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PROCEEDINGS

of the

Biological Society of

Washington

VOLUME 117
2004

Vol. 117(1) published 1 June 2004 Vol. 117(3) published 7 December 2004
Vol. 117(2) published 4 August 2004 Vol. 117(4) published 20 December 2004

WASHINGTON
PRINTED FOR THE SOCIETY

Supplement to the Proceedings
of the Biological Society of
Washington

EDITOR

RICHARD V. STERNBERG
RICHARD C. BANKS

ASSOCIATE EDITORS

Classical Languages Invertebrates

FREDERICK M. BAYER STEPHEN L. GARDINER
CHRISTOPHER B. BOYKO
JANET W. REID

Plants Vertebrates

CAROL HOTTON GARY R. GRAVES
CAROLE C. BALDWIN
EDWARD O. Murpy

Insects Invertebrate Paleontology
WAYNE N. MaTuis GALE A. BISHOP

All correspondence should be addressed to the
Biological Society of Washington,
National Museum of Natural History
Washington, D.C. 20013

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OFFICERS AND COUNCIL
of the
BIOLOGICAL SOCIETY OF WASHINGTON
FOR 2004—2005

OFFICERS

President
ROY W. McDIARMID

President-Elect
W. RONALD HEYER

Secretary
CAROLE C. BALDWIN

Treasurer
T. CHAD WALTER

COUNCIL

Elected Members
MICHAEL D. CARLETON F. CHRISTIAN THOMPSON
W. DUANE HOPE JEFFREY T. WILLIAMS
MARILYN SCHOTTE NEAL WOODMAN

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TABLE OF CONTENTS

Volume 117

Alvarenga, Herculano M. F. and Storrs L. Olson, A new genus of tiny condor from the Pleistocene of
Brazil (Av essavill tii dae) eee merece ae asters eee ee ahehrus wgeeectsues oteees ade cosbece es, Goes
Graves, Gary R., Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 13. An undescribed intra-
generic combination Heliodoxa imperatrix xX Heliodoxa jacula ...............++-+++++++00--
Gill, Anthony C. and Hiroyuki Tanaka, Pholidochromis cerasina, a new species of pseudochromine
dotty back fish from the west Pacific (Perciformes: Pseudochromidae) .......................
Kawai, Tadashi and J. F. Fitzpatrick, Jr. Redescription of Cambaroides japonicus (De Haan, 1841)
(Crustacea: Decapoda: Cambaridae) withallocation of a type locality and month of collection of
(NOES “ao oes Sie'o's-o-o.ple: Gay Geontn to Mua ce: OE n eC Ceo: Creer Re HIE RPS aN er Sree a ee eee
Campos, Martha R. and Diego M. Valencia, Two new species of freshwater crabs of the genus Chaceus
Pretzmann, 1965 from the Serrania de Perija of Colombia (Crustacea: Decapoda:
Escudo thelphusiG ae) mesg tone ere eres Ta sesso ey leone eI ay eueeade send Ges eper suits. haapoaiparnaaas
McLaughlin, Patsy A. and Akira Asakura, Reevaluation of the hermit crab genus Parapagurodes
McLaughlin & Haig, 1973 (Decapoda: Anomura: Paguroidea: Paguridae) and a new genus for
Parapagurodes doederleini (Doflein, 1902) ...........-. 0.0 c ee eee
Asakura, Akira and Takeharu Kosuge, Pseudopaguristes bicolor, a new species of hermit crab
(Crustacea: Decapoda: Diogenidae) from Japan, the third species of the genus .................
Felder, Darryl L. and Brian Kensley, A new species of axiid shrimp from chemosynthetic communities
of the Louisiana continental slope, Gulf of Mexico (Crustacea: Decapoda: Thalassinidea) ........
Moore, Wendy, Description of a new Synidotea species (Crustacea: Isopoda: Valvifera: Idoteidae) from
LAW Aller ee er uate arent oie tore ke Aes See ae mse we ee wee ees aisen eos ES Peta ese ee
Schotte, Marilyn and Richard Heard, A new species of Synidotea (Crustacea: Isopoda: Valvifera) from
themorcherns Gulia i NICxdC Ot enc ate sens erie eee Se hehe eR PGi tae weeeG Bee oan Sem Oe
Ho, Ju-shey and Il-Hoi Kim, A new genus of the Clausidiidae (Copepoda: Poecilostomatoida) associ-
ated with a polychaete from Korea, with discussion of the taxonomic status of Hersiliodes Canu,
IS 5S eereae eee eee cate ates eee a area mr ata nh iene atielvoy eo atie Voeeaien se sinsadi dec ees a ah bce cl Sia Man Se ah eye Shae
Dreyer, Jennifer, Tomoyuki Miura, and Cindy Lee Van Dover, Vesicomyicola trifurcatus, a new genus
and species of commensal polychaete (Annelida: Polychaeta: Nautiliniellidae) found in deep-sea
clamsiromithe BlakesRidserc oldiscc piesa eae een ene eee nna ae ees israel
Cairns, Stephen D. and Frederick M. Bayer, Studies on western Atlantic Octocorallia (Coelenterata:
Anthozoa). Part 4: The genus Paracalyptrophora Kinoshita, 1908 .....................---..--
Wasshausen, D. C. and J. R. I. Wood, Notes on the genus Dicliptera (Acanthaceae) in Bolivia .....
AVOSvAnnualiNicetin geViiMUteS aires eerie sac eee Scan Stay Aedes Shiites eee aoe ae
Asakura, Akira, Pseudopaguristes shidarai, a new species of hermit crab (Crustacea: Decapoda:
Diogenidae) from Japan, the fourth species of the genus ...............---.----+--+--+--+---
L6pez-Mejia, Marilu, Fernando Alvarez, and Luis M. Mejia-Ortiz, A new species of Procambarus
(Crustacea: Decapoda: Cambaridae) from Veracruz, Mexico ..............-.---++-+++-eeeeee:
Lewis, Julian J., Brackenridgia ashleyi, a new species of terrestrial isopod from Tumbling Creek Cave,
Missouri (Isopoda: Oniscidea: Trichoniscidae) .............--222-- eee eee eee eee eee
Markham, John C., New species and records of Bopyridae (Crustacea: Isopoda) infesting species of the
genus Upogebia (Crustacea: Decapoda: Upogebiidae): the genera Orthione Markham, 1988, and
Gyre ComaliasSAbanceni aS Olle. care ies hoe eee ee Oe Oe eR a Hee eee
Duplessis, Kirk and Henry M. Reiswig, Three new species and a new genus of Farreidae (Porifera:
Mexatinellida-gHexaGtim@si cal paeresws rns ten cs een ree ose eis AHS) Si eee, ctroncheira en eaeren She es eos er eue har ewetreus
Meyer, Stephen C., The origin of biological information and the higher taxonomic categories .......
Dickerman, Robert W., A review of the North American subspecies of the Great Blue Heron (Ardea
[QROGHOS) 23 4-0-6 9.0.4 8:80 BS LES OS ELS AS Ol RT TT ee Re as eee ae are ee ae eee
Goodman, Steven M. and Voahangy Soarimalala, A new species of Microgale (Lipotyphla: Tenrecidae:
Oryzorictinae) from the Forét des Mikea of southwestern Madagascar ..................-..-.-
Woodman, Neal, Designation of the type species of Musaraneus Pomel, 1848 (Mammalia:
SOHCOMOLPNAgS OLICIG ae) benaey wei el ruereee a eee uss Slay Sls, Sie yea yee enum asl Geena subieashereeteerenane.
Esselstyn, Jacob A., Peter Widmann, and Lawrence R. Heaney, The mammals of Palawan Island,
LPAONUUY DY OVUNSS aad. OB @, alo Bb eoatyog: Cumgte auctor beacG- cra Ota ee CN Ce Ee RSET TS eS ee
Kraus, Fred and Allen Allison, A new species of Tropidonophis (Serpentes: Colubridae: Natricinae)
from the D’Entrecasteaux Islands, Papua New Guinea ...........------ 22-2 eee eee eee

106
114
140
150
153

169

176

186

199
213

242

Dail

266

271

303

McCranie, James R. and Franklin E. Castafieda, A new species of snake of the genus Omoadiphas
(Reptilia: Squamata: Colubridae) from the Cordillera Nombre de Dios in northern Honduras .... .
Malabarba, Luiz R., Flavio C. T. Lima, and Stanley H. Weitzman, A new species of Kolpotocheirodon
(Teleostei: Characidae: Cheirodontinae: Compsurini) from Bahia, northeastern Brazil, with a new
diagnosisiof the Genus) oa cye te cesussicue.s celtic te easesee snsveae tele Co Seclseeeban: siaaspetee cackenc ie) tare eee ee
Castro, Ricardo M. C. and Richard P. Vari, Astyanax biotae, a new species of stream fish from the Rio
Paranapanema basin, upper Rio Parana system, southeastern Brazil (Ostariophysi: Characiformes:
(Oi nkctes(cis Clo) ere a ce era ic culos doc od ce 6 ocd Soo oucds 66s e500
Benine, Ricardo C., Gabriela Zanon Peligao, and Richard P. Vari, Tetragonopterus lemniscatus
(Characiformes: Characidae), a new species from the Corantijn River basin in Suriname .........
Worsaae, Katrine, Wolfgang Sterrer, and Thomas M. Iliffe, Longipalpa saltatrix, a new genus and
species of the meiofaunal family Nerillidae (Annelida: Polychaeta) from an anchihaline cave in
| 5X¢) 101 (6 - ene nae eee ere ergs eee ete erect etme Sint in oA ci ciniS h Gidis sa cg hele Glos cio 5 0 0-0
Campos, Martha R., Neostrengeria lemaitrei, a new species of freshwater crab from Colombia
(Crustacea: Decapoda: Pseudothelphusidae), and the vertical distribution of the genus...........
Alvarez, Fernando, José Luis Villalobos, and Thomas M.. Iliffe, A new species of Agostocaris (Caridea:
Agostocarididae) from Acklins Island, Bahamas .........................----2.---2.-----
Wicksten, Mary K. and Joel W. Martin, A new species of caridean shrimp of the family Stylodactylidae
from the eastern Pacific'OCeam 4). in bestia lee oe 3 eine ee ope eee eee eee wt
Buhl-Mortensen, L. and W. A. Newman, A new pedunculate barnacle (Cirripedia: Heteralepadidae)
fromthe Northwest Atlamtic= cise cce ccd c iy ar eae ep ene ail 6 enn eee ae
Kornicker, Louis S. and J. A. Rudjakov, Two new species of seven-spined Bathyconchoecia from the
North Atlantic and Indian oceans (Crustacea: Ostracoda: Halocypridae)......................
Yanagi, Kensuke and Marymegan Daly, The hermaphroditic sea anemone Anthopleura atodai n. sp.
(Anthozoa: Actiniaria: Actiniidae) from Japan, with a redescription of A. hermaphroditica .......
Robinson, Harold and Abigail J. Moore, New species and new combinations in Rhysolepis
(Helvantheaés, Asteraceae) five) a0 sean eee tee asec ee uy artnet) Casi oan uo ee oc Re
Cairns, Stephen D. and Frederick M. Bayer, Studies on western Atlantic Octocorallia (Coelenterata:
Anthozoa). Part 5. The genera Plumarella Gray, 1870; Acanthoprimnoa, n. gen.; and Candidella
Bayer V994) ss. cece gists aie Riana muattti wera in, sheen See are aeons arene cee ot Si oe Se aS ate ee ee
Ardelean, Adorian and Daphne Gail Fautin, A new species of the sea anemone Megalactis (Cnidaria:
Anthozoa: Actiniaria: Actinodendridae) from Taiwan and designation of a neotype for the type
Species OL thezSenUs’ ax avs. esau oy wee SAAS eI TE aA RON GEC oT eee cet ee
Vazquez-Bader, Ana Rosa and Adolfo Gracia, A new genus and new species of crab of the family
Xanthidae MacLeay, 1838 (Crustacea: Decapoda: Brachyura) from the southwestern Gulf of Mexico
Sternberg, Richard v. and Marilyn Schotte, A new anchialine shrimp of the genus Procaris (Crustacea:
Decapoda: Procarididae) from the Yucatan Peninsula ............-...--+---+-+--++---+-++:>
Phone, Hla and Hiroshi Suzuki, Macrobrachium patheinense, a new species of freshwater prawn
(Crustacea: Decapoda: Palaemonidae) from Myanmar .................-2---00 ee eee eee eees
Gomez, Samuel, A new species of Enhydrosoma Boeck, 1872 (Copepoda: Harpacticoida: Cletodidae)
fromthe Eastern Tropical! Pacific: 2: 3 4eccsie condenses cle Aen che Sasi oa SO eee
Borrero-Pérez, Giomar Helena and Milena Benavides-Serrato, New record of Ophiosyzygus disacan-
thus Clark, 1911 (Echinodermata: Ophiuroidea: Ophiomyxidae) in the Caribbean Sea ...........
Knapp, Leslie W. and Hisashi Imamura, Sunagocia sainsburyi, a new flathead fish (Scorpaeniformes:
Platycephalidae)itroninorthwestemeAus tralia eee ie nein niente enna eee
Vari, Richard P. and Carl J. Ferraris, Jr., A new species of Nannocharax (Characiformes:
Distichodontidae) from Cameroon, with the description of contact organs and breeding tubercles in
Li SYR e{=) 1b eee names eee nee eRe etn enema ey CSIC OOD OG A.S OU oe oo soe 516 dt
DoNascimiento, Carlos, Francisco Provenzano, and John G. Lundberg, Rhamdia guasarensis
(Siluriformes: Heptapteridae), a new species of cave catfish from the Sierra de Perijé, northwestern
WEMEZUCL A eccd st Semen dilatere ican ie ses ses wins sareecaachni sion cease ee al Se CA poses TALE oe AR Sem
Olson, Storrs L., Taxonomic review of the fossil Procellariidae (Aves: Procellariiformes) described
from Bermuda iby Re We ShULeldt =< gic cnc wre cos erenep aes ne ures ey cre ee crist er nee ae ee oS
Espinasa, Luis and Bethany Burnham, Revision of the genus Squamigera (Insecta: Zygentoma:
Nicoletiidae) with descriptions of two new species ................ 0.000 ee eee eee eee eee eee
Minutesiof the: 20047AnnualiMeeting saaeeisem camo ccn ci ccic ae cctcie sc cteer om ely Stele renee
ConstitutionandiBy laws: sess ee eis a eas ats tinal oe Ren in eee

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| PROCEEDINGS oF THE
BIOLOGICAL SOCIETY
or WASHINGTON

24 JUNE 2004

VOLUME 117
NUMBER 1

.THE BIOLOGICAL SOCIETY OF WASHINGTON

2003-2004
Officers
President: Roy W. McDiarmid Secretary: Carole C. Baldwin
President-elect: W. Ronald Heyer Treasurer: T. Chad Walter
Elected Council

Michael D. Carleton G. David Johnson

Clyde Roper Michael Vecchione

Marilyn Schotte Don Wilson

Custodian of Publications: Storrs L. Olson

PROCEEDINGS

Editor: Richard v. Sternberg

Associate Editors

Classical Languages: Frederick M. Bayer Invertebrates: Stephen L. Gardiner
Plants: Carol Hotton Christopher B. Boyko
Insects: Wayne N. Mathis Janet W. Reid
Vertebrates: Gary R. Graves Invertebrate Paleontology: Gale A. Bishop

Ed Murdy

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SNITHSON 4g

JUL 1 6 2004

( a condor from the Pleistocene of Brazil
(Aves: Vulturidae)

Herculano M. EF Alvarenga and Storrs L. Olson

(HA) Museu de Historia Natural de Taubaté, Rua Colombia 99, Jardim das Nagoes, Taubaté SP,

CEP 12030-520, Brazil
(SLO) Division of Birds, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560, U.S.A.

Abstract.—A new genus and species of Vulturidae (Cathartidae auct.), Win-
gegyps cartellei, is described from Pleistocene cave deposits in the states of
Bahia and Minas Gerais, Brazil. This species is closely related to condors
Gymnogyps and Vultur, particularly the former, as opposed to the smaller ca-
thartid vultures, but is much smaller, being slightly smaller than the smallest
living member of the family, the Lesser Yellow-headed Vulture Cathartes bur-
rovianus. The Vulturidae appears to consist of two basic divisions (condors vs.
other vultures) that differ profoundly in the morphology of the skull. Each
appears to have been more diverse in the past and to contain larger or smaller
species than survived to the present.

Resumo.—Um novo género e nova espécie de Vulturidae (Cathartidae auct.),
Wingegyps cartellei, é descrito dos depositos pleistocénicos de cavernas da
Bahia e Minas Gerais, Brasil. Este é mais relacionado aos condores Gymnogyps
e Vultur, principalmente com o primeiro, do que com os verdadeiros urubus,
embora seja de tamanho reduzido, menor ainda que Cathartes burrovianus, 0
menor membro vivente da familia. Os Vulturidae se constituem de dois grupos
basais, condores e urubus, que diferem entre si basicamente pela morfologia
do cranio (o tamanho nao é fundamental), sendo que ambos parecem ter sido

OCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
117(1):1—9. 2004.

bastante diversificados no passado.

Peter Wilhelm Lund was a Danish natu-
ralist who resided in Brazil from 1832 until
his death in 1880. Between 1835 and 1849
he shipped masses of Quaternary fossils
from the state of Minais Gerais back to
Denmark for study (Voss and Myers 1991).
The great majority of these fossils were of
mammals, including extinct megafauna, but
also rodents and bats (Voss and Myers
1991, Czaplewski and Cartelle 1998, Car-
telle 1999).

The mammals were originally studied by
Herluf Winge, who published his excep-
tionally perceptive findings in a series of
volumes entitled E Museo Lundii from
1887 to 1915. The study of fossil birds
from these deposits fell to his brother Oluf

Winge (1888) who produced a list of some
126 species. Only one of these, the anatid
Chenalopex (now Neochen) pugil, was
named as new, and many were referred to
modern taxa. Others could not be assigned
either for lack of comparative material or
because Winge considered them probably
to represent unknown species that he left
unnamed.

Among the last was a vulture that Winge
(1888: 33) regarded as probably belonging
to a new genus and species (“‘G. sp. indet.
magnitudine Catharistae atrati” [=
gyps atratus|). This was represented by the
distal end of a humerus and an ulna lacking
the distal end. He described these speci-
mens in considerable detail and illustrated

Cora-

7, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

the humerus in comparison with a fossil of
the Black Vulture Coragyps atratus. Noth-
ing further was ever made of this discovery
during the succeeding 115 years.

In identifying fossil bird remains from a
cave in the state of Bahia, we puzzled over
a peculiar ovoid cranium that defied place-
ment to family until we happened to notice
that fossil crania of Gymnogyps from Ran-
cho La Brea, California, seemed to be sim-
ilar in shape, although larger. We identified
two humeri from Bahia as probably belong-
ing to the same species as the cranium, and
a well-preserved distal fragment appeared
to be identical to that illustrated by Winge
as his unidentified new genus. We were able
to borrow Winge’s original material and
confirmed that he was quite correct that a
new genus and species is indicated. This,
however, turns out not to be closely related
to the smaller genera of Vulturi-
dae,Cathartes or Coragyps, but to the much
larger condors, especially Gymnogyps.

Comparative material examined.—Pre-
liminary comparisons were made with al-
most all families of non-passerine birds and
all species of South American vultures in
Museu de Histéria Natural de Taubaté. The
original material of Vulturidae collected by
Lund in Minas Gerais was borrowed from
the Zoological Museum University of Co-
penhagen (ZMUC) and restudied and com-
pared. Modern skeletons examined in the
Division of Birds, National Museum of
Natural History, Smithsonian Institution
(USNM) included: Gymnogyps californi-
anus 3369, 492447; Vultur gryphus
345384, 429839; Sarcoramphus papa
345434, 559318; Coragyps atratus 613353;
Cathartes aura 490864, 612254; C. melam-
brotus 621939, C. burrovianus 431336,
622341.

Systematics

Class Aves
Family Vulturidae

Within the family, there is marked oste-
ological distinction, particularly in the neu-

rocranium, between the two living genera
of condors (Vultur and Gymnogyps) on one
hand, and all of the other genera (hereafter
‘‘vultures’’) on the other. The more salient
of these were first noted by Miller and
Howard (1938) and were further docu-
mented by Fisher (1944). The extinct genus
and species Breagyps clarki was also
shown to belong with the condors based on
cranial characters (Miller and Howard
1938). Cranial differences were detailed
and extended to additional fossil specimens
by Emslie (1988). In the following com-
parisons, “‘condors” includes the new ge-
nus.

Neurocranium.—In dorsal view the neu-
rocranium of condors is relatively longer
and narrower, appearing almost ovoid in
shape; the tranverse nuchal crest is visible
because the attachments of the cervical
musculature extend much farther dorsally
than in the vultures, and the cerebellar
prominence is much larger and more dis-
tinct.

In posterior view, the last two features
are equally distinct. The foramen magnum
is much larger and more elliptical in con-
dors, as opposed to nearly circular in the
vultures. The occipital condyle is more dis-
tinctly stalked (better seen in ventral view)
and is rounded, lacking the notch in the
dorso-posterior surface seen in vultures. In
condors, the dorsolateral margins of the fo-
ramen magnum give rise to distinct crests
that angle ventro-laterally to the extremely
well developed paroccipital processes (su-
praoccipital processes of Suarez and Emslie
2003; opisthotic processes of Fisher 1944;
postauditory processes of Miller & Howard
1938) that parallel the similarly well devel-
oped processes that angle out from the oc-
cipital condyle (occipital processes of Sua-
rez and Emslie 2003; medial basitemporal
processes of Bock 1960; exoccipital pro-
cesses of Fisher 1944). In the vultures these
processes were always much smaller and
differently shaped.

In lateral view, the temporal fossa is
much larger in condors, so that the postor-

VOLUME 117, NUMBER 1

bital and zygomatic processes are farther
apart, and the orbit is relatively smaller than
in the vulture.

Humerus.—Differs from Cathartes or
Coragyps in having the distal end more ex-
panded and the ectepicondylar prominence
situated more distally on the shaft. Sarco-
ramphus differs in having a large pneumat-
ic Opening in the depression between the
entepicondylar prominence and the ulnar
condyle. The entepicondylar prominence is
less developed than in Vultur. In almost ev-
ery respect, down to the slightest detail of
pneumatization, the distal end of the hu-
merus of the new genus is a perfect dupli-
cate in miniature of that of Gymnogyps. The
complete humerus of the new genus is so
worn as to preserve few useful characters,
but it does have on the palmar surface a
slightly pneumatized depression just distal
to the head as in Gymnogyps. This depres-
sion is absent in Cathartes, only slightly in-
dicated in Vultur, and bears a very large
pneumatic foramen in Coragyps and Sar-
coramphus.

Wingegyps, new genus

Type species.—Wingegyps cartellei, new
species.

Diagnosis.—A tiny condor most similar
to Gymnogyps in the narrowness and elon-
gation of the neurocranium, but even nar-
rower, with the braincase in dorsal view be-
ing of nearly uniform width, rather than ex-
panding posteriorly. Muscle scars on either
side of the cerebellar prominence are cor-
respondingly narrower. The foramen mag-
num opens directly posteriorly rather than
partly ventrally. The paroccipital processes
and their associated crests are angled more
ventrally than in Gymnogyps or Vultur. Dif-
fers from other condors in having the entire
occipital condyle and its stalk visible in lat-
eral view (not visible in Breagyps, only par-
tially visible in Gymnogyps and Vultur).

The ulna is like that of condors and Sar-
coramphus in having very little pneumati-

zation in the brachial depression (well de-
veloped in Cathartes and Coragyps) but
differs from all but Vultur in having the
olecranon more distinctly set off from the
margin of the internal cotyla. The olecranon
is narrower, however, than in Vultur.
Etymology.—Winge + Greek gyps, vul-
ture, in commemoration of the perspicacity
of Oluf Winge for recognizing the distinc-
tiveness of this remarkable new genus.

Wingegyps cartellei, new species
Figs. 1-4

Holotype.—Neurocranium lacking the
parasphenoid rostrum and ethmoid region,
with damage to the anterior margin of the
frontals and left otic area, MCL CLA782
(Fig. 1B, 2B).

Type-locality.—Brazil, Bahia State,
Municipio de Morro do Chapeu, Gruta dos
Brejoes (11°00’30"S, 41°26'07’W), eleva-
tion ca. 600 m.

Horizon and age.—Probably late Pleis-
tocene or early Holocene. A radiocarbon
date of 12,200 + 120 radiocarbon years be-
fore present was obtained from a coprolite
of a ground sloth from the type-locality
(Czaplewski and Cartelle 1998). Associated
mammals from caves in Bahia and Minas
Gerais are considered to be of Pleistocene
age (Cartelle 1999).

Measurements (mm) of holotype.—Total
length as preserved 48.6; width at level of
postorbital processes 27.5; width at level of
base of zygomatic processes 29.4; greatest
depth at midline 27.2; width and depth of
foramen magnum; 8.9 X 8.1; width of oc-
cipital condyle 4.1

Paratypes.—Topotypes: Complete but
very worn left humerus MCL CLA670
(Fig. 4C); distal third of left humerus MCL
CLA1678 (Fig. 3B).

Lapa do Tit, Minais Gerais, Brazil: distal
half of right humerus ZMUC 1116 (Fig.
3A); right ulna lacking distal end ZMUC
1118 (Fig. 4A).

Measurements (mm) of paratypes.—Hu-
meri (in the same sequence as above): total

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ioe Ie
B, Wingegyps cartellei, new species, holotype MCL CLA782; C, Cathartes aura USNM 612254. Seale =
2 cm.

length 129.5, —, —; length from head to
distal end of pectoral crest 54.4, —, —;
shaft width and depth at midpoint 10.0 x
8.5, —, 9.7 X 7.6; distal width —, 24.8,
23.2; greatest dimension of brachial depres-
sion 13.4; 11.7, 13.0; greatest dimension of
radial condyle —, 10.8 11.1. Ulna: proxi-
mal width 12.5; proximal depth 15.8; length
of brachial depression 23.4.

Etymology. Dedicated to paleotologist
Castor Cartelle of the Universidade Federal
de Minas Gerais in recognition of his ex-

Neurocrania in dorsal (top) and posterior (bottom) views: A, Gymnogyps californianus USNM 3369;

cavations at Gruta dos Brejoes (Cartelle
1983) and his contributions to the paleon-
tology of Brazil.

Diagnosis—Much smaller than any
known condor; slightly smaller than the
smallest living cathartid vulture (Lesser
Yellow-headed Vulture Cathartes burrovi-
anus).

Discussion.—Wingegyps is indisputably
a condor based on the very distinct features
of the neurocranium and on similarities of
the distal end of the humerus. Its extraor-

VOLUME 117, NUMBER 1 5

Fig. 2. Neurocrania in lateral view: A, Gymnogyps californianus USNM 3369; B, Wingegyps cartellei, new
species, holotype MCL CLA782; C, Cathartes aura USNM 612254. Scale = 2 cm.

6 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3.

dinarily small size is quite unanticipated,
being somewhat smaller than the smallest
living species of the family (Carthartes
burrovianus). The humerus is only slightly
shorter than in females of the Black Vulture
Coragyps atratus from the tropics, which
are smaller than individuals at the temperate
ends of the species’ range (Brodkorb 1944).
But the humerus is proportionately much
shorter in Coragyps atratus than in Cathar-
tes, so that this species is otherwise much
larger than Cathartes burrovianus (1875 g
in a female Coragyps from Panama vs 960
g ina male C. burrovianus form Guyana).

Wingegyps shows that condors were
much more diverse in size in the past. The
family Vulturidae may be viewed as being
divisible into two basic groups: the condors
(Vultur, Gymnogyps, Breagyps, Winge-

Humeri in anconal view: A, Wingegyps cartellei, new species, paratype ZMUC 1116; B, Wingegyps
cartellei, new species, paratype MCL CLA1678; C, Gymnogyps californianus USNM 492447, reduced to the
same size as B; D, same natural size. Scale = 2 cm except for C.

gyps), which appear to be derived (Emslie
1988) and the remaining living genera (Ca-
thartes, Coragyps, Sarcoramphus), which
may be paraphyletic. Both may have been
more diverse at one time and perhaps some
of the larger fossil taxa (Geronogyps, Plio-
gyps, “Sarcoramphus”’ kernense—see Em-
slie 1988), for which cranial material is un-
known, may prove to be more closely re-
lated to the assemblage of smaller vultures
than to condors.

Known only from a rather limited area in
eastern Brazil, Wingegyps doubtless had a
greater range than indicated at present, pos-
sibly much greater. If it has been collected
in fossil deposits elsewhere the material
might easily be overlooked as belonging to
Cathartes or Coragyps because of its small
size.

VOLUME 117, NUMBER 1

Fig. 4.
paratype ZMUC 1118; B, Cathartes burrovianus USNM 431336; C, Wingegyps cartellei, new species, paratype
MCL CLA670; D, Cathartes burrovianus USNM 431336. Scale = 2 cm.

What sort of feeding niche might such a
tiny condor have occupied? The habits of
living species of the family are briefly sum-
marized from Olson et al. (1967), Sick
(1993), and Hertel (1994). The living con-
dors Vultur and Gymnogyps forage by sight
and prefer soft viscera from large carcasses.
Sarcoramphus and Coragyps forage by
sight and are very aggressive at carcasses.
Coragyps takes food in small bits, tearing
even small carcasses such as a frog or
mouse to pieces before eating. The species
of Cathartes are very different in finding
food with their keen sense of smell. Thus,
they specialize in finding caracasses of

Right ulnae (A, B) and left humeri (C, D) in palmar view: A, Wingegyps cartellei, new species,

small animals either before they are located
by sight foragers or detecting food that can-
not be seen at all from above. They are also
very docile and not at all competitive with
other vultures at carcasses.

The small size of Wingegyps would have
placed severe limitations on its ability to
process the majority of carcasses or to com-
pete at carcasses with other species of vul-
tures. If we assume that it was like its clos-
est relatives in lacking the olfactory capa-
bilities of Cathartes, Wingegyps would
have had little success competing with any
of the species of Cathartes for small car-
casses. There does seem to be a potential

8 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

niche in the New World, however, that is
not as fully exploited as it is in the Old
World, viz. palm fruits.

In Africa, the Palm-nut Vulture (Gypho-
hierax, Accipitridae) feeds mainly on the
soft mesocarp of the African oil palm Eleis
guineensis. This palm has been introduced
to Brazil and Sick (1993:149) describes
Turkey Vultures Cathartes aura as being a
“nuisance” in palm plantations in Amazon-
ia, where they consume the fruits. He also
records them as feeding on the native palm
Acrocomia sclerocarpa (= A. aculeata), a
very widespread species occurring through
the West Indies and from Mexico south to
southern Brazil and Paraguay (Henderson et
al. 1995), and overlapping the small known
range of Wingegyps. Although Wingegyps
may possibly have been the New World
ecological equivalent of the unrelated Old
World Palm-nut Vulture, its habits might
also have been like that of the Egyptian
Vulture Neophron percnopterus in subsist-
ing on scraps thrown off of carcasses by
larger vultures. Such habits might better ex-
plain the extinction of Wingegyps, as many
of the larger avian scavengers in the New
World also went extinct at the time of dis-
appearance of much of the mammalian
megafauna (Steadman and Martin 1984).

Acknowledgments

Travel by SLO to Brazil was made pos-
sible by the Alexander Wetmore Endow-
ment Fund, National Museum of Natural
History, Smithsonian Institution. We are
grateful to Castor Cartelle of the Museu de
Ciéncias Naturais (MCL) of the Pontificia
Universidade Catdlica de Minas Gerais,
Belo Horizonte, Brazil, for making the fos-
sil material from Gruta dos Brejoes avail-
able for study, to Jon Fjeldsa and Kim Aar-
is-Sgrensen, of the Zoological Museum
University of Copenhagen (ZMUC), Den-
mark, for lending the material studied by
Oluf Winge. Frederick V. Grady, Smithson-
ian Department of Paleobiology, cleaned
and repaired fossil specimens. Kevin Sey-

mour, Royal Ontario Museum, suggested a
reference. Photographs are by John Steiner,
Smithsonian Center for Scientific Imaging
and Photography, and the figures were ar-
ranged by Brian Schmidt, Division of
Birds, Smithsonian Institution.

Literature Cited

Bock, W. J. 1960. Secondary articulation of the avian
mandible.—Auk 77:19-55.

Brodkorb, P. 1944. Geographical variation in the Black
Vulture.—Papers of the Michigan Academy of
Science, Arts, and Letters 29:115-121.

Cartelle, C. 1983. Tesouro féssil no sertao baiano.—
Ciéncia Hoje 1(5):35-43.

. 1999. Pleistocene mammals of the cerrado
and caatinga of Brazil. Pp. 27-46 in J. EK Ei-
senberg and K. H. Redford, eds., Mammals of
the Neotropics, vol. 3. The Central Neotropics.
Ecuador, Peru, Bolivia, Brazil. Chicago, Uni-
versity of Chicago Press, 609 pp.

Czaplewski, N. J., & C. Cartelle. 1998. Pleistocene
bats from cave deposits in Bahia, Brazil—Jour-
nal of Mammalogy 79:784-803.

Emslie, S. D. 1988. The fossil history and phyloge-
netic relationships of condors (Ciconiiformes:
Vulturidae) in the New World.—Journal of Ver-
tebrate Paleontology 8:212-228.

Fisher, H. I. 1944. The skulls of the cathartid vul-
tures.—Condor 46:272-296.

Henderson, A., G. Galeano, & R. Bernal. 1995. Palms
of the Americas. Princeton, New Jersey, Prince-
ton University Press, 352 pp.

Hertel, E 1944. Diversity in body size and feeding
morphology within past and present vulture as-
semblages.—Ecology 75:1074-1084.

Miller, L. H., & H. Howard. 1938. The status of the
extinct condor-like birds of the Rancho La Brea
Pleistocene.—Publications of the University of
California at Los Angeles in Biological Scienc-
es 1:169-176.

Olson, S. L., H. Loftin, & J. Wiese. 1967. Observa-
tions on the behavior of Black and Turkey Vul-
tures at traps and in captivity.—Bird-Banding
38:75-76.

Sick, H. 1993. Birds in Brazil. Princeton, New Jersey,
Princeton University Press, 703 pp.

Steadman, D. W., & P. S. Martin. Extinction of birds
in the Late Pleistocene of North America. Pp.
768-780 in P. S. Martin and R. G. Klein, eds.,
Quaternary Extinctions. A Prehistoric Revolu-
tion. Tucson, University of Arizona Press, 892
Pp.

Suarez, W., & S. D. Emslie. 2003. New fossil material
with a redescription of the extinct condor Gym-
nogyps varonai (Arredondo, 1971) from the

VOLUME 117, NUMBER 1

Quaternary of Cuba (Aves: Vulturidae).—Pro-
ceedings of the Biological Society of Washing-
ton 116:29-37.

Winge, O. 1888. Fugle fra Knuglehuler i Brasilien.
E Museo Lundii 1 (2): pp. 54 + 5 + 1 plate.

Voss, R. S., & P. Myers. 1991. Pseudoryzomys simplex
(Rodentia: Muridae) and the significance of
Lund’s collections from the caves of Lagoa San-
ta, Brazil—Bulletin of the American Museum
of Natural History 206:414-432.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(1):10-16. 2004.

Diagnoses of hybrid hummingbirds (Aves: Trochilidae).
13. An undescribed intrageneric combination,
Heliodoxa imperatrix x Heliodoxa jacula

Gary R. Graves

Department of Zoology, MRC-116, National Museum of Natural History,
Smithsonian Institution, PO. Box 37012, Washington, D.C. 20013-7012, U.S.A.

Abstract.—An enigmatic specimen collected by Perry O. Simons, presum-
ably on the Pacific slope of the Ecuadorian Andes, is demonstrated to be a
hybrid between Heliodoxa imperatrix and Heliodoxa jacula jamesoni. This
represents the only known instance of intrageneric hybridization in Heliodoxa.
External measurements of the hybrid are consistent with the proposed parental

hypothesis.

At the monthly meeting of the British Or-
nithologists’ Club on the 17 January i900,
Ernst Hartert (1900:39) exhibited a speci-
men of hummingbird, “obtained in Ecuador
by Mr. Simons, combin[ing] in a striking
way the shape and colours of Eugenia [He-
liodoxa] imperatrix and Heliodoxa jacula
jamesoni . . . to be described in detail in the
‘Novitates Zoologicae’.”’ Although Hartert
never published a description or reported a
museum registration number, this brief ex-
hibition notice has been cited in catalogs of
avian hybrids (Gray 1956, Panov 1989).
Here I provide a taxonomic assessment of
the specimen employing the methods and
assumptions outlined in Graves (1990) and
Graves & Zusi (1990), as modified by the
findings of Graves (1998, 1999).

Methods

The specimen, now deposited in the Nat-
ural History Museum (registration number,
1902.3.13.2211), bears two labels, one from
the British Museum marked “P. O. Si-
mons,’ and an older one from the Roths-
child Museum. Perry O. Simons collected
mammals and birds for Oldfield Thomas
(British Museum) from 1898 until his mur-
der near Cuervas, Argentina, in 1901 (Allen
1903, Chubb 1919). Both specimen labels

are marked with Hartert’s taxonomic deter-
mination. Curiously, neither the specimen
labels nor the Natural History Museum cat-
alog indicate when or where the specimen
was collected.

I compared the specimen (Figs. 1, 2) with
all species in the subfamily Trochilinae, the
typical hummingbirds (Zusi & Bentz 1982,
Sibley & Monroe 1990, Bleweiss et al.
1997), deposited in the Natural History Mu-
seum, Tring, and the National Museum of
Natural History, Smithsonian Institution.
The specimen appears to be a male in de-
finitive plumage as judged by the absence
of striations on the maxillary ramphotheca
and the presence of a well-defined, strongly
iridescent gorget and coronal stripe. De-
scriptions in this paper refer to definitive
male plumage. Simons’ specimen is clearly
assignable to the genus Heliodoxa in pos-
sessing a unique combination of characters:
(a) robust, moderately long (22.7 mm),
nearly straight bill (Fig. 1); (b) feathers ex-
tend forward on the bill obscuring the nos-
trils; (c) unmodified regimes; (d) tarsal
feathers extend to the base of toes; (e) mod-
erately forked tail (fork depth = 35.3 mm;
Fig. 2), (f) unspotted rectrices; (g) small
brilliant gorget; and (h) brilliant coronal
stripe.

VOLUME 117, NUMBER 1

Fig. 1.

According to Chubb (1919), Stephens &
Traylor (1983), and Paynter (1993), Si-
mons’ collecting itinerary overlapped the
known range of the genus Heliodoxa on the
Pacific slope of the Ecuadorian Andes
(prov. Azuay, Cafar, Chimborazo, Guayas,
and El Oro), and on the Amazonian slope
of the Andes in Peru (depto. Junin and
Puno) and Bolivia (depto. La Paz). For the
purposes of the hybrid diagnosis, I restrict-
ed the pool of potential parental species
(Graves 1990, Graves & Zusi 1990) to He-
liodoxa aurescens, H. rubinoides, H. lead-
beateri, H. schreibersii, H. branickii, H. im-
peratrix, and H. jacula jamesoni (taxonomy

A probable hybrid, Heliodoxa imperatrix X H. jacula jamesoni (BMNH 1902.3.13.2211).

of Schuchmann 1999). I measured selected
specimens with digital calipers (rounded to
the nearest 0.1 mm): wing chord; bill length
(from anterior extension of feathers); and
rectrix length (from point to insertion of the
central rectrices to the tip of each rectrix)
(Table 1). Pairs of rectrices are numbered
from the innermost (R1) to the outermost
(R5).

I evaluated the color of the breast at the
ventral midline and of the medial vane of
the dorsal surface of R4 (7 mm from tip)
with a calibrated colorimeter (CR-221
Chroma Meter, Minolta Corporation)
equipped with a 3.0 mm aperture. The mea-

Table 1.—Ranges (mean + standard deviation) of measurements (mm) of adult males of Heliodoxa imperatrix,
H. jacula jamesoni, and a probable hybrid, Heliodoxa imperatrix X H. jacula jamersoni (BMNH

1902.3.13.2211).

Heliodoxa imperatrix
(n = 10)

Wing chord 70.4—75.8
(WBA = 1.7)
Bill length 23.3—24.9
(23.4 + 0.8)
Rectrix 1 22.6-26.6
(4S) = 12)
Rectrix 2 27.9-32.9
(29.7 = 1.4)
Rectrix 3 38.7-46.5
(41.3 + 2.3)
Rectrix 4 51.0-60.3
(54.8 += 2.8)
Rectrix 5 62.2-76.2

(68.4 + 5.7)

Heliodoxa jacula jamesoni BMNH
(n = 12) 1902.3.13.2211

73.3-79.0 75.5
(75.9 + 1.8)

21.8-24.4 Del
@32 = 0.7)

32.7—35.5 DISeu
(3-7 25 10)

36.7-40.0 34.3
(38.5 + 1.0)

42.3-46.5 43.7
(44.4 + 1.1)

47.7-51.3 55.6
(CY7/ = 1123)

48.5-54.3 64.0
@ilo7/ 2 21)

12 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

suring head of the CR-221 uses 45° circum-
ferential illumination. Light from the pulsed
xenon arc lamp is projected onto the spec-
imen surface by optical fibers arranged in a
circle around the measurement axis to pro-
vide diffuse, even lighting over the mea-
suring area. Only light reflected perpendic-
ular to the specimen surface is collected for
color analysis. Colorimetric data from iri-
descent feathers are acutely dependent on
the angle of measurement, the curvature of
plumage surfaces in museum skins, and the
degree of pressure applied to the plumage
surface by the Chroma Meter aperture. In
order to reduce measurement variation, I
held the aperture flush with the surface of
the breast plumage or rectrix without de-
pressing it. The default setting for the CR-
221 Chroma Meter displays mean values
derived from three sequential, in situ mea-
surements. I repeated this procedure twice,
removing the aperture between trials. Thus,
each datum summarized in Table 2 repre-
sents the mean of six independent colori-
metric measurements.

Colorimetric characters were described in
terms of opponent-color coordinates (L, a,
b) (Hunter & Harold 1987). This system is
based on the hypothesis that signals from
the cone receptors in the human eye are
coded by the brain as light-dark (L), red-
green (a), and yellow-blue (b). The ratio-
nale is that a color cannot be perceived as
red and green or yellow and blue at the
same time. Therefore “redness” and
““sreenness” can be expressed as a single
value a, which is coded as positive if the
color is red and negative if the color is
green. Likewise, “‘yellowness” or “blue-
ness”’ is expressed by b for yellows and -b
for blues. The third coordinate, L, ranging
from 0 to 100, describes the “lightness” of
color; low values are dark, high values are
light. The more light reflected from the
plumage, the higher the L value will be. Vi-
sual systems in hummingbirds (e.g., Gold-
smith & Goldsmith 1979) differ signifi-
cantly from those of humans and the rele-
vance of opponent color coordinates to col-

ors perceived by hummingbirds is
unknown.

Results and Discussion

I considered hypotheses that the speci-
men represents (7) an undescribed geo-
graphic variant or genetic color morph of
one of the aforementioned species of Helio-
doxa; (ii) a hybrid; or (iii) an undescribed
species of Heliodoxa. Simons’ specimen
does not appear to represent an unknown
color morph or geographic variant of any
described species because of its unique tail
morphology (Table 1). As noted by Hartert
(1900), the specimen combines characters
of Heliodoxa imperatrix and Heliodoxa ja-
cula (Figs. 1-3; Tables 1, 2).

The hybrid diagnosis focuses on the
identification of apomorphic character
states of possible parental species in puta-
tive hybrids (Graves 1990). Complete dom-
inance and polygenic inheritance of plum-
age characters, however, may preclude or
obscure the expression of parental apomor-
phies in hybrids. When parental apomor-
phies are not identifiable, the parentage of
a hybrid may be indicated, although less
conclusively, by the presence or absence of
a suite of plesiomorphic characters.

The pool of potential parental species
may first be narrowed by focusing on the
absence of rufous or buff pigmentation in
the hybrid’s plumage. Because brown and
reddish-brown pigments appear to exhibit
consistent penetrance in hummingbird hy-
brids (Banks & Johnson 1961, Graves &
Newfield 1996), Heliodoxa rubinoides (1u-
fous on inner vanes of secondaries and pri-
maries; cinnamon-buff margins of breast
and abdomenal feathers), H. aurescens (ru-
fous pectoral band), and H. branickii (ru-
fous inner vanes of rectrices) can be elim-
inated from further consideration as paren-
tal species. In a similar fashion, H. schrei-
bersi (black throat, breast, and abdomen)
and H. leadbeateri (brilliant violet coronal
stripe; coppery-bronze hindcrown and
neck) are exceedingly unlikely to be paren-

VOLUME 117, NUMBER 1

Fig. 2.

tal species because they possess characters
not observed in the hybrid. Based on plum-
age characters, the hybrid is most likely the
product of the species, H. imperatrix X H.
jJacula jamesoni. Below, I present a synop-
sis of the essential evidence.

The visual display of iridescence in He-
liodoxa imperatrix and H. jacula has
evolved to be viewed head-on. Both paren-
tal species possess brilliant gorgets and cor-
onal stripes that exhibit metallic irides-
cence. In H. imperatrix, the green coronal
stripe is bluntly triangular in shape, extend-
ing from the base of the bill and narrowing
to a point along the midline of the crown
(even with the anterior edge of the eye).
The bluish-green coronal stripe in H. jacula
extends from the bill to the hindcrown
forming a coronal stripe. The coronal stripe
of the hybrid is intermediate in appearance
between those of H. imperatrix and H. ja-
cula. Heliodoxa imperatrix possesses a
small purplish-pink gorget that appears to
be surrounded by a field of dimly glowing,
greenish-black plumage when viewed head-
on. The blue gorget of H. jacula is sur-
rounded by a field of green plumage, which
is spangled with glowing iridescence when
viewed head-on. In the hybrid, the color

Dorsal surface of the rectrices of a probable hybrid, Heliodoxa imperatrix * H. jacula jamesoni
(BMNH 1902.3.13.2211).

and quality of iridescence exhibited by the
gorget (purple exhibiting pinkish tones at
certain angles) and the surrounding plum-
age are intermediate in appearance between
those of H. imperatrix and H. jacula.

The ventral plumage of Heliodoxa im-
peratrix exhibits brilliant golden-green iri-
descence on the lower breast, flanks, and
abdomen when viewed head-on. The breast
and abdominal plumage is significantly
darker in H. jacula and exhibits far less ir-
idescence than in H. imperatrix. The color
and quality of iridescence in the hybrid is
intermediate between those of the postulat-
ed parental species (Table 2). The rectrices
of H. imperatrix are dark bronzy-olive be-
coming progressively darker from R1 to
RS5, whereas those of H. jacula are bluish-
black (the lateral webs of R1 are tinted with
olive in some individuals). Rectrix color in
the hybrid is roughly intermediate between
that of the postulated parental species (Ta-
ble 2):

As a second step, the parental hypothesis
was tested with an analysis of size and ex-
ternal proportions (Table 1, Fig. 3). Mea-
surements of avian hybrids fall within the
mensural ranges exhibited by their parental
species as a consequence of a polygenic

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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VOLUME 117, NUMBER 1

from negligible to moderate: wing chord
(3.7%), bill length (0.9%), R1 (37.6%), R2
(29.6%), R3 (7.5%), R4 (10.3%), and R5
(32.3%). Measurements of the hybrid fall
within the cumulative range of parental
measurements for all seven characters and
within the parental means for five charac-
ters (wing chord, R1, R2, R3, R5). In sum-
mary, evidence obtained from plumage col-
or and pattern, as well as from external size
and shape, is consistent with the hypothesis
that Simons’ specimen is an intrageneric
hybrid between Heliodoxa imperatrix and
H.. jacula jamesoni.

Simons collected avian specimens on the
Pacific slope of the Ecuadorian Andes in
the provinces of Azuay, Chimborazo,
Guayas, El Oro, and Pichincha from 1 No-
vember 1898 to 12 July 1899 (Chubb 1919,
Paynter 1993). His northernmost collecting
locality, Guaillabamba, Pichincha (0°04’S,
78°21'W), lies in a semi-arid intermontane
valley some 30 km southeast of the zone of
sympatry for Heliodoxa imperatrix and H.
Jacula jamesoni in humid cloud forest on
the Pacific slope (see Ridgely & Greenfield
2001). This suggests one of three possibil-
ities: (1) Simons collected the specimen
along the Quito-Guaillabamba-Gualea road,
but on the Pacific slope; (2) he purchased
the specimen from a third party, possibly a
native collector; or (3) he obtained the spec-
imen at an unknown area of sympatry be-
tween the parental species on the Pacific
slope in west-central or southwestern Ec-
uador. Whatever the source, Simons’ spec-
imen represents the only known instance of
intrageneric hybridization in Heliodoxa.

Acknowledgments

I am grateful to Robert Prys-Jones, Mi-
chael Walters, Mark Adams, Don Smith,
and the Schliisselmeister, Frank Steinhei-
mer, of The Natural History Museum,
Tring, for permission to study Simons’
specimen and for loaning it for long-term
study. I thank Richard C. Banks and Rich-
ard L. Zusi for comments on the manu-

script. Travel was supported by the Re-
search Opportunities Fund, the Alexander
Wetmore Fund, and the Department of Ver-
tebrate Zoology, Smithsonian Institution.

Literature Cited

[Allen, J. A.] 1903. [Perry O. Simons, widely known
as an energetic and careful collector. . .]|—Auk
20:94-96.

Banks, R. C., & N. K. Johnson, 1961. A review of
North American hybrid hummingbirds.—Con-
dor 63:3-28.

Bleweiss, R., J. A. W. Kirsch, & J. C. Matheus. 1997.
DNA hybridization evidence for the principal
lineages of hummingbirds (Aves: Trochili-
dae).—Molecular Biology and Evolution 14:
325-343.

Buckley, P. A. 1982. Avian genetics. Pp. 21-110 in M.
Petrak, ed., Diseases of cage and aviary birds,
2nd ed. Lea and Febiger, Philadelphia, 680 pp.

Chubb, C. 1919. Notes on collections in the British
Museum, from Ecuador, Peru, Bolivia, and Ar-
gentina, part 1. Tinamidae-Rallidae.—Ibis (11th
series):1:1-55.

Goldsmith, T. H., & K. M. Goldsmith. 1979. Discrim-
ination of colors by the black-chinned hum-
mingbird, Archilochus alexandri.—Journal of
Comparative Physiology A 130:209-220.

Graves, G. R. 1990. Systematics of the ““green-throat-
ed sunangels” (Aves: Trichilidae): valid taxa or
hybrids?—Proceedings of the Biological Soci-
ety of Washington 103:6-25.

1998. Diagnoses of hybrid hummingbirds

(Aves: Trochilidae). 6. An intergenetic hybrid,

Aglaiocercus kingi X Metallura tyrianthina,

from Venezuela—Proceedings of the Biological

Society of Washington 111:511-520.

1999. Diagnoses of hybrid hummingbirds

(Aves: Trochilidae). 8. A provisional hypothesis

for the hybrid origin of Zodalia glyceria

(Gould, 1858).—Proceedings of the Biological

Society of Washington 112:491-502.

, & N. L. Newfield, 1996. Diagnoses of hybrid

hummingbirds (Aves: Trochilidae). 1. Charac-

terization of Calypte anna xX Stellula callipoe
and the possible effects of egg volume on hy-
bridization potential. Proceedings of the Biolog-

ical Society of Washington 109:755-763.

, & R. L. Zusi. 1990. An intergeneric hybrid
hummingbird (Heliodoxa leadbeateri X Helian-
gelus amethysticollis) from northern Colom-
bia——Condor 92:754-760.

Gray, A. P. 1958. Bird hyrids. Commonwealth Agri-
cultural Bureaux, Bucks, England, 390 pp.

Hartert, E. 1900. [Mr. Ernst Hartert exhibited two hy-
brids of hummingbirds].—Bulletin of the Brit-
ish Ornithologists’ Club 10:39-40

16 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Hunter, R. S., & R. W. Harold. 1987. The measurement
of appearance, 2nd edition. Wiley, New York,
All pp.

Panov, E. N. 1989. Natural hybridisation and etholog-
ical isolation in birds. Nauka, Moscow, 510 pp.

Paynter, R. A., Jr. 1993. Ornithological gazetteer of
Ecuador, 2nd edition. Museum of Comparative
Zoology, Harvard University, Cambridge, Mas-
sachusetts, 247 pp.

Ridgeley, R. S., & P. J. Greenfield. 2001. The birds of
Ecuador. Volume 1: status, distribution, and tax-
onomy. Cornell University Press, Ithaca, New
York, 848 pp.

Schuchmann, K. L. 1999. Family Trochilidae. Pp.
468-680 in Handbook of the Birds of the

World, vol. 5. Barn-owls to Hummingbirds (J.
del Hoyo, A. Elliott, & J. Sargatal, Eds.). Lynx
Edicions, Barcelona, 759 pp.

Sibley, C. G., & B. L. Monroe, Jr. 1990. Distribution
and taxonomy of birds of the world. Yale Uni-
versity Press, New Haven, Connecticut, 1111
PPp-

Stephens, L., & M. A. Traylor, Jr. 1983. Ornithological
gazetteer of Peru. Museum of Comparative Zo-
ology, Harvard University, Cambridge, Massa-
chusetts, 271 pp.

Zusi, R. L., & G. D. Bentz. 1982. Variation of a muscle
in hummingbirds and swifts and its systematic
implications—Proceedings of the Biological
Society of Washington 95:412—420.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(1):17—22. 2004.

Pholidochromis cerasina, a new species of pseudochromine dottyback
fish from the west Pacific (Perciformes: Pseudochromidae)

Anthony C. Gill and Hiroyuki Tanaka

(ACG) Fish Research Group, Department of Zoology, The Natural History Museum, Cromwell
Road, London SW7 5BD, U.K.:;
(HT) Jinguh Clinic, 2-2-79 Jinguh, Miyazaki, Miyazaki 880, Japan.

Abstract.—Pholidochromis cerasina is described from the 43.9-mm SL ho-
lotype from Talisei Island, off the northern tip of Sulawesi, Indonesia. It is
distinguished from its congener P. marginata (Lubbock) from Papua New
Guinea and the northern Solomon Islands in lacking both dark submarginal
markings on the median fins and prominent dark grey to black spots surround-

ing sensory pores on the head.

Fishes of the Indo-Pacific subfamily
Pseudochrominae were recently revised by
Gill (2003), who recognised 80 species in
10 genera, four of which were newly de-
scribed. One of the newly described genera,
Pholidochromis, was erected to accommo-
date a single species, Pseudochromis mar-
ginatus Lubbock, 1980, and distinguished
from other pseudochromine genera in hav-
ing the following combination of external
characters: lower lip complete (uninterrupt-
ed at symphysis); dorsal-fin rays III, 22;
anal-fin rays III, 13; scales in lateral series
28—32; dorsal and anal fins with well-de-
veloped scale sheaths; and predorsal scales
extending anteriorly to or forward of pos-
terior nostrils. It is also unique among pseu-
dochromid genera in having the following
combination of osteological characters:
three equal-sized supraneural bones; first
dorsal pterygiophore posterior lamina run-
ning most of the length of the bone; and
11—12 consecutive dorsal pterygiophores
inserting in a 1:1 relationship with inter-
neural spaces directly behind neural spine 4.

Gill (2003) recorded Pholidochromis
marginata from the east coast of Papua
New Guinea, Bougainville Island, and off
the northern tip of Sulawesi, Indonesia. The
latter record was based on a single speci-

men (USNM 136954) collected at Talisei
Island in 1909 by the United States Bureau
of Fisheries Steamer Albatross; it bears a
silk tag with the number “‘2038.”’ The spec-
imen differs from other examined speci-
mens (all of which were collected 50 or
more years after the Sulawesi specimen) in
lacking conspicuous dark spots on the head
and dark submarginal stripes on the median
fins. Although no comments on the condi-
tion of these markings were made in his
revision, the first author attributed their ab-
sence to the age of the specimen, with the
assumption that it was badly faded.

In 1995, the first author received a colour
illustration of an aquarium individual of a
pseudochromid from W. E. Burgess (for-
merly of Tropical Fish Hobbyist Publica-
tions Inc.). It was a pale pink, deep-bodied
fish with orange to red spots on the body
and median fins, and a yellow ring around
the eye. The first author was unable to iden-
tify 1t confidently with any known species,
but suggested that it was perhaps an unusu-
ally coloured individual of either Pseudoch-
romis fuscus Miller & Troschel, 1849
(which is often yellow with blue to grey
spots and a similar body shape) or a poor
illustration of P. marshallensis Schultz,
1953 (which, though usually more slender

18 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

with a darker ground coloration, has yellow
to orange or red spots on the body).

In May 2000, the second author sent the
first author a photograph of a pseudochrom-
id from a recent article in the Japanese
aquarium journal Aqualife, as well as ad-
ditional aquarium photographs of the spec-
imen. The fish depicted was very similar in
coloration and shape to the one in Burgess’s
illustration, thus rekindling interest in its
identity. A search of the first author’s col-
lection of pseudochromid photographs re-
vealed an illustration of a similar specimen
collected on the Albatross expedition (orig-
inal housed in the National Museum of Nat-
ural History, Smithsonian Institution). The
number “2038” was written in pencil next
to the illustrated fish.

As Fowler (1931) had reported on pseu-
dochromids collected by the Albatross, his
paper was searched in attempt to locate a
reference to the number “2038.” No such
reference was found, but a colour descrip-
tion closely matching the illustration was
found for a specimen numbered “22731”
from Talisse Island, which Fowler had iden-
tified as Pseudochromis xanthochir Bleeker,
1855 (a junior subjective synonym of P.
fuscus). The number “22731” refers to a
linen tag attached to a 45.0-mm-SL speci-
men of the pseudoplesiopine Pseudople-
siops typus Bleeker, 1858 (now registered
USNM 146624). However, the Albatross il-
lustration (and Fowler’s description) is ob-
viously not based on the specimen of P.
typus. Although P. typus may be pale pink
to pale grey with a ring around the eye
(which is red to black in life), it does not
possess red spots on the body. Moreover,
the illustration depicts a fish with relatively
short, broad pelvic fins, whereas they are
long and slender in specimens P. typus (n-
cluding the specimen in USNM 146624).
The other illustration and photographs of
aquarium specimens are also not referable
to P. typus.

However, the P. typus specimen was col-
lected from Talisse (=Talisei) Island on the
same date as the Albatross Pholidochromis

specimen (presumably from the same sta-
tion), and this, coupled with the close sim-
ilarity in body form and pelvic-fin shape,
led us to question whether the illustration
was of the Pholidochromis specimen, and
whether the “2038” may refer to the silk
tag number on that specimen. We therefore
asked S. L. Jewett and J. T. Williams of the
National Museum of Natural History to
check whether there were further details
that might corroborate this. Jewett consult-
ed the original illustration and responded
(pers. comm., 4 Aug 2000): ‘It not only
says 2038 in pencil, but Leonard Schultz
{former Curator of Fishes at USNM, and
author of a paper on the pseudochromid ge-
nus Labracinus, based mostly on Albatross
specimens] wrote a note in the margin that
says “see USNM 136954.’” She also noted
that there is a small tag in the jar containing
USNM 136954 indicating that the specimen
was drawn.

We therefore conclude that the illustra-
tion is based on the Pholidochromis speci-
men. Clearly, then, the absence of dark
markings in the specimen are not due to
fading, as such markings are not indicated
in the Albatross illustration, nor are they
evident in the illustrations or photographs
of live aquarium specimens. Thus, we con-
clude that the specimens represent a species
distinct from P. marginata, and therefore
describe it as new.

Materials and Methods

Methods of counting, measuring and pre-
sentation follow Gill (2003). Institutional
codes follow Leviton et al. (1985).

Pholidochromis cerasina, new species
Cherry Dottyback
Fig. 1

Pseudochromis xanthochir [non Bleeker,
1855]; Fowler, 1931: 32 (color descrip-
tion).

Pholidochromis marginata [non Pseudoch-
romis marginatus Lubbock, 1980]; Gill,

VOLUME 117, NUMBER 1

Fig. 1.
(Photo by P. Hurst.)

2003:000, fig. 5 (description and distri-
bution in part).

Holotype.—USNM 136954, 43.9 mm
SL, Indonesia, Sulawesi, Talisei (=Talisse)
Island, R/V Albatross, 9 November 1909.

Diagnosis.—Pholidochromis cerasinus
is distinguished from other pseudochromi-
nes in having the following combination of
characters: dorsal-fin rays III, 22; anal-fin
rays III, 13; scales in lateral series 29-30;
dorsal and anal fins with well-developed
scale sheaths; predorsal scales extending
anteriorly to just behind anterior nostrils;
and no prominent dark grey to black spots
surrounding sensory pores on head.

Description.—Dorsal-fin rays IJ, 22, at
least last 18 segmented rays branched (ray
preceding first apparent branched ray dam-
aged); anal-fin rays III, 13, at least last 12
segmented rays branched (anteriormost ray
damaged); pectoral-fin rays 19/19; upper
procurrent caudal-fin rays 6; lower procur-
rent caudal-fin rays 5; total caudal-fin rays
28; scales in lateral series 29/30; anterior
lateral-line scales 23/24; anterior lateral line
terminating beneath segmented dorsal-fin

Pholidochromis cerasina, USNM 136954, 43.9 mm SL, holotype, Talisei Island, Sulawesi, Indonesia.

ray 17/17; posterior lateral-line scales 10 +
0/9 + O; scales between lateral lines 3/3;
horizontal scale rows above anal-fin origin
12 + 1 + 3/13 + 1 + 3; circumpeduncular
scales 16; predorsal scales 21; scales behind
eye 3; scales to preopercular angle 4; gill
rakers 5 + 10; pseudobranch filaments 9;
circumorbital pores 17/18; preopercular
pores 9/8; dentary pores 4/4; posterior in-
terorbital pores 0.

Lower lip complete; dorsal and anal fins
with well-developed scale sheaths; predor-
sal scales extending anteriorly to just be-
hind anterior nostrils; posterior margin of
opercle with 4 inconspicuous serrations;
outer gill rakers of ceratobranchial-1 with
teeth mostly confined to raker tips; anterior
dorsal-fin pterygiophore formula S/S/S + 3/
1 + WI/WI/II/I/I/I/I/1/1/1 + 1; dorsal-
fin spines pungent and moderately slender;
anterior anal-fin pterygiophore formula 3/1/
1 + 1/I/1/1 + 1; anal-fin spines pungent
and moderately slender to stout, second
spine stouter than third; pelvic-fin spine
slender, tip weakly pungent; second seg-
mented pelvic-fin ray longest; caudal fin

20 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

rounded (inferred for holotype from Alba-
tross illustration); vertebrae 10 + 16; epi-
neurals 13; epurals 3.

Upper jaw with 2 pairs of curved, en-
larged caniniform teeth anteriorly, medial
pair smallest, and about 6 (at symphysis) to
2-3 (on sides of jaw) irregular rows of
small conical teeth, outermost of rows of
teeth much larger and more curved than
those of inner rows; lower jaw with 2 pairs
of curved, enlarged caniniform teeth ante-
riorly, medial pair smallest, and about 5 (at
symphysis) to | (on sides of jaw) inner
rows of small conical to caniniform teeth,
those on middle of jaw large and canini-
form; vomer with 1—2 rows of small conical
teeth arranged in chevron; palatine with 2—
3 rows of small conical teeth arranged in
elongate ovoid patch, anterior part of tooth
patch more-or-less contiguous with postero-
lateral arm of vomerine tooth patch; ectop-
terygoid edentate; tongue moderately point-
ed and edentate.

As percentage of SL: head length 28.0;
orbit diameter 9.1; snout length 7.7; fleshy
interorbital width 6.2; bony interorbital
width 3.9; body width 13.9; snout tip to
posterior tip of retroarticular bone 15.9;
predorsal length 35.3; prepelvic length
35.3; posterior tip of retroarticular bone to
pelvic-fin origin 21.0; dorsal-fin origin to
pelvic-fin origin 34.9; dorsal-fin origin to
middle dorsal-fin ray 34.4; dorsal-fin origin
to anal-fin origin 46.2; pelvic-fin origin to
anal-fin origin 27.6; middle dorsal-fin ray
to dorsal-fin termination 26.2; middle dor-
sal-fin ray to anal-fin origin 33.5; anal-fin
origin to dorsal-fin termination 39.0; anal-
fin base length 29.6; dorsal-fin termination
to anal-fin termination 17.1; dorsal-fin ter-
mination to caudal peduncle dorsal edge
10.9; dorsal-fin termination to caudal pe-
duncle ventral edge 19.6; anal-fin termina-
tion to caudal peduncle dorsal edge 19.8;
anal-fin termination to caudal peduncle ven-
tral edge 10.7; first dorsal-fin spine 2.7; sec-
ond dorsal-fin spine 5.2; third dorsal-fin
spine 7.1; first segmented dorsal-fin ray
13.7; fourth from last segmented dorsal-fin

ray broken; first anal-fin spine 3.0; second
anal-fin spine 5.7; third anal-fin spine 7.1;
first segmented anal-fin ray broken; fourth
from last segmented anal-fin ray broken;
third pectoral-fin ray broken (both sides);
pelvic-fin spine 9.8; second segmented pel-
vic-fin ray 22.6; caudal-fin length not de-
termined (ray tips broken).

