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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 



IV 



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 



VI 



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- 



vu 



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 



IX 



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 



XI 



Quechnalia 780 

cardenasii n.c 78 1 

fiilta n.c 78 1 

smithii 781 



PROCEEDING^ 

OF THE 

BIOLOGICAL SOCIETY 

OF 

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 



\^ 



o 



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 



99 



98 



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 



8 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



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

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

Phenetic relationships were assessed us- 
ing correlation and distance matrices 



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

Results 

The diploid number of 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 


A 


A 




B 


SOD-1 


A 


A 




B 


EST-1 


A 


A 




A, B 


EST-2 


A,b 


A 




A, B 


GPI 


A, B 


A 




A 


G3PDH 


A, B 


A, 


b 


A,B 


PGM-1 


A 


A, 


B 


A, B 


6-PGD 


A, B 


A, 


B 


A 


MDH-1 


A 


A 




A, B 


MDH-2 


A, b 


A 




A, B 


SOD-2 


A 


A 




A, B 


AAT-1 


A,b 


A 




A 


IDH-1 


A,C 


B, 


C 


B, C 


IDH-2 


A 


A, 


B 


A 


Pep-2 


A 


A 




A 


ACOH 


A 


A 




A 


CK 


A 


A 




A 


PNP 


A 


A 




A 



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 



10 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



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<u -5 03 til ^ 
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VOLUME 106, NUMBER 1 



13 



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- 



14 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



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



Males 



MB 

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

SB 

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

RB 

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 



5 


21.96 


21.4 


22.3 


4 


22.65 


20.8 


23.9 


7 


23.25 


21.9 


23.9 


13 


23.98 


22.3 


26.7 


11 


24.06 


22.6 


25.7 


5 


24.11 


23.5 


24.7 


5 


16.94 


16.6 


17.4 


7 


17.52 


16.9 


18.0 


4 


17.63 


16.8 


18.5 


11 


18.22 


17.4 


19.5 


5 


1S.63 


18.1 


19.5 


13 


18.69 


17.5 


19.8 


5 


9.19 


8.9 


9.5 


13 


9.37 


8.9 


10.7 


11 


9.43 


9.1 


9.9 


5 


9.74 


8.9 


10.4 


4 


9.95 


9.8 


10.2 


7 


10.17 


9.3 


10.9 


5 


6.90 


6.8 


7.1 


4 


7.48 


7.2 


7.8 


7 


7.61 


7.3 


8.0 


11 


7.74 


7.1 


8.1 


13 


7.80 


7.3 


8.8 


5 


7.92 


7.6 


8.5 


5 


8.49 


8.2 


8.8 


4 


8.63 


8.1 


9.0 


7 


8.91 


8.6 


9.6 


11 


9.36 


9.1 


9.8 


5 


9.36 


9.1 


9.6 


13 


9.43 


8.4 


10.6 


4 


5.87 


5.5 


6.3 


7 


5.88 


5.6 


6.2 


5 


5.96 


5.7 


6.4 


13 


6.18 


5.9 


6.6 


11 


6.40 


6.0 


6.7 


5 


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 



15 



Table 3.— Continued. 



Females 



MB 

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

SB 

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

RB 

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 

Ml 

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 



6 


21.14 


20.5 


21.9 


15 


21.36 


19.9 


22.8 


9 


21.77 


20.8 


23.0 


13 


22.93 


21.7 


24.0 


18 


23.30 


21.6 


24.8 


3 


23.30 


22.8 


24.1 


6 


16.65 


16.3 


17.1 


9 


16.85 


16.5 


17.3 


15 


16.92 


16.0 


17.9 


18 


17.97 


16.9 


18.7 


3 


18.01 


17.7 


18.3 


13 


18.21 


17.3 


18.9 


6 


8.40 


8.0 


8.8 


18 


8.56 


8.2 


9.7 


13 


8.87 


8.1 


9.5 


15 


9.15 


8.5 


9.6 


9 


9.37 


9.0 


10.1 


3 


9.60 


9.0 


10.1 


6 


6.88 


6.5 


7.3 


15 


7.36 


7.0 


7.8 


3 


7.50 


7.4 


7.6 


18 


7.51 


7.0 


8.0 


9 


7.52 


7.1 


8.1 


13 


7.72 


7.2 


8.2 


6 


8.25 


7.8 


8.5 


15 


8.51 


7.9 


9.3 


9 


8.66 


8.1 


9.0 


18 


8.99 


8.5 


9.4 


13 


9.08 


8.5 


9.5 


3 


9.21 


9.0 


9.5 


9 


5.76 


5.5 


5.9 


6 


5.83 


5.5 


6.3 


15 


5.86 


5.4 


6.3 


13 


6.07 


5.8 


6.6 


18 


6.28 


5.6 


6.7 


3 


6.49 


6.2 


6.9 



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



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



16 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



2.0 -■ 



< 0.0 4 



-2.0 -■ 



-4.0 -■ 




G. 


t. bakeri 


♦ 


G. 


p. streckeri 


+ 


G. 


attwoteri 


o* 


G. 


texensis 


DM 



-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 



17 



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. 


% 


Variable 
coeff. 


% 


Variable 
coeff. 


% 








Males 








CBL 


-1.8433 


28.51 


-0.3363 


3.36 


-4.3026 


41.24 


BL 


0.6980 


10.14 


-4.5368 


42.57 


1.8989 


17.09 


PL 


-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 


ZB 


-1.1519 


11.26 


0.6130 


3.87 


0.5727 


3.47 


MB 


2.0305 


17.87 


2.2454 


12.77 


-2.0193 


11.01 


SB 


0.8427 


5.67 


-1.8478 


8.03 


0.3642 


1.52 


RB 


-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 


M3 


0.8382 


2.38 


-1.2829 


2.35 


0.6570 


1.16 


Ml 


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 


BL 


0.4529 


7.94 


-1.7802 


34.17 


-0.2815 


8.76 


PL 


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 


ZB 


0.1029 


1.18 


0.3581 


4.49 


0.5166 


10.49 


MB 


2.4876 


26.45 


-0.5207 


6.06 


0.5198 


9.81 


SB 


-0.3687 


3.06 


0.9749 


8.87 


-1.3019 


19.20 


RB 


-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 


M3 


-0.4172 


1.50 


0.9869 


3.88 


-0.0849 


0.54 


Ml 


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- 



18 



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. attwateri 

G. texensis 

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 



19 



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 



20 



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 



21 



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

gophers.— Journal of Wildlife Management 36: 
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, , & J. C. Patton. 1973. Chromosomal 

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Block, S. B., & E. G. Zimmerman. 1991. Allozymic 
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Dalquest, W. W., & W. Kilpatrick. 1973. Dynamics 
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Plateau of Texas.— The Southwestern Natural- 
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Davis, W. B. 1940. Distribution and variation of 
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Dittmar, G. W., M. L. Deike, & D. L. Richmond. 
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Dowler, R. C. 1 982. Genetic interactions among three 
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Godfrey, C.L.,G.S.McKee,&H.Oakes. 1973. Gen- 
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lege Station, 2 pp. 

Harris, H., & D. A. Hopkinson. 1976. Handbook of 
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North-Holland Publishing Co., Amsterdam, 475 
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Hart, E. B. 1978. Karyology and evolution of the 
plains pocket gopher, Geomys bursarius.— Oc- 
casional Papers of the Museum of Natural His- 
tory, The University of Kansas 71:1-20. 

Hendricksen, R. L. 1972. Variation in the plains 
pocket gopher (Geomys bursarius) along a tran- 
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Transactions of the Kansas Academy of Science 
75:322-368. 

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

, & S. L. Williams. 1982. Genie differentiation 

in pocket gophers of the genus Pappogeomys, 
with comments of intergeneric relationships in 
the subfamily Geomyinae.— Journal of Mam- 
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Kennerly, T. E., Jr. 1954. Local differentation in the 
pocket gopher {Geomys personatus) in southern 



22 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Texas.— The Texas Journal of Science 6:297- 
329. 

. 1959. Contact between the ranges of two al- 

lopatric species of pocket gophers.— Evolution 
13:247-263. 

Murphy, R. W., J. W. Sites, Jr., D. G. Buth, & C. H. 
Haufler. 1990. Proteins I: isozyme electro- 
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sociates, Inc., Sunderland, 588 pp. 

Owen, J. G., & M. A. Chmielewski. 1985. On ca- 
nonical variates analysis and the construction 
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Systematic Zoology 34:366-374. 

Rogers, J. S. 1 972. Measures of genetic similarity and 
genetic distance.— Studies in Genetics, VII, 
University of Texas Publication 7213:145-153. 

Rohlf, F. J. 1988. NT-SYS-pc: numerical taxonomy 
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Exeter Software, Setauket, New York. 

Russell, R. J. 1968. Revision of pocket gophers of 
the genus Pappogeomys. —University of Kansas 
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581-776. 

SAS Institute. 1988a. SAS procedures guide, release 
6.03 edition. SAS Institute, Inc., Cary, North 
Carolina, 441 pp. 

. 1988b. SAS/ST AT user's guide, release 6.03 

edition. SAS Institute, Inc., Cary, North Caro- 
lina, 1028 pp. 

Schmidly, D. J., & F. S. Hendricks. 1 976. Systematics 
of the southern races of Ord's kangaroo rat, Di- 
podomys cr^z7.— Bulletin of the Southern Cali- 
fornia Academy of Science 75:225-237. 

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

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

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

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

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



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

, & . 1978. Morphometric variation in 

the desert pocket gopher (Geomys arenarius). — 
Annals of the Carnegie Museum 47:541-570. 

, & . 1980. Morphological variation in 

the southeastern pocket gopher Geomys pinetis 
(Mammalia: Rodentia).— Annals of the Carne- 
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, & . 1981. Systematic review of the 

Texas pocket gopher, Geomys personatus 
(Mammalia: Rodentia).— Annals of the Carne- 
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 



23 



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 



25 



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 


1 




Falco newtoni 




1 


Coturnix sp. 




2 


Numida meleagris 


2 


4 


Turnix nigricollis 




2 


Coracopsis vasa 




1 


Tyto alba 




2 


Otus rutilus 




2 


Merops superciliosus 




1 


Hypsipetes madagascariensis 


1 


1 


Foudia madagascariensis 




1 



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- 



26 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



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

The most common bird species recovered 
from these surface deposits was Numida. 
Adults of this species weigh over 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 



27 




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 



28 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 







A.>. 





J. 



