Maxilla
small, strongly sexually dimorphic. Maxil-
liped 5 -segmented with terminal claw. Ven-
tral surface of cephalosome sexually di-
morphic, with maxillules and maxillae much
more widely separated in female than in
male, and with transverse arched scleroti-
zation in front of maxillae in female.

Legs 1-4 with 3 -segmented rami. Leg 1
with inner spine on basis. Second and third
segments of exopods of legs 1-3 in both
sexes with small round hyaline flaps. Second
segment of endopods in legs 1-3 with 1 in-
ner seta but this segment in leg 4 with 2
such setae.

Leg 5 with elongate free segment bearing
4 setae in both sexes, but innermost of these
setae sexually dimorphic, short and spini-
form in male, but long and setiform in fe-
male.

Pseudonicothoe branchialis
Avdeev & Avdeev, 1978

Material— 2 92, 5 <55, 3 copepodids from
the pandalid shrimp Heterocarpus sibogae
de Man, in shrimp trawl in 392-400 m,
FRV Soela, station NWS-7, 18°33.2'S,
117°30.9'E, 25 Apr 1983. One aduh 9, 3
adult (55, and 3 copepodids (2 99, 1 6) de-
posited in the Northern Territory Museum,
Darwin, Northern Territory, Australia; 1 9,
1 5, and 1 dissected S in The Natural History
Museum, London, England, BM(NH) Reg.
Nos. 1992.1067-1069.

Male. —Body (Fig. la) with flattened pro-
some subcircular in dorsal view. Length 1 .20
mm (1 .05-1.32 mm) and greatest width 0.6 1
mm (0.5 1-0.66 mm), based on 4 specimens
in lactic acid. Somite bearing first pair of
legs fused with cephalosome. Epimera of
metasomal somites rounded. Tergum of so-
mite bearing leg 3 arched forward medially,
exposing broad tergum of somite bearing leg
4, this tergum crenulated posteriorly and
showing median longitudinal sclerotization.
Ratio of length to width of prosome 1.15:

1. Ratio of length of prosome to that of
urosome 1.45:1.

Somite bearing leg 5 (Fig. lb) 65 x 170
jLim. Genital somite (Fig. Ic) rectangular in
dorsal view, 83 x 135 ixm (length including
leg 6 but not its setae). Four postgenital so-
mites from anterior to posterior 70 x 104,
78 X 92, 78 X 78, and 47 x 81 ixm. Anal
somite indented medially.

Caudal ramus (Fig. Id) elongate, outer
side 91 A^m, inner side 127 )Lim, and width
at midregion 29 )um. Ratio of outer length
to width 3.14:1. Ratio of inner length to
width 4.38: 1. Outer lateral seta, placed dor-
sally, 77 ium, dorsal seta 44 /xm, outermost
terminal seta 67 jum, and innermost ter-
minal seta, placed subterminally, short, 28
Aim, all these setae smooth. Two long me-
dian terminal setae 122 )um (outer) and 890
ixm (inner), both with very small serrations
along their midregions (Fig. le). Rami with
thick sclerotized inner and outer walls.

Rostrum (Fig. 1 f) weakly developed. An-
tennule (Fig. 2a) 1 1 -segmented, 450 jum long,
first 2 segments stout, remaining 9 segments
slender. Lengths of its segments (measured
along their posterior nonsetiferous mar-
gins): 56 (52 jum along anterior margin), 117,
16, 21, 42, 39, 39, 36, 36, 37, and 18 Aim,
respectively. Formula for armature: 3, 14,

2, 2, 2, 2, 2, 2, 2, 2, and 7+1 aesthete. Six
setae on second segment, associated with
conspicuous, incomplete, segmental scler-
otizations, noticeably stronger than other
setae. All setae smooth. In 4 males segmen-
tal sclerotizations in segments 4-11 stronger
in left antennule (Fig. 2b) than in right an-
tennule (Fig. 2a).

Antenna (Fig. 2c) short, 130 ixm. long in-
cluding terminal seta (compare length with
that of antennule, 450 iim). Four segment-
ed, but terminal seta showing trace of sub-
division. First segment (coxa) unarmed.
Second segment (basis) with minute exopod
4 X 4.5 fjLVCv with 1 seta 30 jum and orna-
mented with patch of spinules. Endopod
with large first segment having prolonged
inner distal comer and dense field of small

VOLUME 106, NUMBER 2

317

' i '

Fig. 1. Pseudonicothoe branchialis Avdeev & Avdeev, 1978. Male, a, dorsal (scale A); b, urosome, dorsal
(B); c, genital double somite and first postgenital somite, ventral (C); d, anal somite and caudal ramus, dorsal
(C); e, detail of longest seta on caudal ramus, dorsal (D); f, cephalosome, ventral (E).

318

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Pseudonicothoe branchialis Avdeev & Avdeev, 1978. Male, a, antennule, anterodorsal (scale C); b,
segments 3-11 of antennule, ventral (C); c, antenna, antero-outer (F); d, end view of siphon, ventral (G); e,
mandible, posterior (G); f, maxillule, ventral (G); g, maxilla, ventral (G); h, maxilliped, anterior (H); i, endopod
of maxilliped, anterior and slightly inner (F); j, endopod of maxilliped, posterior (F).

VOLUME 106, NUMBER 2

319

Spines on inner ventral surface; second seg-
ment small with 2 smooth inner setae and

1 terminal seta with slight indication of di-
vision proximally; terminal seta 44 fj.m with
small lateral spinules.

Siphon very short, in ventral view ap-
pearing as round sucking disk 60 ixm in di-
ameter (Fig. 2d). Mandible (Fig. 2e) simple
blade 117 ixm long. Maxillule (Fig. 2f) with

2 lobes, outer lobe with 2 setae, inner lobe
with 3 setae, all setae smooth. Maxilla (Fig.
2g) situated far posterior to maxillule (see
Fig. If). Subrectangular first segment 70 ^m
long and stout. Second segment 52 fim long,
bearing 1 proximal seta; first third partially
and indistinctly separated from second third,
terminal third slightly clawlike and weakly
separated from preceding third. Maxilliped
(Fig. 2h) 5 -segmented, 450 iim long includ-
ing claws. First segment (syncoxa) with 1
smooth inner seta, second segment (basis)
elongate with 1 smooth inner seta and or-
namented with large inner field of long spi-
nules and small spinules along outer surface.
Three endopodal segments, first with outer
seta and 2 small setules, second with 1 spine
18 jum, and third with 1 spine 29 fim, and
terminal claw 90 ixm. (Fig. 2i, j).

Ventral region between maxillipeds and
first pair of legs as in Fig. If.

Legs 1^ (Figs. 3a-c, 4a) with 3-segmented
rami. Formula for armature as follows:

Pi coxa 0-1 basis l-I exp I-l; I-l; 11,1,3

enpO-l;0-l; 1,5
P2 coxa 0-1 basis 1-0 exp I-l; I-l; 11,1,4

enpO-l;0-l; 1,1,4
P3 coxa 0-1 basis 1-0 exp I-l; I-l; 11,1,3

enpO-l;0-l; 1,1,3
P4 coxa 0-1 basis 1-0 exp I-l; I-l; 11,1,4

enp0-l;0-2; 1,1,2

Inner spine on basis of leg 1 minutely
barbed and 34 ^ni long. Outer seta on basis
of legs 1-3 long and feathered, but in leg 4
short and smooth. Terminal element on ex-
opod of leg 1 intermediate between spini-
form and setiform, barbed along outer side,
with long coarse setules along inner side.

Leg 2 with 2 outer elements on third en-
dopodal segment intermediate in form be-
tween spine and seta, with short spinules
along outer side and longer setules along
inner side (Fig. 3b). Exopods of legs 1-3
with small hyaline flaps (suckers?), 1 on seg-
ment 2 and 3 on segment 3 (Fig. 3a-c).

Leg 5 (Fig. 4b, c) with elongate free seg-
ment, in ventral view 86 x 25 fim, ratio
3.4:1, bearing 4 setae, 2 terminal (outer 117
fim, inner 130 ixm) and 2 subterminal; 1
subterminal seta long and barbed (85 ixm),
other subterminal seta (inner) short, 43 iim
stout, smooth, mucronate. Dorsal seta on
body 86 ^ni.

Leg 6 (Fig. 4d) with 3 setae from outer to
inner 65, 92, and 99 iira, innermost seta
with small proximal inner protuberance.

Color of living specimens unknown.

Female. —Body (Fig. 4e) with flattened
prosome (as shown in Avdeev & Avdeev,
1978: fig. 2) more elongate and tapered an-
teriorly than in male. Length 1.93 mm (1 .93-
1 .94 mm) and greatest width 1 .00 mm (1.01-
0.98 mm), based on 2 specimens in lactic
acid. Terga of somites bearing legs 2-4 part-
ly fused. Ratio of length to width of prosome
1.58:1. Ratio of length of prosome to that
of urosome 2.59:1.

Somite bearing leg 5 (Fig. 5a) 47 x 122
fim. Genital double somite 83 fim. long, 100
lim wide in anterior rounded half, 73 /um
wide in abruptly narrowed posterior half
Genital areas located dorsolaterally just an-
terior to junction of 2 halves of double so-
mite. Each area (Fig. 5b) with 2 small del-
icate setae. Three postgenital somites from
anterior to posterior 47 x 65, 50 x 55, and
31 X 49 Mm.

Caudal ramus similar to that of male but
slightly larger. 130 fim long on outer side,
39 fxm wide at midregion, ratio 3.33:1.

Rostrum, antennule, antenna, siphon,
mandible, and maxillule, arranged as in Fig.
5c, like those of male. Maxilla (Fig. 5d) small,
55 fj,Tn long, apparently 2-segmented, with
terminal spine. First segment with 2 small
setae. Maxilliped as in male. Maxillules and

320

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Pseudonicothoe branchialis Avdeev & Avdeev, 1978. Male, a, leg 1 and intercoxal plate, anterior
(scale H); b, leg 2 and intercoxal plate, anterior (H); c, leg 3 and intercoxal plate, anterior (H).

VOLUME 106, NUMBER 2

321

Fig. 4. Pseudonicothoe branchialis Avdeev & Avdeev, 1978. Male, a, leg 4 and intercoxal plate, anterior
(scale H); b, leg 5, dorsal (C); c, leg 5, ventral (C); d, leg 6, ventro-outer (C). Female, e, dorsal (A).

322

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Pseudonicothoe branchialis Avdeev & Avdeev, 1978. Female: a, urosome, dorsal (scale E); b, genital
area, dorsal (C); c, cephalosome, ventral (A); d, maxilla, ventral (G); e, exopod of leg 1, anterior (H); f, leg 2,
anterior (H); g, exopod of leg 3, anterior (H); h, leg 5, dorsal (H).

VOLUME 106, NUMBER 2

323

maxillae widely separated, with transverse
sclerotized bar between them, as in Fig. 5c.

Legs 1-4 segmented and armed as in male.
Legs 1-3 with exopods having small round
hyaline flaps, 1 on second segment and 3 on
third segment (Fig. 5e-g), as in male.

Leg 5 (Fig. 5h) with free segment 133 x
52 jum, ratio 2.56:1. Four setae from inner
to outer 125, 140, 133, and 122 ixm. Dorsal
seta 146 iim.

Leg 6 represented by 2 setae on genital
area (Fig. 5b).

Only 1 somewhat damaged egg sac seen,
separated from female, containing approx-
imately 12 flattened, discoidal, linearly ar-
ranged eggs, measuring 1045 x 330 ^um.

Color unknown.

Remarks. —We have been unable to make
a direct comparison of the specimens from
Heterocarpus sibogae with the type speci-
mens of Pseudonicothoe branchialis. The
types consist of two females, holotype and
paratype, deposited in the Laboratory of
Parasitology of Marine Animals, Pacific
Ocean Scientific Research Institute of Fish-
eries and Oceanography, Vladivostok, Rus-
sia.

Although the description and figures of P.
branchialis published by Avdeev & Avdeev
(1978) conform in major respects to our
specimens from Australia, a few minor dif-
ferences may be observed: (1) few relatively
long setae on the antennule (Avdeev &
Avdeev's fig. 5), (2) the antenna with a
feathered seta on the exopod and the fourth
segment with one of the two small setae
feathered (their fig. 10), (3) the outer branch
of the maxillule with three setae (their fig.
6), (4) the first segment of the maxilla in the
female with one curved claw (their fig. 7),
and (5) the free segment of leg 5 relatively
short, ratio 1.64:1 (their fig. 7). We believe
that these small differences may be attrib-
utable to the relatively small number of
specimens studied, the difficulties of obser-
vation, and to the style of illustration.

The exopods of swimming legs 1-3 of both
male and female P. branchialis are orna-

mented with marginal structures referred to
as hyaline ffaps in the present account, and
as suckers by Avdeev & Avdeev (1978).
When viewed from the side, as in Fig. 3a-
c, these structures appear to be hyaline ex-
tensions of the lateral margin of the exo-
podal segments. When they are reffexed
across the surface of the exopodal segment,
radial surface striations are visible and their
appearance is more suckerlike. Similar
structures were figured on legs 1-3 of male
P. procircularis (Carton) by Carton (1967).

The discovery of the male of P. bran-
chialis prompted us to reconsider the valid-
ity of the genus Pseudonicothoe which was
treated as a subjective synonym of Parani-
cothoe Carton by BoxshaU & Lincoln (1983).
The type species of Paranicothoe, P. cla-
docera, differs from Pseudonicothoe in the
segmentation of the antennules, in the seg-
mentation and armature of the maxilliped,
and in the spine and setal formula of the
swimming legs. According to Huys &
Boxshall (1991) the antennule is 10-seg-
mented in P. cladocera, the distal part com-
prises relatively short segments, and the api-
cal segment is longer than the two preceding
segments combined, whereas in P. bran-
chialis the antennule is 1 1 -segmented, the
distal part comprises relatively long seg-
ments, and the apical segment is markedly
shorter than the subapical segment. The en-
dopod of the maxilliped of P. cladocera is
2-segmented and is armed with one claw on
the first segment and two on the second. In
P. branchialis, by comparison, the endopod
is 3 -segmented, with the first segment car-
rying three short setae, the second segment
bearing one claw, and the third two claws.
Finally, the endopod of leg 4 has a setal
formula of 0-1; 0-2; 1,1,2 in P. branchialis
and 0-1; 0-1; 1,1,2 in P. cladocera.

These differences justify the generic level
separation of Pseudonicothoe and Parani-
cothoe, as represented by its type species,
and we propose to recognize Pseudonico-
thoe as a valid genus again, thereby revers-
ing the proposal made by Boxshall & Lin-

324

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

coin (1983). The type species of
Pseudonicothoe is P. branchialis. Parani-
cothoe is also a valid genus, containing only
the type species, P. cladocera. We propose
to transfer Paranicothoe procircularis (Car-
ton) to Pseudonicothoe, as a new combi-
nation, Pseudonicothoe procircularis (Car-
ton, 1967). This transfer is based on the 11-
segmented condition of the antennule, and
on the spine and setal formula of the swim-
ming legs.

Acknowledgment

We thank Dr. A. J. Bruce, Northern Ter-
ritory Museum, Darwin, Australia, for
sending specimens of the nicothoid to us for
study. The study of the copepods was aided
by a grant (BSR 88 21979) to AGH from
the National Science Foundation of the
United States.

Literature Cited

Audouin, V., & H. Milne Edwards. 1826. Memoire
sur la Nicothoe, animal singulier qui suce le sang
des homards.— Annales de Sciences Naturelles
9:345-358.

Avdeev, G. V., & V. V. Avdeev. 1978. Pseudoni-
cothoe branchialis gen. et sp. n. (Crustacea, Co-
pepoda) from gills of Heterocarpus laevigatus
from the Pacific Ocean.— Zoologicheskii Zhur-
nal 57:1893-1897.

Boxshall, G. A., & R. J. Lincoln. 1983. Some new
parasitic copepods (Siphonostomatoida: Ni-
cothoidae) from deep-sea asellote isopods.—
Journal of Natural History 17:891-900.

Carton, Y. 1967. Description de Mcor/zo^ prodrcw-
laris n. sp. (Crustacea, Copepoda) discussion sur
la forme male. — Videnskabelige Meddelelser fra
Dansk Naturhistorisk Forening 130:143-152.

. 1970a. Description de Paranicothoe n. gen.

un nouveau representant de la famille des Ni-
cothoidae.— Galathea Report 11:239-246.

. 1970b. Le genre Paranicothoe, un nouveau

representant de la famille des Nicothoidae.—
Second International Congress of Parasitology,
Journal of Parasitology 56(II):47-48.

Connolly, C. J. 1 929. A new copepod parasite Choni-
osphaera cancrorum, gen. et sp. n., representing
a new genus, and its larval development.— Pro-
ceedings of the Zoological Society of London
1929:415^27.

Dana, J. D. 1852. Conspectus crustaceorum quae in
orbis terrarum circumnavigatione Carolo Wilkes
e classe reipublicae foederatae duce, lexit e de-
scripsit Jacobus D. Dana. Part 2.— Proceedings
of the American Academy of Arts and Sciences
2:9-61.

Hansen, H.J. 1897. The Choniostomatidae. A family
of Copepoda, parasites on Crustacea Malacos-
traca. Copenhagen, Denmark. Pp. 1-205.

Humes, A. G. 1975. Hadrothoe crosnieri n. gen., n.
sp. (Crustacea, Copepoda), from a penaeid
shrimp (Crustacea, Decapoda) in Madagas-
car. — Zoologischer Anzeiger 195:21-34.

Huys, R., & G. A. Boxshall. 1991. Copepod evolu-
tion. The Ray Society 159:1-468.

Pillai, N. K. 1962. Choniomyzon gen. nov. (Cope-
poda: Choniostomatidae) associated with Pan-
w/z>u5.— Journal of the Marine Biological As-
sociation of India 4:95-99.

Thorell, T. 1859. Till kannedomen om vissa parasi-
tiskt lefvande Entomostracer. — Ofversigt af
Kongliga Akademiens Forhandlingar 1 6(8): 335-
362.

(AGH) Boston University Marine Pro-
gram, Marine Biological Laboratory, Woods
Hole, Massachusetts 02543, U.S.A.; (GAB)
The Natural History Museum, Cromwell
Road, London, England SW7 5BD.