Live coloration (based on a color illus-
tration of holotype and photographs and an
illustration of aquarium specimens).—Head
and body pale pinkish grey to pinkish olive
dorsally, becoming pale pink to pale yellow
or white ventrally; posttemporal pore in
dusky grey spot (not apparent in illustra-
tions); orbital rim yellow to bright orange
or bright red; pale blue to mauve stripe ex-
tending from anteroventral edge of eye to
middle of upper lip (not apparent on illus-
tration of holotype); iris silvery white, blue
dorsally, with grey to blue suboval ring
around pupil; body with small (about half
pupil size) pale orange to bright orange or
bright red spots, these best developed dor-
sally and posteriorly, and more or less ar-
ranged along horizontal scale rows; dorsal
and anal fins pale pink to pale blue with
blue distal margin, and 2—S horizontal rows
of pale orange to bright orange or crimson
spots (crimson spots encircled with pale
pink in photographed individuals); caudal
fin pale pink to pale blue with blue distal
margin and bright red to crimson spots (en-
circled with pale pink in photographed in-
dividuals), these irregularly arranged on
basal part of fin, becoming arranged in con-
vex columns on remainder of fin; pectoral
fins hyaline with pinkish to yellowish hue;
pelvic fins pale pink to pale blue.

Preserved coloration.—Head and body
pale brown, paler ventrally; posttemporal
pore in dusky grey spot; fins whitish hya-
line to plain hyaline.

Habitat and Distribution.—No_ habitat
data are known for the holotype. We also
lack precise locality or habitat information
for aquarium individuals of the species;
however, K. Endoh (pers. comm.) informed

VOLUME 117, NUMBER 1

Fig. 2.
Province, Papua New Guinea. (Photo by P. Crabb; after Gill, 2003:fig. 20.)

us that they were collected in the Philippine
Islands.

Comparisons.—Pholidochromis cerasina
agrees closely with its congener, P. margin-
ata (Fig. 2), in most details, but differs in
lacking conspicuous dark spots around the
sensory pores on the head (only the post-
temporal pore of P. cerasina has an incon-
spicuous dusky grey spot whereas P. mar-
ginata has conspicuous dark grey to black
spots on at least the posterior suborbital,
upper preopercular, anterior interorbital,
posttemporal and parietal pores) and in
lacking dark submarginal markings on the
median fins (present as dark grey to black
convex marking on the caudal fin, and short
dark grey to black stripe on the posterior
part of the dorsal and anal fins in P. mar-
ginata).

Values for 13 morphometric characters of
the holotype of P. cerasina lay at the ex-
treme or outside ranges observed in P. mar-
ginata (15 specimens, 27.2—45.6 mm SL).
Although more specimens are needed to de-
termine whether these are truly diagnostic,
they are at least suggestive. We also docu-
ment these in order to correct Gill’s (2003)

Pholidochromis marginata, CAS 65783, 32.4 mm SL, southern side of Nagada Harbour, Madang

description of P. marginata, as this includ-
ed data from the holotype of P. cerasina.
The characters are as follows (values ex-
pressed as % SL, and given first for P. cer-
asina, followed by P. marginata): fleshy in-
terorbital width (6.2; 5.0—6.1); predorsal
length (35.3; 36.0—39.0); middle dorsal-fin
ray to dorsal-fin termination (26.2; 20.5—
25.2); anal-fin termination to caudal pedun-
cle ventral edge (10.7; 10.8—12.4); first dor-
sal-fin spine (2.7; 2.7—5.1); second dorsal-
fin spine (5.2; 5.1—7.9); third dorsal-fin
spine (7.1; 7.0—10.3); first segmented dor-
sal-fin ray (13.7; 11.0—13.9); first anal-fin
spine (3.0; 3.7—5.7); second anal-fin spine
(5.7; 6.8-9.6); third anal-fin spine (7.1;
7.0—11.1); pelvic-fin spine (9.8; 9.6—13.2);
and second segmented pelvic-fin ray (22.6;
22.6—26.3).

Pholidochromis cerasina might also be
confused with Pseudochromis fowleri Her-
re, 1934, from Sabah and the Philippine Is-
lands, and Pseudochromis fuscus, from
throughout the West Pacific, which it re-
sembles in general body shape. These spe-
cies differ from Pholidochromis cerisina in
having an incomplete lower lip (interrupted

22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

at symphysis) and more segmented dorsal-
fin rays (23-25, usually 24 in fowleri and
25-29 in fuscus versus 22 in cerasina).
Remarks.—The live coloration of P.
marginata is unknown, but, accepting the
dark pigmentation on the head and median
fins, is likely to be similar to P. cerasina.
Moreover, as noted by Gill (2003), some
specimens of P. marginata have pale spots
on the body and median fins, and these pos-
sibly correspond with the red to orange
spots shown by P. cerasina.
Etymology.—tThe specific epithet is from
the Latin cerasinus, meaning “‘of cherry.”
It alludes to the cherry-like bright orange to
red spots on the body and median fins.
Material examined.—See above.

Acknowledgments

We thank S. L. Jewett and J. T. Williams
for lending the holotype for study, and for
their help in checking the history of the AI/-
batross specimen and its illustration. We
thank W. E. Burgess and K. Endoh for
sending an illustration and photographs, re-
spectively, of the species. S. E. Reader as-
sisted with radiographing the holotype.

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117(1):23-—34. 2004.

Redescription of Cambaroides japonicus (De Haan, 1841)
(Crustacea: Decapoda: Cambaridae) with allocation of a type locality
and month of collection of types

Tadashi Kawai and J. E Fitzpatrick, Jr.*

(TK) Hokkaido Nuclear Energy Environmental Research Center, 261-1 Miyaoka, Kyowa,
Hokkaido 045-0123, Japan, e-mail: kawaita@fishexp.pref.hokkaido.jp;
(JFF) Museum of Natural History, Tulane University,
Belle Chasse, Louisiana 70037, U.S.A.

Abstract.—The Japanese crayfish, Cambaroides japonicus (De Haan), is re-
described and illustrated, and details of its distribution and morphological var-
iation are provided. Notable character differences between the populations of
Honshu Island and Hokkaido Island indicate that gene flow between them is
precluded. Analysis of geographical variation demonstrates that the undesig-
nated type locality of the species is in central-western Aomori Prefecture, Hon-
shu. The analysis of the gastrolith weights of the lectotype and possible top-
otypes indicates that the lectotype was collected in June.

The German medical doctor, Philip Franz
von Siebold, was the first to introduce the
natural history of Japan to European aca-
demics (Siebold 1897). He also taught Eu-
ropean medicine to traditional Japanese
practitioners, and on 23 February 1826, at
Shimonoseki City, Yamaguchi Prefecture,
received specimens of a crayfish used as a
Japanese drug from his student, Kosai Ya-
maguchi (Siebold 1897). These were sent
to the Netherlands and were described as
Cambaroides japonicus by De Haan
(1841). The brief description of the species
included no locality or other collection data.
Heretofore, taxonomic studies of C. japon-
icus have been limited to examining cyclic
dimorphism (Kawai & Saito 1999), and the
genus Cambaroides has yet to be the sub-
ject of modern morphological studies.

This paper provides a redescription of C.
japonicus, allocates a type locality based on
an analysis of geographic variation, and
suggests a probable month of collection of

* Deceased 11 July 2002.

the types based on an analysis of monthly
changes in gastrolith weight.

Abbreviations used in the text are: GVM,
geographical variation in morphology;
POCL, postorbital carapace length; RMNH,
Nationaal Natuurhistorisch Museum, Lei-
den; and TCL, total carapace length.

Calculation of GVM: The geographical
variation of each specimen was divided into
three different levels (see Fig. 1), and the
mean of the levels among specimens was
calculated in each collection (Appendix I).
The mean in each collection was classified
into three degrees; 1.0—1.6, 1.7—2.3, 2.4—
3.0.

Cambaroides japonicus (De Haan, 1841)
Fig. 2, Table 1

Diagnosis.—Body pigmented; eyes well
developed, pigmented. Carapace subcy-
lindrical, dorsal and lateral surfaces with
large punctations, without tubercles; cervi-
cal spines absent. Rostrum acuminate,
broadest at base; margins thickened, strong-
ly convergent, lacking spines or tubercles;
median carina present, often very weak;

24 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

A, rostrum

DREMEL RK ote tere es,

B, telson

C, sternal
plates

Fig. 1.

Definition of morphological variations. A-1, median carina present on rostrum; A-2, carina inter-

mediate between A-1 and A-3; A-3, carina almost absent. B-1, caudomedian excavation present on telson; B-2,
excavation intermediate between B-1 and B-3; B-3, excavation absent; C-1, sternal plates closed; C-2, sternal
plates intermediate between C-1 and C-3; C-3, sternal plates open.

acumen comprising 27.5—-59.5% (X =
47.8%, SD = 4.6, n = 200) of rostrum
length, latter consisting 14.7—26.7% OX =
17.3%, SD = 2.5, n = 200) of TCL. Areola
112.3 0% = 1.9%, SD = O2, 1m = BOO)
times as long as broad, constituting 26.3—
AN 1% 0X = 25%, SD = 3.1, 2 = ZOO) oF
TCL and 30.9-46.9% (X = 35.5%, SD =
3.3, n = 200) of POCL. Antennal scale 1.6—
2.8 (X = 2.2%, SD' = 0.3, n = 100) times
as long as broad, widest at midlength, lat-
eral margin thickened, terminating in large,
stout spine. Pleura of somites 2 and 3 with
rounded to subtruncate ventral margins.
Palm of chela of cheliped with scattered

large punctations on dorsal, lateral, and
ventral surfaces, without setae; palm inflat-
ed, width 1.2-1.6 (X = 1.4%, SD = O01, n
= 100) times length of mesial margin.
Large punctations on dorsal, lateral and
ventral surface of fixed finger and dactyl.
Hooks present on ischia of second and
third pereiopods in males, hooks simple and
not reaching basioischial articulation. In
situ gonopods (first pleopods) of adult male
symmetrical, bases not contiguous. In me-
sial aspect (Fig. 2A), apex directed cephal-
odistally nearly 45° to axis of shaft, with
strong endopodite and protopodite; apex
(Fig. 2B) sclerotized, at least distally, ce-

VOLUME 117, NUMBER 1 25

Nese

Fig. 2. Cambaroides japonicus (De Haan, 1841), all figures from lectotype (RMNH 5602, RMNH 5603),
except B (redrawn from Hart 1953), I from paralectotype female (RMNH 2912), and j from paralectotype
male#1: A, mesial view of first pleopod; B, mesial view of distal portion of first pleopod; C, lateral view of
mandible; D, ventral view of ischum of third maxilliped; E, lateral view of first three abdominal segments; F
dorsal view of carapace; G, epistome; H, dorsal view of distal podomeres of right cheliped; I, annulus ventralis;
J, proximal podomeres of pereiopods; K, dorsal view of telson and uropods. Line = 2 mm.

26 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Measurements of type series of Cambaroides japonicus.

Lectotype

Carapace

Total length 31.6

Postorbital length 25.4

Width 16.4

Height 11.1
Areola

Length 98

Width 4.9
Rostrum

Length 6.6

Width 7.3
Chela

Length of mesial margin of palm 10.9

Width of palm 11.4

Length of lateral margin 24.5

Length of dactyl 14.2
Abdomen width 14.7

phalolaterally swollen into straight, sub-
acute, stout, cephalodistally directed mesial
process, cephalic process, and central pro-
jection with blade-like caudal process; 3
subequal spines near mid-width of apex,
length about one-tenth of width of apex.
Proximal part of gonopod subcylindrical in
cross section, becoming subtriangular dis-
tally. Sperm groove along caudomesial face
of gonopod shallow, open between mesial
process and central projection, ending in
relatively blunt tip. Adult male gonopod
with “juvenile suture’’.

Annulus ventralis (Fig. 21) immovable,
symmetrical, rounded in outline, about 1.2
times as long as wide. Preannular plate
transversely subdivided into 2 subtriangular
plates, cephalic margin of anterior part
broadly attached to preceding sternite, mid-
dle section of posterior part with shallow
depression as fossa without sinus. Postan-
nular sclerite subcircular, about 1.7 times as
broad as long, and 0.5 times as wide as an-
nular plate.

Measurements of type specimens provid-
ed in Table 1.

Description of lectotype.—Cephalotho-

Paralectotype Paralectotype Paralectotype

male #1 male #2 female
WM 21.7 30.4
18.6 17.6 25.1
11.1 11.7 15.1
8.7 9.4 11.1
3 7.0 9.4
3.7 2.6 4.2
4.7 43 Voll
4.9 4.5 V3
6.9 5.6 8.9
7.6 733) 8.9
16.4 15.8 20.4
8.3 8.0 10.8
10.6 10.2 17.3

rax (Fig. 2F) subcylindrical, slightly com-
pressed laterally; dorsoventrally depressed
(greatest width of thoracic section 1.5 times
depth); POCL 80.4% of TCL. Areola 2.0
times longer than wide, dense punctate,
length 31.0% of TCL (38.6% of POCL).
Rostrum acuminate, tip barely reaching dis-
tal margin of antennal scale and midlength
of ultimate podomere of antennal pedencle;
acumen comprising 48.4% of rostrum
length, latter consisting 20.9% of TCL;
floor (dorsal surface) of rostrum dense
punctate; median carina nearly absent.

Postorbital ridges poorly defined. Sub-
orbital angle obtuse, without tubercle or
spine. Antennal scale with strong distolat-
eral spine, tip reaching tip of rostrum and
midlength of ultimate podomere of anten-
nular peduncle.

Greatest width of abdomen 89.6% great-
est width of carapace. Proximal podomere
of uropod lacking spine or tubercle on lat-
eral lobe, mesial lobe broadly rounded; me-
sial ramus of uropoda with caudolateral
spine, and submedian dorsal ridge termi-
nating in small caudomedian spine, tip of
which not reaching caudal margin; lateral

VOLUME 117, NUMBER 1

ramus with stout caudolateral spine; lateral
ramus divided into cephalic and caudal sec-
tions, separated by transverse flexure bear-
ing spines. Telson divided into cephalic and
caudal sections, each caudolateral corner
with pair of stout, fixed spines. Caudal mar-
gin of telson with deep median excavation.

Epistome (Fig. 2G) with subovate ce-
phalic lobe bearing prominent cephalome-
dian projection, margins of lobe markedly
elevated; fovea of epistome scarcely visi-
ble; central portion of epistome with pair of
transverse grooves, and deep transverse
grooves along cephalic margin of weakly
arched zygoma. Third maxilliped (Fig. 2D)
with mesial margin bearing 21 denticles;
mesial half of ischium with row of clusters
of long, stiff setae. Incisor ridge of right
margin (Fig. 2C) with 5 corneous denticles.

Palm of right chela (Fig. 2H) subovate in
cross section, moderately depressed dorso-
ventrally. Total chela length 77.5% of TCL.
Palm 2.1 times as long as wide, length of
mesial margin 44.5% of total chela length;
dorsal surface with deep, widely scattered
punctations, which become scarce laterally
and caudolaterally; ventral surface less
punctate. Dorsal surface of both fingers
with poorly defined, longitudinal submedi-
an ridge, and rows of deep punctations; tip
of fingers corneous, subacute. Opposable
surface of fixed finger with row of 9 tuber-
cles, third from base largest. Opposable sur-
face of dactyl with row of 5 tubercles;
length of dactyl 1.3 times length of mesial
margin of palm. Carpus (Fig. 2H) longer
than broad, dorsal surface with prominent
longitudinal furrow, lateral and mesial sur-
face with large, crowded punctations; me-
sial surface with large, blunt subdistal
spine, lateral surface with proximal spine;
ventral surface with oblique furrow, short
longitudinal furrow, and deep punctations.
Merus with row of prominent tubercles on
ventromesial margin, punctuate dorsally
and ventrally.

Gonopods as described in “‘Diagnosis’’.
In addition, tips of gonopods extending be-

yond cephalic margin of coxae of fourth pe-
reiopods.

Description of paralectotype male#].—
Differing from lectotype as follows: great-
est width of thoracic section 1.3 times
depth. POCL 84.2% of TCL. Areola length
33.0% of TCL (39.2% of POCL). Acumen
comprising 46.1% of rostrum length, latter
consisting 21.3% of TCL. Greatest width of
abdomen 95.5% greatest width of carapace.
Total chela length 74.2% of TCL; palm of
right chela 2.2 times as long as wide; length
of mesial margin 42.1% of total chela
length. Dactyl length 1.2 times length of
mesial margin of palm; opposable surface
of fixed finger with 10 tubercles; opposable
surface of dactyl with 6 tubercles.

Description of paralectotype—female.—
Differing from lectotype, except in second-
ary sexual characteristics, as follows: great-
est width of thoracic section 1.4 times
depth. POCL 82.6% of TCL. Areola 2.2
times as long as broad. Areola length 30.9%
of TCL (37.5% of POCL). Acumen com-
prising 50.0% of rostrum length, latter con-
sisting 23.4% of TCL. Opposable surface
of fixed finger with 6 tubercles, proximal
largest; opposable surface of dactyl with 6
tubercles, length of finger 1.2 times length
of mesial margin of palm.

Disposition of types.—All dry and lack-
ing most appendages. Lectotype: 1 male,
RMNH 5602, RMNH 5603. Paralectotypes:
2 males and | female, RMNH 2912. The
lectotype has the mark ““@”’ written on the
areola, but is a male. A milky-white gastro-
lith is included with the lectotype. Its dry
weight (dried at 80°, 48 hr) is 0.0305 g, and
its shape is semi-globular, with the greatest
diameter 4.5 mm, the least diameter 4.2
mm, and the greatest height 1.8 mm.

Type locality.—No type locality for C.
japonicus has ever been designated. In or-
der to establish a type locality, we exam-
ined geographic variation in morphology
(GVM) to identify any unique characters
that might be displayed by the type speci-
mens. Earlier, Fitzpatrck (1995) detected
possible geographically defined races based

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

145°E
4sN “oly
A
Sakhalin Hokkaido
ie ae
( 140 “67 : =
“\ Hokkaido 4 :
‘ a re 10
an ; ——_——= 28
z 40°N- 21 * 11
: ‘é Z L » we 30 29 16 4 YA
~— Korean \ es ie
°° Pen. { Honshu / 343 me :
sh be ~ i
S20 eo
ie ob
, 130 140°E é is
ray a
ES Uae Aomori Pref.
“e330 6064 7\
40° = ee 77 (ee
AkitaePref. ( Iwate Pref.
i | =
oe, fog

Fig. 3.
sampling sites listed in Appendix I.

on a distinct rostral carina and pleural mar-
gin shape of the abdominal segment. Sam-
ples, however, were too small and too wide-
ly scattered for definite conclusions. In our
larger series from far more localities, we
examined the presence or absence of a me-
dian carina on the rostrum, median exca-
vation in the caudal margin of the telson,
and open or closed sternal plates. The GVM
was Classified into three levels (Fig. 1), and
mean GVM in each collection was sum-
marized according to three categories by
previously mentioned calculation (Fig. 4).
Three GVM characters were found to be
common in all the type series specimens,
and similar to characters found only in
specimens from central-western Aomori
Prefecture, Honshu (Figs. 1—4, Appendix I,
55—62). This strongly suggests that the type

Known geographical range of Cambaroides japonicus (De Haan, 1841). Numbers correspond to

specimens were collected in that area. The
central western Aomori Prefecture was de-
signed as a probable locality of the type se-
ries. Kurimi (1811) and Ohtsuki (1817), in
a paper published at the time Siebold re-
ceived specimens of C. japonicus, remarked
that the species commonly inhabited cen-
tral-western Aomori Prefecture. This lends
support to our assumption about the type
locality.

Date of type collection.—Monthly sam-
plings of C. japonicus were made from
April to November 1989 in Iwaki City (Fig.
3, Appendix I, 59), in the central-western
part of Aomori. For each sampling, gastro-
liths were removed from the stomachs of
30 individuals, ranging in size from 17.7
mm to 25.4 mm POCL, which corresponds
to size range of the types. The result indi-

VOLUME 117, NUMBER 1

Aomori

@ Pref.
ree

B, caudomedian

: AkitaePref.
excavation of telson

Iwate Pref.

C, sternal plates

Fig. 4. Geographical variation of (A) carina on rostrum, (B) caudomedian excavation of telson, and (C)
sternal plates. Solid circle, 1.0—1.6 level of mean GVM in collection; semi-open circle, 1.7—2.3; open circle,

2.4—3.0. A, Sapporo City; A’, Kazuno City.

cates that the dry weight (dried at 80°, 48
hr) of the gastrolith from lectotype (0.0305
g) is similar to that of the June sample (Fig.
5). Thus, it is believed the type series was
likely collected in June.

Range and specimens examined.—We
examined a total of 405 specimens from
Hokkaido Islands and four of its larger

nearby islands (Rebun, Rishiri, Teuri, and
Yagishiri), as well as the northern part of
Honshu Island (major parts of Aomori Pre-
fecture, and northern part of Akita and Iwa-
te Prefecture). Information on sampling
sites is provided in Fig. 3 and Appendix I.

As far as known, C. japonicus is endemic
to the entire Japanese Archipelago. How-

30 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

S ‘2
° =)
5 ui

=
°
v9)

Weight of gastrolith (g)
i=)
°

0 ——
A M J

Fig. 5.

J A S O N

Monthly change of dry gastrolith weight in Cambaroides japonicus (De Haan, 1841) from Iwaki

City, Aomori Prefecture (see Fig. 3, 59). Vertical bar indicates SD. Dotted horizontal line shows the dry gastrolith

weight of the lectotype (0.0305 g).

ever, Okada (1933:155—156) mentioned that
“Mr. T. Urita, a director of the Girl’s High
School at Maoka in south part of Sakhalin,
U.S.S.R., informed me that C. japonicus
seems not to occur in the stream and rivers,
and that if it is found in anywhere in Sa-
khalin, it is very rare; however, I examined
Outomari of Sakhalin specimens in the col-
lection of Professor Iijima of Tokyo Impe-
rial University, these are preserved in the
Zoological Institute, Faculty of Sciences,
Tokyo Imperial University”. And Urita
(1942:39) said “I consequently spent con-
siderable time and labour in search of this
species, especially in Outomari and its
neighbourhood, but unfortunately, without
success; presumably, this species does not
exist here in south Sakhalin, not, at any
rate, at present”. On 11 November 2001,
all specimens in the Tokyo Imperial Uni-
versity were transferred to the University
Museum, the University of Tokyo, but no
specimen from the Sakhalin could be found
there.

Size.—The largest lake specimen is a
male from Lake Akan, Hokkaido, measur-
ing 39.2 mm POCL; the largest brook spec-
imen is a male from Hamamasu with a

POCL of 34.3 mm. The smallest ovigerous
female is 17.6 mm POCL.

Variation.—Most variations were noted
in the number and comparative sizes of tu-
bercular ornamentation, particularly on the
cheliped. The caudolateral corner of the ce-
phalic section of the telson bears one to
three fixed spines. The lateral margin of the
telson of most specimens gently tapers to a
rounded caudal margin, but in some the lat-
eral margins are subparallel and the caudal
margin is flat. Some specimens have bosses
between the sternal plates (Fig. 21), but in
most specimens these bosses are nearly ab-
sent.

Color.—Dorsal and lateral surfaces of
cephalothorax, abdomen, chelae, and tail
fan dark brown to chocolate, ventral surface
light brown. Ventral surface of chela dark
orange. Tips of pereiopods dark orange.
Caudal process and three spines of distal
adult male gonopods amber. Background
colors translucent to light brown in freshly
molted individuals. The whitish-blue col-
orations or “blue color phase” (Fitzpatrick
1987) on the dorsal and lateral surfaces of
thoracic carapace, abdomen, chelae, and tail
fan, was found in specimens from Abashiri,

VOLUME 117, NUMBER 1

Obihiro, Iwamisawa, and Hamamasu, Hok-
kaido Prefecture, and in Shichinohe, Ao-
mori Prefecture (see Appendix I).

Crayfish associates and conservation sta-
tus.—During the past decade, local extinc-
tions of C. japonicus have been reported
from throughout its range. In eastern Hok-
kaido its numbers have been declining rap-
idly, while population numbers of the intro-
duced crayfish, Pacifastacus leniusculus
(Dana, 1852) in the same area have been
increasing (Kawai et al. 2002). Also, Kawai
et al. (2002) demonstrated that following
the introduction of P. leniusculus into Lake
Kussharo and Lake Shikaribetsu, C. japon-
icus disappeared. Pacifastacus leniusculus
is known to be a vector of crayfish plague
fungus, Aphanomyces astaci (Schikora), to
which it is resistant, but to which C. japon-
icus 1s highly susceptible (Unestam 1969).
It is possible that Aphanomyces may be a
factor affecting displacement of C. japoni-
cus at some localities, but there is as yet no
investigation of infection to the natural pop-
ulations in Hokkaido. The mechanisms un-
derlying the negative impacts of P. lenius-
culus on C. japonicus required further in-
vestigation.

Cambaroides japonicus was designated
an endangered species by the Japanese
Fisheries Agency in 1995 and by the Jap-
anese Environmental Agency in 2000.

Ecological notes.—Cambaroides japon-
icus appears to be restricted to lentic habi-
tats, either lakes or small brooks in which
current velocity is less than 10.0 cm/sec. In
brooks, the species is found beneath boul-
ders, or burrows in the banks. It appears to
be a secondary burrower, and retreats un-
derground to remain below the frost line in
winter. Females enter burrows prior to ovu-
lation, and remain in them to lay eggs. Most
burrows are Y- or T-shaped, with two open-
ings slightly above or below the water sur-
face.

Reproduction.—Mating in C. japonicus
is unique (Kawai & Saito 2001). The male
moves beneath the female to deposit its
spermatophore, and does not grasp the fe-

male with its chelae. In Hokkaido, mating
pairs were encountered only in September
and October, and ovigerous females during
the subsequent May. Spermatozoa obvious-
ly are stored in the annulus ventralis for a
six-month period during winter. Number of
ova ranges from 50 to 100, and egg diam-
eter is 2.3—2.7 mm.

Name in Japanese.—In Japan, it is usual
for organisms to have one or more local
names. To prevent possible ambiguity in
this pragmatic system, and make it easier to
incorporate taxonomic and distributional in-
formation, the common, Japanese name
Zarigani, is proposed. This name, which re-
fers to an animal that moves backward
(Ohtsuki 1817), is also mentioned in older
papers (e.g., Kurimi 1811). The names “‘Sa-
rugani’’ which is the local name in Aomori
Prefecture, and “‘Sarukani,” the local name
in Akita Prefecture, means “the backward
creeping crab.”’ Two local names are on the
label attached to the specimens of C. ja-
ponicus at Saito Ho-Onkai Museum, Sen-
dai, Japan (Nos. 1039, 1369). Also, the
Ainu people, former occupants of Hokkaido
and northern Honshu, know C. japonicus as
‘““Tekinpekorupe,” alluding to an armed
knight (Ohtsuki 1817).

Discussion

Cambaroides japonicus occurs in north-
ern parts of Honshu and Hokkaido Islands
(Fig. 3). It is likely that populations of the
species inhabiting certain areas of Honshu
were introduced from Hokkaido, but differ-
ences in the GVM (Fig. 4, Appendix I) in-
dicate that the majority of populations on
Honshu are native. An exception is seen in
the GVM of specimens from Kazuno City
(A’), Akita Prefecture, Honshu, which
agrees with that of Sapporo City (A), Hok-
kaido. In 1943, a locality report in a small
stream in Kazuno City originated from in-
troduction of a population in Sapporo City
(Mr. T. Komoriya, Japanese regional report
1978).

The distribution of Asian branchiobdel-

32 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

lidans, which are symbionts on crayfish, in-
cluding C. japonicus, may shed some ad-
ditional light on this issue, since there is a
high degree of endemism in the various
species. Cirrodrilus aomorensis and C. tsu-
garensis occur in Honshu (Gelder & Oht-
aka 2000), while C. inukaii and C. uchidai,
and others occur only in Hokkaido (Ya-
maguchi 1934). There is no overlap in the
natural distributions of these two species.
However, C. inukaii and C. uchidai have
both been found in Kazuno City, Akita Pre-
fecture, northern Honshu, an occurrence
that might be explained by an introduction
of C. japonicus from Hokkaido (Gelder &
Ohtaka 2000).

Acknowledgments

We thank A. Ohtaka, J. E. Cooper, and
Y. Hanamura, who offered many useful
suggestions concerning the present study.
Thanks are extended to C. H. J. M. Fransen,
S. E Mawatari, M. Takeda, K. Sakamoto,
T. Urano, Y. Yabumoto, G. Scholtz, H. Hay-
ashi, Y. Kobayashi, K. Nakata, and T. Ya-
maguchi, who were most generous with
their time, their collections, and their per-
sonal solicitude. Figure | was drawn mostly
by M. Tanaka.

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of the ‘Form’ and further morphological char-
acters for the crayfish genus Cambaroides
(Cambaridae). Pp. 82—89 in M. and M. M. Kel-
ler, B. Oidtmann, R. Hoffmann, and G. Vogt,
eds., Freshwater crayfish 12. Papers from the
Twelfth Symposium of the International Asso-
ciation of Astacology, Weltbild Verlag, Augs-
burg, Germany.

,& . 2001. Observations on the mating

behavior and season, with no form alternation,

of the Japanese crayfish, Cambaroides japoni-
cus (Decapoda, Cambaridae), in lake Koma-
dome, Japan.—Journal of Crustacean Biology

21:885-—890.

, K. Nakata, & T. Hamano. 2002. Temporal
changes of the density in two crayfish species,
the native Cambaroides japonicus (De Haan)
and the alien Pacifastacus leniusculus (Dana),
in natural habitats of Hokkaido, Japan. Pp. 198—
206 in G. Whisson and B. Knott, eds., Fresh-
water crayfish 13. Papers from the Thirteenth
Symposium of the International Association of
Astacology, Curtin Print and Design, Perth,
Australia.

Kurimi, Z. 1811. Senchufu. Kouwa Shuppan, Tokyo,
534 pp. (rewritten in 1982)

Ohtsuki, B. 1817. Ranwantekihou. Kouwa Shuppan,
Tokyo, 524 pp. (rewritten in 1980)

Okada, Y. 1933. Some observations of Japanese cray-
fishes.—Science Reports of the Tokyo Bunrika
Daigaku, Section B 1:155—-158 + pl. 14.

Siebold, Ph. EF von. 1897. Nippon. Archiv zur Bes-
chreibung von Japan und dessen Neben- und
Schutzlandern Jezo mit den stidlichen Kurilen,
Sachalin, Korea und den Liukiu-Inseln, 2nd edi-
tion, vol. 1: 421 pp. + figs. 1-51, front 2 pls.
and 1 map; vol. 2: 342 pp. + figs. 1—47.

Unestam, T. 1969. Resistance to the crayfish plague in
some American, Japanese and European cray-
fishes.—Report, Institute of the Freshwater Re-
search Drottningholm 49:202—209.

Urita, T. 1942. Decapod crustaceans from Saghalien,
Japan.—Bulletin of the Biogeographical Socie-
ty of Japan 12:1—78.

Yamaguchi, H. 1934. Studies on Japanese Brachiob-
dellidae with some revisions on the classifica-
tion.—Journal of the Faculty of Science, Hok-
kaido University, Series VI, Zoology 3:177—
219.

VOLUME 117, NUMBER 1

Appendix I.—Sampling data and geographic variation of morphology in Cambaroides japonicus.

a
OMANI ADUNAFSWNHFKFTOANIADAUMHKWNY

AnmnrnnnbBPR HHH HHH HWWWWWWWWWWNNNNNNNN DN WV
BWNF DOA AIANUAHEWNKTDOAWAIAIADAUNSWNHNHK CTOANIAUANKWNH OS

Sampling site
(No. ref. to Fig. 3)

Rebun
Rishiri
Wakkanai
Nakagawa
Shibetsu
Abashiri
Utoro
Tsubetsu
Maruseppu
Akan
Shikaoi
Memuro
Ikeda
Otofuke
Obihiro
Shintoku
Erimo
Monbetsu
Kamikawa
Biei
Akabira
Furano
Iwamisawa
Takikawa
Ofuyu
Teuri
Yagishiri
Hamamasu
Sapporo
Otaru
Yoichi
Kucchan
Niseko
Kyowa
Rankoshi
Iwanai
Suttsu
Kyogoku
Oshamanbe
Chitose
Eniwa
Soubetsu
Shiraoi
Sahara
Assabu
Kaminokuni
Shikabe
Kikonai
Toi
Todohokke
Fukushima
Matsumae
Imabetsu
Shiura

Date

04 Sep 1968
17 Nov 1992
11 Aug 1998
16 Aug 1990
O01 Sep 1957
04 July 1982
18 Aug 1999
25 Apr 1987
10 Aug 1998
04 Aug 1933
14 May 1987
04 Aug 1986
25 Sep 2000
25 Sep 2000
08 May 1986
09 Aug 1998
19 Sep 2000
20 July 1990
20 Sep 2000
07 Aug 1998
08 Aug 1999
08 Aug 1999
27 May 1959
11 Aug 1992
23 Aug 1992
29 July 2002
29 July 2002
17 June 2000
03 Nov 1990
03 June 1999
28 Aug 2001
07 Oct 2000
08 Oct 2000
10 Oct 1998
21 July 2001
O01 July 2001
13 Sep 2001
28 Sep 2001
19 Sep 2001
02 Aug 1999
04 Noy 2001
02 Aug 1998
12 Aug 1990
O1 Aug 1998
23 June 1997
22 June 1997
23 Aug 1998
22 June 1997
20 Aug 1992
23 June 1997
22 June 1997
22 June 1997
05 Aug 1998
30 Aug 1997

Specimens

33
31
34
31
31
30
33
32
32
Jl
36
30
30
31
31
30
32
34
31
31
31
32
32
31
30
35
35
35
34
35
35
35
35
34
35
36
35
34
33
32
32
31
30
32
34
31
31
32
32
35
33
37
31
32

92
21
21
20
21
22
21
20
al
22
21
21
91
20
20
21
21
20
21
21
20
22
23
21
2 il
25)
25
23
2Q5
26
25
25
25
26
25
24
25)
26
25
Q5
Q?
20
23
91
23)
21
24
27
27
27
27
25
24
20

POCL + SD
AIL3) 22 DS)
22.3 = 4.8
DQ 2 D§3
20.2 = 0.0
16.1 + 2.0
Dpdecd). 2 Pe)
IQ/2 2= i)
ASM 2 Of
21.0 + 4.1
26.8 + 10.8
26.9 + 84
24.5 + 0.0
13.2 = 0.0
1.97 = 0.0
19.2 + 0.0
15.4 + 0.0
18.8 + 0.5
IS}.5) 22 22,33
21.4 +49
D3) = 35
19.6 + 0.0
18.5 + 3.0
18.7 + 1.0
202
20.0 + 0.0
21.6 = 2.0
20.4 + 1.6
20.7 + 3.0
23a = 20
ADY) 2 27
USES =s1kS
Dies) 28 2s
QING 25 Doll
PN 2 Dall
19.@ = 2A
IO.2 2 If
20.5 + 4.2
p,3) 2 Il-9)
I@.7/ = 2D
21.0 = 2.5
/DQ.3} 2 ZS
20.7 + 0.0
18.8 + 1.7
Alls) 22, 20)
AVS 25 25)
Doped 2 M5)
Wp, = 2,1
20.4 = 1.7
19.9 + 1.0
20.7 = 2.5
Ailey 2= 2,
DBA aD,
22.0 + 1.4
20:92 23

Rostrum

1.0 +
IES =
a=

S
I+

0.0
0.7
0.4
0.0

+ 0.0

0.0
0.5
0.0
0.6
0.0
0.0

+ 0.0

0.0
0.0
0.0

+ 0.0

0.0
0.0

+ 0.0

0.0
0.0
0.0

+ 0.0

0.0
0.0
0.4
0.5
0.0
0.4

+ 0.0
+ 0.0

0.0
0.4
0.4
0.5
0.0

+ 0.0
+ 0.5

0.5
0.5

+ 0.0

0.0
0.0
0.6
0.5
0.0
0.5
0.5
0.0
0.3
0.8
0.8

+ 0.8
+14

Telson

Sternal plates

1.0 = 0.0
1.0 + 0.0
1.0 + 0.0
1.0 + 0.0
1.0 + 0.0
2.0 + 0.0
2.0 + 0.0
1.0 + 0.0
1.0 + 0.0
1.0 + 0.0
1.0 + 0.0
1.0 + 0.0

Appendix I.—Continued.

Sampling site
(No. ref. to Fig. 3)

SS) Nakasato

56 Kanagi

57 Goshogawara
58 Kizukuri

59 Iwaki

60 Hirosaki

61 Ikarigaseki
62 Ajigasawa
63 Namioka

64 Aomori

65 Hiranai

66 Tenmabayashi
67 Shichinohe
68 Yokohama
69 Higashidouri
70 Mutsu

71 Ouhata

72 Kawauchi

73 Wakinosawa
74 Tashiro

WD Ohdate

76 Kazuno

Wi Ninohe

—: not measured.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Date

21 Nov 1999
07 Oct 2000

19 Sep 1998

22 Aug 1999
26 July 1998
30 Sep 1997

04 Oct 1931

12 Aug 1998
22 June 2000
03 May 1998
10 Oct 2000

14 July 1994
22 June 1999
23 May 1999
27 Aug 1999
27 Aug 1999
22 June 1999
05 June 1998
17 May 1998
21 June 1991
24 Aug 2002
23 June 1990
22 June 1999

Specimens

31
30
37
38
36
32
32
32
34
37
32
30
32
31
32
31
31
30
33
30
33
37
31

al
92
24
22)
20
92
25
22
ont
23
23
92
91
20
QD
91
25
oD
20
22
27
22
QD

POCL + SD
DNS) = 2,5)
20.3 + 0.4
NZ se 4
DBI] 32 Af
ANS) 2D
W233 a= DD
WYO 25 22
MO.i1 2.3)
ANd} 2B NY)
20.7 + 3.4
IAL se 2a
18.6 + 1.7
IS) 22 ILO
19.4 + 0.0
19.0 + 1.4
23.0 = 0.7
18.3 + 0.8
AMA Se Mei
18.3 + 0.9
73 = 0.3
ANY = ZY
18.9 + 1.1
18.3) 2 12

Rostrum

Ans) 25 (0),7/
3.0 + 0.0
Des) = OLS)
Ap 2 Os)
2.7 + 0.6
3.0 + 0.0
Mell 2 Dei
2.8 = 0.5
2.8 + 0.4
1.9 = 0.6
2.4
3.0
Zoll
3.0 = 0.0
2.5 + 0.6

It Ut 1+ 1 It dt Ut 1 Le IE It it

Telson

1.5
2.0
2.0
De
1.5
2.0
2.6
eS)
2.2
2.8
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
3.0
1.5
3.0
3.0
2.7

0.7
1.4
+ 0.9
xe O.7/
0.8
1.0
0.5
1.0
+ 0.8
0.4
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.6

I+ I+

Ine Ute We |e

It It It It [+ It I+ It It it I+ I+ It I+

Sternal plates

3.0 = 0.0
Dis) 2 (0,7)
2.5 + 0.6
2.0 + 0.0
3.0 + 0.0
3.0 + 0.0
3.0 + 0.0
3.0 + 0.0
2.7 + 0.6
3.0 = 0.0
3.0 + 0.0
3.0 + 0.0
3.0 = 0.0
3.0 = 0.0
2.6 + 0.5
2) 22 (0,7)
3.0 + 0.0
3.0 = 0.0
1.0 + 0.0
3.0 = 0.0

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(1):35—41. 2004.

Two new species of freshwater crabs of the genus Chaceus
Pretzmann, 1965 from the Serrania de Perija of Colombia
(Crustacea: Decapoda: Pseudothelphusidae)

Martha R. Campos and Diego M. Valencia

(MRC) Universidad Nacional de Colombia, Instituto de Ciencias Naturales,
Apartado Aéreo 103698, Bogota, Colombia, S. A, e-mail: mhrocha@ciencias.unal.edu.co;
(DMV) Universidad Nacional de Colombia, Departamento de Biologia, Apartado Aéreo 7495,
Bogota, Colombia, S. A.

Abstract.—Two new species of the genus Chaceus Pretzmann, 1965, C. cu-
rumanensis and C. ibiricensis, are described and illustrated. The description of
these two new species brings to nine the total number of species known in this
genus, distributed in the Sierra de Santa Marta of Colombia, and Serrania de
Perija of Colombia and Venezuela. A key for the identification of the species
based on the morphology of the first male gonopod is presented.

The genus Chaceus Pretzmann, 1965
comprises a group of freshwater crabs dis-
tributed in the Sierra de Santa Marta in Co-
lombia and the Serrania de Perija in Colom-
bia and Venezuela. The systematics, cladis-
tic and biogeography of the genus have
been reviewed by Rodriguez (1982, 1992),
Campos & Rodriguez (1984), Rodriguez &
Campos (1989), Rodriguez & Bosque
(1990), Rodriguez & Viloria (1992) and
Rodriguez & Herrera (1994). With the dis-
covery of two new species, described here-
in, from the western slope of the Serrania
de Perija of Colombia, the genus now con-
tains nine species.

Species of Chaceus are distinguished pri-
marily by characteristics of the efferent
branchial channel, the third maxilliped and
the first male gonopod. The efferent bran-
chial channel is partially closed by the spine
of the jugal angle, and by the produced lat-
eral lobe of the epistome. The exognath of
the third maxilliped is 0.60 to 0.80 times as
long as the ischium. The first male gonopod
usually has the lateral process well devel-
oped, its shape varying according to the
species, and is either subtriangular, elon-
gated, or rounded. The apex is formed by
mesial and caudal processes. A key for the

species of the genus is presented, based ex-
clusively on the morphology of the first
male gonopod. The terminology used for
the different processes of the gonopod is
that established by Smalley (1964), and
Rodriguez (1982).

The shape of the efferent branchial chan-
nel, the length of the exognath of the third
maxilliped, and the structure of the first
male gonopod of the genus Chaceus sug-
gest a close relationship with the genus
Strengeriana Pretzmann, 1971. The first
male gonopods in all species of Chaceus
have the same basic elements as species of
Strengeriana. Rodriguez (1982) has theo-
rized on the possible derivation of the genus
Hypolobocera Ortmann, 1897, from an an-
cestral Chaceus based on the homology of
the finger-like mesial process in the latter,
and the triangular caudal process with the
two papillae found near the spermatic chan-
nel in the former. The morphology of the
first gonopod in C. davidi Campos & Rod-
riguez, 1984, for example, supports this the-
ory since the mesial and caudal processes
are surrounded by a ridge that somewhat
resembles the shape of the apex in species
of Hypolobocera.

The material is deposited in Museo de

36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Historia Natural, Instituto de Ciencias Na-
turales, Universidad Nacional de Colombia,
Bogota (ICN-MHN). The abbreviations cb
and cl, reported as cl X cb, indicate cara-
pace breadth and carapace length, respec-
tively. Color nomenclature follows Smithe
@sr)):

Family Pseudothelphusidae Rathbun, 1893
Tribe Strengerianini Rodriguez, 1982
Genus Chaceus Pretzmann, 1965

Chaceus curumanensis, new species
Fig. 1

Holotype.—Quebrada San Sebastian,
Municipio Curumani, foothill of the Serran-
ia de Perija, Cesar Department, Colombia,
100 m alt., 8 Dec 1978, leg. M. Tiirkay,
male, 14.7 xX 24.5 mm, ICN-MHN-CR
1923).

Paratype.—Same locality data as holo-
type: | male, 13.2 x 23.4 mm, ICN-MHN-
CR 1266.

Type locality.—Quebrada San Sebastian,
Municipio Curumani, foothill of the Serran-
fa de Perijé, Cesar Department, Colombia,
100 m alt.

Diagnosis.—Third maxilliped with ex-
ognath 0.67 times length of ischium. First
gonopod with lateral process elongated,
with distal portion slightly rounded in cau-
dal view, subtriangular in distal view; apex
with needle-shaped mesial process, and tri-
angular caudal process; disto-mesial margin
curving below mesial and caudal processes.

Description of holotype.—Carapace (Fig.
1F) with cervical groove straight, narrow
and shallow distally, wide and deep proxi-
mally, ending some distance from lateral
margin. Anterolateral margin lacking de-
pression behind external orbital angle, but
with slight depression near middle followed
by another near level of cervical groove.
Lateral margin with series of tubercles.
Postfrontal lobes small, oval, delimited an-
teriorly by 2 depressions. Median groove
lacking. Front without distinct upper border,
frontal area regularly sloping downward,
slightly bilobed in dorsal view, lower mar-

gin sinuous in frontal view. Dorsal surface
of carapace smooth, covered by small pa-
pillae, regions not well demarcated. Third
maxilliped with slight depression on distal
half of external margin of merus, exognath
0.67 times length of ischium (Fig. 1H). Or-
ifice of efferent branchial channel partially
closed by spine of jugal angle, and by pro-
duced lateral lobe of epistome (Fig. 1G).
First pereiopods heterochelous; chelae with
palms swollen, and fingers slightly gaping
when closed (Fig. 11). Walking legs (pe-
reiopods 2—5) slender, but not unusually
elongated (total length 1.14 times breath of
carapace).

First male gonopod with lateral process
elongated, with distal portion slightly
rounded in caudal view (Fig. 1A), subtrian-
gular in distal view (Fig. IE); apex with
needle-shaped mesial process, directed ce-
phalically, and triangular caudal process,
directed transversely to mesial process in
caudal and cephalic views, both processes
surrounded by lateral process in distal view;
disto-mesial margin curving below mesial
and caudal processes (Fig. 1C—E); lateral
side of gonopod expanded with irregular
rows of short setae, caudal surface with
long setae proximally (Fig. 1A, B, D).

Color.—The holotype, preserved in al-
cohol, is light brown (near 37, Antique
Brown) with dark specks on the dorsal side
of the carapace. The dorsal and ventral sur-
faces of the chelae and walking legs are
brown (near 139, True Cinnamon). The
ventral surface of the carapace is brown
(near 239, Ground Cinnamon).

Etymology.—The specific name refers to
the type locality, the Municipio Curumani.

Remarks.—Comparison of this new spe-
cies with descriptions and specimens of
other species of the genus revealed that this
new species is most similar to Chaceus
pearsei (Rathbun, 1915). The two can be
distinguished by differences in the gono-
pods. The male first gonopod of C. pearsei
has been described and illustrated by Rod-
riguez (1982:37, fig. 12). The lateral pro-
cess in this new species is elongated, with

VOLUME 117, NUMBER 1 37

UCFinwen

Fig. 1. Chaceus curumanensis, new species, male holotype, ICN-MHN-CR 1993: A, left first gonopod,
caudal view; B, same, lateral view; C, same, cephalic view; D, same, mesial view; E, same, apex, distal view;
E right side of carapace with eye, dorsal view; G, left orifice of efferent branchial channel; H, left third max-
illiped, external view; I, left cheliped, external view. 1, lateral process; 2, mesial process; 3, caudal process.

38 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

the distal portion slightly rounded in caudal
view, whereas it is subtriangular in C. pear-
sei. The mesial process in C. pearsei is fin-
ger-like, blunt, whereas it is needle-shaped
in C. curumanensis. The caudal process in
this new species is slightly parallel to the
mesial process in distal view, whereas it is
recurved at its base in C. pearsei.

Chaceus ibiricensis, new species
Fig. 2

Holotype.—Los Laureles Farm, Vereda
Alto del Tucuy, Corregimiento La Victoria
de San Isidro, Municipio La Jagua de Ibir-
ico, Serrania de Perija, Cesar Department,
Colombia, 1100 m alt., 9°34'35.8’N,
73°6'26.0"W, 7 Mar 1996, leg. M. R. Cam-
pos, male, 13.0 X 21.3 mm, ICN-MHN-CR
IQQy.

Paratypes.—Same locality data as holo-
type: 19 males, size range 8.6 X 13.7 mm
to 13.7 X 22.7 mm, 16 females, size range
8.2 X 12.8 mm to 12.4 X 20.1 mm, ICN-
MHN-CR 1549.

Additional non-paratypic material.—Be-
tween Veredas Alto de las Flores, and Nue-
vo Mundo, Corregimiento La Victoria de
San Isidro, Municipio La Jagua de Ibirico,
Serrania de Perija, Cesar Department, Co-
lombia, 1350-1400 m alt., 7, 8 Mar 1996,
leg. M. R. Campos, 23 males, size range
7.3 X 11.4 mm to 13.0 X 21.4 mm, 15
females, size range 8.4 X 13.1 mm, to 14.4
x 24.8 mm, ICN-MHN-CR 1550, 1552.—
Tucuy River, Vereda Alto de las Flores,
Corregimiento La Victoria de San Isidro,
Municipio La Jagua de Ibirico, Serrania de
Perija, Cesar Department, Colombia, 870 m
alt., 11 Mar 1996, leg. M. R. Campos, 6
males, size range 9.9 X 15.9 mm to 10.9 x
17.6 mm, 3 females, size range 10.8 * 17.2
mm to 11.9 X 20.4 mm, 2 juveniles, ICN-
MHN-CR 1559.—La Sorpresa Farm, Ver-
eda Alto de las Flores, Corregimiento La
Victoria de San Isidro, Municipio La Jagua
de Ibirico, Serrania de Perija, Cesar De-
partment, Colombia, 1280 m alt., 12 Mar

1996, leg. J. V. Rueda, 1 male, 12.4 X 21.1
mm, ICN-MHN-CR 1560.

Type locality.—Los Laureles Farm, Ver-
eda Alto del Tucuy, Corregimiento La Vic-
toria de San Isidro, Municipio La Jagua de
Ibirico, Serrania de Perija, Cesar Depart-
ment, Colombia, 1100 m alt., 9°34’35.8’N,
73°6'26.0"W.

Diagnosis.—Third maxilliped with ex-
ognath 0.72 times length of ischium. First
male gonopod with lateral process hood-
like; mesial process prominent, subcylindri-
cal, semicircular caudally with median con-
striction and subdistal subtriangular papilla
cephalically; caudal process subtriangular;
disto-mesial margin forming semicircular
projection in cephalo-lateral direction.

Description of holotype.—Carapace (Fig.
2F) with cervical groove straight, narrow,
shallow, ending some distance from lateral
margin. Anterolateral margin with shallow
depression behind external orbital angle fol-
lowed by approximately 5 papillae. Lateral
margin with series of approximately 10 tu-
bercles. Postfrontal lobes small, oval, de-
limited anteriorly by 2 depressions. Median
groove shallow, and narrow. Front lacking
distinct upper border, frontal area regularly
sloping downward, bilobed in dorsal view,
lower margin sinuous in frontal view. Dor-
sal surface of carapace smooth, covered by
small papillae, regions not well demarcated.
Third maxilliped with external margin of
merus straight, exognath 0.72 times length
of ischium (Fig. 2H). Orifice of efferent
branchial channel partially closed by spine
of jugal angle, and by produced lateral lobe
of epistome (Fig. 2G). First pereiopods het-
erochelous; palm of larger chela strongly
swollen, fingers gaping when closed (Fig.
21); palm of smaller chela moderately swol-
len, fingers not gaping when closed. Walk-
ing legs (pereiopods 2—5) slender and elon-
gated (total length 1.25 times the breadth of
carapace).

First male gonopod with lateral process
hood-like, lateral and cephalic outer surface
covered with irregular papillae and spinules
(Fig. 2A—C); apex with mesial and caudal

VOLUME 117, NUMBER 1 39

OC Pinzen

m
mn

A-D | G

Fig. 2. Chaceus ibiricensis, new species, male holotype, ICN-MHN-CR 1992: A, left first gonopod, caudal
view; B, same, lateral view; C, same, cephalic view; D, same, mesial view; E, same, apex, distal view; EF right
side of carapace with eye, dorsal view; G, left orifice of efferent branchial channel; H, left third maxilliped,
external view; I, right cheliped, external view. 1, lateral process; 2, mesial process; 3, caudal process.

3
3

40 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

processes; mesial process prominent, sub-
cylindrical, semicircular caudally (Fig. 2A,
B); with median constriction, and subdistal
subtriangular papilla cephalically (Fig. 2C—
E); caudal process subtriangular, both pro-
cesses partially surrounded by lateral pro-
cess in distal view; disto-mesial margin
forming semicircular projection into ce-
phalo-lateral direction (Fig. 2E); lateral ex-
panded side of gonopod with rows of long,
plumose setae, mesial side with row of spi-
nules; caudal surface with conspicuous long
setae proximally (Fig. 2A—D).

Color.—The holotype, preserved in al-
cohol, is brown (near 240, Kingfisher Ru-
fous) on the dorsal side of the carapace. The
dorsal and ventral surfaces of chelae and
walking legs are brown (near 223B, Verona
Brown). The ventral surface of the carapace
is light brown (near 223C, Sayal Brown).

Habitat.—The vegetation of the collec-
tion areas is primary forest. The specimens
were collected in shaded, moist banks of
springs and streams, in soft mud under
rocks.

Etymology.—tThe specific name refers to
the type locality, the Municipio La Jagua
de Ibirico.

Remarks.—Comparison of this new spe-
cies with descriptions and specimens of
other species of the genus revealed that it
is most similar to Chaceus turikensis Rod-
riguez & Herrera, 1994. The two can be
distinguished by differences in the size of
the eyes, and in the gonopod. The male first
gonopod of C. turikensis has been described
and illustrated by Rodriguez & Herrera
(1994:123, fig. 2). In C. turikensis the eyes
do not fill the orbital cavity, whereas in this
new species they do fill the orbital cavity.
In C. ibiricensis the lateral process of the
gonopod is hood-like with the distal portion
directed distally in caudal view (Fig. 2A—
E), whereas the lateral lobe is foliose and
the distal portion is directed transversely to
the main axis of the appendage in C. turi-
kensis. The mesial process is ellipsoidal in
C. turikensis, whereas it is subcylindrical
with a median constriction and subdistal

subtriangular papilla cephalically in C. tbir-
icensis.

Key to Species of Chaceus

1. Lateral process of gonopod well devel-
oped
— Lateral process of gonopod reduced
C. nasutus Rodriguez, 1980
2. Lateral process of gonopod subtriangular
or elongated
— Lateral process of gonopod rounded... 8
3. Lateral process of gonopod with semi-
circular notch on lateral surface
.. C. cesarensis Rodriguez & Viloria, 1992
— Lateral process of gonopod without
semicircular notch on lateral surface .. 4
4. Mesial process of gonopod about same
length as length of caudal process
. C. dayidi Campos & Rodriguez, 1984
— Mesial process of gonopod longer than
caudal process
5. Mesial process of gonopod with median
constriction and subapical subtriangular
papilla cephalically
Peeters: Sos C. ibiricensis, new species
— Mesial process of gonopod without me-
dian constriction and subapical papilla
cephalically
6. Mesial process of gonopod ellipsoidal ...
.. C. turikensis Rodriguez & Herrera, 1994
— Mesial process of gonopod not ellipsoi-
dal aco, To MR eee vi
7. Mesial process of gonopod finger-like,
blunteaee ee ae C. pearsei (Rathbun, 1915)
— Mesial process of gonopod needle-
shaped C. curumanensis, new species
8. Mesial process of gonopod with round-
ed, elongated papilla basally
. C. caecus Rodriguez & Bosque, 1990
— Mesial process of gonopod lacking
rounded, elongated papilla basally
ia a eae C. motiloni Rodriguez, 1980

Acknowledgments

I am indebted to R. Lemaitre, of the Na-
tional Museum of Natural History, Smith-
sonian Insitution, for his corrections and
suggestions to improve the manuscript. I
thank E G. Stiles, and the anonymous ref-
erees for providing useful comments. The
illustrations were prepared by Juan C. Pin-

VOLUME 117, NUMBER 1

zon. The specimens of Chaceus curuma-
nensis were donated by M. Tiirkay, of the
Senckenberg Museum of Frankfurt to the
ICN-MHN collection.

Literature Cited

Campos, M. R., & G. Rodriguez, 1984. New species
of freshwater crabs (Crustacea: Decapoda:
Pseudothelphusidae) from Colombia.—Pro-
ceedings of the Biological Society of Washing-
ton 97:538—543.

Ortmann, A. 1897. Carcinologische Studien.—Zoolo-
gische Jahrbiicher, Abtheilung fiir Systematik,
Geographie und Biologie der Tiere 10:258-372.

Pretzmann, G. 1965. Vorlaufiger Bericht tiber die Fam-
ilie Pseudothelphusidae.—Anzeiger der Oster-
reichischen Akademie der Wissenschaften
Mathematische Naturwissenschaftliche Klasse
(1)1:1-10.

. 1971. Fortschritte in der Klassifizierung der
Pseudothelphusidae.—Anzeiger der Mathema-
tisch Naturwissenschaftliche der Osterreichisch-
en Akademie der Wissenschaften (1)179(1—4):
14-24.

Rathbun, M. J. 1893. Descriptions of new species of
American freshwater crabs.—Proceedings of
the United States National Museum 16(959):
649-661.

. 1915. New fresh-water crabs (Pseudothelphu-
sa) from Colombia.—Proceedings of the Bio-
logical Society of Washington 28:95—100.

Rodriguez, G. 1980. Description préliminaire de
quelques espéces et genres nouveaux de Crabes
d'eau douce de 1’Amérique tropicale (Crusta-
cea: Decapoda: Pseudothelphusidae).—Bulletin

du Muséum nationale d’ Histoire naturelle, Paris

(4) 2 Section A (3):889—-894.

. 1982. Les crabes d’eau douce d’ Amérique.

Famille des Pseudothelphusidae.—Faune Trop-

icale 22:1—223.

. 1992. ‘The freshwater crabs of America. Fam-

ily Trichodactylidae and supplement to the

Family Pseudothelphusidae.—Faune Tropicale

31:1-189.

, & C. Bosque. 1990. A stygobiont crab, Cha-

ceus caecus n. sp. and its related stygophile spe-

cies Chaceus motiloni Rodriguez, 1980, (Crus-

tacea, Decapoda, Pseudothelphusidae) from a

cave in the Cordillera de Perija, Venezuela.—

Mémoires de Bioespéologie XVII:127—134.

, & M. R. Campos. 1989. Cladistic relation-

ships of freshwater crabs of the tribe Strenger-

ianini (Decapoda: Pseudothelphusidae) from the
northern Andes, with comments on their bio-
geography and descriptions of new species.—

Journal of Crustacean Biology 9:141—156.

, & FE Herrera. 1994. A new troglophilic crab,

Chaceus turikensis, from Venezuela, and addi-

tional notes of the stygobiont crab Chaceus cae-

cus Rodriguez & Bosque 1990, (Decapoda:

Brachyura: Pseudothelphusidae)—Mémoires

de Bioespéologie XXI:121—126.

, & A. L. Viloria. 1992. Chaceus cesarensis, a
new species of the fresh-water crab (Crustacea:
Decapoda: Pseudothelphusidae) from Colombia
with a key to the genus.—Proceedings of the
Biological Society of Washington 105:77—80.

Smalley, A. 1964. A terminology for the gonopods of
the American river crabs.—Systematic Zoology
13:28-31.

Smithe, FE B. 1975. Naturalist’s color guide. The Amer-
ican Museum of Natural History, New York.
Part 1: unnumbered pages.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(1):42—56. 2004.

Reevaluation of the hermit crab genus Parapagurodes McLaughlin &
Haig, 1973 (Decapoda: Anomura: Paguroidea: Paguridae) and a

new genus for Parapagurodes doederleini (Doflein, 1902)

Patsy A. McLaughlin and Akira Asakura*

Shannon Point Marine Center, Western Washington University, 1900 Shannon Point Road,
Anacortes, WA 98221-9081B, U.S.A.;
*Natural History Museum and Institute, Chiba 955-2, Aoba-cho, Chuo-ku,
Chiba, 260-8682, Japan

Abstract.—The question of polyphyly in the hermit crab genus, Parapagu-
rodes McLaughlin & Haig, 1973, has been investigated by comparisons of a
series of morphological characters among the eight species presently assigned
to the genus. The results of the analysis have shown that the only mutually
shared characters are an acutely developed rostrum and the presence, in males,
of a short or very short right sexual tube. Consequently, the composition of
Parapagurodes is herein restricted to the two species originally assigned, viz.
P. makarovi McLaughlin & Haig, 1973, and P. laurentae McLaughlin & Haig,
1973. Parapagurodes hartae Mclaughlin & Jensen, 1996, is transferred to the
genus Pagurus, and four species subsequently transferred from Pagurus to
Parapagurodes, viz. P. gracilipes (Stimpson, 1858), P. nipponensis (Yokoya,
1933), P. constans (Stimpson, 1858), and P. imaii (Yokoya, 1939) are returned
to Pagurus. A new genus, Dofleinia, is proposed for the species, Parapagu-

rodes doederleini (Doflein).

When first proposed, the genus Parapa-
gurodes McLaughlin & Haig, 1973, was
characterized, in part, as having 11 pairs of
biserial gills; a moderately well developed,
but not recurved external lobe of the max-
illulary endopod; fifth pereopods with cox-
ae symmetrical; males with a short right
sexual tube, and biramous left pleopods ab-
sent or weakly developed on pleomeres (cf.
Schram & Koenmann 2003) 3—5; females
lacking paired first pleopods, with biramous
left pleopods 2—4 weakly to moderately
well developed, left fifth pleopod weakly
developed or absent; and a telson with ter-
minal margins straight, slightly concave or
slightly oblique. Additionally, the authors
noted that while a right sexual tube was al-
ways present in mature males, its length
and orientation were variable, and in one
specimen both right and left tubes were pre-
sent. Variations also were observed in the

number and development of male and fe-
male pleopods in both P. makarovi Mc-
Laughlin & Haig, 1973, the type species of
the genus, and the second described spe-
cies, P. laurentae McLaughlin & Haig,
1973. In recent years, one new species, P.
hartae McLaughlin & Jensen, 1996, has
been described in the genus, and five Jap-
anese species have been transferred to it
viz.: Pagurus gracilipes (Stimpson, 1858),
P. nipponensis (Yokoya, 1933), P. constans
(Stimpson, 1858), and P. imaii (Yokoya,
1939) by Komai (1998, 1999) and Cata-
pagurus doederleini Doflein, 1902 by
Asakura (2001).

At the time of the establishment of Par-
apagurodes McLaughlin & Haig, 1973,
male sexual tube development had been re-
ported in less than two dozen genera.
McLaughlin & Haig (1973) could relate
Parapagurodes to only two of those genera,

VOLUME 117, NUMBER 1

Pagurodes Henderson, 1888 and Acantho-
pagurus de Saint Laurent, 1969, but cited
several characters by which the three genera
could be separated. In the subsequent 30
years the number of genera with docu-
mented sexual tube development has more
than doubled (cf. McLaughlin 2003). None-
theless, Parapagurodes still can be allied
only to Pagurodes and Acanthopagurus and
more remotely to Catapagurus A. Milne
Edwards, 1880. However, recently the
monophyly of Parapagurodes itself has
come under question (Lemaitre & Mc-
Laughlin 2003b).

In their introductory remarks regarding
Parapagurodes hartae, McLaughlin & Jen-
sen (1996: 841) made the unfortunate state-
ment that “‘... males have a small sexual
tube on the coxa of the right fifth pereopod.
This species, therefore, cannot be attributed
to Pagurus, but must be assigned to Para-
pagurodes ...” The comment was prompt-
ed by the fact that prior to their description
of P. hartae, this taxon had been reported
from California and Washington, USA, and
British Columbia, Canada, as Pagurus sp.
(McLaughlin & Haig 1973, Hart 1982, Jen-
sen 1995). Regrettably, McLaughlin & Jen-
sen’s (1996) remark has been interpreted by
some carcinologists to mean that species
with papillae and/or very short sexual tubes
are automatically excluded from the genus
Pagurus (e.g., Komai 1998, 1999). Accord-
ing to the views of McLaughlin & Lemaitre
(2001) and Lemaitre & McLaughlin
(2003a, 2003b), the presence or absence of
very short male sexual tubes and/or papillae
should not be seen as the single cause to
transfer species from genera to which they
are otherwise morphologically attributable,
or assign species to genera that they are not
otherwise morphologically allied.

McLaughlin & Jensen (1996) justified
their generic assignment on the basis of
morphological and larval similarities
among the three species then assigned to
Parapagurodes. However, they also pointed
out, as McLaughlin & Haig (1973) had for
P. laurentae, that P. hartae had superficial

resemblances to a few northeastern Pacific
species of Pagurus.

Upon the observation of very short sex-
ual tubes in Pagurus gracilipes and P. nip-
ponensis, Komai (1998) provisionally
transferred these two species to Parapagu-
rodes, while noting their close similarities
to species of McLaughlin’s (1974) bernhar-
dus group of Pagurus. Komai (1998) also
pointed out that while Pagurus gracilipes
and P. nipponensis shared the presence of
a small right male sexual tube, both species
differed substantially from Parapagurodes
makarovi, P. laurentae, and P. hartae.

In the subsequent, continuing study of
Japanese species of Pagurus, Komai (1999)
transferred Pagurus constans and P. imaii
to Parapagurodes because he found very
short male sexual tubes on both fifth coxae
in the former species, and a single right
tube in the latter. He also provided a minor
emendation to the generic diagnosis by call-
ing attention to the slight median indenta-
tion, cleft or concavity sometimes seen in
the gill lamellae, and to the occurrence of
the left sexual tube, although he acknowl-
edged that McLaughlin & Haig (1973) sim-
ilarly had reported the rare occurrence of a
left tube in P. makarovi. Unfortunately, Ko-
mai’s (1999) emendation, like McLaughlin
& Jensen’s (1996) brief generic diagnosis,
failed to acknowledge the absence or re-
duction in number of male pleopods and the
usual absence of the fifth left female pleo-
pod in the type species.