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 



29 



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 = 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 


« = 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 


« = 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 



30 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



C 

o 

CO 
CO 

on 
+-• 

CO 

+-• 



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 



31 



berthae 



C 
Q) 

'> 

CD 

a 




• - 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 



32 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



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

Comparative material examined.— Os- 
teological material of Coua spp. is rare in 
collections and skeletons of C delalandei, 
coquereli, cursor, and verreauxii are not 
available for comparison. With the excep- 
tion of delalandei, all of these birds are small 
species, and the absence of comparative 
material did not hamper the analysis. The 
pelvis of C. berthae was compared to mod- 
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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Battistini, R. 1965. Sur le decouverte de V Aepyornis 
dans le Quatemaire de I'Extreme-Nord de Mad- 
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Baumel, J. J. (ed.) 1979. Nomina anatomica avium. 
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Berger, A. J. 1952. The comparative functional mor- 
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. 1953. On the locomotor anatomy of the Blue 

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Decary, R. 1938. Les grottes d'Andranoboka.— Bul- 
letin de I'Academie Malgache, new series 21: 
71-80. 

de Saint-Ours, J. 1953. Etude des grottes d'Andra- 
noboka.— Travaux de Bureau Geologique, An- 
tananarivo, 13 pp. 

, & R. Paulian. 1953. Les grottes d'Andra- 
noboka. rinstitute de Recherche Scientifique, 
Antananarivo, 1 1 pp. 

Dewar, R. E. 1984. Extinctions in Madagascar: the 
loss of the subfossil fauna. Pp. 574-593 in P. S. 
Martin & R. G. Klein, eds., Quaternary ex- 
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Godfrey, L., & M. Vuillaume-Randriamanantena. 
1986. Hapa lemur simus: endangered lemur once 
widespread.— Primate Conservation 7:92-96. 

Goodman, S. M. 1993. A reconnaissance of He Sainte 
Marie, Madagascar: the status of the forest, avi- 
fauna, lemurs and fruit bats.— Biological Con- 
servation (in press). 

James, H. P., & S. L. Olson. 1991. Descriptions of 



VOLUME 106, NUMBER 1 



33 



thirty-two new species of birds from the Ha- 
waiian Islands: Part II. Passeriformes.— Orni- 
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de la faune subfossile de Madagascar. Lemu- 
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Langrand, O. 1990. Guide to the birds of Madagas- 
car. Yale University Press, New Haven, xi + 
364 pp. 

MacPhee, R. 1986. Environment, extinction and Ho- 
locene vertebrate localities in southern Mada- 
gascar.— National Geographic Research 2:441- 
455. 

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

Randriamanantena. 1984. Team finds giant 
lemur skeleton.— Geotimes 29(1): 10-1 1. 

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ossements d'oiseaux provenant des terrains re- 
cents de Madagascar.— Bulletin du Museum 
d'Histoire Naturelle, Paris 1:9-1 1. 

, & . 1879. Histoire physique, naturelle 

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velle d'oiseau de Madagascar. —Bulletin du Mu- 
seum National d'Histoire Naturelle, Paris, 2nd 
series 22:65-66. 

Olson, S. L., & H. F. James. 1991. Descriptions of 
thirty-two new species of birds from the Ha- 
waiian Islands: Part I. Non-Passeriformes. — 
Ornithological Monograph no. 45. 

Ravoavy, F. 1991. Identification et mise en catalogue 
des vertebres non-primates subfossiles et actuels 



des grottes d'Anjohibe, (Majunga), Fouille 1986. 
DEA, Universite d' Antananarivo, Madagascar. 

Simons, E. L., L. R. Godfrey, M. Vuillaume-Randria- 
manantena, P. S. Chatrath, & M. Gagnon. 
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in the Ankarana Mountains of northern Mad- 
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319. 

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of Africa. Vol. II. Academic Press, London, xvi 
+ 552 pp. 

von den Driesch, A. 1976. A guide to the measure- 
ment of animal bones from archaeological 
sites.— Peabody Museum Bulletin no. 1. 

Vuillaume-Randriamanantena, M. 1982. Contribu- 
tions a I'etude des Os Longs des Lemuriens 
Subfossiles Malgaches. Dissertation 3rd cycle, 
Universite d' Antananarivo, Madagascar. 

, L. R. Godfrey, & M. R. Sutherland. 1985. 

Revision of Hapalemur (Prohapalemur) gallieni 
(Standing 1905).-Folia Primatologia 45:89- 
116. 

(SMG) Field Museum of Natural History, 
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tananarivo (101), Madagascar. 



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 



35 



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. 



36 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Description of holotype.— Total length 
(TL), 224 mm; tail length, 7 mm; midbody 
diameter (MBD), 7 mm; body diameter at 
vent, 6 mm; TL/MBD, 32; 318 middorsal 
scale rows; 307 midventral scale rows; 13 
mid-subcaudal scales; 13 mid-dorsocaudal 
scales; 28 scale rows around body anteriorly 
reducing to 24 rows posteriorly; head 
(viewed from above) tapered; in profile, 
snout rounded, projecting anteriorly above 
mouth, without transverse rostral keel; nos- 
trils lateral, near tip of snout; eyes dorso- 
lateral, visible beneath ocular shield im- 
mediately behind posterior edge of 
preocular, diameter of eye 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 



37 




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 



38 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 






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



39 



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40 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



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

Comparisons. —As a member of the 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 



41 



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 



42 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



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

All head scales except the rostral have 
basal glands forming a glandular line which 
is overlapped by the posteriorly projecting 
free edge of the preceding scale; glands lying 
along 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 



43 



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





Sex 


TL 


MD 


MDC 


MV 


MSC 


Tail/TL 




# of scale rows 






Museum # 


28-27 


27-26 


26-25 


25-24 


UF 54186 


F 


244 


457 


12 


447 


11 


0.016 


0.47 (0.52) L 


0.52 (0.55) R 


0.96 R 


0.96 L 


UF 54188 


F 


255 


457 


11 


444 


12 


0.020 


0.91 (0.93) R 


0.92 L 


— 


— 


UF 54189 


F 


255 


460 


11 


448 


10 


0.014 


0.55 (0.75) L 


0.88 R 


— 


— 


UF 54192 


F 


233 


441 


11 


429 


12 


0.017 


0.93 R 


0.93 L 


— 


— 


UF 55644 


F 


227 


458 


11 


448 


10 


0.015 


0.59 (0.65) L 


0.74 (0.78) R 


0.97 


0.97 


USNM 319550 


F 


255 


434 


10 


427 


9 


0.016 


0.58 (0.89) L 


0.93 R 


0.97 R 


— 


UF 55648 


F 


232 


434 


11 


426 


11 


0.017 


0.57 (0.63) L 


0.66 (0.74) R 


— 


— 


UF 52866 


M 


203 


422 


12 


408 


11 


0.020 


— 


0.04 R 


0.91 


0.91 


USNM 319549 


M 


207 


412 


12 


396 


13 


0.024 


0.62 (0.85) R 


0.64 (0.67) L 


— 


— 


UF 55123 


M 


226 


427 


11 


412 


12 


0.018 


0.06 (0.19) R 


0.06 L 


— 


— 


UF 55646 


M 


210 


461 


13 


449 


12 


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 



44 



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 



45 



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 



47 




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 



48 



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 


(5 


26.6 


8.2 


8.8 


13.6 


2.7 


1.5 


3.0 


8.3 


1.4 


1.2 


A-3769 


6 


28.9 


8.6 


9.3 


14.3 


3.1 


1.5 


3.2 


8.6 


1.4 


1.3 


A-3770 


9 


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 



49 



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50 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



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

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 



52 



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 



53 



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- 



54 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



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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 



56 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



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



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 



58 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



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

Methods and materials. —The methods of 
counting and measuring specimens in this 
paper are those outlined in Fink & Weitz- 
man (1974:1-2). Standard length (SL) and 
other body measurements were taken in mm 
and are expressed as percentages of the stan- 
dard length or, in the case of subunits of the 
head, as percentages of the bony head length. 
Ranges of counts include all specimens, with 
the values in square brackets being those of 
the holotype. Counts of total vertebrae are 
from radiographs and include the four ver- 
tebrae of the Weberian apparatus, and the 
fused 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 



59 



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 

44 



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 



7 


6-7 


6-7 


14 


13-14 


13-14 


14 


13-14 


13-14 


14 


13-15 


13-15 


41 


40-41 


40-41 


13 


10-13 


10-13 


7 


6-7 


6-7 



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



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



60 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 1 . 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 



61 



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 



64 



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 



65 



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 


B 


Paratype L-25e 


A ? B * A 


B 


Paratype L-25i 


? A ? * ? 


9 



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 & 



66 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



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

Age and locality. —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 



67 





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. 



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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 



69 





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. 



70 



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 



71 








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



72 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 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:r 


w.r 


Remarks 


13222 


5.5 mm 


34 mm 


2.5 mm 


6:1 


1:2.2 


holotype; DS17 


13223 


8 


26 + 


4.5 


— 


1:1.8 


paratype; DS9 


13224 


7.5 


29+ 


4 


— 


1:1.9 


paratype; DS37 


13225 


5 


24 


2.5 


5:1 


1:2.0 


paratype; DSll, DS24 


13226 


4.5 


15 + 


2 


— 


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 



73 



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 



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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 



75 



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 



76 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



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



Asteroids 



Ophiuroids 



Crinoids 



Primary circlet: 
centrale 



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 



77 



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) 



—(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- 



78 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



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

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 



79 



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- 



80 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



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

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

Oegophiurida Rediagnosed 

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



(cf.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 



81 



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- 



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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 



87 





^^ 



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. 



88 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



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

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

Superfamily Lysiosquilloidea 

Giesbrecht, 1910 
Tetrasquillidae, new family 

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

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

Included genera. —Three: Tetrasquilla 
Manning & Chace, 1990, containing only 
T. mccullochae (Schmitt, 1940), the only 
known pantropical stomatopod (see Man- 
ning & Chace 1990); Tectasquilla Adkison 
& Hopkins, 1984, containing only Tecta- 
squilla lutzae Adkison & Hopkins, 1984, 
known only from off" northwestern 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 



89 




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- 



90 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




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




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



VOLUME 106, NUMBER 1 



91 



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

Literature Cited 

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

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

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

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

Lamarck, J. B. P. A. de. 1818. Histoire naturelle des 
animaux sans vertebres, presentant les carac- 
teres generaux et particuliers de ces animaux, 
leur distribution, leurs classes, leurs families, 
leurs genres, et la citation des principales especes 
qui s'y rapportent; precedee d'une introduction 
offrant la determination des caracteres essentiels 
de I'anomas, sa distinction du vegetal et des 
autres corps naturelles, enfin, I'exposition des 
principes fondamentaux de la zoologie 5:612 
pp., Deterville, Paris. 

Latreille, P. A. 1803. Histoire naturelle, generale et 
particuliere, des Crustaces et des Insectes 3:467 
pp., F. Dufart, Paris. 



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

. 1970. Two new stomatopod crustaceans from 

Australia. — Records of the Australian Museum 
28(4):77-85. 