PROC. BIOL. SOC. WASH.

106(2), 1993, pp. 325-331

BOREOMYSIS OPARVA, A NEW POSSUM SHRIMP

(CRUSTACEA: MYSIDACEA) FROM AN EASTERN

TROPICAL PACIFIC SEAMOUNT

Jennifer Saltzman and Thomas E. Bowman

Abstract.— Boreomysis oparva is described from near-bottom low oxygen
waters surrounding an inactive seamount, Volcano 7. It is characterized by an
upturned rostrum, large eyes without papillae, and a male pleopod 3 exopod
with 3 spiniform setae on each of the last eight segments. It is abundant slightly
below the seamount summit in near-bottom low oxygen water, but absent from
water near the seamount base (with higher oxygen) and from low oxygen pelagic
waters.

Volcano 7 is an inactive seamount 20 km
in diameter in the eastern tropical Pacific at
13°23'N, 102°27'W. It arises from a depth
of 3400 m to a summit at 730 m; the latter
penetrates the pronounced oxygen-mini-
mum zone of the region (Wishner et al. 1 990,
Levin et al. 1991). In November 1988 the
plankton and benthos of Volcano 7 were
sampled from shipboard on the K/V Atlan-
tis II and from the submersible D.S.R.V.
Alvin. The samples collected from the sub-
mersible contained more than 400 speci-
mens of an undescribed species of the mys-
idacean genus Boreomysis, described and
illustrated herein.

Methods

A multiple opening-closing 8 -net system
(183 jLim mesh) mounted on the Alvin
(Wishner & Go wing 1987) was used to col-
lect zooplankton 1-3 m above the bottom
during daytime dives. The 1 -liter Plexiglas
cod-end chambers were equipped with
spring-loaded front and rear doors that were
fixed open during the tow and were snapped
shut at the end of the tow, making the cham-
ber watertight. Simultaneously with the
chamber closing a spring-loaded needle
punctured a rubber balloon within the
chamber, releasing its contents of glutaral-
dehyde and fixing the sample in situ. Plank-

ton samples were also collected from ship-
board with a 1 m^ MOCNESS plankton net
system (333 ixm mesh) in vertically strati-
fied tows to 1 200 m depth.

Order Mysidacea Boas, 1883

Suborder Mysida Boas, 1883

Family Mysidae Dana, 1850

Subfamily Boreomysinae Holt & Tattersall,

1905

Genus Boreomysis G. O. Sars, 1869

Boreomysis oparva, new species

Figs. 1-20

Material. —Sqq TablQ 1.

Types. — Holotype 6,22.5 mm. Dive 2 145,
Net 4, USNM 251923. Paratypes: Dive
2139, Net 8, 14 specimens, USNM 251918;
Dive 2142, Net 8, 100 specimens, USNM
251919; Dive 2143, Net 4, 2 specimens,
USNM 251920; Dive 2144, Net 8, 4 spec-
imens, USNM 251921; Dive 2145, Net 4,
6 specimens, USNM 251922.

Etymology.— From "o," the chemical
symbol for oxygen, plus the Latin "parvus"
(little, slight), referring to the occurrence of
this mysid in low-oxygen water.

Description. —Length up to about 25 mm.
Anterior margin of carapace slightly con-
vex. Rostrum acute, reaching V3 to V2 length
of 1 St segment of antenna 1 , directed dor-
sally at angle of about 45° to lateral axis of

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table \.—Boreomysis oparva collected from Volcano 7 by the submersible D.S.R.V. Alvin in November 1988.
Collections marked with an asterisk are deposited in the National Museum of Natural History, Smithsonian
Institution.

Dive-Net

Juv.

Maximum
length
(mm)

Mid-depth
(m)

Location

2139-1

18.0

790

summit

26.5

852

summit

22.5

790

summit

—

21.2

810

summit

25.4

852

summit

2140-1

—

19.8

804

summit

—

20.0

793

summit

23.0

815

summit

—

18.8

874

summit

—

21.5

804

summit

—

18.3

793

summit

23.9

817

summit

18.7

876

summit

2142-1

23.0

797

summit

—

21.4

782

summit

23.4

788

summit

116

25.0

797

summit

22.2

797

summit

22.5

788

summit

21.5

808

summit

2143-2

22.1

1300

flank

— ,

—

1297

flank

—

1308

flank

—

20.2

1300

flank

22.0

1309

flank

2144-1

—

1289

flank

—

14.3

1179

flank

—

1250

flank

—

20.8

1195

flank

—

17.5

1259

flank

2145-3

—

9.6

1323

flank

22.5

1352

flank

—

20.3

1304

flank

carapace. Antero ventral comer of carapace
acute; cervical groove well developed. Car-
apace covering laterally 1 st segment of ex-
opods of thoracopods and posteriorly about
Vi of pleonite 1 . Eye with rather short, nar-
row stalk and broad cornea, reaching well
beyond midlength of 1 st segment of antenna
1 , without ocular papilla.

Antenna 1 , 1 st segment of peduncle nar-
rower than 2nd and 3rd segments, nearly
3 X as long as wide, with medial papilla at

distal ^/k bearing 4 setae and distolateral pa-
pilla with 5 setae. 2nd segment short, with
2 long dorsal setae, shorter seta on disto-
medial comer, and 3 setae on distolateral
papilla. 3rd segment slightly more than Vi
length of 1 st segment, with cluster of 7 setae
at distomedial comer and 3 setae on quad-
rate process at midwith of distal margin. S
antennular brush very dense and long; if
present, S lobe obscured by setae of brush.
1st segment of lateral flagellum densely

VOLUME 106, NUMBER 2

327

Figs. 1-9. Boreomysis oparxa. 1. Head and thorax, dorsal; 2. Same, lateral: 3. Eye and rostrom. dorsal: 4,
Antenna. 1. 2. dorsal: 5. Peduncle of 5 antenna 1, ventral; 6. Antenna 2. dorsal; 7. Right mandible, gnathal
surface: 8. Left mandible, same, but molar omitted: 9. Maxilla 1. outer ramus.

328

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 10-16. Boreomysis oparva. 10. Apex of mandibular palp; 11, Maxilla 2; 12, Endites of maxilla 1, 1st
endite (below) enlarged: 13, Maxilliped; 14, Thoracopod 2: 15. Thoracopod 5; 16. Penis, lateral.

VOLUME 106, NUMBER 2

329

Figs. 17-20. Boreomysis oparva. 17, Exopod of 3 pleopod 3, distal segments; 18, Telson, dorsal; 19, Apex
of telson, dorsal; 20, Right uropod, dorsal, with enlarged detail of spines of exopod and endopod.

armed with setae about as long as 1st seg-
ment of peduncle; only proximal parts of
setae shown in Fig. 4.

Antenna 2 peduncle not quite reaching
midlength of scale; segment 1 produced dis-
tolaterally into spiniform process. Scale
nearly 4 x as long as broad; terminal spine
reaching beyond nearly truncate apex, which
slants laterally at about 1 5° to axis of scale.

Left mandible with bicuspid incisor and
lacinia; spine-row of 8 spines. Right man-
dible with tricuspid incisor; lacinia dichot-
omous, dorsal ramus a curved pointed tooth,
ventral ramus quadrate, bearing 3 spines;
spine-row formed of a single serrate spine
separated by gap from 9 simple spines with
common base. Palp 2nd segment broad
proximally, narrowing distally, with scat-

tered marginal setae; 3rd segment about 0.7
length of 2nd segment, distal half with close-
set plumose marginal setae and 2 long apical
setae.

Maxilla 1 outer ramus with 7 setae on
surface and 1 5 apical spines.

Maxilla 2 protopod margin with dense
covering of fine setae interspersed with 9
shorter and stouter setae. 1 st endite with 1 6,
lobes of 2nd endite with 1 2 and 1 5 apical
setae respectively. Endopod segments sub-
equal in length; 1 st segment with 9 setae on
medial margin; 2nd segment with about 30
marginal setae. Exopod reaching slightly be-
yond 1st endopod segment, with about 33
marginal setae.

Maxilliped (endopod of thoracopod 1)
endite of basis reaching distal margin of me-

330

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

rus. Carpus nearly as long as propus and
dactyl (excluding nail) combined. Medial
margin of all segments densely setose.

Endopods of thoracopods 2-8 with pro-
pus divided by incomplete suture. Dactyl of
thoracopod 2 without nail. Anterior margin
of propus of thoracopods 3-8 with 3 clusters
of 5-7 setae in which 1 seta is much longer
than others. Exopods of thoracopods with
17-20 segments.

Penis about as long as basiopod of thora-
copod 8, widening distally; posterior margin
with right-angled bend proximal to rounded
setose apex.

3rd 5 pleopod with 18 segmented endo-
pod and longer 23 segmented exopod. On
exopod paired plumose setae of proximal
segments replaced on last 8 segments by 3
simple spiniform setae on each segment, ex-
cept 2 such setae on apical segment; 2 of the
3 setae at distomedial comer, 1 at distolater-
al comer.

Telson slightly more than 3 x as long as
width at base, slightly longer than pleonite
6, posterior half with slightly concave mar-
gins. Apical cleft about Vk length of telson,
each side with about 30 teeth, margins grad-
ually diverging posteriorly; base not dilated
but with narrow slit. Each lobe of apex armed
with long spine flanked laterally by 1 spine
and medially by 2 spines; flanking spines
half length of long spine. Lateral margins of
telson each armed with 20-23 spines, some
shorter than others but not arranged in reg-
ular pattern.

Endopod of uropod slightly longer than
telson, with 2 spines (0-1 in immatures) in-
serted ventrally near medial margin adja-
cent to statocyst. Exopod % longer than en-
dopod, lateral margin naked for proximal
^7, 2 spines on outer margin at distal end of
naked part.

Comparisons. —Nearly 40 species ofBor-
eomysis are recognized currently, but it is
uncertain how many of them will prove
eventually to be valid. Some are based only
on females or immature males, hence the
structure of the mature male exopod of

pleopod 3, a character of high taxonomic
value, is unknown. The condition in B.
oparva, the 8 distal segments each with 3
simple setae, is at present unique; other spe-
cies in which this pleopod has been de-
scribed have 2 such setae. This feature,
combined with the upturned rostrum, the
large eyes lacking papillae, and the 2 spines
on both the endopod and exopod of the uro-
pod, readily distinguish B. oparva from sim-
ilar species.

Birstein & Tchindonova (1958) key out
27 species and 2 varieties of Boreomysis,
and li (1964) gives a key to the 10 species
and 2 varieties that he recognized from the
northwestern Pacific.

Ecology. —The summit of Volcano 7 (730
m) lies in the oxygen minimum zone (Wish-
ner et al. 1 990), and B. oparva was not found
there. Slightly deeper at the lower summit
(ca. 790 m), associated with the increase in
oxygen concentration from 0.08 to 0.88 ml/
liter (Levin et al. 1991), B. oparva became
abundant, reaching a maximum of 1 14/m^.
At the flank of the seamount (1185-1310
m) B. oparva was less abundant and absent
from most of the samples. It was absent
from near-bottom waters at the base of the
seamount (3400 m) and from pelagic waters
(0-1200 m) surrounding the seamount. The
abundance peak at the lower summit was
similar to that found for many infaunal taxa
(Levin etal. 1991).

i^oo^. — Gowing & Wishner (1992) found
dense numbers of gram-positive bacteria-
like bodies in the guts of some specimens
of this mysid, suggesting that they oppor-
tunistically consume bacterial aggregates or
mats at the lower boundary of the oxygen
minimum zone.

Acknowledgments

We thank Dr. Karen F. Wishner for pro-
viding us with the specimens of the new
species of Boreomysis and for reviewing the
manuscript. Ship time for the collection of
the specimens was provided by NSF grants

VOLUME 106, NUMBER 2

331

OCE 87-16564 to Dr. Wishner and OCE
87-0193 to Dr. Marcia M. Gowing, and
ONR contract N00014-84-K-0081 to Dr.
Lisa A. Levin.

Literature Cited

Birstein, Ya. A., & Yu. G. Tchindonova. 1958. Deep-
sea mysids from the northwestern part of the
Pacific Ocean.— Trudy Instituta Okeanologii 27:
258-355. [in Russian]

Boas, J. E. V. 1883. Studien iiber die Verwandt-
schaftsbeziehungen der Malakostraken.— Mor-
phologisches Jahrbuch 8:485-579.

Dana, J. D. 1850. Synopsis generum crustaceorum
ordinis "Schizopoda" J. D. Dana elaboratus, et
descriptiones specierum hujus ordinis quae in
orbis terrarum circumnavigatione, Carolo Wilkes
e Classe Reipublicae Faederatae Duce, auctore
lectae (pars I).— American Journal of Sciences
and Arts (2) 9:129-133.

Gowing, M. M., & K. F. Wishner. 1992. Feeding
ecology of benthopelagic zooplankton on an
eastern tropical Pacific seamount.— Marine Bi-
ology 112:451-467.

Hoh, E. W. L., & W. M. Tattersall. 1905. Schizop-
odous Crustacea from the north-east Atlantic
slope. — Report on the Sea and Inland Fisheries
of Ireland, 1902-1903, part 2, Appendix 4:99-
152, pis. 15-25.

li, N. 1964. Fauna Japonica, Mysidae. Biogeograph-
ical Society of Japan, Tokyo, 610 pp.

Levin, L. A., C. L. Huggett, & K. F. Wishner. 1991.
Control of deep-sea benthic community struc-
ture by oxygen and organic matter gradients in
the eastern Pacific Ocean.— Journal of Marine
Research 49:763-800.

Sars, G. O. 1869. Undersogelser over Christiania-
Qorden Dybvandsfauna anstillede paa en i Som-
meren 1868 foretagen Zoologisk Reise.— Nytt
Magasin for Naturvidenskapene 16:305-362.

Wishner, K. F., & M. M. Gowing. 1987. In situ fil-
tering and ingestion rates of deep-sea benthic
boundary-layer zooplankton in the Santa Cat-
alina Basin. — Marine Biology 94:357-366.

, L. Levin, M. Gowing, & L. Mullineaux. 1 990.

Multiple roles of the oxygen minimum in ben-
thic zonation on a deep seamount.— Nature 346:
57-59.

(JS) Graduate School of Oceanography,
University of Rhode Island, Narragansett,
Rhode Island 02882-1197, U.S.A.; (TEB)
Department of Invertebrate Zoology (Crus-
tacea), National Museum of Natural His-
tory, Smithsonian Institution, Washington,
D.C. 20560, U.S.A.

The sequence of the authors is according
to the recency of their birthdates.

PROC. BIOL. SOC. WASH.

106(2), 1993, pp. 332-338

THE IDENTITY OF TALITROIDES ALLUAUDI

(CHEVREUX) (CRUSTACEA: AMPHIPODA: TALITRIDAE)

WITH NOTES ON A NEW LOCALITY

Hiroshi Morino and Reuven Ortal

Abstract. —A lectotype of Talitroides alluaudi (Chevreux) is designated from
the type-series and described to remove taxonomic confusion on the identity
of this species. The discovery of this species in Israel is briefly discussed.

Examination of terrestrial talitrids col-
lected from southern parts of Israel revealed
two genera and three species, one of which
proved to be Talitroides alluaudi. This spe-
cies is one of the most widespread landhop-
pers, known from the tropics to warm-tem-
perate regions and hothouses in Europe and
North America (Friend & Richardson 1986),
though so far not known from the Levant.
Although some authors have remarked on
various characters, especially concerning the
pleopods, of this species (Medcof 1940, Pal-
men 1949, Andersson 1962), no full de-
scriptions or figures have been published
since the original description of Chevreux
(1896, 1901). Close examination of material
from Israel, with reference to Chevreux' s
descriptions, revealed a few minor but dis-
tinct discrepancies between them as well as
between previous descriptions of T. alluau-
di.

Chevreux (1896) described this species
three times from two localities. He gave a
brief original description of this species on
the basis of material procured from hot-
houses at the Paris Museum ("les serres du
Museum de Paris"). In this paper he men-
tioned that the Paris material was identical
to that collected from the Seychelles. Al-
though he did not specify the locality of the
type, the title of his paper suggests that Paris
is the probable type locality. Shoemaker
(1936), however, assumed that the Sey-
chelles was the type locality. In 1901, Chev-
reux treated the Seychelles material in de-

tail. The Paris material was described again
in 1925 (Chevreux & Page 1925) with il-
lustrations. A study of these three descrip-
tions discloses the following apparent un-
conformity in some characters: The Paris
1896 and Seychelles material have well-de-
veloped brood plate on female gnathopod
2, though this is lacking in the Paris 1925
material, and; the Seychelles material has
1 -articulated and 3 -articulated inner ramus
on pleopods 1 and 2, respectively, though
the Paris 1925 material has 4 -articulated
and 1 -articulated inner ramus, respectively
(no description of pleopods for the Paris
1896 material). If these descriptions are
taken authentically, we could recognize three
species in the Chevreux' s concept of T. al-
luaudi: two from Paris and one from the
Seychelles. And if the Paris 1896 material
was from the Seychelles, as assumed by
Shoemaker (1936), two species may be dis-
cernible, from Paris and the Seychelles, re-
spectively. Medcof (1940) has also pointed
out some confusion in the earlier descrip-
tions of this species, which he attributed to
geographical variation ("based on exami-
nations of specimens collected from areas
sometimes widely separated"). Since Med-
cof s analysis of variation is limited to the
pleopod structure, he did not notice the
variation in the brood plate. In addition,
material from Israel at hand displays a pe-
culiar feature of the pereopod dactyl, which
was described in the Seychelles material dif-
ferently, and was not mentioned for the Par-

VOLUME 106, NUMBER 2

333

is material. Thus it is necessary to deter-
mine the original concept of T. alluaudi
through direct reference to the type-series.
The type material of Talitrus alluaudi de-
posited in Museum National d'Histoire Na-
turelle in Paris consists of preserved spec-
imens from four localities (MNHN Paris
Am 4500, 4501, 4502, 4503) and slides
(MNHN Paris Am 4504). The label on the
slides ("Serre du Museum") suggests that
they comprise the type specimen. Unfor-
tunately the condition of the slides is so bad
that it is not possible to discriminate the
pertinent characters, and all the preserved
specimens are from other than the Paris
Museum. Thus the lectotype is selected from
the locality nearest to the Paris Museum and
described to resolve the problem of T. al-
luaudi. Material from Israel and Hawaii is
also examined.