In his review of the genus Catapagurus
A. Milne-Edwards, 1880, Asakura (2001)
redescribed Catapagurus doederleini and
found it to be markedly divergent from all
other species that had been assigned to Ca-
tapagurus. Asakura transferred Doflein’s
(1902) taxon to Parapagurodes stating that
it agreed with all the diagnostic characters
proposed by McLaughlin & Haig (1973) for
their genus; however, it was primarily the
presence of a very short right sexual tube
that prompted his action. He quite correctly
acknowledged the dimorphic second pereo-
pods of P. doederleini, as well as the lack

44 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

of corneous spines on the ventral margins
of the second right and both third pereo-
pods.

As previously indicated, Lemaitre &
McLaughlin (2003b) expressed the opinion
that Parapagurodes, as presently constitut-
ed, represented a polyphyletic taxon. To
evaluate the merits of their conclusion, we
have critically reviewed the descriptions of
each of the assigned taxa. We have supple-
mented these reviews by reexamining spec-
imens of Parapagurodes laurentae and P.
hartae in the first author’s personal collec-
tions (PMcL). Additionally we have ex-
amined representatives of P. constans, P.
doederleini, P. gracilipes, P. imaii, and P.
nipponensis from the collections of the Nat-
ural History Museum and Institute, Chiba
(CBM-ZC), the Hilgendorf collection from
the Museum fiir Naturkunde, Berlin, Ger-
many (ZMB), and specimens donated to
one of the authors by Dr. M. Imafuku, Kyo-
to University. From our reviews and ex-
aminations, we present the comparative di-
agnoses of the eight species we have used
to determine the validity of the current ge-
neric assignments.

Animal size is indicated by shield length
(sl) as measured from the tip of the rostrum
to the midpoint of the posterior margin of
the shield. Reported sexual tube length cor-
responds to the criterion of McLaughlin
(2003): very short (<1 coxal length), short
(1-2 coxal lengths), moderate (>2-—5 coxal
lengths). The reference by McLaughlin &
Haig (1973) to the fourth pereopod being
subchelate or not subchelate is interpreted
here according to McLaughlin (1997) who
recognized three conditions in the propodal-
dactyl articulation of this appendage.
McLaughlin & Haig’s (1973) “‘subchelate”’
is viewed by McLaughlin (1997) as being
semichelate, whereas McLaughlin & Haig’s
(1973) “‘not subchelate” is now considered
to actually be subchelate. The abbreviation
Ovig. indicates ovigerous female. Previous-
ly published illustrations used in this man-
uscript are of specimens in the collections
of the Los Angeles Country Natural His-

tory Museum (LACM) [transferred to that
Museum from the Allan Hancock Founda-
tion (AHP)], Los Angeles, California; the
Royal British Columbia Provincial Museum
(RBCPM), Victoria, British Columbia; and
Zoologische Staatssammlung Miinchen
(ZSSM), Munich.

Review and Reexamination

Parapagurodes makarovi McLaughlin &
Haig, 1973
Figs. 1A, 2A, 3A, 4A, B, 5A

Description by McLaughlin & Haig
(1973:119—120, figs. 4a, 5—8). No supple-
mental material available.

Diagnosis.—Gill lamellae essentially
biserial but with or without very weak dis-
tal indentation or concavity. Rostrum acute-
ly triangular. Maxillule with somewhat pro-
duced endopodal external lobe, not re-
curved. Right cheliped elongate, more so in
large individuals; dorsal surface of palm
distinctly convex; dorsal surface of carpus
with row of spines mesiad of midline. Left
cheliped elongate dorsal surface of palm
convex, elevated in midline proximally.
Ambulatory legs similar, somewhat later-
ally compressed; dactyls as long or longer
than propodi, slender, in dorsal view
straight, each with row of corneous spines
on ventral margin; carpi each with dorso-
distal spine. Fourth pereopods usually sub-
chelate, occasionally weakly semichelate;
preungual process very small; propodal
rasp with 1—3 irregular rows of corneous
scales. Sternite of third pereopods (sixth
thoracomere) semi- or subsemicircular.
Sternite of fifth pereopods (eighth thora-
comere) separated into two broad lobes by
weak median depression; coxae of fifth pe-
reopods symmetrical. Males with short sex-
ual tube developed from coxa of right fifth
pereopod; left gonopore sometimes with pa-
pilla, occasionally with short tube. No
paired pleopods in either sex. Males usually
without, occasionally with weakly biramous
left pleopods on pleomeres 3 and 4. Fe-
males with weakly developed, biramous left

VOLUME 117, NUMBER 1

Fig. 1.

Coxae and sternite of fifth pereopods. A, B, Parapagurodes makarovi McLaughlin & Haig, 1973,
3 (sl = 3.5 mm), 6 (sl = 3.6 mm), LACM; C, D, P. laurentae McLaughlin & Haig, 1973, ¢ (sl = 3.4 mm),
3 (sl = 3.4 mm), LACM; E, P. hartae McLaughlin & Jensen, 1996, ¢ (sl = 3.2 mm), RBCPM 974-00368-
22; E P. gracilipes (Stimpson, 1858), ¢ (sl = 9.0 mm), CBM-ZC 2977; G, P. nipponensis (Yokoya, 1933), 3
(sl = 7.3 mm), CBM-ZC 1162; H, P. constans (Stimpson, 1858), d (sl = 8.7 mm), CBM-ZC 59; I, P. imaii
(Yokoya, 1939), 5 (sl = 2.5 mm), CBM-ZC 2699; J, P. doederleini (Doflein, 1902), 3 (sl = 8.6 mm), ZSSM

274/1. A—D redrawn from McLaughlin & Haig (1973); E redrawn from McLaughlin & Jensen (1996); J from
Asakura (2001).

46 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2.

Endopod of maxillule. A, Parapagurodes makarovi McLaughlin & Haig, 1973, 6 (sl = 3.5 mm),

LACM; B, P. laurentae McLaughlin & Haig, 1973, 3 (sl = 3.4 mm), LACM; C, P. hartae McLaughlin &
Jensen, 1996, ¢ (sl = 3.2 mm), RBCPM 974-00368-22; D, P. gracilipes (Stimpson, 1858), ¢ (sl = 9.0 mm),
CBM-ZC 2977; E, P. nipponensis (Yokoya, 1933), ¢ (sl = 7.3 mm), CBM-ZC 1162; E P. constans (Stimpson,
1858); 3d (sl = 8.7 mm), CBM-ZC 59; G, P. imaii (Yokoya, 1939), ¢ (sl = 2.5 mm), CBM-ZC 2699; H, P.
doederleini (Doflein, 1902), d (sl = 8.6 mm), ZSSM 274/1. A, B redrawn from McLaughlin & Haig (1973);
C redrawn from McLaughlin & Jensen (1996); H from Asakura (2001).

pleopods on pleomeres 2—4, pleopod 5 ab-
sent or rudimentary. Telson with posterior
lobes separated by very small, shallow me-
dian cleft; terminal margins straight or
somewhat concave, each with several to nu-
merous spinules or small to very small
spines.

Parapagurodes laurentae McLaughlin &
Haig, 1973
Figs. 1B, 2B, 3B, 4C, D, 5B

Description by McLaughlin & Haig
(1973:129—134, figs. 4b, 9-11).

Supplemental material examined.—
U.S.A.: 3 6 (sl = 1.4-2.9 mm), 3 @ (sl =
1.7—2.9 mm), 2.5 mi SE Seal Rocks, Santa
Catalina I, CA, 159-174 m, 25 Oct 1941,
PMcL.

Diagnosis.—Gill lamellae essentially
biserial but with or without very weak dis-
tal indentation or concavity. Rostrum acute-
ly triangular. Maxillule with moderately
well developed endopodal external lobe,
not recurved. Right cheliped usually elon-
gate, more so in large individuals; dorsal
surface of palm convex; dorsal surface of

VOLUME 117, NUMBER 1

carpus with row of spines mesiad of mid-
line. Left cheliped moderately long, dorsal
surface of palm convex. Ambulatory legs
similar, somewhat laterally compressed,
dactyls as long or longer than propodi, slen-
der, in dorsal view usually straight, with
row of corneous spines on ventral margin;
carpi each with dorsodistal spine. Fourth
pereopods usually semichelate, occasional-
ly subchelate; preungual process very
small; propodal rasp with 1-3 irregular
rows of corneous scales. Sternite of third
pereopods (sixth thoracomere) subsemicir-
cular. Sternite of fifth pereopods (eighth
thoracomere) separated into two broad
lobes by weak to moderate median depres-
sion; coxae of fifth pereopods symmetrical.
Males with short or very short sexual tube
developed from coxa of right fifth pereo-
pod; left gonopore sometimes with papilla.
No paired pleopods in either sex. Males
usually with weakly biramous left pleopods
on pleomeres 3 and 4, occasionally without
unpaired pleopods. Females usually with
moderately well developed, biramous ple-
opods on pleomere 2—4; pleopod 5 rudi-
mentary, rarely absent. Telson with poste-
rior lobes separated by very shallow me-
dian cleft; terminal margins concave or
slightly oblique, each with row of very
small spinules and 1—4 small spines at pos-
terolateral angles.

Parapagurodes hartae McLaughlin &
Jensen, 1996
PigsmlC2C3C4E 5C

Description by McLaughlin & Jensen
(1996:844—847, figs. 1-4).

Supplemental material examined.—Can-
ada: 1 ¢ (sl = 1.1 mm), 1 2 (sl = 1.5 mm),
Taylor Inlet, Barkley Sound, British Colum-
bia, 10 m, 10 Jun 1994, PMcL.

large males; dorsal surface of palm convex;
dorsal surface of carpus with row of spines
mesiad of midline. Left cheliped with dac-
tyl and fixed finger short and broad in small
males and females, longer in large males;
dorsal surface of palm convex. Ambulatory
legs similar; dactyls slightly shorter to
slightly longer than propodi, moderately
slender, laterally compressed, in dorsal view
straight, with row of corneous spines on
ventral margin; carpi each with dorsodistal
spine and row of low, sometimes spinulose
protuberances on dorsal surface, rarely 1
dorsoproximal spines (second pereopods).
Fourth pereopods usually semichelate, oc-
casionally subchelate; preungual process
very small; propodal rasp with 2—4 irregu-
lar rows of corneous scales. Sternite of third
pereopods (sixth thoracomere) subsemicir-
cular to subrectangular. Sternite of fifth pe-
reopods (eighth thoracomere) separated into
two broad lobes by weak to moderate me-
dian depression; coxae of fifth pereopods
symmetrical. Males often with very short
sexual tube developed from coxa of right
fifth pereopod; left gonopore without papil-
la. No paired pleopods in either sex. Males
with unequally biramous left pleopods on
pleomeres 3—5. Females with moderately
well developed, left biramous pleopods on
pleomeres 2—4; pleopod 5 as in male. Tel-
son with posterior lobes separated by shal-
low, U-shaped median cleft; terminal mar-
gins rounded or slightly oblique, each with
row of very small spinules and lor 2 small
spines at posterolateral angles.

Parapagurodes gracilipes (Stimpson,
1858)
Figs. 1D, 2D, 3D, 4E 5D

Redescription by Komai (1998:268—275,
figs. LA, 2—5, 7).

Supplemental material examined.—Ja-
pan: 2 ¢ (sl = 5.4, 9.0 mm), 1 2 (sl = 6.7
mm), off Choshi, Chiba, 10—20 m, 3 Sep
1996, CBM-ZC 2977.

Diagnosis.—Gill lamellae biserial. Ros-
trum acutely triangular. Maxillule with

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

VOLUME 117, NUMBER 1

somewhat produced endopodal external
lobe, slightly to distinctly recurved. Right
cheliped moderately (small specimens) to
considerably elongate in large individuals;
dorsal surface of palm weakly convex but
with dorsomesial portion somewhat elevat-
ed; dorsal surface of carpus with row of
spines mesiad of midline. Left cheliped
with dorsal surface of palm somewhat flat-
tened, dorsomesial and dorsolateral margins
slightly elevated. Ambulatory legs similar;
dactyls longer than propodi, strongly twist-
ed; moderately broad, each with row of nu-
Merous corneous spines on ventral margin;
carpi each with single or double row of
multifid spines. Fourth pereopods semiche-
late; no preungual process; propodal rasp
with several rows of corneous scales. Ster-
nite of third pereopods (sixth thoracomere)
subquadrate, weakly skewed, sulcate me-
dially. Sternite of fifth pereopods (eighth
thoracomere) separated into two subovate
lobes by shallow median groove; coxae of
fifth pereopods symmetrical. Males with
very short sexual tube developed from coxa
of right fifth pereopod; left gonopore with-
out tube or papilla. No paired pleopods in
either sex. Males with unequally biramous
pleopods on pleomeres 3—5. Females with
well developed, biramous pleopods on
pleomeres 2—4; pleopod 5 with endopod
noticeably reduced. Telson with posterior
lobes separated by very small, or indistinct
median cleft; terminal margins nearly hor-
izontal, each with eight to ten small spines
and two or three larger spines at postero-
lateral angles, lateral margins occasionally
with spinules.

Parapagurodes nipponensis (Yokoya,
1933)
Figs. 1E, 2E, 3E, 4G, 5E

Redescribed by Komai (1998:275—279,
figs 1B, 6, 7) only as similar to P. gracili-
pes with certain noted differences.

Supplemental material examined.—Ja-
pan: 4 6 (sl = 6.3-7.6 mm), 1 2 (sl = 5.3
mm), Kumano Nada, 50 m, Sep 1981,
PMcL; 2 ¢ (sl = 8.0, 9.2 mm), off Kashi-
ma, Irakaki, 65 m, 24 Apr 1991, CBM-ZC
50; 1 ¢ (sl = 7.3 mm), off Kii Minabe, Kii
Peninsula, 80—100 M, 24 Mar 1995, CBM-
ZC 1162.

Diagnosis.—Gill lamellae biserial. Ros-
trum acutely triangular. Maxillule with
somewhat produced external lobe, slightly
to distinctly recurved. Right cheliped mod-
erately (small specimens) to considerably
elongate in large individuals; dorsal surface
of palm weakly convex but with dorsome-
sial marginal area somewhat elevated; dor-
sal surface of carpus with row of spines me-
siad of midline. Left cheliped with dorsal
surface of palm somewhat flattened, dor-
somesial and dorsolateral margins slightly
elevated. Ambulatory legs similar; dactyls
longer than propodi, strongly twisted; mod-
erately slender to moderately broad; each
with prominent longitudinal sulcus on lat-
eral face and row of numerous very tiny
corneous spines on ventral margin; carpi
each with single or double row of multifid
spines. Fourth pereopods semichelate; no
preungual process; propodal rasp with sev-
eral rows of corneous scales. Sternite of
third pereopods (sixth thoracomere) sub-

Fig. 3.

Ambulatory dactyls. A—H, dactyl of left third pereopod (A-C lateral view, D-H, mesial view); I,

dactyl of left second pereopod (mesial view); J, dactyl of right second pereopod (mesial view). A, Parapagurodes
makarovi McLaughlin & Haig, 1973, 6 (sl = 3.5 mm), LACM; B, P. laurentae McLaughlin & Haig, 1973, 3
(sl = 3.2 mm), LACM; C, P. hartae McLaughlin & Jensen, 1996, 3 (sl = 2.8 mm), RBCPM 974-00368-22;
D, P. gracilipes (Stimpson, 1858), d (sl = 9.0 mm), CBM-ZC 2977; E, P. nipponensis (Yokoya, 1933), 3 (sl
= 7.3 mm), CBM-ZC 1162; E P. constans (Stimpson, 1858), ¢ (sl = 8.7 mm), CBM-ZC 59; G, H, P. imaii
(Yokoya, 1939), 5 (sl = 2.5 mm), CBM-ZC 2699, ovig. 2 (sl = 1.6 mm), CBM-ZC 1911; I, J, P. doederleini
(Dofiein, 1902), 5 (sl = 8.6 mm), ZSSM 274/1. A, B redrawn from McLaughlin & Haig (1973); C redrawn
from McLaughlin & Jensen (1996); I, J from Asakura (2001).

50 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4.

Dactyl and propodus of left fourth pereopod (lateral view). A, Parapagurodes makarovi McLaughlin

& Haig, 1973, 3 (sl = 3.5 mm), LACM; B, P. laurentae McLaughlin & Haig, 1973, ¢ (sl = 3.4 mm), LACM;
C, P. hartae McLaughlin & Jensen, 1996, d (sl = 3.2 mm), RBCPM 974-00368-22; D, P. gracilipes (Stimpson,
1858), 5 (sl = 9.0 mm), CBM-ZC 2977; E, P. nipponensis (Yokoya, 1933), 3d (sl = 7.3 mm), CBM-ZC 1162;
EP. constans (Stimpson, 1858), ¢ (sl = 8.7 mm), CBM-ZC 59; G, P. imaii (Yokoya, 1939), 3 (sl = 2.5 mm),
CBM-ZC 2699; H, P. doederleini (Doflein, 1902), 3 (sl = 8.6 mm), ZSSM 274/1. A, B redrawn from Mc-
Laughlin & Haig (1973); C redrawn from McLaughlin & Jensen (1996); H redrawn from Asakura (2001).

quadrate to subrectangular. Sternite of fifth
pereopods (eighth thoracomere) separated
into two subovate lobes by shallow median
groove; coxae of fifth pereopods symmet-
rical. Males with very short sexual tube de-
veloped from coxa of right fifth pereopod;
left gonopore without tube or papilla. No
paired pleopods in either sex. Males with
unequally biramous pleopods on pleomeres
3-5. Females with well developed, bira-
mous pleopods on pleomeres 2—4; pleopod
5 with endopod noticeably reduced. Telson
with posterior lobes separated by very small
median cleft; terminal margins oblique,
each with 8 or 9 small spines and 1 larger
spine at posterolateral angles.

Parapagurodes constans (Stimpson, 1858)
Figs. 1E 2E 3K 4H, 5E G

Redescription by Komai (1999:80-88,
figs. 1-4).

Supplemental material examined.—Ja-
pan: Hilgendorf collection, 1 ovig. 2 (sl =
5.5 mm), ZMB 8650; 1 6 (sl = 11.1 mm),
1 ovig. 2 (sl = 10.2 mm), Sagami Bay,
ZMB 17800; 1 3 (sl = 8.7 mm), off Tone
River mouth, Choshi, Chiba, 60 m, 21 Oct
1991, CBM-ZC 59; 2 3 (sl = 6.3, 10.7
mm), Hakodate Bay, 10—20 m, 17 Mar
1995, CBM-ZC 2362.

Diagnosis.—Gill lamellae biserial. Ros-
trum triangular. Maxillule with moderately
well developed external lobe, not recurved.
Right cheliped somewhat suboval in dorsal
view; dorsal surface of palm weakly con-
vex; dorsal surface of carpus with scattered
spines mesiad of midline. Left cheliped
with dorsal surface of palm weakly convex.
Ambulatory legs similar; dactyls slightly
longer than propodi, moderately slender,
laterally compressed, in dorsal view slightly
to prominently twisted, with longitudinal

VOLUME 117, NUMBER 1

sulcus on lateral face and row of corneous
spines on ventral margin; carpi each with
dorsodistal spine and row of low protuber-
ances on dorsal surface. Fourth pereopods
semichelate, preungual process apparently
absent; propodal rasp with several rows of
corneous scales. Sternite of third pereopods
(sixth thoracomere) subrectangular. Sternite
of fifth pereopods (eighth thoracomere) sep-
arated into two somewhat flattened, round-
ed lobes by shallow median depression;
coxae of fifth pereopods symmetrical.
Males with papilla or very short sexual tube
developed from coxa of both right and left
fifth pereopods. No paired pleopods in ei-
ther sex. Males with unequally biramous
left pleopods on pleomeres 3—5. Females
with moderately well developed, biramous
pleopods left on pleomeres 2—4; pleopod 5
reduced. Telson with posterior lobes sepa-
rated by shallow median cleft; terminal
margins broadly rounded, each unarmed or
with few very small spinules adjacent to
cleft.

Parapagurodes imaii (Yokoya, 1939)
Figs. 1G, 2G, H, 3G, 41, 5H

Redescription by Komai (1994:33-38,
figs. 1-3).

Supplemental material examined.—Ja-
pan: 1 ¢ (sl = 2.1 mm), 1 ovig. @ (sl =
1.6 mm), Funakoshi Bay, Iwate, Sanriku,
66 m, 25 May 1995, CM-ZC 1911; 1 ¢ (al
= 2.5 mm), off Takeoka, Boso Peninsula,
ca 80 m, 2 Mar 1995, CBM-ZC 2699.

Diagnosis.—Gill lamellae biserial, but
with slight terminal concavity, cleft or de-
pression. Rostrum triangular. Maxillule
with moderately well developed external
lobe, not recurved. Right cheliped elongate
in large males, dorsal surface of palm con-
vex; dorsal surface of carpus with two lon-
gitudinal rows of spines. Left cheliped with
dorsal surface of palm elevated in midline.
Ambulatory legs somewhat dissimilar, third
sexually dimorphic; dactyls of second and
third right slightly shorter to slightly longer
than propodi, moderately slender, laterally

compressed, in dorsal view barely twisted,
with row of corneous spines on ventral mar-
gin, third left of females broadened, pro-
podus with prominent ventral spine; carpi
each with dorsodistal spine and row of low,
sometimes spinulose protuberances on dor-
sal surface. Fourth pereopods semichelate;
preungual process absent; propodal rasp
with 2 or 3 rows of corneous scales. Ster-
nite of third pereopods (sixth thoracomere)
subcircular to subovate, slightly skewed.
Sternite of fifth pereopods (eighth thora-
comere) separated into two somewhat flat-
tened, rounded lobes by shallow median de-
pression; coxae of fifth pereopods symmet-
rical. Males with very short sexual tube de-
veloped from coxa of both right and left
fifth pereopods. No paired pleopods in ei-
ther sex. Males with unequally biramous
left pleopods on pleomeres 3—5. Females
with moderately well developed, biramous
left pleopods on pleomeres 2—4; pleopod 5
reduced. Telson with posterior lobes sepa-
rated by shallow median cleft; terminal
margins oblique, each with 3 or 4 moderate
to strong spines.

Parapagurodes doederleini (Doflein,
1902)
Figs. 1H, 21, J, 3H, 4J, 5I

Redescription by Asakura (2001:885—
888, figs. 45-47).

Supplemental material examined.—Ja-
pan: 2 6 (sl = 8.1, 9.2 mm), off Kochi,
Tosa Bay, 190 m, 10 Aug 1991, CBM-ZC
184.

Taiwan: 1 3 (sl = 9.3 mm), 1 @ (sl =
7.8 mm), Su-Aou, 100—200 m, 6 Aug 1996,
CBM-ZC 2922.

Diagnosis.—Gill lamellae biserial. Ros-
trum triangular. Maxillule with moderately
well developed endopodal external lobe,
not recurved. Right cheliped stout, dorsal
surface of palm slightly convex; dorsal sur-
face of carpus with covering of spines and
spinulose tubercles. Left cheliped with dor-
sal surface of palm very slightly elevated
in midline. Ambulatory legs dissimilar, dac-

52 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5.

Telson (A-E H, I, dorsal view, G, ventral view of posterior portion). A, Parapagurodes makarovi

McLaughlin & Haig, 1973 6 (sl = 3.5 mm), LACM; B, P. laurentae McLaughlin & Haig, 1973, d (sl = 3.4
mm), LACM; C, P. hartae McLaughlin & Jensen, 1996, 3 (sl = 3.2 mm), RBCPM 974-00368-22; D, P.
gracilipes (Stimpson, 1858), d (sl = 9.0 mm), CBM-ZC 2977; E, P. nipponensis (Yokoya, 1933), 3 (sl = 7.3
mm), CBM-ZC 1162; E G, P. constans (Stimpson, 1858); ¢ (sl = 8.7 mm), CBM-ZC 59; H, P. imaii (Yokoya,
1939), 3 (sl = 2.5 mm), CBM-ZC 2699; I, P. doederleini (Doflein, 1902), 5 (sl = 8.6 mm), ZSSM 274/1. A,
B redrawn from McLaughlin & Haig (1973); C redrawn from McLaughlin & Jensen (1996); I from Asakura

(2001).

tyls longer than propodi, strongly twisted,
second left with row of 40—60 well devel-
oped, comb-like corneous spines on ventral
margin; second right and third each with
longitudinal row of short transverse rows of
setae; carpi each with row of spines on dor-
sal surface. Fourth pereopods subchelate;
preungual process absent; propodal rasp
with 3 or 4 rows of corneous scales. Ster-
nite of third pereopods (sixth thoracomere)
rectangular. Sternite of fifth pereopods

(eighth thoracomere) as narrow rod with
pair of rounded lobes anteriorly; coxae of
fifth pereopods asymmetrical. Males with
very short sexual tube developed from coxa
of right fifth pereopod, left sometimes with
papilla. No paired pleopods in either sex.
Males with unequally biramous left pleo-
pods on pleomeres 3—5. Females with mod-
erately well developed, biramous left pleo-
pods on pleomeres 2—4; pleopod 5 reduced.
Telson with posterior lobes separated by

VOLUME 117, NUMBER 1

broad, deep median concavity; terminal
margins oblique, each with 1—5 moderate to
strong corneous spines.

Results

In her discussion of significant generic
characters, de Saint Laurent-Dechancé
(1966) listed three that are pertinent to our
investigation: sexual tube development, de-
velopment of the external lobe of the max-
illulary endopod, and pleopod number and
development. Perusal of the abbreviated di-
agnoses of the eight species currently as-
signed to Parapagurodes shows that attri-
butes of these three characters are not uni-
versally shared.

While sexual tube length (Fig. 1) varies
from very short to short in P. makarovi
(Figs. 1A, B) and P. laurentae (Fig. 1C, D)
only very short tubes develop in the other
six species (Figs. 1E—J), and occasionally
are not apparent at all. However, recent
studies have shown that sexual tube devel-
opment is known to vary within genera
(e.g., McLaughlin 1997, 2003; McLaughlin
& Lemaitre 2001; Lemaitre & McLaughlin
2003a, 2003b). Nevertheless, P. doederleini
is more importantly distinguished from the
other seven species because in addition to
the very short right sexual tube, the coxae
of the male fifth pereopods are asymmetri-
cal (Fig. 1J).

The external lobe of the maxillulary en-
dopod (Fig. 2) is moderately well devel-
oped in all eight species, but is slightly to
distinctly recurved only in P. gracilipes and
P. nipponensis (Figs. 2D, E).

Parapagurodes was initially character-
ized as having unpaired male pleopods
varying from reduced on pleomeres 3—5 to
completely absent, and female unpaired
pleopods often being reduced on pleomeres
2—4 and absent on pleomere 5. All subse-
quently assigned taxa are described as hay-
ing at least moderately well developed un-
paired, unequally biramous pleopods on
male pleomeres 3—5 and on female pleo-
meres 2—5.

Several other characters frequently in-
cluded in generic diagnoses also have been
examined. Rostral development, for exam-
ple is generally similar among species with-
in a single genus. All eight species have an
acutely developed rostrum, but then so do
many species assigned to other genera.

With the exception of P. constans, all of
the species under consideration herein are
described as having an elongate right che-
liped, at least in large males. In P. hartae
and P. imaii this elongation is considered a
sexually dimorphic character (McLaughlin
& Jensen 1996, Komai 1999), whereas in
P. gracilipes and P. nipponensis apparently
the elongation is growth related (Komai
1998). Similar lengthening of the left che-
liped is reported for these species. In con-
trast, the chelipeds are typically elongate re-
gardless of sex or size in P. makarovi, P.
laurentae and P. doederleini. That cheliped
elongation is comparable among the eight
taxa is doubtful.

Major differences among the eight spe-
cies can be observed in the shape and ar-
mature of the dactyls of the ambulatory legs
(Fig. 3). In P. makarovi and P. laurentae
the dactyls (Figs. 3A, B) are moderately
long, slender, laterally compressed, and in
dorsal view appear straight; the dorsal sur-
faces of the carpi are armed only with a
dorsodistal spine. The dactyls are similarly
straight in P. hartae (Fig. 3C), but vary in
length from shorter to only slightly longer
than the propodi; the carpi each have a row
of low protuberances on the dorsal surface
in addition to the dorsodistal spine. The
dactyls of P. gracilipes and P. nipponensis
(Figs. 3D, E), although moderately long
and laterally compressed, are moderate to
broad and strongly twisted, the ventral mar-
gins of each are provided with a row of
numerous small corneous spines; the dorsal
surfaces of the carpi are provided with one
or more rows of small spines. In contrast,
while the dactyls of P. constans (Fig. 3F)
are longer than the propodi and slightly to
noticeably twisted, the ventral margins each
are armed with fewer and much larger cor-

54 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

neous spines; each carpus is armed only
with a row of low protuberances in addition
to the dorsodistal spine. The dactyls of P.
imaii and P. doederleini are dimorphic, but
do not represent comparable conditions. As
reported by Komai (1999), the third left
dactyl and propodus of females of P. imaii
differ from those of males. In males the
dactyl and propodus of the third left (Fig.
3G) are moderately long and slender as they
are on the second and third right. The fe-
male dactyl (Fig. 3H) is broad and promi-
nently flattened; a well developed calcare-
ous spine is present on the ventrodistal mar-
gin of the propodus. The dimorphism in P.
doederleini involves the dactyls of the sec-
ond pereopods. The left is provided with a
ventral row of closely-spaced, corneous
spines that present a comb-like appearance
(Fig. 31); the right, and the dactyls of the
third pereopods completely lack spines, and
instead are provided with short transverse
rows of setae over the entire length of the
mesial faces (Fig. 3J).

The shape of the anterior lobe of the ster-
nite of the third pereopods and the config-
uration of the sternite of the fifth pereopods
have been proposed as generic or at least
group characters (e.g., McLaughlin 1981,
2003; Lemaitre et al. 1982). The anterior
lobe of the sternite of the third pereopods
is subsemicircular in P. makarovi, P. lau-
rentae, and P. hartae, semicircular or su-
bovate in P. imaii, but subquadrate to sub-
rectangular in P. gracilipes and P. nippo-
nensis and subrectangular in P. constans
and P. doederleini. The sternites of the fifth
pereopods are less clearly definable in these
eight taxa.

The fourth pereopods (Fig. 4) are sub-
chelate or only very weakly semichelate in
P. makarovi and P. laurentae and P. doe-
derleini, but semichelate in the remaining
species. The number of rows of corneous
scales making up the propodal rasps of
these appendages exhibit overlapping intra-
specific variation in all eight species.

The telsons of P. makarovi and P. lau-
rentae (Figs. 5A, B) have straight to weakly

concave or very slightly oblique terminal
margins that are armed with small spines or
spinules. Similar conformation and arma-
ture are seen in P. gracilipes (Fig. 5D) and
to a lesser extent in P. nipponensis (Fig.
SE). In contrast, the terminal margins of the
telsons of P. hartae (Fig. 5C) and P. con-
stans (Fig. 5K G) are broadly rounded and
unarmed or only weakly armed. The telson
of P. imaii (Fig. 5H) differs in having dis-
tinctly oblique terminal margins, each
armed with prominent spines, and the telson
of P. doederleini (Fig. 51) is plainly differ-
ent from the other seven.

Conclusions

From the evidence presented, there can
be little doubt that Parapagurodes, as pres-
ently constituted, represents a heteroge-
neous collection of taxa. Consequently, we
restrict Parapagurodes to the two species
initially assigned, P. makarovi and P. lau-
rentae. Parapagurodes hartae is herein
transferred to Pagurus and the four species
formerly included in Pagurus are returned
to it.

We concur with Komai (1998) that P.
gracilipes and P. nipponensis are closely al-
lied to the bernhardus group of Pagurus,
and undoubtedly should be included in that
group. We do not advocate separating the
bernhardus group from the admittedly
polyphyletic Pagurus at this time, as Pa-
gurus bernhardus (Linnaeus, 1758) is the
type species of the genus. To remove P.
bernhardus and its allied species would
leave the remaining 80 or so species with-
out generic union. Consequently, until such
time as all species currently assigned to Pa-
gurus have been thoroughly recognized and
defined, this genus necessarily must remain
a “‘catch-all’’. In contrast, there is ample
justification to establish a new genus for the
very distinctive P. doederleini as is done
herein.

Dofleinia gen. nov.

Catapagurus: Doflein 1902:624 (in part).—
Miyake 1978:78 (key, in part), 141 (Gn

VOLUME 117, NUMBER 1

part); 1982:232 (key, in part); 1991:232
(key, in part); 1999:232 (key, in part).
Parapagurodes: Asakura 2001:885 (in

part).

Diagnosis.—Gills biserial; 11 pairs. Ros-
trum well developed, acute. Antennal pe-
duncles with supernumerary segmentation.
Maxillule with external lobe of endopod
moderately well developed, not recurved.
Third maxilliped with well developed crista
dentata, 1 accessory tooth. Sternite of third
maxillipeds unarmed. Chelipeds subequal,
right stronger but not necessarily longer.
Second pereopods dimorphic, left with row
of closely-spaced comb-like corneous teeth
on ventral margin, right with ventral margin
unarmed. Third pereopods similar; sternite
with subrectangular anterior lobe. Fourth
pereopods subchelate; dactyl with well de-
veloped preungual process; propodal rasp
consisting of 3 or 4 rows of corneous
scales. Fifth pereopods chelate; coxae of
males asymmetrical. Males with very short
sexual tube developed from right gonopore,
papilla frequently produced from left.

Abdomen well developed, twisted; colu-
mellar muscle usually prominent. Males
without paired first or second pleopods;
with unequally biramous unpaired left ple-
opods 3—5. Females without paired first ple-
opods, with subequally biramous, unpaired,
left pleopods 2—4, pleopod 5 as in male.
Uropods asymmetrical. Telson with distinct
lateral indentations; posterior lobes separat-
ed by very broad median cleft.

Type species.—Catapagurus doederleini
Doflein, 1902.

Etymology.—Named after E Doflein who
first described the type species; gender fem-
inine.

Acknowledgements

The authors acknowledge, with thanks,
the gift of specimens to the first author by
Dr. M. Imafuku, Kyoto University, and the
loan of specimens by Dr. C. O. Coleman,
Naturhistorisches Forschungsinstitut Muse-
um ftir Naturkunde zu Berlin. This work

has been supported in part by a Grant-in-
Aid for Scientific Research (C) from the
Ministry of Education, Science, Culture and
Sports of Japan to Akira Asakura (No.
14540654). This, in part, is also a scientific
contribution from the Shannon Point Ma-
rine Center, Western Washington Universi-

ty.

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Milne-Edwards, A. 1880. Report on the results of
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. 1982. Japanese crustacean decapods and sto-

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(1):57—67. 2004.

Pseudopaguristes bicolor, a new species of hermit crab (Crustacea:
Decapoda: Diogenidae) from Japan, the third species of the genus

Akira Asakura and Takeharu Kosuge

(AA) Natural History Museum and Institute, Chiba. 955-2, Aoba-cho, Chuo-ku,
Chiba 260-8682, Japan, asakura@chiba-muse.or.jp
(TK) Ishigaki Tropical Station, Seikai National Fisheries research Institute, 148-446, Fukai Ota,
Ishigaki, Okinawa, Japan

Abstract.—Pseudopaguristes bicolor, a new species of the recently estab-
lished diogenid genus Pseudopaguristes McLaughlin, is described and illus-
trated from Okinawa, Japan. This is the third species assigned to this genus.

The recently established diogenid genus
Pseudopaguristes McLaughlin, 2002, is
characterized by eight functional gills, male
chelipeds with the right larger than the left
and dissimilar in armature, female chelipeds
similar from left to right, fourth pereopods
with a clump of long capsulate setae on the
carpi, and the paired first and second ple-
opods modified as gonopods. The type spe-
cies, P. janetkae McLaughlin, 2002, was re-
corded from Guam, the Mariana Islands. A
second species, P. bollandi Asakura &
McLaughlin, 2003, was recorded from Oki-
nawa, tropical Japan. The present authors
recently found the third species of this ge-
nus, again from Okinawa. The new species
is very easily separated from both P. janet-
kae and P. bollandi by its characteristic col-
oration and morphology of the male cheli-
peds.

The holotype is deposited in the Natural
History Museum and Institute, Chiba
(CBM-ZC). The terminology used follows
McLaughlin (1974, 2002) with the excep-
tion of the fourth pereopods as defined by
McLaughlin (1997), gill structure by
McLaughlin & de Saint Laurent (1998),
and the posterior carapace by McLaughlin
(2000). Abbreviations used are; coll., col-
lector; and SL, shield length as measured
from the tip of the rostrum to the posterior
margin of the shield.

Pseudopaguristes bicolor, new species
Figs. 1-8

Material.—Holotype: male, SL = 2.65
mm, 78 m, 24°25.5'’N, 124°03.3’E, off Yar-
abu-zaki, Ishigaki-jima Island, Okinawa, 21
Nov. 2002, coll. T. Kosuge, CBM-ZC 6759.

Description.—Eight functional pairs of
quadriserial, phyllobranchiate gills (Fig.
1A). Shield (Fig. 1B) 1.30 times longer
than broad; anterior margin between ros-
trum and lateral projections concave; lateral
projections triangular, with strong submar-
ginal spine; anterolateral angles each with
strong corneous spine; lateral margins con-
vex; posterior margin truncate; dorsal sur-
face slightly convex, with elevated area pre-
sent on each anterolateral portion; scattered
tufts of short setae. Rostrum (Fig. 1B)
prominent, triangular, reaching nearly to
apices of ocular acicles, with terminal
spine. Posterior carapace lateral elements
(Fig. 1B) small, well calcified, unarmed.
Branchiostegites (Fig. 1C) each with row of
spines on dorsal margin anteriorly.

Ocular peduncles (Fig. 1B) moderately
long, 0.75 length of shield. Corneas (Fig.
1B, C) very slightly dilated. Ocular acicles
(Fig. 1B) each terminating in strong, bifid
cormeous spines; separated basally by more
than breadth of rostrum.

Antennular peduncles (Fig. 1D) stout,
with few setae on each segment; when fully

58 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

SA
SS

Fig. 1. Pseudopaguristes bicolor, new species: holotype male (CBM-ZC 6759), SL = 2.65 mm, off Yarabu-
zaki, Ishigaki-jima Is., Okinawa. A, arthrobranch gill lamella; B, shield and cephalic appendages, dorsal; C,
distal half of cephalothorax and cephalic appendages, right, lateral; D, right antennule, lateral; E, right antennal
peduncle, ventral; H, right antennal flagellum. Scales equal 0.5 mm (A) and 1 mm (B-F).

extended, distal margins of ultimate seg-
ments reaching distal margins of corneas;
ultimate segments unarmed; penultimate
segments with ventral margins each bearing
acute spine; basal segments with ventrod-

istal angles each bearing acute spine and
dorsolateral margins each bearing acute
subdistal spine.

Antennal peduncles (Fig. 1B, C, E) mod-
erately long, when fully extended, reaching

VOLUME 117, NUMBER 1

Fig. 2.

Pseudopaguristes bicolor, new species: holotype male (CBM-ZC 6759), SL = 2.65 mm, off Yarabu-

zaki, Ishigaki-jima Is., Okinawa. Right mouthparts: A, mandible, internal; B, maxillule, external; C, same,
endopod; D, maxilla, external; E, first maxilliped, internal; E second maxilliped, external; G, third maxilliped,

external; H, same, internal.

distal 0.30 of ocular peduncles, scarcely se-
tose; fifth segments with dorsal margins
each bearing acute subproximal spine;
fourth segments with dorsodistal margins
each bearing acute spine and ventrodistal
margins each bearing another acute spine;
third segments with prominent spine at ven-
trodistal margin; second segments with dor-
solateral distal angles produced, terminating
in prominent bifid spine, dorsomesial distal

angles each with acute corneous spine; first
segment unarmed. Antennal acicles mod-
erately long, straight; dorsomesial margins
each with 3 (right) or 4 (left) spines; dor-
solateral margins each with 2 strong spines;
distal margins each with 2 strong spines.
Antennal flagella (Fig. 1F) consisting of
about 18 articles, each article with several
short setae.

Mandible (Fig. 2A) without distinguish-

Fig. 3.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Pseudopaguristes bicolor, new species: holotype male (CBM-ZC 6759), SL = 2.65 mm, off Yarabu-

zaki, Ishigaki-jima Is., Okinawa. Right cheliped: A, dorsal; B, mesial; C, lateral.

ing characters. Maxillule (Fig. 2B, C) with
external lobe of endopod well developed,
articulated, and recurved; internal lobe with
2 bristles. Maxilla (Fig. 2D) with moder-
ately narrow scaphognathite. First maxilli-
ped (Fig. 2E) with well developed, setose
epipod. Second maxilliped (Fig. 2F) with-
out distinguishing characters. Third maxil-
liped (Fig. 2G, H) with carpus bearing dor-
sodistal spine; merus with dorsodistal spine,

ventral margin bearing 4 spines; ischium
with strong ventrodistal spine, crista dentata
well-developed, no accessory tooth; basis
with 2 sharp spines.

Chelipeds subequal; right (Fig. 3) larger
than left. Dactyl as long as palm; terminat-
ing in broad corneous claw; dorsal face flat,
with scattered large tubercles; cutting edge
with several calcareous teeth. Fixed finger
terminating in corneous claw; dorsal face

VOLUME 117, NUMBER 1

Fig. 4. Pseudopaguristes bicolor, new species: holotype male (CBM-ZC 6759), SL = 2.65 mm, off Yarabu-
Zaki, Ishigaki-jima Is., Okinawa. Left cheliped: A, dorsal; B, mesial; C, lateral.

flat, with scattered large tubercles; cutting
edge with several calcareous teeth. Palm
1.07 length of carpus; dorsal surface flat,
with scattered large tubercles; dorsomesial
margin with row of very strong spines; dor-
solateral margin of palm and fixed finger
with row of strong spines. Carpus 0.50
length of merus; dorsal face with scattered
large tubercles, dorsolateral margin with
row of strong conical-shaped spines, dor-
somesial margin with row of very strong

spines. Merus with dorsal face bearing 2
distal spines, subdistal transverse row of
several spines, and row of spines on re-
mainder of dorsal margin, tips semitrans-
parent; ventromesial margin with 3 widely-
separated strong spines, tips semitranspar-
ent, ventrolateral margin with row of spines
or tubercles. Ischium unarmed. Coxa with
acute spine ventromesially.

Left cheliped (Fig. 4) slenderer than
right. Dactyl with dorsal face without tu-

62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5.

Pseudopaguristes bicolor, new species: holotype male (CBM-ZC 6759), SL = 2.65 mm, off Yarabu-

zaki, Ishigaki-jima Is., Okinawa. Second left pereopod: A, lateral; B, propodus, ventral; C, dactyl, propodus,

and carpus, mesial; D, merus, mesial.

bercles; number of tubercles or spines on
dorsal faces of palm and carpus fewer; mer-
us with ventromesial and ventrolateral mar-
gins bearing 6 and 5 spines, respectively;
other surfaces similar to right.

Second pereopods (Fig. 5) with armature
similar from left to right; right 1.10 length
of left. Basically, spines on ambulatory pe-
reopods with semitransparent tips. Dactyls

1.10 (eft) or 1.25 (ight) length of propodi,
each terminating in strong corneous claw;
dorsal margins each with row of strong
spines; ventral margins each with row of 9
strong corneous spines and, on left, accom-
panied with 2 tiny corneous spines mesial-
ly. Propodi 1.60 (left) or 1.55 (right) length
of carpi, each with row of 10 strong spines
on dorsal margin; ventral faces each with 2

VOLUME 117, NUMBER 1

E 0.5 mm

|) Imm

Fig. 6. Pseudopaguristes bicolor, new species: holotype male (CBM-ZC 6759), SL = 2.65 mm, off Yarabu-
zaki, Ishigaki-jima Is., Okinawa. Third left pereopod: A, lateral; B, dactyl and propodus, mesial (propodus
slightly ventral view); C, carpus, merus and ischium, lateral. Fourth left pereopod: D, dactyl, propodus and

carpus, lateral; E, ventral setae of carpus.

irregular rows of widely-separated, tiny
corneous spines, ventromesial distal mar-
gins with | (right) or 2 (left) acute corneous
spines. Carpi 0.55 (left) or 0.60 (right)
length of meri, each with strong, corneous
or corneous-tipped, slender spine at dorso-

distal angle and row of 5 slender spines on
dorsal face mesially. Meri with ventral mar-
gins each with row of slender spines and,
on left, accompanied with 2 small spines
mesially; dorsal margins each with row of
spines. Ischia each with few, slender cor-

64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 7. Pseudopaguristes bicolor, new species: holotype male (CBM-ZC 6759), SL = 2.65 mm, off Yarabu-
zaki, Ishigaki-jima Is., Okinawa. A, right fifth pereopod. Left first pleopod: B, external; C, internal; D, distal
portion, internal, enlarged. Left second pleopod: E, external; FE distal portion, enlarged, external; G, same, mesial.
H, third pleopod. I, telson. Scales equal 1 mm (A, H, I) and 0.2 mm (B-E).

VOLUME 117, NUMBER 1

Fig. 8.

Pseudopaguristes bicolor, new species: holotype male (CBM-ZC 6759), SL = 2.65 mm, off Yarabu-

zaki, Ishigaki-jima Is., Okinawa. A, dorsolateral view; B, right cheliped, mesial; C, left second pereopod, lateral;
D, same, mesial; E, left third pereopod, lateral; K same, mesial. Photo by A. Asakura.

neous-tipped spines dorsally and small
spine at ventromesial distal angle. Coxae
unarmed.

Third pereopods (Figs. 6A—C) with ar-
mature similar from left to mnght, nght 1.05
length of left. Dactyls 1.20 (left) or 1.25
(right) length of propodi, each terminating
in strong corneous claw; mesial faces each

with row of small corneous spines ventrally
and 4 (left) or 2(right) small spines dorsal-
ly; dorsal margins with few tiny spines on
proximal 0.25; ventral margins each with
row of 9 strong corneous spines. Propodi
1.60 length of carpi; dorsal faces unarmed
(left) or row of small tubercles or spines
(right); ventral faces each with row of

66 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

small, widely-separated corneous spines,
ventromesial distal angles each with 1 (left)
or 2 (right) acute corneous spines. Carpi
0.70 (eft) or 0.80 (right) length of meri,
each with strong spine at dorsodistal angle;
dorsal margin unarmed (left) or with minute
subproximal spine (right). Meri with ventral
margins each bearing 3 (left) or 2 (right)
small spines; dorsal margins each with row
of spines. Ischia each with small dorsodistal
spine and another small ventrodistal spine.
Coxae unarmed.

Sternite of third pereopods with anterior
lobe rectangular, unarmed.

Fourth pereopod (Fig. 6D) subchelate.
Dactyl terminating in strong corneous claw;
prominent preungual process present at
base of claw; ventral face with 1 corneous
spine laterally. Propodal rasp with 2 rows
of corneous scales. Carpus with large dor-
sodistal spine; ventral face with clump of
long capsulate setae (Fig. 6E).

Fifth pereopod (Fig. 7A) chelate; dactyl
and propodus with well-developed rasps.

Male first pleopods (Fig. 7B—D) paired,
modified as gonopods; basal lobe bearing
few setae at superior mesial angle; inferior
lamella with distal margin bearing row of
short, hooked spines, and lateral margin
with row of setae; internal lobe with row of
setae on mesial margin; external lobe dis-
tinctly exceeding inferior lamella in distal
extension. Second pleopods (Fig. 7E—G)
paired, modified as gonopods; basal seg-
ment naked; endopod with several long se-
tae; appendix masculina twisted; lateral and
distal margins and inferior face with mod-
erately long setae. Third (Fig. 7H) to fifth
left pleopods each with exopod well devel-
oped, endopod reduced.

Uropods asymmetrical, left larger than
right; rasps of exopods and endopods well
developed; protopods each with row of
spines posteriorly.

Telson (Fig. 71) with lateral constrictions;
anterior portion unarmed; posterior lobes
separated by deep median cleft, left lobe
larger than right, terminal margins fringed
with spines.

Female unknown.

Color in life (Fig. 8).—Shield white; an-
tennules with flagella and ultimate segment
yellow, setae on flagella blue, penultimate
and basal segments red; antennas with fla-
gella bearing alternative red and white
bands, fifth segment with middle red band,
proximal half of third segment red, second
segment red except for white distal spines,
first segment red except for ventral face, an-
tennal acicle with subdistal red band, other
surfaces of antennas white; ocular pedun-
cles yellow, each with red band on proximal
0.25; ocular acicles red except for white
distal spines; third maxillipeds with propo-
dus, carpus, and merus and penultimate
segment of exopod each bearing middle red
band, other surfaces white; second maxil-
lipeds with middle red band on penultimate
segment of exopod. Both chelipeds and sec-
ond through fifth pereopods with irregular
red area on each segment.

Etymology.—From the Latin bicolor, two
colors, in reference to the alternating red
and white color bands on the pereopods
characteristic to this species.

Distribution.—Known only from the
type locality.

Remarks.—Despite their general similar-
ities in morphology, the new species, P. bi-
color, is readily distinguished from both P.
jJanetkae and P. bollandi by differences in
coloration in life. The chelipeds and the
second and third pereopods in P. bicolor
have alternating red and white bands. These
appendages are uniformly red in P. bollan-
di, and, in P. janetkae, the meri and carpi
and proximal half of palm of the chelipeds
are cranberry-red and the carpi, propodi and
dactyls of the second and third pereopods
are light cream, tinged with yellow.

Morphologically, P. bicolor is similar to
both P. janetkae and P. bollandi, but some
differences are seen among them. The most
striking difference that separates P. bicolor
from both P. janetkae and P. bollandi is the
degree of dissimilarity in the chelipeds in
males. In male P. janetkae and P. bollandi,
the chelipeds are very unequal, and arma-

VOLUME 117, NUMBER 1

tures are much stronger in the right than in
the left. However, in male P. bicolor, the
chelipeds are subequal and the dissimilarity
of the armature is not so large as in those
in P. janetkae and P. bollandi. Furthermore,
the dorsal surfaces of the chelae are provid-
ed with tubercles in P. bicolor and P. bol-
landi but with spines in P. janetkae.

Other minor difference includes the fact
that, although a preungual process is absent
in P. bollandi, both P. bicolor and P. ja-
netkae have a very prominent preungual
process developed at the base of the claw,
giving the dactyl a quasi-chelate appear-
ance. However, so few specimens have
been reported in any of these species (four
specimens in P. janetkae, one in P. bollandi
and one in P. bicolor), it is not possible to
evaluate variability. Thus, we expect future
collection efforts to provide more precise
information on morphological descrimina-
tion between the species.

Acknowledgements

The authors are most grateful to Dr. Patsy
A. McLaughlin (Shannon Point Marine
Center, Western Washington University) for
her elaborate review of the manuscript and
the captain Higa Koei (Okinawa) for the
successful cruise to collect this important
material. The comments by Jacques Forest
(Muséum national d’ Histoire naturelle, Par-
is), D. L. Rahayu (Research Center for
Oceanography, Indonesia) and an anony-
mous reviewer greatly improve the final

draft of the manuscript. This work was part-
ly supported by a Grant-in-Aid for Scien-
tific Research (C) from the Ministry of Ed-
ucation, Science, Culture and Sports of Ja-
pan awarded to Akira Asakura (No.
14540654).

Literature Cited

Asakura, A., & P. A. McLaughlin. 2003. Pseudopa-
guristes bollandi, new species, a distinct hermit
crab (Crustacea: Decapoda: Diogenidae) from
Japan.—Proceedings of the Biological Society
of Washington 116:453—463.

McLaughlin, P. A. 1974. The hermit crabs (Crustacea,
Decapoda, Paguridae) of northwestern North
America.—Zoologische Verhandelingen 130:1—
396.

. 1997. Crustacea Decapoda: hermit crabs of

the family Paguridae from the KARUBAR

cruise in Indonesia. In A. Crosnier & P. Bouch-
et, eds., Résultats des Campagnes MUSOR-

STOM, 16.—Mémoires du Muséum national

d’Histoire naturelle 172:433-572.

. 2000. Crustacea Decapoda: Porcellanopagu-

rus Filhol and Solitariopagurus Viirkay (Pagur-

idae), from the New Caledonian area, Vanuatu
and the Marquesas: new records, new species.

In A. Crosnier, ed., Résultats des Campagnes

MUSORSTOM, volume 21.—Mémoires du

Muséum national d’Histoire naturelle, Paris,

volume 184:389—414.

. 2002. Pseudopaguristes, a new and aberrant

genus of hermit crabs (Anomura: Paguridea: Di-

ogénidae).—Micronesica 34:185—-199.

, & M. de Saint Laurent. 1998. A new genus

for four species of hermit crabs formerly as-

signed to the genus Pagurus Fabricius (Deca-
poda: Anomura: Paguridae)—Proceedings of

the Biological Society of Washington 111:158—

187.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
117(1):68-75. 2004.

A new species of axiid shrimp from chemosynthetic communities of
the Louisiana continental slope, Gulf of Mexico
(Crustacea: Decapoda: Thalassinidea)

Darryl L. Felder and Brian Kensley

(DLF) Department of Biology, University of Louisiana at Lafayette, Louisiana 70504, U.S.A.,
e-mail: DLF4517 @louisiana.edu;
(BK) Department of Systematic Biology, National Museum of Natural History, Smithsonian
Institution, Washington, D.C. 20560-0163, U.S.A., e-mail: kensley.brian@nmnh.si.edu*

Abstract.—Calaxius carneyi, new species (Axiidae), is described from two
male specimens collected by manned submersibles working near hydrocarbon
seeps in deep waters (544 m) on the continental slope off Louisiana, in the
northern Gulf of Mexico. Both specimens were taken adjacent to communities
of clams that comprise a major constituent of chemosynthetic assemblages at
the collection site. The new species is characterized in part by ventrally truncate
abdominal pleura, as opposed to the acutely triangular or broadly rounded
pleura found in other known members of Calaxius, only one of which is known
to occur in the Atlantic Ocean. The new species is readily distinguished from
its congeners by unique dentition of its heavy triangular rostrum and postrostral
carapace, its short eyestalks and antennal acicle, the absence of well-defined
teeth on the massive chelipeds, and the narrow, subtriangular telson. Chelipeds,
pleopods and uropods of the two known specimens herewith described are
covered extensively by long setae, many of which are plumose and densely
fouled by flocculent debris.

Recent investigations of methane cold
seeps in the Gulf of Mexico have discov-
ered a number of previously undescribed
taxa associated with chemosynthetic com-
munities in deep waters of the continental
slope (e.g., Gustafson et al. 1998). How-
ever, some collections from these unique
habitats consist of single specimens, and
comparative studies have been deferred
pending recovery of additional materials.
One such case was presented by the collec-
tion of a single, somewhat fragmented,
molted integument from the male of an ap-
parently undescribed axiid mud_ shrimp,
collected in 1988 during a dive of the
manned submersible Pisces IJ. A second,
smaller, intact male specimen, was obtained

* Deceased.

from an adjacent site in 1992 with a shal-
low core sampler deployed by the Johnson
Sea-Link manned submersible. Collections
on subsequent dives by submersibles and
vessel-based box coring in this area have
brought no additional materials to our at-
tention.

While the female of this species remains
unknown, it is readily apparent that the spe-
cies is undescribed. The marked size dif-
ference between the intact specimen and the
earlier recovered exuvia provides a glimpse
of ontogenetic variation in characters, and
allows us to select diagnostic characters that
should apply to a wide size range. Also,
from familiarity with typical sexual dimor-
phism in congeneric species, we expect that
the description here provided will serve ad-
equately for identification of female speci-
mens, if encountered.

VOLUME 117, NUMBER 1

Upon arrival at the surface, specimens
were fixed in 10% formalin (with Rose
Bengal stain for only the 1992 collection),
transferred to 80% ethyl] alcohol, and finally
archived in 70% ethyl alcohol. Carapace
length (CL) was measured from the poste-
rior margin of the orbit to the posterior mar-
gin of the carapace midline. Total length
(TL) was measured from the tip of the ros-
trum to the tip of the extended telson. Spec-
imens are archived in the National Museum
of Natural History (USNM), Smithsonian
Institution, Washington, D.C.

Family Axiidae Huxley, 1879

Calaxius Sakai & de Saint Laurent, 1989
Calaxius carneyi, new species
Figs. 1 & 2

Material examined.—Holotype, USNM
1009165, male, CL 10.1 mm, TL 26.5 mm,
Johnson Sea-Link submersible sta 3269,
box core B, deployed from submersible
about 2 m from chemosynthetic mussel
community, Bush Hill site, 544 m, Louisi-
ana continental slope, northern Gulf of
Mexico, 27°46.904'N, 91°30.286'W, 11
Aug 1992. Paratype, USNM 1009166, exu-
via of male, CL 18.3 mm, TL 50.5 mm,
submersible Pisces I sta 880031 (8831), lo-
cation and depth same as for holotype, Aug
1988.

Diagnosis.—Rostrum heavy, triangular,
extending slightly more than twice length
of eyes, bearing flattened, upturned terminal
spine and pair of similar upturned subter-
minal spines. Antennal acicle short, over-
reaching proximal third of penultimate
(fourth) peduncular article. Chelipeds mas-
sive, lacking well defined teeth. Carapace
bearing dentate lateral and submedian ca-
rina. Pereopodal epipods and pleurobranchs
present. Abdomen with pleura 3—5 ventrally
truncate, bearing small anterior and poste-
rior marginal denticles; lacking male pleo-
pod 1; appendices internae on pleopods 2—
5; uropodal exopod bearing tranverse su-
ture; terminated by narrow, subtriangular
telson.

Description of holotype.—Integument
firm but pliable, with numerous clumps of
elongate, plumose, fouled setae, often ob-
scuring underlying structures on chelae,
pleopods, uropods, and telson; calcification
most heavy in carapace teeth and chelae.
Carapace with posterior midline elevated,
bracketed on either side by paired setose
punctae, midline elevation becoming a
rounded crest in cardiac region where sur-
mounted by a slight but distinct prominence
or tubercle, marked dorsally by translucent
or worn area (Fig. 1a); rostrum heavily cal-
cified, triangular, slightly more than twice
length of eyes, terminal spine subacute, up-
turned, dorsoventrally flattened, triangular
in dorsal view; subterminal pair of spines
similar to terminal, also upturned, imparting
concave appearance to flattened dorsal sur-
face of rostrum (Fig. 1b, c); supraocular
spines (lacking in the paratype, a larger
specimen) and supraorbital spines strong,
similar in calcification, shape and orienta-
tion to subterminal pair; lateral carina orig-
inating from supraorbital spine, diminishing
immediately anterior to second spine or
tooth, continuing as a low ridge toward pos-
terior; submedian carina originating from
posteriormost of two slightly offset sub-
median teeth, becoming ill-defined toward
posterior; median carina a weak crest bear-
ing a worn tubercle near its posterior end,
and otherwise lacking ornamentation.

Sternite of fourth pereopods (seventh
thoracic somite) with deep median slit, tho-
racic shield produced to form acute, mar-
ginally sinuate, triangular flange to either
side; 3-branched carina set between articu-
lations of fourth pereopods (Fig. 2a). Ab-
dominal pleuron | narrowed, acute ventral-
ly (Fig. la); pleuron 2 ventrally broad, with
an angular tooth or acute corner at the pos-
teroventral end; pleura 3—5 ventrally trun-
cate, with small acute tooth on anteroven-
tral margin and another on posteroventral
margin; pleuron 6 with small acute tooth on
anteroventral margin and broad triangular
flange at posteroventral margin.

Eyestalks small, subcylindrical, reaching

70 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

\ a, f c, d e b

Fig. 1. Calaxius carneyi, new species (where setation is shown, setules and flocculent coating of plumose
appendages not fully illustrated). a—-e, holotype male, USNM 1009165: a, carapace, abdomen, left pereopods 1
and 3—5 in lateral view, with right pereopod 2 internal surface; b, anterior carapace, eyes and antennal peduncles,
right side, lateral view, setation not shown; c, anterior of carapace in dorsal view; d, right pereopod 1 or major
cheliped, internal surface; e, right pereopod 2, external surface. f, paratype male, USNM 1009166: right pereopod
1 or major cheliped, external surface. Scale bars indicate 2.0 mm.

VOLUME 117, NUMBER 1

Fig. 2.

Calaxius carneyi, new species, holotype male, USNM 1009165 (where setation is shown, setules and

flocculent coating of plumose appendages not fully illustrated). a, posterior thoracic sternites and coxae of
pereopods 3—5, ventral view, setation not shown; b, left maxilliped 3, external surface; c, endopod of left
maxilliped 3, internal surface; d, pereopod 3, external surface; e, pereopod 4, external surface; f, pereopod 5,
external surface; g, abdominal somite 6, telson and uropods, dorsal surface. Scale bars indicate 2.0 mm.

almost to midlength of rostrum (Fig. la, b,
c). Cornea terminal, slightly globose, di-
ameter equal to or slightly exceeding that
of eyestalk.

Antennular peduncle reaching well be-
yond rostrum (Fig. 1b). Antennal peduncle
bearing produced nephridiopore proximov-
entrally, second article bearing dorsodistal
spine overreaching much of acicle, acicle
short, not bifid, overreaching proximal third
of penultimate (fourth) peduncular article,
third peduncular article distally bearing
strong ventromesial spine. Maxilliped 3 ba-
sis bearing short, acute mesial spine (Fig.
2b); ischium of endopod with strong, dis-
tally elevated crista dentata on internal sur-
face, bearing about 16 spines, distalmost of

which are largest and most strongly direct-
ed mesiad (Fig. 2c); merus with two mesial
spines, one near or just short of midlength,
the other larger and in distal third; carpus
with short triangular, tooth at distal extreme
of flexor margin; all articles of endopods
bearing fields of long setae, many dense
and heavily plumose, especially on mesiad
and internal surfaces.

Pereopods 1—4 bearing epipods. Pereo-
pods 2—4 with pleurobranchs above coxae
(on thoracic somites 5—7).

Chelipeds (pereopod 1) similar in form
and ornamentation on the 2 sides, right the
heaviest (Fig. la, d); ischium with single
well-defined spine on inferior margin; mer-
us with single flattened spine near mid-

72 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

length of inferior margin, which is weakly
marked by adjacent sinuation or serration,
distal corner of flexor margin on external
side forming short, heavy raised spine (Fig.
la); carpus very short, bearing numerous
patches of long setae on external surface,
dorsal margin weakly tuberculate, terminat-
ing distally in blunt tooth, ventral carina of
external surface forming flange distally
which terminates in flattened, weakly
hooked tooth; propodus very thick and
heavily calcified, lacking well-defined teeth
on weakly tuberculate dorsal margin, bear-
ing numerous patches of long setae on ex-
ternal surface, including among tubercles of
dorsal surface, along well-marked carina of
ventral margin, below cutting edge of fixed
finger, and proximal to gape, external sur-
face proximal to dactylus with scattered
low tubercles and granules, fixed finger
bearing two erect teeth on proximal half
and single less erect tooth in distal half, ter-
minus spiniform, internal surface with weak
carina adjacent to and slightly below cut-
ting edge; movable finger very thick and
heavy, with dense patches of long setae ex-
ternally and dorsally, cutting edge bearing
rounded to lobiform tooth in proximal third,
broad ill-defined tooth or sinuous lobe in
distal two-thirds, terminus weakly hooked
and subacute, a carina above cutting edge
on internal surface. Pereopod 2 (Fig. le)
merus lacking marginal spines, combined
length of ischium and merus about equal to
combined length of carpus and propodus,
length of dactyl about half total length of
propodus, opposable cutting edges of fin-
gers corneous, finely pectinate, distinctly
spooned distally. Pereopod 3 (Fig. 2d) mer-
us lacking marginal spines, external surface
of propodus bearing five sets of corneous
spiniform setae, set individually or in short
transverse rows of two or three near inferior
margin, fewer sets and few such setae near
superior margin, falcate dactylus with three
distinct corneous spiniform setae on exter-
nal surface, distally with two more very
small ones set near flexor margin, and a
sharp corneous spine forming terminus. Pe-

reopod 4 (Fig. 2e) merus lacking marginal
spines, external surface of propodus bearing
six sets of corneous spiniform setae, set in-
dividually or in transverse rows of two to
four near inferior margin, four such sets of
one to three setae near superior margin, fal-
cate dactylus with five distinct corneous
spiniform setae on external surface, distally
with additional very small corneous seta,
sharp corneous spine forming terminus. Pe-
reopod 5 (Fig. 2f) merus and propodus
lacking marginal spines, propodus bearing
stiff bristles at distal inferior end of pro-
podus, concealed by dense distal fields of
setae; lanceolate dactylus twisted laterally,
opposed to terminal bristles of propodus
when flexed.