. 1 980. The superfamilies, families, and genera 

of Recent stomatopod Crustacea, with diagno- 
ses of six new families.— Proceedings of the Bi- 
ological Society of Washington 93:362-372. 

,& A. J. Bruce. 1984. Erythrosquillamegalops, 

a remarkable new stomatopod from the western 
Indian Ocean.— Journal of Crustacean Biology 
4:329-332. 

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

Milne Edwards, A. 1869. Rade de Saint- Vincent du 
Cap-Vert (supplement). Pp. 136-138, pi. 17 in 
L. de Folin & L. Perier, 1867-1872, Les fonds 
de la mer, etude intemationale sur les particu- 
larites nouvelles des regions sous-marins, 1, 
Bordeaux. 

Moosa, M. K. 1986. Stomatopod Crustacea. Resul- 
tats du Campagnes MUSORSTOM I & II Phil- 
ippines, 2.— Memoires du Museum National 
d'Histoire Naturelle, Paris (Series A, Zoology) 
133:367-414, pi. 1. 

. 1991. The Stomatopoda of New Caledonia 

and Chesterfield Islands. Pp. 149-219 in B. 
Richer de Forges, ed., Le benthos des fonds 
meubles des lagons de Nouvelle-Caledonie, 1, 
Editions de I'ORSTOM, Paris. 

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

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



93 



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 



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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 



95 




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



96 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



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

Juxtafabia muliniarum (Rathbun, 1918), 
new combination 

Pinnotheres muliniarum Rathbun, 1918:81, 
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 



97 




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



mm. 



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

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



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

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




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



98 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




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 



99 




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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contenant leur description et leurs moeurs, avec 
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is, 1:1-258, pis. 1-8. 



Bousquette, G. D. 1980. The larval development of 
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Biirger, O. 1895. Ein Beitrag zur Kenntniss der Pin- 
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Campos, E. 1989a. Comments on taxonomy of the 
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. 1989b. Tumidotheres a new genus for Pin- 
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. 1990. Calyptraeotheres, dinQvj gtnxxs of Vin- 

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, & H. Griffith. 1990. Clypeasterophilus, a new 

genus to receive the small-palped species of the 
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Campos-Gonzalez, E. 1988. New molluscan hosts for 
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Christensen, A. M., & J. J. McDermott. 1958. Life- 
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Dana, J. D. 1851. On the classification of the Crus- 
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De Man, J. G. 1889. Ueber einige oder seltene Indo- 
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Glassell, S. A. 1933. Description of five new species 
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. 1934. Affinities of the brachyuran fauna of 

the Gulf of California.— Journal of Washington 
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. 1935. New or little known crabs from the 

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Green, T. M. 1985. Pinnotheres jamesi synonymizcd 
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Holmes, S. J. 1894. Notes on the West American 
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Konishi, K. 1983. Larvae of pinnotherid crabs (Crus- 
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Linnaeus, C. 1767. Systema naturae per regna tria 
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Lockington, W. N. 1877. Description of a new genus 
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Rathbun, M. J. 1893. Scientific results of explora- 
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Sakai, T. 1934. Species of the genus Pinnixa (Pin- 
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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, 



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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 



no 



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 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




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 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



(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 



116 



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 



118 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




A: 0.20mm 



B: 0.10mm 
C-G: 0.20mm H, I: 0.20mm 



mm 



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 



120 



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. 

Literature Cited 

Arimoto, I. 1976. Taxonomic studies of caprellids 
(Crustacea, Amphipoda, Caprellidae) found in 
the Japanese and adjacent waters. — Special 
Publications from the Seto Marine Biological 
Laboratory, Series III:i-v + 1-229. 

. 1978. Nippon no Warekara [The Caprellidea 

of Japan]. —Dobutsu to Shizen [Animal and Na- 
ture] 8:10-15 (in Japanese). 

Boeck, A. 1872. Bidrag til Califomiens Amphipo- 
defauna.— Forhandlinger i Videnskabs-selska- 
bet i Christiania: 32-51. 

Dougherty, E. C, & J. E. Steinberg. 1953. Notes on 
the skeleton shrimps (Crustacea: Caprellidae) of 
California.— Proceedings of the Biological So- 
ciety of Washington 66:39-49. 

International Commission on Zoological Nomencla- 
ture. 1985. Pp. i-xx + 1-338 z>7 W. D. L. Ride 
et al., eds., International code of zoological no- 
menclature. Third edition adopted by the XX 
general assembly of the International Union of 
Biological Sciences. University of California 
Press, Berkeley and Los Angeles, U.S.A. 

Kim, H. S., & K. S. Lee. 1975. Faunal studies on the 
genus Caprella (Crustacea: Amphipoda: Ca- 



VOLUME 106, NUMBER 1 



121 



prellidae) in Korea.— Korean Journal of Zool- 
ogy 18:115-126. 

Laubitz, D. R. 1970. Studies on the Caprellidae 
(Crustacea, Amphipoda) of the American North 
Pacific— National Museums of Canada, Pub- 
lications in Biological Oceanography 1:1-89. 

. 1972. The Caprellidae (Crustacea, Amphip- 
oda) of Atlantic and Arctic Canada.— National 
Museums of Canada, Publications in Biological 
Oceanography 4:1-82. 

Lee, K. S. 1988. Fauna of Caprellidae (Amphipoda) 
of Cheju Island and its adjacent waters, Ko- 
rea.— The Korean Journal of Systematic Zool- 
ogy, Special Issue 2:97-106. 

Marelli, D. C. 1981. New records for Caprellidae in 
California, and notes on a morphological vari- 
ant of Caprella verrucosa Boeck, 1871.— Pro- 
ceedings of the Biological Society of Washington 
94:654-662. 

Martin, D. M. 1977. A survey of the family Caprel- 
lidae (Crustacea, Amphipoda) fi-om selected sites 
along the northern California Coast.— Bulletin 
of the Southern California Academy of Sciences 
76:146-167. 

McCain, J. C. 1968. The Caprellidae (Crustacea: Am- 
phipoda) of the Western North Atlantic. —Unit- 
ed States National Museum Bulletin 278:i-vi + 
1-147. 

, & J. E. Steinberg. 1970. Amphipoda I. Ca- 

prellidea I. Fam. Caprellidae. Pp. 1-78 in H.- 
E. Gruner & L. B. Holthuis, eds., Crustaceorum 
Catalogus, Pars 2. Dr. W. Junk Publishers, The 
Hague, The Netherlands. 



Takeuchi, I. 1989. Taxonomic and ecological studies 
of the Caprellidea (Crustacea, Amphipoda) in- 
habiting the Sargassum zone. Doctoral thesis. 
Faculty of Agriculture, The University of To- 
kyo, Tokyo, 244 pp (in Japanese). 

, & R. Hirano . 1991. Growth and reproduction 

of Caprella danilevskii (Crustacea: Amphipoda) 
reared in the laboratory.- Marine Biology 1 10: 
391-397. 

Utinomi, H. 1943. Caprellids obtained in Onagawa 
Bay, northern Japan. — Science Reports of To- 
hoku University, Series 4. Biology 17:271-279. 

. 1947. Caprellidae of Japan and adjacent wa- 
ters.— Seibutsu Supplement 1:68-82 (in Japa- 
nese). 

. 1964. Caprellidea. Pp. 11-15, pis 1-3 in T. 

Kikuchi, ed., Fauna and flora of the sea around 
the Amakusa Marine Biological Laboratory. Part 
V. Amphipod Crustacea. Amakusa Marine Bi- 
ological Laboratory, Kyusyu University, Ku- 
mamoto, Japan (in Japanese). 

Vassilenko, S. V. 1974. [Caprellids of the seas of the 
USSR and adjacent waters.] — Opredeleliteli po 
Faune SSSR 107:1-288 (in Russian). 

Otsuchi Marine Research Center, Ocean 
Research Institute, The University of To- 
kyo, Akahama, Otsuchi, Iwate 028-11, Ja- 
pan. 



PROC. BIOL. SOC. WASH. 
106(1), 1993, pp. 122-130 

A NEW SPECIES OF KALLIAPSEUDES 
(CRUSTACEA: TANAIDACEA: KALLIAPSEUDIDAE) 

FROM TRINIDAD 

Roger N. Bamber 

Abstract. —A new species of kalliapseudid tanaidacean, Kalliapseudes sonia- 
dawnae, is described from three specimens collected from the silty-clay benthos 
in 8-10 m of water in the Gulf of Paria, off the northwest coast of Trinidad. 
The new species is similar to congeners from the Bahamas and the Pacific coast 
of Mexico, but can be distinguished by its stouter limb and antennal articles, 
and elongate dactyli of pereopods 2 and 3. It is unique in bearing a stout 
pectinate spine on the basis of pereopod 1 . 



Kalliapseudid tanaidaceans have been 
described previously from the Atlantic 
coasts of Central and South America. Lang 
(1956) established the family and described 
several species from Brazil, while most re- 
cently Sieg (1982) described Kalliapseudes 
(Mesokalliapseudes) bahamensis from the 
Caribbean. In Trinidad waters, Bacescu & 
Gutu (1975) described Discapseudes suri- 
namensis from the Caroni Swamp on the 
northern west coast. The present material 
was collected in 1982 during a study of the 
benthos off the Port-of-Spain coastal area, 
Trinidad, in the Gulf of Paria (Agard 1984). 
Three specimens of a hitherto undescribed 
kalliapseudid were taken from two separate 
sites. 

With this limitation on the quantity of 
material, only one specimen was dissected 
and mounted for microscopical examina- 
tion; the holotype (which was missing its 
antennae) and one male paratype were ex- 
amined whole; consequently features of the 
detailed anatomy (e.g., the mouthparts) re- 
late only to the success of the single dissec- 
tion and details of, for example, the epig- 
nath must await further material. The 
conspicuous distinction of the new species 
does not, however, depend on any such sub- 
tleties. 

The specimens had little sclerotization and 



long setae were commonly not straight. All 
figures were drawn with the aid of the cam- 
era lucida and represent the anatomy as it 
exists, rather than any stylized symmetrical 
interpretation. 

The type material is lodged at the Na- 
tional Museum of Wales, Cardiff, UK 
(NMW). 