Systematic Account

Talitroides alluaudi (Chevreux, 1896)
Figs. 1, 2

Talitrus Alluaudi Chevreux, 1896:112, figs.

1-4; 1901:389, figs. 1-6. -Chevreux &

Page, 1925:270, figs. 280-281.
Orchestia senni Menzel, 191 1:438, figs. 4-9
Talitroides alluaudi. — Palmen, 1 949:6 1 , figs.

1-12.— Andersson, 1962:21 1, figs. 1-3.—

Bousfield, 1984:210.

Material examined. —T^o females (5.5
mm— lectotype, 5.0 mm— paralectotype),
from Serres de la Ville de Paris, Boulogne
sur Seine, France (MNHN Paris Am 4500);
1 female (5.0 mm), from lies Sechelles,
Mahe, Auct. det 1901 (MNHN Paris Am
4502); 2 females (6 mm and 7 mm), from
Ponta Delgada, lie S. Michel, Agores, Aug
1930, M. Mequignon collector, (MNHN
Paris Am 4503); 5 females (up to 5.5 mm),
from Ben Gurion Univ. campus, Beersheba,
Israel, bamboo stands, 10 Dec 1987, Y.
Margalit collector (Morino Cat. No. T.766),
4 Mar 1989, R. Ortal collector (Morino Cat.
No. T836); 1 female (4.5 mm), from Hal-

awa, Molokai Is., Hawaii, 9 Mar 1968, Si-
nonaga collector (Morino Cat. No. T.868);
1 female (4.8 mm), from Kokee, Kauai Is.,
Hawaii, 11-12 Mar 1968, Sinonaga collec-
tor (Morino Cat. No. T.869).

Description of lectotype.— Eyt small in
size, subround. Inferior antennal sinus me-
dium deep. Head longer than deep.

Antenna 1 exceeding mid-point of pe-
duncular article 5 of antenna 2, peduncular
articles subequal in length; flagellum a little
shorter than peduncle, 6 -articulated. An-
tenna 2: peduncle weakly spinose, article 5,
1.79 times as long as article 4; flagellum
subequal to peduncle in length, 1 1 -articu-
lated.

Upper lip: as wide as deep. Mandible: in-
cisor chitinized strongly, brown in color,
5 -dentate; left lacinia chitinized as incisor,
4-dentate; right lacinia not strongly chitin-
ized, with 3 cusps; spine rows of 2 broad
plumose bristles. Lower lip normal. Maxilla
1: innermost spine of outer plate leaning
medially, outermost 2 spines without den-
ticles; palp small, joint of article 2 indistinct.
Maxilla 2: inner plate distolaterally pro-
duced into pointed tip, several medio-distal
setae plumose. Maxilliped: inner plate api-
cally truncate, with 3 weak spine-teeth, in-
creasing in size laterally, and with several
plumose stiff" setae; outer plate weakly ar-
cuate with rounded apex, weakly setose; palp
articles 2 and 3 slender, with a few stiff" setae
distally, article 2 with remnant of medial
lobe represented by elongate setae, article 4
partially fused to article 3.

Gnathopod 1 : coxa truncate distally, low-
er margin spinose, inner shelf weak, with a
spine; basis broadened distally; carpus 1.50
times as long as propod; propod simple,
gradually narrowed to base of dactyl, pos-
terior margin with 3 stiff* long spines and
several shorter submarginal spines on inner
and outer sides, anterior margin with 2 spine
clusters; dactyl cuspate anteriorly, base with
a stiff* seta posteriorly, nail as long as base.
Gnathopod 2: coxa cuspate posteriorly, as
deep as wide, lower margin rounded and

334

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. I. Talitroides alluaudi (Chevreux). Boulogne sur Seine, France. The lectotype, MNHN Am 4500-1
(female, 5.5 mm body length). A. lateral view; B, antenna 1; C, antenna 2; D, left mandible; E, right mandible;
F. maxilla 2; G, maxilla 1: H. maxilliped; I. distal part of right palp of maxilliped; J, upper lip; K, lower Up.

weakly spinulose; basis slender, anterior
margin spinulose; merus and carpus with
tumescence posterioriy, carpus as long as
propod; propod anterior margin weakly se-
tose, posterior tumescent lobe well devel-
oped, exceeding dactyl by half of propod
length.

Coxae 3 and 4 shallower than wide, with

a stiff seta posteroventrally, and a few spi-
nules ventrally. Coxa 5 anterolobate, ante-
rior lobe deeper than that of pereopod 4.
Coxa 6: posterior lobe truncate distally, an-
terior margin of the lobe vertical. Coxa 7
shallow.

Pereopods 3-7 cuspidactylate, propod
lacking hinge spine, dactyl base with a stiff

VOLUME 106, NUMBER 2

335

Fig. 2. Talitroides alluaudi (Chevreux). Boulogne sur Seine. France. A. gnathopod 1; B, dactyl of gnathopod
1; C, gnathopod 2; D. coxal gill of gnathopod 2; E. pereopod 3; F & G. dactyl of pereopod 3; H, pereopod 4; I,
dactyl of pereopod 4; J. pereopod 5: K. dactyl of pereopod 5; L, pereopod 6; M, coxal gill of pereopod 6; N,
dactyl of pereopod 6; O, pereopod 7; P. dactyl of pereopod 7: Q. abdominal side plates 1-3; R, pleopod 1; S,
pleopod 2; T, pleopod 3; U, uropod 3: V. uropod 2: W. uropod 1: X. telson. G = Azores, MNHN Am 4503-1
(female 6.0 mm); M = Boulogne sur Seine, France, paralectotype MNHN Am 4500-2 (female 5.0 mm); others
= Boulogne sur Seine. France, lectotype MNHN AM 4500-1 (female 5.5 mm).

336

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

seta. Pereopod 3: dactyl base weakly
pinched, nail with slight prominence at
middle on posterior margin. Pereopod 4
shorter than pereopod 3, basis weakly ar-
cuate, dactyl base pinched, nail with sharp
dentition posteriorly. Pereopod 5 longer than
pereopod 4, basis narrow with a submar-
ginal spine at posterodistal comer, lacking
posterodistal lobe. Pereopod 6 similar to
but much shorter than pereopod 5, dactyl
nail almost straight. Pereopod 7 slightly lon-
ger than pereopod 6, basis expanded pos-
teriorly, as deep as wide.

Coxal gill of gnathopod 2 as long as basis,
distal margin with crenulations. Coxal gills
of pereopods 3-5 small, constricted at mid-
dle. Coxal gill of pereopod 6 longer than
basis, reverse L-shaped. Brood plates con-
fined to pereopods 3-5, small and slender
with 2-3 simple setae apically.

Abdominal side plate 1 rounded postero-
ventrally. Plates 2 and 3 bluntly pointed
posteroventrally, posterior margins weakly
spinulose. Pleopod 1: peduncle weakly ar-
cuate, with 3 plumose setae on outer mar-
gin, with 2 retinaculae; outer ramus
7 -articulated; inner ramus shorter than half
of outer ramus, 2 -articulated. Pleopod 2 a
little shorter than pleopod 1 , peduncle outer
margin with a plumose seta, outer ramus
6 -articulated, inner ramus 1 -articulated with
apical setae. Pleopod 3 small and slender,
0.4 times as long as peduncle of pleopod 2,
with 2 spinules subapically, lacking rami.

Uropod 1 : peduncle with marginal spines
on both edges, distolateral spine strong,
closely set to distomarginal spine; rami sub-
equal in length and shorter than peduncle,
with elongate apical spines; outer ramus
marginally bare; inner ramus with 3 mar-
ginal spines. Uropod 2: peduncle with mar-
ginal spines distally; rami subequal to each
other and to peduncle in length, with elon-
gate apical spines; outer ramus marginally
bare; inner ramus with spines proximally.
Uropod 3 very small, not reaching tip of
telson, subtriangular in shape; peduncle
broad at base with a strong spine ventrally;

ramus small, conical, partially fused to pe-
duncle, with a spinule at tip. Telson broad,
apically rounded with apical and 3-4 lateral
spines.

Male. —Not known.

Variations. —Eye in the Israeli material is
somewhat larger than that of the lectotype.
Number of flagellar articles ranges from 4
to 6 on antenna 1 and from 8 to 9 on antenna
2 in the material examined, tending to in-
crease with body size. Marginal setae on
propod anterior margin of gnathopod 2 also
show variation in number: Seychelles and
Hawaiian Islands material lacks them, Is-
raeli material with 1-2 setae, Azores ma-
terial with 0-2. Swiss material (Menzel 1911,
as Orchestia senni) also has one seta. Inner
rami of pleopods 1 and 2 fluctuate in the
number of articles: from 1 to 4 (mostly 2)
articles in pleopod 1 ; 1 , exceptionally 3 (in
Azores material, 7 mm body length) articles
in pleopod 2. Only this larger specimen from
the Azores has reduced ramus in pleopod
3, otherwise lacking them. For each speci-
men examined, article number of inner ra-
mus of pleopod 1 is higher than, rarely same
as that of pleopod 2. This range and pattern
of variation in pleopods is in accord with
those in the other geographical populations:
Basel, Switzerland (Menzel 1911); Urbana,
Illinois, U.S.A. (Medcof 1940); Finland
(Palmen 1949) and the Canary Islands (An-
dersson 1962). Dactyl nail of pereopod 3
has usually very weak prominence at the
posterior middle. The Azores material (6.0
mm) has, however, distinct prominence (Fig.
2G) which is rather similar to the dentition
of pereopod 4.

Discussion

Most of the variations mentioned above
occur within a given population rather than
between geographically separated popula-
tions. Thus it is not possible to recognize
more than one species amongst the material
examined. All the material lacks a brood
plate on female gnathopod 2, and has dac-

VOLUME 106, NUMBER 2

337

tylar dentition on pereopod 4 (rarely also
on pereopod 3). Chevreux (1901) men-
tioned the dactylar dentition on pereopod
5, not on pereopod 4, in his Seychelles ma-
terial. At the same time, in our specimens,
the inner ramus of pleopod 1 has more ar-
ticles than that of pleopod 2, and never less,
as by Chevreux (1901). Thus it is possible
that Chevreux treated an exceptional spec-
imen or more probably some confusion oc-
curred during the preparation of his mate-
rial.

The present species is most closely related
to Talitroides topitotum (see Bousfield 1984).
Friend & Lam (1985) listed several discrim-
inating characters between T. alluaudi and
T. topitotum. In addition, the following ones
are to be mentioned: T. alluaudi has gnatho-
pod 2 without brood plate, strongly chitin-
ized mandible, maxilla 2 with distally
pointed inner plate, and pereopod 4 with
dactylar dentition. These features are rather
peculiar among landhoppers. Talitroides al-
luaudi also resembles Arcitalitrus sylvaticus,
a non-cuspidactylate landhopper, in having
simple gnathopod 1 , weakly setose anterior
margin in gnathopod 2, reduced and plu-
mose-setose peduncle of pleopods, similar
spination in uropods 1 and 2, etc. But the
latter is distinguished from the former by
the possession of a strongly arcuate outer
plate and less setose maxilliped. It is inter-
esting that T. alluaudi has a weakly arcuate
inner plate.

The distribution of T. alluaudi in conti-
nental areas is believed to be due to syn-
anthropic dispersal (Friend & Lam 1985),
and the occurrence is thought to be limited
by the humidity (Andersson 1962). Thus it
may be astonishing that T. alluaudi has been
discovered in an arid part of Israel with pre-
cipitation as little as 200 mm annually. So
far in Israel the present species is known
solely from the bamboo stands growing on
a small "island" in the middle of a small
pool in the University campus at Beersheba.
The bamboo, now commonly utilized for
gardening in Israel, is an exotic plant for

this country, most of which were imported
from West Europe. Thus it is highly prob-
able that the population of T. alluaudi in
Beersheba was introduced accidentally from
European countries with the bamboo and
has survived on this small "island," where
a humid microclimate may have prevailed.

Acknowledgments

We are grateful to D. Defaye in Museum
National d'Histoire Naturelle, Paris for the
loan of the type material treated here.
Thanks to J. Aoki in Yokohama National
University who provided us with compar-
ative material. Thanks also to K. Iwatsuki
of The University of Tokyo who helped lo-
cate the type material.

Literature Cited

Andersson, A. 1962. On a collection of Amphipoda
of the family Talitridae from the Canar>' Is-
lands. -Arkiv for Zoologi 15(1 1):2 11-218.

Bousfield, E. L. 1984. Recent advances in the sys-
tematics and biogeography of landhoppers (Am-
phipoda: Talitridae) of the Indo-Pacific re-
gion. -Pp. 171-210 in F. J. Radovsky, P. H.
Raven & S. H. Sohmer. ed., Biogeography of
the tropical Pacific, Bishop Museum Special
Publication 72.

Chevreux, E. 1896. Recherches zoologiques dans les
serres du Museum de Paris IV. — Sur un Am-
phipode terrestre exotique. Talitrus Alluaudi now
sp., acclimate dans les serres du Jardin des
Plantes de Paris. — Feuille des Jeunes Natural-
istes 26:112-113.

. 1901. Mission scientifique de M.Ch. Alluaud

aux lies Sechelles. Crustaces Amphipodes.—
Memoires de la Societe Zoologique de France
14:388-438.

. & L. Fage. 1925. Faune de France 9. Am-
phipodes. Paul Lechevalier. Paris. 486 pp.

Friend, J. A., & P. K. S. Lam. 1985. Occurrence of
the terrestrial amphipod Talitroides topitotum
(Burt) on Hong Kong Island.— Acta Zootaxon-
omica Sinica 10(l):27-33.

. & A. M. M. Richardson. 1986. Biology of

terrestrial amphipods. — Annual Review of En-
tomology 31:25^8.

Medcof. J. C. 1940. Variations in the pleopod struc-
ture of the terrestrial amphipod Talitrus alluau-
di Chevreux. -Lloydia 3(l):79-80.

Menzel, R. 1911. Exotische Crustaceen im botan-

338

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ischen Garten zu Bazel. —Revue Suisse de Zool-

ogie 19:433-444.

Palmen, E. 1949. Talitroides alluaudi (Chevreux)
(Amphipoda, Talitridae) in Finnland gefun-
den.— Archivum Societatis Zoologicae Botani-
cae Fennicae 'Vanamo' 2:61-64.

Shoemaker, C. R. 1936. The occurrence of the ter-
restrial amphipods, Talitrus alluaudi and Tali-
trus sylvaticus, in the United States.— Journal of
the Washington Academy of Sciences 26(2): 60-
64.

(HM) Department of Biology, Ibaraki
University, Mito 310, Japan; (RO) Depart-
ment of Ecology, Systematics and Evolu-
tion, The Hebrew University of Jerusalem,
Givat Ram, Jerusalem 91904, Israel.

PROC. BIOL. SOC. WASH.

106(2), 1993, pp. 339-345

A DESCRIPTION OF A NEW SPECIES OF

MACROBRACHIUM FROM PERU, AND DISTRIBUTIONAL

RECORDS FOR MACROBRACHIUM BRASILIENSE (HELLER)

(CRUSTACEA: DECAPODA: PALAEMONIDAE)

Guido A. Pereira S.

Abstract. —A systematic study is made on a collection of freshwater shrimps
(Palaemonidae) from northern Peru collected by Dr. R. W. McDiarmid during
the years 1977-1980. A new species is described Macrobrachium depressi-
manum, and Macrobrachium brasiliense (Heller) is reported. The diversity of
palaemonids in the Amazon region is discussed.

Resiimen.—Se realiza un estudio sistematico sobre una coleccion de cama-
rones de agua dulce del norte del Peru, colectados por el Dr. R. W. McDiarmid
durante los afios de 1 977 y 1 980. Se describe una nueva especie, Macrobrachium
depressimanum, y se reporta la especie Macrobrachium brasiliense (Heller). Se
discute acerca de la alta diversidad de camarones palaemonidos en la region
del Amazonas.

The present work is based on collections
made primarily by Dr. R. W. McDiarmid
in northern Peru during 1977 and 1980. The
specimens were collected during field re-
search in ethnobiological anthropology,
conducted in the region of the Alto Mara-
iion River by scientists from the University
of California. The collection has many sam-
ples of two species of the freshwater shrimp
genus Macrobrachium Bate, 1868 and sev-
eral species of crabs of the families Tricho-
dactylidae and Pseudothelphusidae. Only
the shrimps are reported in this paper. De-
scriptions are based on adult males. Total
length (TL) is taken from the tip of the ros-
trum to the posterior border of telson. Car-
apace length (CL) is taken from the ocular
margin to the posterior border of carapace.
Only partial synonymy is given for M. brasi-
liense (Heller, 1862). All material is depos-
ited in the National Museum of Natural
History, Smithsonian Institution, Washing-
ton D.C., U.S.A. (USNM).

Macrobrachium depressimanum,
new species
(Figs. 1, 2)

Type material— Holotype, 1 5, USNM
230032, 35.5. mm (TL), 6.2 mm (CL), Peru,
Amazonas Department, Santiago River, La
Poza, 4°01'S, 77°47'W, 26 Jan 1980, coll.
R. W. McDiarmid. Paratypes, 2 9(1 ovig.),
USNM 230030; 4 9 (2 ovig.), 14 <5, USNM
230031 (locality and date, same as for ho-
lotype).

Additional material examined. — \0^ 9,
133 (5, from the following localities in Peru:
Amazonas Department, Santiago River, La
Poza, 4°01'S, 1T47'W; Caterpiza River,
3°55'S, 77°42'W; Cenepa River near Huam-
pami, 4°28'S, 78°10'W; Santiago River at
Pinglo, at confluence between Santiago Riv-
er and Maraiion River, 4°26'S, 77°39'W;
Marafion River across from Santa Maria de
Nieva, and at confluence of Nieva River
with Marafion River, 4°35'S, 77°77'W;

340

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1 mm

Fig. 1 . Macrobrachium depressimanum, new species. A, carapace and cephalic appendages; B, carapace; C,
posterior region of telson; D, telson; E, appendix masculina and appendix interna; F, first pereiopod; G, second
pereiopod; H, third pereiopod; I, fifth pereiopod.