Pleopod 1 absent, posterior pleopods all
bearing dense cover of long, plumose,
heavily fouled setae; appendix interna pre-
sent on pleopods 2—5. Uropodal exopod
(Fig. 2g) bearing four spines along external
margin and an articulated spine where this
margin meets the transverse suture, five ad-
ditional spines along transverse suture, and
no spines on dorsal surface, long setae
forming dense fringe on margins, but on
dorsal surface limited to few patches near
external margin; endopod with single
strong spine at distal end of external margin
and another small spine overreaching distal
margin at end of weak median ridge, long
setae forming dense fringe on margins, and
distributed in patches on dorsal surface near
external margin and along medial ridge.
Telson length distinctly greater than its bas-
al width, tapering toward posterior, widest
at lateral lobes in proximal one-quarter of
length, single pair of strong fixed dorsal
spines in anterior half, two to four fixed
marginal serrations or spines posterior to
proximal lobes, and two pairs of short, ar-
ticulated marginal spines in distal third of
lateral margins, distal margin evenly con-
vex, densely setose.

Variations.—Paratype: Spination and tu-
berculation in the exuvia of this larger spec-
imen differ in several minor ways from or-
namentation in the holotype. There are few-

VOLUME 117, NUMBER 1

er spines on the margin of the rostrum, as
the supraoculars are not present. The mar-
gin of the rostrum is somewhat broadly
concave in this region on the paratype, al-
though it retains an overall triangular shape.
Dorsal tuberculation of the propodal palm
is less evident than in the holotype. Spines
on the cutting edge of the major chela differ
slightly in shape from those on the holo-
type, but the pattern and placement of this
spination is conserved (Fig. If). Granula-
tion on the internal surface of the propodus
in the paratype is stronger than that in the
holotype. The external margin of the uro-
podal exopod in the paratype male bears
five rather than four fixed spines, while the
external margin of the endopod bears three
spines rather than a single one. Lateral mar-
gins the telson bear up to five serrations or
fixed lateral spines in the paratype, and the
pairs of articulated marginal spines are rel-
atively smaller than in the holotype and
very difficult to discern. Small angular,
acute corners or teeth on the anteroventral
margins of abdominal pleurae 3—5 are also
more difficult to discern in the holotype.
These appear to be somewhat worn or
smoothed off in the paratype, although the
acute posteroventral margins remain readily
evident.

Etymology.—The species is named for
Robert S. Carney, Louisiana State Univer-
sity, Baton Rouge, who oversaw collections
of the specimens and made these materials
available for our study. His own work on
hydrocarbon vent communities of the Gulf
of Mexico (see Carney 1994) has brought
needed attention to these remarkable assem-
blages of marine organisms.

Remarks.—Yen species were listed in a
recent review of Calaxius by Kensley &
Hickman (2001). The present description
accounts for the eleventh known member
and only the second species to be found in
the Atlantic Ocean. There seems little doubt
as to the generic placement of this new spe-
cies, given the dentate rostrum twice as
long as the eyestalks, the dentate carapace
carinae, the transverse suture on the uro-

podal exopod, presence of pereopodal epi-
pods and pleurobranchs, the absence of ple-
opod | in the male, and the presence of
appendices internae on pleopods 2-5 (see
Poore 1994:97). The specimens lack strong
dentition on the dorsal surface of the first
chelipeds, as seen in Acanthaxius Sakai &
de Saint Laurent, 1989, which they some-
what resemble.

In contrast to the known congeners and
the original generic definition (Sakai and de
Saint Laurent 1989:84), the rostrum of C.
carneyi is heavier and more broadly trian-
gular, the eyestalks and antennal acicle are
comparatively shorter, and the palm of the
cheliped lacks well-separated and defined
teeth dorsally, having at most a covering of
low tubercles, and the telson is distinctly
narrowed posteriorly or subtriangular. The
abdominal pleura of C. carneyi resemble
neither the acutely triangular plates seen in
four other previously described species nor
the broadly rounded plates seen in five oth-
er species; rather, pleura 3—5 are ventrally
truncate with small anterior and posterior
marginal denticles.

The only species of Calaxius previously
reported from the broad geographic area of
the Gulf of Mexico and contiguous regions
is C. oxypleura (Williams, 1974), recorded
from the Straits of Florida. This species has
abdominal pleura 3—5 ventrally angular or
acute, rather than truncate, and a narrow
dentate rostrum unlike that of C. carneyi.

Ecology.—A dense cover of flocculent
materials on plumose setae of both the ho-
lotype and paratype exuvia (much of which
has disintegrated over time in alcohol, or
which was brushed free in the course of
morphological studies) may derive from the
unique environments inhabited by these an-
imals, but nothing is known of the burrow
structure or feeding behavior. The floccu-
lent coatings on the axiid setae could be a

‘passive result of these animals’ trophic ties

to members of the chemosynthetic com-
munity, at a primary or secondary consum-
er level. Accumulations of mussels and
worms near hydrocarbon seeps in the north-

74 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ern Gulf of Mexico do in varied ways de-
pend upon methanotrophic or sulfur-oxidiz-
ing bacteria for metabolic resources (see
Van Dover 2000:363—366). These bacteria
occur in animal tissues as endosymbiotic
cells or as scattered mats immediately sur-
rounding the seeps. They may directly ex-
ploit methane as both a carbon and energy
source or, much as in hydrothermal vent en-
vironments, directly oxidize rich cold-seep
sources of sulfides for metabolic energy.
Precipitates and bacteria might simply ac-
cumulate on or among the setules of highly
plumose setae, perhaps as these axiids
move about or ventilate burrows in these
matted settings. While the flocculent mate-
rials could simply mask movements among
bacterial mats or mussel communities, we
cannot rule out that the axiids themselves
may directly consume accumulations of
chemosynthetic bacteria, either by access-
ing exposed mats or undertaking behaviors
that favor the forming of such accumula-
tions within and along walls of their bur-
rows. It is suspected that other thalassini-
deans engage in burrow-modulated feeding
behaviors that are microbially based, albeit
in reduced interstitial waters of shallow
hypoxic environments (Felder 2001) where
at least one species lives in apparent asso-
ciation with lucinid bivalves harboring che-
mosynthetic gill bacteria.

Even if C. carneyi could be shown to de-
pend upon the chemosynthetic community
as a nutritional resource, perhaps by stable
isotope measurements, this would not nec-
essarily confirm its restriction to occurrence
with chemosynthetic communities of hy-
drocarbon seeps. As has recently been re-
ported for infaunal worms associated with
methane seeps off California (Levin et al.
2000), infaunal thalassinideans are also
likely pre-adapted to organic-rich, reducing
environments, and may in fact be widely
distributed forms that do not strictly exhibit
chemosynthesis-based trophic specializa-
tions. Owing to very limited general sam-
pling for infaunal macrocrustaceans from
slope environments in the Gulf of Mexico

to date, its occurrence in sediments other
than those near cold seeps cannot be ruled
out.

Acknowledgements

For providing the specimens, we thank
R. Carney and his associates at Louisiana
State University, Baton Rouge. Support for
the present study was furnished to DLF un-
der U.S. Department of Energy grant no.
DE-FG02-97ER12220 for studies of ende-
mism in the northern Gulf of Mexico. We
are grateful to S. Brooke, C. Allen, and C.
Young for field assistance and sharing ship
time funded under support of NSF grant no
0243688-OCE (to C. Young), in the course
of our continuing studies of hydrocarbon
vent infaunal decapods. This is contribution
no. 98 from the University of Louisiana
Laboratory for Crustacean Research.

Literature Cited

Carney, R. S. 1994. Consideration of the oasis analogy
for chemosynthetic communities at Gulf of
Mexico hydrocarbon vents—Geo-Marine Let-
ters 14:149-159.

Felder, D. L. 2001. Diversity and ecological signifi-
cance of deep-burrowing macrocrustaceans in
coastal tropical waters of the Americas (Deca-
poda: Thalassinidea).—Interciencias 26:2—12.

Gustafson, R. G., R. D. Turner, R. A. Lutz, & R. C.
Vrijenhoek. 1998. A new genus and five new
species of mussels (Bivalvia, Mytilidae) from
deep-sea sulfide/hydrocarbon seeps in the Gulf
of Mexico.—Malacologia 40:63-113.

Huxley, T. H. 1879. On the classification and distri-
bution of the crayfishes.—Proceedings of the
Zoological Society of London 1878:752-788.

Kensley, B., & C. P. Hickman, Jr. 2001. A new species
of Calaxius Sakai & Saint Laurent, 1989, from
the Galapagos Islands (Crustacea: Decapoda:
Axiidae).—Proceedings of the Biological Soci-
ety of Washington 114:484—488.

Levin, L. A., D. W. James, C. M. Martin, A. W. Rath-
burn, L. H. Harris, & R. H. Michener. 2000. Do
methane seeps support distinct macrofaunal as-
semblages? Observations on community struc-
ture and nutrition from the northern California
slope and shelf.—Marine Ecology Progress Se-
ries 208:21—39.

Poore, G. C. B. 1984. A phylogeny of the families of
Thalassinidea (Crustacea: Decapoda) with keys

VOLUME 117, NUMBER 1 75

to families and genera.—Memoirs of the Mu- Van Dover, C. L. 2000. The Ecology of Deep-Sea Hy-

seum of Victoria 54:79—120. drothermal Vents. Princeton University Press,
Sakai, K., & M. de Saint Laurent. 1989. A check list Princeton, New Jersey. 424p.

of Axiidae (Decapoda, Crustacea, Thalassini- Williams, A. B. 1974. Two new axiids (Crustacea: De-

dea, Anomura), with remarks and in addition capoda: Thalassinidea: Calocaris) from North

descriptions of one new subfamily, eleven new Carolina and the Straits of Florida.—Proceed-

genera and two new species.—Naturalists 3:1— ings of the Biological Society of Washington

104. 87(39):45 1-464.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(1):76-87. 2004.

Description of a new Synidotea species (Crustacea: Isopoda:
Valvifera: Idoteidae) from Hawaii

Wendy Moore

University of Arizona, Department of Entomology, Tucson, Arizona 85721-0036 U.S.A.,
email, wmoore @ag.arizona.edu

Abstract.—This paper provides the first description of a Hawaiian isopod of
the genus Synidotea, S. oahu n. sp. This species is most similar to S. laevi-
dorsalis (Miers, 1881) and S. harfordi Benedict, 1897. A list of Synidotea
species described to date with biogeographic information, and a list of all
marine isopods described from the Hawaiian Islands, are provided.

This paper provides the first description
of a Synidotea species from the Hawaiian
Islands. The isopod genus Synidotea Har-
ger, 1878 currently contains 57 species, in-
cluding the species herein described (see
Table 1). The following characters define
this genus: penes fused forming penial
plate, fifth oostegites absent, and sexually
dimorphic mouthparts (Poore 2001). In ad-
dition, Synidotea species possess the fol-
lowing combination of characters: antennae
2 flagellum multiarticulate, maxillipedal
palp triarticulate, pleon with one partial su-
ture, pereonites 2—4 coxal plates not visible
in dorsal aspect and (unlike most other val-
viferan genera) pereonites 5—7 tergite-coxal
plate sutures can be either present or absent.

The Californian species of Synidotea
were reviewed by Menzies & Miller (1972),
who also included a biogeographic account
of the genus that, at the time, contained 36
species. The phylogeny and biogeography
of the 22 idoteid genera, including Syni-
dotea, were discussed by Brusca (1984).
Poore (2001) redefined and inferred the
phylogeny of the families within the Val-
vifera.

Most Synidotea species occur in the Arc-
tic and in boreal waters (39 of the 57 de-
scribed species); 13 species have been de-
scribed from tropical/subtropical waters. To
date, only one other Synidotea species has
been described from the islands of the trop-

ical Pacific, S. pacifica Nobili, 1906 from
the Tuamotu Islands. Synidotea oahu n. sp.
is one of only 29 marine isopods known
from the Hawaiian islands (see Table 2).
The only other known Hawaiian valviferan
is Colidotea edmondsoni Miller, 1940.
Nine species in this genus belong to the
Synidotea hirtipes species-group (Monod
1931, Menzies & Miller 1972): S. hirtipes
(H. Milne Edwards, 1840), S. laevidorsalis
(Miers, 1881), S. laticauda Benedict, 1897,
S. harfordi Benedict, 1897, S. marplatensis
Giambiagi, 1922, S. brunnea Pires & Mor-
eira, 1975, S. keablei Poore & Lew Ton,
1993, S. grisea Poore & Lew Ton, 1993,
and §. oahu n. sp. Members of the S. hir-
tipes species-group share the following dis-
tinguishing characters: pereon smooth,
frontal margin of head entire or slightly ex-
cavate, and posterior border of pleotelson
with median excavation. Because S. oahu
n. sp. possesses these characters I herein
consider it a member of this group. Species
boundaries within the S. hirtipes group have
been disputed in the literature. Chapman &
Carlton (1991, 1994) argued that S. laevi-
dorsalis is a widespread species, which has
been widely introduced to many coastlines
from Japan by the shipping industry. Chap-
man & Carlton (1991, 1994) have thus sug-
gested the synonymy of seven of the nine
species within this group. However, their
taxonomic justification for the synonymies

VOLUME 117, NUMBER 1

Fig. 1.

Holotype, dorsal view.

was weak, based entirely on an analysis of
length-width ratios of various body parts of
the dorsal aspect of these species. Poore
(1996) refuted the synonyms; through care-
ful comparison of the pleotelson, penial
plate and pereopod 1, he clearly demon-
strated that the populations descibed from
various Indo-Pacific coastlines represent
valid and separate species. He also noted
that the species boundaries are further sup-

ported by different ecological distributions
of the species in this group. This case un-
derscores the importance of detailed, accu-
rate taxonomy in the pursuit of successfully
identifying translocated species. Taxono-
mists are accustomed to the challenging
task of recognizing species boundaries
within groups that contain many similar
species; oftentimes differences between
species, although solid and obvious once
made explicit, are not apparent to the un-
trained eye.

Order Isopoda Latreille, 1817
Suborder Valvifera Sars, 1882
Family Idoteidae Samouelle, 1819
Genus Synidotea Harger, 1878
Synidotea oahu, new species
Figs. 1-6

Type material examined.—Holotype,
ovigerous female, USNM 1009176. Ha-
wail: Oahu Is., 0.8 km from town of Kailua,
collected from small batches of seaweed by
Ray Greenfield, August 20, 1950. Paratype,
female, USNM 99384. Hawaii: Oahu Is.,
Ewa Beach, 32 km from Honolulu, collect-
ed from seaweed by Ray Greenfield, Au-
gust 1, 1954.

Etymology.—The specific epithet oahu
derives from the poetic vowel-rich Hawai-
ian language, providing this binomen, Syn-
idotea oahu, with every vowel in the En-
glish alphabet. In Hawaiian, oahu means
“the gathering place.’ Oahu is also the
name of the second largest island in the Ha-
waiian archipelago and the type locality of
this species. This word is used here as a
noun in apposition.

Diagnosis.—Cephalon dorsal surface
with a weak, transverse depression in front
of eyes. Pereonites 1—7 with mesial, broad-
ly rounded grooves on dorsal surface. Max-
illa 1 mesial lobe with two unique, stout,
distally-serrate robust setae with mesial set-
ules. Mandibles (both right and left) with
four-toothed incisors and four-toothed laci-
nia mobili with additional large serrate
spine-like process. Ratio of head width to

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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VOLUME 117, NUMBER 1

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B
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Fig. 2. Paratype. A, lateral view; B, dorsal view.

pereonite 4 width is 0.69. Pleotelson (fused
pleonites and telson) 1.26 times longer than
wide (measured along lateral margin, from
posterior margin of coxa of pereonite 7 to
distal-most tip of telson).
Description.—Body length: ovigerous
female holotype, 8 mm, non-ovigerous fe-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

male paratype, 7.5 mm. Body yellowish tan
in alcohol.

Cephalon dorsal surface with a weak,
transverse depression in front of eyes. Fron-
tal margin straight. Eyes bulge outward,
forming part of contour of lateral margin of
head. Ratio of head width to pereonite 4

VOLUME 117, NUMBER 1 81

Fig. 3. Holotype. A, right antennna 1; B, left maxilla 2 close-up of inner lobe; C, left maxilla 2: D, head,
ventral view; E, head, lateral view; E left mandible, dorsal view: G, left mandible, mesial view: H, left mandible,
ventral view, I, right antenna 2.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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“SpUPISs] ULIIEMLH{ OY) SUIPUNOLINS s1o}eM dy} UI 1N990 0} poytoder spodosr sue [fe JO ISI] Y—Z 2IGe.L

VOLUME 117, NUMBER 1

Table 2.—Continued.

Recorded distribution

Species and author

Suborder
family

Hawaiian Islands, Samoa, Japan, Christmas Is., Andaman

Paralimnoria andrewsi (Calman, 1910)

Limnoriidae

Is., Aldabra Atoll, Caribbean

Hawaiian Islands

Cymodocella hawaiiensis Bruce, 1994
Neonaesa rugosa Harrison & Holdich, 1982

Sphaeromatidae

Hawaiian Islands, Society Islands, New Guinea,

Queensland

VALVIFERA

Hawaiian Islands

Idoteidae

Colidotea edmondsoni Miller, 1940

Synidotea oahu n. sp.

Hawaiian Islands

width is 0.69. Antenna 1 with triarticulate
peduncle and uniarticulate flagellum with
six pairs of jointed aesthetascs. Antennae 2
extended to third pereonite; with five-artic-
ulate peduncle, article 5 at least twice as
long as any other peduncular article; fla-
gellum with 15—17 articles, terminal two ar-
ticles very small.

Maxilliped with a triarticulate maxilli-
pedal palp, single coupling hook on the left
maxilliped only (holotype). The paratype
has one coupling hook on both left and
right maxilliped. Maxilla | mesial lobe with
two stout distally-serrate robust setae with
mesial setules; lateral lobe with ten serrate
robust setae and many simple setae along
lateral and mesial margins. Maxilla 2 with
plumose, simple and comb setae as figured.
Mandibles with four-toothed incisors and
large molar processes with short spines sur-
rounding margins. Lacinia mobilis of left
and right mandible four-toothed with an ad-
ditional large serrate spine-like process.

Pereonites 1-7 with mesial, broadly
rounded grooves on dorsal surface, other-
wise dorsal surface and lateral margins
smooth, without rugae, tubercules, or
scales. Pereonite 3 widest. Lateral margins
of pereonite 1-3 evenly convex, 4-7
straighter but not sharply angulate. Pereo-
pods setose. Pereopod 1 with dactyl as long
as propodus; two stout setae arise from base
of unguis; distal lateral surface of propodus
covered with serrate setae. Pereopods 2—7
with setation patterns as figured.

Pleotelson 1.26 times longer than wide;
dorsal surface evenly convex; posterior bor-
der with median excavation. Pleopods 1 and
2 with plumose marginal setae on endopods
and exopods, both rami without sutures.
Pleopods 1—3 with coupling setae on mesial
margin of peduncle. Pleopods 3—5 with plu-
mose marginal setae on exopods only; the
number of setae decrease from pleopods 3
to pleopods 5; exopods with partial sutures
on lateral margins, Uropod with an oblique
ridge and 3 plumose setae at mesial junc-
tion of protopod and exopod. Uropod exo-
pod length to width ratio is 0.96.

84 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4.

Discussion.—Synidotea oahu males are
unknown. This species superficially resem-
bles other members of the S. hirtipes spe-
cies-group, particularly S. laevidorsalis and
the widely distributed species S. harfordi.
These three species are possibly closely re-
lated, however, a phylogenetic analysis of
this large genus is needed to test this hy-
pothesis.

Synidotea oahu differs from S. harfordi
and S. laevidorsalis most strikingly in its
smaller body size (S. oahu, 7.5—8.0 mm; S.
laevidorsalis, 12.3-—35 mm, S. harfordi, 18
mm). Menzies & Miller (1972) noted that
Synidotea species follow a general trend of
increasing body size with increasing lati-
tude. Wallerstein & Brusca (1982) showed
the same trend for all intertidal idoteids oc-
curring in the northeast Pacific. Species
within Synidotea range in length from the
3 mm tropical Pacific S. pacifica, to the 32
mm S. bicuspida and 35 mm S. laevidor-
salis from Arctic and boreal waters. Syni-
dotea oahu fits this pattern, with a body size

Holotype. A, left maxilliped; B, left maxilla 1.

of 7.5-8 mm, the average body size for
tropical Synidotea (Menzies & Miller
IQ7/2)).

S. oahu also differs from other members
of the S. hirtipes species-group in the fol-
lowing characters: S. oahu has unique stout
distally-serrate robust setae with mesial set-
ules on the mesial lobe of maxilla 1 and a
four-toothed mandibular incisor, whereas S.
harfordi and S. laticauda both have a two-
toothed mandibular incisor. Synidotea oahu
also differs from S$. harfordi in its broadly
rounded median dorsal impressed lines on
pereonites 2—4, whereas in S. harfordi these
lines are distinctly triangulate. Also, the
dactyl of pereopod 1 in S. oahu is nearly as
long as the propodus, whereas in S. harfordi
it is much longer than the propodus.

Acknowledgments

I am grateful to Marilyn Schotte for loan-
ing specimens from the USNM collections.
I also thank R. C. Brusca, G. C. B. Poore,

VOLUME 117, NUMBER 1 85

Fig. 5. Holotype. A, right pereopod 5; B, right pereopod 7; C, right pereopod 6; D, right pereopod 1; E,
right pereopod 1, close-up of propodus and dactyl; EK right pereopod 2; G, right pereopod 3; H, right pereo-
pod 4.

86 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Holotype, A, right pleopod 5; B, right pleopod 4; C, right pleopod 3; D, right uropod; E, right
pleopod 1; E right pleopod 2.

VOLUME 117, NUMBER 1

B. Kensley, M. Schotte and 2 anonymous
reviewers for commenting on the manu-
script. Chip Griffin did the illustrations. Bri-
an Kensley, Marilyn Schotte and Steve
Schilling’s World List of Marine, Freshwater
and Terrestrial Crustacea Isopoda website fa-
cilitated the compilation of both tables.

Literature Cited

Benedict, J. E. 1897. A revision of the genus Synido-
tea.—Proceedings of the Academy of Sciences
of Philadelphia 1897:387-404.

Brusca, R. C. 1984. Phylogeny, evolution and bioge-
ography of the marine isopod Subfamily Ido-
teinae (Crustacea: Isopoda: Idoteniae).—Trans-
actions of the San Diego Society of Natural His-
tory 20(7):99-134.

Chapman, J. W., & J. T. Carlton. 1991. A test of cri-
teria for introduced species; the global invasion
by the isopod Synidotea laevidorsalis (Miers,
1881).— Journal of Crustacean Biology 11:386-

400.
& . 1994. Predicted discoveries of the
introduced isopod Synidotea laevidorsalis

(Miers, 1881).—Journal of Crustacean Biology
14:700-7 14.

Giambiagi, D. 1922. Cuatro nuevos isopodos de la Ar-
gentina.—Physis Buenos Aires 5(20):230-244.

Harger, O. 1878. Descriptions of new genera and spe-
cies of Isopoda, from New England and adja-
cent regions.—American Journal of Science
15(3):373-379.

Latreille, P. A. 1817. Les Crustaces, les Arachnides, et
les Insectes. In G.L.C.ED. Cuvier Le Regne
Animal, distribue d’ apres son organisation, pour
servrir de base a l’histoire naturelle des ani-
maux et d’introduction a l’anatomie comparee.
Volume 3 Paris.

Menzies, R. J., & M. A. Miller. 1972. Systematics and
zoogeography of the genus Synidotea (Crusta-
cea: Isopoda) with an account of Californian

species—Smithsonian Contributions to Zoology
102:1-33.

Miers, E. J. 1881. Revision of the Idoteidae, a family
of sessile-eyed Crustacea.—Journal of the Lin-
nean Society of London 16:1-88.

Miller, M. A. 1940. The isopod Crustacea of the Ha-
waiian Islands (Chelifera and Valvifera)—Oc-
casional Papers of the Bernice P. Bishop Mu-
seum, Honolulu 15(26):295-361.

Milne Edwards, H. 1840. Histoire Naturelle des Crus-
taces, comprenant Il’anatomie, la physiologie et
la classification de ces animaux. Paris: Roret.

Monod, T. 1931. Tanaidaces et Isopodes subantarctique
de la collection Kohl-Larsen du Senckenberg
Museum.—Senckenbergiana 13(1):10-30.

Nobili, G. 1906. Diagnoses preliminaires de Crustaces,
Decapodes et Isopodes nouveaux recueillis par
M. le Dr. G. Seurat aux iles Tuamotou.—Bul-
letin du Museum National d’ Histoire Naturelle,
Paris 12:256-270.

Pires, A. M. S., & P. S. Moreira. 1975. Two new spe-
cies of Synidotea (Crustacea, Isopoda, Valvi-
fera) from Brazil.—Boletim do Instituto Ocean-
ografico 24:45-67.

Poore, G. C. B. 1996. Species differentiation in Syni-
dotea (Isopoda: Idoteidae) and recognition of
introduced marine species: a reply to Chapman
and Carlton.—Journal of Crustacean Biology
16:384-394.

. 2001. Isopoda Valvifera: diagnoses and rela-

tionships of the families—Journal of Crusta-

cean Biology 21:213-238.

, & H. M. Lew Ton. 1993. Idoteidae of Aus-
tralia and New Zealand (Crustacea: Isopoda:
Valvifera).—Invertebrate Taxonomy 7:197-278.

Samouelle, G. 1819. The entomologists’ useful com-
pendium, or an introduction to the knowledge
of British Insects. London: Boys. 496 pp.

Sars, G. O. 1882. Oversigt af Norges Crustacea.—For-
handlinger 1 Videnskaps-selskabet 1 Christiania
18:1-124.

Wallerstein, B. R., & R. C. Brusca. 1982. Fish preda-
tion: a preliminary study of its role in the zoo-
geography and evolution of shallow water ido-
teid isopods (Crustacea: Isopoda: Idoteidae).—
Journal of Biogeography 9:135-150.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(1):88—94. 2004.

A new species of Synidotea (Crustacea: Isopoda: Valvifera) from the
northern Gulf of Mexico

Marilyn Schotte and Richard Heard

(MS) Department of Systematic Biology, National Museum of Natural History, Smithsonian
Institution, Washington, D.C. 20013-7012, U.S.A.;
(RH) Department of Coastal Sciences, Gulf Coast Laboratory, PO. Box 7000, Ocean Springs,
Mississippi 39566, U.S.A.

Abstract.—Synidotea fosteri, n. sp., the sixth known member of the genus
Synidotea from the western Atlantic Ocean, is described from shallow waters
(1—2 m) adjacent to open beaches in the northern Gulf of Mexico. Its current
range extends from western Florida westward to Texas. The new species is
distinguished from other related species by small size, fairly straight lateral
margins of first pereonite, having the posterior margin of pleotelson straight to
very slightly emarginate and by details of the appendix masculina. A key to
the known western Atlantic species of the genus Synidotea is also given.

Introduction

The presence of an undescribed species
belonging to the valviferan genus Synidotea
has been known from the Gulf of Mexico
for over 20 years. Although there are only
two published records listed as “Synidotea
sp.’ and “Synidotea sp. A’ from Texas and
Florida, respectively (Clark & Robertson
1982, Rakocinski et al. 1996), it has also
been observed in beach habitats at Grand
Island, Louisiana and Gulf Shores, Ala-
bama (R. Heard, pers. obs.). More recent
collections of Synidotea made near Panama
City, Florida, have made possible the de-
termination of a new species, which is the
subject of this report. In the most recent dis-
cussion of the 56 nominal world species of
Synidotea, Poore (1996) lists the relevant
characters used to differentiate several spe-
cies in this genus which closely resemble
S. laevidorsalis (Miers, 1881), as does the
present new species.

Family Idoteidae Samouelle, 1819
Genus Synidotea Harger, 1878

Synidotea Harger, 1878:374; Richardson,
1905:376; Rafi and Laubitz, 1990:2672;
Poore and Lew Ton, 1993:261—262.

Diagnosis.—Body about twice as long as
wide, integument sometimes setose or with
sculpturing; cephalon narrower than per-
eonite 1; body width greatest at pereonite
4. Pleon lacking articulating pleonites,
pleonite 1 indicated by single, small ventro-
lateral suture; apex acute, rounded or ex-
cavate. Antenna 2 multiarticulate. Mandible
with secondary tooth on lacinia mobilis.
Maxillipedal palp, with articles 2 and 3
fused, 4 and 5 also fused. Coxae 2—4 with-
out dorsal coxal plates; coxae 5—7 with ex-
panded dorsal plates. Penes fused complete-
ly and swollen distally, attached to posterior
margin of pleonite 1. Oostegites forming
brood-pouch on pereonites 1—4.

Key to the Species of Synidotea from the
Western Atlantic Region

la. Pleotelson tapers to narrowly rounded,
produced apex S. nodulosa
lb. Pleotelon faintly to deeply emarginate
at apex, not produced
2a. Cephalon bearing two convexities sep-
arated by narrow groove and 2 small,
medial tubercles anteriorly; lateral mar-
gins of pereonites 1—4 angular ....
S. littoralis

VOLUME 117, NUMBER 1

2b. Cephalon smooth, lacking sculpturing;
lateral margins of pereonites not angu-
EVR a6 Tate eset ce SN ere ee ree ere one eee 3
3a. Lateral margins of pleotelson almost
parallel for first 4 of length; angled me-
dially in distal one-third with broad,
shallow emargination on distal margin
3 Sea Betirera peeetin eee sey cee eee ears S. brunnea
3b. Lateral and distal margins of pleotelson
NOLASTADONE Se en hake ee POR op eee 4
4a. Cephalon with deep medial notch; pleo-
telson tapering to very narrow, emar-
GMAW BOOK ocooacocccses S. marmorata
4b. Cephalon without deep medial notch,
faintly emarginate at most; pleotelson
not tapering to narrow apex
5a. Antennal flagellum with 13—20 articles;
body length of mature male ca. 12.5
mm; appendix masculina of male not
extending beyond apex of endopod of
IODOG! D> socssaaacacac S. marplatensis
5b. Antennal flagellum with 7—8 articles;
body length of mature male ca. 4.2 mm;
appendix masculina extending beyond
apex of endopod of pleopod 2
S. fosteri n.sp.

Synidotea fosteri, n. sp.
Figs. 1, 2

Synidotea sp. Clark & Robertson, 1982:46
(key), 49-50 (Table & Text), 57 (Fig. 5);
Rakocinski, et al. 1996:351 (Table).

Material examined.—Holotype male
USNM 1022910, TL 6.5 mm, from sea
grass clumps (origin unknown) in surf/
swash zone, “Bid-a-wee’’ Beach, Panama
City) Beachy sElonday 30712727 N-
085°52.5’'W, sal. 34 ppt., coll. R. Heard and
J. Foster, 23 Nov 1996; Allotype female
USNM 1022911, TL 6.0 mm, same data.
Paratypes: 7 males, 6 ovig. females, 53 fe-
males, 1 juv., USNM 1022912, same local-
ity data.

Other material: | male, 7 ovig. females,
2 females, 2 juvs., open gulf off Santa Rosa
Beach, northwest Florida, 1—1.5 m, coarse
sand with detritus, Sargassum and algae,
coll. R. Heard, 15 July 1991.

Description.—Male: body length 2.7

times greatest width (at pereonite 3) with
minutely spinulose integument (Figs. 3C,
D) and faint dorsolateral sculpturing on all
tergites. Cephalon with faint curved ante-
rior groove above slight dome and faint lat-
eral grooves, transverse posterior groove
deeper; anterior margin of cephalon
straight, sometimes with minute medial
emargination. Width of head to width of
pereonite 4 ratio 0.79. Eyes prominent. Lat-
eral angles of first pereonite nearly straight;
lateral angles of pereonites 2—4 convex and
5-7 nearly straight, making continuous
margin. Lunette on pereonites 2—4 with
broadly rounded posterior margin. Coxal
plates not discernible dorsally. Sutures sep-
arating tergites from coxae faintly visible
on tergites 2—4. Pleotelson length to width
ratio 1.29; length of pleotelson 0.32 times
body length, lateral margins tapering slight-
ly to broad apex with straight or very
slightly emarginate posterior margin. Uro-
podal peduncle with single, oblique ridge;
length to width ratio of exopod 0.94, with
curve between lateral margin and truncate
apex.

Antenna 1 with 4 articles, terminal article
bearing aesthetascs; antenna 2, single distal
plumose seta on 5th peduncle article; fla-
gellum with 8 articles. Mouthparts typical
of genus, with secondary tooth noted on la-
cinia mobilis of mandible.

Pereopod 1, palmate propodus bearing
many pectinate spine-like setae; end of dac-
tyl reaching carpus-merus suture. Pereo-
pods 2—4 similar with posterior margins of
propodi, carpi, and meri bearing several
long and short simple setae. Pereopod 7
with spine-like setae, some pectinate on an-
terodistal margins of propodus, carpus and
merus. Pereopods lacking dense pads of se-
tae.

Pleopod 1, peduncle with 5 coupling
hooks, 23 and 25 plumose marginal setae
on endopod and exopod, respectively. Ple-
opod 2, appendix masculina parallel-sided
through 90% of its length, tapering to near-
ly acute apex, curving laterad, extending
slightly beyond apex of endopod; endopod

90 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. A, male habitus; B, female habitus; C, antenna; D, antennule; E, lateral aspect of male; EK mght
mandible; G, left mandible; H, maxilliped; I, second maxilla; J, first maxilla. Scale = 1 mm.

bearing 9 plumose marginal setae, exopod
bearing 22. Pleopods 3—5 with partial suture
on exterior margin of exopod; exopods
bearing few setae, endopods none. Fused
penial plate weakly waisted, widening

somewhat distally with apex evenly, broad-
ly rounded.

Ovigerous female.—As in male except
for sexual characters and length/width pro-
portions. Length of body 2.3 times width.

VOLUME 117, NUMBER 1 91

Fig. 2. A, ventral pleon; B, pereopod 1; C, pereopod 2; D, pereopod 4; E, pereopod 7; E uropod; G, penial
papilla; H, pleopod 1; I, pleopod 2 of male; J, apex of appendix masculina.

Length of pleotelson 0.29 times body Etymology—The species is named for
length. Length to width ratio of pleotelson Mr. John M. Foster of Gulf Coast Labora-

de 32. tory, who collected the new species in the
Color.—Specimens in preservation a company of the second author.
light red-brown color, pigmentation subtly Remarks.—The Synidotea species S. hir-

reticulated overall. tipes H. Milne Edwards, 1940, S. brunnea

92 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3.

Pires & Moreira, 1975, S. marplatensis
Giambiagi, 1922, S. laticauda Benedict,
1897, S. harfordi Benedict, 1897, S. laevi-
dorsalis Miers, 1881, S. keablei Poore and
Lew Ton 1993, and S. fosteri n. sp. resem-
ble each other closely. Based on morpho-
logical differences, Poore, 1996 concluded
that S. hirtipes, S. laticauda and S. laevi-
dorsalis, all from Indo-Pacific coasts, are
valid and separate species, not synonyms of
the earliest described member of the group
(S. laevidorsalis), and do not represent a
global invasion thereof, as suggested by
Chapman and Carlton, 1991. Poore and
Lew Ton, 1993 described S. keablei from
Australia, which also superficially resem-
bles S. laevidorsalis. But consistently dif-
ferent character states again allowed these
authors to call into question the conclusion
of Chapman and Carlton and their resulting
synonymies.

Of the western Atlantic species, S. fosteri

Scanning Electron Micrographs: A, dorsal view of cephalon, pereonites | and 2; B, lateral margins
of pereonites 1—3; C, integument of dorsal pereon; D, close-up of integument.

most resembles S. marplatensis and S.
brunnea, neither of which were available
for direct observation. S$. marplatensis and
S. fosteri can be separated by the number
of articles in the antennal flagellum (7—8 in
S. fosteri, 13-20 in S. marplatensis); rela-
tive length of the appendix masculina (ex-
tending beyond apex of pleopodal endopod
in S. fosteri, shorter than the apex in S. mar-
platensis); and the larger size of mature
male specimens, e.g., 12.5 mm in the latter
vs. 6.5 mm in the new species. Chief dif-
ferences separating S. brunnea from S. fos-
teri include 13 articles in the antennal fla-
gellum (7-8 in S. fosteri), convex margin
of pereonite | lateral margin (nearly straight
in the new species) and distinct difference
in shape of the pleotelson. In S. brunnea
these lateral margins are nearly straight then
angled medially in the distal third, joined
by a broad but shallowly emarginate apex
on the distal margin. In S. fosteri the pleo-

VOLUME 117, NUMBER 1

Table 1.—Comparison of two additional Synidotea species from North America, as addendum to Poore, 1996.

Data from our own observations.

Maximum length of ovigerous fe-
male

Maximum length of adult male

Color in alcohol

Pleotelson length : width in males
(number of specimens)
Pereon margin

Frontal margin of cephalon; dorsal
sculpture

Head width: pereonite 4 width

Pereopod 1 of male

Setation of ischium-propodus of
pereopods of female

Setation of ischium-propodus of
pereopods of male

Fused penial plate

Uropodal peduncle
Uropodal exopod: length/width

S. fosteri

6.0 mm

6.5 mm
red-brown, reticulated

1.29 (6)

pereonite 1 nearly straight; 2 and
3 convex, 4—7 straight

straight; weak depression in front
of eyes

0.79

palm of propodus concave; dactyl
reaching carpus-merus suture

long and short setae along lower
margins

weakly waisted; length: width
1.85; broadly rounded apically

1 oblique ridge

curve between lateral margin lat-
eral and truncate apex; 0.94

S. harfordi

17.0 mm

blotchy yellow-brown; darker me-
dial stripe on pereon

1.21 (1)

pereonite 1 with subtle curved an-
gle; 2—7 making continuous
line

straight; obvious depression in
front of eyes

0.62

palm of propodus concave; dactyl
reaching carpus-merus suture

dense pads of short setae

not waisted; length: width 2.14;
rounded apically

no oblique ridge

curve between lateral margin and
truncate apex; 0.88

telson is broadly rounded apically with little
or no emargination. The male of S$. brunnea
is as yet unknown.

S. fosteri can be distinguished from all
others in this group by the combination of
the nearly rectilinear lateral margins of the
first pereonite, which are rounded or convex
in most of the others. It is readily separated
from S. laevidorsalis by the shape of the
fused penial plate, and the longer, narrower
pleon in the latter. Mature males of the new
species measure 4.2 to 6.5 mm in length,
whereas Miers’ type specimens of S. lae-
vidorsalis (also male) are longer than 25
mm. Table 1, patterned after Poore’s 1996
comparison of five Synidotea species Indo-
Pacific coasts, lists the same morphological
data for S. fosteri and S. harfordi to help
distinguish this group of similar animals.

Ecological notes.—Synidotea fosteri was
collected on sand substrata at depths of 1—
2m. All of our records came from sites
adjacent to high energy beaches facing the

open Gulf of Mexico. Specimens collected
and observed during our study occurred be-
tween the beach and first or second seaward
sand bar. The specimens were always found
associated with unattached macro-plant de-
tritus or algae. Other peracarids commonly
found associated with S. fosteri included the
amphipods Micropotopus raneyi Wigely
and Atylus urocarinatus Mc Kinney.

Acknowledgments

We wish to thank John Foster, Sara
LeCroy, and Jerry McClelland for making
material available for study. Our sincere ap-
preciation goes to Scott D. Whittaker, SEM
Lab Manager in the Laboratories of Ana-
lytical Biology, National Museum of Nat-
ural History for technical assistance with
the Scanning Electron Micrographs. We
also thank Dr. Brian Kensley of the Nation-
al Museum of Natural History and two
anonymous reviewers for helpful comments
on the manuscript.

94 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Literature Cited

Benedict, J. E. 1897. A revision of the genus Synido-
tea.—Proceedings of the Academy of Sciences
of Philadelphia 1897:387—404.

Chapman, J. W., & J. T. Carlton. 1991. A test of cri-
teria for introduced species: the global invasion
by the isopod Synidotea laevidorsalis.—Journal
of Crustacean Biology 11(3):386—400.

Clark, S. T., & R. B. Robertson. 1982. Shallow water
marine isopods of Texas.—Contributions in
Marine Science 25:45—59.

Giambiagi, D. 1922. Cuatro nuevos isopodos de la Ar-
gentina.—Physis 5:230—244.

Harger, O. 1878. Descriptions of new genera and spe-
cies of Isopoda, from New England and adja-
cent regions.—American Journal of Sciences
and Arts (series 3) 15:373—-379.

Miers, E. J. 1881. Revision of the Idoteidae, a family
of sessile-eyed Crustacea.—Journal of the Lin-
nean Society 16(89):1—87.

Milne Edwards, H. 1840. Histoire Naturelle des Crus-
tacés, comprenant |’anatomie, la physiologie et
la classification de ces animaux, vol. 3. Paris.

Pires, A. M. S., & P. S. Moreira. 1975. Two new spe-
cies of Synidotea (Crustacea, Isopoda, Valvi-

fera) from Brazil.—Boletim Instituto Oceano-
grafico da Universidade de Sao Paulo 24:45—
67.

Poore, G. C. B. 1996. Species differentiation in Syni-
dotea ({sopoda: Idoteidae) and recognition of
introduced marine species: a reply to Chapman
and Carlton.—Journal of Crustacean Biology
16:384—394.

Poore, G. C. B., & H. Lew Ton. 1993. Idoteidae of
Australia and New Zealand (Crustacea: Isopo-
da: Valvifera).—Invertebrate Taxonomy 7:197—
278.

Rafi, EF, & D. R. Laubitz. 1990. The Idoteidae (Crus-
tacea, Isopoda, Valvifera) of the shallow waters
of the northeastern Pacific Ocean.—Canadian
Journal of Zoology 68:2649—2689.

Rakocinski, C., R. W. Heard, S. E. LeCroy, J. A.
McClelland, & T. Simmons. 1996. Responses
by macrobenthic assemblages to extensive
beach restoration at Perdidi Key, Florida,
U.S.A.—Journal of Coastal Research 12:326—
35/2).

Richardson, H. 1905. A monograph on the isopods of
North America.—Bulletin of the United States
Museum 54:1—727.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(1):95—-105. 2004.

A new genus of the Clausidiidae (Copepoda: Poecilostomatoida)
associated with a polychaete from Korea, with discussion of the

taxonomic status of Hersiliodes Canu, 1888

Ju-shey Ho and Il-Hoi Kim

(JSH) Department of Biological Sciences, California State University, Long Beach, California

90840-3702, U.S.A., e-mail: jsho@csulb.edu;

(IHK) Department of Biology, Kangreung National University, Kangreung, Kangwondo, 210-702,

Korea, e-mail: ihkim@kangnung.ac.kr

Abstract.—A new genus and new species of the Clausidiidae (Copepoda:
Poecilostomatoida), Hemadona clavicrura, is described based on the specimens
obtained from the washings of the polychaete, Dasybranchus caudatus Grube,
collected from Namhae-do Island in Korea. The new genus is characteristic in
having (1) the 3rd segment of the antenna drawn out to form a sharp claw, (2)
a 3-segmented maxilliped in female, and (3) an armature formula of IJ-4 on
the third endopodal segment of leg 1. Phylogenetic analysis on the genera of
the Clausidiidae shows that Hersiliodes can not be relegated to a synonym of
Hemicyclops as proposed in the recent past. It is a sister-taxon with the new
genus, Hemadoma, and separated from Hemicyclops in having a 6-segmented
antennule, an armature formula of IJ,4 on the distal segment of the endopod
of leg 1, and a medial protrusion on the proximal segment of the male max-
illiped. Interestingly, the phylogenetic analysis shows also that the three genera
(Conchyliurus, Leptinogaster, and Pholadicola) living in bivalve mollusks are

monophyletic.

Poecilostome copepods of the family
Clausidiidae are known to live largely in
symbiosis with various marine inverte-
brates, such as alcyonarians, polychaetes,
mollusks, and callianassid crustaceans. Cur-
rently, the family comprises nearly 80 spe-
cies in 9 genera. One of its genera, Hersi-
liodes Canu, 1888, living in association
with polychaetes and bivalves, has been
considered almost impossible to separate
from Hemicyclops Boeck, 1872 by Bocquet
et al. (1963) and Vervoort & Ramirez
(1966). Furthermore, Gooding (1963) as
well as Humes & Huys (1992) had even
advocated the doubtfulness of keeping the
genus Hersiliodes as a valid taxon in the
Clausidiidae. Nevertheless, in his book on
the copepods associated with the marine in-
vertebrates of the British Isles, Gotto (1993)
treated Hersiliodes as a valid genus of the

Clausidiidae and, furthermore, in their re-
port on a new species of Hersiliodes from
Korea, Kim & Stock (1996) alleged that the
genus differs from Hemicyclops in bearing
a 6-segmented (instead of 7-segmented) an-
tennule and an armature formula of II,4 Gn-
stead of I,5) on the third endopodal segment
of leg 1. However, it should be pointed out
that the former character state is also found
in one (out of 38) species of Hemicyclops
and the latter, in all nine species of Con-
chyliurus.

Recently, one of us (IHK) discovered,
during his general survey of the symbiotic
copepods on Namhae-do Island in Korea
Strait, a new genus and species of clausidiid
associated with a polychaete. The new form
carries, interestingly, some characteristic
features of both Conchyliurus and Hersili-
odes. Thus, in this paper, in addition to de-

96 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

scribing this new clausidiid, a phylogenetic
analysis of the ten genera of the Clausidi-
idae will be conducted to investigate the
taxonomic status of the genus Hersiliodes.

Materials and Methods

The polychaetes, Dasybranchus caudatus
Grube (Capitellidae), were dug out from the
mud flat and were placed in a plastic bag
and fixed with 70% ethanol. Back at the
laboratory, water was added into the bag
containing the worm fixed in alcohol and
then shaken hard to dislodge the copepods.
The water together with the sediment and
debris were examined under a dissection
microscope for associated copepods. The
copepods were removed and preserved in
70% ethanol. In studying the preserved co-
pepods, the specimens were cleared in lac-
tic acid, dissected on a wooden slide (Hu-
mes & Gooding 1964), and examined under
a compound microscope. All drawings were
made with the aid of a camera lucida. For
formula of armature, ““A’’ represents aes-
thete; Roman numeral, spine; and Arabic
numeral, seta.

Seven genera were recognized by Humes
& Huys (1996) in the family Clausidiidae.
They are Hemicyclops Boeck, 1873; Clau-
sidium Kossmann, 1874; Hippomolgus G.
O. Sars, 1917; Leptinogaster Pelseneer,
1929; Conchyliurus Bocquet & Stock,
1957; Doviella Rocha, 1986; and Hyphal-
ion Humes, 1987. However, several chang-
es have been made since then; two new
genera (Foliomolgus Kim and Pholadicola
Ho & Wardle) were added, respectively, by
Kim (2001) and Ho & Wardle (1992), Her-
siliodes Canu, 1888 was suggested to be
resurrected by Kim & Stock (1996), and
Doviella was relegated to a synonym of a
clausiid genus by Ho & Kim (in press).
Thus, including a new genus to be de-
scribed below, there are now 10 genera in
the Clausidiidae to be considered.

The data used in the character analysis to
prepare for construction of a matrix were
taken from the type species of each of the

10 clausidiid genera. Since Ho’s (1992)
phylogenetic analysis of the Poecilostoma-
toida shows that Erebonasteridae Humes,
1987 occurs in sister-taxa relationship with
a monophyletic clade comprising Clausidi-
idae + (Oncaeidae + Paralubbockiidae),
Erebonasteridae was accordingly employed
as an outgroup to polarize the 14 characters
selected and also to root the cladogram(s)
in reconstruction of the phylogeny. Al-
though Centobnaster Huys & Boxshall,
1990 is generally considered the most prim-
itive erebonasterid copepod (Huys &
Boxshall 1990), some features in Tychidion
Humes, 1973 were found to be even more
primitive. Therefore, both Centobnaster
and Tychidion were used as outgroup in the
polarization of the selected characters
shown in Appendix A. Also, in coding mul-
tistate characters, when a transformation se-
ries containing a single basal bifurcation
(dichotomous transformation) was encoun-
tered, the method of “‘internal rooting” pro-
posed by O’Grady & Deets (1987) was em-
ployed. In this case, as shown in Characters
3, 4 and 8 in Appendix B, the coding of
“0” indicates apomorphy, not plesiomor-
phy.

The computer program HENNIGS86 Ver-
sion 1.5 (Farris 1988) was employed to an-
alyze the phylogenetic relationships among
the genera of the Clausidiidae. The com-
mand “‘ie*’’ (implicit enumeration) was
used to produce multiple, shortest trees
through performance of exhaustive search
and use all available tree space to find all
shortest trees. In order to avoid predeter-
mination of the topology of the resultant
cladogram(s), all multistate characters were
changed to nonadditive (unordered) before
employing the command to reconstruct the
phylogeny.

Description
Order Poecilostomatoida Thorell, 1859
Family Clausidiidae Embleton, 1901
Hemadona, new genus
Diagnosis.—Body elongate, 9-segment-
ed in female and 10-segmented in male.

VOLUME 117, NUMBER 1

First pediger fused to cephalosome. Anten-
nule short, 6-segmented, with 2nd and 3rd
segments incompletely separated. Antenna
4-segmented, with 3rd segment (middle
segment of endopod) drawn out into a large
claw, distal segment tipped with 7 elements.
Labrum well-developed. Mandible tipped
with 2 large spiniform elements and 2 setae.
Paragnath a lobe with spinules. Maxillule
bilobate distally, both lobes tipped with se-
tae. Maxilla 2-segmented, with armature
formula of 2, 4. Maxilliped 3-segmented in
female and 4-segmented in male; proximal
segment in male with medial outgrowth.
Legs 1—4 biramous with 3-segmented rami;
armature formulae generally as in Hemicy-
clops, except 3rd segment of leg 1 endopod
with II,4 and 3rd segment of leg 4 exopod
with III,I,5. Leg 5 2-segmented, armature
formula as in Hemicyclops. Basal segment
of leg 5 in male fused to pediger. Leg 6 in
male a single seta on genital operculum.
Caudal ramus with usual 6 elements. Egg
sac elongate, multiseriate.

Etymology.—TYhe generic name Hema-
dona is an anagram of the island Namhae-
do located in the Korean Strait from where
the new genus was discovered. Gender fem-
inine.

Type species.—Hemadona clavicrura
new species.

Hemadona clavicrura, new species
Figs. 1-3

Material examined.—3 22 and 7 dd
collected from washings of Dasybranchus
caudatus Grube collected from intertidal
mud flat on Nambhae-do Island (34°49'N
128°03’E) in Korea Strait on 22 July 2001.
Holotype 2 (USNM 1013731), allotype ¢
(USNM 1013732), and 6 paratypes (USNM
1013733, including 1 2 and 5 6 <) are de-
posited in the U.S. National Museum of
Natural History in Washington, D.C. Dis-
sected paratypes (1 2 and | ¢G) are kept in
the author’s ([HK) collection.

Female.—Body (Fig. 1A) elongate, 6.34
mm long (excluding setae on caudal rami).

Cephalothorax semicircular and containing
Ist pediger. Second pediger widest of body,
1.04 mm; width of 3rd and 4th pedigers de-
creasing only slightly from that of 2nd pe-
diger. Urosome 5-segmented, 2.37 times
longer than prosome. Genital double somite
longer than wide, 877 X 693 wm, with ali-
form dorsolateral protrusion in anterior half
of somite covering area of egg sac attach-
ment (Fig. 1A). Abdomen 3-segmented,
with all segments longer than wide, 833 X
553 pm, 798 X 508 wm, and 880 X 430
yum. Caudal ramus (Fig. 1C) 4.44 times lon-
ger than wide (720 X 162 wm), armed with
1 short, outer seta at about midlength of
lateral margin, | short, medial, subterminal
seta, and 2 short and 2 long terminal setae;
longest terminal seta (830 wm) 1.15 times
as long as ramus. Egg sac greatly elongated
(7.05 mm), longer than body and cylindri-
cal.

Rostrum subquadrate in dorsal view, pro-
duced forward, and well demarcated from
cephalothorax (Fig. 1A). Antennule (Fig.
1B) short and robust, 6-segmented; formula
of armature: 5, 16, 10, 4, 2 + A, and 7 +
A. Antenna (Fig. 1C) 4-segmented; first
segment (coxobasis) longer than wide, with
long outer-distal seta; second segment (Ist
endopodal segment) shorter than proximal
segment, with small subterminal seta; third
segment (2nd endopodal segments) drawn
out into a large uncinate claw, with basal
patch of spinules on outer surface and 2 un-
equal setae plus | blunt tip, bent, spiniform
seta bearing terminal row of spinules on
medial margin; terminal segment 2.75 times
longer than wide, tipped with 3 unequal se-
tae and 4 spiniform setae structured as that
one on 3rd segment. Labrum (Fig. 1D)
well-developed, with submarginal, inner,
central process, and 2 disjunct, marginal
rows of spinules on either side of this pro-
cess. Gnathobase of mandible (Fig. 1E)
armed terminally with 1 stout, pinnate ele-
ment, | stout, spinulose element, and | pin-
nate and | naked setae. Paragnath (Fig. 1F)
an obtuse lobe fringed with spinules on dis-
tal margin. Maxillule (Fig. 1G) bilobate,

98 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Hemadona clavicrura, new genus, new species, female. A, habitus, dorsal; B, antennule; C, antenna;
D, labrum; E, mandible; E paragnath; G, maxillule; H, maxilla. Scale bars: A, 1 mm; B, C, 0.02 mm; D, E 0.02
mm; E, G, H, 0.05 mm.

VOLUME 117, NUMBER 1

small outer lobe tipped with | long and 2
short setae and larger inner lobe with 5 un-
equal setae. Maxilla (Fig. 1H) 2-segmented;
robust proximal segment (syncoxa) armed
with | large spiniform and 1 small pinnate
setae; distal segment (allobasis) tipped with
2 spiniform elements bearing spinules on
one side and 2 pinnate setae. Maxilliped
(Fig. 2A) 3-segmented; proximal segment
(syncoxa) with 2 unequal medial setae;
middle segment (basis) greatly expanded
laterally and carrying 2 unequal medial se-
tae; terminal segment (endopod) tiny, bear-
ing | spiniform and 2 setiform elements.

Legs 1—4 (Figs. 2B—D, 3A) biramous,
with 3-segmented rami. Formula of spines
and setae as follows:

Coxa Basis Exopod Endopod

Leg 1 O-1 1-1 I-0; 1-1; O-1; 0-1;
I,1,4 11,4

Leg 2 O-I 1-0 [-0; I-1; O—1; 0-2;
II,15 I1,1,3

Leg 3 O-I 1-0 I-0; I-1; O-1; 0-2;
III,15 IL,0,2

Leg 4 O-1 1-0 1J1-0; I-1; O-1; O-2;
I,1,5 ILU,1

Outer surface of all segments on rami
fringed with spinules. Outer spines on all
legs club-shaped, with swollen tip covered
with fine denticles. Leg 5 (Fig. 3B) 2-seg-
mented; proximal segment small, carrying
simple, outer seta; distal segment elongate,
about 4 times longer than wide (750 X 187
zm), armed with 3 club-like spines and 1
thin, simple seta.

Male.—Body (Fig. 3C) elongate as in fe-
male, 3.40 mm long (excluding setae on
caudal rami). Cephalothorax semi-ellipsoid
shaped and containing Ist pediger. Second
pediger widest of body, 532 wm wide;
width of 3rd and 4th pedigers decreasing
only slightly from that of 2nd pediger. Uro-
some 6-segmented, 1.51 times longer than
prosome. Ventrally, proximal segment of
leg 5 indistinctly separated from its pediger
(Fig. 3D). Genital somite slightly longer
than wide, 310 X 300 um; genital opercu-
lum (Fig. 3D) small. Abdomen 4-segment-

ed, with following measurements (proceed-
ing from anterior to posterior): 295 x 282
pm, 366 X 275 pm, 317 X 254 pm, and
423 X 246m. Caudal ramus 4.08 times
longer than wide (408 Xx 100 wm) and
armed as in female. Maxilliped (Fig. 3E) 4-
segmented; proximal segment (syncoxa)
with large, medial protrusion tipped with 3
sharp tines; second segment (basis) largest,
armed with small patch of subterminal den-
ticles on lateral surface, a seta in distome-
dial corner followed by a row of spinules
on medial margin; third segment (lst en-
dopodal segment) smallest and naked; distal
segment (2nd endopodal segment) drawn
out into a long claw with accessory tine and
2 simple setae on medial surface of basal
region.

Etymology.—The species name is a com-
bination of Latin, clava (= a club) and crus
or cruris (= leg), alluding to the club-
shaped outer and terminal spines on all five
pairs of legs.

Remarks.—The general appearance of
Hemadona clavicrura resembles the species
of Conchyliurus in having an elongated
(non-cyclopiform) body. They are further
alike in having a 6-segmented antennule, an
armature formula of II,4 on the terminal
segment of the endopod of leg 1, and a
prominent medial, basal protuberance on
the proximal segment (syncoxa) of the male
maxilliped. These four features are also
shared with Hersiliodes. However, H. clay-
icrura cannot be placed in Conchyliurus
due to the presence of the following char-
acter states: (1) the hook on the 3rd seg-
ment of the antenna is completely fused to
its segment proper, (2) the gnathobase of
the mandible carries four (instead of three)
terminal elements, (3) the proximal seg-
ment (syncoxa) of the maxilla bearing two
(instead of none) elements at outer-distal
corner, and (4) the maxilliped in female is
3-segmented (instead of 2-segmented).
Moreover, 10 species of Conchyliurus are
known and, unlike H. clavicrura living in
association with polychaetes, they were all

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

100

LORD

ere >
VIE
£ SS IWS

SRS

QS Ja
=

SN if

»
IANS

ZEKXYWNN x

Uy Za WN SS KX

Ye NOs
= \ YY \
= MOO

IOS
MON
a

SSS
SY QS S
SH. ;

4 —

55S

<2
ZEISS

SOS

SEB SSS NNN

OES OSS

g 2; D, leg 3.

Hemadona clavicrura, new genus, new species, female. A, maxilliped; B, leg 1; C, le

Fig. 2.
Scale bars: A, 0.05 mm; B—D, 0.2 mm.

101

VOLUME 117, NUMBER 1

Weta

ae.

SS.
OO
SKE
SS ss
ers

KS
GE:

SSS

&
<<

SRR

SSS
Z
SRE

SSS

SSX

SoS
oss

LE
x
Ss

NY,

Hemadona clavicrura, new genus, new species. Female: A, leg 4; B, leg 5. Male: C, habitus, dorsal:
D, first three somites of urosome, ventral; E, maxilliped. Scale bars: A, B, D, 0.2 mm; C, 0.5 mm: E, 0.05 mm.

Fig. 3.

102

Tree 1

f— Foliomolgus
j=1 64 f= Clausidium
L144 Hemicyclops
f— Hemadona
| lk-124— Hersiliodes
| f= Hippomolgus
Tease f= Hyphalion
f— Conchyliurus
L134 j= Leptinogaster
114— Pholadicola

Tree 9

f— Hyphalion

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Tree 7

{(— Hippomolgus

f=184| (= Conchyliurus
| 16 [= Leptinogaster
194| L141 Pholadicola

| f= Foliomolgus

L174] [= Hemicyclops
1 54| f= Clausidium
134 f= Hyphalion
124) f= Hemadona

414— Hersiliodes

(= Pholadicola

pel2i Leptinogaster

| f=144_ Conchyliurus
| j=-16 Hippomolgus

L154

7 f= Foliomolgus
f— Clausidium

L134 Hemicyclops

Fig. 4.

[= Hemadona

L314— Hersiliodes

Clausidiid phylogeny produced through analysis of nonadditive (unordered) coding. Showing three

representatives from three patterns of phylograms. (Other 15 phylograms are available from JSH upon request.)

reported from the mantle cavities of the bi-
valve mollusks.

Of the four differences mentioned above
between H. clavicrura and Conchyliurus,
only items (1) and (4) also apply to the dis-
tinction between it and Hersiliodes. So far
two species of Hersiliodes are known from
either a polychaete (Bocquet et al. 1963) or
a bivalve (Kim & Stock 1996). Thus, it
seems Hemadona is closer to Hersiliodes
than to Conchyliurus.

In general, H. clavicrura is most char-
acteristic in having an unusually long uro-
some (2.37 times longer than its prosome)
and club-shaped outer and/or terminal
spines on all five pairs of legs.

Phylogenetic Analysis

A total of 18 equally parsimonious trees
(cladograms, phylograms) were obtained
with a length of 37 steps, a consistency in-
dex (CI) of 64 and a retention index of 62.

A close comparison of these 18 trees shows
that there are three patterns of tree accord-
ing to the grouping of the 10 genera. In
Pattern I, as Tree 1 in Fig. 4, the 10 genera
are separated into two clades, with one
clade (Clade 16) containing Clausidium,
Foliomolgus, Hemadona, Hemicyclops, and
Hersiliodes and the other clade (Clade 17),
Conchyliurus, Hippomolgus, Hyphalion,
Leptinogaster, and Pholadicola. There are
10 phylograms belonging to this catego-
ry—Tree 1, 2, 3, 4, 5, 6, 11, 12, 13 and 14
(authors’ enumeration; unpublished data).
Phylograms in Pattern I, as Tree 9 in Fig.
4, have Hyphalion set aside on a clade of
its own and the remaining nine genera di-
vided into two groups, with Clausidium,
Foliomolgus, Hemadona, Hemicyclops, and
Hersiliodes in one clade (Clade 15) and
Conchyliurus, Hippomolgus, Leptinogaster,
and Pholadicola in the other clade (Clade
16). There are six phylograms belonging to

VOLUME 117, NUMBER 1

this category—Tree 8, 9, 10, 16, 17 and 18
(authors’ enumeration; unpublished data).
One of the two remaining phylograms, Tree
7, belonging to Pattern III, is shown in Fig.
4. It has the 10 clausidiid genera divided
into two groups, with one comprising Con-
chyliurus, Hippomolgus, Leptinogaster, and
Pholadicola and another one, Clausidium,
Foliomolgus, Hemadona, Hemicyclops, Hy-
phalion and Hersiliodes.

The difference among the three patterns
mentioned above is chiefly due to the in-
consistent positions of Hyphalion. In Pat-
tern I (see Tree 1 in Fig. 4), it is a member
of the group comprising Conchyliurus, Hip-
pomolgus, Leptinogaster, and Pholadicola;
in Pattern II (see Tree 9 in Fig. 4) it is by
itself; and in Pattern III (see Tree 7 in Fig.
4) it is a member of the group comprising
Clausidium, Foliomolgus, Hemadona,
Hemicyclops, Hyphalion and Hersiliodes,
which is entirely different from the one that
it is affiliated with in Pattern I.

There are two monophyletic taxa that
maintain identical relationships in all 18
phylograms. They are Hemadona + Her-
siliodes and Conchyliurus + (Pholadicola
+ Leptinogaster). The former two genera
are held together by sharing characters 1
(with 6-segmented antennule), 12 (proximal
segment of male maxilliped with medial
protrusion) and 13 (with an armature for-
mula of II,4 on distal segment of leg 1 en-
dopod), and the latter three genera, by shar-
ing characters 3 (with 2 elements on 3rd
segment of antenna), 5 (mandible tipped
with 3 elements) and 10 (with 3-segmented
maxilliped in female). It is noteworthy that
both Hemadona and Hersiliodes are char-
acteristic in having a 6-segmented anten-
nule (Character 1). They were not placed in
the same group (clade) with Conchyliurus
+ Leptinogaster + Pholadicola on any of
the 18 phylograms. In other words, the two
constant monophyletic taxa are remotely re-
lated. It is interesting to point out that the
latter three genera comprise parasites of bi-
valve mollusks, while species of Pholadi-
cola inhabit in the host’s intestine, and

103

those of Conchyliurus and Leptinogaster
are found in the host’s mantle cavities.

Five of the 18 obtained phylograms con-
tain a clade with trichotomy, three of these
phylograms (Trees 1, 5 and 12) are in Cat-
egory I and the other two (Trees 9 and 17),
in Category II. Hemicyclops appears as one
of the three terminal clades in all five phy-
lograms showing trichotomy. In four of
these five phylograms, 1.e., Trees 1, 5, 12
and 9, the branch embracing Hemadona +
Hersiliodes appears as another terminal
clade, with either Clausidium or Foliomol-
gus as the third terminal clade. In addition,
Trees 2 and 13 in Category I have a topol-
ogy showing Hemicyclops in a sister-taxon
relationship with Hemadona + Hersiliodes.
These six phylograms indicate that both
Hemadona and Hersiliodes are closely af-
filiated with Hemicyclops. However, since
none of the 18 phylogram shows Hersili-
odes in a sister-taxa relationship with Hem-
icyclops, the former, accordingly, cannot be
relegated to a synonym of the latter. Thus,
the present phylogenetic analysis supports
Kim and Stock’s (1996) notion that Hersi-
liodes is a valid genus in the Clausidiidae
and cannot be synonymized with Hemicy-
clops.

Key to the Genera of the Clausidiidae

A key to the genera of the Clausidiidae
was provided by Humes and Huys (1992).
Since only six of the ten genera currently
recognized were dealt with in that key, a
new key is provided below.