Description 

Order Tanaidacea Hansen, 1895 

Suborder Apseudomorpha Sieg, 1980 

Family Kalliapseudidae Lang, 1956 

Kalliapseudes {Mesokalliapseudes) 

soniadawnae, new species 

Material.— One 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 



132 



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 


Endopodite longer 


In the first, second 


Endopodite approxi- 


Endopodite: 


than exopodite 


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 


Equal 


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 


With apparent 




from urosome 


articulations 


articulations 


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 


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 


1:5 


1:4 


4:5 


bidens, kodiakensis. 










reedi, wilsonae 


1:5 


1:6 


1:4 


4:5 


Males 










shawangunkensis 


1:3 


— 


0:3 


0:3 


subgracilis 


1:3 


— 


9 


0:4 


amabilis 


1:3 


— 


0:3 


0:4 


reedi, wilsonae 


1:4 


— 


0:3 


0:4 


bidens 


1:4 


— 


2- 


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 



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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- 



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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, 



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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). 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




300jim 




^1 



W/ V-f I 

1 10^ 




m 



D 



. ,.,,^. 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, 



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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 



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■C-e: 

09 






P'C 



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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 



8- 

1- 



QOO 



o 



I I I I — I I I I I I — I — r 

10 15 



20 



-I— I — I I I I I I 
25 



30 



35 



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 



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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^: . ^ 


L 




1 

r * -^ 


%^i^'sfe^^ 


rw 


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 



8- 

7- 












o o° 


^-^6- 














E 












o 


£5- 










o 














o 




.C 4- 










o 




ts- 








o 


o^ 
















0) 














_J 2- 














0- 














c 


1 1 1 1 1 1 1 
5 


1 1 1 1 1 

10 


1 1 1 1 1 
15 


1 1 1 1 

20 


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. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




WW 


^-^^H 


^^feb ' j^^sSMMHt^ 


• \&- § jM*'^ ^^^^1 


^'^g 


9^ 


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 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




35 



40 



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 



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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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Seas of the U.S.S.R.-Academiya Nauk SSSR, 
Keys to the Fauna of the SSSR 56:1-433 (trans- 
lated from Russian by the Israel Program for 
Scientific Translations, Jerusalem 1965). 



VOLUME 106, NUMBER 1 



181 



Webster, H. E., & J. E. Benedict. 1887. The Annelida 
Chaetopoda, from Eastport, Maine.— Report of 
the United States Commissioner of Fisheries 
1885:707-755. 

Willey, A. 1902. Polychaeta. Pp. 262-283 in Report on 
the collections of natural history made in the Ant- 
arctic regions during the voyage of the "Southern 
Cross." William Qowes and Sons, Limited, Lon- 
don, 344 pp. 

Swedish Museum of Natural History, 
Stockholm, and (postal address): Tjamo 
Marine Biological Laboratory, PL 2781, 
S-452 96 Stromstad, Sweden. 

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. 





SH 


HBW 


sw 


AH 


AW 


WH 


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 




b 




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. 



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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, 



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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 



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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- 



I 



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. 



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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) 



10 


36 


-0.85 


-2 


1^ 


4 


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- 



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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 

INTERNATIONAL COMMISSION ON ZOOLOGICAL 

NOMENCLATURE 

% The Natural History Museum 
Cromwell Road 
London, SW7 5BD, U.K. 
Tel. 071-938 9387 

Applications published in the Bulletin of Zoological Nomenclature 

The following Applications were published on 30 September 1992 in Vol. 49, 
Part 3 of the Bulletin of Zoological Nomenclature. Comment or advice on these 
Applications is invited for publication in the Bulletin, and should be sent to the 
Executive Secretary, I. C.Z.N. , % The Natural History Museum, Cromwell Road, 
London SW7 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. 



INFORMATION FOR CONTRIBUTORS 

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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 



1- 



PROCEEDINGS 

OF THE 

BIOLOGICAL SOCIETY 

OF 

WASHINGTON 



VOLUME 106 
1 1 JUNE 1993 



NUMBER 2 



ISSN 0006-324X 



THE BIOLOGICAL SOCIETY OF WASHINGTON 

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President-elect: Janet W. Reid 



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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 



o 









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 

Diagnosis. —Prepharynx shorter than 
pharynx* (21); one cecum atrophied* (26); 
vitelline follicles not extending anteriorly to 
posterior margin of seminal vesicle* (32); 
subterminal mouth* (36); no oral spines* 
(38); no prepharynx (40); secondary group 
of vitelline follicles surrounding testes (41). 

Acknowledgments 

We gratefully acknowledge the loan of 
specimens from Doctora Margarita Ostrow- 
ski de Nunez, Buenos Aires, Argentina. This 
study was supported by funds from oper- 
ating grant A7696 from the Natural Sci- 
ences and Engineering Research Council of 
Canada (NSERC) to DRB. 

Literature Cited 

Archie, J. W. 1989. Homoplasy excess ratios: new 
indices for measuring levels of homoplasy in 
phylogenetic systematics and a critique of the 
consistency index.— Systematic Zoology 38:253- 
269. 

Baugh, S. C. 1957. Contributions to our knowledge 
of digenetic trematodes. II. — Proceedings of the 
National Academy of Science of India 26:295- 
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VOLUME 106, NUMBER 2 



219 



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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 



1 








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 
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Dall, W. H. 1908. [Reports on the dredging opera- 
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in the Gulf of California, in charge of Alexander 
Agassiz, carried on by the U.S. Fish Commis- 
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Commander Z. L. Tanner, U.S.N.. command- 
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224 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Commission Steamer "Albatross," from Octo- 
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L. M. Garrett, U.S.N., commanding. XIV] The 
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Kaicher, S. D. 1978. Card catalogue of world-wide 
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1859-1964. 

Olsson, A. A., & F. M. Bayer. 1972. Gerdiella, a. new 
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Thiele, J. 1925. Gastropoda der Deutschen Tiefsee- 
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Department of Invertebrate Zoology, 
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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 



*» 




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 




B 



/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 




B 



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. 



I 



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 












I 



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 




19 

C. branchiophilus 




eps 





4 

8 
9 



CD 




13 



14 



17 



18 



21 



fl 



22 



Figs. 2-22. 2-5, Cambarincola acudentatus Holt, 2, ventro-lateral view; 6-10, Cambarincola alienus Holt; 
11-14, Cambarincola barbarae Holt; 15-18, Cambarincola bobbi Holt; 19-22, Cambarincola branchiophilus 
Holt. 



VOLUME 106, NUMBER 2 



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 




35 

C. demissus 




C. dubius 





25 



26 





31 



32 




39 




40 



45 



46 



27 
28 



33 
34 



41 
42 



47 



^ 



48 



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 



<] 



52 




57 




58 




63 



64 




74 



a 



tp 



54 



c^ 



59 




60 




65 




66 




70 



75 



O 



76 



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 





81 



<7 



82 




C. holostoma 




C. hoiti 




C. illinoisensis 





85 




86 




eps 




C. ingens 





d 



87 



92 



93 



X7 



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 




V 



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 



O 



147 




133 



^ 



134 






148 



t3 



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 







£>> 



58 



^ 



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 




O 



174 176 



175 




^ 



177 




C. restans 




179 —^ /-^ 




C. serratus 




C. sheltensis 



185 




182 



^ 



186 



183 O^ 



^J 188 



< 



< 



xO 



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 



Q 



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 




t? 



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 




d 



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 



D 



280 





283 



284 







y 



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. 



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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 



<i 





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 



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S. veracruzicus 



349 

S. wardinus 




T. friaufi 



340 





345 346 

S. villalobosi 





350 





354 



355 



341 



338 







Cb 



343 



/::> 



342 344 



347 



348 



V 



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 



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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 



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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 

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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Cambarincola Ellis. 1912. (Oligochaeta. Bran- 
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chiobdellidae (Oligochaeta).— American Mid- 
land Naturalist 64:169-176. 
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-. 1967b. Status of the genera Branchiobdella 
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-. 1968c. The genus Pterodrilus (Annelida: 
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-. 1973a. Epigean branchiobdellids (Annelida: 
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-. 1973c. Branchiobdellids (Annelida: Clitel- 
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-. 1978. The reassignment of Cambarincola 
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-. 1981b. New species oi Sathodrilus HoU, 1 968, 
(Clitellata: Branchiobdellida) from the Pacific 
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-. 1982. A new species of the genus Camba- 
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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) 



J 



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 



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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 



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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.). 



304 



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- 



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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 





25 




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 



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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 








1 


4th joint: ventral 


1 


1 


2 


6th joint 


1 








7th joint: ventral 


1 


1 


2 


Mandible 








Basale 


13 


12? 


11 


Endopodite 








2nd joint: ventral 


4 


3 


4 


2nd joint: dorsal 


2 


3 


2 


3rd joint 


7 


6 


7 


Maxilla 








Basale: dorsal 





1? 


1 


Basale: ventral 


1 


2? 


1 


Endopodite 








1st joint: anterior 


4 


5 


5 


1st joint: posterior 


2 


2 


2-3 


Fifth limb 








Epipodite 


13 


nd 


14 


Protopodite + 








endopodite 


17 


nd 


19 


Exopodite 








1st joint: ventral 


8 


nd 


9 


Sixth limb 








Epipodite 


ca. 12 


nd 


15 


Furca 








(no. of short claw- 








like processes) 


7 


7 


6-7 



tral, 2 dorsal; terminal endopodial joint of 
male without hook-like process. Mandible: 
one of male endopodite terminal bristles 
wavy bent and with long spine-like pro- 
cesses in the middle. Maxilla: second en- 
dopodite joint with 6 bristles. Fifth limb: 
second exopodial joint without terminal 
bristle on ventral margin; third exopodial 
joint with 3 bristles. Sixth limb: process on 
dorsal 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 



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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 




1 

1 






E 


' 


E 

o 


, 




' i ' 


E 



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). 



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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 



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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 



326 



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 


s 


2 


Juv. 


Maximum 
length 
(mm) 


Mid-depth 
(m) 


Location 


2139-1 








7 


18.0 


790 


summit 


4 


1 


2 


1 


26.5 


852 


summit 


5 


1 


7 


2 


22.5 


790 


summit 


6 


2 


4 


— 


21.2 


810 


summit 


8* 


5 


8 


1 


25.4 


852 


summit 


2140-1 


2 


2 


— 


19.8 


804 


summit 


2 


— 


4 


— 


20.0 


793 


summit 


3 


8 


4 


3 


23.0 


815 


summit 


4 


1 


2 


— 


18.8 


874 


summit 


5 


1 


4 


— 


21.5 


804 


summit 


6 


— 


3 


4 


18.3 


793 


summit 


7 


4 


7 


7 


23.9 


817 


summit 


8 


1 


12 


13 


18.7 


876 


summit 


2142-1 


3 


4 


3 


23.0 


797 


summit 


2 


1 


4 


— 


21.4 


782 


summit 


3 


10 


17 


4 


23.4 


788 


summit 


4 


26 


72 


116 


25.0 


797 


summit 


5 


6 


8 


6 


22.2 


797 


summit 


7 


6 


18 


6 


22.5 


788 


summit 


8* 


9 


56 


35 


21.5 


808 


summit 


2143-2 


2 


1 


1 


22.1 


1300 


flank 


3 


— , 


3 


3 


— 


1297 


flank 


4* 


— 


1 


1 


— 


1308 


flank 


5 


— 


3 


2 


20.2 


1300 


flank 


8 


3 


6 


6 


22.0 


1309 


flank 


2144-1 


— 


1 


— 


— 


1289 


flank 


3 


— 


1 


1 


14.3 


1179 


flank 


4 


1 


1 


— 


— 


1250 


flank 


7 


1 


— 


— 


20.8 


1195 


flank 


8* 


— 


1 


3 


17.5 


1259 


flank 


2145-3 


— 


— 


2 


9.6 


1323 


flank 


4* 


2 


4 


1 


22.5 


1352 


flank 


8 


1 


1 


— 


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 



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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). 