Ucayali, Tahuallo River, Pucalpa, 8°23'S,
74°32'W; Ucayali, Tahuallo River, Pucalpa,
Neshuya, 8°23'S, 74°32'W; Madre de Dios,
Tambopata Natural Reservoir, La Torre
River, 11°58'S, 69°11'W.
Diagnosis — KdulX males with second pe-

reiopod smooth; ratio palm length/finger
length about 0.6; palm flattened anterodor-
sally (Fig. IG).

Description. —Nostrum nearly straight,
slightly arched over eyes, reaching (or slight-
ly overreaching) distal border of scaphoce-

VOLUME 106, NUMBER 2

341

H^S^^k.

MV v-*-^i

SBl^^^w

^Pwi

\^yf jiff'

^p^^^EI

TjpB^N^^J^^^^B

S! a K u 5 u

! ' /- -

Fig. 2. Detail of appendix masculina. A, Macrobrachium depressimanum, new species, anterior view; B,
Macrobrachium jelskii (Miers, 1877), lateral view. Same scale for A and B.

342

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

rite with apex, tip pointing upwards. Upper
border bearing 1 1 (9-1 1 in paratypes) teeth,
2 proximal teeth more spaced behind the
orbit. Lower margin bearing 5 (4-6 in para-
types) teeth (Fig. lA, B). Carapace smooth.
Scaphocerite 2.5 times longer than wide.
Abdomen smooth, posteroventral angle of
fifth pleuron acute; sixth abdominal seg-
ment 1.5 times length of fifth, same length
as telson. Telson with 2 pairs of dorsal spines
situated at % and ^/^ its length from base.
Posterior margin, tapering abruptly to me-
dian apex bearing 2 pairs of lateral spines.
Inner pair of spines overreaching median
apex. Four plumose setae present on pos-
terior margin between inner spines (Fig. 1 C,
D). First pereiopods slenderer than second
pair, reaching scaphocerite with tips of fin-
gers; palm cylindrical, 0.9 times length of
dactyl. Carpus 3.1 times length of palm, 1.1
times that of merus (Fig. IF). Second pair
of pereiopods equal, smooth, reaching an-
terior border of scaphocerite with distal car-
pus. Fingers thin, closing over entire length
of chela, without conspicuous teeth. Palm
flattened anterodorsally, 2.1 times longer
than high, 0.6 times length of dactyl. Carpus
1.5 times length of palm, 0.9 times that of
merus (Fig. IG). Third pair of pereiopods
failing to reach distal border of scaphocer-
ite. Propodus with longitudinal row of 4
spines on inner margin, 1.3 times length of
dactyl, 1.7 times that of carpus (Fig. IH).
Fifth pair of pereiopods reaching distal bor-
der of scaphocerite with tip of dactyl. Prop-
odus with longitudinal row of 6 spines on
inner margin, 2 times length of dactyl, 1.7
times length of carpus (Fig. II). Appendix
masculina 2.1 times longer than appendix
interna (Fig. IE), with acute apex, and 18
short, smooth spines.

Fecundity. —One ovigerous female, TL =
38 mm, CL = 7.1 mm, with 23 eggs, 1.2
mm largest diameter.

Distribution.— T)r. Celio Magalhaes from
Instituto Nacional de Pesquisas da Ama-
zonia (INPA), Brazil, has informed me that

there are specimens of this species in the
invertebrate collection of his Institute, from
the following localities in Brazil: Amazonas
State, Tefe River, Solimoes, 3°35'S,
64°47'W; Solimoes River, Catalao coast,
near Manaus, 3°8'S, 60°1'W; Acre State,
Tarauaca River, Tarauaca, 6°42'S, 69°48'W;
Rondonia State, Madeira River, fall of Teo-
tonio, 3°22'S, 58°45'W; Mamore River, near
Guajara-Mirim, 10°23'S, 65°23'W; Madei-
ra River, near Guajara-Mirim, 10°23'S,
65°23'W. Based on these data the species is
distributed over the upper Amazon Basin,
from Peru to northeastern Brazil.

Etymology. —The specific name is de-
rived from a composite of the Latin words
depressus meaning dorsoventrally flattened
and manus meaning hand. It stresses the
typical shape of the anterior part of the palm
in this species.

Remarks.— The present species is mor-
phologically related to Macrobrachium jel-
skii (Miers, 1877). M. depressimanum has
more teeth on the dorsal border of the ros-
trum. The formula for M. jelskii is 6-9 (1
post-ocular) whereas it is 10-12 (2 post-oc-
ular) in M. depressimanum. The relative
proportion of segments in the second pe-
reiopods is very distinctive, the ratio be-
tween palm length/dactyl length is 0.6 in M.
depressimanum whereas it is 1.5 in M. jel-
skii. The anterodorsally flattened shape of
the palm is a typical feature of M. depres-
simanum.

The shape of the appendix masculina is
not of common usage in the taxonomy of
Macrobrachium. However, the appendix
masculina of M. depressimanum is very dis-
tinctive. It is 2. 1 times longer than the ap-
pendix interna, whereas in M. jelskii it is
about 1.5 times longer. The appendix in-
terna of M. depressimanum is also more
slender compared to that of M. jelskii (Fig.
2). The two species are allopatric but there
is a possibility of some overlap since M.
jelskii was reported by Holthuis (1966) as
occurring in north-east Brazil.

VOLUME 106, NUMBER 2

343

Macrobrachium brasiliense (Heller, 1862)

Palaemon brasiliensis. —Heller, 1862:419.
Macrobrachium brasiliense. — Holthuis,

1952:79, pi. 19. -Holthuis, 1959:93.-

Rodriguez, 1982:382.

Material examined. — 63 9, 114 5, 1 1 2 ju-
veniles, from the following localities in Peru,
Amazonas Department: Santiago River, La
Poza, 4°01'S, 77°47'W; Castillo Creek
(mouth enters Santiago River at 1.5 km
south of La Poza), 4°02'S, 77°47'W; at Ping-
lo, confluence at Santiago River and Mara-
non River, 4°26'S, 77°39'W; way to Galilea,
creek in the forest, 4°15'S, 77°49'W; Galilea,
4°01'S, 77°49'W; Yutapis River, Shiringa,
4°01'S, 77°51'W; Caterpiza River, 3°55'S,
77°43'W; Cenepa River, 3°55'S, 77°43'W.

Diagnosis. —Adult males with short spines
on carapace, specially on anterolateral bor-
ders. The larger of the second pair of pereio-
pods has a finger/palm length ratio of 2.0-
2.3. Fixed finger bears a row of 4 to 11
tubercles along inner border.

Discussion

Since the revision of the Palaemonidae
by Holthuis (1952), several new species of
palaemonid shrimps have been described
from the Amazon Region (Tiefenbacher
1978; Ramos-Porto 1979; Kensley & Walk-
er 1982; Rodriguez 1982; Pereira 1985,
1986, 1991). The latter authors report spe-
cies that, judged by the small brood and
somewhat large size of the eggs, can be con-
sidered to have abbreviated larval devel-
opment and a strictly land-locked fresh-
water habitat (Sollaud 1923a, 1923b;
Shokita 1973, 1977, 1979; Gamba 1984;
Magalhaes 1985, 1986, 1988; Chong &
Khoo 1987a, 1987b). The Amazon Region
has the largest number of strictly freshwater
species (24) of palaemonids in the world
(Pereira 1989). This agrees with the tradi-
tional view of taxonomists and biogeogra-

phers that consider the Amazon Region as
an area with high endemism and diversity,
probably as a result of both environmental
heterogeneity and the long and complex pa-
leogeographic history of the basin (Prance
1982). The upper Amazon Basin (east and
northeast portion) was formed most re-
cently during Pliocene times, whereas the
middle basin was formed during the Cre-
taceous (Putzer 1984, Rodriguez & Pereira
1992). The complexity and long history of
the basin led to the emergence of strictly
freshwater palaemonids in at least four gen-
era, Macrobrachium, Palaemonetes Heller,
1869, Euryrhynchus Miers, 1877, and Pseu-
dopalaemon Sollaud, 1911, according to
Magalhaes & Walker (1988), and Pereira
(1989). The latter two genera are endemic
to South America, and all four are strictly
freshwater with abbreviated larval devel-
opment.

An interesting argument arises when con-
sidering the phylogenetic history of the fam-
ily (Pereira 1989). There is evidence of an
early freshwater origin of this group because
the closest relatives of the primitive genus
Euryrhynchus occur in West African fresh-
waters (Powell 1976). This raises questions
about the tempo and mode of evolution of
the family and their invasion into the ma-
rine and freshwater habitat (Pereira 1989).
Magalhaes & Walker (1988) considered ab-
breviated development as a derived con-
dition. They concluded that there has been
massive intergeneric convergence toward
this type of life cycle in the Palaemonids
mainly because of selective pressure to live
in plankton-poor waters. However, a dif-
ferent point of view arises if the abbreviated
development is considered the primitive
condition. It could be argued that the life
cycle with complete larval development, and
acquired independently only by Macrobra-
chium and Palaemonetes, is the derived
condition. The complete development
eventually permitted these freshwater
shrimps to be successful in colonizing more

344

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

productive environments like estuaries and
mixohaline waters.

Acknowledgments

I wish to thank Dr. M. Reaka, Dr. R. W.
McDiarmid, and Dr. C. Magalhaes for re-
viewing the manuscript. Special thanks to
Dr. R. B. Manning for his advice and pro-
viding working space at the Smithsonian
Institution, Division of Crustacea, during
1984-1988. Thanks to Dr. C. Magalhaes
from INPA, Brazil, and Lie. H. Ortega from
Universidad de San Marcos, Peru, for gen-
erously providing additional data on the
species. Sheila Pauls kindly helped me in
the final editing of this manuscript. Support
for present research was provided by Con-
sejo de Desarrollo Cientifico y Humanisti-
co, Universidad Central de Venezuela.

Literature Cited

Bate, C. S. 1868. On a new genus, with four new
species, of freshwater prawns.— Proceedings of
the Zoological Society of London 1868:363-368.

Chong, S. S. C, & H. W. Khoo. 1987a. Abbreviated
larval development of the freshwater prawn,
Macrobrachium pilimanus (De Man, 1879),
(Decapoda, Palaemonidae), reared in the labo-
ratory.— Journal of Natural History 21:763-774.

. 1987b. The abbreviated larval development

of the freshwater prawn, Macrobrachium ma-
layanum (Roux, 1934), (Decapoda, Palaemon-
idae), reared in the laboratory.— Crustaceana 53:
29-42.

Gamba, A. L. 1984. Different egg-associated and lar-
val development characteristics of Macrobra-
chium jelskii and Macrobrachium amazonicum
(Arthropoda: Crustacea) in a Venezuelan con-
tinental lagoon.— International Journal of In-
vertebrate Reproduction and Biology 7: 1 35-142.

Heller, C. 1862. Beitrage zur naheren Kentniss der
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. 1869. Zur naheren kenntniss der in den siis-

sen gewassem des siidlichen Europa vorkom-
mendem Meerescrustaceen.— Zeitschrift wis-
senschaffliche Zoologie 19:156-162.

Holthuis, L. B. 1952. A general revision of the Pa-
laemonidae (Crustacea, Decapoda, Natantia) of
the Americas. II. The subfamily Palaemoni-
dae.— Occasional Papers, Allan Hancock Foun-
dation Publications, 12, 396 pp.

. 1959. The Crustacea Decapoda of Suriname

(Dutch Guiana).— Zoologische Verhandlingen
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. 1966. A collection of freshwater prawns

(Crustacea, Decapoda, Palaemonidae) from
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lier.— Bulletin Institute Royale Sciences Natu-
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Kensley, B., & I. Walker. 1982. Palaemonid shrimps
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Magalhaes, C. 1985. Desenvolvimento larval obtido
em laboratorio de palaemonideos da Regiao
Amazonica. I. Macrobrachium amazonicum
(Heller, 1862) (Crustacea, Decapoda).— Ama-
zoniana 9:247-274.

. 1986. The larval development of the palae-
monid shrimps from the Amazon region reared
in the laboratory. IV. Abbreviated development
of Palaemonetes ivonicus Holthuis, 1950 (Crus-
tacea: Decapoda).— Amazoniana 10:63-78.

. 1988. The larval development of Palaemonid

shrimps from the Amazon Region reared in the
laboratory. II. Extremely abbreviated larval de-
velopment in species of the genus Euryrhynchus
Miers, 1877 (Crustacea, Decapoda, Euryrhyn-
chinae).— Crustaceana 55:39-52.

, & I. Walker. 1988. Larval development and

ecological distribution of central amazonian Pa-
laemonid shrimps (Decapoda, Caridea).— Crus-
taceana 55:279-292.

Miers, E. J. 1877. On a collection of Crustacea, De-
capoda and Isopoda, chiefly from South Amer-
ica, with descriptions of new genera and spe-
cies.— Proceedings of the Zoological Society of
London 1877:653-679.

Pereira, G. 1985. Freshwater shrimps from Vene-
zuela III: Macrobrachium quelchi (De Man) and
Euryrhynchus pemoni n. sp. (Crustacea, Decap-
oda, Palaemonidae) from La Gran Sabana.—
Proceedings of the Biological Society of Wash-
ington 98:615-621.

. 1986. Freshwater shrimps from Venezuela I:

seven new species of Palaemoninae.— Proceed-
ings of the Biological Society of Washington 99:
191-213.

. 1989. Cladistic, taxonomy, biogeography and

the evolutionary history of the shrimp family
Palaemonidae (Crustacea, Decapoda, Caridea).
Unpublished Ph.D. Dissertation, University of
Maryland, College Park, 417 pp.

. 1991. Camarones de agua dulce de Venezuela

II: neuvas adiciones en las familias Atydae y
Palaemonidae (Crustacea, Decapoda, Carid-
ea).— Acta Biologica Venezuelica 13:75-88.

Prance, G. T. (ed.). 1982. Biological Diversification

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345

in the Tropics. Columbia University Press, New
York, xvi + 714 pp.

Powell, C. B. 1 976. Two new freshwater shrimps from
West Africa: the first euryrhynchinids (Decap-
oda, Palaemonidae) reported from the old
world. — Revue de Zoologie Africaine 91:649-
674.

Putzer, H. 1 984. The geological evolution of the Am-
azon Basin and its mineral resources.: Pp. 14-
46 in H. Sioli, ed., The Amazon. Limnology and
landscape ecology of a mighty tropical river and
its basin. Dr. W. Junk Publisher, The Nether-
lands, ix + 763 pp.

Ramos-Porto, M. 1979. Pseudopalaemon amazo-
nensis, especie nova de camarao de Bacia Ama-
zonica (Crustacea, Decapoda, Palaemonidae).—
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cia, 31a Reuniao Anual, Resumes, Suplemento
de Ciencia e Cultura 31:693.

Rodriguez, G. 1 982. Fresh-water shrimps (Crustacea,
Decapoda, Natantia) of the Orinoco Basin and
the Venezuelan Guayana.— Journal of Crusta-
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, & G. Pereira. 1992. New species, cladistic

relationships, and biogeography of the genus
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tacean Biology 12:298-311.

Shokita, S. 1973. Abbreviated larval development of
the freshwater prawn, Macrobrachium shokitai
Fujino et Baba (Decapoda, Palaemonidae) from

Iriomote Island of the Ryukyus.— Annotationes
Zoologicae Japonenses 46:1 1 1-126.

. 1977. Abbreviated metamorphosis of land-
locked freshwater prawn, Macrobrachium as-
perulum (Von Martens) from Taiwan.— Anno-
tationes Zoologicae Japonenses 50:1 10-122.

. 1979. Early life history of the genus Mac-
robrachium (Decapoda, Palaemonidae). — Uni-
versity of the Ryukyus 1:9-17.

SoUaud, E. 1911. Pseudopalaemon bouvieri, nouveau
genre, nouvelle espece, de la famille des Palae-
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turelle, Paris 17:12-16.

. 1923a. Le developpement larvaire des Pa-

laemoninae. I. Partie descriptive. La conden-
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510-603.

. 1923b. Recherches sur I'embryogenie des

Crustaces Decapodes de la sous-famille des Pa-
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et de la Belgique, supplement 5, 234 pp.

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Institute de Zoologia Tropical, Univer-
sidad Central de Venezuela, Aptdo. 47058,
Caracas 1041 -A, Venezuela.

PROC. BIOL. SOC. WASH.

106(2), 1993, pp. 346-352

A NEW SPECIES OF CRAYFISH (DECAPODA:

CAMBARIDAE) BELONGING TO THE GENUS CAMBARUS,

SUBGENUS HIATICAMBARUS, FROM THE UPPER

ELK RIVER DRAINAGE OF WEST VIRGINIA

Raymond F. Jezerinac and G. Whitney Stocker

Abstract. —A new species of crayfish, Cambarus (Hiaticambarus) elkensis, is
described from the upper Elk, Holly, and Birch rivers of West Virginia. The
species is most closely related to C. (H.) chasmodactylus but differs from it in
having shorter fingers, a much narrower gape between the fingers, and moderate
impressions at the base of the fixed finger, especially on the ventral surface.

While conducting a crayfish survey of the
state of West Virginia in 1988, we captured
what we thought to be C. (//.) chasmodac-
tylus James (1966) from the upper Elk River
drainage. We reported this (Jezerinac &
Stocker 1989:3) as a new drainage record
for the species. Additional collections were
made in 1989 from the upper Elk basin, the
Elk River and its tributaries below Sutton
Lake, as well as in surrounding watersheds,
notably the Greenbriar, Gauley, Cherry, and
Cranberry rivers. Representatives of the
species were caught only in the upper Elk
River drainage. With the additional mate-
rial, we noted that these crayfish differ in
several respects from C. (//.) chasmodac-
tylus and are described herein as members
of a new species.

Cambarus {Hiaticambarus) elkensis,

new species

Fig. 1, Table 1

Cambarus bartonii veteranus. —Faxon,
1914:389 [in part, Elk River, Cogar's
Mills, West Virginia].

Cambarus (Hiaticambarus) chasmodacty-
lus.— J QzennsLC & Stocker, 1989:3 [in
part].