1. Formula of armature on terminal seg-
ment of leg 1 endopod I,5........... 2
— Formula of armature on terminal seg-

ment of leg 1 endopod otherwise ..... 5
2. Antenna 3-segmented ........ Ayphalion
— Antenna 4-segmented .............. 3

3. Maxilliped in female 4-segmented ..
airieaeacdescn: «nea eo aan «ces a Hemicyclops
— Maxilliped in female reduced or absent 4
4. Antennule 7-segmented; 3rd segment of
antenna with 4 elements ..... Foliomolgus
— Antennule 6-segmented; 3rd segment of
antenna with 2 elements .... Leptinogaster

104 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

5. Endopods of legs 1—4 with sucking
discs; middle exopodal segment of leg |
without inner seta Clausidium

— No sucking discs on legs; middle exo-
podal segment of leg 1 with inner seta

ATMO SO Oe oon Oe oo hoe OS 6

6. Proximal segment of maxilla armed with
SCtAG! fn, ary 3 oe eR Mi hocle seen entrees 7
— Proximal segment of maxilla unarmed.. 8

7. Maxilliped in female 4-segmented and
well-developed; armature formula for
terminal segment of leg 1 exopod I,I,5

EES ROR: TRAINS oe Hersiliodes

— Maxilliped in female 3-segmented and
reduced; armature formula for terminal
segment of leg] exopod III,I,5 .. Hemadona

8. Armature formula for terminal segment
of female leg 1 endopod II,4 ......
Se Oe LenS otto ate Conchyliurus

— Armature formula for terminal segment
of female leg 1 endopod otherwise.... 9

9. Maxilliped in female rudimentary ...
RNA. ee ENE REAP EN NE ORE 2, Pholadicola

— Maxilliped in female well developed, at
least 3-segmented Hippomolgus

Acknowledgment

Studies on this project were aided by a
grant from the Paramitas Foundation to the
senior author (JSH) and from the Korea
Science and Engineering Foundation (2000-
1-20200-003-3) to the junior author (HK).

Literature Cited

Bocquet, C., J. H. Stock, & G. Kleeton. 1963. Copé-
podes parasites d’invertébrés des cotes de la
Manche. X. Cyclopoides poecilostomes asso-
ciés aux Annélides polychetes dans la région de
Roscoff.—Archives de Zoologie expérimentale
et générale 102 (Notes et Revue 1):20—40.

Farris, J. S. 1988. Hennig86 Reference, Version 1.5.—
Published by the author, Port Jefferson, New
York, 18 pp.

Gooding, R. U. 1963. External morphology and clas-
sification of marine poecilostome copepods be-
longing to the families Clausidiidae, Clausiidae,
Nereicolidae, Eunicicolidae, Synaptiphilidae,

Catiniidae, Anomopsyllidae, and Echiurophili-
dae.—Unpublished Ph.D. thesis, University of
Washington, Seattle, 276 pp.

Gotto, V. 1993. Commensal and parasitic copepods as-
sociated with marine invertebrates (and
whales).—The Linnean Society (London), 264
pp.

Ho, J.-s. 1992. Phylogeny of Poecilostomatoida: a ma-
jor order of symbiotic copepods.—Bulletin of
Plankton Society of Japan, Special Volume Pp.
25-48.

, & I.-H. Kim. (2003). New clausiid copepods

(Poecilostomatoida) associated with poly-

chaetes of Korea, with cladistic analysis of the

family Clausiidae.—Journal of Crustacean Bi-

ology 22:568-581.

, & W. J. Wardle. 1992. Pholadicola intestin-
alis, new genus and species, a clausidiid cope-
pod parasitic in a deep-burrowing clam from
Texas.—Bulletin of Marine Science 51:37—44.

Humes, A. G., & R. U. Gooding. 1964. A method for
studying the external anatomy of copepods.—
Crustaceana 6:238—240.

, & R. Huys. 1992. Copepoda (Poecilostoma-
toida and Siphonostomatoida) from deep-sea
hydrothermal vent areas off British Columbia,
including Amphicrossus altalis, a new species
of Erebonasteridae, with notes on the taxonomic
position of the genus Tychidion Humes.—Ca-
nadian Journal of Zoology 70:1369—1380.

Huys, R., & G. A. Boxshall. 1990. Discovery of Cen-
tobnaster humesi, new genus, new species (Er-
ebonasteridae), the most primitive poecilosto-
matoid copepod known, in New Caledonian
deep waters.—Journal of Crustacean Biology
10:504—519.

Kim, I.-H. 2001. Foliomolgus cucullus, a new genus
and species of Clausidiidae (Crustacea: Copep-
oda: Poecilostomatoida) associated with a poly-
chaete in Korea.—Proceedings of the Biological
Society of Washington 114:660—666.

, & J. H. Stock. 1996. A new species of Clau-
sidiidae (Copepoda, Poecilostomatoida) associ-
ated with the bivalve Ruditapes philippinarum
in Korea. Cahiers de Biologie marine 14:1—6.

O’Grady, R. T., & G. B. Deets. 1987. Coding multi-
state characters, with special reference to the
use of parasites as characters of their hosts.—
Systematic Zoology 36:268—279.

Vervoort, W., & E Ramirez. 1966. Hemicyclops thal-
assius nov. spec. (Copepoda, Cyclopoida) from
Mar del Plata, with revisionary notes on the
family Clausidiidae.—Zodlogische Mededelin-
gen (Leiden) 41:195—220.

VOLUME 117, NUMBER 1 105

Appendix 1.—Characters and character states used in the phylogenetic analysis of the Clausidiidae. Numbers
in parentheses denote the numerical coding of the character states. Coding in Characters 3, 4, and 8 employed
“‘internal rooting’? proposed by O’Grady & Deets (1987). Centobnaster and Tychidion were utilized as the
outgroup in polarization of the character state transformations.

Ol 3rd and 4th segments of antennule separated (0) or fused (1)
02 Aesthetasc on antepenultimate segment of antennule absent (0) or present (1)
03 3rd segment of antenna with 3 elements (1), 4 elements (0) or 2 elements (2)

04 Terminal segment of antenna with 6 elements (1), 7 elements (0), 5 elements (2) or 4 elements (3)
05 Mandible tipped with 4 elements (0) or 3 elements (1)

06 Inner lobe of maxillule carrying 2 elements (0) or 3 elements (1)

07 Outer lobe of maxillule carrying 3 elements (0) or 4 elements (1)

08 Proximal segment of maxilla without seta (0), with 1 seta (1), 2 setae (2) or 3 setae (3)

09 Distal segment of maxilla with 4 elements (0), 3 elements (1), 2 elements (2) or 1 element (3)
10 Maxilliped in female 4-segmented (0), 3-segmented (1), 2-segmented (2) or absent (3)

11 Proximal segment of female maxilliped with 2 setae (0), 1 seta (1) or none (2)

12 Proximal segment of male maxilliped without protrusion (0) or with medial protrusion (1)
13 Distal segment of endopod on leg 1 with an armature formula of I, 5 (O) or Il, 4 (1)
14 Distal segment of exopod on leg 4 with an armature formula of II, I, 5 (O) or II, I, 5 (1)

Appendix 2.—Data matrix of 14 characters and their states in ten genera of Clausidiidae as used in the
cladistic (phylogenetic) analysis. The question mark “*?’’ indicates an unknown state. Due to the application of
“internal rooting’’ (O'Grady & Deets, 1987) those characters coded with “‘1°’ in the outgroup are treated as
plesiomorphic and *‘O”’ in the ingroup, apomorphic.

Characters

Genus (OTU) 1 5 10 14
Outgroup 0) 0) 1 1 0 0) 0) 1 0 0) 0 0) 0) 0)
Clausidium 0 0 0 0 1 1 1 3 0 0) 0) 0 ? 0
Conchyliurus 1 1 2 0) 1 1 1 0) 0 2 1 ] 1 1
Foliomolgus 0) 1 0) (0) 0) 1 1 1 1 3 ey 0) 0) 0)
Hemadona ! 0 0) 0) 0) 1 1 2 0 I 0) 1 1 1
Hemicyclops 0 1 0) 0 0) 1 1 2 0) 0) 0) 0) 0) 0)
Hersiliodes 1 1 0) 0) 0) 1 1 D 0 0) 0) 1 1 0)
Hippomolgus 1 1 i 0 0) i 1 0) 0) 0) 1 ? 0) 0)
Hyphalion 1 0 2 v 0) ? ? 1 0 2 2 0) O 0)
Leptinogaster 1 1 2 2 1 0 0) 0 D 3 ? 0) 0 0)
Pholadicola 1 0) D 3 1 0 0 0 3 3 ? 0 ? ?

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(1):106—113. 2004.

Vesicomyicola trifurcatus, a new genus and species of commensal
polychaete (Annelida: Polychaeta: Nautiliniellidae) found in deep-sea

clams from the Blake Ridge cold seep

Jennifer Dreyer, Tomoyuki Miura, and Cindy Lee Van Dover

(JD, CLVD) The College of William and Mary, Department of Biology, Millington Hall,
Williamsburg, Virginia 23187, U.S.A., email: jcdrey@wm.edu; clvand@wm.edu

(ITM) Department of Biological Production and Environmental Science, Faculty of Agriculture,

Miyazaki University, 1-1 Gakuen-Kibanadai-Nishi, Miyazaki 889-2192, Japan, email:
miura@cc.miyazaki-u.ac.jp

Abstract.—A new genus and species of deep-sea polychaete belonging to
the family Nautiliniellidae is described from the Blake Ridge cold seep off the
coast of South Carolina at a depth of 2155 m. This species is commensal within
the mantle cavity of ~60% of the vesicomyid clams collected at the seep site.
Vesicomyicola trifurcatus is distinguished from previously described nautili-
niellid genera and species by the presence of two pairs of tentacular cirri and
up to seven trifurcate hooked chaetae on the posterior parapodia. The new
species resembles [heyomytilidicola tridentatus in having trifurcate hooks, but
the arrangement and number of chaetae differs. Only two types of chaetae are
present in V. trifurcatus: four to seven stout, simple hooks anteriorly to mid-
body, and up to seven trifurcate hooks posteriorly. In contrast, there are three
types of chaetae in /. tridentatus: up to five stout hooks per parapodium, each
with a minute projection on cutting edge of the main fang, 10—20 simple,
slender tridenate chaetae, and numerous minute mucronate chaetae. A key to
species of Nautiliniellidae is included.

The Nautiliniellidae is a small group of
deep-sea polychaetes that live in the mantle
cavity of a clam or mussel host. Nautili-
niellids have been collected from chemo-
synthetically based deep-sea habitats, in-
cluding cold seeps and hydrothermal vents.
Since nautiliniellids were first reported by
Miura & Laubier (1989), 10 genera and 14
species have been described (Table 1). Two
undescribed species have also been report-
ed, one from a cold seep at Barbados
Trench (4960 m; Olu et al. 1996) and one
off the Pacific coast of Mexico (3221 m;
Olu, pers. comm.).

An additional genus, Santelma, has been
assigned to the family Nautiliniellidae
(Blake 1993, Glasby 1993), but its affilia-
tion with the Nautiliniellidae remains ques-
tionable. The only known species, Santelma

miraseta (Fauchald, 1972), was first placed
in the family Pilargidae and the genus Pi-
largis. Blake (1993) redescribed the species
and assigned it to Santelma, a new nautili-
niellid genus, based on chaetal similarities.
Unlike nautiliniellids, S. miraseta has ex-
truded neuroaciculae, a median antenna (or
its trace), and it lacks neuropodial hooks
and parapodial cirri. Based on these fea-
tures, S. miraseta fits better within the orig-
inal family Pilargidae (Salazar- Vallejo,
pers. comm.). We follow the precedent of
Miura & Hashimoto (1996) and exclude S.
miraseta from the Nautiliniellidae.
Nautiliniellids have reduced and simpli-
fied body structures that are associated with
a commensal or parasitic life. These modi-
fications include a less developed anterior
region, the presence of only simple hooked

VOLUME 117, NUMBER 1

Table 1—Family Nautiliniellidae: list of genera and species, host bivalve genus and family, collection depth of type specimen, location where type specimen was

collected and author reference.

Reference

Location

Depth
(m)

Host bivalve Genus
(Family)

Genus and species

Blake (1993)

Florida Escarpment
Okinawa Trough

3303

Unknown

1 Flascarpia alvinae

Miura & Hashimoto (1996)

Blake (1993)
Blake (1993)

1395
3243

Bathymodiolus (Mytilidae)

(Mytilidae)
Unknown

2 Iheyomytilidicola tridentatus
3 Laubierus mucronatus

4 Miura spinosa

Florida Escarpment
Santa Maria Basin
_ Okinawa Trough

565
625
701

1114

Miura & Hashimoto (1993)
Miura & Hashimoto (1993)
Miura & Laubier (1990)

near Adula (Mytilidae)

5 Mytilidiphila enseiensis

Okinawa Trough
Sagami Bay

Bathymodiolus (Mytilidae)
Solemya (Solemyidae)

6 Mytilidiphila okinawaensis
7 Natsushima bifurcata

Miura & Hashimoto (1996)
Miura & Laubier (1989)

Blake (1990)

Kagoshima Bay
Japan Trench

98
5650:
3700

Solemya (Solemyidae)

8 Natsushima graciliceps

Calyptogena (Vesicomyidae)

Thyasira (Thyasiridae)

9 Nautiliniella calyptogenicola

10 Petrecca thyasira

Laurentian Fan

Miura & Ohta (1991)

Okinawa Trough
Sagami Bay

1400
1170

625
1160
2155

Calyptogena (Vesicomyidae)
Calyptogena (Vesicomyidae)
Calyptogena (Vesicomyidae)
Conchocele (Thyasiridae)

11 Shinkai longipedata
12 Shinkai sagamiensis
13 Shinkai semilonga

Miura & Laubier (1990)

Miura & Hashimoto (1996)
Miura & Hashimoto (1996)

Present study

Okinawa Trough
Sagami Bay

14 Thyasiridicola branchiatus

Blake Ridge Diapir

Vesicomya (Vesicomyidae)

15 Vesicomyicola trifurcatus

107

chaetae modified to grasp host tissue, and
the absence of anal cirri on the pygidium.
Diagnostic characters of the family include
the number of prostomial appendages, num-
ber of tentacular cirri, and chaetal mor-
phology and number. These characters are
specific to each genus but are useful for
species identifications since seven of the ten
nautiliniellid genera are monospecific.
Based on these morphological characters,
we determined that the specimens collected
from the Blake Ridge cold seep belong to
a new genus and species described herein.

Material and Methods

Biological samples were collected at the
Blake Ridge Diapir site (ODP Site 996;
32°30'N, 76°11'W; 2155 m) on 25 to 28
Sep 2001, using the DSV Alvin. A descrip-
tion of the study site can be found in Van
Dover et al. (2003). Although geological
and chemical properties of this site have
been explored during the past decade, the
Alvin 2001 samples represent the first col-
lections of megafauna and macrofauna from
this area.

Host clams were collected using a suc-
tion sampler. The clams were identified as
a new genus and species in the Family Ves-
icomyidae, based on morphological char-
acters, molecular differences in comparison
to described species, and geographic and
bathymetric location (E. Kryolora, pers.
comm.). Clams were dissected and nautili-
niellids were removed and placed into ei-
ther 10% buffered formalin or 3% glutar-
aldehyde and 0.1 M phosphate buffer with
0.25 M sucrose (pH 7.4). After 24 hours,
formalin-fixed nautiliniellids were rinsed
and stored in 70% ethanol.

Photographs of the external morphology
were taken with a compound light micro-
scope (LM) and a scanning electron micro-
scope (SEM). Specimens for LM were
mounted in glycerol and ethanol and ob-
served with a Zeiss Axioskop 2 binocular
compound microscope. Specimens for SEM
were dehydrated through a graded series of

108 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ethanol, terminating with 100% ethanol.
Samples were then critical-point dried, gold
sputter coated (20 nm thick), and observed
with an Amray SEM 1810. Images were
captured using a Spot camera (Diagnostic
Instruments) or a DPI11 digital camera
(Olympus). Line illustrations were prepared
using a camera lucida attached to a Wild
Heerbrugg compound microscope.

Systematics

Family Nautiliniellidae Miura & Laubier,
1989
Vesicomyicola, new genus

Type species.—Vesicomyicola trifurca-
tus, new species, by present designation.

Diagnosis.—Body with strong dorsal
arch, ventrally flattened. Prostomium with
one pair of palps, without eyes. Tentacular
segment fused with prostomium, with dor-
sal and ventral cirri, neuroacicula, and neu-
ropodial hooked chaetae. Parapodia sub-bi-
ramous, with dorsal and ventral cirri. Noto-
and neuropodia each with one embedded
acicula. Chaetae absent on notopodia. Two
types of chaetae present on neuropodia:
simple hooked chaetae on anterior segments
(some with single subapical tooth present
on anterior to mid-body segments), and tri-
curcate hooked chaetae on posterior seg-
ments. Pygidium cylindrical, without anal
cirri.

Gender.—Masculine.

Etymology.—TYhe generic name is de-
rived from the name of the host vesicomyid
clams these polychaetes inhabit.

Vesicomyicola trifurcatus, new species
(Figs. 1—4)

Type material.—Holotype (ODP Site
Woe S2SOMIN, WO We QiSS ma, Bs} Seo
2001, Alvin Dive 3712; USNM 1016220)
and five paratypes (USNM 1016221) from
same dive and date were deposited in the
collections of the National Museum of Nat-
ural History, Smithsonian Institution,
Washington, District of Columbia. An ad-

ditional five paratypes, each from the same
dive and date, were deposited in the Mu-
seum National d’Histoire Naturelle, Paris
(MNHN POLY TYPE 1405) and the Na-
tional Science Museum, Tokyo (NSMT—
Pol P 458).

Additional material.—Voucher speci-
mens were retained in the collection of
CLVD in the Department of Biology at The
College of William and Mary.

Description.—Holotype female, oviger-
ous, measuring 8.4 mm long, 1.3 mm wide,
including parapodia, with 37 segments.
Paratypes ranging from 4.4—12.7 mm long,
0.8—1.6 mm wide, including parapodia, and
with 28—41 segments. Body flattened ven-
trally, arched dorsally. Some live specimens
with green pigment in parapodia, others
with pale pink color; preserved specimens
in alcohol pink to white in color. Some pre-
served females pale green; internal oocytes
evident through transparent parapodial epi-
dermis. Preserved holotye and paratypes
curled (Fig. 1A).

Prostomium rounded, with palps (Fig.
2A-C). Eyes absent. Tentacular segment
fused with prostomium, with one pair of
dorsal and ventral cirri, neuroacicula, and
neuropodial hooked chaetae (Fig. 2C).
Foregut with well-developed muscular re-
gion (Fig. 2A—C). Pygidium rounded, with-
out anal cirm (Fig. 2D).

Parapodia subbiramous, with dorsal and
ventral cirri. Dorsal cirri with inflated base
and tapering tip, twice as long as ventral
cirri. Notopodia with single embedded acic-
ula, lacking chaetae (Fig. 3A). Neuropodia
with a single bent acicula and hooked chae-
tae (Fig. 3B).

Neuropodial hooks of two types. Ante-
rior neuropodia with simple stout hooks
with recurved tips, four to seven on each
parapodium (Fig. 4A, B), some anterior to
mid-body chaetae with single small apical
tooth near tip, appearing slightly bifid (Fig.
4C). Posterior neuropodia with thinner, sim-
ple hooks with trifurcate tips, up to seven
per neuropodium (Fig. 4D, E).

Etymology.—The specific name comes

VOLUME 117, NUMBER 1

Fig. 1.
body.

from fri- = three times, + furcatus =
forked, in reference to the trifurcate chaetae
present on the posterior segments.
Biology.—The mantle cavities of ~60%
of the Blake Ridge clams sampled contained
one to five nautiliniellid polychaetes. Carbon
and nitrogen stable isotope compositions of
worm and clam tissues were consistent with
a parasitic life-style for the worm, but the
sulfur isotope composition of the worms was
so distinct from that of the clams that an
alternative diet must be inferred (Van Dover
et al. 2003). Van Dover et al. (2003) pro-
posed a feeding strategy whereby ciliary ac-
tivity of the clam gills moves sufficient vol-
umes of seawater to allow the polychaetes
to collect and consume suspended organic
particles either from gill mucus or from a
worm-generated mucus net.

Discussion

Vesicomyicola trifurcatus resembles spe-
cies in the genera Nautiliniella, Natsushi-

109

Vesicomyicola trifurcatus new genus, new species. Scanning electron micrograph (SEM) of whole

ma, Shinkai, and Thyasiridicola, based on
shared characters of the tentacular segment,
which in these four genera includes dorsal
and ventral cirri and neurochaetae (with the
exception of the genus Thyasiridicola,
which lacks neurochaetae). The genus Ves-
icomyicola differs from these four genera in
the number and morphology of the neuro-
podial chaetae.

Vesicomyicola trifurcatus resembles
Iheyomytilidicola tridentatus Miura &
Hashimoto, 1996 based on the trifurcate
chaetal morphology, but the arrangement
and number of chaetae on the parapodia dif-
fers. There are only two types of chaetae
present in V. trifurcatus: stout, simple
hooks (four to seven; sometimes bifid) on
the anterior to mid-body parapodia, and tri-
furcate hooks (up to seven) on the posterior
parapodia. In contrast, there are three types
of chaetae in /. tridentatus: stout hooks (up
to five), each with a minute projection on
the cutting edge of the main fang; simple,

110

Fig. 2.
view. B. Drawing of anterior end, dorsal view. C. Drawing of anterior end, ventral view. D. LM of pygidium,
dorsal view.

slender tridentate chaetae (10—20); and nu-
merous minute chaetae with mucronate tips
(Miura & Hashimoto 1996).

Based on its unique set of morphological
characters, we consider V. trifurcatus to be
a new genus and species. A key to nautili-
niellid species is provided to aid in identi-
fication; most species are location and host
specific.

The terminology and interpretation of
prostomial appendages in this family is the
subject of some debate (Blake 1993, Miura
& Hashimoto 1996), suggesting the need
for a re-evaluation and revision of this fam-
ily and its genera once a consistent diag-
nosis of prostomial appendages can be ap-
plied.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

100pm B

SF Vi
O cae e
Fy me

SSS

100pm G

Vesicomyicola trifurcatus new genus, new species. A. Light micrograph (LM) of anterior end, dorsal

Color dimorphism was a distinctive char-
acter of live V. trifurcatus, but on preser-
vation the color variation was lost. Poly-
chaetes with green parapodia in new col-
lections (2003) were all gravid females. In
other nautiliniellid species, color dimor-
phism corresponds to sexual dimorphism
(Miura & Hashimoto 1996, Miura 1998).
We have yet to confirm that the pale colored
specimens are males. With the discovery of
each new species in the Nautiliniellidae, we
learn more about the ecology of these
worms and their relationship with their host
bivalves; we still know little about the in-
ternal anatomy, reproductive biology and
larval characteristics, or the trophic ecology
of this polychaete family.

VOLUME 117, NUMBER 1

Fig. 3.

100pm A

Vesicomyicola trifurcatus new genus, new species. A. Drawing of mid-body parapodium with em-
bedded aciculum and dorsal cirrus; lateral view. B. Drawing of mid-body neuropodium and ventral cirrus; ventral
view.

Key to the species of Nautiliniellidae

la.

1b.

2a.

2b.

3a.

3b.

Aa.

Ab.

Sa.

Sb.

Prostomial appendages (palps or an-
tennae) absent
BECUASE cid dtten ee sei Miura spinosa Blake, 1993
One or two pairs of prostomial ap-

pendages present ................ 2
Tentacular segment with only one pair
GUGTIUO A Se Bian GG Eee Orono olan rets 3
Tentacular segment with one pair of
dorsal and ventral cirri ........... 7
Tentacular segment with or without
neurochaetae; all neuropodial hooks
SIEM SLE a a en eet ae alee tis le eis 4
Tentacular segment without neuro-
chaetae; some neuropodial hooks stout

ote Graken ero ot orem acne ee 5

Neurochaetae 220 (up to 35) per
parapodium; neurochaetae with inflat-
ed, subdistal stems and _ slightly
curved, pointed distal ends ......
Mytilidiphila enseiensis
Miura & Hashimoto, 1993
Neurochaetae =20 per parapodium;
neurochaetae with rounded tips and
slightly curved, distal ends
Mytilidiphila okinawaensis
Miura & Hashimoto, 1993
Only one type of neurochaeta present:
large, stout hooks
Two types of neurochaetae present:

Yon

6a.

6b.

7a.
7b.
8a.

8b.

8c.

111

100pm B

One to two large, stout hooks and 15—

20 small, mucronate tipped chaetae Gin
crows of 2) ..... Laubierus mucronatus
Blake, 1993

Three types of neurochaetae present:

<five stout hooks with minute projec-

tion on cutting fang, 10—20 tridentate

chaetae, and numerous minute, slen-

der chaetae with single mucronate
Gouin cesoac Theyomytilidicola tridentatus
Miura & Hashimoto, 1996

Maximum of one to two stout hooks
per parapodium Petrecca thyasira
Blake, 1990

Maximum of seven to eight stout
hooks per parapodium
Flascarpia alvinae Blake, 1993
One type of neurochaetae present... 8
Two types of neurochaetae present... 9

One large, stout hook per parapodium
Nautiliniella calyptogenicola
Miura & Laubier, 1989

Maximum of four stout hooks per

parapodium, and branchiae-like noto-
podial projections present
.... Thyasiridicola branchiatus Miura &
Hashimoto, 1996

Number of anterior stout hooks vari-

able (2—25) and notopodial branchiae-
like projections absent

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig.

9a.

Ob.

4. Vesicomyicola trifurcatus new genus, new species. A. SEM of simple anterior neuropodial hooks.
B. LM of anterior neuropodial hooks. C. SEM of mid-body chaetae with small apical tooth near tip, appearing
slightly bifid. D, E. SEM of posterior trifurcate chaetae.

Neurochaetae with two to three stout
hooks and numerous bifurcate simple
CHACtAC yea Steuaen rake Cheer Cee 1]
Neurochaetae with four to seven stout
hooks per parapodium present anteri-
orly and five to seven trifurcate hooks
posteriorly ... Vesicomyicola trifurcatus,
new genus and species

10a. Notopodia in middle regions espe-

cially elongate; middle to posterior

10b.

10c.

neuropodia with a single hook with
strongly curved distal fang .... Shinkai
longipedata Miura & Ohta, 1991

Notopodia not elongate in any re-

gions; middle to posterior neuropo-

dia with a single hook, strongly

curved on distal end with knob on tip
Reber 18s AY Shinkai sagamiensis
Miura & Laubier, 1990

Notopodia in middle regions slightly

VOLUME 117, NUMBER 1

elongated; middle to posterior neu-

ropodia with =five, slightly curved
Nooksy FF: een ce Shinkai semilonga
Miura & Hashimoto, 1996

Short, conical notopodia on middle
segments Natsushima bifurcata
Miura & Laubier, 1990

Elongate notopodia on middle seg-
ments Natsushima graciliceps
Miura & Hashimoto, 1996

lla.

11b.

Acknowledgments

We thank Captain Silva, the crew of R/
V Atlantis, Expedition Leader Dudley Fos-
ter, the pilots and technicians of DSV Alvin,
and members of the science party for their
assistance at sea, and Karine Olu and Dan-
iel Desbruyeres for loaning us nautiliniellid
specimens. We are grateful to Joe Scott,
Jewel Thomas and Megan Ward for help
with illustration preparations and layout and
Dr. Norman Fashing for use of his camera
lucida. The manuscript benefited from re-
views of Brigitte Hilbig, Stephen Gardiner
and one anonymous reviewer. This research
was supported by National Oceanic & At-
mospheric Administration’s National Un-
dersea Research Program (University of
North Carolina NC-Wilmington National
Undersea Research Center) and Ocean Ex-
ploration Program. The Carol Woody In-
ternship Program (College of William and
Mary) and the Lerner Gray Memorial Fund
of the American Museum of Natural His-
tory provided support to JD for collabora-
tion with TM in Japan.

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.

. 1993. New genera and species of deep-sea

polychaetes of the Family Nautiliniellidae from

the Gulf of Mexico and the Eastern Pacific.—

Proceedings of the Biological Society of Wash-

ington 106:147—157.

113

Dall, W. H. 1886. Reports on the results of dredging,
under the supervision of Alexander Agassiz, in
the Gulf of Mexico, and in the Caribbean Sea,
1877-79, by the U.S. coast survey steamer
Blake XXIV. Report on the Mollusca. Part 1.
Brachiopoda and Pelecypoda.—Bulletin of the
Museum of Comparative Zoology at Harvard
University 12:171—318.

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.

Glasby, C. J. 1993. Family revision and cladistic anal-
ysis of the Nereidoidea (Polychaeta: Phyllodo-
cida).—Invertebrate Taxonomy 7:1551—1573.

Miura, T. 1998. Annelida Polychaeta (in part). Pp. 70—
75 in D. Desbruyéres & M. Segonzac, eds.
Handbook of deep-sea hydrothermal vent fauna.
French Research Institute for the Exploitation
of the Sea (IFREMER), France, pp. 279.

, & J. Hashimoto. 1993. Mytilidiphila, a new

genus of nautiliniellid polychaete living in the

mantle cavity of deep-sea mytilid bivalves col-
lected from the Okinawa Trough.—Zoological

Science 10:169—174.

, & . 1996. Nautiliniellid polychaetes

living in the mantle cavity of bivalve molluscs

from cold seeps and hydrothermal vents around

Japan.—Publications from the Seto Marine

Laboratory 37(316):257—274.

, & L. Laubier. 1989. Nautilina calyptogeni-

cola, anew genus and species of parasitic poly-

chaete on a vesicomyid bivalve from the Japan

Trench, representative of a new Family Nautil-

inidae.—Zoological Science 6:387—390.

, & . 1990. Nautiliniellid polychaetes

collected from the Hatsushima cold-seep Site in

Sagami Bay, with descriptions of the new gen-

era 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
8:83-387.

Olu, K., M. Sibuet, E Harmegnies, J.-P. Foucher, & A.
Fiala-Medioni. 1996. Spatial distribution of di-
verse cold-seep communities living on various
diapatric structures of the southern Barbados
prism.—Progress in Oceanography 38:347-376.

Van Dover, C. L. et al. 2003. Blake Ridge methane
seeps: characterization of a soft-sediment, che-
mosynthetically based ecosystem.—Deep-Sea
Research I 50:281—300.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(1):114—139. 2004.

Studies on western Atlantic Octocorallia (Coelenterata: Anthozoa).
Part 4: The genus Paracalyptrophora Kinoshita, 1908

Stephen D. Cairns and Frederick M. Bayer

Department of Systematic Biology (Invertebrate Zoology), National Museum of Natural History,
Smithsonian Institution, PO. Box 37012, Washington, D.C. 20013-7012, U.S.A.,
e-mail: cairns.stephen@nmnh.si.edu

Abstract.—Previously undocumented from the western Atlantic, three new
species of Paracalyptrophora are described from this region. In order to fa-
cilitate comparisons, all six species in the genus are diagnosed, illustrated,
included in a dichotomous key, and compared in a table of distinguishing
characteristics. P. kerberti is herein designated the type species of Paracalyp-
trophora. Additional specimens are reported of all six species. Paracalyptro-
phora is now know to occur in the central and South Pacific and both sides of
the North Atlantic at depths of 150—1480 m.

Kinoshita (1908:58), in his report on
Primnoidae from Japanese waters, recog-
nized the sharp distinction between species
of Calyptrophora having the large sclerites
of the body of the polyp inseparably fused
to form solid rings, as in the type species
C. japonica Gray, and those in which the
large sclerites encircling the body of the
polyp remain separable and unfused. For
the latter he established the subgenus Par-
acalyptrophora including Calyptrophora
kerberti Versluys, C. mariae Versluys, and
C. josephinae (Lindstrém). This subgenus
was not recognized by subsequent authors
until it was elevated to generic status in
keys but without further description (Bayer
1981:937; Bayer & Stefani 1989:455).

Dredging and trawling in the western At-
lantic by several research vessels, including
the USFC steamer Albatross and R/V Ger-
da, obtained many specimens referable to
three species of Paracalyptrophora, which
are described herein.

Material and Methods

Most of the specimens reported in this
paper were collected by the R/V Gerda, a
vessel operated by the University of Miami,
the specimens later deposited at the USNM.

Other specimens were collected by the: Al-
batross, Oregon, Silver Bay, Chalcal II
(MNHN), and Atlantis (MCZ).

Designation of polyp scales follows the
terminology used by Versluys (1906) as
amplified by Bayer et al. (1983). Synony-
mies are purported to be complete. The
SEM photomicrographs were taken by the
authors on a variety of instruments in the
SEM Lab at the NMNH.

The following abbreviations are used:
Alb—USFWS Albatross; G—R/V_ Gerda;
H:W—height to maximum width of an
opercular scale; [L—inner-lateral opercular
scale; JSL-I—Johnson Sea-Link-I, MCZ—
Museum of Comparative Zoology, Harvard,
Cambridge; MNHN—Muséum national
d’Histoire naturelle, Paris; MOM—Musée
Océanographique, Monaco; NMNH—Na-
tional Museum of Natural History, Smith-
sonian, Washington, D.C.; O—R/V
Oregon; SB—R/V Silver Bay; OL—outer-
lateral opercular scale; SEM—Scanning
Electron Microscope stub number (unpre-
faced number in Bayer series, Cairns series
prefaced with a C); USNM—uUnited States
National Museum (now the NMNH);
ZMA—Zoologisch Museum, Amsterdam;
ZMB—Zoologisches Museum, Berlin.

VOLUME 117, NUMBER 1

Subclass Octocorallia
Order Alcyonacea
Suborder Calcaxonia
Family Primnoidae Gray, 1858
Genus Paracalyptrophora Kinoshita, 1908

Calyptrophora.—Versluys, 1906:104
(part).—Ktikenthal, 1919:468 (part);
1924:317 (part).—Aurivillius, 1931:301
(part)—Deichmann, 1936:171 (part).—
Bayer, 1956: F221 (part)—Tixier-Duri-
vault, 1987:171 (part).

Calyptrophora (Paracalyptrophora) Ki-
noshita, 1908:58.

Paracalyptrophora Bayer, 1981:937,
946.—Bayer & Stefani, 1989:455 (in key
only).—Bayer, 2001:367.

Type species.—Calyptrophora kerberti
Versluys, 1906, here designated.

Diagnosis.—Primnoidae with verticillate
polyps directed downward, enclosed in two
pairs of large abaxial scales (1.e., basal and
buccal) extending around body to form
rings, a pair of smaller infrabasals, and in
one species a variable number of small ad-
axial buccals. The two pairs of large body
wall scales are never inseparably fused,
sometimes not even meeting at adaxial side
of body. When present, sclerites of tentacles
are few and small, but usually absent en-
tirely. Branching dichotomous, in one or
two fans.

Description.—Colonies are dichoto-
mously branched in one plane or in two
parallel fan-shaped planes, with polyps al-
ways arranged in whorls and directed
downward. The polyps are armed with two
pairs of large abaxial scales that nearly or
completely encircle the body. They may be
so firmly wedged together by the complex
sculpture along the abaxial midline that a
few pairs may remain joined through clean-
ing and preparation, but they are not insep-
arably fused abaxially or adaxially to form
solid rings; in many cases the members of
the buccal pair do not even meet adaxially.
One pair of curved infrabasal scales lies be-
tween the large basal scales and the scler-
ites of the coenenchyme. Eight roughly tri-

115

angular scales/plates fold over the retracted
tentacles to form an operculum covering the
retracted tentacles and closing the buccal
aperture. In one species, small adaxial buc-
cal scales may be developed below the ad-
axial opercular scales. The tentacles are ei-
ther without sclerites, or have extremely
small scales in such small numbers as to be
easily overlooked. The axis is stiff, brittle,
heavily calcified, weakly grooved longitu-
dinally, brownish or blackish and some-
times with metallic luster; the holdfast is
calcareous, irregularly discoidal, attached to
solid substrate.

Distribution.—Southwestern Pacific (Ti-
mor Sea, Norfolk Ridge), Japan, Hawaii,
and the North Atlantic; 150—1480 m.

Remarks.—So far as known, the corre-
lation of downward facing polyps and two
pairs of large, unfused body scales is
unique for this genus. Although the mem-
bers of basal and buccal abaxial body scale
pairs are separate and unfused, they some-
times are so tightly interlocked by the com-
plex tubercular sculpture of the margins
that meet along the abaxial suture that they
remain attached even after maceration in
sodium hypochlorite solution. The adaxial
processes of the basal pair may meet but
are not permanently united, and the abaxial
symphysis usually separates during manip-
ulation for mounting.

The following key begins with a deter-
mination of the gross colony form; how-
ever, if only branch fragments are available,
this can be problematic. In that case, the
tabular key (Table 1) can be used to distin-
guish all six species. In fact, the shape, size,
and ornamentation of the buccal scales
alone are probably adequate to distinguish
the six species.

Key to the Six Species of
Paracalyptrophora
(Atlantic species in bold face)

1. Colonies in the shape of a single large
fan; mature colonies over 40 cm in
height

1’. Colonies in the shape of two rounded,

116 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

parallel fans; mature colonies usually
less than 30 cm in height ........... 3
2. Distal margin of buccal scales only
slightly flared, revealing most of the
opercular scales in abaxial view; dorso-
lateral edge of buccal scales ridged ...
Shite ae P. josephinae (Lindstr6m, 1877)
2’. Distal margin of buccal scales strongly
flared, obstructing view of most of the
underlying opercular scales; dorso-lat-
eral margin of buccal scales not ridged
4 AEE {ERE UND Wo: P. simplex, n. sp.
3. Dorso-lateral margin of buccal scales
ridged; coenenchymal scales also ridged

ug uoaiSy 3 Gyios sche Rene UBM Se) ae) eee ae 4
3’. Dorso-lateral margin of buccal scales
granular or smooth, but not ridged; coe-
nenchymal scales granular, but not
THEBES. Foose eee s S eee epee ewe 5

4. Dorso-lateral margin of basal scales not
ridged; dorso-lateral margin of buccal
scales with one low ridge; tentacular
sclerites present P. carinata, n. sp.

4’. Dorso-lateral margin of basal scales
having 3 or 4 prominent ridges; dorso-
lateral margin of buccal scales with 4 or
5 prominent ridges; tentacular scales ab-
SCNtwy ae P. mariae (Versluys, 1906)

5. Polyps small (1.0—1.2 mm in length);
adaxial buccals absent; sclerites uni-
formly granular; tentacular sclerites ab-
sent P. duplex, n. sp.

5’. Polyps large (over 2 mm); 1—5 adaxial
buccals usually present; sclerites
smooth with only slight indication of
granularity; tentacular sclerites present

P. kerberti (Versluys, 1906)

Paracalyptrophora duplex, new species
Figs. 1A—D, 2A—D, 3, 4 A-K

Primnoa regularis Duchassaing & Michel-
otti, 1860:17, pl. 1, figs. 12-13 (see Re-
marks herein).

Material examined/Types.—Straits of
Florida off Cape Canaveral: 28°08'N,
80°04'W, 49 m (depth suspect), O-5191, 14
Jan 1965, one small colony lacking holdfast
and most of main stem, USNM 52755,
paratype.

Northwest of Little Bahama Bank:
27°37.65'N, 78°58.74'W, 404 m, JSL-I-

3572, 10 Aug 1993, one large colony lack-
ing holdfast, USNM 93960, paratype.

North of Little Bahama Bank: 27°29.5'N,
78°37.5'W, 485-496 m, G-252, 5 Feb 1964,
one dry specimen with commensal galath-
eid crab, USNM 52747 (SEM 1744), para-
type.

West of Little Bahama Bank: 27°21’'N,
79°15"W, 439-503 m, SB-440, 29 Dec
1958, 3 damaged colonies, and detached
branches, USNM 51264 (SEM C1045),
paratypes.

Off Southwest Point, Grand Bahama:
26°35'N, 78°25'W, 329-421 m, G-692, 21
Jul 1965, one branch, USNM 52748; one
nearly complete small colony lacking hold-
fast, USNM 52749; one colony, USNM
52752; 3+ broken branches, USNM 52753,
paratypes.

Off Southwest Point, Grand Bahama:
26°29'N, 78°39'W, 247-374 m, G-697, 22
Jul 1965, one nearly complete small colony
with part of holdfast, USNM 52745, para-
type.

Off Southwest Point, Grand Bahama:
26°28'N, 78°37'W, 555-575 m, G-695, 22
Jul 1965, one branch, USNM 52756, para-
type.

Off Southwest Point, Grand Bahama:
26°27'N, 78°43'W, 522—489 m, G-706, 22
Jul 1965, one young colony lacking hold-
fast, USNM 52746 (SEM 1752); 3 more or
less complete colonies and detached
branches, USNM 52754 (SEM 263, 1755,
1756), paratypes.

Off Southwest Point, Grand Bahama:
26°27'N, 78°43'W, 384-403 m, G-533, 4
Mar 1965, one colony, USNM 52751 (SEM
C1046—47), holotype; 2 damaged colonies,
one denuded incomplete axis, and detached
branches, USNM 52750 (SEM 1753, 1754,
C1042); one young colony, USNM 100773,
paratypes.

Off Havana: 23°10'39"N, 82°20'21"W,
389 m, Alb-2350, 20 Jan 1885, one intact
colony and many damaged branches,
USNM 17314, paratype.

South of Great Inagua Island: 20°43'N,
73°29'W, 448 m, O-5416 24 May 1965, one

VOLUME 117, NUMBER 1

F no

: aearcgaestteten a
ees vee lore —
a ee

. ¢ a .
retett eee cenett”

<e
=

Fig. 1. A—D, Paracalyptrophora duplex (A, paratype from G-252, USNM 52747; B, C, holotype of Primnoa
regularis, Turin Coel. 275; D, holotype, USNM 52751): A, base of double fan showing enclosed galatheid crab,
X 0.36; B, upper part of colony with broken branch in place, X 0.25; C, lower part of colony showing calcified
holdfast, x 0.25; D, complete holotype, X 0.31. E, P. simplex, holotype USNM 52767, xX 0.15. E P. josephinae,
Atlantis 23-152, USNM 100788, branch fragment, X 1.0. G, P. carinata, holotype, USNM 49948, complete
colony and holdfast, X 0.33. H, P. mariae, Chalcal I], CP25 (MNHNP), colony, X 0.25. I, P. kerberti, Alb-
5093, USNM 30105, X 1.0.

118 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

colony without holdfast (dry), USNM
1008871, paratype.

Holotype of Primnoa regularis, Guade-
loupe, Museo Regionale di Scienze Natur-
ali, Turin, Coel. 275 (ex. 175), 1 complete
dry colony and several broken branches, all
polyps lost (SEM C1043—44).

Type locality.—26°27'N, 78°43'W (off
Southwest Point, Grand Bahama), 384—
403 m.

Description.—Colonies consist of a ro-
bust, vertical main stem and a pair of par-
allel, dichotomously branching fans. The
stem is anchored in a dense, white, calcar-
eous, semi-hemispherical holdfast, the larg-
est known 32 mm in diameter. Most dam-
aged specimens are broken above the hold-
fast; only four of the specimens reported
herein are complete in this regard. The
main stem is inflexible, 7-10 cm in height,
up to 8 mm in diameter, and usually round
in cross section, although in large speci-
mens the stem is slightly compressed in a
direction perpendicular to the fan. In large
colonies, the main stem constitutes about
35% of the height of the colony. The un-
derlying stem axis is golden or black-
brown, faintly longitudinally striate, and,
when dried, often splits longitudinally to re-
veal a lighter colored central core. The first
bifurcation of the main stem, which results
in two branches, is in the plane of the even-
tual fans; the second series of bifurcations,
which results in four branches, is perpen-
dicular to the fans; and the third and all
remaining bifurcations are in the plane of
the fans. The length of the internode be-
tween the first and second bifurcations is
quite short (e.g., 1.5 mm) and in most col-
onies, except for small ones, this internode
is subsumed into the second internode, such
that it would appear as though the first di-
vision of the main stem is into four robust
branches that are oriented perpendicular to
the fans. All higher order branching is di-
chotomous and equal such that both branch-
es are of the same diameter, neither one
seeming to dominate (and thus not lyri-
form). In some cases, one half of a dichot-

omy remains simple or divides at a much
wider interval than usual so that adjacent
branches do not interfere with one another,
but in general, most of the branching occurs
within 5 cm of the top of the main stem,
resulting in many elongate, unbranched ter-
minal branches up to 11—13 cm in length.
Rarely are there more than 7 nodes leading
to a terminal branch, the most highly divid-
ed branches being those on the margin of
the colony fans. A large colony might have
23-30 terminal branches per fan, or 46—60
terminal branches in the colony. The dis-
tance between adjacent branches of a fan is
usually only 2—4 mm, whereas the distance
between the two parallel fans is 12-15 mm.
A fully developed fan is usually wider than
tall, large fans measuring 21—25 cm across
and 15—16 cm tall, thus occupying the top
*; of the colony height. The holotype is 23.5
cm tall and 14 cm wide, with a main stem
length (broken) of 7.6 cm, but the largest
known specimen (holotype of P. regularis)
is 27 cm tall, 25 cm wide, with a complete
main stem length of 10 cm.

Polyps are arranged in regular whorls
and directed downward. Usually the whorls
are composed of three polyps, but whorls
of 4 or 5 may occur on the proximal part
of the branches; in some colonies, whorls
of 4 or even 5 predominate; 14—20 whorls
occupy 3 cm of axial length, but the vari-
ation in any one colony is not usually so
great. In general, polyps are well spaced, in
that polyps within a whorl do not touch one
another and there is a distance of 0.40—0.45
mm between adjacent whorls. Polyps are
present on the stalk in small colonies, but
in large specimens they are absent from
both stalk and the lower part of major
branches; polyps are also often missing
from the side of the branches that face the
opposite fan. Polyps are 1.0—1.2 mm in
length and 0.65—0.80 mm in width.

Each polyp is protected by two pairs of
large abaxial body scales and a smaller pair
of infrabasals. The infrabasal scales are the
smallest of the body wall sclerites, only
about 0.20 mm in maximum height, cres-

VOLUME 117, NUMBER 1 119

Fig. 2. Paracalyptrophora duplex (A, B, D, paratype from G-706, USNM 52754; C, holotype, USNM
52751): A, abaxial stereo view of polyp; B, lateral stereo view of polyps; C, opercular view of a polyp; D,
lateral view of a polyp with jagged-edged buccal scales. Scale bars 0.5 mm.

120

cent-shaped, and anchor the polyp to the
branch coenenchyme. The basal scales are
much larger, up to 1.1 mm, and project per-
pendicular to the branch. Each basal bears
a serrate, projecting spine at its dorso-lat-
eral margin, the spine variable in shape
ranging from short and broad to tall and
slender, the latter constituting slightly over
half the height of the scale. These spines
usually bear one finely serrate ridge on their
outer side, which is continuous with a ridge
on the dorso-lateral margin of the basal
scale and which extends only about half
way to the base of the basal scale. Tall,
slender basal spines also have three more
ridges separated by 90°, whereas broad bas-
al spines have 8—10 small parallel ridges on
their inner face. The upper, inner face of the
basals has a small ridge that hinges with the
straight proximal margin of the adjacent
buccal scale. The buccal scales are slightly
shorter (0.9-1.0 mm) but much broader
than the basals, and have a free, flared distal
margin (0.15—0.25 mm) that encloses the
opercular scales and obstructs a view of the
operculars from the adaxial side. The pro-
jecting buccal margin, which is translucent
due to a thinning of the scale as well as a
replacement of the inner tubercles with
short spines, may be evenly rounded, pro-
duced as a broad lobe on each side (Figs.
2A—C), or divided into 2 or 3 more or less
acute, flat lobes, the latter condition more
common in young colonies (Fig. 2D). This
character varies to a considerable extent
even in a single specimen. Whereas the ab-
axial margins of the basal scales meet as a
sharp, raised crest along the abaxial suture,
the buccal scales overlap one another at the
abaxial midline, often in one direction
along half the length, and in the opposite
direction along the other half (Fig. 2A). The
dorso-lateral margins of the buccals are
evenly rounded, not ridged. The outer sur-
faces of the body scales and operculars are
uniformly covered with small (8-10 wm di-
ameter), rounded to sharp granules, and
their inner surfaces by crowded, complexly
ornamented tubercles also 8-10 pm in di-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ameter. The opercular scales are triangular
in shape, decreasing in size from the abax-
ials (length = 0.48 mm, H:W 1.55) to the
adaxials (length = 0.29 mm, H:W = 1.1).
As is typical for many primnoids, the ad-
and abaxials are symmetrical in shape,
whereas the outer- and inner-laterals are
asymmetrical, each class of operculars be-
ing more developed on their abaxial margin
and thus having an off-centered keel. All
operculars bear a prominent keel on their
distal, inner surface as well as a field of
crowded tubercles that concentrate on the
central and basal regions. The lateral re-
gions under the opercular scales are bare or
covered with short spines and opercular
margins are usually finely dentate, each
equilateral triangular tooth being about 3 °m
in height (Fig. 4E). The upper surface of
the operculars is covered with smooth gran-
ules like the body wall scales. The tentacles
appear to be devoid of sclerites.
Coenenchymal scales are polygonal to
elongate in shape, ranging from 0.15 to
0.80 mm in length, but on average about
0.4 mm. Those on the main stem occur as
two layers, a lower layer of flattened scler-
ites, and an upper layer of thicker (0.06—
0.10 mm), rotund scales that are fitted in a
closely abutted, mosaic pattern (Fig. 4G—
H). The coenenchymal scales of the branch-
es occur in one layer and are flattened (0.02
mm thick), with slightly overlapping mar-
gins. Both types of coenenchymal scales are
covered exteriorly with small (10-12 wm
diameter) granules, most of which are in-
dependent but occasionally are linked in
short rows that appear to radiate outward
from near the center of the scale, but ridges
are never present. Their inner surfaces are
covered with complex tubercles 8-12 wm
in diameter. Coenenchymal scales also cov-
er the basal holdfast. The black axial back-
ground gives the translucent coenenchymal
scales a milky white color.
Etymology.—Latin duplex = double or
twofold, an allusion to the double fan-shape
of the colonies.
Comparisons.—Paracalyptrophora du-

VOLUME 117, NUMBER 1

Fig. 3.

Distribution of Paracalyptrophora duplex.

plex is compared to P. simplex in the ac-
count of that species and to other conge-
nerics in Table 1.

Distribution.—Straits of Florida from off
Cape Canaveral to Havana; Bahamas
(Grand Bahama Island and south of Ina-
gua); Lesser Antilles (Guadeloupe) (Fig. 3);
374-555 m.

Remarks.—TYhe convex space between
the two parallel fans appears to provide an
ideal niche for galatheid crabs, one of
which in each colony may place its abdo-
men in the region of dense branching at the
top of the main stem, and orient its claws
along the branching orientation of the fans
(Fig. 1A). Coral and crab appear to be the
same color.

Examination of the dry, somewhat dam-
aged holotype (deposited at the Turin Mu-
seum) of Primnoa regularis Duchassaing &
Michelotti, 1860 (Figs. 1B—C), which was

121

designated as the type species of the genus
Narella by Gray (1870), shows it to be con-
specific with P. duplex. Even though this
specimen no longer retains any polyps or
polyp sclerites, the branching of the colony
and the coenenchymal sclerites are perfect-
ly consistent with this species, and thus log-
ically would have nomenclatural priority.
However, following strict nomenclatural
priority would cause widespread confusion
within primnoid taxonomy. For instance,
because P. regularis was chosen as the type
of Narella, the three species heretofore
placed in Paracalyptrophora would now be
placed in the genus Narella, and the 25 spe-
cies heretofore placed in Narella would
have to be transferred to the next available
generic name, i.e., Calypterinus Studer,
1887. Furthermore, the morphological re-
lationship implied by the names Calyptro-
Phora and Paracalyptrophora would be

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

122

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VOLUME 117, NUMBER 1

lost. To avoid this widespread changing of
generic combinations and the confusion that
it would cause, we will suggest to the ICZN
that the type of Primnoa regularis be sup-
pressed and a neotype be designated (ICZN,
1999: article 75.6, conservation of prevail-
ing usage by a neotype), a specimen that is
consistent with the current understanding of
the genus Narella and with the species N.
regularis as described by Cairns & Bayer
(2003).

Paracalyptrophora simplex, new species
Figs. 1E, 4L—-T, 5A-C, 9

Material examined/Types.—North of Lit-
tle Bahama Bank: 27°34.5'N, 78°49'W,
488-516 m, G-254, 6 Feb 1964, broken
branches probably of a single colony,
USNM 52769, paratype.

North of Little Bahama Bank: 27°29.5'N,
78°37.5'W, 485—496 m, G-252, 5 Feb 1964:
one large dry colony (holotype), USNM
52767 (SEM 1757, 1758, C1049-51); 5

branches, USNM 52757; one colony,
USNM 52763; one large dry colony,
USNM 52764; one large dry colony,
USNM 52765; one large dry colony,

USNM 52766; four dry branches, USNM
52768, paratypes.

Off Settlement Point, Grand Bahama:
26°45'N, 79°05'W, 494-530 m, G-1125, 13
Jun 1969, 6 branches and fragments,
USNM 52762 (SEM 265, 267, 1731), para-
types.

Straits of Florida: 26°38’N, 79°02’ W, 516
m, G-1312, 31 Mar 1971, detached branch-
es, USNM 57556, paratypes.

Off Southwest Point, Grand Bahama:
26°31'N, 78°51'W, 366 m, G-503, 4 Feb
1965, 3 dry colonies, USNM 52770 (SEM
1760, 1761, 1771), paratypes.

North of North Bimini, Bahamas:
25°59'N, 79°19'W, 439-458 m, G-633, 30
Jun 1965, 5 broken branches, USNM 52759
(SEM 1721, 1722, 1747), paratypes.

North of North Bimini, Bahamas:
25°56'N, 79°22'W, 402 m, G-798, 12 Sep

128

1966, 2 incomplete colonies, USNM 52760
(SEM C1048), paratypes.

Off Havana, Cuba: 23°09'10"N, 82°23’W,
706 m, Alb-2152, 30 Apr 1884, 2 branches,
USNM 7166, paratypes.

Yucatan Channel: 20°59’N, 86°23’00’W,
305 m, bottom temp. 17.1°C, Alb-2353, 22
Jan 1885, 2 dichotomous branches in poor
condition, USNM 50087, paratypes.

Arrowsmith Bank, Yucatan: 20°57’N,
86°34'W, 165-140 m, G-899, 10 Sep 1967,
one colony, USNM 52761, paratype.

Type locality.—27°29.5'’N, 78°37.5'W
(north of Little Bahama Bank), 485—496 m.

Description.—Colonies consist of a ro-
bust, vertical main stem, which gives rise
to a fan that is uniplanar and consists of
dichotomously branching elements. The
stem is anchored in a white calcareous
holdfast, although only one specimen was
collected with the base intact. The main
stem is inflexible, up to 11 cm in height and
9.6 mm in basal diameter, in large speci-
mens compressed in the branching plane. In
large colonies, the main stem constitutes
about 22% of the height of the colony. The
stem axis is golden or dark brown with a
slightly lighter colored core, and faintly
longitudinally striate. All branching is di-
chotomous and equal, except for the two
outermost branches of the fan of larger col-
onies, which are often twice the diameter of
the inward branching stems as well as being
straight, which confers a lyrate shape to the
colony. Although long end branches up to
12 cm length occur, in general, branching
occurs throughout the fan at intervals of
about every 1.5 cm, sometimes resulting in
terminal branching that have resulted from
15 previous bifurcations. The distance be-
tween adjacent branches is about 4—5 mm.
The fan is roughly the same height as
width. The largest colony (the holotype) is
41 cm tall, with a fan 38 cm in height and
34 cm in width, and a broken main stem
only 3 cm long.

Polyps are arranged in regular whorls
consisting of 4—8 polyps (usually 6); 14—
20 whorls occur in 3 cm of axial length,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

a Scale 1
oe = SCale 2

eee Scale 3
ames Scale 4

Fig. 4. A—K, Paracalyptrophora duplex (A, B, E, FE holotype; C, D, paratype, SB-1440; G, J, K, paratype,
G-533; H, I, holotype of Primnoa regularis, Turin Museum): A, basal scale; B, buccal scale; C, D, main stem
coenenchymal scales in situ; E, distal margin of opercular scale; EK ad-, OL-, IL- and adaxial operculars: G, H,
side views of coenenchymal scales from stalk; I, top of coenenchymal scale from branch; J, K, upper granular
and lower warty sides of coenenchymal scales. L—-T, Paracalyptrophora simplex (L, M, P=T, holotype; N, O,
paratype from G-798): L, basal scale; M, buccal scale; N, O, main stem coenenchymal scales in situ; P, ad-,
OL, IL, and adaxial operculars; Q, infrabasal scale; R, S, lower and upper views of coenenchymal scales;

VOLUME 117, NUMBER 1

and although this range may be present
within a single colony, 17 seems to be the
predominant number. In general, polyps are
closely spaced, i.e., adjacent polyps in a
whorl are usually directly adjacent or even
overlapping, and the distance between ad-
jacent whorls is quite small (0.10—0.25
mm), such that the tip of the buccal scales
of the polyps of one whorl almost touch the
buccal spines of the polyp of an adjacent
whorl. Polyps occur on the main stem of
small colonies. Polyps are 1.3—1.5 mm in
length and 0.8—0.95 mm in width.

Each polyp is protected by two pairs of
large abaxial body scales and a pair of nar-
row, curved infrabasal scales situated be-
tween the coenenchymal sclerites and the
basal pair. The body wall scales are virtu-
ally identical in shape and ornamentation to
those described for P. duplex, differing pri-
marily in size, those of P. simplex being
slightly larger, i.e., the infrabasals are up to
0.33 mm in height, the basals up to 1.15
mm, and the buccals up to 1.05 mm, the
latter with a flared distal margin 0.25 mm
in extent, which, like that of P. duplex, may
be produced as a single broad lobe (Figs.
4M, 5C) or divided into 2—4 acute teeth.
Furthermore, the dorso-lateral margins of
the basals bear only short ridges (Fig. 5B),
whereas the dorso-lateral margins of the
buccals often bear parallel, aligned rows of
surface granules (Fig. 5A). The opercular
scales are also similar in shape but slightly
larger, the abaxial operculars up to 0.56 mm
in length and the adaxials 0.34 mm in
length, but all operculars having slightly
serrate margins and a H:W ratio of 1.4—1.6,
similar to that of P. duplex. The coenen-
chymal sclerites are also quite similar in
size and shape to those of P. duplex; how-
ever, the surface granules are somewhat
larger, up to 18 wm in diameter.

Etymology.—Latin simplex = simple,

<—

125

single, or onefold, an allusion to the colo-
nies in the shape of a single fan.
Comparisons.—The shape of the polyps
of P. simplex is virtually identical with
those of C. duplex, differing primarily in
having slightly larger (20—25%) sclerites
and thus larger polyps. But, even though
the polyps are larger, both species have the
same range of polyps per cm, this because
the distance between polyp whorls of P.
simplex is shorter. In general, the polyps of
P. simplex are more crowded, having more
polyps per whorl as well as having more
closely spaced whorls, these characters
serving to distinguish isolated branches.
Characters at the grosser (colonial) level
that distinguish P. simplex from P. duplex
are that it produces only one fan, it attains
a larger colony size, branching occurs
throughout the fan with as many as 15
nodes, and large colonies tend to have a
lyrate branching pattern (Table 1).
Distribution.—Known only from the in-
sular side of the Straits of Florida from the
Yucatan Channel to north of Littke Bahama
Bank, Bahamas (Fig. 9); 165-706 m.

Paracalyptrophora josephinae
(Lindstrom, 1877)
Figs. 1K 6A—C, 7A—G

Calyptrophora josephinae Lindstrom,
1877:6, pl. 1, figs. 1-3 (Josephine
Bank).—Versluys, 1906:109 (re-exami-
nation of type and Studer’s specimen).—
Kiikenthal, 1919:474 (diagnosis); 1924:
319 (diagnosis and key).—Thomson,
1927:29 (Alice Bank).—Aurivillius,
1931:301, fig. 60, pl. 6, fig. 5 (re-descrip-
tion of type, key to species in genus).—
Deichmann, 1936:172 (remarks).—Gras-
shoff & Zibrowius, 1983:119, pl. 1, figs.
5, 6 (Josephine Bank).—Carpine & Gras-
shoff, 1985:33 (MOM _ deposition).—

T, lower view of coenenchymal scale. Scale bar 1: R = 0.20 mm; 2: E G, I, PR. Q, S = 0.25 mm, H = 0.083
mm; 3: E, J, K, T = 0.05 mm; 4: A-C, L-N = 0.25 mm.

126 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Paracalyptrophora simplex (A, C, holotype; B, paratype from G-633, USNM 52759): A, abaxial
stereo view of a whorl; B, lateral stereo view of a polyp; C, opercular stereo view of a polyp. Scale bars 0.5 mm.

VOLUME 117, NUMBER 1

Grasshoff, 1985a:305 (Talisman speci-
mens from Biskaya); 1985b:73 (Jose-
phine and Great Meteor Banks); 1986:27
(remarks).

Stachyodes Josephinae.—Studer, 1901:41
(Azores).

Calyptrophora (Paracalyptrophora) jose-
phine.—Kinoshita, 1908:58 (taxonomic
reassignment).

Paracalyptrophora josephinae.—Bayer,
1981:938, text-fig. 77 (new comb.).
Grasshoff, 1989:219 (listed).—Bayer,
2001:367 (mentioned).

Material examined.—Atlantis Seamount:
34°0S5'N, 30°15'’W, 293 m, R/V Atlantis
cruise 152, station 23, 26 Aug 1948, 2 di-
chotomous branches (MCZ, in alcohol;
fragment USNM 100788) (SEM 1719).

Fragment of holotype (SEM C1052-S54).

Types.—The holotype is deposited at the
Swedish Museum of Natural History
(1113). Type Locality: Josephine Bank
(36°46'’N, 14°07’W), 201-214 m, station
36n.

Diagnosis.—This species has been de-
scribed three times before, the first being
the detailed original description of Lind-
strom, the second by Versluys (1906), and
the third and by far the most detailed by
Aurivillius (1931), all three based on the
holotype or fragments of it. We have also
examined a small fragment of the holotype
but can add little to the previous descrip-
tions except for what can be illustrated by
SEM. Thus, only a diagnosis for this spe-
cies is presented herein:

Colonies uniplanar, one of the largest
(the holotype) 55 cm in height. Branching
dichotomous, occurring throughout colony
at intervals of 20—35 mm. Stem axis bronze
to golden yellow. Polyp whorls consist of
4—7 polyps (the average being 6); 13-17
(usually 14) whorls occur over 3 cm axial
length; adjacent whorls separated by 0.4—
0.6 mm, depending on branch diameter.
Polyps 1.3—1.5 mm in length (not 1.6 mm,
as stated by Lindstrém) and 0.75—0.90 mm
in width. Infrabasals typically crescent

127

shaped, 0.25—0.30 mm in height. Basals
0.75—0.90 mm in height, the distal 0.17—
0.20 mm (20%) being a short, quite broad,
blunt distal ““spine’’, which on the exposed
interior face is covered with 10—12 parallel,
serrate ridges (Figs. 7A—B). Each basal
scale also bears one prominent ridge on its
exterior dorso-lateral margin. Buccal scales
up to 1.0 mm in length, having very slightly
flared, evenly rounded distal margins that
envelop only the proximal 0.07—0.09 mm
of the opercular scales; however, the dorso-
lateral margins of buccal scales usually bear
2—4 low ridges. Opercular scales typical in
shape for the genus, the abaxial up to 0.57
mm, the outer- and inner laterals equal to
or longer than the abaxials (0.54—0.69 mm),
and adaxials up to 0.43 mm in length, all
operculars having a H:W of 1.4—1.5. Coe-
nenchymal sclerites irregular in shape, up
to 0.76 mm in length, but mostly 0.4 mm
in length. These scales, like those of the
polyps, bear small (10 wm diameter) blunt
granules exteriorly, which are occasionally
linked in short rows but never formed into
ridges. Inner faces of coenenchymal scales
as well as those of polyp scales bear com-
plexly ornamented tubercles about 10 wm
in diameter. Tentacular sclerites not ob-
served.

Comparisons.—C. josephinae is quite
similar to P. simplex, as can be seen in the
comparison of characters in Table 1, but
differs from both P. simplex and P. duplex
in having less flared and less projecting
buccal scales (Figs. 6A—C), which allows a
view of most of the opercular scales in ab-
axial view. P. josephinae also has fewer
polyp whorls per cm because the average
spacing between whorls is higher. The dor-
so-lateral ridges on the buccal and basal
scales are more prominent than those of P.
simplex. Finally, the short, broad basal
“spines” of P. josephinae may be unique,
these spines more accurately called a flat-
tened lobe.

Remarks.—Apart from having “‘erect and
regularly dichotomizing branches” (Lind-
strom 1877:6), the form of Lindstrém’s

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Paracalyptrophora josephinae, holotype, Swedish Museum of Natural History 1113: A, abaxial stereo
view of a polyp; B, lateral stereo view of a polyp; C, adaxial stereo view of a polyp. Scale bar 0.5 mm.

VOLUME 117, NUMBER 1

Fig. 7. A-—G, Paracalyptrophora josephinae, holotype: A, B, inner side of basal scale, B showing the finely
ridged projecting spine; C, buccal scale; D, ad-, OL, IL, and adaxial operculars; E, G, opercular and coenen-
chymal tubercles on lower side of opercular and coenenchymal scales, respectively; EK upper and lower faces of
3 coenenchymal scales. H-O, Paracalyptrophora carinata, holotype: H, basal scale; I, buccal scale; J, whorl of
4 polyps; K, ab-, OL, IL, and adaxial opercular scales; L, infrabasal scale; M, 3 coenenchymal scales showing
ridging; N, tentacular scale; O, margin of opercular scale. Scale bar 1: E, G, N, O = 25 wm; 2: D, F = 0.25
mm; J = 0.75 mm; L = 0.125 mm; 3: A, C, I, K, M = 0.25 mm; B = 0.083 mm; 4: H = 0.25 mm.