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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 






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 


V; 


^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 
Macrouren.— Sitzungs Berichte der Akademie 
Wissenschaften in Wiens 45:389^26. 

. 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 
44:1-296. 

. 1966. A collection of freshwater prawns 

(Crustacea, Decapoda, Palaemonidae) from 
Amazonia, Brazil, collected by Dr. G. Mar- 
lier.— Bulletin Institute Royale Sciences Natu- 
relle, Belgian 42:1-1 1. 

Kensley, B., & I. Walker. 1982. Palaemonid shrimps 
from the Amazon Basin, Brazil (Crustacea: De- 
capoda: Natantia). — Smithsonian Contribu- 
tions to Zoology 362:iii + 1-28. 

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 



VOLUME 106, NUMBER 2 



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).— 
Sociedade Brasileira para o Progresso da Cien- 
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- 
cean Biology 2:378-391. 

, & G. Pereira. 1992. New species, cladistic 

relationships, and biogeography of the genus 
Fredius (Decapoda: Brachyura: Pseudothelphu- 
sidae) from South America.— Journal of Crus- 
tacean Biology 12:298-311. 

Shokita, S. 1973. Abbreviated larval development of 
the freshwater prawn, Macrobrachium shokitai 
Fujino et Baba (Decapoda, Palaemonidae) from 



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- 
monidae. — Bulletin de Museum de Histoire Na- 
turelle, Paris 17:12-16. 

. 1923a. Le developpement larvaire des Pa- 

laemoninae. I. Partie descriptive. La conden- 
sation progressive de I'ontogenese. — Bulletin 
Biologique de la France et de la Belgique 57: 
510-603. 

. 1923b. Recherches sur I'embryogenie des 

Crustaces Decapodes de la sous-famille des Pa- 
laemoninae.— Bulletin Biologique de la France 
et de la Belgique, supplement 5, 234 pp. 

Tiefenbacher, L. 1978. Zur systematik und Verbrei- 
tung der Euryrhynchinae (Decapoda, Natantia, 
Palaemonidae). — Crustaceana 35:177-189. 

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 




6 



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) 








BV 


^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 




95 



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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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 


19 


20 


21 


22 


23 


24 


25 


26 


27 


28 


29 


30 


n 


X 


H. meeki 






























MA to GA 




2 


- 


- 


- 


2 


11 


53 


80 


33 


8 




189 


26.8 


E. coast FL 




2 


4 


8 


2 


2 


11 


29 


40 


10 


6 




114 


25.9 


W. penin. FL 












12 


50 


60 


40 


14 


1 




177 


26.0 


FLA pan. to TX 








1 


4 


11 


26 


44 


53 


10 


5 


2 


156 


26.2 


Yucatan 










1 


3 


6 


10 


5 


2 






27 


25.5 


Species total 




4 


4 


9 


6 


30 


104 


196 


218 


69 


20 


2 


662 


26.2 


H. unifasciatus 






























Florida 




2 


7 


10 


18 


17 


5 












59 


22.9 


West Indies 


2 


- 


3 


11 


25 


46 


37 


11 


5 


1 






141 


24.1 


Cent. America 






2 


5 


10 


11 


6 


- 


1 








35 


23.5 


Carib. S. Am. 








6 


27 


33 


23 


24 


1 








114 


24.3 


South America 






5 


16 


31 


33 


21 


2 


1 








109 


23.5 


Yucatan 






1 


1 


5 


4 


1 












12 


23.3 


Bermuda 




5 


34 


37 


17 


5 


2 












100 


21.9 


Species total 


2 


7 


52 


86 


133 


149 


95 


37 


8 


1 






570 


23.5 



head length with 83% of 240 specimens 
greater than or equal to 1.0. Ratio of LJL 
to SL 0.20 to 0.35 with 95% of 240 speci- 
mens 0.22 to 0.30. PREORB to ORB ratio 
0.61 to 1.0. usually 0.70 to 0.90 (92% of 
265 specimens). Distance from base of an- 
teriormost pelvic ray to caudal base extends 
anteriorly to mid-eye, usually between pos- 



terior portion of eye and posterior margin 
of opercular membrane. Dorsal- and anal- 
fin bases about equal, ratio of ABASE to 
DBASE 0.83 to 1.07, mean of 0.96 for 265 
specimens. Origin of dorsal fin over that of 
anal fin. Bases of dorsal and anal fins cov- 
ered with scales. Distance from anterior- 
most pectoral ray to origin of pelvic fin less 



Table 3.— Numbers of dorsal-fin rays in populations of Hyporhamphus meeki and H. unifasciatus. 



Population 


12 


13 


14 


15 


16 


17 


n 


X 


H. meeki 


















MA to GA 




2 


100 


56 


1 


1 


160 


14.4 


E. coast FL 




8 


40 


34 


3 




85 


14.4 


W. penin. FL 






52 


95 


10 




157 


14.7 


FL pan. to TX 


1 


5 


90 


65 


4 




165 


14.4 


Yucatan 






15 


12 






27 


14.5 


Species total 


1 


15 


297 


262 


18 


1 


594 


14.5 


H. unifasciatus 


















Florida 






14 


28 


6 




48 


14.8 


West Indies 




1 


42 


77 


7 




127 


14.7 


Cent. America 






6 


35 


3 




44 


14.9 


Carib. S. Am. 






16 


92 


10 




118 


14.9 


South America 




1 


32 


76 


11 




120 


14.8 


Yucatan 






1 


9 


2 




12 


15.1 


Bermuda 




5 


48 


27 


2 




82 


14.3 


Species total 




7 


159 


344 


41 




551 


14.8 



376 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 4.— Numbers of anal-fin rays in populations of Hyporhamphus meeki and H. unifasciatus. 



Population 


14 


15 


16 


17 


18 


n 


X 


H. meeki 
















MA to GA 


1 


18 


107 


36 


1 


163 


16.1 


E. coast FL 


1 


31 


43 


10 




85 


15.7 


W. penin. FL 




13 


98 


45 


1 


157 


16.2 


FL pan. to TX 


10 


59 


80 


16 




165 


15.6 


Yucatan 




3 


21 


3 




27 


16.0 


Species total 


12 


124 


349 


110 


2 


597 


15.9 


H. unifasciatus 
















Florida 






25 


22 


1 


48 


16.5 


West Indies 




15 


83 


29 


1 


128 


16.1 


Cent. America 




4 


27 


13 




44 


16.2 


Carib. S. Am. 




8 


63 


47 




118 


16.3 


South America 


1 


9 


63 


45 


1 


119 


16.3 


Yucatan 






10 


2 




12 


16.2 


Bermuda 




2 


21 


46 


12 


81 


16.8 


Species total 


1 


38 


292 


204 


15 


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 


9 


10 


11 


12 


13 


n 


X 


H. meeki 
















MA to GA 




2 


75 


59 


3 


158 


11.4 


E. coast FL 




19 


70 


17 




106 


11.0 


W. penin. FL 






49 


98 




147 


11.7 


FL pan. to TX 




1 


77 


41 


1 


120 


11.4 


Yucatan 






18 


9 




27 


11.3 


Species total 




22 


289 


224 


4 


558 


11.4 


H. unifasciatus 
















Florida 




19 


45 


5 




69 


10.8 


West Indies 




39 


94 


3 




136 


10.7 


Cent. America 


2 


2 


35 


2 




41 


10.9 


Carib. S. Am. 




9 


108 


3 




120 


11.0 


South America 




14 


104 


4 




122 


10.9 


Yucatan 






11 


1 




12 


11.1 


Bermuda 


3 


104 








107 


10.0 


Species total 


5 


187 


397 


18 




607 


10.7 



VOLUME 106, NUMBER 2 



377 



Table 6.— Summary of morphometric data in percent standard length in populations oi Hyporhamphus meeki, 
except for SL in mm. 









Atlantic 










( 


Gulf 






%SL 


n 


Min 


Max 


Mean 


SD 


SE 


n 


Min 


Max 


Mean 


SD 


SE 


SL 


169 


48.1 


179 


96.7 


27.16 


2.09 


124 


63.6 


176 


128.9 


23.81 


2.14 


P>-P2 


169 


31.8 


37.8 


34.5 


1.01 


0.08 


124 


31.7 


38.1 


34.6 


1.09 


0.10 


P.-C 


169 


39.2 


46.2 


43.0 


1.04 


0.08 


124 


41.3 


45.7 


43.6 


0.91 


0.08 


LJL 


161 


20.6 


35.1 


27.0 


2.32 


0.18 


107 


20.5 


29.8 


23.9 


1.88 


0.18 


HDL 


169 


19.1 


34.4 


24.4 


1.27 


0.10 


124 


21.9 


26.0 


23.7 


0.76 


0.07 


UJL 


169 


2.2 


4.8 


4.2 


0.32 


0.02 


124 


3.7 


4.8 


4.3 


0.23 


0.02 


UJW 


169 


4.3 


6.2 


5.2 


0.32 


0.02 


124 


4.7 


5.8 


5.2 


0.21 


0.02 


BD-P, 


169 


8.6 


13.1 


11.7 


0.79 


0.06 


124 


10.1 


13.3 


11.9 


0.62 


0.06 


BD-P2 


169 


7.2 


15.9 


11.4 


1.62 


0.12 


124 


9.0 


15.3 


12.9 


1.20 


0.11 


ABASE 


169 


12.3 


17.3 


14.4 


0.79 


0.06 


124 


12.5 


16.0 


13.9 


0.72 


0.06 


DBASE 


169 


13.3 


16.9 


14.8 


0.67 


0.05 


124 


13.2 


16.4 


14.6 


0.67 


0.06 


P,L 


161 


12.3 


16.4 


14.6 


0.72 


0.06 


109 


13.1 


16.3 


14.5 


0.64 


0.06 


ORB 


169 


4.8 


7.1 


5.9 


0.41 


0.03 


124 


5.0 


6.9 


5.7 


0.39 


0.03 


PREORB 


169 


3.4 


6.1 


4.5 


0.28 


0.02 


124 


3.4 


5.2 


4.2 


0.30 


0.03 



have been described (as Hyporhamphus sp.) 
from Chesapeake Bay, which may indicate 
utihzation of estuarine waters as nursery ar- 
eas (Hardy & Johnson 1974). Larvae were 
collected along the Gulf coast of Florida 
most frequently during spring and summer 
in less than 30 m of water (Houde et al. 
1979). The particulars of spawning are un- 
known. Eggs have been attached to floating 
Zostera blades (by their adhesive filaments) 
over vegetated habitats during summer 
months in Chesapeake Bay (Olney and 
Boehlert 1988). 