Diagnosis.— Body pigmented, eyes well
developed. Carapace subovate, dorsoven-
trally flattened. Rostrum with weakly con-

vergent, slightly concave, thickened mar-
gins, lacking spines or tubercles and
terminating in upturned corneous tubercle.
Areola 3.5 to 5.5 (X = 4.6) times as long as
wide and comprising 35.1 to 38.1% (X =
36.6%) of entire length of carapace, and
bearing 5 to 7 punctations across narrowest
part. Cervical spine absent or represented
by small tubercle. Suborbital angle obtuse
to obsolete. Postorbital ridges terminating
cephalically in sharp spine or tubercle.
Branchiostegal spine very small. Antennal
scale 2.5 times longer than broad, with me-
sial and lateral margins subparallel near and
at midlength; basiopodite with very small
spine; ischiopodite lacking spine. Epistome
subtriangular, zygoma with about 120° arch.
Chela smooth and bearing one row of very
indistinct tubercles along mesial margin of
palm; lateral margin of fixed finger smooth;
both fingers with very poorly defined dor-
somedian longitudinal ridges; fixed finger
moderately impressed at base especially on
ventral surface; dactyl 1.8 to 2.3 (X = 2.0)
times longer than mesial margin of palm;
palm length 28.4 to 32.3% {X = 30.8%) of
chela length; elongated setae at base of fixed
finger very sparse or absent (best seen on
young specimens); gape of fingers subequal
to width of dactyl in first form males, less
so in second form males, and almost non-
existent in females. Mesial margin of carpus

VOLUME 106, NUMBER 2

347

of chela with large spiniform tubercle and
smaller conical one proximally; ventral sur-
face with conical knob on distal articular
rim. Ventrolateral ridge of merus with 2 to
4 (X = 2.5) spines; ventromesial one with
6 to 12 (X= 9) spines. Ischium of only third
pereiopod with simple hook not reaching
tubercle on corresponding basis. Boss on
ischium of fourth pereiopod prominent.
First pleopod of Form I male (Fig. IB, F)
with short terminal elements; corneous cen-
tral projection truncate distally, recurved at
greater than 90° to main shaft of appendage,
with subapical notch; mesial process in-
flated, tapering, rounded to acute distally,
directed caudolaterally at angle slightly
greater than 90° to main shaft of appendage.
Female with annulus ventralis shallowly
embedded in sternum, asymmetrical, and
subrhomboidal. (Additional morphometric
and meristic data, such as simple descrip-
tive statistics, ratios, and regression analy-
sis, may be obtained from the authors or
the library at The Ohio State University at
Newark.)

Holotypic male, Form /. — Cephalothorax
(Fig. lA, J) subovate in cross section, dor-
soventrally compressed. Abdomen narrow-
er than thorax (21.0 mm and 17.4 mm);
greatest width of carapace distinctly greater
than height at caudodorsal margin of cer-
vical groove (21.0 mm and 15.2 mm). Are-
ola 4.5 times as long as broad with 5 to 7
crowded punctations across narrowest part;
length of areola 37.5% of total length of car-
apace. Rostrum with thickened, weakly
convergent, elevated, concave margins de-
void of spines or tubercles; dorsal surface
of rostrum slightly concave with many
punctations, rather sparse punctations on
and at base of indistinctly delineated acu-
men. Subrostral ridges well developed and
evident in dorsal aspect to base of acumen.
Suborbital angle obtuse. Postorbital ridge
moderately prominent, grooved dorsolater-
ally, and terminating cephalically in acute
corneous tubercule. Branchiastegal spine
represented by a spiniform tubercle. Cer-

vical spine reduced to small rounded tu-
bercle on left side, absent on right. Carapace
densely punctate dorsally except in gastric
region, distinctly sculptured over attach-
ment of mandibular muscle; lateral surface
with many small granules in branchiostegal
region; larger granules in mandibular and
ventral half of hepatic region.

Abdomen slightly shorter than carapace
(38.6 mm and 40.0 mm respectively); pleu-
ra rounded to subtruncate ventrally with an-
gular caudoventral extremities on third
through fifth segments. Cephalic section of
telson with 2 spines on each caudolateral
comer, mesial spines movable; borders of
caudal section evenly rounded. Proximal
podomere of uropod with both lobes ter-
minating in corneous-tipped blunt spine;
mesial ramus with low submedian dorsal
keel ending in small, blunt, premarginal
spine not extending beyond distal edge of
ramus.

Cephalomedian lobe of epistome (Fig. II)
broadly rounded with small cephalomedian
projection, margin somewhat thickened and
elevated ventrally; main body with distinct
median groove and paired slitlike grooves
immediately cephalic to arched epistomal
zygoma. Ventral surface of proximal podo-
mere of antennular peduncle with small
spine at base of distal third. Antennal pe-
duncle with strong lateral spine on basis,
remaining podomeres lacking spines. An-
tennal scale (Fig. IG) about 2.5 times as
long as broad with mesial and lateral mar-
gins subparallel for some distance proximal
and distal to midlength; strong distolateral
spine reaching beyond midlength of ulti-
mate podomere of antennular peduncle.
Ventral surface of ischium of third maxil-
liped with broad, longitudinal band of long
setae laterally and with submarginal lateral
row of smaller plumose ones, few additional
short plumose setae in area between; mesial
margin with 2 1 denticles.

Right chela (detached) (Fig. IL) 2.4 times
as long as broad, mesial margin of palm
30.6% of chela length; dactyl 2. 1 times palm

348

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

length; mesial margin of palm with one row
of 7 indistinct tubercles; remainder of palm
with widely spaced large, deep punctations
on mesial region, more crowded on lateral
region, lateral surface rounded. Gape be-
tween fingers subequal to dactyl width, and
proximal half of opposable surface of fixed
finger without tufts of setae; both fingers
with poorly defined median longitudinal
ridge on dorsal surface, absent on ventral
surface, both with conspicuous deep punc-
tations; opposable margin of fixed finger with
row of 1 1 small (seventh enlarged) rounded
corneous tubercles, single row of minute
denticles present slightly ventral to tuber-
cular row along distal fourth of finger; op-
posable margin of dactyl with row of 12
tubercles, minute denticles forming single
row below level of tubercles along distal
third. Lateral surface of fixed finger and me-
sial surface of dactyl non-costate, punctate.

Carpus of cheliped (Fig. IL) longer than
broad with deep dorsal furrow flanked by
sparse punctations mesially and laterally;
mesial surface with 1 spiniform tubercle and
more proximally situated smaller conical
one; ventral surface very sparsely punctate
and bearing 1 tubercle on distal articular
rim. Merus with 1 premarginal tubercle dor-
sally; ventrolateral ridge with 2, ventrome-
sial ridge with 9 tubercles. Ventromesial
margin of ischium with 2 very small tuber-
cles.

Hook on ischium of third pereiopod only
(Fig. IK), hook overreaching basioischial
articulation and opposed by small tubercle
on basis. Coxa of fourth pereiopod with
prominent caudomesial boss ventrally dis-
posed, and somewhat flattened caudally.
Boss on coxa of fifth pereiopod vestigial.
First pleopods (Fig. IB, C) reaching coxae
of third pereiopods, symmetrical, with very
small gap between bases (Fig. ID). (See "Di-
agnosis" for descriptions of the pleopods
and Fig. IB, F.)

Allotypic y^ma/^.— Excluding secondary
sexual characteristics, differing from holo-
type in following respects: areola 5.0 times

as long as broad and constituting 36.6% of
total length of carapace; very small cervical
tubercle present on both sides; mesial mar-
gin of palm of right chela 30.9% of chela
length; gape between fingers about V4 width
of dactyl; few plumose setae present at base
of fixed finger; no enlarged tubercle on op-
posable surface of fixed finger; 1 (right) and
1 1 (left) tubercles on opposable margin of
dactyl; 2 dorsal premarginal tubercles on
merus; ventrolateral ridge of merus with 3
(right) and 2 (left) tubercles; ventromesial
ridge of merus with 1 1 (right) and 1 2 (left)
tubercles.

Annulus ventralis subrhomboidal (Fig.
IH), wider than long, slightly movable, with
caudal wall weakly developed. Postannular
sclerite about half as wide and about ¥3 as
long as annulus. First pleopods reaching
midlength of annulus when abdomen flexed.

Morphotypic male. Form //.—Differing
from holotype in following respects: areola
4.2 times as long as broad; areola length
36.2% of carapace length; cervical tubercle
absent on both sides; spine on ventral sur-
face of proximal podomere of antennuar pe-
duncle near distal margin; spine on right
antennal scale damaged, antennal scale 2.3
times longer than broad; right chela regen-
erated; left chela 2.3 times as long as broad;
mesial surface of palm occupying 29.7% of
chela length; dactyl length 2.2 times palm
length; gape about % width of dactyl; setae
at base of fixed finger; ventrodistal margin
of carpus with 2 tubercles; merus with 2
premarginal tubercles dorsally, 2 large and
1 small (left) and 4 (right) tubercles on ven-
trolateral ridge of merus; ventromesial ridge
with 9 (right) and 7 (left) tubercles; ven-
tromesial margin of ischium with 4 (right)
and 3 (left) tubercles; hook on ischium of
third pereiopod much reduced, not over-
reaching basioischial articulation and op-
posed by small tubercle on basis; first pleo-
pod (Fig. IC, E) with juvenile suture on shaft,
central projection inffated.

Color notes.— ^2isic color of dorsal sur-
faces of chela, carpus, merus, legs, and car-

VOLUME 106, NUMBER 2

349

Fig. 1 . Cambarus {Hiaiicambarus) elkensis. new species. All from holou pe male. Form I. except C. E. from
morphotype male. Form II. and H. from allotype female): A. lateral view of carapace: B. C. mesial view of first
pleopod: D. caudal view of first pleopods: E. F. lateral view of first pleopod: G. antennal scale: H. annulus
venlralis: 1. episiome: J. dorsal view of carapace; K, proximal podomeres of third, fourth, and fifth pereiopods:
L. dorsal view of distal ix>domeres of cheliped. (See Table 1 for precise measurements.)

350

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

apace brown; abdomen darker brown; tu-
bercles on opposable margins of fingers and
mesial spiniform tubercle of corpus yellow-
ish; distinct black band lining cervical
groove; mandibular adductor region with
black reticulated pattern; rostral margins and
dorsal tubercles on chela reddish; under-
surface cream.

Types. —The holotype, allotype, and
morphotype (USNM 260038, 260039, and
260040, respectively) are deposited in the
National Museum of Natural History,
Smithsonian Institution, Washington D.C.
A small series of paratypes (1 5 I, 8 6 II, 10
9) is at The Ohio State University at Newark
Crayfish Museum.

Type locality.— ThQ Laurel Fork of the
Left Fork of the Holly River (Holly River
-^ Elk River -^ Kanawha River drainage)
at Holly River State Park campground,
Webster County, Hacker Valley District,
West Virginia. (1.1 (air) km NNE of Hacker
Valley.) At this site, the stream is about 10
m wide, 0.5 m deep, and flows over a sub-
strate of sandstone boulders, cobbles, grav-
els, and sand. The specimens were captured
from under cobbles lying on sands and grav-
els where there was considerable current.
The adjacent terrestrial vegetation included
hemlock (Tsuga), birch (Betula), alder (Al-
nus), and rhododendron (Rhododendron).

Range and specimens examined. —The
species is endemic to the upper Elk Basin,
specifically in the Elk River above Sutton
Lake, and in the Holly and Birch rivers. The
collectors were R. F. Jezerinac, G. W. Stock-
er and T. Jones (Coll 1) and GWS and TJ
(Coll 2) unless otherwise stated.

We have examined 22 collections con-
taining a total of 95 specimens— 6 (Form 1)
males, 46 (Form II) males, and 43 females
from the following localities: Nicholas
County: Birch River Intersection (Inters)
County Road (Co Rd) 10 and Co Rd 1/9
(0.8 (air) km E of Birch River), 10 Sep 1988,
Coll 1 (1 6 II); Pocahontas County: Slaty
Fork (Fk.) United States Route (U S Rte)

219, 0.3 km (0.4 mi) S of Co Rd 219/12
(0.3 (air) km SE of Slaty Fk.), 21 Aug 1988,
Coll 2 (1 6 I, 6 9); Old Field Fk. U S Rte
250, 12.8 km (8.0 mi) NW of Co Rd 219/
25 (at Marys Chapel), 19 Jul 1989, Coll 1
(2 $ II, 2 9); Old Field Creek (Ck.) Inters U
S Rte 219 and Co Rd 219/1 (3.0 (air) km S
of Slaty Fk.), 27 May 1989, GWS, RFJ (1
9); Webster County: Laurel Fk. Holly River
State Park Campground (1.1 (air) km NNE
of Hacker Valley), 17 Jun 1988, GWS, RFJ
(4 S II, 5 9); same locality 23 Aug 1988, Coll
2 (I 5 1, 3 <5 II, 3 9) ; and 20 Jun 1990, GWS,
Chelsey Stocker, Vicky Stocker (I S I, 2 S
II, 3 9); Right Fk. Inters St Rte 20 and Co
Rd 5 (at Diana), 3 Sep 1988, GWS, RFJ (1
9); 20 Jul 1989, Coll 1 (1 $ II, 1 9); (10) Birch
R. Co Rd 30, 6.1 km (3.8 mi) N of St Rte
20 (4.6 (air) km RW of Cowen), 23 Aug

1988, Coll 2 {I 6 II, 1 9); Elk R. Co Rd 26,
5.6 km (3.5 mi) NE of Co Rd 15 (3.0 (air)
km NW of Bergoo), 22 Aug 1988, Coll 2 (4

5 II, 4 9); Back Fk. Co Rd 24, 5.1 km (3.2
km) E of St Rte 20 (2.2 (air) km E of Webster
Springs), 20 Jul 1989, Coll 1 (1 <5 I, 6 (5 II,

6 9); Left Fk. Co Rd 3 just W of St Rte 20
(1.3 (air) km SW of Hacker Valley), 21 Jul

1989, Coll 1 (3 S II, 1 9); Right Fk. Inters
Co Rd 5/1 and Co Rd 15 (at Guardian), 21
Jul 1989, Coll 1 (2 S II); Left Fk. Inters Co
Rd 3 and Co Rd 8 (at Polling), 21 Jul 1989,
Coll 1 (3 S II); Elk R. Inters Co Rd 26/1
and Co Rd 26/7 (1.9 (air) km NE of Bergoo),
20 Jul 1989, Coll 1 (1 <5 I, 4 5 II, 1 9); Elk
R. Co Rd 26, 5.6 km (3.5 mi) NE of Co Rd
1 5 (3.0 (air) km NW of Bergoo), 20 Jul 1 989,
Coll 1 (2 5 II, 3 9, 1 9 ovig.); Elk R. Co Rd
15/3, 0.8 km (0.5 mi) E of St Rte 15 (0.5
(air) km SW of Curtin), 20 Jul 1989, Coll 1
(1 S II); Birch R. Inters Co Rd 30 and Co
Rd 40 (at Boggs), 21 Jul 1989, Coll 1 (2 6
II, 2 9); Leatherwood Cr. Co Rd 26/4, 0.8
km (0.5 mi) S of Co Rd 26 (1.1 (air) km S
of Bergoo), 20 Jul 1989, Coll 1 (3 6 II, 3 9);
Elk R. Co Rd 7 at Webster Springs Water
Treatment Plant (2.4 (air) km W of Webster
Springs), 22 Jul 1989, Coll 1 (1 $ II); USNM

VOLUME 106, NUMBER 2

351

43706 Elk R., Cogar's Mills, 30 Jul 1899,
Collectors unknown (16 1) [locality un-
known, Braxton County on USNM label].

Variations.— Most specimens examined
have concave rostral margins that taper to
the acumen and the rostral length is greater
than the rostral width. Some specimens have
subparallel rostral margins, others have
margins that constrict rather sharply to form
the acumen, and some have rather broad
rostra. The suborbital angle varies from ob-
solete to obtuse. These variations are not
restricted to any specific watershed.

Size.— ThQ largest specimen examined is
a Form II male from the Birch River in
Webster County with a carapace length (CL)
of 45.6 mm. The largest female has a 39.0
mm CL and those of the largest and smallest
Form I males are 41.2 mm and 36.7 mm,
respectively. For measurements of the types
see Table 1.

Life history notes.— Form I males were
captured on 20 June (1 specimen), 20 July
(2), and 21 and 23 August (1 each). The only
ovigerous female was caught on 2 1 July and
carried 112 eggs having diameters of 2.2-
2.6 mm. This female has a CL of 37.6 mm.
Specimens are not available for the other
months of the year.

Ecological notes. —As has been observed
with other members of the subgenus Hia-
ticambarus (Hobbs, 1981:147), C. (H.) elk-
ensis is found under loose rocks in riffles,
or pools that have currents. The species was
sought for unsuccessfully in small, head-
water streams. In these smaller tributaries
C. (Cambarus) bartonii carinirostris Hay
(1914) was abundant. We also sampled large
rivers, especially below Sutton Lake, and
failed to capture it. Cambarus {Puncticam-
barus) robustus Girard (1852) was more
common in these bodies of water. Since the
species is found in the Birch River below
Sutton Lake, we suspect that suitable hab-
itat for this species was probably destroyed
when Sutton Lake was constructed.

Relationships. — Cambarus (//.) elkensis

Table 1 . — Measurements (in mm) of types of Cam-
barus (Hiaticambarus) elkensis, new species.

Character

Holotype

Allotype

Morpho-
type

Carapace

Height
Width
Length

15.2
21.0
40.0

12.3
16.1
32.8

14.2
18.7
37.3

Areola

Length
Width

15.0
3.3

12.0
2.4

13.5
3.2

Rostrum

Length

Length to anterior
postorbital ridges
Width between eyes

9.3

7.4
4.5

8.0

6.5

3.7

8.8

7.6
4.6

Postorbital ridge

Width

9.0

7.5

8.3

Chela (right)

Length lateral margin
Length mesial margin

of palm
Width of palm
Length of dactyl
Thickness of palm

40.8

12.5
17.1
26.0
10.2

26.9

8.3
11.0
17.1

6.7

31.7

9.4
14.0
20.5

8.4

Abdomen

Length
Width

38.6

17.4

34.0
13.9

38.2
16.1

Gonopod

Length

8.3

—

7.8

Antennal scale

Length
Width

6.6
2.6

5.4
2.2

6.2*
2.7*

* Left antennal scale.

appears to be most closely related to C (H.)
chasmodactylus but differs from it by hav-
ing the dactyl length/palm length ratio <2.3,
less gaping fingers, and moderately strong
impressions at the base of the fixed finger
of the chela, especially on the ventral sur-
face.