130

eastern Atlantic holotype and subsequently
reported colonies has not been described.
Evidently the “splendid specimen” 5.5
decimeters long was not available to Ver-
sluys (1906) or Aurivillius (1931:301) for
their re-description of Lindstr6m’s type, as
Aurivillius reported only “‘a number of
fragments, about 5-8 mm long’, and we
received on loan only a small branch for
comparison. The size stated by Aurivillius
must be centimeters rather than millimeters,
as some of the pieces had one or two bi-
furcations. Nonetheless, Lindstrém’s allu-
sion to the diameter of the ““basis”’ (=main
stem) indicates that he probably had a com-
plete colony, and, had it been a biplanar
colony, Lindstrom surely would have men-
tioned this fact. Observation of the type
specimen by Stockholm curator Bj6rn Soh-
lenius (pers. comm., 2002) confirms thatthe
holotype is uniplanar. Furthermore, accord-
ing to M. Grasshoff (pers. comm., 2002),
most of the specimens he collected and ob-
served in situ (see synonymy) were unipla-
nar.

Distribution.—Eastern and mid-Atlantic:
Bay of Biscay; Josephine, Great Meteor,
and Atlantis Seamounts; Azores (south of
Flores and Alice Bank); 214-1480 m. The
undocumented references of P. josephinae
from the western Atlantic (Grasshoff
1985b, 1986) probably pertain to the types
of P. carinata.

Paracalyptrophora carinata, new species
Figs. 1G, 7H—O, 8A-C, 9

Calyptrophora josephinae.—Grasshoff,
1985b:73 (in part: western Atlantic ref-
erence); 1986:27 (in part: western Atlan-
tic reference).

Material examined/Types.—Lesser Antil-
les, southwest of St. Lucia: 13°34’N,
61°04W, 514 m, black sand, bottom tem-
perature 8.4°C, Alb-2752, 4 Dec 1886, one
colony (holotype) with part of holdfast, and
5 detached branches, USNM 49948 (SEM
264, C1055—56, 63), paratypes.

Lesser Antilles, between St. Lucia and

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

St. Vincent: 13°34’N, 61°03 W, 514 m, black
sand, bottom temperature 8.4°C, Alb-2753,
4 Dec 1886, one incomplete colony with
part of holdfast and tangled with hemp fi-
bers from the tangle-bar, USNM 49968,
paratype.

Type _locality.—13°34'N, 61°04'W
(southwest of St. Lucia, Lesser Antilles),
514 m.

Description.—Colonies consist of a ro-
bust, vertical main stem, which gives rise
to a pair of parallel, dichotomously branch-
ing fans. The main stem is anchored in a
dense, white, irregularly-shaped calcareous
mass—the holdfast—the largest of the two
observed being 18 mm in width. The main
stem of the larger specimen (the holotype)
is inflexible, 5 cm in height, and 4.4 mm in
maximum diameter, supporting fans up to
18 cm in height and 8 cm broad, the entire
colony being 23 cm in height. The stem
axis is golden-yellow to bronze in color and
faintly longitudinally striate. Branching is
uniformly dichotomous (but not lyrate), the
first two internodes being quite short, the
remaining internodes, which may number
up to 10 for certain terminal branches, are
spaced fairly uniformly at intervals of 18—
21 mm throughout the colony. Occasionally
terminal branches are up to 8 cm in length.
A slight irregularity in the branching pat-
tern of the paratype has led to three of the
first four branches contributing to one fan,
the opposite, parallel fan being smaller,
originating from only one of the original
four branches.

Polyps are arranged in whorls and di-
rected downward, each whorl consisting of
4-8 polyps (usually 6); 12—/4—-16 whorls
occupy 3 cm of axial length. In general,
polyp whorls are well spaced, such that
each whorl is separated by 0.60—0.65 mm.
Polyps are common on the main stem, often
arranged in spirals around the axis. Individ-
ual polyps are 1.50—1.75 mm in length and
0.80—0.92 mm in width.

Each polyp is protected by two pairs of
large abaxial body wall scales and a pair of
smaller crescent-shaped infrabasals, which

VOLUME 117, NUMBER 1

Fig. 8. Paracalyptrophora carinata, holotype, USNM 49948: A, abaxial stereo view of polyps; B, lateral
stereo view of a polyp; C, adaxial stereo view of a polyp. Scale bar 0.5 mm.

132

are about 0.30 mm in height and typical in
shape for the genus. The basal scales are
the largest sclerites, up to 1.2 mm in height,
the distalmost 0.45—0.50 mm consisting of
a prominent, pointed, finely serrated spine.
Each spine is covered with several rows of
closely spaced teeth. The dorso-lateral mar-
gins of the basal scales are not ridged, but
acutely curved to cover the lateral sides of
the polyp. The buccal scales are 0.9—1.0
mm long and have fairly straight, finely ser-
rate (apices of triangles about 6 pm tall)
distal margins that are not flared and over-
lap the basal margins of the operculars by
only 0.10—0.15 mm, which exposes most of
the opercular scales in lateral or adaxial
views (Figs. 8A—B). There is a slight swell-
ing on the center of the proximal third of
each buccal from which a low ridge origi-
nates and continues along the dorso-lateral
margin of the sclerite (Fig. 8A). The oper-
cular scales are triangular in shape, and, in
general, decrease in size and H:W ratio
from ab- to adaxial direction. Of the two
abaxial operculars, usually only one is sym-
metrical, the other being more developed on
the adaxial side. These operculars are up to
0.85 mm in height, the symmetrical one
having a H:W of 1.58, the asymmetrical of
1.9. The outer-lateral operculars are of
equal height but similar to the asymmetrical
abaxials in shape. The inner-lateral oper-
culars are also asymmetrical in shape but
slightly smaller and squatter in shape, only
up to 0.7 mm in height, having a H:W of
1.7-1.8. The adaxial operculars are sym-
metrical, rarely over 0.55 mm in height, and
have a broad base with a H:W of 1.2-1.4.
Tentacular sclerites are very rare, shaped as
flattened rods up to 82 wm in length and 26
um in width.

Coenenchymal sclerites are elongate to
irregular in shape, up to 0.87 mm in max-
imum length. The exterior surface is cov-
ered by small granules (14—15 wm in di-
ameter) and prominent longitudinal or retic-
ulately arranged ridges (Fig. 7M). The inner
surface of the coenenchymal scales, as well
as those of the polyps, are covered with

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

complexly ornamented tubercles 10-12 1m
in diameter.

Etymology.—Latin carinata = keeled, an
allusion to the ridged coenenchymal scales.

Comparisons.—Paracalyptrophora cari-
nata is easily distinguished from the two
other western Atlantic species by its polyp
morphology: having non-flared, straight-
edged buccal sclerites that cover only the
bases of the opercular scales. In addition,
P. carinata has larger polyps and thus less
whorls per axis length, non-ridged basals,
ridged coenenchymal scales, and small ten-
tacular scales (see Table 1). Tentacular
scales are also present in Japanese material
of P. kerberti (Versluys) but the taxonomic
significance of this character in Paracalyp-
trophora has yet to be determined.

Paracalyptrophora carinata 1s most sim-
ilar to the eastern Atlantic P. josephinae,
especially in polyp morphology, both spe-
cies having very similarly-shaped buccal
scales with non- or only slightly flared,
straight distal margins. However, P. cari-
nata differs in having a biplanar colony and
having larger polyps with consequently
larger opercular scales. It also has much
taller basal spines and a lesser developed
dorso-lateral ridge of the basal scales. Fur-
thermore, P. carinata has non-flared buccal
scales, whereas those of P. josephinae are
slightly flared, and the operculars of P. car-
inata are pointed outward, whereas those of
P. josephinae are usually pointed down-
ward toward the branch axis. Each of these
differences taken separately might indicate
range of variation or perhaps a subspecies
of P. josephinae, but taken together these
consistent differences are considered to
warrant differentiation as a different spe-
cies.

Distribution.—Known only from south-
west of St. Lucia, Lesser Antilles (Fig. 9);
514 m.

Paracalyptrophora mariae
(Versluys, 1906)
Figs. 1H, 10A—C

Calyptrophora mariae Versluys, 1906:107—
109, pl. 9, fig. 25, text-figs. 140—145 (Ti-

VOLUME 117, NUMBER 1

Gulf of Mexico

Fig. 9. Distribution of Paracalyptrophora simplex (circles) and P. carinata (square).

mor Sea).—Kitikenthal, 1919:474 (diag-
nosis); 1924:317, 318-319 (key, diagno-
sis).—Aurivillius, 1931:301 (key).—van
Soest, 1979:103 (type deposition).
Calyptrophora (Paracalyptrophora) mar-
iae.—Kinoshita, 1908:58 (listed).
Paracalyptrophora mariae.—Bargibant in
Forges, Grandperrin & Laboute, 1987:34
(listed).—Bayer, 2001:367 (listed).

Material examined.—Chalcal II, CP25
(HGP-44), 23°38.6'S, 167°43.12’E (Stylas-
ter Bank, on Norfolk Ridge just southeast
of New Caledonia), 418 m, | large colony
(NMNH) and SEM stubs 1202-1204
(USNM).

Types.—A fragment of the holotype is
deposited at the ZMA (Coel. 7414), but the
larger colony is missing (van Soest 1979).
Type Locality: 10°39’S, 123°E (Roti Strait
between Timor and Roti), 520 m.

Diagnosis.—Colonies biplanar, the larg-
est of the two known specimens (the holo-
type) 27.5 cm in height, consisting of a
main stem 5.5 cm in height and two parallel
fans, each about 22 cm in height and 16 cm
in width. Branching dichotomous (equal,
not lyrate), occurring every 2—3.5 cm in the
lower half of fan, the distal branches often
over 10 cm in length and rarely the result
of more than 5 bifurcations. Stem axis
black; branches often a metallic gold. Polyp
whorls consist of 4—7 polyps, the larger
number on thicker branches; 11—15 whorls
occur over 3 cm branch axial length; adja-
cent whorls separated by 0.5—1.0 mm, de-
pending on branch diameter. Polyps 1.4—1.8
mm in length and about 0.9 mm in width.
Infrabasal scales crescent shaped and about
0.15 mm in height, each bearing one prom-
inent longitudinal ridge. Basals about 0.85

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 10. Paracalyptrophora mariae, Chalcal Il, CP 25, USNM Stub 1202, 1204: A, abaxial stereo view of
a polyp; B, lateral stereo view of a polyp; C, adaxial stereo view of a polyp. Scale bars 0.5 mm.

VOLUME 117, NUMBER 1

mm in height, the distal 0.20 mm being a
robust, projecting spine. Dorso-lateral mar-
gins of basals bear 3 or 4 prominent, serrate
ridges (Fig. 10B). Buccal scales about 0.85
mm in length and have a slightly flared and
slightly projecting dorso-lateral distal mar-
gin, which nonetheless covers only the bas-
al part (about 0.2 mm) of the opercular
scales. Dorso-lateral margin of each buccal
scale bears 4 or 5 prominent ridges (Fig.
10A). Operculars typical for the genus, the
abaxial operculars up to 0.83 mm in height
and adaxials only 0.36 mm, but most oper-
culars maintaining a H:W of 1.5—1.7. Ten-
tacular scales not noted. Coenenchymal
branch sclerites irregular in shape, rarely
more than 0.5 mm in maximum length, and
covered externally with granules and prom-
inent ridges (Fig. 10B).

Comparisons.—Paracalyptrophora mar-
iae is distinguished from all other species
by having prominently and multiply-ridged
body wall scales (Table 1), including the
infrabasals, as well as ridged coenenchymal
scales.

Remarks.—Despite a moderate synony-
my, this species is known from only two
specimens, the holotype and the specimen
listed without comment by Bargibant
(1987), illustrated herein, who must also be
credited with the new combination. The
New Caledonian specimen is similar to the
description of the holotype, differing pri-
marily in having slightly smaller polyps
(1.4 mm vs. 1.6—-1.8 mm) and thus more
whorls per 3 cm (14—15 vs. 11-12).

Distribution.—Timor Sea and southeast
of New Caledonia; 418—520 m.

Paracalyptrophora kerberti
(Versluys, 1906)
Figs. 1I, 11A—-C, 12A—J

Calyptrophora japonica.—Studer, 1878:
642 (Japan).

Calyptrophora kerberti Versluys, 1906:
105-107, text-figs. 134-139 (Japan).—
Nutting, 1912:59 (Japan).—Ktikenthal,
1919:472—473, text-figs. 223-226 (Ja-

135

pan); 1924:318, text-fig. 173 (key, diag-
nosis).—Aurivillius, 1931:301 (key).—
van Soest, 1979:103 (type deposition).—
Utinomi, 1979:1011—1013, fig. 2a—1 (Sa-
gami Bay).

Calyptrophora (Paracalyptrophora) ker-
berti.—Kinoshita, 1908:58, 63-65, pl. 4,
fig. 29 (Japan).

Calyptrophora (Paracalyptrophora) Ker-
beti (sic).—Kinoshita, 1909:8—9, pl. 1, fig.
2, 2 text-figs. (Japan).

Paracalyptrophora kerberti.—Bayer, 2001:
367 (listed, new comb.).

Material examined.—Japan: Alb-5093, 1
colony, USNM 30105 (reported by Nutting,
1912), SEM C1058-62, 1064.

Types.—The holotype is deposited at the
ZMA (Coel. 2294) (van Soest 1979). The
second specimen described by Versluys,
also from Japan (Hilgendorf collection), is
interpreted as a paratype, and is deposited
at the ZMB (2065). Type Locality: “‘Ja-
pan”, depth unknown (Bloemhoff collec-
tion), although Utinomi (1979) suggests
that the specific type locality is Sagami
Bay.

Diagnosis.—Colonies biplanar, the larg-
est known colony (Kiikenthal 1919) 24 cm
in height and 11 cm in fan width; greatest
stem length (Versluys 1906) 9 cm. Branch-
ing dichotomous (equal, not lyrate), most
branching occurring in lower half of fan,
the distal branches rarely over 6.5 cm in
length are rarely the result of more than 6
or 7 bifurcations. Stem axis brown, black,
or golden. Polyp whorls consist of 4—8
(usually 5) polyps; 8-13 (usually 10)
whorls occur over 3 cm branch length; ad-
jacent whorls widely spaced, 0.5—2.0 mm.
Polyps 2.0—3.0 mm in length and about 1.2
mm in width. Infrabasals crescent shaped,
about 0.35 mm in height. Basals 1.4—1.5
mm in height, the distalmost 0.6—0.8 mm a
prominent sharp spine, which bears one
finely serrate ridge on its outer surface; oth-
erwise the basal scales are unridged and
fairly smooth. Buccal scales 1.3—1.9 mm in
length, the longer scales those having a dis-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 11. Paracalyptrophora kerberti, Alb-5093, USNM 30105: A, abaxial stereo view of a polyp with

spinose buccal scales; B, lateral stereo view of a polyp with straight-margins buccal scales; C, adaxial stereo

view of a polyp. Scale bar 0.5 mm.

VOLUME 117, NUMBER 1

137

Fig. 12.

Paracalyptrophora kerberti, Alb-5093, USNM 30105: A, B, inner and outer view of a basal scale;

C, buccal scale with a small distal, triangular distal margin; D, buccal scale with a prominent distal spine; E,
ab-, OL, IL, and adaxial operculars; FE infrabasal scale; G, 2 tentacular scales; H—I, upper and lower views of
coenenchymal scales; J, 3 adaxial buccal scales. Scale bar 1: H, I = 0.25 mm; 2: G = 25 wm; 3: J = 0.125

mm; 4: A-F = 0.25 mm.

tal spine; only slightly flared at distal mar-
gin, which covers only the basal part of the
opercular scales; and relatively smooth,
without any ridges and with only sparse
granulation. Distal margin of buccals may
be straight (Fig. 11C), jagged (Figs. 11B,
12C), or bear a prominent, serrate spine up
to 0.35 mm in length projecting from the
dorso-lateral margin (Figs. 11A, 12D), all
variations present on the same colony. One
to five small (up to 0.47 mm in length and
0.22 mm in width), flat, elliptical to oval-
shaped adaxial buccal scales often present
between the interior, adaxial ridge of the
buccal scales and the adjacent adaxial and
inner-lateral operculars. These scales usu-

ally are not paired. Abaxial operculars sym-
metrical, up to 1.10 mm in height, having
a H:W of 1.4—1.7. Outer-lateral operculars
equal in height but usually slightly narrow-
er than abaxials and asymmetrical, having
a H:W of 1.7—2.3. Inner-laterals up to 0.92
mm in height, asymmetrical; H:W = 1.8.
Adaxial operculars almost equilateral in
shape (H:W = 1.1—1.2), and much smaller
(only up to 0.7 mm in length). Distal mar-
gins of the abaxials and outer-laterals are
coarsely serrate. All operculars bear prom-
inent keels on their distal, inner surfaces,
those on the larger operculars (e.g., abaxials
and outer-laterals) sometimes divided into 3
or 4 parallel crests (Fig. 12E). Outer faces

138 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

of all operculars fairly smooth, like the oth-
er body wall sclerites; inner surface tuber-
culate, but tubercles restricted to the central
region, the margins fairly smooth. Tentac-
ular sclerites flattened rods up to 92 4m in
length and 26 1m in width. Coenenchymal
sclerites elongate but irregular in shape, up
to 1.0 mm in length but usually only about
0.5 mm. Their upper surfaces are covered
with low granules 12—14 wm in diameter;
there are no ridges.

Comparisons.—Paracalyptrophora ker-
berti is the most distinctive species in the
genus, having several unique characters. It
is the only species known to have adaxial
buccal scales. It is also distinctive in having
the largest polyp size and thus the smallest
number of polyp whorls per cm (Table 1).
Furthermore, as mentioned by Nutting
(1912), it is distinctive in often, but not al-
ways, having one prominent spine on the
distal dorso-lateral margin of each buccal
scale. Finally, the exterior sculpture of all
scales is extremely reduced, the body wall
scales almost appearing as smooth.

Remarks.—Although Versluys (1906)
described the holotype as being uniplanar,
he qualified his description as being based
on a small damaged specimen, and also re-
ported a paratype that was biplanar. It was
Kinoshita’s (1908:63) opinion, based on
*““several’’ specimens, that the species bears
two parallel fans, and all subsequent re-
cords of this species were based on biplanar
colonies.

Distribution.—Off Honshu, Japan; 150—
731 m.

Acknowledgments

We wish to thank Bjorn Sohlenius
(Swedish Museum of Natural History) for
the loan of a fragment of the holotype of
Calyptrophora josephinae, and Lisa Levi
(Museo Regionale di Scienze Naturali, Tu-
rin) for the loan of the type of Primnoa re-
gularis. We are also grateful to Manfred
Grasshoff for sharing his knowledge of
eastern Atlantic Paracalyptrophora. Final-

ly, we thank Marilyn Schotte and Linda
Cole for their technical assistance in trans-
lation and constructing the distribution
map, respectively.

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. 1989. Die Meerenge von Gibraltar als Fau-
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. 1870. Catalogue of the lithophytes or stony
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139

. 1924. Coelenterata: Gorgonaria. Das Tierreich
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(1):140-149. 2004.

Notes on the genus Dicliptera (Acanthaceae) in Bolivia

D. C. Wasshausen and J. R. I. Wood

(DCW) Department of Systematic Biology—Botany, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560-0166, U.S.A.,
e-mail: wasshausen.dieter @nmnh.si.edu;
(JRIW) Department of Botany, University of Oxford, South Parks road, Oxford, OX1 3RB, U.K.,
e-mail: jriwood @hotmail.com

Abstract.

Taxonomic notes on Dicliptera are presented in preparation for

the authors’ forthcoming annotated and illustrated checklist of Bolivian Acan-
thaceae. Two new species (D. palmariensis and D. purpurascens) are described
and illustrated. Infraspecific variation of D. squarrosa is discussed. A key to
all of the recognized species of Dicliptera from Bolivia is also provided.

Dicliptera is one of the most difficult
genera taxonomically in the Acanthaceae.
Like Dyschoriste the genus often lacks
clear-cut characters to distinguish among
species. Most authors have depended on
bract characters to differentiate among spe-
cies but in fact the bracts are quite variable
within most species so this character needs
to be used with caution and in conjunction
with other characters such as corolla size,
which is often useful. We have taken a
broad view of each species both in this pa-
per and in our planned treatment of Boli-
vian Acanthaceae. By doing this it seems
that the species we recognize make some
geographical and ecological sense, although
the variation found in almost every case is
quite extreme. This applies both to local en-
demic species such as Dicliptera cocha-
bambensis Lindau and to the more wide-
spread species such as D. squarrosa Nees
and D. jujuyensis Lindau.

Seven species of Dicliptera are presently
recognized in Bolivia. Two are new, one of
which (D. palmariensis) is endemic to Bo-
livia. Of the other five, one (D. cochabam-
bensis) 1s endemic to Bolivia, two extend
to northern Argentina (D. jujuyensis and D.
scutellata Griseb.) and two (D. sexangular-
is (L.) Juss. and D. squarrosa Nees) are
widespread in South America. The seven

species can be separated by the following
key.

Key to Species of Dicliptera in Bolivia

1. Inflorescence of naked spikes forming a
panicle of spikes; bracts minute, oblan-
ceolate, < 5 mm long .... D. sexangularis

1. Inflorescence varied but if paniculate,
leafy; bracts usually > 8 mm long, not
oblanceolate

2. Cymule bracts ovate, elliptic or obovate,
scarcely broader than long, not leaf-like;
corollaideepy pina eee eee 3

2. Cymule bracts linear to linear-(ob-)lan-
ceolate, sometimes with a leaf-like apex,
always much broader than long; corolla
red, orange or yellow

3. Cymule bracts ovate, 6-15 mm wide;
leaves pubescent below D. scutellata

3. Cymule bracts elliptic to obovate, 3-6
mm wide; leaves almost glabrous below

D. cochabambensis

4. Cymule bracts usually leafy at the apex;
flowers usually in dense, sessile clusters
or heads in the leaf axils .... D. squarrosa

4. Cymule bracts not leafy at the apex;
flowers in pedunculate, axillary cymes,
often forming a leafy panicle

5. Cymule bracts narrowly linear-elliptic,
broadest in the middle; leaves softly pu-
bescent above D. palmariensis

5. Cymule bracts linear-lanceolate, broad-

VOLUME 117, NUMBER 1

est at the base; leaves soon glabrescent
above
6. Inflorescence branches leafless, usually
short; bracts linear-oblong, acute; corolla
lobes almost half as long as the tube ....
Schatten Amare de eike a) ea ein D. jujuyensis
6. Inflorescence branches often with sub-
tending leaves, often well-developed;
bracts lanceolate, finely acuminate; co-
rolla lobes less than one fourth as long
as the tube D. purpurascens

Dicliptera palmariensis Wassh. & J. R. I.
Wood, sp. nov.
Fig. 1 A-H

Quoad formam bractearum cymulorum
Diclipteram garciae Leonard tangit, ob fo-
lia pilis lanatis induta, bracteas acutas, non
apiculatas ab ea removendum.

Ascending or weakly erect, much-
branched, probably perennial herb to 0.75
m; stems dark purplish-green, obscurely
ridged with paler vertical lines along the de-
pressions, densely pilose with long, patent,
straggly, multicellular trichomes; leaves
petiolate, the petioles 0.3—1.6 cm long, pi-
lose, the blades ovate or elliptic, acute at
apex, narrowed to the base and = attenuate
on the petiole, 3—9 cm long, 1—4 cm wide,
both surfaces pilose with large-celled tri-
chomes, especially on the veins, cystoliths
abundant above, the margin entire or ob-
scurely repand, ciliate; inflorescence of pe-
dunculate cymes in the axils of the upper
leaves, the cymes typically few-flowered
and the flowers often aborting, the inflores-
cence thus rather lax and open; peduncles
1-10 cm long, subtending bracts leaf-like,
shortly petiolate, the petioles 3—5 mm long,
the blades lanceolate or lanceolate-elliptic,
acute, 0.7—2 cm long; cymules pedicellate,
the pedicels ca. 0.5 mm long; cymule bracts
slightly unequal, 8-15 mm long, 3 mm
wide, narrowly oblong-elliptic, acute at
both ends, pilose, the base often pale; inner
bracts 6—10 mm long, lanceolate, ciliate on
the upper margins; bracteoles lanceolate,
ca. 4 mm long; calyx 2.5—3 mm long, 5-
lobed to just above the base, the lobes

141

equal, ca. 2 mm long, lanceolate, acute, cil-
iolate; corolla red, 25—28 mm long, cylin-
drical from a slightly bulbous base, gradu-
ally widened to ca. 3 mm, pilose without,
2-lipped, the lips ca. 3 mm long; anthers
equaling the corolla; filaments 14 mm long,
sparsely pilose, inserted ca. 14 mm above
the base of the corolla, anther thecae at dif-
ferent heights, glabrous, ca. 1.25 mm long;
ovary pubescent; style ca. 25 mm long,
with a few scattered trichomes; capsule 6
mm long, obovoid, pubescent, 2-seeded;
seeds with a few, short trichomes, lenticu-
lar, ca. 2.25 mm wide.

Type.—BOLIVIA: Tiraque, 1—2 km
above El Palmar along the old road from
the Chaparé to Cochambamba, 900 m, 6 Jul
1997, J. R. I. Wood 12403 (holotype K!;
isotypes LPB, US!).

Additional specimens.—BOLIVIA: Ti-
raque, 1—2 km above El Palmar along the
old road from the Chaparé to Cochabamba,
1200 m, 6 Jun 1998, J. R. I. Wood 13674
(K, LPB, US); El Palmar, 155 km along old
road from Cochabamba to Villa Tunari
[LSS GAB WO tn, 2 Seo ISIS
Kessler et al. 8115 (GOET, LPB, US).

The only possible Bolivian species D.
palmariensis could be confused with is D.
purpurascens but it can readily be distin-
guished by its diffuse ascending habit, very
pubescent indumentum, pedunculate cymes,
smaller corollas and above all by the short-
er, narrowly oblong-elliptic bracts, broadest
in the middle and narrowed to both an acute
apex and base. However, there are two col-
lections from San Martin Department in
Central Peru (Schunke 3349 and 4370, both
at K and F), which are in many ways in-
termediate between D. palmariensis and D.
purpurascens. The bracts are similar in
shape to those of D. palmariensis but the
apex is acuminate and apiculate and the
specimens lack the distinct pilose indumen-
tum of D. palmariensis. Given the wide
variation found in many species of Diclip-
tera these specimens might suggest D. pal-
mariensis should be included in a very var-
lable D. purpurascens but they are far re-

142 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

\\ y
NY
y
q
WY j
ves PA
WN IPNYIVE
\ i
a
AWA
NWA
We ~
I: NV

LEZ

Fig. 1. A-H, Dicliptera palmariensis (J. R. I. Wood 12403). A, Habit; B, Pedunculate cymes; C, Calyx and

corolla; D, Inner bract, bracteoles, calyx and aborted flower; E, Calyx and pistil; K Calyx lobes and nectar disk;
G, capsule; H, Capsule dehisced.

VOLUME 117, NUMBER 1

moved geographically from D. palmarien-
sis and are not exactly intermediate
between the two recognized species. It
seems best, therefore, to recognize the two
species particularly as there are no inter-
mediates in Bolivia.

Dicliptera purpurascens Wassh. & J. R. I.
Wood, sp. nov.
Fig. 2 A-F

Species nova plerumque purpurascens
bracteis longis (usque 2.5 mm) lanceolatis,
long-acuminatis bene distincta.

Annual or short-lived perennial herb,
0.5—2.5 m high, usually erect in open situ-
ations but commonly ascending or even de-
cumbent in moister, shady conditions; stems
stout, somewhat woody below, strongly an-
gled, usually purplish, scurfy-pubescent,
much branched; leaves petiolate, the peti-
oles 0.5—4 cm long, scurfy-pubescent, the
blades equal or nearly so, ovate or ovate-
elliptic, 4-15 cm long, 2—7 cm wide, acute
or shortly acuminate at apex, tapering at the
base, entire, often purplish, darker green
above than below, glabrous except for the
usually ciliolate margins and a few scat-
tered, usually multicellular trichomes, es-
pecially on the veins, cystoliths scattered on
both surfaces; inflorescence of shortly pe-
dunculate or subsessile, axillary and ter-
minal cymes, these becoming very dense on
older plants with 1—3 cymes arising from
each axil, commonly purplish and glandu-
lar-pilose but sometimes greenish and very
thinly pilose; peduncles 0-3 cm long,
scurfy-pubescent; subtending bracts leaf-
like, petiolate, the petioles 0-3 mm long,
the blades typically narrowly oblong-ellip-
tic, to 3 cm long; cymules pedicellate, the
pedicels 0-4 mm long; cymule bracts
slightly unequal, 20—25 mm long, 2-5 mm
wide, lanceolate, long-acuminate; inner
bracts linear-acuminate, 12-15 mm long;
bracteoles similar but only to 10 mm long;
calyx 3—4 mm long, 5-lobed to just above
the base, the lobes subulate, minutely cili-
olate; corolla orange-red, 34-40 mm long,

143

cylindrical from a slightly bulbous base,
gradually widened to 3—4 mm, sparsely pi-
lose and minutely gland-dotted without, 2-
lipped, the lips ca. 4 mm long; anthers
equaling the corolla; filaments 17 mm long,
sparsely pilose, inserted ca. 13 mm above
base of corolla, anther thecae at different
heights, glabrous, ca. 1.5 mm long; ovary
pubescent; style ca. 29 mm long, glabrous;
stigma globose; capsule 7 mm long, 4 mm
wide, obovoid, pubescent, 2-seeded; seeds
papillose, lenticular, ca. 1.25 mm wide.
Type.—BOLIVIA: Carrasco, ca. 5 km E
of Valle de Sajta on main road from Chi-
moré to Santa Cruz, 240 m, 29 May 1996,
Wasshausen, Brummitt, Wood & Ritter
2067 (holotype US!; isotypes K!, LPB).
Habitat and distribution.—Dicliptera
purpurascens is locally frequent in moist
lowland rain forest between 200 and 600 m
in Bolivia and Peru. It is essentially a plant
of the SW basin of the Amazon River with
an outlying population in a very moist area
of the Andean foothills in Bolivia. It has
not yet been found in Brazil but is likely to
occur in Acre as well. This disjunct distri-
bution is shared with a number of other
Acanthaceae species, notably Pachystachys
spicata (Ruiz & Pavon) Wassh., Ruellia in-
flata Rich., R. yurimaguensis Lindau, Jus-
ticia megalantha Wassh. & J. R. I. Wood
(in press), J. pilosa (Ruiz & Pavon) Lindau
and J. riedeliana (Nees) V. A. W. Graham
and appears to be a common pattern.
Additional specimens.—BOLIVIA: San-
ta Cruz, Ichilo, by track from Escuela Ichilo
to Campamento Ichilo on E side of Rio Ich-
ilo, Ambor6 Park, 400 m, 27 Jul 1999, J.
R. I. Wood 14943 (K, LPB); Cochabamba,
Carrasco, Valle de Sajta, 1 Jul 1988, Hen-
sen 6 (BOL, US); km 228, Santa Cruz road,
Rio Murillo, Valle de Sajta, 212 m, 18 Jul
1990, Sigle 510 (US); Experimental Sta-
tion, Valle de Sajta, 280 m, 11 Aug 1990,
I. Vargas 673 (LPB, USZ); Valle de Sajta,
ca. 235 km NW of Santa Cruz, 400 m, J.
R. I. Wood 10072 (K, LPB, US); 0.5 km E
of Valle de Sajta, 250 m, 29 May 1996, J.
R. I. Wood 11178 (K, LPB); 12 de Julio,

144 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

———

EE
2

= <<
J

Fig. 2. A-K Dicliptera purpurpascens (Wasshausen 2067). A, Habit; B, Pedunculate cymes; C, Inner bracts,
bracteoles, calyx and pistil; D, Calyx and pistil; E, Corolla; R Nectar disk, pistil and calyx lobes.

VOLUME 117, NUMBER 1

ca. 9 km S of Israel, on E side of Rio Sajta,
400 m, 24 Jul 1999, J. R. I. Wood 14893
(K, LPB); Zona del Arroyo de 6 de Agosto,
Cerro de la Concordia, E bank of Rio Ichoa,
600 m, 27 Jul 1999, J. R. I. Wood 14935
(K, LPB); Pando, Abuna, Nacebe, Rio Or-
ton, 11 Oct 1989, Beck et al. 19283 (LPB,
US); Gentry et al. 77583 (MO, US). PERU;
Cuzco, Convencion, along Rio Pichari, 2
km E of Colonizacion Pichari, 620 m, 13
Jun 1975, Wasshausen & Encarnacion 544
(K, US); Paucartambo, Kosfipata District,
along trail behind and W of Pilcopata, 580
m, 26 Jun 1975, Wasshausen & Encarna-
ci6n 583 (K, US); Quispicanthis, 3 km E of
Quincimil, 960 m, 7 Oct 1976, Wasshausen
& Encarnacion 735 (US); Madre de Dios,
Manu, Adan Rajo, km 225, Shintuya-Pil-
copata, 520 m, 26 Jun 1975, Wasshausen
& Encarnacion 575 (K, US); Talmamanu,
Chilifas, km 18 on Iberia-IMapari road, 1
Jun 1978, Encarnacion 1169 (K, US); near
Shintuya, along Alto Rio Madre de Dios,
450 m, 13 Oct 1979, Gentry et al. 26736
(MO, US); Manu, Parque Nacional de
Manu, Est. Cocha Cashu [11°50’S,71°25’
W], 350 m, 4 Aug 1984, Foster 9746 (MO,
US); Explorer’s Inn, near confluence of Rio
Tambopata and Rio La Torre, 29 km SW of
Puerto Maldonado [12°50’S, 69°20’W], 9
Jul 1987, Smith, Smith & Condon 938 (K,
US); Rio Tambopata [12°48’S, 69°17'W],
200 m, 9 Jul, 1998, Michelangeli 477 (US);
Ayacucho, La Mar, on trail between Santa
Rosa and Sanabamba along Rio Santa Rosa,
700 m, 9 Jun 1975, Wasshausen & Encar-
nacion 531 (K, US).

There is considerable variation in the in-
dumentum and color of Dicliptera purpur-
ascens. The purple colored form is the only
form found in central Bolivia in the De-
partments of Cochabamba and Santa Cruz
while only green forms are known from
Pando. Both forms occur in Peru but the
purple one is a good deal more common.
All plants from Bolivia are densely glan-
dular-pilose. In Peru plants are more com-
monly glandular-pilose but thinly pilose
forms also occur.

145

Dicliptera purpurascens is obviously re-
lated to D. palmariensis but the two species
are immediately distinguished by the dif-
ferent bracts.

Dicliptera squarrosa Nees

Dicliptera squarrosa Nees, in Mart., Fl.
Bras. 9:161. 1847. Type: Brazil, Minas Ger-
ais, Reidel 34 (lectotype, here chosen,
GZU!; isolectotype NY!); sin loc., Schiich
S.n. (Syntype W, not seen).

Dicliptera sericea Nees, in Mart., FI.
Bras. 9:162. 1847. Type: Brazil, Sao Paulo,
Sorocoba, Riedel & Lund 1984 (lectotype,
here chosen, LE!; isolectotype NY!).

Dicliptera pohliana Nees, in Martt., FI.
Bras. 9:162. 1847. Type: Brazil, Minas Ger-
ais, Tazenda de Roma, Pohl 2973 (lecto-
type, here chosen, W!).

Dicliptera tweediana Nees, in DC.,
Prodr. 11:482. 1847. Type: Uruguay, Porto
Alegre, Sellow 13 (d585) (syntype B, de-
stroyed); ibid, Sellow 16 (d531) (syntype B,
destroyed); Argentina, Buenos Aires, 7Twee-
die s.n. (syntype K!).

Dicliptera niederleiniana Lindau, Bot.
Jahrb. 19, Beibl. 48:18. 1894. Type: Argen-
tina, Entre Rios, Primer Misionero de Her-
nandez, Puck & Fernandez 42 (holotype B,
destroyed ?).

Dicliptera imminuta Rizzini, Arquiv.
Jard. Bot. Rio de Janeiro, 8:348. 1948.
Type: Brazil, Santa Catarina, Reitz, C861
(holotype RB).

Dicliptera rauhii Wassh., Beitr. Biol.
Pflanzen 63:425. 1988. Type: Peru, Cuzco,
prov. Urubamba, Machu Picchu, Rauh &
Hirsch P804 (holotype HEID!).

Dicliptera squarrosa is an exceptionally
widespread species extending from Brazil
south of the Amazon region westward to
the eastern slopes of the Andes in Bolivia
and then southward to Uruguay and central
Argentina. Its occurrence further north is
uncertain although we feel that Dicliptera
rauhii Wassh. from Peru belongs to this
species and probably also several species
described by Leonard from Colombia. D.

146

squarrosa is very variable with a welter of
different forms throughout its range all in-
tergrading with each other and forming no
discrete units except perhaps at a very local
level. We can make out the following rather
imprecise geographical forms:

Form 1J.—Plants from Argentina, Uru-
guay and Paraguay corresponding to the
types of D. tweediana and D. pohliana have
glabrous, narrowly lanceolate, obtuse
leaves and relatively few-flowered axillary
cymes, which become congested above into
a terminal thyrse. This form does not occur
in Bolivia but some Argentinian plants, es-
pecially from the Tucuman region have
broader leaves which approach form 4 (be-
low) found in Bolivia although the leaves
always appear to be glabrous.

Form 2.—Fig. 3 A—G. Some populations
in the Rio Unduavi Valley along the road
from La Paz to Sud Yungas appear very
distinct. These plants have subglabrous
leaves and a relatively long inflorescence of
axillary cymes forming many distinct pseu-
doverticels, which are not confluent above.
The corollas are yellow and the cymule
bracts are oblong, gray-pubescent and cili-
ate-margined with distinct squarrose tips.
Collections corresponding to this form in-
clude: BOLIVIA: La Paz, Nor Yungas, on
N side of Rio Unduavi valley, on road to
Sud Yungas, 2200-2400 m, 9 Jul 1974,
Wood 8596 (K, LPB, US); ibid, 2400 m, 1
Jul 1995, Wood 9952 (K, LPB); ca. 2 km
above El Velo de la Novia on the Sud Yun-
gas road, 2400 m, 14 Jun 1998, Wood
13716 (K, LPB); Sud Yungas, km 66 on
Sud Yungas road to Puente Villa ca. 50 m
from El Castillo, 1830 m, 12 Jun 1996,
Wasshausen & Brummitt 2123 (CAS,
GOET, K, LPB, US).

However, forms similar to form 2 but
with reddish-orange corollas and bracts
with few or no cilia occur elsewhere in the
La Paz region and also in Peru. All these
forms are difficult to distinguish from Di-
cliptera scandens Leonard from Colombia
except that they bear no field notes to sug-
gest they are scandent. Even D. scandens

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

itself is not always scandent. Collections
which conform to this more broadly-de-
fined form 2 include: BOLIVIA: La Paz,
Tamayo, on descent into Rio Yuyo, ca. 60
km S of Apolo on road to Charazani, 1150
m, 12 Jun 2000, Wood & Wendleberger
16438 (K, LPB); Murillo, 29.3 km NE of
the summit along Zongo Valley, 2200—2300
m, Solomon, Luteyn & Dorr 19068 (LPB,
MO, US); Zongo Valley, 1900 m, 28 Jun
1997, Wood 12349 (K, LPB); Sud Yungas,
ca. 15 km from Huancané on road to San
Isidro, 2300 m, | Jul 1995, Wood 9964 (K,
LPB, US); 2 km E of Puente Villa, 1200
m, 12 Jun 1996, Wasshausen & Brummitt
2124 (K, LPB, US); Inquisivi, Lewis 39127
(LPB, MO, US); Cochabamba, Ayopaya, 1
km above Independencia, 2500 m, 13 May
2000, Wood & Zaraté 16339 (K, LPB); Co-
chabamba, Ayopaya, 4 km S of Saila Pata,
Kessler 12364 (LPB, US). PERU: La Mer-
ced-Oxapampa, 2300 m, 17 Aug 1976,
Palmer 44 (K); San Martin, Zepalacio near
Moyobamba, 1200-1600 m, Mar 1934,
Klug 3601 (& K).

Form 3.—In the northern Bolivian An-
des, mostly at lower altitudes and particu-
larly in areas of high rainfall, there is an-
other form. This also has glabrous leaves
but the bracts are relatively broad, leaf-like
and mucronate, usually elliptic or obovate,
never ciliate or squarrose but commonly
pubescent to subglabrous. The axillary
cymes are relatively few-flowered. Speci-
mens that conform to this form include:
BOLIVIA: Pando, W bank of Rio Madeira,
3 km above Riberao, 27 Jul 1968, Prance
et al. 6539 (K, NY, US); Beni, Ballivian,
10 km S of Rurrenabaque, 250 m, 29 Jul
1998, Wasshausen & Wood 2162 (US,
LPB); La Paz, Caranavi, 2 km up road be-
hind Caranavi, 640 m, 10 Jun 1996, Was-
shausen et al. 2118 (K, LPB, US); Sud
Yungas, Santa Ana de Alto Beni, 580 m, 20
Aug 1963, Holliday 26 (K); 7.5 km N of
end of Road to San José, 26.5 km along
road to La Asunta, 1040 m, 5 Aug 1991,
Acevado et al. 4451 (K, US); stream at bot-
tom of ascent to Huancané, ca. 5 km from

147

VOLUME 117, NUMBER 1

———- AVY 4 <X
Sw 5 Za 2

SS =
SSS
Y, A

S—

Fig. 3. A-G, Dicliptera squarrosa Form 2 (Wasshausen 2133). A, Habit; B, Inflorescence; C, Cymes; D
Corolla; E, Inner bract, bracteoles and pistil; EK Inner bract, bracteoles, calyx lobes and pistil; G, Nectar disk,

pistil and calyx lobes.

148 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Puente Villa, 1200-1300 m, 10 Jul 1994,
Wood 8616 (K, LPB); 0.5 km from Puente
Villa along Rio Unduavi, 1200 m, 14 Jun
1996, Wasshausen & Brummitt 2130 (K,
LPB, US); 1 km above Puente Villa market
in side valley, 1100 m, 14 Jun 1998, Wood
13709 (K, LPB); Cochabamba, Chaparé, 15
km W of Villa Tunari along road to Cocha-
bamba, 800 m, 19 Jun 1994, Wood 8528
(K, LPB); Carrasco, 6 km W from main
road at Bulo Bulo, 500 m, 2 Nov 1997,
Wood 12784 (K, LPB); Santa Cruz, Ichilo,
4 km S of Huaytu towards San Rafael de
Amboro6, 500 m, 21 May 1995, Wood 9838
(K, LPB).

This form occurs over quite a wide area
and is not uniform in the size or shape or
indumentum of the bracts. The obovate
bracts of Holliday 26, for example, are very
different from the long, elliptic to subrhom-
boid bracts of Wasshausen et al. 2118. Sim-
ilarly the pilose bracts of Prance et al. 6539
are rather different from the subglabrous to
thinly pubescent bracts more commonly
seen. The common elements are the nearly
glabrous, distinct and few-flowered pseu-
doverticels and large bracts.

Form 4.—This form is characterized by
its pubescent leaves and distinct inflores-
cence. The flowers are mainly in the up-
permost leaf axils and the uppermost ver-
ticels support many-flowered cymes, which
are confluent into a dense, terminal thyrse.
This is essentially a plant of bushy stream
gullies in the Tucuman-Bolivian forest area
extending from around Pojo in the Siberia
area south to Tarija, where it perhaps inter-
grades with Form J, which differs in little
more than the glabrous leaves. It also ex-
tends east into the Chuiqutania plains where
it intergrades with Form 5. It is also similar
to some plants from Peru including Diclip-
tera rauhii and two collections from the
Macchu Pichu area [Ugent 5339 (K) and
Stafford 790 (K)], which seem to differ
only in having glabrous leaves. Specimens
that conform to this form include: BOLIV-
IA: Cochabamba, Carrasco, on ascent from
Pojo to Siberia, 2300 m, 2 Feb 1996, Wood

& Ritter 1O515 (K, LPB); Santa Cruz, Val-
legrande, 35 km SE of Vallegrande on road
to Masicuri, 1750 m, 24 May 1996, Was-
shausen, Brummitt & Wood 2039 (K, LPB,
US); Florida, 2 km W of Samaipata, 1600
m, 15 May 1994, Wood 8639 (K, LPB,
US); La Yunga de Mairana, 2300 m, 18 Sep
1994, Wood S674 (K, LPB, US); ca. 5 km
above Bermejo towards Samaipata, 1100 m,
17 Jul 1995, Wood 9995 (K, LPB, US); Ich-
ilo, Rio Suruttu, 400 m, 2 Aug 1924, J.
Steinbach 6312 (US); Guarayos, ca. 5 km
from Ascension on road to Perseverancia,
300 m, 19 Jul 1995, Wood 9999 (K, LPB,
US); Chavez, between Perseverancia and E]
Arroyan, 300 m, 22 Jul 1995, Wood 10048
(K, LPB, US); Chuquisaca, Azurduy, 4 km
N of Mollini, Sopachuy-Azurduy road,
2000 m, 15 Feb 1999, Wood & Serrano
14510 (&, LPB, US); Boeto, 10 km N of
Villa Serrano, 2300 m, 16 Apr 1995, Wood
9753 (K, LPB); 1 km below Nuevo Mundo
towards Rio Grande, 2100 m, 17 Mar 1996,
Wood 10867 (K, LPB, US); Tomina, gorge
of Rio Sillani, 3 km W of Padilla, 2200 m,
13 Feb 1994, Wood 7946 (K, LPB); 10 km
W of Padilla, 2300 m, 9 Apr 1994, Wood
8218 (K, LPB, US); on ridge between Pa-
dilla and Monteagudo, 2500 m, 10 Apr
1994, Wood 8229 (K, LPB); Rio Limon
Valley, ca. 5 km above Thiu Mayo, 1300
m, 15 Jun 1997, Wood 12305 (K, LPB,
US); Siles, 12 km E of Monteagudo to-
wards Camiri, 1300 m, 14 Apr 1995, Wood
9685 (K, LPB); Calvo, Rio Taperillas valley
between Monteagudo and Muyu Pampa,
1300 m, 14 Apr 1995, Wood 9722 (K, LPB,
US); Serrania Inca Huasi, 8 km from Muyu
Pampa towards Lagunillas, 1500 m, 8 Mar
1998, Wood, Goyder & Serrano 13255 (K,
LPB, US); Tarija, Los Pinos near Tarija,
2200 m, 11 Mar 1904, Fiebrig 3133 (K).
Form 5.—This form is essentially the
same as the previous form except that the
flowers are clearly in axillary pseudoverti-
cils rather than in a terminal thyrse and so
somewhat intermediate with Form 3. It is
apparently local in relatively open grassy
habitats in the Santa Cruz region. It is not

VOLUME 117, NUMBER 1

clear whether it is simply an adaptation to
open situations or differs genetically in
some way. Specimens that confrom to this
form include: BOLIVIA: Santa Cruz, Ichi-
lo, km 27 on old road to Cochabamba up
side road to Los Espejillos, 500 m, 20 Jul
1994, Wood 862] (K, LPB); Chavez, San
Javier, 500 m, 23 Jul 1995, Wood 10062 (K,
LPB); 15—20 km W of Concepcion on road
to San Javier, 500 m, 4 Aug 1997, Wood
12540 (K, LPB).

Form 6.—This form appears to be re-
stricted to the Tarija area. It is characterized
by having some inflorescences borne on
long, axillary peduncles. The specimen that

149

conforms to this form: BOLIVIA: Tarija,
O’ Connor, 5—6 km W of Entre Rios, 3 Jun
2000, Wood 16384 (K, LPB).

Acknowledgments

Our special thanks to Alice Tangerini
who skillfully prepared the line drawings.

Literature Cited

Ezcurra, C. 1993. Acanthaceae. Pp. 278-359 in A. Ca-
brera, ed., Flora de la Provincia de Jujuy (Re-
publica Argentina) 9, Col. Cient. INTA 13,
Buenos Aires.

Wood, J. R. I. 1988. Colombian Acanthaceae—some
new discoveries and some reconsiderations.—
Kew Bull. 43:1—51.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
117(1):150—151. 2004.

BIOLOGICAL SOCIETY OF WASHINGTON

130th Annual Meeting, 20 May 2003

President Roy McDiarmid called the meeting to order at 11:00 a.m. in the Waldo
Schmitt Room, National Museum of Natural History (NMNH). Council members present:
Marilyn Schotte and Don Wilson (Elected Council), Rafael Lemaitre (President Elect),
Richard Sternberg (Editor), Chad Walter (Treasurer), Carole Baldwin (Secretary), and
Richard Banks, Stephen Cairns, Bruce Collette, Brian Kensley, David Pawson, and Storrs
Olson (Past Presidents).

Minutes of the 129th Annual Meeting were summarized by Secretary Baldwin. Those
minutes did not appear in the Proceedings last year as usual but will appear in Volume
116(2). Following approval of the minutes, McDiarmid called on Chad Walter for the
Treasurer’s Report (Table 1). Income for the period | January 2002 to 31 December 2002
was $119,712.34, and expenses for the same period were $103,855.39. Total Society assets
as of 15 April 2003 were $89,614.17. The value of the endowment fund decreased by
$9,513.60 for the calendar year, a loss that includes fees paid to buy into the American
Funds Investment Company of America as well as a net loss from stock-market fluctua-
tions. The Audit Committee, Brian Kensley and Richard Banks, indicated that they had
reviewed the books and ledgers of the Treasurer and found all financial records to be
accurate and in good order. The Treasurer’s report was approved.

Editor Richard Sternberg reported that four issues of Volume 115 were published com-
prising 69 papers and 909 pages. As of 20 May 2003, there were 27 submissions, down
from 42 in 2002 but close to the 33 submissions at the same date in 2001. There continues
to be no backlog for papers accepted in the Proceedings.

Sternberg announced that beginning with Volume 116(1), the table of contents and
abstracts for the Proceedings can be viewed at www.allenpress.com in their “‘apt.online”’
section. Sternberg also noted that during the past year, a few authors had pulled manu-
scripts from the Proceedings at late stages in the publication process when they discovered
they would have to pay page charges. Those authors indicated that they were transferring
their papers to Zootaxa, a taxonomic journal available both online and in printed form
that does not require contributors to pay page charges. President McDiarmid announced
that he will appoint a special committee to investigate the effects Zootaxa and other largely
online journals with rapid publication rates might have on the Proceedings. The Editor’s
report was accepted.

The Finance Committee (Stephen Cairns, Oliver Flint, Frank Ferrari, and Chad Walter)
reported that three of the four recommendations approved last year by the Council had
been put into effect (increasing the cost of reprints, increasing the cost of library sub-
scriptions, and re-investing $55,000 of the Society’s endowment funds into the American
Funds Investment Company of America). Gift-fund categories for potential benefactors
have not yet been defined.

Custodian of Publications Storrs Olson announced that the organized separates have
been distributed to appropriate NMNH Divisions, and a plan to decrease the stock of
bound issues of the Proceedings will be formulated once the Society’s Web site is online.
Steve Gardiner, Associate Editor for Invertebrates for the Proceedings, has been working
on the Web site on behalf of the Society. He projected sample pages from the site for
comments and discussion.

VOLUME 117, NUMBER 1 151

In response to suggestions made at the 2002 annual meeting that a single annual meeting
be held in the future that combines the Council meeting and the annual meeting, a single
meeting was held this year with Council members meeting fifteen minutes prior to the
commencement of the annual meeting. The annual meeting was adjourned at 12:20 p.m.

Respectfully submitted,
Carole C. Baldwin

Secretary
Summary Financial Statement for 2002

General Endowment Total

Fund Fund Assets
Assets: January 1, 2002 (6,991.95) 75,790.67 68,798.72
Total Receipts for 2002 111,624.46 8,087.882 119,712.34
Total Disbursements for 2002 86,253.91 17,601.48° 103,855.39
Assets: December 31, 2002 18,378.60 66,277.07 84,655.67
Net Changes in Funds 25,370.55 (9,513.60) 15,856.95

4 Annual gain in value of Endowment.
> Annual loss in value of Endowment.

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INFORMATION FOR CONTRIBUTORS

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Front cover—from this issue, p. 70.

CONTENTS

A new genus of tiny condor from the Pleistocene of Brazil (Aves: Vulturidae)
Herculano M. F. Alvarenga and Storrs L. Olson
Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 13. An undescribed intrageneric combination

Heliodoxa imperatrix x Heliodoxa jacula Gary R. Graves
Pholidochromis cerasina, a new species of pseudochromine dottyback fish from the west Pacific
(Perciformes: Pseudochromidae) Anthony C. Gill and Hiroyuki Tanaka

Redescription of Cambaroides japonicus (De Haan, 1841) (Crustacea: Decapoda: Cambaridae) with
allocation of a type locality and month of collection of types

Tadashi Kawai and J. F. Fitzpatrick, Jr.

Two new species of freshwater crabs of the genus Chaceus Pretzmann, 1965 from the Serrania de Pera

of Colombia (Crustacea: Decapoda: Pseudothelphusidae)

Martha R. Campos and Diego M. Valencia

Reevaluation of the hermit crab genus Parapagurodes McLaughlin & Haig, 1973 (Decapoda:

anomura: Paguroidea: Paguridae) and a new genus for Parapagurodes doederleini (Doflein, 1902)

Patsy A. McLaughlin and Akira Asakura

Pseudopaguristes bicolor, a new species of hermit crab (Crustacea: Decapoda: Diogenidae) from

Japan, the third species of the genus Akira Asakura and Takeharu Kosuge
A new species of axiid shrimp from chemosynthetic communities of the Louisiana continental slope,
Gulf of Mexico (Crustacea: Decapoda: Thalassinidea) Darryl L. Felder and Brian Kensley

Description of a new Synidotea species (Crustacea: Isopoda: Valvifera: Idoteidae) from Hawaii

Wendy Moore

A new species of Synidotea (Crustacea: Isopoda: Valvifera) from the northern Gulf of Mexico

Marilyn Schotte and Richard Heard

A new genus of the Clausidiidae (Copepoda: Poecilostomatoida) associated with a polychaete from
Korea, with discussion of the taxonomic status of Hersiliodes Canu, 1888

Ju-shey Ho and I/-Hoi Kim

Vesicomyicola trifurcatus, a new genus and species of commensal polychaete (Annelida: Polychaeta:
Nautiliniellidae) found in deep-sea clams from the Blake Ridge cold seep

Jennifer Dreyer, Tomoyuki Miura, and Cindy Lee Van Dover

Studies on western Atlantic Octocorallia (Coelenterata: Anthozoa). Part 4: The genus

Paracalyptrophora Kinoshita, 1908 Stephen D. Cairns and Frederick M. Bayer

Notes on the genus Dicliptera (Acanthaceae) in Bolivia D. C. Wasshausen and J. R. I. Wood

2003 Annual Meeting Minutes

Table of Contents and Abstracts available online: www.apt.allenpress.com/aptonline

WU) A

Si/

106

114
140
150

a PROCEEDINGS oF TH:
aap IOLOGICAL SOCIETY
or WASHINGTON

4 AUGUST 2004

VOLUME 117
NUMBER 2

THE BIOLOGICAL SOCIETY OF WASHINGTON
2003-2004
Officers

President: Roy W. McDiarmid Secretary: Carole C. Baldwin
President-elect: W. Ronald Heyer Treasurer: T. Chad Walter

Elected Council

Michael D. Carleton G. David Johnson
Clyde Roper Michael Vecchione
Marilyn Schotte Don Wilson

Custodian of Publications: Storrs L. Olson

PROCEEDINGS

Editor: Richard v. Sternberg

Associate Editors

Ed Murdy

BIOLOGICAL SOCIETY OF WASHINGTON
P.O. BOX 1897
LAWRENCE, KANSAS 66044, U.S.A.

EDITOR, RICHARD BANKS

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(2):153-168. 2004.

Pseudopaguristes shidarai, a new species of hermit crab
(Crustacea: Decapoda: Diogenidae) from Japan,

the fourth species of the genuis.\\ 34)

Akira Asakura

‘
AUG 20 2004

Abstract.—Pseudopaguristes shidarai, a new species of the recently estab-
lished diogenid genus Pseudopaguristes McLaughlin, is described and illus-
trated from Okinawa, Japan. This is the fourth species assigned to this genus.

The diogenid genus Pseudopaguristes
McLaughlin, 2002, was established for P.
janetkae McLaughlin, 2002, on the basis of
specimens from Guam, the Mariana Islands.
The genus is characterized by eight func-
tional gills, male chelipeds with the right
larger than the left and dissimilar armature,
female chelipeds similar from left to right,
fourth pereopods with a clump of long cap-
sulate setae on the carpi, and the paired first
and second pleopods modified as gonopods.
The second species, P. bollandi Asakura &
McLaughlin, 2003, and the third species P.
bicolor Asakura and Kosuge, 200x, were
recorded from Okinawa, tropical Japan.
Through the courtesy of Mr. Hiroyuki Shi-
dara, the author recently obtained the fourth
species of this genus, which was again col-
lected from Okinawa. This new species is
separated from all of the described species
by coloration and morphology of antenna
and telson.

The holotype is deposited in the Natural
History Museum and Institute, Chiba
(CBM-ZC). The terminology used follows
McLaughlin (1974, 2002) with the excep-
tion of the fourth pereopods as defined by
McLaughlin (1997), gill structure by
McLaughlin & de Saint Laurent (1998),
and the posterior carapace by McLaughlin
(2000). Abbreviation used is: SL, shield
length as measured from the tip of the ros-
trum to the posterior margin of the shield.

Pseudopaguristes shidarai, new species
Figs. 1-12

Material.—Holotype: male, SL = 2.55
mm, 20-25 m, SCUBA diving, Miyako-
jima Island, Okinawa, Feb. 2003, CBM-ZC
6814. Paratypes: 2 males, SL = 1.85, 2.05
mm, 1 female, SL = 2.65 mm, data same
as holotype, CBM-ZC 6815.

Description of holotype and paratype
males.—Eight functional pairs of quadri-
serial, phyllobranchiate gills (Fig. 1A): no
pleurobranchs on fifth and eighth thoracic
somites, arthrobranchs of third maxillipeds
and chelipeds vestigial (Fig. 1B). Shield
(Fig. 1C) 1.25—1.35 times longer than
broad; anterior margin between rostrum and
lateral projections concave; lateral projec-
tions triangular, with small submarginal
spine; anterolateral angles each with blunt-
tipped corneous spine, not visible dorsally;
lateral margins nearly straight, somewhat ir-
regular; posterior margin truncate; dorsal
surface slightly convex, with some elevated
areas bearing anterior row of spines and se-
tae laterally. Rostrum (Fig. 1C) prominent,
triangular, produced, with terminal spine.
Posterior carapace lateral elements (Fig.
1C, arrow) small, well calcified, unarmed.
Branchiostegites (Fig. 1D) each with row of
spines on dorsal margin anteriorly.

Ocular peduncles (including corneas)
(Fig. 1C) moderately long, 0.75—0.85
length of shield. Corneas (Fig. 1C) very

154 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Pseudopaguristes shidarai, new species: holotype male (CBM-ZC 6814), SL = 2.55 mm, off Miyako
Is., Okinawa. A, arthrobranch gill lamella on fourth pereopod; B, vestigial gills on third maxilliped and cheliped;
C, shield and cephalic appendages; D, left lateral view of distal half of cephalothorax and antenna. Color pattern
indicated in C. Scales equal 0.5 mm (A) and | mm (B-D).

VOLUME 117, NUMBER 2 155

F,G,I-L

Fig. 2. Pseudopaguristes shidarai, new species: A—D, F—M: holotype male (CBM-ZC 6814), SL = 2.55
mm, off Miyako Is., Okinawa. E: paratype male (CBM-ZC 6815), SL = 2.05 mm, same locality. Left antennule:
A, lateral. Left antennal peduncle: B, lateral; C, dorsal; D, mesial. Right antennal peduncle: E, mesial. Left
mouthparts: E mandible, internal; G, maxillule, external; H, same, endopod; I, maxilla, internal; J, first maxil-
liped, internal; K, second maxilliped, internal; L, third maxilliped, external; M, same, proximal portion, internal.
Scales equal 1 mm.

slightly dilated. Ocular acicles (Fig. IC) scarcely setose; when fully extended, distal
each with 2 or 3 strong spines on distal margins of ultimate segments reaching dis-
margin; separated basally by breadth of ros- tal margins of corneas, spines onto seg-
trum. ments all with semitransparent tips; ulti-

Antennular peduncles (Fig. 2A) stout, mate segments unarmed; penultimate seg-

156 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3.
Is., Okinawa. Right cheliped: A, dorsal; B, mesial; C, lateral; D, dactyl, dorsal. Color pattern indicated in A—C.
Scales equal 1 mm.

ments with ventromesial margins each bear-
ing acute spine; basal segments with
ventromesial and ventrolateral distal angles
each bearing acute spine and dorsolateral
margins each bearing acute subdistal spine.

Antennal peduncles (Figs. 1C, D, 2B—E)
moderately long, when fully extended,
reaching basal portions of corneas; fifth

Pseudopaguristes shidarai, new species: holotype male (CBM-ZC 6814), SL = 2.55 mm, Miyako

segments with dorsal and ventral margins
each bearing 2 or 3 small spines; fourth
segments with dorsodistal margins each
bearing acute spine and ventrodistal mar-
gins each also bearing acute spine; third
segments each with prominent spine at ven-
trodistal margin; spines on fourth and third
segments often with semitransparent tips:

VOLUME 117, NUMBER 2

IZ—_
LA

jw Lz

MDW -

157

Fig. 4. Pseudopaguristes shidarai, new species: holotype male (CBM-ZC 6814), SL = 2.55 mm, Miyako
Is., Okinawa. Left cheliped: A, dorsal; B, mesial; C, lateral; D, dactyl and fixed finger, dorsal. Color pattern

indicated in A—C. Scales equal 1 mm.

second segments with dorsolateral distal an-
gles produced, bearing strong bifid spine
dorsally and 3 or 4 strong spines laterally,
dorsomesial distal angles each with blunt-
tipped, small spine; first segment unarmed.
Antennal acicles moderately long, straight,

terminating in strong spine; dorsomesial
margins each with 4—6 strong spines; dor-
solateral margins each with 2 or 3 strong
spines; ventral margins each with row of 9—
11 acute spines. Antennal flagella scarcely

setose.

158 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Pseudopaguristes shidarai, new species: holotype male (CBM-ZC 6814), SL = 2.55 mm, Miyako
Is., Okinawa. Right second pereopod: A, lateral; B, dactyl, propodus, and carpus, mesial; C, merus and ischium,
mesial. Left second pereopod: D, carpus, merus and ischium, mesial; E, merus and ischium, lateral. Color pattern
indicated in A—E. Scale equals 1 mm.

VOLUME 117, NUMBER 2 159

Fig. 6. Pseudopaguristes shidarai, new species: holotype male (CBM-ZC 6814), SL = 2.55 mm, Miyako
Is., Okinawa. Left third pereopod: A, lateral; B, dactyl, propodus, and carpus, mesial; C, merus and ischium,
mesial. Right third pereopod: D, carpus, merus and ischium, mesial; E, merus and ischium, lateral. Color pattern
indicated in A—-E. Scale equals 1 mm.

160 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Ag LY

Fig. 7.

Mandible (Fig. 2F) without distinguish-
ing characters. Maxillule (Fig. 2G, H) with
external lobe of endopod well developed
and recurved, internal lobe with 2 bristles.
Maxilla (Fig. 21) with moderately narrow
scaphognathite. First maxilliped (Fig. 2J)
with well developed, setose epipod. Second
maxilliped (Fig. 2K) without distinguishing
characters. Third maxilliped (Fig. 2L, M)
with carpus bearing dorsodistal spine; mer-
us with dorsodistal spine, ventral margin

Zp»,

Pseudopaguristes shidarai, new species: holotype male (CBM-ZC 6814), SL = 2.55 mm, Miyako
Is., Okinawa. Left fourth pereopod: A, lateral; B, distal portion of dactyl, dorsal; C, ventral setae of carpus. Left
first pleopod: D, external; E, internal; EK distal portion, internal, enlarged. Scales equal 1 mm (A) and 0.2 mm
(B-E).

bearing 4 or 5 spines; ischium with strong
ventrodistal and dorsodistal spines, crista
dentata well-developed, no accessory tooth;
basis with 2 sharp spines.

Male with both chelipeds bearing dense
setae covering dorsal faces of dactyls, fixed
fingers, palms and carpi and sometimes de-
tritus densely accumulating on them result-
ing into yellow or yellowish brown colored
appearance. Right cheliped (Fig. 3) stouter
than, and dissimilar from, left; dactyl as

VOLUME 117, NUMBER 2

Fig. 8.