Distribution. —Atlantic coast of the Unit- 
ed States from Miami, Florida to Cape Cod, 
Massachusetts and rarely north to Cham- 
cook, Passamaquoddy Bay, New Brunswick 
(Leim & Day 1959), and in the Gulf of Mex- 
ico from the Everglades to Galveston, Texas 
(Fig. 1). Also occurs in Yucatan. The ob- 
served water temperature range is 13.7 to 
34.9°C, so H. meeki has a subtropical to 
temperate distribution. Sympatric with H. 
unifasciatus on the east coast of Florida from 
St. Lucie Inlet south to Miami and on the 
west coast from the Everglades to Tampa 
Bay. 

Etymology . —^diVnQd after Seth E. Meek 
who first separated the two species we rec- 



ognize here (Meek and Goss 1884:223) . . . 
"all the specimens . . . thus far taken on the 
Atlantic coast of the United States north of 
the Florida Keys . . . belong to a species 
differing from the West Indian unifasciatus, 
in the slendemess of body and in the greater 
length of the lower jaw," but misapplied the 
name H. roberti to the northern species. 

Comparisons. — Meristic characters allow 
for statistical separation of H. meeki from 
H. unifasciatus (alpha = 0.05; Tables 8-12), 
though infraspecific variation exists within 
populations of both species. Hyporhamphus 
meeki is discemable from the southern spe- 
cies H. unifasciatus in usually having more 
gill rakers on both the first and second arch- 
es (Tables 1 and 2). Ninety two percent of 
the 780 H, meeki specimens examined have 
total RGR, counts from 33 to 40, whereas 
in H. unifasciatus 91% of 657 specimens 
examined have total RGR, counts from 28 
to 32. Second arch gill rakers in 76% of 662 
specimens of//, meeki range from 26 to 29, 
whereas 92% of 570 specimens of//, uni- 
fasciatus range from 19 to 25. Pectoral fin 
usually with 11 or 12 rays, 10 or 11 in //. 
unifasciatus. 

Regressions of morphometries also allow 
us to distinguish the two species. This is best 



378 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



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f~~ P-, r-, —I — i — — _- 



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nC f^. r*". — fN — — — ' 



nCnC>c— "NCNCNCNCNOmvCf^lNOm 

ococococococococococccocococ 



f^, [^moNfN-^iri-<;tTt(Nr-(N-^m 

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— mrj-fNfN — — _,___ 

f^i f^J O O nC oc oc m — • r^, in r~- f^i 

m od r~" — nC •^ m rn in r- nC nC nc' -"^ 

ONen^renf^j _______ 



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y: — Cl. _: Z _^ ^ 



^ - T < < 

Z^ r\ /^ f-^ ^ 

-' ~ S < C 



illustrated in the preorbital on orbit rela- 
tionship (Fig. 3). The slopes of the regres- 
sions are significantly different {P > F > 
.001). Ratio of preorbital length to orbit di- 
ameter is usually greater than 0.70 in H. 
meeki (92% of 265 specimens examined), 
but less than 0.70 in H. unifasciatus (75% 
of 224 specimens). Lower jaw length on 
standard length tends to be greater in H. 
meeki (Fig. 4). however, there is consider- 
able infraspecific variation within popula- 
tions of the two species especially in H. un- 
ifasciatus. 

Geographic variation in morphology is 
not reported in detail herein. Analysis to 
date indicates that morphology of Atlantic 
and Gulf populations of H. meeki differs 
slightly. Southern Florida and Yucatan ap- 
pear to be areas of sympatry between H. 
meeki and H. unifaciatus. Collection data 
indicate that sympatr>' of the two species in 
Rorida may be largely avoided temporally, 
because both species migrate northward up 
the Florida coasts when the waters warm 
during summer and autumn; consequently 
H. unifasciatus moves into areas occupied 
by H. meeki in the winter. The dynamics 
of their sympatn,' in Yucatan is probably 
different than in peninsular Rorida. Indi- 
vidual collections from Yucatan (UMMZ 
143085 and MCZ 32881) contain both spe- 
cies. Yucatan H. yneeki have a preorbital/ 
orbit ratio tvpical of H. unifasciatus. Yu- 
catan may be an area of hybridization or 
introgression. and needs further study. 

Comparisons of meristic means by the 
Tukey-Kramer method indicate some geo- 
graphic variability in populations of both 
H. meeki and H. unifasciatus. There ap- 
pears to be little consistency in trends in H. 
meeki populations across the different me- 
ristic characters. The only trend with any 
consistency is the position of the west coast 
of the peninsula of Florida population. It is 
usually at the extreme of the range of means, 
being closest to mean values of H. unifas- 
ciatus populations. The one trend that stands 
out in populations oiH. unifasciatus is that 



VOLUME 106, NUMBER 2 



379 



Table 8.— Comparison of means for dorsal-fin ray counts for populations of Hyporhamphus meeki and H. 
unifasciatus, and between H. meeki and H. unifasciatus. Populations joined by the same line are not significantly 
different by the Tukey-Kramer method, alpha = 0.05. 



Population 



TK 



H. meeki 






West peninsula of FL 


157 


14.7 1 


Yucatan 


26 


14.5 1 


FL panhandle to TX 


165 


14.4 


Atlantic coast of FL 


85 


14.4 


MA to GA 


160 


14.4 


H. unifasciatus 






Yucatan 


12 


15.1 1 


Caribbean South America 


118 


14.9 1 1 


Central America 


44 


14.9 1 


Rorida 


48 


14.8 1 


South America (Brazil) 


120 


14.8 1 1 


West Indies 


127 


14.7 1 


Bermuda 


82 


14.3 


H. meeki 


594 


14.5 1 


H. unifasciatus 


551 


14.8 1 



Bermuda is always at an extreme of the range 
of means, usually at the lower extreme. 

Material examined. — 33S specimens of 
H. meeki (48.1-179 mm SL) from 50 col- 
lections, and 230 specimens of H. unifas- 



ciatus (62.4-205) from 54 collections, with 
almost complete morphometric and meris- 
tic data are listed. Additional specimens used 
mainly for meristic data are in the second 
author's files. 



Table 9.— Comparison of means for anal-fin ray counts for populations of Hyporhamphus meeki and H. 
unifasciatus, and between H. meeki and H. unifasciatus. Populations joined by the same line are not significantly 
different by the Tukey-Kramer method, alpha = 0.05. 



Population 



TK 



H. meeki 






West peninsula of FL 


157 


16.2 


MA to GA 


163 


16.1 


Yucatan 


26 


16.0 


Atlantic coast of FL 


85 


15.7 


FL panhandle to TX 


165 


15.6 


H. unifasciatus 






Bermuda 


81 


16.8 


Florida 


48 


16.5 


Caribbean South America 


118 


16.3 


South America (Brazil) 


119 


16.3 


Central America 


44 


16.2 


Yucatan 


12 


16.2 


West Indies 


128 


16.1 


H. meeki 


597 


15.9 


H. unifasciatus 


550 


16.4 



380 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



50 



£ 
E 



40 



O 30 

z 

LU 



< 20 

LU 



10 








^i ^ 







50 



100 



150 



200 



250 



STANDARD LENGTH (mm) 



Fig. 4. Relationship of lower jaw length to standard length in Hyporhamphus meeki (squares) and two 
populations of//, unifasciatus, Central and South America (triangles) and West Indies (diamonds). 



Hyporhamphus meeki 

Holotype. -VSNM 294369 (1, 160) 
Morehead City, NC; 5 Sept 1964. 

ATLANTIC U.S. (paratypes): USNM 
132257 (4, 55.7-115) Woods Hole, MA; 2 
Sep 1876. USNM 68368 (5, 54-65.8) Great 
South Bay, NY; 1898. USNM 187214 (2, 
73.6-98.9) Deep Cove, Indian R., DE; 7 
Sept 1956. VIMS-CBL 173 (2, 108-123) 
CBL Pier, MD; 21 Jul 1936. VIMS-CBL 
1523 (11, 73.6-91.1) Broomes I., Patuxent 
R., MD; 13 Aug 1958. USNM 90798 (1, 
179) Cape Charles, VA. VIMS 61 (21, 87.6- 
123) Lynnhaven Inlet, Broad Bay, VA; 29 
Sept 1954. VIMS 607 (7, 78.5-159) York 
R., VA; 18 Jul 1970. USNM 131146 (1, 
147) Cape Charles, VA. USNM 131 146 (4, 
102-1 17) Morehead City, NC; 5 Sept 1964; 



taken with the holotype. VIMS-CBL 3571 
(1, 51.5) Morehead City, NC; 9 Jul 1967. 
USNM 51878(1, 153) Beaufort, NC. VIMS 
7873 (3, 71.9-115) Carteret Co., Atlantic 
Ocean beach, NC; 12 Sept 1976. USNM 
294396 (1, 150) SC, 32°26'N, 79°50'W; 6 
Aug 1953. USNM 149969 (1, 107) Georgia 
coast. USNM 294377 (2, 99.7-104) St. Si- 
mons I., GA; 15 Mar 1956. USNM 294426 
(1, 119) St. Simons I., GA; 14 Apr 1960. 
USNM 294427 (1, 105) St. Simons I., GA; 
16 Nov 1955. USNM 294441 (1, 160) St. 
Simons I., GA; 11 Nov 1956. USNM 
294435 (1, 55.6) off Savannah, GA, 31°4rN, 
80°35'; 21 Oct 1953. 

Other material examined: ATLANTIC 
U.S.: VIMS uncat. (56, 48. 1-146), Glouces- 
ter Pt., York R., VA; 19 Jul 1989 to 17 Sept 
1989. 



VOLUME 106, NUMBER 2 



381 



Table 10.— Comparison of means for F>ectoraI-fin ray counts for populations of Hyporhamphus meeki and H. 
unifasciatus, and between H. meeki and H. unifasciatus. Populations joined by the same line are not significantly 
different by the Tukey-Kramer method, alpha = 0.05. 