Crayfish associates.— CoW^cXQd with C
(//.) elkensis at one or more sites were Or-
conectes (Crockerinus) sanbornii sanbornii
(Faxon, 1884), O. (Procericambarus) spi-

352

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

nosus (Bundy, 1877), C. (C) b. carinirostris,
and C. (P.) robustus.

Etymology. —We name this crayfish after
the Elk River of West Virginia to which it
appears to be endemic.

Acknowledgments

We thank all individuals who helped in
the collection of specimens for this study,
especially T. Jones. Special thanks are ex-
tended to Dr. H. H. Hobbs, Jr. of the Smith-
sonian Institution, Dr. J. F. Fitzpatrick, Jr.,
of the University of South Alabama, Dr. H.
H. Hobbs, III of Wittenberg University and
two anonymous reviewers for their com-
ments concerning the manuscript. Funding
for this project was provided, in part, by the
West Virginia Division of Natural Re-
sources, Non-game Program. Dave Dennis
and Susan Hendrix of The Ohio State Uni-
versity prepared the illustrations and funds
for them were provided by The Ohio State
University, Department of Zoology.

Literature Cited

Bundy, W. F. 1877. On the cambari of northern In-
diana.— Proceedings of the Academy of Natural
Sciences of Philadelphia 29:171-174.

Faxon, W. 1884. Descriptions of new species of Cam-
barus, to which is added a synonymical list of
the known species of Cambarus and Astacus. —

Proceedings of the American Academy of Arts
and Sciences 20:107-158.

. 1914. Notes on the crayfishes in the United

States National Museum and the Museum of
Comparative Zoology with descriptions of new
species and subspecies to which is appended a
catalogue of the known species and subspe-
cies.— Memoirs of the Museum of Comparative
Zoology at Harvard College 40(8):35 1-427.

Girard, C. 1852. A revision of the North American
astaci, with notes on their habits and geographic
distribution.— Proceedings of the Academy of
Natural Sciences of Philadelphia 6:87-91.

Hay, W. P. 1914. Cambarus bartonii carinirostris Hay.
In Walter Faxon, Notes on the crayfishes in the
United States National Museum and Museum
of Comparative Zoology.— Memoirs of the Mu-
seum of Comparative Zoology at Harvard Col-
lege 40(8):384-385.

Hobbs, H. H., Jr. 1981. The crayfishes of Georgia. —
Smithsonian Contributions to Zoology 318:viii
+ 549.

James, H. A. 1966. Range and variations of the sub-
species of Cambarus longulus (Decapoda, As-
tacidae).— Proceedings of the United States Na-
tional Museum 119(3544):1024.

Jezerinac, R. F., & G. Whitney Stocker. 1989. Dis-
tribution of the stream crayfishes of the genus
Cambarus (Decapoda: Cambaridae) in West
Virginia.— Ohio Journal of Science 89(2):2-3.

(RFJ) Department of Zoology, The Ohio
State University at Newark, University
Drive, Newark, Ohio 43055, U.S.A.; (GWS)
13773 Bodle Road, Newark, Ohio 43055,
U.S.A.

PROC. BIOL. SOC. WASH.
106(2), 1993, pp. 353-358

TWO NEW SPECIES OF GLIRICOLA (PHTHIRAPTERA:

GYROPIDAE) FROM THE SPINY TREE RAT,

MESOMYS HISPID US, IN PERU

Roger D. Price and Robert M. Timm

Abstract.— Two new species of Gliricola, G. woodmani and G. halli (Phthir-
aptera: Gyropidae), are described and illustrated for specimens from the spiny
tree rat, Mesomys hispidus (Rodentia: Echimyidae), in Peru.

Resumen.—SQ describe e ilustra dos nuevas especies de Gliricola, G. wood-
mani y G. halli (Phthiraptera: Gyropidae) que fueron encontradas en una rata
espinosa arborea, Mesomys hispidus (Rodentia: Echimyidae), en Peru.

Thirty-three species of chewing lice of the
genus Gliricola Mjoberg currently are rec-
ognized, these being found on members of
the rodent families Echimyidae, Caviidae,
and Capromyidae. The majority, or 23, of
these louse species occur on echimyids, with
5 each on caviids and capromyids. In the
most recent work on Gliricola, Emerson &
Price (1975) describe five of these taxa as
new, provide illustrations and brief reviews
for seven described earlier, and give liter-
ature citations that include all previously
known taxa.

We recently obtained two important se-
ries of Gliricola from the spiny tree rat, Me-
somys hispidus (Desmarest) (Rodentia:
Echimyidae), in Peru. Our study of these
specimens confirms that two species are
present and their unique features indicate
that the specimens we have represent two
undescribed species. It is our purpose here
to describe and illustrate these new species.

The locality of capture for the host, Me-
somys hispidus, is Reserva Cuzco Amazon-
ico, 14 km E of Puerto Maldonado, Dept.
Madre de Dios, in extreme southeastern
Peru at an elevation of 200 m [12°33'S,
69°03'W], Reserva Cuzco Amazonico is a
national wildlife reserve located on the north
bank of the Rio Madre de Dios, approxi-
mately 300 km east-northeast of the city of
Cuzco. For details of the habitat, climate,

and history of the reserve see Duellman &
Koechlin (1991). Woodman et al. (1991)
provide an annotated listing of the mam-
mals at this site.

Gliricola woodmani, new species
Figs. 1-5

Type host.— Mesomys hispidus (Desma-
rest).

Female. —As in Fig. 1. Head longer than
wide, with numerous short dorsal setae.
Thorax as shown; mesonotum with row of
22 short setae. Marginal abdominal tergal
setae: I, 21; II-V, 25-26; VI-VII, 23; VIII,
14; with markedly longer group of setae lat-
erally on IV- VI. Anterior abdominal tergal
setae: I, 2; II, 14; III-V, 20-23; VI-VII, 17-
18; VIII, 14. Last tergite with total of 8 an-
terior setae and marginally each side with
medium setae flanking pair of very long se-
tae. Pleura II- VII each with 14-16 marginal
and anterior setae, including medium to long
seta at outer comer; pleuron VIII with 6-9
setae, including single very long comer seta.
Large spiracles on pleura III-VIL Marginal
abdominal stemal setae: II, 6; III-VII, 9-
1 1. Anterior abdominal stemal setae: II, 5;
III-VII, 6-10. Stema II-III as in Fig. 3. Ster-
num VIII with total of 10 setae. Ventral
terminalia (Fig. 4) with posterior margin
bearing 3 + 2 short setae on each side and

354

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

anteriorly with three prominent lobes, each
of outer pair of lobes with two slender spat-
ulate setae and inner lobe with medium seta
laterad of shorter broader spatulate seta.

Male. —As in Fig. 2. Much as for female,
except as follows. Mesonotum with row of
22-23 setae. Marginal abdominal tergal se-
tae: I, 19-20; II-V, 23-29; VI-VII, 20-23;
VIII, 13-14; all setae short and of essentially
similar length. Anterior abdominal tergal
setae: II, 16-18; III-V, 25-27; VI-VII, 21-
30; VIII, 14-17. Last tergite with total of
8-9 anterior setae and marginally each side
with single very long seta flanked by several
short setae. Pleura II- VII each with 12-17
marginal and anterior setae, including me-
dium seta at outer comer; pleuron VIII with
9-11 setae, including very long comer seta.
Marginal abdominal sternal setae; II, 6; III-
VIII, 7-10. Anterior abdominal sternal se-
tae: II-VII, 8-12; VIII, 12-14. Chaetotaxy
of subgenital plate as shown. Genitalia (Fig.
5) relatively simple, with straight parameres
slightly swollen basally, endomeral plate
posteriorly flattened to slightly rounded,
transverse bridge near paramere base, slen-
der tapered basal apodeme, and small light-
ly spiculate sac.

Dimensions (in mm).— Temple width, fe-
male 0.23, male 0.21-0.22; prothorax width,
female 0.20, male 0.19; metathorax width,
female 0.28, male 0.26; abdominal width at
V, female 0.43, male 0.31-0.33; head length,
female 0.27, male 0.26; total length, female
1.68, male 1.39-1.41; male genitalia width
0.08, length 0.40-0.41, paramere length
0.07-0.08.

Type material. —Hololype male, ex Me-
somys hispidus, Peru: Dept. Madre de Dios,
14 km E Puerto Maldonado, Reserva Cuzco
Amazonico, el. 200 m, 14 June 1989; in the
Snow Entomological Collection, University
of Kansas, Lawrence. Paratypes, 1 female,
1 male, same data and depository as holo-
type.

Rem.arks.—T]i\s species is readily sepa-
rated from all other species of Gliricola on
the basis of its large dimensions, relatively

simple male genitalia, the unique arrange-
ment of the spatulate setae on the female
ventral terminalia, the longer lateral mar-
ginal tergal setae on female abdominal seg-
ments IV-VI, the single very long seta on
pleuron VIII and the medium to long seta
on each of pleura II-VII, the pair of very
long setae on each side of the last female
tergum and a single such seta on the male,
and the large number of setae on sterna II-
III. Although some other Gliricola may share
features similar to some of the above, none
has all of them. In fact, G. woodmani differs
so significantly from all other congeneric
species that it is difficult to select even a
closely related species.

Etymology.— T\iis species is named in
honor of Neal Woodman, University of
Kansas, in recognition of his efforts in ob-
taining these and other valuable specimens
of ectoparasites and their hosts from the
Neotropics, and his efforts in unraveling
systematic problems in Neotropical mam-
mals.

Gliricola halli, new species
Figs. 6-9

Type host.—Mesomys hispidus (Desma-
rest).

Female.— l:^Qdid and thorax near to G.
woodmani. Abdomen as in Fig. 6. Meso-
notum with row of 20-2 1 short setae. Mar-
ginal abdominal tergal setae: I, 18-26; II-
VI, 29-37; VII, 25-30; VIII, 21-22; with
lateral setae only slightly longer than me-
dian setae. Anterior abdominal tergal setae:
II, 9-17; III-V, 16-24; VI-VII, 21-28; VIII,
18-23. Last tergite with total of 6-14 an-
terior setae and marginally each side with
medium setae ffanking pair of very long se-
tae. Pleural marginal and anterior setae, in-
cluding longer seta at outer comer: II, 11-
13; III, 9-15; IV-VII, 11-14; VIII, 7-10
setae, including 2 very long posterior setae.
Large spiracles on pleura III- VII. Marginal
abdominal sternal setae: II, 0; III, 6-8; IV-
V, 9-11; VI-VII, 11-14. Anterior abdom-

VOLUME 106, NUMBER 2

355

Figs. 1-9. 1-5, Gliricola woodmani: ( 1 ) female; (2) male; (3) female sterna II-III; (4) female ventral terminalia;
(5) male genitalia; 6-9, Gliricola halli: (6) female abdomen; (7) female stema II-III; (8) male genitalia; (9) female
ventral terminalia.

356

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

inal sternal setae: II, 4-5; III-IV, 1-8; V, 4-
8; VI, 6-8; VII, 11-14. Sterna II-III as in
Fig. 7. Sternum VIII with total of 10-17
setae. Ventral terminalia (Fig. 9) with each
side of posterior margin bearing two minute
setae medially and five slender spatulate se-
tae laterally, each side anteriorly with two
medium setae laterad and two short setae
mediad of lobe bearing single seta and broad
spatulate seta.

Male. —Much as for female, except in cer-
tain quantitative chaetotaxy. Marginal ab-
dominal tergal setae: I, 23; II-VI, 28-32;

VII, 27; VIII, 16. Anterior abdominal tergal
setae: II, 6; III-IV, 16-17; V-VII, 20-23;

VIII, 17. Last segment with five anterior
setae. Genitalia (Fig. 8) much as for G.
woodmani, but with evenly rounded en-
domeral plate and shorter parameres.

Dimensions (in mm).— Temple width, fe-
male 0.20-0.2 1 , male 0.20; prothorax width,
female 0. 1 7-0. 1 9, male distorted; metatho-
rax width, female 0.21-0.27, male distort-
ed; abdominal width at V, female 0.33-0.40,
male distorted; head length, female 0.22-
0.24, male 0.22; total length, female 1.35-
1.51, male 1.24; male genitalia width 0.08,
length 0.38, paramere length 0.05-0.06.

Type material. — Holotype female, ex
Mesomys hispidus, Peru: Dept. Madre de
Dios, 14 km E Puerto Maldonado, Reserva
Cuzco Amazonico, el. 200 m, 14 June 1989;
in the Snow Entomological Collection, Uni-
versity of Kansas, Lawrence. Paratypes, 3
females, 1 male, same data and depository
as holotype.

Remarks.— ^Y^n though G. halli occurs
on the same host as G. woodmani, the for-
mer is easily separated by its smaller di-
mensions, the arrangement of the spatulate
setae on the female ventral terminalia, the
shorter lateral marginal tergal setae on fe-
male abdominal segments IV- VI, the pair
of very long setae on pleuron VIII, the small
number of setae on sterna II-III, and the
generally larger number of tergal setae and
small number of pleural setae.

A number of other Gliricola species have
a similar type of setal configuration asso-

ciated with the female ventral terminalia,
but they differ significantly in other aspects.
Probably the closest species morphologi-
cally to G. halli is G. humilis Wemeck from
Proechimys albispinus (I. Geoffroy); how-
ever, the latter has smaller dimensions, a
different length and shape of the male gen-
italic parameres, a markedly truncate fe-
male abdomen, as well as a different ab-
dominal chaetotaxy.

Etymology.— This species is named in
honor of the late E. Raymond Hall, Uni-
versity of Kansas, in recognition of his nu-
merous contributions to mammalogy and
conservation and his establishment of an
outstanding research collection and library
at the University of Kansas for the study of
mammalogy.

Discussion

The Neotropical rodent family Echimyi-
dae is the most speciose and ecologically
diverse of the living caviomorph rodents. It
includes some 15 genera and 70 to 100 spe-
cies. The family was already diverse by the
Oligocene (Reig 1986). The genus Meso-
mys, the spiny tree rats, is one of the poorest
known genera in the family. Four species of
Mesomys currently are recognized; how-
ever, these names are based on few speci-
mens and the relationships of the known
populations in the genus are in need of re-
view (Emmons & Feer 1990, Nowak 1991).
In a recent preliminary analysis of the re-
lationships of several groups of echimyids,
Patton (1989) found Mesomys [hispidus] to
occupy a basal but somewhat ambiguous
position with respect to the other lineages.
However, Mesomys is clearly an old and
distinct lineage within the Echimyidae. The
most widespread species within the genus
is Mesomys hispidus, which is found in the
northern and western Amazon Basin, oc-
curring in southern Colombia and Vene-
zuela, eastern Ecuador and Peru, and west-
em Brazil.

Although the occurrence of two conge-
neric species of chewing lice on the same

VOLUME 106, NUMBER 2

357

host taxon is not typical, it is also not un-
usual. Within the Gliricola, a number of host
taxa share two or more species of lice. This
may in part be due to confusion in the host
taxonomy, but there are sufficient examples
of co-occurrence that we are not suspicious
of the material we are using as the basis of
the description of our two new species. Ad-
mittedly, longer series of the lice would be
preferable, but the differences between the
two are clearcut and there is no way they
can be confused.

The discovery of two congeneric species
of chewing lice on a single host individual
confirms our suspicions that species of Glir-
icola can be truly sympatric. How these two
congeners are distributed on the host and
precisely what they are feeding upon remain
to be documented.

To date, 42 species of chewing lice in 4
genera (Gliricola, Gyropus Nitzsch, Harri-
sonia Ferris, and Hoplomyophilus Mendez)
have been described from 25 host species
representing 8 genera of echimyids. Within
these Gliricola, we find that each genus of
host has its own fauna of lice and there are
,no shared species with any other host genus.
Species of Gliricola have been described
from Diplomys (1 species), Echimys (3 spe-
cies), Kannabateomys (1 species), Euryzy-
gomatomys (2 species), Hoplomys (1 spe-
cies), Isothrix (1 species), and Proechimys
(14 species). The discovery of these two new
species (G. woodmani and G. halli) on Me-
somys is consistent with this, as no lice have
been described previously from this host
genus.

Given the diverse radiation of the echi-
myids in South America and the paucity of
lice available from these rodents, we strong-
ly suspect that numerous new species of
Gliricola have yet to be collected and much
remains to be learned about the systematics
and host relationships in this diverse genus.

Acknowledgments

Field work at Cuzco Amazonico by RMT
was supported by National Geographic So-

ciety Grant 4016-89 and the Museum of
Natural History. Blgo. Jose Purisaca P. of
the Direccion General Forestal y de Fauna,
Ministerio de Agriculture, Lima, issued per-
mits for our work. Jose E. Koechlin, of Cuz-
co Amazonico Lodge, provided excellent fa-
cilities and support for our studies there.
Anthony B. Luscombe and the Asociacion
de Ecologia y Conservacion (ECCO) were
most instrumental in assisting with logis-
tics, equipment, and supplies in Peru and
contributed immeasurably to making our
work successful. William E. Duellman and
Philip S. Humphrey's efforts in securing
funds and aiding in logistics are most ap-
preciated. Rosa Arana, Errol D. Hooper,
Cecilia Pacheco, Victor Pacheco, Cheryl A.
Schmidt, and Neal Woodman provided
outstanding assistance with the fieldwork.
We thank Gloria Arratia for translating our
abstract into Spanish for the resumen, and
Barbara L. Clauson for her constructive
comments on this manuscript. Our study
has been partially supported by Project No.
Min-17-015 of the Minnesota Agricultural
Experiment Station and has been assigned
paper no. 20,157 of the Scientific Journal
Series.

Literature Cited

Duellman, W. E., & J. E. Koechlin. 1991. The Re-
serva Cuzco Amazonico, Peru: Biological in-
vestigations, conservation, and ecotourism.—
Occasional Papers of the Museum of Natural
History, The University of Kansas, 142:1-38.