161

Pseudopaguristes shidarai, new species: holotype male (CBM-ZC 6814), SL = 2.55 mm, Miyako

Is., Okinawa. Left second pleopod: A, external; B, same, distal portion, enlarged; C, internal. D, telson. Scales

equal 0.2 mm (A-C) and | mm (D).

long as palm, terminating in strong corne-
ous claw; dorsal face flat, with 2 rows of
large tubercles, dorsomesial margin with
row of large tubercles; mesial face with 2
rows of tubercles; cutting edge with nu-
merous corneous teeth on distal half and
broad calcareous tooth medially. Fixed fin-
ger terminating in corneous claw; dorsal
face flat, with scattered large tubercles; cut-
ting edge with few corneous teeth distally.
Palm 1.40-1.50 length of carpus; dorsal
surface flat, with scattered large tubercles;
dorsomesial margin with row of strong
spines; dorsolateral margin of palm and
fixed finger with row of strong spines. Car-
pus 0.40—0.50 length of merus; dorsal face
flat, with few large tubercles or spines, dor-
solateral and dorsomesial margins each
with row of spines. Merus with dorsal face
bearing large distal spine accompanied me-
sially with 1—3 small spines, subdistal trans-
verse row of 3 or 4 spines, and, posterior
to it, with dorsal longitudinal row of slender

semitransparent-tipped spines; ventromesial
margin with 3 or 4 strong spines, ventro-
lateral margin with row of 4 slender, semi-
transparent-tipped spines. Ischium un-
armed. Coxa with spine ventromesially.
Left cheliped (Fig. 4) slender. Dactyl
1.25-1.35 length of palm, terminating in
strong corneous claw; dorsal face with only
few tubercles, dorsomesial margin with few
tubercles; mesial face with few spiniform
tubercles; cutting edge with numerous cor-
neous teeth on distal half. Fixed finger ter-
minating in corneous claw; dorsal face flat,
with scattered large tubercles; cutting edge
with several corneous teeth distally. Palm
as long as carpus; dorsal surface flat, with
scattered large tubercles; dorsomesial mar-
gin with row of strong spines; dorsolateral
margin of palm and fixed finger with row
of large tubercles. Carpus 0.45—0.55 length
of merus; dorsal face flat, with few large
tubercles or spines, dorsolateral and dor-
somesial margins each with row of spines

162 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

or tubercles; lateral face with several tuber-
cles. Merus with dorsal face bearing large
distal spine, subdistal transverse row of 3
or 4 spines, and, posteriorly, dorsal longi-
tudinal row of spines; ventromesial margin
with 3 or 4 strong spines, ventrolateral mar-
gin with row of 3—5 slender spines. Ischium
unarmed. Coxa with spine ventromesially.

Second and third pereopods with dense
setae present on dorsal margins of each seg-
ment and sometimes detritus densely accu-
mulating on them resulting in yellow stripe-
like appearance.

Second pereopods (Fig. 5) with armature
similar from left to right; right 1.10 length
of left. Dactyls 0.90 (left) or 1.00 (right)
length of propodi, each terminating in
strong corneous claw; dorsal margins each
with row of strong spines, larger proximal-
ly; ventral margins each with row of 9 or
10 strong corneous spines. Propodi 1.85—
1.90 (left) or 1.80—1.85 (right) length of
carpi, each with row of strong spines on
dorsal margin; ventromesial distal margins
with or without spine. Carpi 0.40—0.50
length of meri, dorsal margins each with
row of strong spines. Meri each with row
of spines on ventral margin; dorsal margins
each with row of spines on proximal half.
Ischia each with or without small spine dor-
sally. Coxae unarmed.

Third pereopods (Fig. 6) with armature
similar from left to right, right 1.10 length
of left. Dactyls 0.95—1.00 length of propodi,
each terminating in strong corneous claw;
dorsal margins unarmed; ventral margins
each with row of 8 or 9 strong corneous
spines. Propodi 1.75—1.80 (left) or 1.90—
2.00 (right) length of carpi; dorsal faces un-
armed; ventromesial distal angles each with
1 or 2 acute spines. Carpi 0.60—0.65 (left)
or 0.55—0.65 (right) length of meri, each
with strong spine at dorsodistal angle; dor-
sal margin with 3 or 4 small spines. Meri
with ventral margins each bearing few
small spines or unarmed; dorsal margins
each with row of slender, semitransparent-
tipped spines. Ischia each with | or 2 small
dorsodistal spines. Coxae unarmed.

Sternite of third pereopods with anterior
lobe rectangular, unarmed.

Fourth pereopod (Fig. 7A) subchelate.
Dactyl terminating in strong corneous claw;
prominent preungual process present at
base of claw (Fig. 7B); ventral face with 1
or 2 corneous spines laterally. Propodal
rasp with | or 2 rows of corneous scales.
Carpus with acute dorsodistal spine; ventral
face with clump of long capsulate setae
(Fig. 7C).

Fifth pereopod chelate; dactyl and pro-
podus with well-developed rasps.

Male first pleopods (Fig. 7D—F) paired,
modified as gonopods; basal lobe with sev-
eral setae at superior mesial angle; inferior
lamella with distal margin bearing row of
short spines, and lateral margin with several
setae; internal lobe with row of setae on
mesial margin; external lobe exceeding in-
ferior lamella in distal extension. Male sec-
ond pleopods (Fig. 8A—C) paired, modified
as gonopods; basal segment with scattered
setae proximally and few setae distally; en-
dopod with several long setae; appendix
masculina twisted; lateral and distal mar-
gins and inferior face with moderately long
setae. Third to fifth left pleopods each with
exopod well developed, endopod reduced.

Uropods asymmetrical, left larger than
right; rasps of exopods and endopods well
developed; protopods each with row of
spines posteriorly.

Telson (Fig. 8D) with lateral constric-
tions; anterior portion unarmed; posterior
lobes separated by deep median cleft, left
lobe larger than right, terminal margins
with 5 or 6 spines (left) or 2 or 3 spines
(right).

Description of female paratype.—Female
paratype differs from holotype and paratype
males as follows: Chelipeds (Figs. 9, 10)
subequal, right very slightly larger; arma-
ment generally similar. Dactyl as long as
(right) or 1.10 length (left) of palm, termi-
nating in strong corneous claw; dorsal faces
of both chelipeds each with 6 large, spini-
form tubercles (right) or 1 small spine (left),
dorsomesial margins each with row of large

VOLUME 117, NUMBER 2

~ SW NYS,
oN SAIN \

163

Fig. 9. Pseudopaguristes shidarai, new species: paratype female (CBM-ZC 6815), SL = 2.65 mm, Miyako
Is., Okinawa. Right cheliped: A, dorsal; B, mesial; C, lateral. Color pattern indicated in A—C. Scales equal

1 mm.

(right) or small (left) spines; mesial faces
with several spiniform tubercles or spines;
cutting edges with numerous corneous teeth
on distal halves. Fixed fingers each termi-
nating in corneous claw; cutting edges with
several corneous teeth distally. Palms 1.10
length of carpi; dorsal surfaces of palms
and fixed fingers flat, each with row of
strong spines, right, accompanied mesially

by small tubercle on palm and row of 3
tubercles on fixed finger, dorsolateral mar-
gins of palms and fixed fingers each with
row of strong spines, dorsomesial margins
of palms each with row of strong spines;
mesial faces each with few spines (right) or
unarmed (left); lateral faces each with row
of tubercles (right) or few spines (left). Car-
pus 0.50 (right) or 0.40 (left) length of mer-

164

\ ANY

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 10. Pseudopaguristes shidarai, new species: paratype female (CBM-ZC 6815), SL = 2.65 mm, Miyako
Is., Okinawa. Left cheliped: A, dorsal; B, mesial; C, lateral. Color pattern indicated in A—C. Scales equal 1 mm.

us; dorsal faces flat, each with row of spines
on midline, dorsolateral and dorsomesial
margins each with row of strong spines; lat-
eral faces with several spines or tubercles;
mesial faces unarmed. Meri with dorsodis-
tal spine developed (right) or vestigial
(left); dorsal margins each with row of
spines; ventromesial and ventrolateral mar-
gins each with row spines. Ischia each with
row of small spines or tubercles on ventro-
mesial margin. Coxae each with spine ven-
tromesially.

Coxa of only left third pereopod with
gonopore (Fig. 11B).

First abdominal somite with paired uni-
ramous pleopods modified as gonopods
(Fig. 11B); second through fourth abdomi-
nal somites each with unequally biramous
left pleopod (Fig. 11C); fifth with exopod
well developed, endopod rudimentary;
brood pouch represented by row of setae.

Color in life (Fig. 12).—Shield cream,
rostrum red; antennules with flagella semi-
transparent red, other surfaces uniformly

VOLUME 117, NUMBER 2

165

Fig. 11.

Pseudopaguristes shidarai, new species: paratype female (CBM-ZC 6815), SL = 2.65 mm, Miyako

Is., Okinawa. A, left second pereopod, mesial. B, coxae and sternites of third to fifth pereopods, first abdominal

somite, and first pleopods. C, second pleopod.

red; antennas with flagella bearing alterna-
tive red and white bands, fifth segment with
proximal 0.30—0.40 lighter red, other sur-
faces uniformly red; ocular peduncles uni-
formly red or with lighter red band on prox-
imal 0.20; ocular acicles red except for
lighter red distal spines; second and third
maxillipeds uniformly red. Both chelipeds
generally cream in males, but orange in fe-
male; meri red except for distal 0.10—0.25.
Second pereopods generally cream; meri
red except for distal 0.20—0.40; ischium
lighter red. Third pereopods generally
cream; meri with lateral face uniform cream
or small light red patch, mesial face uni-
form red except for distal 0.30—0.40; ischi-
um with lateral face uniform cream or very
faint red, mesial face uniform lighter red.

Etymology.—This species is named for
Mr. Hiroyuki Shidara, an amateur hermit
crab collector and marine aquarist, who
kindly made the specimens available for
this study.

Distribution.—Known only from the
type locality.

Remarks.—Despite their general similar-
ities in morphology, the new species, P. shi-
darai, differs from the other three species
of the genus in shape of telson. In P. shi-
darai, the terminal margins of the telson are
horizontal, and each posterior lobe is armed
with at most only 6 (left) or 3 (right) spines
(Fig. 8D). In contrast, the terminal margins
of the telson of the holotype of P. bollandi,
and only known specimen, are oblique and
the posterior lobes each is armed with 16

166

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

}
'

j
Wf

Pseudopaguristes shidarai, new species: A—B, paratype male (CBM-ZC 6815); C—D, holotype male
(CBM-ZC 6814); E, female. A, anterior half, dorsal. B, whole animal, ventral. C, right cheliped, lateral. D, left

cheliped, mesial. E, living individual of female. Photos by Akira Asakura (A—D) and Kiyohiko Sakuma (E).

Fig. 12.

VOLUME 117, NUMBER 2

167

Table 1.—Color difference between Pseudopaguristes shidarai, new species and P. janetkae McLaughlin.

Character P. shidarai

P. janetkae

Shield anterior margin Cream

Ocular peduncles

Antennule penduncles

Ultimate segment Red
Penultimate segment Red
Basal segment Red

Antennal peduncles

Fifth segment
0.40 lighter red

Fourth segment Red
Third segment Red
Second segment Red
First segment White

Chelipeds F

Dactyl and fixed finge Cream (male)

Orange (female)

Uniform red or with lighter
red band on proximal 0.20

Red, with proximal 0.30—

Cranberry red
Cranberry red on proximal 0.25—0.35, re-
mainder yellow-orange

Red-orange

Cranberry red on proximal 0.5, remain-
der red-orange

Cranberry red

Yellow
Yellow
Yellow
Cranberry red, with yellow produced

dorsolateral distal angle
Cranberry red

Tan tinged with cranberry red
Cranberry red, becoming lighter distally

Cranberry red

Palm Cream (male)

Orange (female)
Carpus Cream (male)

Orange (female)
Merus Red, with distal 0.15—0.25

cream

Third pereopods

Merus lateral face

(eft) or 13 (right) spines. Although Mc-
Laughlin (2002) made no mention of num-
ber of the telsonal terminal spinules in P.
Janetkae, it is 25 (left) or 13 (right) in the
illustration of the holotype (McLaughlin
2002: Fig. 20). The terminal margins of the
telson of P. bicolor are strongly oblique,
and armed with 9 (left) or 6 (right) spines.

The armament of the antennas can sep-
arate P. shidarai from both P. bicolor and
P. janetkae. Produced dorsolateral distal an-
gles of the second segments of the antennas
each has strong, dorsal, bifid spine and 3 or
4 strong lateral spines in P. shidarai (Figs.
1D, 2B). The same portions of both P. bi-
color and P. janetkae are only armed with
a dorsal bifid spine and have no lateral
spines. The ventral margins of antennal aci-
cles are each armed with a row of numerous

Uniformly cream or with
only small, faint red patch

Cranberry red

Cranberry red except cream distal por-
tion

spines in P. shidarai (Figs. 1D, 2D, E), but
they are unarmed in P. bicolor and P. ja-
netkae.

By differences in coloration in life, P.
shidarai 1s readily distinguished from both
P. bicolor and P. bollandi. The chelipeds
and the second and third pereopods are uni-
formly red in P. bollandi and have alter-
nating red and white bands in P. bicolor. In
P. shidarai, these appendages are generally
cream, with red areas on the meri.

Although the coloration of P. shidarai is
somewhat similar to that of P. janetkae in
having cream colored ambulatory pereo-
pods with red proximal portions, many mi-
nor but apparent differences are seen as in
Table 1. The chelipeds of males of this spe-
cies are generally cream, with proximal red
portions (Fig. 12A—D). However, those of

168 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

females are generally orange except for the
proximal red portions (Fig. 12E). This may
exhibit sexually dimorphic color of cheli-
peds in this species. However, since only a
few specimens are examined, future collec-
tion effort will be needed to evaluate this
point. McLaughlin (2002) made no mention
on sexual difference in coloration of P. ja-
netkae.

In addition to the differences in the telson
and antennas as mentioned above, P. shi-
darai differs morphologically from P. ja-
netkae in several important characters. The
propodus of the second left pereopod of the
female P. janetkae differs from the right
and also from the propodi of the males in
having scattered spinules on the mesial
face. However, in P. shidarai, the armature
of the propodi of the second pereopods is
similar from left to right (Fig. 11A) and
also similar to those of the males. In the
female P. shidarai, the dorsal face of the
palms of the chelipeds each bears a longi-
tudinal row of spines. The same surfaces of
P. janetkae are armed with widely-spaced
and somewhat scattered, small spines.

Since so few specimens of each species
have been collected in each species, future
collection efforts will be needed to evaluate
intraspecific variation and interspecific dif-
ference more precisely.

Acknowledgements

The author is most grateful to Messrs.
Hiroyuki Shidara and Shimosato Kazuhiro
who provided the specimens of this impor-
tant species and Mr. Kiyohiko Sakuma for
a beautiful photograph of the female spec-
imen. My special thanks are due to Dr. Pat-

sy A. McLaughlin (Shannon Point Marine
Center, Western Washington University) for
her elaborate review of the manuscript. The
final draft was greatly improved by the
comments from two anonymous reviewers.
This work was partly supported by a Grant-
in-Aid for Scientific Research (C) from the
Ministry of Education, Science, Culture and
Sports of Japan awarded to Akira Asakura
(No. 14540654).

Literature Cited

, & T. Kosuge. 2004. Pseudopaguristes bicolor,
a new species of hermit crab (Crustacea: De-
capoda: Diogenidae) from Japan, the third spe-
cies of the genus.—Proceedings of the Biolog-
ical Society of Washington 117:68—83.

McLaughlin, P. A. 1997. Crustacea Decapoda: hermit
crabs of the family Paguridae from the KA-
RUBAR cruise in Indonesia. In A. Crosnier &
P. Bouchet, eds., Résultats des Campagnes MU-
SORSTOM, 16.—Mémoires du Muséum na-
tional d’ Histoire naturelle 172:433-572.

. 2000. Crustacea Decapoda: Porcellanopagu-

rus Filhol and Solitariopagurus Tirkay (Pagur-

idae), from the New Caledonian area, Vanuatu
and the Marquesas: new records, new species.

In A. Crosnier, ed., Résultats des Campagnes

MUSORSTOM, volume 21.—Mémoires du

Muséum national d'Histoire naturelle, Paris,

volume 184:389—414.

. 2002. Pseudopaguristes, a new and aberrant

genus of hermit crabs (Anomura: Paguridea: Di-

ogenidae).—Micronesica 34:185—199.

, & M. de Saint Laurent. 1998. A new genus

for four species of hermit crabs formerly as-

signed to the genus Pagurus Fabricius (Deca-
poda: Anomura: Paguridae).—Proceedings of

the Biological Society of Washington 111:158—

187.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(2):169-175. 2004.

A new species of Procambarus (Crustacea: Decapoda: Cambaridae)
from Veracruz, Mexico

Marilt. Lopez-Mejia, Fernando Alvarez and Luis M. Mejia-Ortiz

(MLM & LMMO) Universidad Autonoma Metropolitana—Xochimilco (UAM-X),
Departamento El Hombre y su Ambiente, Laboratorio de Fisiologia y Comportamiento Animal,
Calzada del Hueso 1100, Col. Villa Quietud, C.P. 04960, México D.F, México,
e-mail: Imejiam @cueyatl.uam.mx
(FA) Coleccion Nacional de Crustdceos, Instituto de Biologia, Universidad Nacional Aut6noma
de México, Apartado Postal 70-153, México 04510, D.E, México

Abstract.—Procambarus (Villalobosus) chacalli is a new species of crayfish
from ponds at Manantial de Dejigui, Huayacocotla County, Veracruz, Mexico.
It can be placed in the Erichsoni Group of the subgenus because the gonopod
(first pleopod) of the first form male has a flared, broadly curved caudal pro-
cess. Within the Group it is most similar to P. (V.) erichsoni Villalobos and P.
(V.) contrerasi (Creaser). It can be distinguished from these and other members
of the subgenus by a combination of gonopod characters that includes a short
mesial process with a slightly flattened, caudodistally directed tip; a cephalic
process that originates on the caudal process and is longer than the other ter-
minal elements, and a platelike caudal process with a strong fold on the cau-
dolateral surface. Another distinctive character is the subovate annulus ven-
tralis, with two stronger ventral crests that form a deep submedian depression,
and a sinus that extends to the caudomedian margin.

The subgenus Villalobosus Hobbs, 1972,
of the genus Procambarus Ortmann, 1905,
as defined by Hobbs (1972), includes 10
species with the following characters: pres-
ence of hooks on the fourth pereiopods, and
rarely vestigial hooks on the third pereio-
pods, of males; and asymmetrical gonopods
that reach the coxae of the second pereio-
pods, with a tuberculiform or acute central
projection. No new species has been de-
scribed in this subgenus since Hobbs (1982)
described Procambarus (Villalobosus)
cuetzalanae Hobbs, 1982, from a series of
springs, caves and deep holes in the envi-
rons of Cuetzalan, Puebla.

The species of P. (Villalobosus) inhabit
the southern portion of the Huasteca region,
within the states of Hidalgo, Puebla and Ve-
racruz. This is a very mountainous region,
with a number of narrow valleys and can-

yons that play an important role in isolating
crayfish populations (L6pez-Mejia 2001).
The members of this subgenus have been
found in rivers, small streams, impound-
ments, and springs, and in subterranean en-
vironments as stygophiles (Villalobos 1955;
Hobbs 1975, 1982, 1984). The new species
described herein has been found only in
three ponds at the type locality, Manantial
de Dejigui.

The specimens studied are deposited in
the Coleccion Nacional de Crustaceos, In-
stituto de Biologia, Universidad Nacional
Autonoma de México (CNCR), and in the
Colecci6n de Crustaceos de Referencia,
Universidad Autonoma Metropolitana—
Unidad Xochimilco (CCR-UAMX). Other
abbreviation used is: TCL, total carapace
length.

170

Procambarus (Villalobosus) chacalli,
new species
Figs. 1, 2

Diagnosis.—Body pigmented, eyes nor-
mally developed, facets well defined. Ros-
trum reaching distal border of third anten-
nular article, length 15.8 to 20.8% (x =
17.9%, n = 26) of TCL, without marginal
spines (Fig. 1A). Areola 4.9 to 7.2 (x = 5.9,
n = 26) times as long as wide, length 32.3
to 38.6% (x = 35.1%, n = 26) of TCL, 39.7
to 46.1% (x = 42.8%, n = 26) of postor-
bital carapace length, with 2 or 3 punctua-
tions across the narrowest part. Cervical
and infraorbital spines absent, branchioste-
gal spine present. Antennal scale 1.8 to 2.3
(x = 2.1, n = 26) times longer than wide,
with longitudinal groove throughout whole
length, groove shallow anteriorly, becoming
deeper posteriorly. Chelipeds shorter than
total body length; mesial surface of palm of
chela with 7 tubercles in irregular row,
based on holotypic male form I; all tuber-
cles with small tufts of short setae anteri-
orly; fingers as long as palm, both fingers
with 3 longitudinal ridges along ventral and
dorsal surfaces (Fig. 1C). Ischium of fourth
pereiopod with hook extending beyond bas-
ioischial articulation (Figs. 1D—E), ischium
of third pereiopod with vestigial hook. Ce-
phalic lobe of epistome approximately hex-
agonal, with margins undulating slightly, ir-
regular and asymmetrical anteriorly; lateral
angles well defined, devoid of setae (Fig.
1)

First pleopods of form I male asymmet-
rical, reaching coxae of second pereiopods,
with 2 rows of scattered setae running
throughout whole length, setae more abun-
dant and longer proximally. Mesial process
short, slightly truncated distally, directed
distolaterally; central projection triangular,
divided into 2 sections, caudocephalically
oriented; cephalic process spiniform, di-
rected distocephalically, longer than rest of
terminal elements, originating on caudal
process; caudal process in cephalic position
platelike, corneous, wrapping around cen-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

tral projection, with strong fold on caudo-
lateral margin (Figs. 2A—D, F—G). Prean-
nular plate of female with 2 strong lateral
crests extending laterally and surrounding
annulus ventralis, with scattered short setae
on posterior portion of plate (Fig. 1G).
Preannular plate and annulus in loose con-
tact. Annulus approximately circular, with
2 strong crests anteriorly, forming deep, V-
shaped depression; posterior half of dextral
crest curved laterally, becoming less de-
fined; sinistral crest curving laterally to
form tonguelike expansion; rectangular pro-
jection on medial posterior section, forming
margin of sinus. In ventral view, postan-
nular plate ovoid, in caudal view approxi-
mately conical; apical surface bearing small
punctations with short setae; plate not in
contact with annulus. First pleopods present
in females.

Measurements of types.—Provided in Ta-
ble 1.

Holotypic male, form I.—Body and eyes
pigmented. Cephalothorax becoming thick-
er posterior to cervical groove, maximum
width at posterior margin, 0.97 times length
of abdomen. Areola 6.2 times as long as
wide, 32.3% of TCL, with 3 punctations
across narrowest part, with slight median
crest; branchiocardiac grooves well defined.
Surface of carapace densely punctate, punc-
tations increasing in density laterally. Ros-
trum excavated dorsally, margins conver-
gent, without spines; anterior width 2.9
mm, posterior width 3.8 mm. Acumen
reaching distal border of third article of an-
tennular peduncle, slightly shorter than an-
tennal scale, tip oriented dorsally, length of
acumen 28.2% of rostrum length, ventral
keel without spines. Postorbital ridge
straight, moderately strong, with very small
cephalic tubercle. Suborbital angle acute,
branchiostegal spine present on both sides
of carapace, directed anteriorly. Cervical
groove describing acute angle over hepatic
region, cervical spine absent (Fig. 1B).

Abdomen slightly longer than carapace.
Surface of somites covered with regularly
distributed punctations. Uropods with pro-

VOLUME 117, NUMBER 2 171

Fig. 1. Procambarus (Villalobosus) chacalli, new species, all from holotypic male, form I, except G from
allotypic female. A, carapace, dorsal view; B, carapace, lateral view; C, distal podomeres of right cheliped; D,
basal podomeres of left second, third and fourth pereiopods; E, detail of basis and ischium of left fourth

pereiopod; EF epistome, cephalic lobe; G, annulus ventralis. Scale bars represent 3 mm (A, B, C), 2 mm (D, E),
and 1 mm (E G).

N72 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Measurements (mm) of type specimens of Procambarus chacalli, new species.

Total length
Carapace
Total length
Postorbital length
Width
Height
Areola
Length
Width
Rostrum
Length
Width
Antennal scale
Length
Width
Cheliped
Length of mesial margin of palm
Width of palm
Length of lateral margin of propodus
Length of dactyle
Length of carpus
Length of merus
Abdomen

Length
Width

Holotypic Allotypic Morphotypic
male, form I female male, form II
Sil2 55.5 48.7
24.1 26.9 23.6
19.6 22.6 19.3
11 12 10.2
10 12.1 9.3
7.8 10.4 8.6
2 1.7 1.7
4.5 4.25 4.2
3.8 A2 4.8
3.9 41 3.5
2.1 2.4 DP
10.5 9.5 10.5
7.5 Vel Vel

18.9 18.2 19
10.2 915 10.1
6.8 6.6 7.3
10.7 10.7 10.3
27.1 28.6 Dyk
10.4 11.3 9.8

topodite bearing short spines; endopodite
with dorsal median ridge ending posteriorly
in small spine, and well developed distola-
teral spine. Telson covered with tufts of
short setae, loosely forming 4 longitudinal
rows; cephalic portion with 4 spines on pos-
terolateral angle, two lateral ones larger,
second one articulated.

Cephalic lobe of epistome irregular, with-
out cephalomedian extension; distal half
asymmetrical, with central depression. An-
tennule with prominent ventral spine on
basal podomere, with setae on its base; an-
tenna shorter than total body length. Anten-
nal scale 1.8 times longer than wide, lateral
margin ending in acute spine, maximum
width at distal half (Fig. 2E). Third maxil-
liped reaching distal border of third article
of antennal peduncle; internal margin of is-
chium with array of 26 irregular spines; all

segments of third maxilliped with small
tufts of short setae.

Chelae 1.2 times shorter than TCL, ro-
bust, ovate, 2.5 times longer than wide.
Palm 1.4 times longer than wide, surface
covered with small, blunt tubercles each
with small tuft of setae; irregular row of
tubercles along mesial surface. Movable
finger with subsquamate tubercles anteri-
orly. Opposable margins of fingers with
small tufts of setae; opposable surface of
movable finger with 10 tubercles, that of
fixed finger with 7 tubercles, third from
base largest; both fingers ending in corne-
Ous tip.

Carpus of cheliped short, approximately
conical, dorsal surface with scattered tuber-
cles; lateral and ventral surfaces with small
subsquamate tubercles, with tufts of short
setae, distal margin with blunt spine on in-

VOLUME 117, NUMBER 2

173

Fig. 2. Procambarus (Villalobosus) chacalli, new species, all from holotypic male, form I, except H from
morphotypic male, form II. A, left gonopod, mesial view; B, detail of apex of left gonopod, mesial view: C, left
gonopod, lateral view; D, detail of apex of left gonopod, lateral view; E, antennal scale; K caudal view of

gonopods;
G, detail of apex of left gonopod, caudal view; H, caudal view of gonpods. Scale bars represent 1 mm

174

ternal surface. Merus slightly tuberculate;
dorsal surface with large, strong, subdistal
tubercle, and other smaller tubercles; ven-
tral surface with 2 longitudinal rows of
blunt tubercles, distal margin with strong
tubercle. Ischium with dorsal and ventral
surfaces punctate, and row of 6 blunt tu-
bercles along ventromesial margin, increas-
ing in size distally.

Ischium of third pereiopod with vestigial
hook, left and right sides different in size.
Ischium of fourth pereiopod with strong,
thick, cylindrical hook, extending beyond
basioischial articulation, reaching mid-
length of basis. Coxa of fourth pereiopod
with prominent acute boss on caudomesial
ventral angle.

Gonopods as described in Diagnosis.

Allotypic female.—Similar to holotype,
differing in following characters: telson
bearing 2 movable spines on left caudola-
teral angle of cephalic portion and 1 on
right side. Areola 6.2 times as long as wide.
Antennal scale 1.8 times longer than wide.
Eight tubercles on opposable margin of
movable finger. Rostrum reaching first third
of third article of antennular peduncle. Ce-
phalic lobe of epistome asymmetrical as in
holotype, distal border with small variations
with respect to holotype. Annulus ventralis
as described in Diagnosis.

Morphotypic male, form II.—Differing
from holotype in following characters: first
pleopod with apical elements poorly devel-
oped, cephalic process conical and reduced,
central projection small, caudal process sur-
rounding central projection, mesial process
undefined. Areola 4.9 times as long as
wide. Antennal scale 1.6 times longer than
wide. Chelae with irregular row of four tu-
bercles on surface of palm. Protuberance on
ischium of third pereiopod extremely re-
duced. Ischium of fourth pereiopod with
small hook, not surpassing basioischial ar-
ticulation. Rostrum shorter, acumen reach-
ing middle part of third podomere of anten-
nal peduncle.

Type locality.—Nacimiento de Dejigui
(altitude 1675 m), 4 km east of Huayaco-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

cotla, Municipio de Huayacocotla, Vera-
cruz, Mexico (20°32'6"N, 98°26’ 15”W).

Disposition of types.—Holotypic 6 form
I, CNCR 20529; allotypic 2, CNCR 20530;
and morphotypic ¢ form II, CNCR 20531.
Raratypes: 26 1, 2°36 lly SeSeeNek
20532; 5 6 I, 6 2 CCR-UAMX 1001.

Material examined.—All from type lo-
cality: 1 d form I (CNCR 20529), 14 Nov
1999, coll. M. Lopez-Mejia, L. M. Mejia-
Ortiz; 1 2 (CNCR 20530), same date and
coll. as holotype; 1 ¢ form If (CNCR
20531), 2.6 foum 2 S tornmliesees
(CNCR 20532), 5 3d form II, 6 2 (CCR-
UAMX 1001), 2 Nov 2001, coll. M. Lopez-
Mejia, L. M. Mejia-Ortiz, M. Signoret, J.
A. Viccon-Pale, J. Cruz-Hernandez, H. So-
lis.

Etymology.—The specific epithet “‘cha-
calli” is taken from the nahuatl word “‘cha-
calli’, common name used for the crayfish-
es in northern Hidalgo, Mexico.

Remarks.—Procambarus (Villalobosus)
chacalli, new species, can be placed in the
Erichsoni Group due to the presence of a
platelike caudal process on the male gono-
pod (Villalobos 1955). The new species is
morphologically similar to Procambarus
(Villalobosus) contrerasi (Creaser, 1931),
and Procambarus (Villalobosus) erichsoni
Villalobos, 1950, from which it may be dis-
tinguished by the following characters: a
longer rostrum, with tip reaching the distal
border of the third antennular article; a wid-
er areola with a slight median crest; and an
epistome bearing a cephalic lobe with an
undulated surface. Regarding the gonopod
morphology of the form I male, P. chacalli
exhibits the following unique characters: a
short and slightly truncated mesial process,
directed distolaterally; a cephalic process
which is the largest of the terminal ele-
ments, originating on the caudal process;
and a platelike caudal process with a strong
fold on the margin. In P. erichsoni and P.
contrerasi the rostrum reaches the distal
part of the second antennular article, the
areola is narrower, and the surface of the
espitome is smooth; their gonopods bear

VOLUME 117, NUMBER 2

shorter cephalic processes, that originate
between the central projection and the me-
sial process, and the caudal process is
slightly folded. The annulus ventralis of P.
chacalli differs from those of P. erichsoni
and P. contrerasi in the extension of the
lateral projections of the preannular plate,
the shape of the crests and sinus in the an-
nulus, and the size and shape of the postan-
nular plate.

Procambarus (Villalobosus) chacalli has
been collected only at the type locality,
where specimens were captured in three
small, shallow ponds next to the spring. The
largest pond was 8 m/?, and the deepest one
was 0.4 m. The recorded water tempera-
tures ranged from 18.9 to 20.4°C.

Acknowledgments

Our thanks to J. Cruz-Hernandez, M.
Signoret, H. Solis and J. A. Viccon-Pale for
their help during field work, and to Rolando
Mendoza for producing the drawings. We
are also grateful to Drs. J. E. Cooper, R.
Lemaitre and two anonymous reviewers,
for their suggestions.

Literature Cited

Creaser, E. P. 1931. Three new crayfishes (Cambarus)
from Puebla and Missouri.—Occasional Papers

175

of the Museum of Zoology, University of Mich-
igan, 224:1—10, plates I-V.

Hobbs, H. H., Jr. 1972. The subgenera of the crayfish
genus Procambarus (Decapoda: Astacidae).—
Smithsonian Contributions to Zoology 117:1-
DD.

. 1975. New crayfishes (Decapoda: Cambari-

dae) from the southern United States and Mex-

ico.—Smithsonian Contributions to Zoology

201:1-34.

. 1982. A new crayfish (Decapoda: Cambari-

dae) from the state of Puebla, Mexico, with new

locality records for Procambarus (Villalobosus)
xochitlanae and entocytherid ostracod symbi-
onts.—Association for Mexican Cave Studies

Bulletin (8):39—44.

. 1984. On the distribution of the crayfish genus
Procambarus (Decapoda: Cambaridae).—Jour-
nal of Crustacean Biology 4(1):12—24.

Lo6pez-Mejia, M. 2001. Nuevos registros de distribu-
cidn de las especies del subgénero Villalobo-
sus—Hobbs, 1972 (Cambaridae: Procambarus)
en los limites de los estados de Hidalgo, Puebla
y Veracruz. Informe Final de Servicio Social
(Tesis de licenciatura). Universidad Aut6noma
Metropolitana Xochimilco, México, D.F 32 p.
[Unpublished thesis].

Ortmann, A. E. 1905. Procambarus, a new subgenus
of the genus Cambarus.—Annals of the Car-
negie Museum 3(3):435—442.

Villalobos, A. 1950. Contribucién al estudio de los
cambarinos mexicanos, IX: estudio taxonémico
de un grupo de especies del género Procam-
barus.—Anales del Instituto de Biologia, Uniy-
ersidad Nacional Aut6énoma de México, 21(2):
367-413.

1955. Cambarinos de la fauna mexicana

(Crustacea: Decapoda). Tesis Doctoral, Facultad

de Ciencias, UNAM. México, D.EF, 290 p.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(2):176-185. 2004.

Brackenridgia ashleyi, a new species of terrestrial isopod from
Tumbling Creek Cave, Missouri (Isopoda: Oniscidea: Trichoniscidae)

Julian J. Lewis

J. Lewis & Associates, Biological Consulting, 217 W. Carter Avenue,
Clarksville, Indiana 47129 U.S.A.

Abstract.—Brackenridgia ashleyi, a new species of troglobitic trichoniscid
isopod, is described and illustrated from Tumbling Creek Cave, Taney Co.,
Missouri. The genus Brackenridgia was previously known only from the west-
ern United States and Mexico. The discovery of B. ashleyi in the Ozark Pla-
teaus physiographic province extends the range of the genus over 800 kilo-
meters to the northeast. This species is closest geographically to B. cavernarum
and B. reddelli in Texas, but is morphologically more similar to B. heroldi in
California. Despite the presence of large amounts of bat guano in the cave, B.
ashleyi is not guanophilic. By not using the guano microhabitat B. ashleyi
avoids many of the predators present in the cave community.

In the eastern United States troglobitic
trichoniscid isopods have long been known
from the karst areas of the Appalachian
Valley and Interior Low Plateaus. From the
caves of this region have been described six
species of Amerigoniscus (or Caucasone-
thes, from which Amerigoniscus was split
by Vandel 1950) and four species of Mik-
toniscus (Vandel 1950, 1965, 1978; Jass &
Klausmeier 2001). Other genera of trichon-
iscids occurring in the eastern U.S. are An-
droniscus, Haplophthalmus, Hyloniscus,
Trichoniscus, and Trichoniscoides (Leisti-
kow & Wagele 1999, Jass & Klausmeier
2001), but within these the only species of
significance in caves is Haplophthalmus
danicus. This Mediterranean species is a
widely introduced exotic in the U.S., where
it frequents caves (Vandel 1965).

In the Ozark Plateaus Craig (1975) and
Gardner (1986) reported undescribed tri-
choniscids in Missouri caves including
Tumbling Creek Cave. This cave is inhab-
ited by a diverse assemblage of troglobites,
although several of the species remain un-
described. Motivation to describe these taxa
was presented by a decline within the
cave’s ecosystem leading to listing of the

endemic hydrobiid Tumbling Creek Cave-
snail Antrobia culveri as a federal endan-
gered species (U.S. Fish & Wildlife Service
2001). Given this situation, there was a
need for characterization of the fauna and
specimens of the trichoniscid were collect-
ed for the purpose of preparing a descrip-
tion of the species.

Because of reports (Aley 1975, Craig
1975, Gardner 1986, U.S. Fish & Wildlife
Service 2001) listing this species as Amer-
igoniscus (or Caucasonethes) the material
considered herein was received with the
presumption that it represented a species in
that genus. Examination of the isopod
proved this assumption was wrong as evi-
denced by: (1) antenna 1 with short, stout
aesthetascs on the distal article, (2) pereo-
pod 7 propodus with distinct distal tuft of
setae, and (3) male pleopod 1| with vestigial
endopod. These characteristics eliminated
the species from Amerigoniscus (Vandel
1953, 1965, 1978; Schultz 1982, 1994).

Other North American genera for consid-
eration were listed variously in the First Di-
vision Trichoniscinae by Vandel (1965) or
Tribe Typhlotricholigiodini (Tabacaru 1993,
Schultz 1994): Brackenridgia, Typhlotri-

VOLUME 117, NUMBER 2

choligioides, Cylindroniscus and Mexicon-
iscus. The morphological characteristics
listed above for the Tumbling Creek Cave
isopod are exhibited by Brackenridgia and
Cylindroniscus. Schultz (1994) listed the
following characteristics of Cylindroniscus
that separate the genus from Brackenridgia:
(1) an elongate, semi-cylindrical body, (2)
antenna 2 that projects from the front of the
cephalon, and (3) uropod basis with elon-
gate endopod and exopod. Cylindroniscus
has been reported from Cuba and Mexico,
but is not known to occur in the United
States (Schultz 1970). Although the sepa-
ration of Brackenridgia and Cylindroniscus
is not entirely distinct, the Tumbling Creek
Cave trichoniscid seems most appropriately
placed in Brackenridgia. The ten species
now recorded from this genus occur from
Missouri to California and into Mexico
(Rioja 1950, 1951, 1955; Vandel 1965;
Schultz 1984; Dearolf 1953) as presented
in Fig. 1.

Materials and methods.—The isopods
were placed on a glass slide in a drop of
glycerin and appendages were dissected di-
rectly into the glycerin to produce tempo-
rary mounts. All drawings were made on a
Leica compound microscope with an opti-
cal drawing tube. After completion all ap-
pendages were then replaced in microvials
and stored in 70% ethanol. The geographic
coordinates for the type-locality were re-
corded with a Garman Map76 GPS. Tem-
perature readings in Tumbling Creek Cave
were taken with a Taylor digital thermom-
eter.

Family Trichoniscidae
Brackenridgia Ulrich 1902
Brackenridgia ashleyi, new species
Figs. 2—5

Caucasonethes.—Aley, 1975:1.

Caucasonethes sp.—Craig,
part].

Caucasonethes n. sp.—Gardner, 1986:15 [in
part, Tumbling Creek Cave record only];

1975:4 [in

177

U.S. Fish & Wildlife
66804.

Service, 2001:

Material examined.—Missouri: Taney
Co., Tumbling Creek Cave, 22 Apr 2001,
Catherine and Thomas Aley, 2.8 mm ho-
lotype 6 (USNM 1008288); same locality
and collectors, 2.3 mm paratype ¢ (USNM
1008289), 25 Nov 2001; same locality, Jul-
ian J. Lewis and David Ashley, 2.7 mm
paratype ¢ (USNM 1014382), 23 Apr
2003, same locality 21 Feb 1998, William
Elliott, 2.2 mm paratype d. The first three
specimens are deposited in the National
Museum of Natural History, Smithsonian
Institution, Washington, D.C. under catalog
numbers as noted. The Elliott collection is
deposited in the Enns Entomological Mu-
seum, University of Missouri, Columbia.

Description of male.—Eyeless, unpig-
mented, longest 2.8 mm. Body about 2.8
as long as wide, dorsal surface covered with
short, stout triangular spine-like setae.
Head, anterior margin biconcave, rostral
area broadly rounded. Pereonites with lat-
eral margins with short, stout triangular
spine-like setae and short setae; pereonite |
directed cephalad, 5—7 directed caudad.
Lateral margins of pleonites contiguous,
telson with posterior margin produced,
broadly rounded, with 4 small setae.

Antenna | with 5—6 short, stout aesthe-
tascs on distal article. Antenna 2 flagellum
with 5 indistinctly demarcated articles, dis-
tal article with longitudinally striated apical
organ. Mandibles with well developed mo-
lar process. Right mandible with plumose
setae between molar process and incisor, la-
cinia mobilis apically lobed. Left mandible
with two plumose setae between pars inci-
siva/lacinia mobilis and molar process.
Maxilla 1, endopod with 3 bladelike setae;
exopod with 8 spine-like setae (4 dominant
robust + 4 smaller). Maxilliped palp seg-
mentation indistinct, basal article relatively
narrow, not obscuring endite.

Pereopod 1, propodus with 6 subtrian-
gular spine-like setae along outer margin.
Pereopods 5—7 with row of stout spine-like

178 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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MEXICO

Distribution of Brackenridgia species and source of records: (1) B. ashleyi, (2) B. cavernarum (Texas

records—Vandel 1965, Mitchell and Reddell 1971; New Mexico record—Dearolf 1953, needs confirmation);
(3) B. reddelli (Vandel 1965, Mitchell & Reddell 1971); (4) B. sphinxensis (Schultz 1984); (5) B. heroldi (Vandel
1965); (6) B. bridgesi (Vandel 1965, Reddell 1981); (7) B. palmitensis (Mulaik 1960); (8) B. villalobosi (Vandel
1965, Reddell 1981); (9) B. acostai (Vandel 1965, Reddell 1981).

setae along distal margin of carpus and
merus (scales of Vandel 1965 or Schultz
1994). Pereopod 7 propodus with spine-like
setae in row On outer margin leading to
dense tuft of setae adjacent to junction of
dactyl.

Pleopod 1, exopod with small spinules
along lateral margin, tip simple, produced

into subtriangular structure with low, slight-
ly produced knobs subapically; endopod
vestigial, reduced to a subtriangular flange.
Pleopod 2 endopod thin, elongate, tapering
to a point; exopod about 0.3X length of en-
dopod, small, subovate with one apical set-
ule. Pleopods 3-5 undifferentiated. Uro-
pods about 1.5X length of telson, about

VOLUME 117, NUMBER 2

179

Fig. 2.
labrum and antennae, (c) antenna 1 with aesthetascs, (d) antenna 2, (e) uropod; B. reddelli, male from Valdina
Farms Sinkhole, Medina Co., Texas: (f) antenna 1 in situ.

0.3X length of pleon, endopod about 0.6
length of exopod, rami with 2—3 elongate
apical setae.

Etymology.—The species is named in
honor of Dr. David Ashley, of Missouri
Western State College, in recognition of his
years of outstanding effort monitoring the
ecosystem of Tumbling Creek Cave.

Distribution and ecology.—Brackenridg-
ia ashleyi is known only from the type-lo-
cality in the Springfield Plain of the Ozark
Plateau physiographic province in south-

Brackenridgia ashleyi, male from Tumbling Creek Cave, Taney Co., Missouri: (a) habitus, (b) head,

western Missouri. Tumbling Creek Cave
has 2,815 meters of mapped passages
formed in the predominantly dolomitic Jef-
ferson City Formation of Ordovician age.
The cave is on the property of the Ozark
Underground Laboratory and has two en-
trances: (1) the natural entrance marked
Bear Cave on the U.S. Geological Survey
Protem Quadrangle map at N36.55471,
W92.80275; and (2) an artificial entrance
marked Tumbling Creek Cave _ at
N36.54951, W92.80807. Tumbling Creek

180

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Brackenridgia ashleyi, male from Tumbling Creek Cave, Taney Co., Missouri: (a) mandibles (b)
maxilla 1, (c) maxilliped; B. heroldi, Crystal Cave, Sequoia National Park, California: (d) distal elements of
maxilliped.

Cave was dedicated by the U.S. Department
of the Interior as a Natural Landmark in
1980.

Tumbling Creek Cave has been the site
of detailed ecological studies concerning
the community associated with the bat gua-
no of the Gray bat Myotis grisescens (Mar-
tin 1980, Fletcher 1982). Martin (1980) re-
ported the air temperature of the cave was
14.4°C (+1°) and relative humidity at or
near 100%. Temperature measurements by
Lewis and Ashley on 23 April 2003 of the
substrate of the isopods were 14.7°C in the
East Passage and 14.3°C in the main stream
passage.

Martin (1980) reported 54 invertebrate
taxa occurring on guano, bat carcasses and
wood in Tumbling Creek Cave. Typical of
caves with Gray bat maternity colonies,
most surfaces between the bat roosts and

entrance are peppered with bat guano and
piles as much as a meter deep occur in
some areas. Martin found 49 taxa inhabiting
these guano piles, several of which were
predators, including the pseudoscorpion
Hesperochernes occidentalis, an unidenti-
fied harvestman, and beetles Platynus ten-
uicollis, Bembidion sp. and Atheta sp. In
contrast, the terrestrial isopods were never
found on guano nor bat carcasses, only
wood. Of the 14 species recorded from
wood by Martin, only B. ashleyi and the
millipeds Pseudopolydesmus pinetorum and
Scoterpes s. latu dendropus did not also oc-
cur on guano or carcasses. This habitat par-
titioning by these species excluded them
from the richest food source in the cave, but
eliminated significant predation pressure as
well. A fact not appreciated by Martin, who
did not identify the terrestrial isopods, was

VOLUME 117, NUMBER 2

181

Fig. 4. Brackenridgia ashleyi, male from Tumbling Creek Cave, Taney Co., Missouri: (a) pereopod 1, (b)

pereopod 7, (c) pereopod 7 propodus and dactylus.

that the exotic Haplophthalmus danicus
was also living on the wood and therefore
possibly competing with B. ashleyi.

The wood on which the isopods were
most easily observed consists of pieces of
pine boards placed in the cave to attract in-
vertebrates for viewing purposes (the cave
is operated among other things as an edu-
cational facility). The isopod presumably
feeds on microbial decomposers occurring
on the wood. As the presence of the wood
is artificial, under natural circumstances
Brackenridgia must live on the ubiquitous
mud banks. All of the isopods collected for
study herein were found associated with
wood on relatively dry mud banks along the
main stream passage or in the upper level
East Passage. When observed, Bracken-

ridgia moved about within an area of a few
square centimeters, with its antennae ac-
tively probing the environment when walk-
ing.

Life history.—Nothing is known of the
reproduction of B. ashleyi. In examining
material of other Brackenridgia two ovi-
gerous females of B. heroldi were found in
a collection from Hurricane Crawl Cave,
Sequoia National Park, California (collec-
tion date unknown). One was a 3.2 mm
specimen with four embryos in the brood
pouch. The other was 3.5 mm in length and
was carrying one 1.1 mm juvenile.

Discussion.—With the exception of the
humicolous B. heroldi, the species of
Brackenridgia are troglobites restricted to
karst areas isolated from one another by

182 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Brackenridgia ashleyi, male from Tumbling Creek Cave, Taney Co., Missouri: (a) pleopod 1, (b)
pleopod 2, (c) pleopod 3, (d) pleopod 4, (e) pleopod 5.

VOLUME 117, NUMBER 2

hundreds of kilometers of non-cavernous
rocks. Dispersal and gene flow between
populations is thus very unlikely, leaving
little doubt as to the speciation of these an-
imals in their respective karst islands. For
example, B. ashleyi in the Ozarks is sepa-
rated from its nearest relatives inhabiting
the karst of the Balcones Fault Zone in Tex-
as (B. cavernarum and B. reddelli) by over
800 kilometers (Fig. 1).

That not withstanding, the morphological
expression of this speciation is conservative
among the members of the genus. The sub-
triangular shape of the male first pleopod
exopod is similar in all species of the genus.
The species of Brackenridgia can be sepa-
rated from one another by relatively small
differences in the structures present at the
tip of this exopod. In the most primitive
species, e.g., B. cavernarum or B. sphinx-
ensis, the exopod tip is undifferentiated. A
variety of specializations into spinose or
digitiform structures occur in B. reddelli, B.
ashleyi and B. villalobosi, with the bi-spi-
nose pleopod | exopod tip (Vandel 1965)
of B. bridgesi presumably apomorphic. Al]-
though B. ashleyi is closest geographically
to B. cavernarum and B. reddelli in Texas,
the slight modification of the pleopod | ex-
opod tip is more similar to that of B. heroldi
in California.

Much work remains in the systematics of
Brackenridgia. In B. reddelli and B. sphinx-
ensis none of the mouthparts have been il-
lustrated. The latter species is known from
a single tiny dissected specimen so there is
little hope for a better understanding of the
species without additional collecting. Sim-
ilarly, B. palmitensis (Mulaik 1960) re-
mains unidentifiable (Vandel 1965) and at-
tempts to collect a male have been unsuc-
cessful (Reddell 1981). The regional vari-
ation reported by Vandel (1965) for B.
reddelli is suggestive of a cluster of closely
related species inhabiting the caves associ-
ated with the Balcones Fault Zone in Texas.
Confusion has also been created by Van-
del’s (1965) interpretation of the lateral
margin of the male pleopod 1 protopod of

183

B. villalobosi (illustrated by Rioja 1950 fig.
44) as the exopod tip.

Vandel (1965) published a key to the spe-
cies of the genus known at the time, while
Rioja (1955) included only the Mexican
fauna. I have updated these works to en-
compass nine members of the genus, in-
cluding the addition of B. sphinxensis and
B. ashleyi, but excluding B. palmitensis.
The identity of B. palmitensis remains ob-
scure, although Mulaik (1960 fig. 43) illus-
trated 8 short, stout aesthetascs on the distal
article of antenna 1. This characteristic sep-
arates this species from the majority of
Brackenridgia species, including B. ashleyi.

Key to Species of the Genus
Brackenridgia

la. Pereonites with prominent tubercles (fig.
6a), antenna | with elongate aesthetascs
B. acostai (caves, Chiapas, Mexico)
1b. Pereonites without prominent tubercles
(fig. 2a)
2a. Male pleopod 1 exopod tip undifferen-
tiated (fig. 6b)
2b. Male pleopod 1 exopod modified, taper-
ing to 1—2 processes (figs. 5a, 6c, d) (4)
3a. Length excluding antennae and uropods
<2 mm; antenna | distal article with 5
aesthetascs B. sphinxensis
(Sphinx Cave, Cochise County, Arizona)
3b. Length excluding antennae and uropods
>4 mm; antenna 1 distal article with
Isr GESINEAHSES 5 cccccnnn B. cavernarum
(caves, southcentral Texas, unconfirmed
in southeastern New Mexico)
4a. Male pleopod 1 exopod tapering to a
single point (figs. 5a, 6c)
4b. Male pleopod | exopod tip with two
processes (fig. 6d)
5a. Male pleopod 1 exopod tapering to a
digitiform process, antenna | aesthe-
tascs 8 B. villalobosi (caves, Veracruz)
5b. Male pleopod 1 exopod tapering to a
point (figs. 5a, 6c), antenna | aesthe-
tascs 5—6 (fig. 2c)
6a. Maxilliped palp basal segment broad,
obscuring endite (fig. 3d); male pleo-
pod 1 exopod without subapical knobs
(fig. 6c) B. heroldi
(humus and caves, southern California)

184 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6.

Brackenridgia acostai, Cueva del Ticho, Chiapas: (a) head and pereonites 1-3; B. cavernarum,

Kappelman Salamander Cave, Comal Co., Texas: (b) pleopod 1 exopod tip; B. heroldi, Crystal Cave, Sequoia
National Park, California: (c) pleopod 1 exopod tip; B. reddelli, Valdina Farms Sinkhole, Medina Co., Texas:

(d) pleopod | exopod tip.

6b. Maxilliped palp basal segment narrow,
not obscuring endite (fig. 3c); male ple-
opod 1 exopod with low, subapical
MOOS (UG, SA) scoososccvcvcs B. ashleyi
(Tumbling Creek Cave, Missouri)
7a. Male pleopod 1 exopod tapering to two
acute spines, antenna | aesthetascs 14
.. B. bridgesi (caves, northeastern Mexico)
7b. Male pleopod 1 exopod tapering to an
acute spine and a second brush-like
process (6d); antenna | aesthetascs 9—
10 (fig. 2f)
B. reddelli (caves, southcentral Texas)

Acknowledgments

I thank Mr. Thomas Aley, Dr. David
Ashley, Dr. William Elliott, Dr. John R.
Holsinger, Dr. Joan Jass, Dr. Brian Kensley,
Dr. William Muchmore, Dr. Christian
Schmidt, Ms. Marilyn Schotte, Dr. George

Schultz and Dr. Stefano Taiti for reading the
manuscript and making suggestions for its
improvement. The loan of specimens from
the collections of the Smithsonian Institu-
tion was kindly provided by Ms. Marilyn
Schotte and Dr. Brian Kensley. The descrip-
tion of B. ashleyi was funded by the U.S.
Fish & Wildlife Service, The Nature Con-
servancy and the Ozark Underground Lab-
oratory.

Literature Cited

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ratory Newsletter, 7(4)/8(1):1.

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1-72.

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Martin, B. J. 1980. The community structure of ar-
thropods on bat guano and bat carcasses in
Tumbling Creek Cave. Unpublished M.S. the-
sis, University of Illinois at Chicago Circle, 178
pp.

Mitchell, R. W., & J. R. Reddell. 1971. The inverte-
brate fauna of Texas caves. Pp. 35—90 in E. L.
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Mulaik, S. B. 1960. Contribucion el conocimiento de
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de Historia Natural 21(1):79—292.

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of Mexico, Guatamala, and Belize. Bulletin 27,
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coniscidos cavernicolas de México del género
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. 1951. Estudios Carcinol6gicos. XXVI. Des-

cripcion de Protrichoniscus acostai n. sp.

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del Instituto de Biologia 22(1):181—189.

. 1953. Estudios Carcinologicos. XXIX. Un

nuevo género de isOpodo triconiscido de la Cue-

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doba, Ver.—Anales del Instituto de Biologia

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1955. Triconiscidae cavernicolas de Méxi-

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. 1957. Estudios Carcinolo6gicos. XXXVI. Des-
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Schultz, G. A. 1970. Cylindroniscus vallesensis sp.
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. 1982. Amerigoniscus malheurensis, new spe-

cies, from a cave in western Oregon (Crustacea:

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Biological Society of Washington 95(1):89—92.

. 1984. Brackenridgia sphinxensis n. sp. from

a cave with notes on other species from Arizona

and California (Isopoda, Oniscoidea).—South-

western Naturalist 29(3):309-319.

. 1994. Typhlotricholigioides and Mexiconiscus
from Mexico and Cylindroniscus from North
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763-770.

Tabacaru, I. 1993. Sur la classification des Trichonis-
cidae et la position systématique de Thauma-
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cea, Isopoda, Oniscidea).—Travaux Institute
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fauna of Texas.—Transactions of the American
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Vandel, A. 1950. Campagne spéleologique de C. Bo-
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(1928). 14. Isopodes terrestres recueillis par C.
Bolivar et R. Jeannel (1928) et le Dr. Henrot
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. 1953. A new terrestrial isopod from Oregon,

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. 1965. Les Trichoniscidae cavernicoles (Iso-

poda Terrestria: Crustacea) de L- Amérique du

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. 1978. Les espéces appartenant au genre Amer-

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Naturelle de Toulouse 113:303-—310.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(2):186-198. 2004.

New species and records of Bopyridae (Crustacea: Isopoda) infesting
species of the genus Upogebia (Crustacea: Decapoda: Upogebiidae):
the genera Orthione Markham, 1988, and Gyge Cornalia &
Panceri, 1861

John C. Markham

Arch Cape Marine Laboratory, Arch Cape, Oregon 97102-0105 U.S.A.

Abstract.—Two bopyrid genera whose species parasitize only Upogebia spp.
are reviewed and revised. Orthione Markham, 1988, heretofore known only
from its type-species, O. furcata (Richardson, 1904), is rediagnosed and en-
larged. Orthione griffenis, new species, infests Upogebia pugettensis (Dana,
1852) in Oregon, U.S.A. Orthione mesoamericana, new species, infests U.
spinigera (Smith, 1871) on the Pacific coasts of Costa Rica and Colombia. The
genus Metabopyrus Shiino, 1939, is incorporated into Gyge Cornalia and Pan-
ceri, 1861, which is rediagnosed, and a key is given to the four known species.
Gyge ovalis (Shiino, 1939), formerly Metabopyrus ovalis Shiino, 1939, is re-
described on the basis of new material found infesting U. edulis Ngoc-Ho &
Chan, 1992, in Taiwan, a new host and geographical record.

In a recent compilation of the known bo-
pyrid parasites of thalassinidean decapod
crustaceans throughout the world (Mark-
ham 2001), I listed 26 species of the genus
Upogebia known to harbor a total of 27
species of bopyrid isopods. Since then, ad-
ditional material of parasites of species of
Upogebia has become available for exami-
nation or has been reliably reported to me.
It includes two new species of Orthione
Markham, 1988, described herein, as well
as new host and geographic records for Me-
tabopyrus ovalis Shiino, 1939, which is re-
described and reassigned to the genus Gyge
Cornalia & Panceri, 1861.

Materials and Methods

Host specimens bearing parasites, or par-
asites that had been removed from their
hosts, have become available for study from
various sources over a period of several
years. Some were already in scientific col-
lections, while others are being newly do-
nated to institutions housing such collec-
tions. Those institutions are indicated thus:

Museo de Zoologia, Universidad de Costa
Rica, MZUCR; Naturhistorisches Museum,
Wien, Austria, NHMW; Naturhistoriska
Rijksmuseet, Sweden, SMNH; and Natural
History Museum, Smithsonian Institution,
USNM.

Results

Family Bopyridae Rafinesque-Schmaltz,
1815
Subfamily Pseudioninae Codreanu, 1967
Genus Orthione Markham, 1988

Type-species, by original designation,
Pseudione furcata Richardson, 1904. Num-
ber of previously known species: 1, O. fur-
cata (Richardson, 1904), infesting Upoge-
bia affinis (Say, 1818), Massachusetts to
North Carolina, U.S.A.

Revised generic diagnosis, based on
three known species.-Female. Body outline
oblong, about twice as long as wide, sides
nearly parallel, axis only slightly distorted,
all body regions and segments distinct dor-
sally. Head deeply set into pereon, its an-

VOLUME 117, NUMBER 2

terior margin completely covered by frontal
lamina and forming continuous curve with
pereon; maxilliped lacking palp; barbula
with single prominent lanceolate process on
each side, rarely minute process lateral to
it. Pereopods slightly to much larger pos-
teriorly; oostegites generously enclosing
brood pouch, first one with prominent but
unadorned internal ridge, no posterolateral
point. Pleon of six pleomeres, much broad-
er than long, final pleomere deeply enclosed
by fifth; five pairs of biramous pleopods
and similar uniramous uropods completely
covering lateral margins and all but center
of dorsal surface of pleon, endopodites of
first pair larger and medially extended.

Male. Body oblong, at least three times
as long as wide; all body regions and seg-
ments distinct. Head nearly semicircular;
second antennae prominently extended. Pe-
reopods relatively small, though overall
larger and with smaller dactyli posteriorly,
all clustered medially. Pleon about % of to-
tal body length, of six pleomeres; pleopods
absent or as low incomplete oval uniramous
flaps; final pleomere largely surrounded by
fifth, ending in pair of uniramous flaplike
uropods.

Hosts. All in genus Upogebia.

Key to Three Species of Orthione, Based
on Mature Females

1. Pereopods with propodal cups receiving
tips of dactyli, bases produced into large
carinae; pleopodal rami somewhat ovate

O. griffenis new species [Oregon, U.S.A.].

—. Pereopods lacking propodal cups to re-
ceive tips of dactyli, bases lacking cari-
nae; pleopodal rami lanceolate

2. Head much broader than long, with mi-
nute barbular projection lateral to main
process; final pleomere visible dorsally

... O. mesoamericana new species [Pacific
coast from Costa Rica to Colombia].

—. Head about as broad as long, only single
process on each side of barbula; final
pleomere more or less hidden dorsally

O. furcata (Richardson, 1904)
[Atlantic coast of U.S.A.].

187

Orthione griffenis, new species
Figs. 1-3

““New species .. . [of] . . . Orthione.” —Da-
vid, 2001:6.

Material examined.—Infesting Upogebia
pugettensis (Dana, 1852). Collected and
hosts det. by B. D. Griffen. Mudflats, Idaho
Inlet, Yaquina Bay, Oregon, USA, 44°
35.4'N, 124°01.5'W, unspecified date,
2000: 1 2, holotype, USNM 1008784, 1 6,
allotype, USNM 1008785. Same locality,
23 June 2001, 9 22, 7 Sd, paratypes,
USNM 1008786. Collected and hosts det.
by T. H. DeWitt: Riverbend, Yaquina Bay,
Oregon, 12 November 1999, Sample
S59MDF-U31: 1 @, dextral, 1 35, paratypes,
USNM 1008787. Idaho Flat, Yaquina Bay,
Oregon, 24 September 1999, Sample
59UU104M: 1 2°, sinistral, immature, para-
type, USNM 1008788.

Description.—Holotype female (Fig. 1).
Length 11.0 mm, maximal width 9.2 mm,
head length 2.2 mm, head width 2.1 mm,
pleon length 2.9 mm; distortion 15°, dex-
trally. Outline oval, nowhere abruptly
broader or narrower; all body regions and
segments distinct. No pigmentation (Fig.
1A, B).

Head almost square, deeply set into pe-
reon, its anterior edge continuous with per-
eonal margin. Distinct rather long frontal
lamina extending completely across front of
head but not beyond its sides. First anten-
nae long and extended beyond margin of
head, of 5 articles, distal two terminally se-
tose; second antennae greatly reduced, of 1
to 3 articles (Fig. 1C). Barbula (Fig. 1D)
with single long falcate process on each
side, central region entire. Maxilliped (Fig.
1E) subtriangular, lacking palp, with plec-
tron short and blunt; anterior article nearly
rectangular, much longer than triangular
posterior article.

Pereon widest across pereomeres 4—5.
Pereomere | curved strongly around head,
it and pereomere 2 markedly concave an-
teriorly; pereomeres 3—4 nearly straight
across; pereomeres 5—7 concave posterior-

188 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1.
side of barbula. E. Right maxilliped. EF Right oostegite 1, external. G. Same, internal. H. Right pereopod 1. I.
End carpus and dactylus of same. J. Right pereopod 7. K. End carpus and dactylus of same. Scale: 3.60 mm
for A, B, E—G; 1.20 mm for C; 2.40 mm for D; 1.00 mm for H, J; 0.4 mm for I, K.

ly. Pereomere | shortest, all others about
same length. Shallow elongate depression
near each side of dorsal surface of pereo-
meres 2—5. Pereomeres 1—6 bordered by
coxal plates on long side of body, those on
pereomeres 5—6 with crenulate margins;
smaller coxal plates on short sides of per-
eomeres | and 5. First oostegite (Fig. 1E
G) subcircular, its articles of about same
size, separated by deep but narrow groove
externally; internal ridge smoothly curved
and lacking ornamentation. Oostegites 2—5
all long and relatively slender, each reach-
ing about % of distance across brood pouch
and together completely enclosing it. Fifth
oostegite with fringe of long setae along
posterior margin. Pereopods (Fig. 1H, J)
with all articles distinct, more than doubling
in size posteriorly; all bases produced into
broadly rounded carinae; short comma-

Orthione griffenis, new species. Holotype female. A. Dorsal. B. Ventral. C. Right antennae. D. Right

shaped dactyli (Fig. 11, K) with sharp tips
fitting into lip-like receptacles on distal cor-
ners of propodi; all carpi densely setose dis-
tally.

Pleon of 6 pleomeres, all sharply concave
posteriorly, its posterior margin almost
straight across sides of all pleomeres. Ven-
trally, sides of pleon completely covered by
5 pairs of overlapping lanceolate uniramous
lateral plates and uropods, and its middle
region equally covered by 5 pairs of bira-
mous pleopods, all of their rami of size and
structure similar to lateral plates, except
that endopodites of first pleopods much
larger than others and crossing each other
in middle of pleon.

Allotype male (Fig. 2). Length 8.0 mm,
maximal width 3.0 mm, head length 1.1
mm, head width 1.9 mm, pleon length 2.7
mm. Body straight on both sides, rounded

VOLUME 117, NUMBER 2

189

Fig. 2.
antenna 2. E. Right pereopod 1. EK End carpus and dactylus of same. G. Right pereopod 7. H. End carpus and
dactylus of same. I. End of pleon, ventral. Scale: 2.00 mm for A, B; 0.84 mm for E, G; 0.24 mm for C, D,
EH, I.

at both ends. All body regions and seg-
ments distinctly separated (Fig. 2A, B). No
pigmentation.

Head almost semicircular, markedly nar-
rower than any pereomeres. Antennae (Fig.
2C, D) of 3 and 5 articles, respectively,
both pairs directed laterally; first antennae
distally setose; second antennae extending
beyond margins of head.