Population 



TK 



H. meeki 






West peninsula of ¥L 


147 


11.7 


MA to GA 


158 


11.4 


PL panhandle to Texas 


120 


11.4 


Yucatan 


26 


11.3 


Atlantic coast of FL 


106 


11.0 


H. unifasciatus 






Yucatan 


12 


11.1 


Caribbean South America 


120 


11.0 


South America (Brazil) 


122 


10.9 


Central America 


41 


10.9 


Florida 


69 


10.8 


West Indies 


136 


10.7 


Bermuda 


107 


10.0 


H. meeki 


558 


11.4 


H. unifasciatus 


607 


10.7 



ATLANTIC FLORIDA: UF 62140 (1, Smyrna Bch., 29°00'N, 80°32'W; 14 Oct 



125) 4.5 miles N. of Jupiter Inlet; 21 Aug 
1964. UF 77037 (2, 111-127) Matheson 
Hammock; 11 Jul 1970. UF 83999 (20, 
59.4-97.8) Indian R., Brevard Co.; 16 Aug 
1976. USNM 294365 (4, 55.8-141) offNew 



1953. USNM 294397 (19, 52.3-119) 
29°40'N, 81°06'W; 15 Oct 1953. USNM 
294494 (16, 91.4-145) 27°52'N, 80°26'W; 
20 Jan 1961. 
GULF COAST FLORIDA: UF 1010 (9, 



Table 1 1 .—Comparison of means for total first arch gill raker counts for populations of Hyporhamphus meeki 
and H. unifasciatus, and between H. meeki and H. unifasciatus. Populations joined by the same line are not 
significantly different by the Tukey-Kramer method, alpha = 0.05. 



Population 



TK 



H. meeki 






Yucatan 


26 


35.3 


FL panhandle to TX 


207 


35.2 


Atlantic coast of FL 


101 


35.1 


MA to GA 


255 


34.5 


West peninsula of FL 


194 


33.6 


H. unifasciatus 






Caribbean South America 


119 


31.1 


West Indies 


170 


31.1 


South America (Brazil) 


124 


30.6 


Central America 


53 


30.5 


Yucatan 


12 


30.0 


Florida 


66 


30.0 


Bermuda 


113 


29.5 


H. meeki 


780 


34.6 


H. unifasciatus 


657 


30.6 



382 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 12. — Comparison of means for total second arch gill raker counts for populations of Hyporhamphus 
meeki and H. unifasciatus, and between H. meeki and H. unifasciatus. Populations joined by the same line are 
not significantly different by the Tukey-Kramer method, alpha = 0.05. 



Population 



TK 



H. meeki 






MA to GA 


189 


26.8 


FL panhandle to TX 


156 


26.2 


West peninsula of FL 


177 


26.0 


Atlantic coast of FL 


114 


25.9 


Yucatan 


38 


25.5 


H. unifasciatus 






Caribbean South America 


114 


24.3 


West Indies 


141 


24.1 


South America (Brazil) 


109 


23.5 


Central America 


35 


23.5 


Yucatan 


12 


23.3 


Florida 


59 


22.9 


Bermuda 


100 


21.9 


H. meeki 


662 


26.2 


H. unifasciatus 


570 


23.5 



63.6-88.1) Cedar Key; 24 Jul 1948. UF 
51083 (5, 71.8-135) Alligator Harbor; 7 Sept 
1954. UF 52066 (5, 146-170) Alligator 
Harbor; 16 May 1954. UF 68483 (6, 114- 
148) St. Andrew's Bay, Panama City; 8 Oct 
1968. UF 76904 (24, 50.1-74.2) 1 mile N 
ofLee-CollierCo.line; 18 Jun 1966. USNM 
125446(2, 139-1 5 3) Tarpon Springs; 5 Nov 
1896. USNM 184268 (4, 133-146) Johns 
Pass, Madeira Boh.; 29 Sept 1958. USNM 
294431 (5, 103-136) Sarasota; 24 Aug 1967. 
USNM 294452 (16, 1 19-134) Sanibel I.; 11 
Aug 1959. USNM 294489 (6, 149-171) 
Sanibel I.; 8 Aug 1964. 

GULF COAST U.S., ALABAMA-TEX- 
AS: USNM 187122 (2, 15 1-1 52) S.Mobile, 
AL; 7 Jul 1960. VIMS 5095 (1, 113) Pelican 
Bay, Dauphin L, AL; 29 Aug 1974. USNM 
147781 (1, 176) Mississippi Gulf coast; 
1948. USNM 187120 (2, 123-128) inside 
Chandeleur Sound, LA; 19 Aug 1959. 
USNM 187123 (1, 73.9) Grande Isle, LA; 
21 Jul 1930. USNM 94546 (1, 141) Corpus 
Christi, TX. USNM 103390 (1, 69.1) near 
Corpus Christi, TX; 1937. USNM 120056 
(2, 128-155) Galveston, TX; 1941. USNM 



1 87 1 1 9 (2, 1 68-1 72) Corpus Christi, Sham- 
rock Cove, TX; 7 Apr 1 927. USNM 294440 
(26, 100.5-152) Aransas Pass, Institute of 
Marine Science pier, TX; Mar 1959. 

YUCATAN: UMMZ 143085 (28, 94.2- 
145) west of Progreso; 28 Mar to 1 Apr 
1936. UMMZ 143087 (15, 64.2-152) 
Chicxulub, near Progreso; 1 Apr 1936. MCZ 
32881 (3, 142-183) Yucatan; 1906. 

Hyporhamphus unifasciatus 

FLORIDA: UF 56209 (1, 143) Virginia 
Key; 26NOV 1959. UF62140(11, 108-134) 
N. of Jupiter Inlet; 21 Aug 1964. USNM 
34999 (4, 182-192) Key West; Dec 1883. 
USNM 158069 (3, 108-129) Snipe and 
Content Keys; June 1956. USNM 38544 (3, 
138-144) Key West; 15-27 Apr 1884. 
USNM 187121 (4, 75.0-79.9) St. Joseph's 
Bay; 2 Feb 1959. 

WEST INDIES: USNM 5802 (2, 146- 
186) Barbados. USNM 5847 (1, 103) Ja- 
maica. USNM 8803 (1, 111) Jamaica. 
USNM 10730 (2, 79.5-89.9) Bahia Honda, 
Cuba. USNM 34938 (2, 118-123) St. 



VOLUME 106, NUMBER 2 



383 



Thomas. USNM 38537 (2, 120-127) Ja- 
maica; 1-11 Mar 1884. USNM 38601 (6, 
1 19-158) Curacao; 10-18 Feb 1884. USNM 
501 1 1 (1, 194) San Juan Mkt., Puerto Rico; 
14 Jan 1899. USNM 82366 (3, 77.2-86.3) 
Los Arroyos, Cuba; 19 May 1914. USNM 
94077 (4, 112-139) Jamaica. USNM 
107428 (2, 135-165) Bahia Honda An- 
chorage, Cuba; 5 Apr 1937. USNM 130652 
(1, 195) Cuba. USNM 132524 (3, 109-147) 
Port-au-Prince, Haiti; 22 Oct 1945. USNM 
294364 (5, 90.5-129) Sable Bay, Dominica; 
13 Nov 1964. USNM 294493 (6, 78.6-1 19) 
Jobes Harbor, Greater Antilles; 20 Feb 1 966. 
USNM294515 (11, 81.9-167) 17°56'30"N, 
66°13'12"W; 18 Feb 1966. 

BERMUDA: AMNH 18711 (2, 87.3- 
90.2) Bermuda. ANSP 96626 (2, 1 14-117) 
Somerset, Bermuda; 12 Jun 1952. ANSP 
109562 (15, 82.5-135) Somerset, Bermuda; 
4 Jun 1952. ANSP 123715 (4, 100-141) 
Bermuda; Jun 1930. MCZ 34890 (5, 110- 
168) Bermuda. MCZ 40757 (6, 106-118) 
Bermuda; 1872. UMMZ 172321 (3, 99.2- 
154) Reach at Bio. Sta., Bermuda; 31 May 
1951. UMMZ 172369 (3, 81.6-97.9) St. 
George's I., Bermuda; 5 Jun 1951. UMMZ 
172418 (13, 87.9-145) St. George's I., Ber- 
muda; 8 Jun 1951. UMMZ 175957 (2, 85.9- 
108) Reach at Bio. Sta., Bermuda. UMMZ 
175967 (3, 124-140) Jetty at Bio. Sta., Ber- 
muda; 19 Mar 1957. UMMZ 175974 (1, 
105) Ferry Reach at Bio. Sta., Bermuda; 21 
Mar 1957. UMMZ 175981 (1, 79.7) Ferry 
Reach at Bio. Sta., Bermuda; 24 Mar 1957. 
UMMZ 176014(1, 148) Ferry Reach at Bio. 
Sta., Bermuda; 14 Apr 1957. UMMZ 
176154 (1, 114) Ferry Reach at Bio. Sta., 
Bermuda; 4 Jun 1957. USNM 294439 (7, 
87.7-110) Bermuda Harbor, Bermuda; 9 
Mar 1963. 

CENTRAL AMERICA: MCZ 32881 (1, 
183) Yucatan, Mexico; 1906. UF 7107 (2, 
165-181) Veracruz, Mocambo, Mexico; 20 
Jan 1958. UMMZ 143085 (10, 94.2-140) 
W. of Progreso, Yucatan, Mexico; 28 Mar- 
1 Apr 1936. UMMZ 143087 (1,71) Chicxu- 
lub, Yucatan, Mexico; 1 Apr 1936. USNM 



79658 (1, 160) Colon Mkt., Panama; 23 Jan 
1912. USNM 187843 (9, 62.4-85.5) off 
Bluefields, Nicaragua; 3-4 Jun 1962. 

SOUTH AMERICA: USNM 203826 (11, 
104-121) Gulf of Uraba, Colombia; 11 Jul 
1966. USNM 206658 (17, 121-205) Baru 
I., Colombia; 26 Sept 1969. USNM 38574 
(1, 135) Sabanilla, Colombia; 16-22 Mar 
1884. USNM 94764 (1, 188) near Puerto 
Colombia, Barranquilla, Colombia. USNM 
128286 (3, 121-129) Gulf of Venezuela, 
Venezuela; 5 Apr 1925. USNM 198404 (3, 
86.1-108) off French Guiana; 30 Jul 1956. 
MZUSP 5206 (7, 131-148) Ubatuba, Bra- 
zil; 1967. MZUSP 41092 (3, 184-191)Praia 
de Itapenia, Brazil; Jul 1965. MZUSP 41094 
(12, 131-179) Pontal, Ilheus, Brazil; 25 Oct 
1971. USNM 107220 (1, 151) Recife, Bra- 
zil; 1932. 