Emerson, K. C, & R. D. Price. 1975. Mallophaga of
Venezuelan mammals. — Brigham Young Uni-
versity Science Bulletin, Biological Series, 20,
No. 3:1-77.

Emmons, L. H., & F. Peer. 1990. Neotropical rain-
forest mammals: a field guide. University of
Chicago Press, Chicago, 281 pp.

Nowak, R. M. 1991. Walker's mammals of the world.
Vol. II. 5th edition. Johns Hopkins University
Press, Baltimore, pp. 643-1629.

Patton, J. L., & O. A. Reig. 1989. Genetic differen-
tiation among echimyid rodents, with emphasis
on spiny rats, genus Proechimys. Pp. 75-96 in
K. H. Redford and J. F. Eisenberg, eds., Ad-
vances in neotropical mammalogy. Sandhill
Crane Press, Inc., Gainesville.

Reig, O. A. 1986. Distribution patterns and differ-

358

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

entiation of high Andean rodents. Pp. 404-439
in F. Vuilleumierand M. Monasterio, eds.. High
ahitude tropical biogeography. Oxford Univer-
sity Press, Oxford.
Woodman, N. et al. 1 99 L Annotated checklist of the
mammals of Cuzco Amazonico, Peru.— Occa-
sional Papers of the Museum of Natural History,
The University of Kansas, 145:1-12.

(RDP) Department of Entomology, Uni-

versity of Minnesota, St. Paul, Minnesota
55108, U.S.A. (Current address) 4622 Kin-
kead Ave., Fort Smith, Arkansas 72903,
U.S.A.; (RMT) Museum of Natural History
and Department of Systematics and Ecol-
ogy, University of Kansas, Lawrence, Kan-
sas 66045-2454, U.S.A. (direct reprint re-
quests to RMT).

PROC. BIOL. SOC. WASH.

106(2), 1993, pp. 359-365

SPADELLA JAPONICA, A NEW COASTAL
BENTHIC CHAETOGNATH FROM JAPAN

Jean-Paul Casanova

Abstract.— Spadella japonica, a. new benthic chaetognath, is described from
the coastal waters of Japan. It differs from all known species of the genus
Spadella, mainly by the opening of the female genital orifices at the bottom of
a cupel and the small number of hooks. The distribution of the three Spadella
species living in Japanese waters is presented.

When Dr. Taichiro Goto (Mie Univer-
sity, Japan) gave me specimens of a new
Paraspadella during the first workshop on
chaetognaths (University of Surrey, Sep-
tember 1 988), P. gotoi Casanova, 1 990, there
were also two individuals of a Spadella that
he thought to be new. They were collected
in Misaki on 6 August 1987. In November
1991, I received 16 additional individuals
caught in Kominato on 12 September 1991
which, with the other two, differ by many
characters from the species of Spadella hith-
erto described.

Spadella japonica, new species
Figs. 1-3, Table 1

The holotype and two paratypes are de-
posited with the National Science Museum
Tokyo (NSMT-Ch. 20 and 21-22 respec-
tively). Three other paratypes are deposited
in the Museum national d'Histoire Natu-
relle, Paris (UC 366) and three other ones
in the National Museum of Natural History,
Washington, D.C. (USNM 157572). All are
from Kominato.

Description. — Eighteen specimens stud-
ied. Body stumpy when adult (Fig. 1) and
transparent. Length up to 3.75 mm not in-
cluding tail fin. Tail constitutes 48.6 to
51.8% of total length.

Number of hooks increasing with age,
from six (small specimen: 3.25 mm) to eight
(larger ones: 3.60-3.75 mm). Anterior teeth
three or four, short (Fig. 2a, b). No posterior

teeth. Pigment cell of eyes more enlarged in
the second set of specimens received (Fig.
3a). Corona ciliata on neck, oval and trans-
versely elongated (Figs. 1, 3b). Collarette
very wide at level of posterior part of head
and neck, then narrower on trunk (Figs. 1 ,
3b). Sensory tufts symmetrically arranged
on whole body (Fig. 1). Numerous adhesive
papillae on ventral part of body (Fig. 3c, d),
from head to tail, and on both ventral and
dorsal sides of fins. Gut with small intestinal
diverticula at level of neck, not always vis-
ible on preserved specimens. Transverse
musculature thin, stretching from neck al-
most to transverse septum. Ventral ganglion
about middle third of trunk.

Lateral fins beginning on posterior part
of trunk (about 15-19% of the trunk length)
and reaching posteriorly to seminal vesicles
(Fig. 1). Tail fin spatulate. Rayless zone
wholly absent on all fins. Left lateral fin of
a small specimen bearing dorsally a small
area with tiny papillae (Fig. 3e) such as those
found on adhesive organs of Paraspadella
gotoi. Ovaries reach to about midlength of
ventral ganglion; their aperture lateral, at
bottom of brown colored elongate cupel (Fig.
3f, g), of which largest diameter (0.20-0.25
mm) is the same as that of mature ova.
These cupels are the sole colored parts of
the body. Ovaries with from one to five ma-
ture ova and other smaller ones. Seminal
vesicles small, hook-shaped when empty, in
close contact with both lateral and tail fins,

360

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. \. Dorsal view of Spadella japonica, new spe-
cies. CC = corona ciliata, SV = seminal vesicles, VG
= ventral ganglion.

Opening at posterolateral extremity (Fig. 3h,

i).

Remarks.— Two specimens of Spadella
japonica are interesting. One has a curious

"papillated organ" arising from the epi-
dermis on the ventral right side of the tail,
not far from the transverse septum. It is 0. 1 5
mm long and constructed of a short stalk
ending in a swelling provided with papillae
(Fig. 3c, d). A second one shows two of these
"organs" budding on the head and neck: the
papillated swellings are visible but there are
no stalks (Fig. 3j, k).

These observations are pertinent to the
taxonomic position oi Spadella moretonen-
sis Johnston & Taylor, 1919. Indeed, since
the revision of the benthic family Spadel-
lidae Tokioka, 1965 by Bowman & Bieri
(1989), the genus Spadella is restricted to
species lacking adhesive organs, those pro-
vided with such organs constituting the ge-
nus Paraspadella. It has been demonstrated
that these organs are modified parts of fins
(Casanova 1990). Spadella moretonensis w3is
described from East Australia from a single
specimen having two club-shaped papillat-
ed bodies on the posterior half of the tail,
situated ventrally on the right side. But the
authors added: "Though they became
stained like the tissues of the animal [when
using haematoxylin] yet their asymmetrical
arrangement and general appearance sug-
gest that they are foreign bodies— perhaps
of an algal nature." Later on, Tokioka &
Pathansali (1964) wrote: "Species of schi-
zoptera-gTOup l=Paraspadella] are easily
distinguishable by their characteristic ad-
herent organ, while S. moretonensis ... is
devoid of any complete palm-shaped ad-
herent organ ... It is possible this might
rather be a form of schizoptera-gvoup bear-
ing no complete adhesive organs out rudi-
mentary ones." Alvariiio (1 98 1) agreed with
this view. According to Salvini-Plawen
(1986), who first proposed splitting the large
genus Spadella into three smaller ones, some
uncertainty concerns this species with re-
spect to the asymmetry of these bodies.
Nevertheless he placed it in the genus Ge-
phyrospadella (now included in Paraspa-
della). Lastly, because of this uncertainty,
this species was not compared with other

VOLUME 106, NUMBER 2

361

Fig. 2. SEM photographs of Spadella japonica, new species: a, Ventral part of head ( x 1 80); b, Details of
teeth (X 500).

ones in recent papers, neither with Paraspa-
della nor with Spadella (Casanova 1990,
1991).

It appears that the bodies described in S.
moretonensis are the same as those found
in S. japonica. Because in the latter they are
present in only two specimens and on the
head and neck of one of them, they cannot
be considered rudimentary adhesive organs.
Thus S. moretonensis is not a Paraspadella
but a Spadella. According to Goto's obser-
vations (pers. comm.), during culture the
epidermis oi^S. japonica sometimes changes
in appearance, probably as a result of food
or age. Especially when fed with Artemia
nauplii, which are not very good for Spa-
della although they eat them, the epidermis
becomes thin and seems to be deformed.
Similarly, the papillae-like structures oc-
curring in a few specimens living in the sea
might be the result of bad environmental
conditions.

Comparisons with other species.— T\iq
main differential characteristics of the eight
species of Spadella known before the pres-
ent study have been given recently (Casa-
nova 1991). Four of them always have pos-

terior teeth: the cave species Spadella
ledoyeri Casanova, 1986, the two deep spe-
cies Spadella birostrata Casanova, 1987 and
S. equidentata Casanova, 1987, and S. ant-
arctica Casanova, 1991. The other four, as
well as S. moretonensis, may or may not
have posterior teeth. In addition, as they
live in neritic temperate or tropical waters,
comparisons will be made only with them.

The unusual structure of the area sur-
rounding the female genital opening easily
separates Spadella japonica from all other
species of the genus. In more particular re-
gard to those being compared: S. cephal-
optera Busch, 1851 has a prominent cement
gland close to each opening; S. angulata To-
kioka, 1951 and 5. gaetanoi AXwdivmo, 1978
are devoid of this gland; the opening is not
described in 5. bradshawi Bieri, 1974 and
thus there is probably a simple orifice as in
the two latter species; as for S. moretonen-
sis, the aperture is trilobed and situated on
a well-marked prominence. Furthermore,
all these species have more numerous hooks
than S. japonica (Table 1).

Other main features show the specificity
of the new Spadella by comparison with

pi)

^Wn'V^K

SjrffM.^^v*' it^r

^ .-/.iWfi

sysoitv I

W'^^-M

j|^(^-' \

»/,/'/;."'W

I^OPk ■ :i ■

^: ., *

Fig. 3. Spadella japonica, new species: a, Right eye; b, Anterior part of body in dorsal view showing the
corona ciliata (arrow); c, Anterior part of the tail of an original specimen in ventral view, showing a curious
"papillated organ" (arrow) and adhesive papillae; d, Enlargement of the "papillated organ" area; e, Area with
tiny papillae on the ventral side of the left lateral fin (other original specimen); f and g, Two aspects of the female
genital opening (arrow) in dorsal view; h, Empty seminal vesicles; i, Mature seminal vesicles; j, Head and neck
of another original specimen with two "papillated organs" (arrows) in dorsal view; k, Enlargement of the organ
on neck (arrow), just under a sensory spot. Scale bars: 0. 1 mm (b, c, e-j), 0.05 mm (d) and 0.02 mm (a, k).

VOLUME 106, NUMBER 2

363

1. Tomioka

2. Aitsu

3. Mitsu

4. Yashima

5. Setozaki

6. Misaki

7. Kominato

8. Otsuchi

9. Nanao
10. Yoron I.

# Spadella cephaloptera

# Spadella angulata

A Spadella japonica n. sp

Fig. 4. Distribution of the three species of Spadella known from neritic waters of Japan.

each of the five species mentioned above.
Indeed, it differs from:

—Spadella cephaloptera, in which lateral fins
begin at level of the transverse septum
(Japanese specimens) or slightly before
(some Mediterranean specimens) and
seminal vesicles are more or less rounded;

—Spadella angulata, which has a special
orange-brown color pattern on the ventral
and dorsal sides of the body and on the
digestive tract (Tokioka & Bieri 1966,
Bieri et al. 1987); according to Goto and
Thuesen (pers. comm.) who observed the
two species in the Japanese waters (spec-
imens alive as well as preserved ones), S.
angulata is opaque (creamy white) while
S. japonica is transparent;

—Spadella bradshawi, in which the corona
ciliata is massive and lateral fins do not
touch the seminal vesicles;

—Spadella gaetanoi, which bears a pair of
cup-like structures between tip of tail and
seminal vesicles, and moreover, the latter
are reniform;

— Spadella moretonensis, in which the ven-
tral ganglion is very short (less than 25%
of the trunk length).

The characteristics of the six species of
Spadella which are always or sometimes de-
void of posterior teeth are summarized in
Table 1.

Distribution.— Three species of Spadella
live in the neritic waters of Japan (Fig. 4).
Spadella cephaloptera has been found in
Misaki, Mitsu and Yashima. According to
Yosii & Tokioka (1939), the two minute
tentacles on the head are found in small
specimens but not on large ones, whereas
in European specimens they also exist on
adults. Spadella angulata has been men-

364

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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VOLUME 106, NUMBER 2

365

tioned in many localities: Otsuchi, Nanao,
Setozaki, Tomioka, Aitsu and near Okina-
wa (Yoron Island, southwest of Japan). Jap-
anese specimens, first described by Tokioka
(1951) as S. cephaloptera forma angulata,
differ from Malay specimens (Tokioka &
Pathansali 1964) in having, usually, one
posterior tooth. Spadella japonica, accord-
ing to Goto (pers. comm.), has been col-
lected among the Zostera belt in Misaki and
in tide pools of Kominato and Tomioka and
is more common than S. angulata.

Acknowledgments

I thank very sincerely Dr. Taichiro Goto
of Mie University (Japan) who gave me the
specimens of the second new benthic species
he sampled in Japan.

Literature Cited

Alvarino, A. 1978. Spadella gaetanoi, a new benthic
chaetognath from Hawaii. — Proceedings of the
Biological Society of Washington 91:650-657.

. 1981. Spadella legazpichessi, 3. new benthic

chaetognath from Enewetak, Marshall Is-
lands. — Proceedings of the Biological Society of
Washington 94:107-121.

Bieri, R. 1974. A new species of Spadella (Chaetog-
natha) from California.— Publications of the Seto
Marine Biological Laboratory 21:281-286.

, M. Terazaki, E. V. Thuesen, & T. Nemoto.

1987. Colour pattern of Spadella angulata
(Chaetognatha: Spadellidae) with a note on its
northern range extension. —Bulletin of Plankton
Society of Japan 34(l):83-84.

Bowman, T. E., & R. Bieri. 1989. Paraspadella anops,
new species, from Sagittarius cave. Grand Ba-
hama Island, the second troglobitic chaeto-
gnath.— Proceedings of the Biological Society of
Washington 102:586-589.

Busch, W. 1851. Beobachtungen iiber Anatomic und
Entwickelung einiger wirbellosen Seethiere.
Chaetognatha. -Berlin 4:93-100.

Casanova, J. -P. 1986. Spadella ledoyeri, chaeto-
gnathe nouveau de la grotte sous-marine ob-
scure des Tremies (calanques de Cassis).— Rap-
ports de la Commission Internationale pour
I'Exploration Scientifique de la mer Mediter-
ranee 30(2): 196.

. 1987. Deux chaetognathes benthiques nou-

veaux du genre Spadella des parages de Gibral-
tar. Remarques phylogenetiques.— Bulletin du
Museum national d'Histoire naturelle, Paris, 4e
ser., 9, section A(2):375-390.

. 1990. A new species of Para^paJc/Za (Chae-
tognatha) from the coastal waters of Japan. —
Proceedings of the Biological Society of Wash-
ington 103:907-912.

. 1991. The first record of a benthic polar chae-
tognath: a new Spadella from the Antarctic —
Journal of Natural History 25:1355-1362.

Johnston, T. H., & B. B. Taylor. 1919. Notes on
Australian chaetognatha.— Proceedings of the
Royal Society of Queensland 31:28-41.

Salvini-Plawen, L. Von. 1986. Systematic notes on
Spadella and on the chaetognatha in general.—
Zeitschrift fiir Zoologische Systematik und Evo-
lutionsforschung 24(2): 122-1 28.

Tokioka, T. 1951. Pelagic tunicates and chaetognaths
collected during the cruises to the New Yamato
Bank in the Sea of Japan.— Publications of the
Seto Marine Biological Laboratory 2:1-25.

. 1965. The taxonomical outline of Chaetog-
natha.— Publications of the Seto Marine Biolog-
ical Laboratory 12:335-357.

, & D. Pathansali. 1 964. Spadella cephaloptera

forma angulata raised to the rank of species. —
Publications of the Seto Marine Biological Lab-
oratory 12:145-148.

, & R. Bieri. 1966. The colour pattern of Spa-
della angulata Tokioka. — Publications of the
Seto Marine Biological Laboratory 14:323-326.

Yosii, N., & T. Tokioka. 1939. Notes on Japanese
Spadella (Chaetognatha).— Annotationes Zoo-
logicae Japonenses 18:267-273.

Laboratoire de Biologic animale (Plane-
ton), Universite de Provence, 13331 Mar-
seille Cedex 3, France.

PROC. BIOL. SOC. WASH.

106(2), 1993, pp. 366-368

CAUDINA INTERMEDIA, A NEW SPECIES OF SEA

CUCUMBER FROM THE SOUTH CHINA SEA

(ECHINODERMATA: HOLOTHUROIDEA: MOLPADIIDA)

Yulin Liao and David L. Pawson

Abstract. — Caudina intermedia, new species is described. The bodywall os-
sicles of this species are exclusively tables; knobbed buttons or plates are absent.
The tables are typical of the genus Caudina Stimpson, 1853 in the strict sense,
but the absence of other ossicle types suggests some affinities with the genus
Hedingia Deichmann, 1938. A revised key to the known species of Caudina
is presented.

Subsequent to the publication of our re-
cent paper on the molpadiid sea cucumbers
of China (Pawson & Liao 1992), an addi-
tional new molpadiid species was found in
the collections of the Institute of Oceanol-
ogy, Academia Sinica, Qingdao (lOAS).

Family Caudinidae Heding, 1931
Caudina Stimpson, 1853

For a summary of the caudinids of China,
see Pawson & Liao (1992).

Caudina intermedia, new species
Fig. lA-G

Material examined. —HoloXype, lOAS
E1056. off eastern Guangdong, 21°45'N,
115°30'E, 9 Jan 1960, 107 m, muddy sand
bottom. Paratype, lOAS El 057, same lo-
cality as Holotype.

Diagnosis. — Ossicles of body wall exclu-
sively tables with solid spires derived from
four pillars fused together, terminating in a
few teeth. Knobbed buttons, perforated
plates, and phosphatic deposits absent.

Description. —Body more or less barrel-
shaped, with conspicuous narrow tail. Ho-
lotype 1 9 mm long, 1 mm in diameter, tail
7 mm long; Paratype approximately the
same size. Body wall thin, translucent. Anus
surrounded by five minute anal teeth. Color
in alcohol dirty gray to whitish. Radial piec-

es of calcareous ring with short bifid pos-
terior projections; interradial pieces slightly
wider than radials (Fig, IG). Stone canal
and polian vesicle single.

Body wall ossicles exclusively tables, dif-
fering slightly in various parts of the body.
In anterior body wall, disc more or less cir-
cular or irregular in outline, 130-180 iim. in
diameter (A^ 1 48 ^ni, SD 9.3), with four large
central perforations and 5-10 peripheral
ones. Spire tall, average height 150 ixm, sol-
id, derived from four converging and fused
pillars, terminating in three or four spines
(Fig. lA, B). In median and posterior body
wall ossicles, disc more or less square in
outline, average diameter 150 ij,m, com-
monly with four large perforations; spire
solid, average height 1 00 fxm, ending in three
blunt teeth (Fig. IC, D). Tables in tail (Fig.
1 E, F) smaller, with numerous perforations
and small knobs on periphery and elsewhere
on disc. Spire low, solid; in a few cases a
single crossbar present.

Remarks.— This new species is distinct
from other Caudina species in lacking ac-
cessory ossicles in the form of buttons or
plates (see key below). In possessing only
tables in the body wall, C. intermedia ap-
proaches Hedingia Deichmann, 1938, but
the tables of Hedingia are usually consid-
erably larger (200-300 fim), have more per-
forations, and typically have two-pillared or

VOLUME 106, NUMBER 2

367

Fig. 1 . Caudina intermedia, new species. A, tables from anterior body wall; B, same in profile view; C, tables
from median and posterior body wall; D, same, in profile view; E, tables from tail; F, same, in profile view; G,
radial and interradial pieces of calcareous ring. The scale measures 200 ^m for A-D, 100 ^m for E-F, and 1.2
mm for G.

368

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(more commonly) three-pillared spires. Both
specimens of C. intermedia are only 1 9 mm
long. It is conceivable that buttons or plates
may develop with further growth, but not
likely, for in other species of Caudina the
full complement of ossicle types is reached
at an early age and small size.

Key to the Known Species of Caudina
(modified after Pawson & Liao 1992)

1. Tables inconspicuous, scarce, re-
duced to rods or plates, or lacking

arenicola (Stimpson, 1853)

- Tables conspicuous, numerous, with
well-developed spire and disk .... 2

2. Tables accompanied by knobbed
buttons or perforated plates 3

- Only tables present; buttons and
plates absent . . intermedia, new species

3. Tables accompanied by knobbed
buttons 4

- Tables accompanied by perforated
plates; no buttons 5

4. Buttons not abundant, weakly
knobbed, usually oval in outline,
with two large elliptical and two
small circular holes

arenata Gould, 1841

- Buttons abundant, strongly
knobbed, usually with irregular out-
line, the four holes more or less alike
in size and form

similis (Augustin, 1908)

5. Discs of tables 180-280 ixm in di-
ameter; perforated plates very vari-
able in form and in number of holes,
lacking knobs

atacta Pawson & Liao 1992

- Discs of tables 150-180 ^ni in di-
ameter; perforated plates vary only
slightly in form and in number of
holes, often possessing a few low

knobs

. . . zhejiangensis Pawson & Liao 1992

Literature Cited

Augustin, E. 1908. Uber Japanische Seewalzen.—
Abhandlungen der Koniglichen Bayehschen
Akademie der Wissenschaften. II. Supplement
i:1^4.

Deichmann, E. 1938. New holothurians from the
western coast of North America and some re-
marks on the genus Caudina.— VioctQdmgs of
the New England Zoological Club 16:103-1 15.

Gould, A. A. 1841. Report on the Invertebrata of
Massachusetts. Cambridge, Massachusetts, xiii
+ 372 pp.

Heding, S. G. 1931. On the classification of the mol-
padids. — VidenskabeUge Meddelelser fra Dansk
naturhistorisk Forening i Kjobenhavn 92:275-
284.

Pawson, D. L. 1977. Molpadiid sea cucumbers (Echi-
nodermata: Holothuroidea) of the Southern At-
lantic, Pacific and Indian Oceans.— Biology of
the Antarctic Seas VI. Antarctic Research Series
26:97-123.

, & Y. Liao. 1992. Molpadiid sea cucumbers

of China, with description of five new species
(Echinodermata: Holothuroidea). — Proceed-
ings of the Biological Society of Washington 105:
373-388.

Stimpson, W. 1853. Synopsis of the marine inver-
tebrates of the Grand Manan. — Smithsonian
Contributions to Knowledge 5:6-17.

(YL) Institute of Oceanology, Academia
Sinica, 7 Nan-Hai Road, Tsingtao, People's
Republic of China; (DLP) Department of
Invertebrate Zoology, National Museum of
Natural History, Smithsonian Institution,
Washington, D.C. 20560, U.S.A.

PROC. BIOL. SOC. WASH.

106(2), 1993, pp. 369-384

HYPORHAMPHUS MEEKI, A NEW SPECIES OF

HALFBEAK (TELEOSTEI: HEMIRAMPHIDAE) FROM

THE ATLANTIC AND GULF COASTS OF THE

UNITED STATES

Heidi M. Banford and Bruce B. Collette

Abstract.— Hyporhamphus meeki is described from the Atlantic and Gulf of
Mexico coasts of the United States. It has been confused previously with H.
unifasciatus (Ranzani) which occurs from southern Florida, Bermuda, the West
Indies, and Mexico south to southern Brazil. Hyporhamphus meeki has more
gill rakers, usually 33-39 on the first gill arch and 26-29 on the second arch
compared to 28-32 on the first arch and 19-25 on the second arch in H.
unifasciatus. Pectoral-fin rays are usually 1 1 or 1 2 vs. 10 or 1 1 in //. unifasciatus.
The ratio of preorbital length to orbit diameter is usually greater than 0.70 in
H. meeki, less than 0.70 in H. unifasciatus.

There has been considerable confusion
regarding the taxonomy and systematics of
the New World halfbeaks (Meek & Goss
1884, Miller 1945). Although the status of
some species has been clarified, that of Hy-
porhamphus unifasciatus (Ranzani, 1842),
the common inshore halfbeak, has been
questioned for more than 100 years (Meek
& Goss 1884, Collette 1978) but not re-
solved. The name Hyporhamphus unifas-
ciatus (Ranzani) has been used for inshore
halfbeaks in the western Atlantic, eastern
Atlantic, in several parts of the Indo-West
Pacific, and the eastern Pacific. The eastern
Atlantic Hyporhamphus were shown to be
H. picarti (Valenciennes, 1846) by Collette
(1965); the Indo-West Pacific halfbeaks H.
limbatus (Valenciennes, 1846) by Parin et
al. (1980). This paper presents morpho-
metric and meristic analysis of western At-
lantic populations of halfbeaks referred to
H. unifasciatus, with the description of a
new species. Our objective is to describe this
new species of Hyporhamphus, and com-
pare it with the true H. unifasciatus. The
new species will be included in further study
of New World halfbeaks and other publi-

cations, particularly the halfbeak section for
"Fishes of the western North Atlantic."

Populations referred to H. unifasciatus
range in the western Atlantic from Uruguay
in the south, northward along the coast of
the Americas, through the Caribbean (Jor-
dan & Evermann 1896), Gulf of Mexico
(Hoese & Moore 1977), and around Ber-
muda (Beebe & Tee- Van 1933) to Cape Cod,
Massachusetts. Strays have been collected
as far north as Chamcook, Passamaquoddy
Bay, New Brunswick (Leim & Day 1959).
In the eastern Pacific they range from Peru
(Hildebrand 1946) to Baja California and
around the Galapagos Islands (Meek & Hil-
debrand 1923). The range of//, unifasciatus
s. s. (type locality, Brazil) is from Bermuda
and peninsular Florida southward through
the Caribbean to Uruguay. Those popula-
tions referred to as //. unifasciatus from out-
side this range constitute superficially sim-
ilar undescribed species of Hyporhamphus
(Collette 1978). This paper deals only with
the western Atlantic populations of the //.
unifasciatus species group. The eastern Pa-
cific populations will be addressed in future
publications.

370

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

80° 65° 50° 35° 20°

Fig. I. Distribution of Hyporhamphus meeki, n. sp. (closed circles) and H. unifasciatus (open circles) based
on material examined.

Materials and methods. —A total of 568 specimens were examined only for meristic

specimens of western Atlantic Hyporham- characters. The number of specimens ex-

phus was examined for 24 morphometric amined for each character varies due to the

and meristic characters; an additional 1088 condition of material. Due to the poor con-

VOLUME 106, NUMBER 2

371

dition of some specimens only some char-
acters could be observed reliably. Material
was chosen to represent the entire geograph-
ical range of what has been considered H.
unifasciatus in the western Atlantic (Fig. 1).
The majority of material examined was ob-
tained from the following institutions (ab-
breviations from Leviton et al. 1985):
AMNH, ANSP, CBL (Chesapeake Biolog-
ical Laboratory, specimens now at VIMS),
CAS, MCZ, MZUSP, SIO, UF, VIMS,
UMMZ, and USNM. Additional material
was collected by the first author in the York
River at Gloucester Pt., Virginia, and is
housed at VIMS. Following the description
of the new species only the material that
was examined for both morphometries and
meristics is listed.

Most characters examined follow Collette
(1965) and Parin et al. (1980). Measure-
ments were made to the nearest tenth of a
millimeter (mm). Abbreviations and de-
scriptions of characters examined are as fol-
lows: SL (standard length); LJL (lower jaw
length, tip of upper jaw to tip of lower jaw);
HDL (head length, from tip of upper jaw to
posterior margin of opercle membrane); UJL
(upper jaw length, from tip of upper jaw to
where upper jaw bends); UJW (upper jaw
width, where upper jaw bends); P1-P2 (dis-
tance from base of upper pectoral ray to base
of anterior pelvic ray); P2-C (distance from
base of anterior pelvic ray to caudal base);
P2-CX (P2-C distance extended anteriorly
from base of anterior pelvic ray to a point
on the body or head); BD-Pj (body depth
at origin of pectoral fin); BD-P2 (body depth
at origin of pelvic fin); ABASE (length of
anal-fin base); DBASE (length of dorsal-fin
base); PjL (pectoral-fin length, distance from
base of uppermost pectoral ray to tip of lon-
gest ray); ORB (soft orbit length); PREORB
(preorbital length, from comer of mouth to
anterior margin of orbit); ANA (number of
anal-fin rays); DOR (number of dorsal-fin
rays); P, (L, R, number of pectoral-fin rays);
PRED (number of predorsal scales in me-
dian row in front of dorsal fin); RGRi (num-

ber of gill rakers on first arch (upper + lower
= total); RGR2 (number of gill rakers on
second arch (upper + lower = total). VERT
(number of precaudal plus caudal vertebrae,
including the hypural plate = total number
of vertebrae).

Statistical analyses utilized SAS software
(SAS Institute, Inc. 1985). Frequency dis-
tributions of counts were compared be-
tween geographic populations and are pre-
sented in summary tables. If two populations
in close geographic proximity did not have
significantly different counts, counts were
combined to form a single population in
subsequent statistical analysis. Infraspecific
geographic variation is discussed herein.
Analysis of Variance (ANOVA) was per-
formed on five data sets of meristic char-
acters. If the F value for an ANOVA was
significant {P < .05), Tukey's Studentized
Range Test (Tukey-Kramer method) (SAS
Institute, Inc. 1985) was performed to de-
termine which means were significantly dif-
ferent from the others.

Values of morphometric characters were
first plotted against SL, and then plotted
against one another to inspect visually for
separation between populations. Only those
plots that proved to be diagnostic for the
species are presented. Regression equations
were generated for the plots of morpho-
metric characters. Residual plots were in-
spected for homogeneity of variance. Due
to heteroscedasticity of variance, all mor-
phometric data were log transformed for re-
gression analysis.

Hyporhamphus meeki, new species
Fig. 2A

Synonymy.— Ax least 10 different com-
binations of names have been used for this
species. A complete synonymy will be in-
cluded in the halfbeak section of "Fishes of
the western North Atlantic." The common-
est names used have been Hyporhamphus
(or Hemiramphus) roberti (about 30 refer-

372

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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VOLUME 106, NUMBER 2

373

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ORBIT DIAMETER (mm)

Fig. 3. Relationship of preorbital length to orbit length in Hyporhamphus meeki (squares) and H. unifasciatus
(triangles).

ences, mostly 1862 to 1892) or unifasciatus
(about 95 references, 1870 up to the pres-
ent). Bruce (1986) prematurely used the
name H. meeki as a nomen nudum in his
treatment of isopod parasites of the genus
Mothocya.

Diagnosis.— A member of the subgenus
Hyporhamphus distinguished from H. rob-
erti (Valenciennes) by having the dorsal and
anal fins covered with scales. This species
is distinguished by the following combina-
tion of characters: gillrakers on the first arch
31-40 (Table 1); gill rakers on the second
arch 20-30 (Table 2); pectoral-fin rays 10-
1 3 (Table 5); ratio of preorbital length to
orbit diameter is usually greater than 0.70
(>0.70 in 92% of 265 specimens examined;
Fig. 3).

Description.— GiW rakers on upper limb

of first arch 8 to 12, usually 9 to 11, mean
9.8; lower limb 20 to 29, usually 24 to 27,
mean 25.1; total of upper and lower limbs
3 1 to 40, usually 33 to 37, mean 34.6 (Table
1). Gill rakers on upper limb of second arch
2 to 6, usually 4 or 5, mean 4.3; lower limb
20 to 26, usually 22 or 23, mean 22.3; total
20 to 30, usually 25 to 28, mean 26.2 (Table
2). Dorsal-fin rays 12 to 17, usually 14 or
15, mean 14.5 (Table 3); anal-fin rays 14 to
18, usually 15 to 17, mean 15.9 (Table 4);
and pectoral-fin rays 10 to 13, usually 1 1 or
12, mean 11.4 (Table 5). Predorsal scales
{n = 87) 34 to 39, usually 35 to 37, mean
36.1. Vertebrae {n = 88) 31-35 precaudal
+ 16-19 caudal = 49-53 total.

Morphometric data for H. meeki is sum-
marized in Table 6, for H. unifasciatus in
Table 7. Lower jaw length 0.79 to 1.54 of

374

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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VOLUME 106, NUMBER 2

375

Table 2.— Numbers of total gill rakes on second arch in populations of Hyporhamphus meeki and H. unifas-
ciatus.

Population

H. meeki

MA to GA

189

26.8

E. coast FL

114

25.9

W. penin. FL

177

26.0

FLA pan. to TX

156

26.2

Yucatan

25.5

Species total

104

196

218

662

26.2

H. unifasciatus

Florida

22.9

West Indies

141

24.1

Cent. America

23.5

Carib. S. Am.

114

24.3

South America

109

23.5

Yucatan

23.3

Bermuda

100

21.9

Species total

133

149

570

23.5

head length with 83% of 240 specimens
greater than or equal to 1.0. Ratio of LJL
to SL 0.20 to 0.35 with 95% of 240 speci-
mens 0.22 to 0.30. PREORB to ORB ratio
0.61 to 1.0. usually 0.70 to 0.90 (92% of
265 specimens). Distance from base of an-
teriormost pelvic ray to caudal base extends
anteriorly to mid-eye, usually between pos-

terior portion of eye and posterior margin
of opercular membrane. Dorsal- and anal-
fin bases about equal, ratio of ABASE to
DBASE 0.83 to 1.07, mean of 0.96 for 265
specimens. Origin of dorsal fin over that of
anal fin. Bases of dorsal and anal fins cov-
ered with scales. Distance from anterior-
most pectoral ray to origin of pelvic fin less

Table 3.— Numbers of dorsal-fin rays in populations of Hyporhamphus meeki and H. unifasciatus.

Population

H. meeki

MA to GA

100

160

14.4

E. coast FL

14.4

W. penin. FL

157

14.7

FL pan. to TX

165

14.4

Yucatan

14.5

Species total

297

262

594

14.5

H. unifasciatus

Florida

14.8

West Indies

127

14.7

Cent. America

14.9

Carib. S. Am.

118

14.9

South America

120

14.8

Yucatan

15.1

Bermuda

14.3

Species total

159

344

551

14.8

376

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 4.— Numbers of anal-fin rays in populations of Hyporhamphus meeki and H. unifasciatus.

Population

H. meeki

MA to GA

107

163

16.1

E. coast FL

15.7

W. penin. FL

157

16.2

FL pan. to TX

165

15.6

Yucatan

16.0

Species total

124

349

110

597

15.9

H. unifasciatus

Florida

16.5

West Indies

128

16.1

Cent. America

16.2

Carib. S. Am.

118

16.3

South America

119

16.3

Yucatan

16.2

Bermuda

16.8

Species total

292

204

550

16.4

than the distance from the pelvic fin origin
to the caudal base. Median pore of preor-
bital canal usually posterior, rarely medial.

Color. —Coloration in life is a silvery light
tan-green. The fleshy tip of the lower jaw is
a bright orange red.

iSzz^.— Adults attain a maximum size of
179 mm SL (USNM 90798, Cape Charles,
Virginia).

Habitat.— W\ specimens observed were

collected near the surface of inshore or es-
tuarine waters. Specimens collected by the
first author in Chesapeake Bay and the Gulf
of Mexico (Florida) were in areas with a
sandy substrate and in proximity of sub-
merged aquatic vegetation (eel grass, Zos-
ter a). As is often the case in estuarine con-
ditions, the water inhabited by H. meeki is
generally turbid.
Early life history . —L^wslq of H. meeki

Table 5.— Numbers of pectoral-fin rays in populations of Hyporhamphus meeki and H. unifasciatus.

Population

H. meeki

MA to GA

158

11.4

E. coast FL

106

11.0

W. penin. FL

147

11.7

FL pan. to TX

120

11.4

Yucatan

11.3

Species total

289

224

558

11.4

H. unifasciatus

Florida

10.8

West Indies

136

10.7

Cent. America

10.9

Carib. S. Am.

108

120

11.0

South America

104

122

10.9

Yucatan

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