Acknowledgments 

For access to specimens mentioned in this 
paper, we thank E. B. Bohlke and W. F. 
Smith Vaniz (ANSP), G. H. Burgess (UF), 
W. N. Eschmeyer and D. Catania (CAS), 
M. N. Feinberg (AMNH), K. E. Hartel 
(MCZ), N. M. Menezes (MZUSP), D. W. 
Nelson (UMMZ), and R. Rosenblatt (SIO). 
We thank the staff of the many institutions 
that house the material we have examined 
for their help, Mildred Carrington for draw- 
ing the figures of Hyporhamphus (Fig. 2), 
and Janet Nestlerode for assistance with 
summarizing data and for plotting the dis- 
tribution map (Fig. 1). Carole Baldwin, Bar- 
ry Chemoff, Thomas Munroe, and Lynne 
Parenti made valuable comments on drafts 
of the manuscript. The second author thanks 
Frederick Berry, formerly of the National 
Marine Fisheries Service, for an introduc- 
tion to the study of halfbeaks and co-dis- 
covery of this new species in the early 1 960's. 
This is Contribution No. 1776 from the Vir- 
ginia Institute of Marine Science. 

Literature Cited 

Beebe, W.. & J. Tee-Van. 1933. Field book of the 
shore fishes of Bermuda. G. P. Putman's Sons, 
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384 



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Bigelow, H. B., & W. C. Schroeder. 1953. Fishes of 
the Gulf of Maine. -U.S. Fish & Wildlife Ser- 
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Bruce, N. L. 1986. Revision of the isopod crustacean 
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Hardy, J. D., Jr., & R. K. Johnson. 1974. Descrip- 
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Hildebrand, S. F. 1946. A descriptive catalog of the 
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Hoese, H. D., & R. H. Moore. 1977. Fishes of the 
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Houde, E. D., J. C. Leak, C. E. Dowd, S. A. Berkeley, 
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Jordan, D. S., & B. W. Evermann. 1896. The fishes 
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Leim, A. H., & L. R. Day. 1959. Records of uncom- 
mon and unusual fishes from eastern Canadian 
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Leviton, A. E., R. H. Gibbs, Jr., E. Heal, & C. E. 
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PROC. BIOL. SOC. WASH. 

106(2), 1993, pp. 385-389 

A NEW SPECIES OF OEDIPINA 

(AMPHIBIA: CAUDATA: PLETHODONTIDAE) FROM 

NORTHERN HONDURAS 

James R. McCranie, Larry David Wilson, and Kenneth L. Williams 

Abstract. —A new species of Oedipina from a cloud forest locality in northern 
Honduras is described and illustrated. The new species, O. gephyra, shows 
several morphological characteristics that bridge those of the two species groups 
of Oedipina recognized in the most recent revision of the genus. However, 
based on derived features such as the uniform dorsal coloration, short limbs, 
small feet, and long tail, the new species fits more readily into the uniformis 
group than the parvipes group. 



Two species of Oedipina (cyclocauda 
Taylor 1952 and stuarti Brame 1968) have 
heretofore been recorded from Honduras 
(Brame 1968, Meyer & Wilson 1971). Re- 
cently, we collected salamanders of this ge- 
nus from a single hardwood cloud forest 
locality in the western portion of the Cor- 
dillera Nombre de Dios in northern Hon- 
duras. An examination of these specimens 
demonstrated that the population repre- 
sents an undescribed species. 

Methods 

All measurements are in millimeters, 
made to the nearest tenth with dial calipers 
with the aid of a dissecting microscope. 
Measurements taken, method of expressing 
ratios, and the way of counting costal groove 
numbers follow the methods of Brame 
(1968). Abbreviations used are SVL (snout 
vent length [=standard length of Brame]), 
MVZ (Museum of Vertebrate Zoology, Uni- 
versity of California at Berkeley), and 
USNM (National Museum of Natural His- 
tory). An X-ray of the holotype of the new 
species (USNM 316535) confirmed the 
number of trunk vertebrae. Maxillary and 
vomerine tooth row counts are both sides 
summed. Comparative data for the other 
species of Oedipina were taken from Brame 
(1968) and Brame & Duellman (1970), 



however, data for O. cyclocauda were sup- 
plemented by those from the following re- 
cently collected Honduran specimens: AT- 
LANTIDA: mountains behind La Ceiba, 
260 m elev., USNM 316539. YORO: 6.6 
km S Yoro, ca. 1000 m elev., MVZ 171078; 
32.0 km W Yoro, MVZ 167772. 

Oedipina gephyra, new species 
Fig. 1 

Holotype. -USNM 316535, an adult fe- 
male, from 2.5 airline km NNE La Fortuna 
(15°26'N, 87°18'W), 1690 m elev., Cordille- 
ra Nombre de Dios, Departamento de Yoro, 
Honduras, collected 14 Aug 1991 by James 
R. McCranie, Kenneth L. Williams, and 
Larry David Wilson. Original number LDW 
9597. 

Paratypes. — USNM 316536-37, adult fe- 
males and USNM 316538, adult male, col- 
lected 14-16 Aug 1991 at the type locality, 
1690-1810 m. 

Diagnosis. — Oedipina gephyra can be dis- 
tinguished from all other species of Oedi- 
pina by the following combination of char- 
acteristics: 1 7 or 18 costal grooves per side; 
18 or 19 trunk vertebrae; 10-11 costal folds 
not covered by adpressed limbs; adult SVL 
52.3-57.8 {X= 55.6); feet small, adult hind 
foot width 1.4-1.6 (i^= 1.5), SVL/hind foot 



386 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. L Oedipina gephyra, new species, holotype, USNM 316535, SVL 56.7 mm. 



width 34.9-39.6 (Z= 37. 1); feet extensively 
webbed (Fig. 2); coloration uniform black- 
ish-brown with tiny iridophores present on 
all surfaces. 

Description of holotype.— Adult female 
with SVL of 56.7. Snout bluntly rounded in 
dorsal aspect and in profile; head width 5.4; 
SVL/head width 10.50; head length 8.1; 
SVL/head length 7.00; nostrils small, situ- 
ated near tip of snout; distinct nasolabial 
groove extending from lower-posterior 
margin of each nostril to the lip; labial pro- 
tuberances absent; canthus rostralis mod- 
erately arched; eyes not protuberant, not 
visible beyond margin of jaw when viewed 
from below; suborbital groove distinct; 
postorbital groove shallow, extending pos- 
teriorly from eye before turning sharply 
ventrally to connect with the gular fold, an- 
other branch proceeding sharply ventrally 
just posterior to mandible, extending irreg- 
ularly across throat anterior to gular fold; 



no dermal glands on head or body; 43 max- 
illary teeth, extending posteriorly to a point 
two-thirds distance through length of orbit; 
two premaxillary teeth, located posterior to 
lip; 22 vomerine teeth, in long, single, arched 
series, extending beyond outer edge of in- 
ternal nares; axilla-groin length 37.3; 17 
costal grooves per side; 1 8 trunk vertebrae; 
tail length 123.0; SVL/tail length 0.46; tail 
thick, nearly round at base, somewhat lat- 
erally compressed for last half of its length, 
barely constricted at base; tail width 3.6; 
SVL/tail width 15.75; tail depth 3.7; SVL/ 
tail depth 15.32; postiliac gland round, 
prominent; limbs short, 1 1 costal folds not 
covered when limbs adpressed to sides of 
trunk; hindlimb length 9.4; SVL/hindlimb 
length 6.03; hind foot width 1.5; SVL/hind 
foot width 37.80; digits one and two and 
three and four on forelimbs fused and digits 
one and two and four and five on hindlimbs 
fused, rest of digits on both fore- and hind- 



VOLUME 106, NUMBER 2 



387 



limbs fully webbed; digits on forelimbs in 
order of decreasing length 3-2-4-1, those on 
hindlimbs 3-4-2-5-1. 

Coloration in life: Head, body, and limbs 
blackish-brown throughout; tail black on all 
surfaces; tiny iridophores (visible under 
magnification) present on all surfaces. 

Variation. —Variation in morphology and 
coloration of the paratypes is minimal. The 
premaxillary teeth pierce the lip, labial pro- 
tuberances are weakly developed, and a 
mental gland is slightly indicated in the adult 
male (USNM 316538). There are 17 costal 
grooves per side in two and 18 in one 
(USNM 316537). Both females have 1 1 cos- 
tal folds not covered by the adpressed limbs, 
whereas the male has 10. Digits four and 
five on the hindlimbs of USNM 3 1 6538 are 
not fused. Other variation in the entire se- 
ries (including the holotype) is as follows 
(characteristics for the male separated by a 
comma from that of the females; means in 
parenthesis): SVL 55.5-57.8 (56.7), 52.3; 
axilla-groin length 35.6-38.6 (37.2), 33.8; 
head width 5.2-5.5 (5.4), 5.4; SVL/head 
width 10.50-10.67 (10.56), 9.69; head length 
8.1-8.4 (8.2), 8.4; SVL/head length 6.61- 
7.05 (6.89), 6.23; tail length 112.0-129.0 
(121.3), 115.0; SVL/tail length 0.45-0.50 
(0.47), 0.45; tail width 3.4-3.8 (3.6), 3.5; 
SVL/tail width 15.21-16.32(15.76), 14.94; 
tail depth 3.4-3.7 (3.6), 3.2; SVL/tail depth 
15.21-16.32(15.62), 16.34; hind foot width 
1.4-1.6 (1.5), 1.5; SVL/hind foot width 
36.13-39.64 (37.86), 34.87; hindlimb length 
7.6-9.4 (8.6), 8.4; SVL/hindlimb length 
6.03-7.30 (6.66), 6.23; maxillary teeth 43- 
54 (49.0), 48; vomerine teeth 16-22 (18.7), 
18; premaxillary teeth two in all. 

Natural history notes. — Oedipina gephyra 
was collected from inside rotten logs and 
stumps and underneath a plank within 
hardwood cloud forest (Lower Montane Wet 
Forest formation of Holdridge 1967) from 
1690 to 1810m elev. Several specimens of 
another salamander, Nototriton barbouri 
(Schmidt 1936), were also found inside rot- 
ten logs at the type locality while another 




Fig. 2. Right hind foot of a paratype of Oedipina 
gephyra (USNM 316538). Line equals 1.0 mm. 

salamander, Bolitoglossa conanti McCranie 
& Wilson 1993, was common in arboreal 
situations. 

Comparisons. — Brame (1968) recognized 
two species groups of Oedipina, the parvipes 
group with four species and the uniformis 
group with 1 1 species and Brame & Duell- 
man (1970) described an additional species 
belonging to the uniformis group. Oedipina 
gephyra shows several characteristics dis- 
tinctive of each group, and therefore cannot 
be easily placed in either group. Oedipina 
gephyra agrees with the parvipes group and 
can be distinguished from each species in 
the uniformis group by having 1 7 or 1 8 cos- 
tal grooves per side, 18 or 19 trunk verte- 
brae, and extensively webbed feet (19-22 
costal grooves per side, 20-23 trunk ver- 
tebrae, feet usually not extensively webbed 
in the uniformis group). On the other hand, 
O. gephyra resembles the uniformis group 
s