Pereon broadest across pereomere 6, but
only slightly so. Most pereomeres deeply
separated by anterolateral notches. Low
broad middorsal ridge along full length of
pereon. Pereopods (Fig. 2E, G) relatively
small, clustered medially under body; all of
about same size, but their dactyli smaller
posteriorly. All propodi bearing corneous
ridges on surfaces met by folded dactyli
(Fig. 2F H); distal corner of propodus of

Orthione griffenis, new species. Allotype male. A. Dorsal. B. Ventral. C. Right antenna 1. D. Left

pereopod 7 extended into receptacle for end
of dactylus (Fig. 2G, H); all carpi distally
setose.

Pleon of 6 distinct pleomeres, each nar-
rower than that before it. Five pairs of dis-
tinct but sessile oval pleopods, progressive-
ly smaller posteriorly. Final pleomere (Fig.
21) deeply set into fifth, produced into pair
of stubby pointed uniramous uropods, their
margins bearing many short setae.

Remarks on paratypes (Fig. 3).—Of the
nine paratype females, five are dextrally
distorted, as is the holotype, three are sinis-
tral, and one is too immature for assess-
ment. They range in length from 6.2 mm to
18.8 mm and in width from 2.7 to 13.3 mm
(Fig. 3A—G). Most of the mature females
have the endopodites of the first pair of ple-
opods prominently visible (Fig. 3A). One

190 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3.
pleon, ventral. C. End of pleon, ventral. D. Immature, dorsal. E. Same, ventral. E Late larva, dorsal. G. Same,
with male attached, ventral. H. Pleon, ventral. I. Pleon, ventral. J. Immature, end of pleon, ventral. Scale: 4.20
mm for C; 2.10 mm for A, B, D, E; 1.00 mm for F—J.

has long slender extended uropods (Fig.
3B); one mature female has the fifth pleo-
mere deeply separated from the preceding
one (Fig. 3C), as does one immature female
(Fig. 3D) In a very immature female (Fig.
3E G), all oostegites are absent, and pleo-
pods are only uniramous flaps. At a slightly
later stage, an immature female (Fig. 3D,
E) has rudimentary oostegites and small but
distinctly pleopods, the endopodites of the
first pair already pointing medially.

The seven paratype males (Fig. 3G—J) are
6.1 to 10.7 mm in length, and 2.0 to 3.2
mm in width. All are very similar to the
allotype, but one has a more elongate pleon,
its pleomeres deeply separated (Fig. 3H);
one has a deformed fifth pleomere (Fig. 31);
and one has only very tiny traces of pleo-
pods (Fig. 3J).

Etymology.—The name griffenis, geni-
tive singular of a name regarded as a Latin

Orthione griffenis, new species. Paratypes. A-G, females. H—J, males. A. Pleon, ventral. B. End of

third-declension noun, is selected to honor
Blaine D. Griffen, who, in the course of an
ecological study of the host, Upogebia pug-
ettensis, collected most of the material,
called it to my attention and furnished it
and collection data for this description.
Remarks.—Upogebia pugettensis has
been reported many times as the host of
Phyllodurus abdominalis Stimpson, 1857,
which attaches to its abdomen throughout
the host’s geographic range from British
Columbia to central California (Williams
1986, Markham 1992), though the coast of
Oregon remains a gap in the known distri-
bution of P. abdominalis. This is the first
record of infestation of U. pugettensis by a
branchial bopyrid. So far this new species,
Orthione griffenis, is known only from Ya-
quina Bay, Oregon, in the middle of the
range of U. pugettensis, but there it appears
to be fairly common. Upogebia pugettensis

VOLUME 117, NUMBER 2

191

Fig. 4.
D. Right maxilliped. E. Plectron of same. FE Right side of barbula. G. Right oostegite 1, external. H. Same,
internal. J. Right pereopod 1. I. J. Right pereopod 7. Scale: 2.00 mm for A, B, D, F—H; 1.00 mm for E; 0.36
mm for C, I, J.

attains densities of up to 300 burrows per
square meter in Yaquina Bay and occurs
throughout the lower region of that estuary
(Griffen 2002). For comparison of Orthione
griffenis with other species in the genus, see
the remarks on the following species.

Orthione mesoamericana, new species
Figs. 4, 5

““Bopyrid Isopod.’’—Holthuis, 1952:9
[Buenaventura, Colombia; infesting Upo-
gebia spinigera (Smith)].

Material examined.—Infesting Upogebia
spinigera (Smith, 1871). Puerto Jiménez,
Golfo Dulce, Puntarenas, Costa Rica,
08°32'30"N, 83°18'20"W, 13 January 1977.
1 2, holotype, 1 3d, allotype, MZUCR
2194-04. Lund University Chile Expedi-
tion. Buenaventura, Colombia, 03°77'N,
77°02'W, on beach, under lump of clay, 30

Orthione mesoamericana, new species. Holotype female. A. Dorsal. B. Ventral. C. Right antennae.

August 1948. H. Brattstrom and E. Dahl,
colls., L. B. Holthuis det. of host: 1 ¢, para-
type, SMNH 5325.

Description.—Holotype female (Fig. 4).
Length 6.5 mm, maximal width 5.4 mm,
head length 1.3 mm, head width 1.8 mm,
pleon length 2.2 mm. Body axis distortion
4°, dextrally. Body nearly oval, all regions
and segments distinct. No pigmentation ex-
cept for dark eyespots (Fig. 4A, B).

Head slightly convex anteriorly, nearly
semicircular posteriorly, deeply embedded
into pereon. Frontal lamina very short but
extending completely across anterior of
head but not beyond. Eyes as prominent
slender slashes near anterolateral corners of
head. Antennae (Fig. 4C) well-developed,
of 3 and 6 articles, respectively, first ones
extending plainly beyond front margin of
head. Barbula (Fig. 4F) with two projec-
tions on each side, outer one minute, inner

192 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

one extended and curved, both with entire
margins; no decoration in middle of bar-
bula. Maxilliped (Fig. 4D) suboval, lacking
palp but with notch in anterior margin;
plectron (Fig. 4E) small and slender, point-
ing anteriorly.

Pereon broadest across pereomere 4, all
pereomeres distinctly separated laterally.
Tergal plates on both sides of pereomeres
1—4, though only faint on first one. Ooste-
gite 1 (Fig. 4G, H) with nearly parallel
sides, slightly convex ends, both segments
about equally wide and long, separated by
deep external groove, no posterolateral pro-
jection; internal ridge unornamented, pro-
duced into long right-angled flap. Oostegi-
tes 2—5 overlapping and reaching nearly
across and completely enclosing brood
pouch. Pereopods (Fig. 41) more than dou-
bling in size posteriorly, but none extending
beyond body margin; all dactyli short and
fairly blunt and retracting into anterior
notches of distally extended propodi; carpi
all sparsely setose distally, meri and carpi
of anterior pereopods fused.

Pleon of 6 distinct pleomeres, final one
deeply embedded in preceding one. Five
pairs of biramous pleopods, uniramous lat-
eral plates and uniramous uropods. Endop-
odites of first pair of pleopods large, inflat-
ed and extending medially, touching each
other and overlapping fifth oostegites an-
teriorly. Other pleopodal rami, lateral plates
and uropods all similar to each other, all as
lanceolate flaps with entire edges, com-
pletely covering margins of pleomeres.

Allotype male (Fig. 5).—Length 3.3 mm,
maximal width 1.2 mm, head length 0.3
mm, head width 0.9 mm, pleon length 1.0
mm. Sides of body nearly parallel, rounded
at each end. All body regions and segments
distinctly separated. No pigmentation (Fig.
5A, B).

Head semicircular, lacking eyes. Anten-
nae (Fig. 5C) prominent, first ones of 3 ar-
ticles, second ones of 6 or 7 articles; second
antennae extending far beyond margins of
head.

Pereomeres separated by anterior notches

reaching inward nearly % of body width.
Pereopods (Fig. 5D, E) all of nearly same
size, all their articles distinct; dactyli of pe-
reopods | and 2 long and sharply pointed,
others short and blunt; carpi of pereopods
5—7 much longer than others.

Pleon of 6 pleomeres, first one as wide
as pereon, others tapering rapidly posteri-
orly. Pleopods (Fig. 5F) as sessile plates
fairly conspicuous on pleomere 1, much
fainter on pleomeres 2—5, absent behind.
Sixth pleomere embedded in fifth, produced
posteriorly into blunt clublike uropods ex-
tending rearward unequal distances, both
sparsely fringed by minute setae.

Comparison of paratype male.—The oth-
er male is considerably larger, with these
dimensions: length 4.2 mm, maximal width
1.4 mm, head length 0.6 mm, head width
0.9 mm, pleon length 1.7 mm. It is the same
as the allotype in all respects except that
both second antennae are 7-articled, and its
uropods are equal in length.

Etymology.—Adjective mesoamericana
selected to indicate the known range of the
new species, along the Pacific coast of Cen-
tral America.

Remarks.—The paratype male from Co-
lombia is the unidentified bopyrid reported
by Holthuis (1952). Because there was no
female accompanying it, it could not be
identified until the allotype was described
here. The hosts are the same species in both
collections.

Comparison of three known species of
Orthione.—Females. One important addi-
tion to the original generic diagnosis
(Markham 1988) is that the endopodites of
the first pair of pleopods are markedly en-
larged and medially directed in a manner
highly distinctive for Orthione. This is the
case in the type-species, O. furcata, as well
as in the two new species, but I did not
recognize its importance as a diagnostic
character earlier. Of the two new species,
O. mesoamericana is much more similar to
O. furcata than is O. griffenis. Females of
the first two species have heads wider than
long, bearing slit-shaped eyes, and the in-

VOLUME 117, NUMBER 2

193

Fig. 5.
Right pereopod 1. E. Right pereopod 7. E Right pleopods 1. G. Pleomeres 5, 6, ventral. Scale: 1.4 mm for A,
B; 0.5 mm for C-G.

ternal ridge of the first oostegite is produced
into a broad angled flap reaching far pos-
teriorly, though that of O. mesoamericana
is less extended. Also, females of both of
those species have very slender pleopodal
appendages, those of O. mesoamericana be-
ing relatively somewhat broader. All pereo-
pods of O. furcata are about the same size,
while those in the two new species more
than double in size posteriorly. The minute
flap lateral to the large projection on the
barbula is unique to O. mesoamericana. Fe-
males of O. griffenis are distinctive in hav-
ing heads longer than broad, margins of
coxal plates crenulate rather than entire,
pereopodal bases carinate, and pleopodal
appendages ovate rather than lanceolate.
Males. All three species are very similar.
Pereopods of O. furcata are much longer
posteriorly, while those of the two new spe-
cies are of nearly the same size throughout.

Orthione mesoamericana, new species. Allotype male. A. Dorsal. B. Ventral. C. Right antennae. D.

Similarly, uropods of O. furcata are much
smaller than in either of the other two spe-
cies. In O. mesoamericana, second anten-
nae are greatly extended, and pleopods are
mostly absent, in contrast with the other
two species. The head of O. furcata has a
medial region extending posteriorly, that of
O. griffenis is smoothly rounded posterior-
ly, and that of O. mesoamericana is straight
posteriorly.

Genus Gyge Cornalia & Panceri, 1861

Type-species, by monotypy, Gyge bran-
chialis Cornalia & Panceri, 1861.

Revised diagnosis.—Female. Body oval
to squarish, at least %4 as broad as long,
body axis only slightly distorted either dex-
trally or sinistrally, angle of distortion far
forward. Head deeply set into pereon, its
sides diverging slightly to greatly anterior-

194

ly, frontal lamina completely covering an-
terior, its posterior end rounded to pointed.
Eyes usually absent. Antennae reduced.
Barbula with two lateral processes on each
side, they and middle region with deeply
digitate margins. Maxilliped usually nearly
straight across anterior margin, lacking
palp, with slender forward-pointing plec-
tron and at most only small posterior point.
Pereon broadest across pereomere 4,
smoothly rounded both ways, sides of first
4 or 5 pereomeres covered by conspicuous
coxal plates. Pereopods all equally small,
anterior ones with meri and carpi fused, ba-
ses large and often carinate. First oostegite
produced onto long slender and usually
curved posterior point. Other oostegites
narrowly pointed and incompletely enclos-
ing brood pouch. Pleon of 5 pleomeres, fi-
nal one usually notched posteriorly. Three
or four pairs of reduced biramous (or, intra-
specifically, uniramous) pleopods not ex-
tending beyond pleonal edges, their leaflike
rami all of same size and generally with
digitately divided margins. Uniramous uro-
pods tiny to quite large, flaplike, extended
posteriorly and with entire margins.

Male. Body long and slender, its head
well-extended and separated from pereon,
with or without eyes. Antennae well-devel-
oped. Pereopods uniformly small, first two
with proportionately longer dactyli, all with
fused meri and carpi. No midventral tuber-
cles. Pleon of 6 pleomeres, final one em-
bedded in fifth. Pleopods absent or as ses-
sile oval scars. No uropods.

Hosts: In genus Upogebia. Four species
known, from Britain through Mediterranean
to Black Sea; New Zealand; Japan and Tai-
wan; and Thailand.

Discussion.—I am hereby incorporating
Metabopyrus Shiino, 1939, and its two spe-
cies, the type-species M. ovalis Shiino,
1939, and M. irregularis Markham, 1985,
into Gyge Cornalia & Panceri, 1861, which
contained two species, G. branchialis Cor-
nalia & Panceri, 1861, and G. angularis
Page, 1985. The new diagnosis above is
based on all four species. Bourdon (1968:

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

151) observed that “Ce genre [Gyge] res-
semble beaucoup, en vue dorsale, a Meta-
bopyrus Shiino (1939), également parasite
d’Upogebia ...” Similarly, Page (1985:
196) asserted that “... Gyge and Metabo-
pyrus should be united,” but he did not for-
mally make such a combination. In addition
to the four species herein included in Gyge,
two originally in Gyge and two in Meta-
bopyrus, four other species have been cited
as members of the genus, but all of them
are either synonyms of G. branchialis or
now considered to belong to other genera.

The date of publication of the paper by
Cornalia & Panceri (1861), in which they
established the genus Gyge and described
its type-species G. branchialis, has been
subject to some confusion. Bate & West-
wood (1868) and Richardson (1905) listed
the date as 1861, while Bonnier (1900) and
Bourdon (1968) cited it as 1858. Reference
to the original publication indicates that,
while the volume in which the report ap-
peared was for the year 1858, it actually
appeared in 1861. Thus I am citing the date
of publication for both the genus Gyge and
its type-species Gyge branchialis as 1861.

Key to Four Species of Gyge Cornalia &
Panceri, 1861, Based on Mature Females

1. Body smoothly rounded, oval; body axis
distorted more than 30°; final pleomere
extending farther rearward than any oth-
er pleomeres

—. Body with indistinct corners, subrectan-
gular; body axis distorted less than 15°;
final pleomere at least partly embedded
in fifth pleomere and exceeded by one or
more of other pleomeres

2. Long sides of pereomeres distinctly set
apart by extended posterolateral angles;
posterior margin of pleon entire, large
uropods not visible in dorsal view ....
PRT ce ee ere whee oak G. irregularis

(Markham, 1985), n. comb. [Thailand]

—. Long sides of pereomeres continuously
curved; posterior margin of pleon deeply
cleft, revealing minute uropods in dorsal
VIEW") 3 Payee tee eens Sie G. branchialis

Cornalia & Panceri, 1861 [Europe]

VOLUME 117, NUMBER 2

3. Body segments all distinctly separated;
margins of barbula projections digitately
subdivided; internal ridge of oostegite |
cisitatem ere G. ovalis (Shiino, 1939),

n. comb. [Japan, Korea, Taiwan]

—. Body segments only obscurely separat-
ed; margins of barbula projections
smooth; internal ridge of oostegite |
smooth G. angularis Page, 1985

[New Zealand]

Gyge branchialis Cornalia & Panceri,
1861

Abbreviated synonymy. (See Bourdon,
1968, for complete synonymy to 1968.)

Gyge branchialis Cornalia & Panceri, 1861:
87-111; plates I, II [Estuary of Venice,
Italy; infesting Upogebia pusilla (Petag-
na, 1792)].—Bourdon, 1968:147, 151—
159, 169, 322, 410; figs. 28-32; tables
23, 24 [synonymy, summaries of previ-
ous accounts, including records from
Britain and Channel Islands through
France to Adriatic and Black Seas, in-
festing U. deltaura Leach, 1815, U. pus-
illa and U. stellata (Montagu, 1808); re-
description. Arcachon, France, and Na-
poli, Italy; infesting U. pusilla. Roscoff,
France, and Plymouth, England; infesting
U. deltaura. Roscoff, France; infesting U.
stellata. Jersey, Channel Islands; no
host].—Restivo, 1968:506 [Napoli,; in-
festing U. pusilla|.—Restivo, 1975:152,
153, 161-163; table | [Golfo di Napoli;
infesting U. pusilla; study of hyperpara-
sitism by Paracabirops marsupialis (Car-
oli, 1953)].—Dworschak, 1988:68 [Gra-
do, north Adriatic Sea, Italy; near Trieste,
Italy; Rovinj, Slovenia; infesting U. pus-
illa|.—Astall et al., 1996:821—823: table
1 [Clyde Sea, Scotland, and Irish Sea; in-
festing U. deltaura and U. stellata. Ar-
cachon Basin, France; infesting U. pus-
illa).

Gyges [sic] branchialis.—Grube, 1864:77
[Lussin Island, Croatia, Adriatic Sea; in-
festing U. pusilla).

Gyge galatheae Bate & Westwood, 1868:
225-229 [Guernsey, Channel Islands, in-

195

festing Galathea squamifera Leach, 1814
{subsequently reidentified as Upogebia
stellata by Norman, 1905:86}].

Not Gyge branchialis var. arcassonensis
Carayon, 1943:46—47 [=Progebiophilus
euxinicus (Popov, 1929)].

Material.—All identified and reported by
Peter Dworschak.—Infesting Upogebia
pusilla. Punta Spin, Grado, Adriatic Sea, It-
aly, 45°40'’N, 13°23’E, D. Abed-Navandi
coll., August 2000. 1 2 (ovigerous), 1 d,
NHMW 19521. Infesting U. tipica (Nardo,
1868), off Isola Rossa, Rovinj, Croatia,
Adriatic Sea, 45°05'N, 13°40’E, 18 m, D.
Abed-Navandi coll. 4 July 2000, P. Dwor-
schak det., | 2, NHMW 19523.

Remarks.—This is the first record of bo-
pyrid infestation of Upogebia tipica, and
thus a new record for Gyge branchialis.
Gyge branchialis is already known from the
Croatian coast of the Adriatic Sea, and it
does not need further redescription beyond
the detailed accounts presented by Bonnier
(1900) and Bourdon (1968). As indicated in
the synonymy above, G. branchialis has
been reported many times from Britain
through the Mediterranean to the Black Sea
as a parasite of three other species of Upo-
gebia.

Gyge ovalis (Shiino, 1939), new
combination
Fig. 6

Metabopyrus ovalis Shiino, 1939:88—91;
figs. 7, 8 [Hakata Bay, Kytsyt, Japan;
infesting Upogebia major (de Haan,
1839) {subsequently corrected to U. is-
saeffi Balss, 1913}].—Shiino, 1958:48—
49, fig. 10 [unknown specific locality, Ja-
pan; infesting unknown host; further de-
scriptive notes].—Codreanu, 1941:
140.—Codreanu, 1961:140; fig. 1.—
Codreanu & Codreanu, 1963:283.—
Shino, 1972:7.—Markham, 1982:340.—
Markham, 1985:14.—Page, 1985:196.—
Kim & Kwon, 1988:199, 201—203, 220;
fig. 2 [Komso, southwest Korea; infesting
U. major|.—Markham, 2001:198, 201;

196 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

at aS)
G

‘a
i.

{2
eo
cae
3
N

Fig. 6. Gyge ovalis (Shiino, 1939), new combination. A—L, female. M—S, male. A. Dorsal. B. Ventral. C.
Left antennae. D. Left side of barbula. E. Left maxilliped, external. EK Plectron of same. G. Right pereopod 1.
H. Distal end of same. I. Left pereopod 7. J. Distal end of same. K. Left oostegite 1, external. L. Same, internal.
M. Dorsal. N. Ventral. O. Left antenna 1. P. Left antenna 2. Q. Left pereopod 1. R. Right pereopod 7. S. End
of pleon, ventral. Scale: 4.00 mm for A, B, K, L; 1.93 mm for D, E, M, N; 1.00 mm for C, F; 0.88 mm for G,

I; 0.35 mm for H, J, Q—S; 0.18 mm for O, P.

tables 1, 2.—Itani et al., 2002:72; fig. la9
[Yamaguchi Bay, Seto, Inland Sea, Japan;
infesting U. major, study of response to
host’s molting].

Material examined.—Infesting Upogebia
edulis Ngoc-Ho & Chan, 1992. Shan-kong
mudflat, Chang-Hua County, southwest Tai-
wan, 24°06'25"N, 120°25'30"E, Tin-Yam
Chan collector and det. of host: 1 2, 1 d,
USNM 1008790.

Descriptive notes.—Gyge ovalis has now
been collected five times, but no lot has
been large. The original description (Shiino
1939) consisted of two pairs, the next two
collections (Shino 1958, Kim & Kwon
1988) were single females, and the present
material is one pair. The size of the most
recent Japanese collection (Itani et al. 2002)
was not indicated; the photograph in that

report, derived from a frame of a videotape
which was published only to show the fe-
male’s orientation on its host, lacks recog-
nizable details. Variations among the spec-
imens are slight, but all are noticeably dif-
ferent. The present female (Fig. 6A—L)
most resembles the figured syntype in pro-
portions and shapes of body parts, though
it lacks the prominent tergal plates seen on
the long side of pereomeres 1-3 of all pre-
viously recorded females (Fig. 6A, B). The
body of one female (Shiino 1958) is pro-
portionately shorter, and another (Kim &
Kwon 1988) lacks the posterior notch on
the final pleomere. The barbula (Fig. 6D) is
the same as in other females. The maxilli-
ped (Fig. 6E) is less distinctly segmented
than previously seen. The propodus of the
first pereopod (Fig. 6G, H) is produced into

VOLUME 117, NUMBER 2

a helmet-like shape not previously seen.
The first oostegite (Fig. 6K, L) is very
much like that reported by Shiino (1958),
while the one from Korea (Kim & Kwon
1988) was straight posteriorly. The male
(Fig. 6M-—S) has a much more extended
head and an embedded final pleomere (Fig.
6S), in contrast to the figured syntype (Shi-
ino 1939), whose head was little longer
than any pereomere, and whose final pleo-
mere was extended behind the preceding
one.

Remarks.—The present material repre-
sents both a new host, Upogebia edulis, and
new locality, Taiwan, for Gyge ovalis, al-
though Tin-Yam Chan (pers. comm.) re-
ports that it is commonly collected there.
Gyo Itani (pers. comm.) informs me that he
has found Gyge ovalis infesting five differ-
ent species of Upogebia in Japan, although
so far there are published records of only
two host species there.

Acknowledgments

Tin-Yam Chan, National Taiwan Ocean
University, provided material of Gyge oval-
is that he had collected and information
about it. Peter C. Dworschak, NHMW, pro-
vided information about his collections of
G. branchialis and granted me permission
to report on them. Christer Erséus and Kar-
in Sindemark, SMNH, lent the paratype
male of Orthione mesoamericana, to which
Lipke B. Holthuis, Nationaal Natuurhisto-
risch Museum, The Netherlands, had re-
ferred me. Blaine D. Griffen and Theodore
H. DeWitt, Oregon State University, col-
lected and furnished the type material of O.
griffenis and provided information on its
collection. Gyo Itani, Seto Marine Labora-
tory, Japan, sent me reprints of his papers
and provided information about collections
he had made. Becky S. Jordan, Iowa State
University Library, confirmed the date of
publication of the paper by Cornalia & Pan-
ceri (1861). Nguyen Ngoc-Ho, Muséum
National d’Histoire Naturelle, Paris, con-
firmed the current usage of names of Upo-

197

gebia spp. Marilyn Schotte, USNM, provid-
ed essential curatorial services and infor-
mation on collections, lent much material
for examination and description, furnished
elusive references and information about
them. Rita Vargas, MZUCR, lent type-spec-
imens of O. mesoamericana and furnished
details of their collection. Three anonymous
reviewers provided helpful remarks.

This report is a scientific contribution of
the Arch Cape Marine Laboratory (number
27) and of the College of Oceanic and At-
mospheric Sciences, Oregon State Univer-
sity.

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Three new species and a new genus of Farreidae (Porifera:
Hexactinellida: Hexactinosida)

Kirk Duplessis and Henry M. Reiswig*

(KD) Redpath Museum, McGill University, Montreal, Quebec, H3A 2K6 Canada,
e-mail: olodrin@hotmail.com;

(HMR) Department of Biology, University of Victoria, and Natural History Section, Royal British
Columbia Museum, Victoria, British Columbia, V8W 3N5 Canada, e-mail: hmreiswig@shaw.ca

Abstract.—Two new species of Farrea and a single new species of a closely
related new genus, Asceptrulum, all members of the hexactinosan family Far-
reidae, are described from three widely distant locations. Farrea herdendorfi,
new species, with a type series of eight specimens obtained from 2200 m off
Charleston, South Carolina, U.S.A., NW Atlantic Ocean, is distinguished by
two anchorate clavule forms (umbellate and thimblate or thimble-shaped), both
without spiral arrangement of claws. Farrea seiri, new species, represented by
3 fragments of a single specimen from 1450 m near the South East Indian
Ridge, mid Indian Ocean, is characterized by only anchorate clavules of thim-
blate form, a moderate proportion of which have claws spiralled. Asceptrulum
axialis, new genus, new species, is represented by several fragments from a
single specimen collected from 2387 m on the Juan de Fuca Ridge, northern
Oregon, U.S.A., NE Pacific Ocean. It is distinguished by the combination of
complete absence of sceptrules and a one-layered farreoid framework. The

diagnosis of Farreidae is emended to encompass the new genus.

Although recognized 132 years ago, hex-
actinellid sponges, more familiarly referred
to as “glass sponges’’, are still obscure
members of the deep sea invertebrate fauna.
Members of the dictyonine family Farreidae
Gray are among the most commonly en-
countered hexactinellids, distributed mainly
on continental shelves and slopes, but ex-
tending into deep water to over 5200 m
depth. This hexactinosan family presently
includes 21 species distributed unevenly in
5 genera, 17 of those in the genus Farrea.
The most recent family diagnosis (Reiswig
2002) focused on one class of free spicules,
sceptrules, which here consist of at least
one form of clavule or sarule, with or with-
out lonchiole or aspidoscopule. Forms with
narrow-head scopule were excluded from
the family. A second common feature of
most, but not all, members of the family,
not included in that recent diagnosis, is the

minimal, one-layered dictyonal framework
at the growing margin.

Specimens of three forms, obtained from
moderately deep water and submitted to our
laboratory for identification, have proven to
be undescribed species of this family. Two
of them are easily incorporated in the spe-
ciose genus Farrea Bowerbank, 1862, but
the third entirely lacks sceptrules and thus
cannot be assigned to a family on the basis
of present diagnoses. Its assignment to a
new genus erected within Farreidae requires
emendation of that family diagnosis, pro-
posed here.

Materials and Methods

Most submitted specimens (all F. her-
dendorfi, new species, and Asceptrulum ax-
ialis, new genus, new species) were col-
lected by robot submersible and were ac-

200

companied by videotape of the collection
process. Only F. seiri new species, was col-
lected by dredge.

Small fragments of the sponge body wall
were either whole-mounted in Canada bal-
sam for light microscopy (LM), or were
dissolved in hot nitric acid. The acid-
cleaned skeletal frameworks were removed,
rinsed and dried; the remaining spicule sus-
pensions were filtered through 25 mm di-
ameter, 0.2 mm-pore-size, nitrocellulose fil-
ters; filters with spicules were thoroughly
rinsed with distilled water, dried, cleared
with xylene, and mounted in Canada bal-
sam on microscope slides. Characters of
frameworks and spicules were measured by
computer using a microscope-coupled dig-
itizer. Data are reported as mean + standard
deviation (range, number of measure-
ments). Spicules for scanning electron mi-
croscopy (SEM) were similarly nitric acid
cleaned, rinsed in distilled water and then
directly deposited onto cover glasses
mounted on SEM stubs. Acid-cleaned and
rinsed fragments of body wall skeletal
frameworks were mounted directly on stubs
with epoxy. Following gold-palladium coat-
ing, specimens were viewed and photo-
graphed with a JEOL JSM-840 SEM. Spic-
ule drawings were the made by importing
LM or SEM images into a computer image-
processing program, and then tracing on
screen. Type specimens of the new species
have been deposited in the National Mu-
seum of Natural History, Smithsonian In-
stitution, Washington D.C. (USNM).

Family Farreidae Gray, 1872

Diagnosis (emended).—Hexactinosida
typically with sceptrules in the form of cla-
vules, or their derivatives, sarules, lonchi-
oles or aspidoscopules, and typically with a
farreoid dictyonal framework. Where scep-
trules are lacking the framework is farreoid.
Where the framework is euretoid, sarules
are present.

Remarks.—Emendation of the diagnosis

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

of Farreidae is required for inclusion of the
new genus Asceptrulum as proposed below.

Genus Farrea Bowerbank, 1862

Type species.—Farrea occa Bowerbank,
1862, by monotypy.

Farrea herdendorfi, new species
(Figs. 1-3; Table 1)

Holotype.—USNM 1001596; S.S. “Cen-
tral America’ wreck, 300 km S. of Charles-
ton, S.C., 31.5°N, 77°W, 12 Sep 1989, 2200
m depth, coll. C.E. Herdendorf, R/S
‘Nemo’ from R/V ‘Arctic Explorer’, dive
UA.

Paratypes.—All from above sampling
location and vessels; USNM 1001597,
USNM 1001600, USNM 1001601, USNM
1001602, USNM 1001603, all 12 Sep 1989,
col. C.E. Herdendorf, Dive UA; USNM
1001598, USNM 1001599, 21 Sep 1990,
col. B. Evans, dive AC.

Diagnosis.—Farrea with only anchorate
clavules, the heads of which vary between
two extreme forms—an umbellate (hemi-
spherical) form and a thimblate (nearly cy-
lindric) form with straight claws nearly par-
allel to shaft. Shafts of all clavules are
rough but lack conspicuous spines.

Description.—Size and shape: The ho-
lotype is 30 cm in height, with a branching
element arising 16 cm above the lower end
(base is missing), extending 11 cm at 60°
from the primary axis (Fig. 1E). At widest
points, the main body and lateral branch
are, respectively, 5.0 and 4.2 cm thick. The
central body of the sponge is cryptically bi-
laterally symmetical (see below), composed
of an original flat blade or stipe incorporat-
ed as one side of an axial tube by medial
fusion of lateral undulations or ruffles.
Through further lateral extension and tight
curvature, the lateral ruffles fuse to form
tubes appended onto the axial tube. The di-
ameter of outer exposed tubular apertures
of the holotype are 6.7 + 2.0 mm (range
5-10 mm, n = 12).

A sequence of age/maturation stages is

VOLUME 117, NUMBER 2 201

oat.

nat

Suber

Fig. 1. Farrea herdendorfi, new species; body form. A. Paratype USNM 1001601 in lateral view. B. Same
in frontal view. C. Paratype USNM 1001602 in frontal view, asterisks indicate fusion points of lateral pleats to
form axial tube. D. Same viewed from distal end with axial tube closed by fusion series at asterisk. E. Holotype

USNM 1001596, lateral view. EK Paratype USNM 1001597, with thickened walls, in frontal view. Scale bars =

2 cm.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2.

apparent in the type series as inferred from
wall thickness and degree of expansion and
fusion of lateral edges—from a simple
blade to the complex fused structure of the
lateral tubes. The simplest paratype avail-
able (Figs. 1A, B) is an axial blade, 18.6
cm long, with undulatory expansion of lat-
eral margins as ruffles but without fusion
between them. A more complex and older
but shorter specimen, 8.2 cm long (Fig.
IC), exhibits logitudinal fusion between
ruffles along one side to form an axial tube
on the axial blade (Fig. 1D). The holotype
(Fig. 1E) exhibits the next stage with fusion
and branching of ruffles to form a complex
bed of lateral tubes supported upon the ax-
ial tube. Beyond the branching point, the
axial blade and tube of both extremities
arise de-novo from the lateral ruffles—con-
tinuity does not exist between axial com-
ponents of the basal axis and the two distal
shoots. By wall thickness, paratype CA6,
14.7 cm long, is next oldest of the series
(Fig. 1F) but its gross morphology has ap-
parently been simplified by abrasion. It con-
sists of the axial tube bearing only the
thick-walled bases of lateral tubes in alter-
nating offset pairs. The marginal ruffles and
tubes have apparently been torn off during

Farrea herdendorfi, new species, framework (SEM). A. Single-layer primary framework in marginal
area of paratype USNM 1001601, bearing long, usually straight spurs. B. Thickened main body area of holotype
USNM 1001596 with irregular dictyonal layers added on dermal side. Scale bars = 1 mm.

collection. The oldest specimen, not fig-
ured, is an extremely thick-walled and
dense basal part, 6.6 X 4.9 cm, of a much
larger specimen. That this constructed se-
ries of body shapes represents a sequence
in growth of a single species is confirmed
by identical loose spicules in all specimens.

Framework (Fig. 2, Table 1): The frame-
work is an unchannelized dictyonal lattice,
consisting of, in the thinnest-walled
(young) specimens and at growing edges of
thick-walled (older) specimens, a one-layer,
two-dimensional lattice of somewhat irreg-
ular rectangular meshwork with easily rec-
ognizable longitudinal dictyonal strands
(Fig. 2A). Most of the framework of all but
the thinnest specimens is augmented by ad-
dition of secondary dictyonalia in irregular-
mesh network mainly on dermal, but in
some places on the atrial side to form a
three-dimensional framework (Fig. 2B).
Small hexactins attached to beams of pri-
mary and secondary dictyonalia are abun-
dant. Wall thickening and increasing rigid-
ity of the framework with aging is due to
addition of both more secondary dictyona-
lia and small hexactins, but only slightly
accompanied by thickening of primary dic-
tyonal strands. Spurs are long, thin, rough

VOLUME 117, NUMBER 2

203

Fig. 3.
B. Thimblate anchorate clavule, whole and head. C. Umbellate anchorate clavule head. D. Uncinate, whole and
magnified segment. E. Oxyhexaster. EF Onychexaster with magnified ray tip.

and spine-like, usually straight but occa-
sionally slightly curved.

Spicules (Fig. 3, Table 1): Pentactins
(Fig. 3A), lining both dermal and atrial sur-
faces, have tangential rays heavily spined
on outer surfaces, and long proximal ray
with heavy spination only on upper third.
Uncinates are very long, exceptionally thin
and moderately barbed. Two forms of an-
chorate clavules occur intermixed on both
surfaces, both forms having a thin, smooth
shaft ending in a slightly rough, bluntly
pointed tip. The thimblate (thimble-shaped)

Farrea herdendorfi, new species, spicules of holotype USNM 1001596 (SEM). A. Surface pentactin.

form (Fig. 3B) has approximately 15 spines
projecting down from a discoid cap, flaring
slightly outward at the lower edge. The an-
chorate form (Fig. 3C) has approximately
10 spines projecting out and down from a
smoothly rounded cap, continuing on the
angle of curvature without reflexion. Mi-
croscleres consist of two types of smooth
hexasters distributed throughout the speci-
men. Oxyhexasters (Fig. 3E) have six long
primary rays, each bearing 2—3 secondary
rays ending in sharp tips. Onychexasters
(Fig. 3F), alternately interpretable as dis-

204

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Spicule and framework dimensions (in 1m) of Farrea herdendorfi, new species, holotype USNM

1001596, from off South Carolina, USA.

Item Mean

A. Surface pentactin:

tangential ray length 231
tangential ray width 13.1
proximal ray length 298
proximal ray width 2
B. Thimblate clavule length 353
head length SOP
head diameter 34.9
C. Umbellate clavule length 454
head length 36.9
head diameter 48.2
D. Uncinate length 1368
width 6.1
E. Oxyhexaster diameter 120
primary ray length 30.7
secondary ray length 33.1
F. Onychexaster diameter 114
primary ray length 27.9
secondary ray length XO}
G. Framework beam length 352
beam width 28.2
H. Framework spur length 287.0

cohexasters with reduced discs, have 3—4
irregularly lumpy secondary rays (without
sharp spines) each ending in a flat disc with
2—4 short blunt claws.

Etymology.—The species is named after
the collector of the holotype, Prof. Charles
E. Herdendorf, who also served as coordi-
nator of the Adjunct Science and Education
Program, S.S. ‘Central America’ Project,
Columbus-America Discovery Group. Gen-
der of the species name is female.

Remarks.—Herdendorf et al. (1995) re-
ported this form as ““Farrea new species”
(p. 86) and figured the specimen designated
here as holotype being collected (their Fig.
45). The species differs from all known
Farrea in the form of its clavules. The dis-
tinctive thimblate clavule head is most sim-
ilar to that of: form B of F. kurilensis (Oka-
da, 1932), but those clavules have coarsely
thorned shafts and are. accompanied by a

St. dey. Range N
29 179-299 50
2.9 5.2—20.0 50
81 141-464 50
DED, 6.6—17.2 50
WD 130—490 50
35) 20-45 50
eS 19.7-56.2 50
55 313-581 50
4.4 29-49 50
5.3 32-66 50
308 830-2086 50
1.4 3.5-9.9 50
13 86-146 50
4.6 21.6-43.1 50
47 20.7-44.3 50
12 87-141 50
3.6 22.2-39.7 50
4.0 19.3-37.8 50
129 125-748 50
9.2 14.5-57.7 50
110 124-727 50

pileate clavule not present in F. herden-
dorfi.

Farrea seiri, new species
(Figs. 4, 5; Table 2)

Holotype.—USNM_ 1001594, Southeast
Indian Ridge, Indian Ocean, 39°12.83’S,
77°52.88'W, 22 Mar 1996, 1450 m depth,
coll. D.S. Scheirer and K. Johnson, Boo-
merang Expedition, Leg 6, R/V ‘Melville’,
biosample #7, Site 48, Dredge 58, dive#
BMRG 06 MV.

Diagnosis.—Farrea with only anchorate
clavules, all thimblate in form without shaft
spines. The most abundant clavule type has
straight claws while the less common form
has claws spiralled either dextrally or si-
nestrally.

Description.—Size and shape: The ho-
lotype and only sample consists of three

VOLUME 117, NUMBER 2 205

Fig. 4. Farrea seiri, new species, body form and framework of holotype USNM 1001594. A. Body form
with epirhyses evident in magnified inset. B. Outer surface with shallow epirhyses indicated by arrowheads
(SEM). C. Probable primary layer in middle frontal layer of framework exposed by dissection (SEM). D. Two
transverse sections of body wall showing thickened main longitudinal strands deep within framework (SEM).
Scale bars = 0.5 mm.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5.
Straight thimblate anchorate clavule, whole and head. C. Spiral umbellate anchorate clavule head. D. Uncinate,
whole and magnified segment. E. Onychexaster with magnified ray tips.

very stout fragments from the basal part of
a single specimen (Fig. 4A). The specimen
was severely damaged during dredge col-
lection, all distal parts with thin body wall
having been lost. The largest fragment mea-
sures 9.6 X 4.3 cm, the second largest 3.9
x 1.5 cm, and the smallest 1.8 X 1.7 cm.
All fragments are white in colour, with fair-
ly thick walls (for Farreidae), 2.08 + 1.04
mm (range 0.95—3.80 mm, n = 10), though
all are quite fragile, easily crushed and
crumbled.

The main fragment is composed of two
fused tubes, the younger attached obliquely
along the side of the older. The older tu-
bular element provided basal attachment for
the specimen and was dead at the time of
collection. The younger tube has a series of
lateral openings arranged in sets of two in
alternating offset pairs, aperture length 9.2

Farrea seiri, new species, spicules of holotype USNM 1001594 (SEM). A. Surface pentactin. B.

+ 0.9 mm (range 8.3—-10.4 mm, n = 5)
width 5.1 + 0.9 mm (range 4.1—7.0 mm, n
= 5). Attempts to map dermal and atrial
surface topology showed that there is no
consistent distinction between surfaces rel-
ative to tubular walls.

Framework (Fig. 4, Table 2): The outer
layers of the rigid dictyonal framework
(Fig. 4B), are composed of a highly irreg-
ular mesh of hexactins, many with polyra-
dial nodes, outlining shallow extradictyonal
epirhyses and aporhyses as surface pits
(Figs. 4A insert, 4B). Pits have ovoid ap-
ertures, length 0.33 + 0.038 mm (range
0.26—0.38 mm, n = 8), width 0.24 + 0.045
mm (range = 0.18—0.32 mm, n = 8). Dis-
tances between pits 0.66 + 0.18 mm (range
0.33-1.20 mm, n = 32).

Beam thickening has occurred through-
out the entire specimen, and there is no sin-

VOLUME 117, NUMBER 2

207

Table 2.—Spicule and framework dimensions (in 1m) of Farrea seiri, new species, holotype USNM 1001594,

from mid Indian Ocean.

Item Mean

A. Surface pentactin:

tangential ray length 155
tangential ray width 8.1
proximal ray length 193
proximal ray width eo)
B. Straight clavule length 243
head length D3)
head diameter 17.0
C. Spiral clavule length 265
head length D3).7)
head diameter 21.4
D. Uncinate length 802
width 6.5
E. Onychexaster diameter 91
primary ray length . 26.2
secondary ray length 21.9
F. Framework beam length 287
beam width 51

gle layer which can be identified as a buried
farreoid two-dimensional grid (Fig. 4C).
Long stretches of smooth dictyonal strands
located deep within the wall (Fig. 4D) are
hypersilicified, obscuring original hexac-
tins. Both outer and inner meshes are fur-
ther obscured by large numbers of small in-
tercalated hexactins. Spurs are moderately
common on both surfaces, and within the
internal meshwork, but these are not di-
rectly comparable to spurs on the primary
dictyonalia of other farreids.

Spicules (Fig. 5, Table 2): Pentactins
have strong spination on outer surface of
tangential rays, extending almost to the tips
(Fig. 5A). The proximal ray is heavily
spined near the centrum, and entirely rough
throughout its length. Clavules are all an-
chorate and thimblate in form and occur in
two types, a straight thimblate type (Fig.
5B) and spiro-thimblate type (Fig. 5D).
Both have a thin, smooth shaft, ending in a
bluntly pointed tip. The head of the straight
thimblate type has approx. 25 claws pro-
jecting down from a discoid cap, either

St. dev. Range N
37 67-253 50
2.6 3.9-12.5 50
68 99-379 50
2.3 3.1-13.3 50
36 184-347 50
4.9 16.3-39.3 50
3.8 12.8-29.5 50
36 182-345 50
43 21.0-39.3 50
3.4 13.4-28.1 50
376 520-1610 8
DS) 3.3-10.1 8

11 Wag 50
4.0 IQIBSS 50
3.7 13.2—29.0 50
107 89-503 50
16 22-96 83

straight and parallel or flaring slightly out-
ward. The spiro-thimblate type has similar
cap and claws, but claws curve distally ei-
ther to the left (sinistral) or right (dextral).
Pentactins and both clavule types occur on
all surfaces without distinction. Uncinates
are typically long and thin with moderately
developed barbs but without a distinguish-
able centrum (Fig. 5D). The only micros-
clere type is an onychexaster (Fig. 5E) dis-
tributed evenly throughout the specimen.
The finely rough primary rays each bear
four similarly rough secondary rays, each
of which ends in a button margined by 3—
6 short, slightly reclined claws.
Etymology.—The species name, seiri, is
formed from the acronym of its collection
locale, the Southeast Indian Ridge. Gender
of the species name is female.
Remarks.—This species is most similar
and closely related to F. herdendorfi de-
scribed above, but differs in having only
thimblate anchorate clavules and lacking
oxyhexaster microscleres. The unavailabil-
ity of distal portions of the specimen, and

208

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 3.—Spicule and framework dimensions (in wm) of Asceptrulum axialis, new genus, new species, ho-

lotype USNM 1001604, from NE Pacific.

Item Mean

A. Surface pentactin:

tangential ray length 217
tangential ray width 16.8

proximal ray length 446
proximal ray width 15.1

B. Uncinate length 1641
width 10.4

C. Discohexaster diameter 66
primary ray length 10.0
» secondary ray length 25.4

D. Framework beam length* 346
beam width* 43.8

E. Spur length? 335

* In marginal areas of framework.

its presumable two-dimensional farreoid
framework do not seriously compromise its
placement in Farrea since its spiculation is
compelling evidence of its relationship to
other Farrea species. Thickness of the basal
skeleton and presence of shallow epirhyses
and aporhyses are rarely encountered in the
genus but are not unique to this species.

Genus Asceptrulum, new genus

Type species.—Asceptrulum axialis, here
designated.

Diagnosis.—Farreidae lacking sceptru-
les.

Etymology.—The genus name, Asceptru-
lum, is formed from Greek a = without,
plus sceptrula, from Greek skeptron and
Latin sceptrum (neuter) = royal wand or
staff, in allusion to the absence of sceptrule
spicules. Gender of the genus name is neu-
ter.

Remarks.—See under species below.

St. dev. Range N
29 141-279 50
4.4 7.9-25.1 50
108 261-752 50
4.5 8.6—27.1 50
250 1173-1967 12
Zell 5§.5-15.2 23

7 53-81 50
1.7 6.2—13.5 50
3.0 19.8-31.2 50
75 204-584 50
9.1 28.4—77.7 50
719 147-528 50

Asceptrulum axialis, new species
(Figs. 6, 7; Table 3)

Holotype.-—USNM_ 1001604: North
CoAxial segment, Juan de Fuca ridge,
northern Oregon, 46°29.83'N, 129°35.79'W,
19 Jul 1993, 2387 m depth, coll. V. Tun-
nicliffe, R/S ‘“ROPOS’ dive HYS 221.

Diagnosis.—Asceptrulum with axial con-
densation of its farreoid framework.

Description.—Size and shape: The single
specimen encountered and recorded in situ
by video, was broken during collection;
about one-half was recovered as four frag-
ments (Fig. 6A). The intact specimen was
attached to hard substrate in a region of re-
cently formed basalt blocks sparsely
clothed in bacterial mats and strands. Be-
fore collection, the organism was frond-like
or Y-shaped, 14 cm tall, with a branch point
9 cm from the basal attachment. The four
recovered pieces are all thin ribbons or
blades with clear axial thickening, thin mar-

Fig. 6.

Asceptrulum axialis, new genus, new species, body form and framework of holotype USNM 1001604.

A. Body form of recovered fragments with cross-section of one fragment. B. Frontal view of single-layer primary
framework in marginal area (LM). C. Same in slightly oblique transverse view showing long, straight spurs

VOLUME 117, NUMBER 2 209

(SEM). D. Cross-section of axial region of blade, atrial surface down, with longitudinal primary strands seen at
extreme left (SEM). E. Atrial surface of axial region showing thickened primary longitudinal strands (SEM). FE
Dermal surface of same (SEM). Scale bars of B—F = 0.5 mm.

210

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 7.

ginal fringes with low-amplitude undulation
or ruffles. At one point the curved lateral
margins are extended and undergo self-fu-
sion resulting in a short lateral tube 1.4 cm
in diameter. Intact areas of blades are 9.2—
26.2 mm wide and 1.63 + 0.32 mm (range
0.86-1.88 mm, n = 10) thick at axis cen-
ters.

Framework (Fig. 6, Table 3): The pri-
mary framework is a typical farreoid one-
layer, two-dimensional mesh of smooth
beams (Figs. 6B, C) most obvious in the
marginal areas. Blade thickness increases
gradually toward the axis by addition of up
to nine layers of secondary dictyonalia in
irregular arrangement on one side, assumed
dermal, of the primary framework (Fig.
6D). On the two surfaces of the blade axes,
beams are twice as thick on the atrial side
with exposed old primary frame (Fig. 6E),
81.3 + 31.2 pm (range 54-134 wm, n =
5), than on the dermal side (Fig. 6F), 41.1

Asceptrulum axialis, new genus, new species, spicules of holotype USNM 1001604 (SEM). A.
Surface pentactins. B. Uncinate, whole and magnified segment. C. Discohexaster with magnified ray tips.

+ 14.8 wm (range 28.0—64.5 wm, n = 5).
Small hexactins fused to framework beams
are present but sparse. Spurs of the primary
frame are long and straight on both surfaces
(Fig. 6C), but while those on the atrial side
are rough, those of the dermal side are
smooth and often extended and variable in
texture. Many of the dermal spurs are fused
to centres of secondary dictyonalia or tips
of their rays. The secondary structures are
a mixture of true and false nodes, with con-
nections occurring between grid levels by
synapticula. Channelization is absent.
Spicules (Fig. 7, Table 3): The species
has both low diversity and density of loose
spicules. Megascleres consist of large, ro-
bust pentactins (Fig. 7A) and long, thin un-
cinates (Fig. 7B). Pentactins, present on
dermal and gastral surfaces, have tangential
rays with heavy spination on outer and lat-
eral surfaces and proximal rays with coarse
tubercles on the upper third of the ray and

VOLUME 117, NUMBER 2

very sparse low spines over the remainder.
Uncinates are typical with well-developed
barbs, brackets, and no detectable central
tyle. The only microsclere type is a rela-
tively scarce, robust discohexaster (Fig.
7C), distributed evenly throughout the wall.
The six primary rays are short, thick and
smooth, each supporting three heavily
spined secondary rays which end in hemi-
spherically arched discs bearing 5—6 re-
curved marginal spines.

Etymology.—The species name, axialis,
is originally derived from Latin axis = rod
or pole. It is here formed from the English
adjective axial to preserve its euphonious
spelling and reflect the easily visible, dense,
axial skeletal framework. Gender of the
species name is neuter.

Remarks.—Absence of sceptrules in this
specimen cannot be attributed to patholog-
ical condition, damage during collection or
inadequate sampling of spicules. The spec-
imen was almost certainly alive at collec-
tion, with surface pentactins arrayed in the
normal rectangular lattice in places. It is ex-
tremely unlikely that disease or the collec-
tion process would result in loss of only the
one spicule type had sceptrules been pre-
sent. Occasionally sceptrules may be diffi-
cult to obtain in very small samples of far-
reids, but the use of filtration for spicule
collection from cm-size fragments has nev-
er failed to find scopules. When the first
searches for sceptrules in this specimen
proved negative, the entire set of fragments
was eventually extracted for spicules and
examined; not one part of a sceptrule was
found. We are very confident that sceptrules
were neither lost nor overlooked. They
must have been intrinsically absent.

Since sceptrules are lacking in Asceptru-
lum, its assignment to Farreidae rather than
Euretidae is based its one-layered farreoid
framework as its primary dictyonal skele-
ton. A farreoid framework (Reid 1964) con-
sists of a two-dimensional primary grid-like
scaffold with dictyonalia, fused in parallel
longitudinal strands, cross-linked to adja-
cent strands by tangential rays fused side-

211

to-side, resulting in a grid-like layer of
fused framework. It is the single layer, or
two-dimensional, character of this structure
that is considered by some authors to be
distinctive for the family Farreidae. This al-
ternate definition of Farreidae is extremely
important for paleontologists, since loose
spicules are unavailable in fossil material.
Reiswig (2002) did not include the farreoid
framework as a diagnostic feature of Far-
reidae since it is absent in one of its five
extant genera, Sarostegia Topsent, 1904
(with euretoid framework). He did, how-
ever, note that it has historically been an
important diagnostic feature, and thus it is
included here in the emended diagnosis.

The alternative assignment of Asceptru-
lum to Euretidae is poorly supported by
similar overall spiculation (excepting scep-
trule) and presence of a farreoid framework
in one of its genera, the monospecific ge-
nus, Bathyxiphus Schulze, 1899. Position of
Bathyxiphus cannot be used to support as-
signment of Asceptrulum to Euretidae since
its (Bathxiphus) sceptrule type is not known
with complete certainty and its own assign-
ment is both provisional and precarious.
Based upon the firm relationship of the far-
reoid framework with Farreidae, Asceptru-
lum is best assigned to that family.

Within Farreidae, A. axialis has no ob-
vious close relatives. Axial thickening of a
blade-form body is unknown in the family
and absence of oxy-tip microscleres (pres-
ence of only disc- or onych-tip forms) is
known only in four Farrea, all of which
have a body form of branching and usually
anastomosing tubes: F. woodwardi Kent,
1870; F. sollasi Schulze, 1886; F. weltneri
Topsent, 1901; F. occa polyclavula Ta-
bachnick, 1988. None of these are likely an-
cestral forms which could have given rise
to A. axialis through the one-step loss of
sceptrules.

Acknowledgments

We thank the following for providing ac-
cess to the specimens, permission for their

AND

processing, and/or collection data: Drs. C.
E. Herdendorf, R. D. Evans, V. Tunnicliffe,
M. Tsurumi, R. Toll, R. W. Embley, S. K.
Juniper, D. Scheirer, M. K. Harper, and the
Columbus-America Discovery Group, Inc.
This work was supported by a Research
Grant from the Natural Sciences and En-
gineering Research Council of Canada to
HMR.

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Bowerbank, J. S. 1862. On the anatomy and physiol-
ogy of the Spongiadae, part III. On the generic
characters, the specific characters, and on the
method of examination.—Philosophical Trans-
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1087-1135, pls 72-74.

Gray, J. E. 1872. Notes on the classification of the
sponges.—Annals and Magazine of Natural
History (4) 9(54):442—461.

Herdendorf, C. E., T. G. Thompson, & R. D. Evans.
1995. Science on a deep-ocean shipwreck.—
Ohio Journal of Science 95(1):4—224.

Kent, W. S. 1870. On the Hexactinellidae or hexradiate
spiculed siliceous sponges taken in the “Norna’
Expedition off the coast of Spain and Portugal
with description of new species and revision of
the order—Monthly Microscopical Journal 4:
241-252, pls 63-65.

Okada, Y. 1932. Report on the hexactinellid sponges

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

collected by the United States Fisheries steamer
‘Albatross’ in the northwestern Pacific during
the summer of 1906.—Proceedings of the Unit-
ed States National Museum 81(12):1—118, pls
1-6.

Reid, R. E. H. 1964. A monograph of the Upper Cre-
taceous Hexactinellida of Great Britain and
Northern Ireland, part I1V.—Paleontographical
Society Monographs 117(3):cxlix—cliv.

Reiswig, H. M. 2002. Family Farreidae Gray. Pp.
1332-1340 in J. N. A. Hooper & R. M. W. van
Soest, eds., Systema Porifera: A Guide to the
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ers, New York, 1708 pp.

Schulze, F E. 1886. Uber den Bau und das System der
Hexactinelliden.—Abhandlungen der Ko6nig-
lichen Akademie der wissenschaften zu Berlin
(Physikalisch-Mathematisch Classe) 1886:1—97.

. 1899. Amerikanische Hexactinelliden nach
dem materiale der Albatross-Expedition bear-
beitet. Gustav Fischer, Jena, 126 pp.

Tabachnick, K. R. 1988. Hexactinellid sponges from
the mountains of West Pacific. Pp. 49—64 in P.
P. Shirshov, ed., Structural and Functional Re-
searches of the Marine Benthos. Academy of
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Topsent, E. 1901. Eponges nouvelles des Agores.—
Mémoires de la Société zoologique de France
14:448—466.

. 1904. Sarostegia oculata, Hexactinellide nou-

velle des iles du Cap-Vert.—Bulletin du Musée

Océanographique de Monaco 1904(10):1—8.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(2):213—239. 2004.

The origin of biological information and the
higher taxonomic categories

Stephen C. Meyer

Palm Beach Atlantic University, 901 S. Flagler Dr., West Palm Beach, Florida 33401
e-mail: stevemeyer@discovery.org

Introduction

In a recent volume of the Vienna Series
in Theoretical Biology (2003), Gerd B.
Miiller and Stuart Newman argue that what
they call the “origination of organismal
form” remains an unsolved problem. In
making this claim, Miiller and Newman
(2003:3—10) distinguish two distinct issues,
namely, (1) the causes of form generation
in the individual organism during embryo-
logical development and (2) the causes re-
sponsible for the production of novel or-
ganismal forms in the first place during the
history of life. To distinguish the latter case
(phylogeny) from the former (ontogeny),
Miiller and Newman use the term “origi-
nation”’ to designate the causal processes by
which biological form first arose during the
evolution of life. They insist that “the mo-
lecular mechanisms that bring about biolog-
ical form in modern day embryos should
not be confused”? with the causes respon-
sible for the origin (or “‘origination”’) of
novel biological forms during the history of
life (p. 3). They further argue that we know
more about the causes of ontogenesis, due
to advances in molecular biology, molecu-
lar genetics and developmental biology,
than we do about the causes of phylogen-
esis—the ultimate origination of new bio-
logical forms during the remote past.

In making this claim, Miiller and New-
man are careful to affirm that evolutionary
biology has succeeded in explaining how
pre-existing forms diversify under the twin
influences of natural selection and variation
of genetic traits. Sophisticated mathemati-
cally-based models of population genetics

have proven adequate for mapping and un-
derstanding quantitative variability and
populational changes in organisms. Yet
Miiller and Newman insist that population
genetics, and thus evolutionary biology, has
not identified a specifically causal expla-
nation for the origin of true morphological
novelty during the history of life. Central
to their concern is what they see as the in-
adequacy of the variation of genetic traits
as a source of new form and structure. They
note, following Darwin himself, that the
sources of new form and structure must pre-
cede the action of natural selection (2003:
3)—that selection must act on what already
exists. Yet, in their view, the ““genocentric-
ity’’ and “‘incrementalism”’ of the neo-Dar-
winian mechanism has meant that an ade-
quate source of new form and structure has
yet to be identified by theoretical biologists.
Instead, Miiller and Newman see the need
to identify epigenetic sources of morpho-
logical innovation during the evolution of
life. In the meantime, however, they insist
neo-Darwinism lacks any “theory of the
generative” (p. 7).

As it happens, Miiller and Newman are
not alone in this judgment. In the last de-
cade or so a host of scientific essays and
books have questioned the efficacy of se-
lection and mutation as a mechanism for
generating morphological novelty, as even
a brief literature survey will establish.
Thomson (1992:107) expressed doubt that
large-scale morphological changes could
accumulate via minor phenotypic changes
at the population genetic level. Miklos
(1993:29) argued that neo-Darwinism fails
to provide a mechanism that can produce

214

large-scale innovations in form and com-
plexity. Gilbert et al. (1996) attempted to
develop a new theory of evolutionary
mechanisms to supplement classical neo-
Darwinism, which, they argued, could not
adequately explain macroevolution. As they
put it in a memorable summary of the sit-
uation: “starting in the 1970s, many biol-
ogists began questioning its [neo-Darwin-
ism’s] adequacy in explaining evolution.
Genetics might be adequate for explaining
microevolution, but microevolutionary
changes in gene frequency were not seen as
able to turn a reptile into a mammal or to
convert a fish into an amphibian. Microevo-
lution looks at adaptations that concern the
survival of the fittest, not the arrival of the
fittest. As Goodwin (1995) points out, “the
origin of species—Darwin’s problem—re-
mains unsolved’ ”’ (p. 361). Though Gilbert
et al. (1996) attempted to solve the problem
of the origin of form by proposing a greater
role for developmental genetics within an
otherwise neo-Darwinian framework,' nu-
merous recent authors have continued to
raise questions about the adequacy of that
framework itself or about the problem of
the origination of form generally (Webster
& Goodwin 1996; Shubin & Marshall
2000; Erwin 2000; Conway Morris 2000,
2003b; Carrol 2000; Wagner 2001; Becker
& Loénnig 2001; Stadler et al. 2001; Lonnig
& Saedler 2002; Wagner & Stadler 2003;
Valentine 2004:189—194).

What lies behind this skepticism? Is it
warranted? Is a new and specifically causal
theory needed to explain the origination of
biological form?

This review will address these questions.
It will do so by analyzing the problem of
the origination of organismal form (and the

' Specifically, Gilbert et al. (1996) argued that
changes in morphogenetic fields might produce large-
scale changes in the developmental programs and, ul-
timately, body plans of organisms. Yet they offered no
evidence that such fields—if indeed they exist—can
be altered to produce advantageous variations in body
plan, though this is a necessary condition of any suc-
cessful causal theory of macroevolution.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

corresponding emergence of higher taxa)
from a particular theoretical standpoint.
Specifically, it will treat the problem of the
origination of the higher taxonomic groups
as a manifestation of a deeper problem,
namely, the problem of the origin of the
information (whether genetic or epigenetic)
that, as it will be argued, is necessary to
generate morphological novelty.

In order to perform this analysis, and to
make it relevant and tractable to systema-
tists and paleontologists, this paper will ex-
amine a paradigmatic example of the origin
of biological form and information during
the history of life: the Cambrian explosion.
During the Cambrian, many novel animal
forms and body plans (representing new
phyla, sub-phyla and classes) arose in a
geologically brief period of time. The fol-
lowing information-based analysis of the
Cambrian explosion will support the claim
of recent authors such as Miiller and New-
man that the mechanism of selection and
genetic mutation does not constitute an ad-
equate causal explanation of the origination
of biological form in the higher taxonomic
groups. It will also suggest the need to ex-
plore other possible causal factors for the
origin of form and information during the
evolution of life and will examine some
other possibilities that have been proposed.

The Cambrian Explosion

The “‘Cambrian explosion”’ refers to the
geologically sudden appearance of many
new animal body plans about 530 million
years ago. At this time, at least nineteen,
and perhaps as many as thirty-five phyla of
forty total (Meyer et al. 2003), made their
first appearance on Earth within a narrow
five- to ten-million-year window of geolog-
ic time (Bowring et al. 1993, 1998a:1,
1998b:40; Kerr 1993; Monastersky 1993;
Aris-Brosou & Yang 2003). Many new sub-
phyla, between 32 and 48 of 56 total (Mey-
er et al. 2003), and classes of animals also
arose at this time with representatives of
these new higher taxa manifesting signifi-

VOLUME 117, NUMBER 2

cant morphological innovations. The Cam-
brian explosion thus marked a major epi-
sode of morphogenesis in which many new
and disparate organismal forms arose in a
geologically brief period of time.

To say that the fauna of the Cambrian
period appeared in a geologically sudden
manner also implies the absence of clear
transitional intermediate forms connecting
Cambrian animals with simpler pre-Cam-
brian forms. And, indeed, in almost all cas-
es, the Cambrian animals have no clear
morphological antecedents in earlier Ven-
dian or Precambrian fauna (Miklos 1993,
Erwin et al. 1997:132, Steiner & Reitner
2001, Conway Morris 2003b:510, Valentine
et al. 2003:519—520). Further, several re-
cent discoveries and analyses suggest that
these morphological gaps may not be mere-
ly an artifact of incomplete sampling of the
fossil record (Foote 1997, Foote et al. 1999,
Benton & Ayala 2003, Meyer et al. 2003),
suggesting that the fossil record is at least
approximately reliable (Conway Morris
2003b:505).

As a result, debate now exists about the
extent to which this pattern of evidence
comports with a strictly monophyletic view
of evolution (Conway Morris 1998a, 2003a,
2003b:510; Willmer 1990, 2003). Further,
among those who accept a monophyletic
view of the history of life, debate exists
about whether to privilege fossil or molec-
ular data and analyses. Those who think the
fossil data provide a more reliable picture of
the origin of the Metazoan tend to think
these animals arose relatively quickly—that
the Cambrian explosion had a “short fuse.”
(Conway Morris 2003b:505—506, Valentine
& Jablonski 2003). Some (Wray et al. 1996),
but not all (Ayala et al. 1998), who think
that molecular phylogenies establish reliable
divergence times from pre-Cambrian ances-
tors think that the Cambrian animals evolved
over a very long period of time—that the
Cambrian explosion had a “long fuse.”’ This
review will not address these questions of
historical pattern. Instead, it will analyze
whether the neo-Darwinian process of mu-

215

tation and selection, or other processes of
evolutionary change, can generate the form
and information necessary to produce the
animals that arise in the Cambrian. This
analysis will, for the most part,? therefore,
not depend upon assumptions of either a
long or short fuse for the Cambrian explo-
sion, or upon a monophyletic or polyphyletic
view of the early history of life.

Defining Biological Form and Information

Form, like life itself, is easy to recognize
but often hard to define precisely. Yet, a rea-
sonable working definition of form will suf-
fice for our present purposes. Form can be
defined as the four-dimensional topological
relations of anatomical parts. This means that
one can understand form as a unified arrange-
ment of body parts or material components
in a distinct shape or pattern (topology )—one
that exists in three spatial dimensions and
which arises in time during ontogeny.

Insofar as any particular biological form
constitutes something like a distinct ar-
rangement of constituent body parts, form
can be seen as arising from constraints that
limit the possible arrangements of matter.
Specifically, organismal form arises (both
in phylogeny and ontogeny) as possible ar-
rangements of material parts are con-
strained to establish a specific or particular
arrangement with an identifiable three di-
mensional topography—one that we would
recognize as a particular protein, cell type,
organ, body plan or organism. A particular

?If one takes the fossil record at face value and
assumes that the Cambrian explosion took place within
a relatively narrow 5—10 million year window, explain-
ing the origin of the information necessary to produce
new proteins, for example, becomes more acute in part
because mutation rates would not have been sufficient
to generate the number of changes in the genome nec-
essary to build the new proteins for more complex
Cambrian animals (Ohno 1996:8475—8478). This re-
view will argue that, even if one allows several hun-
dred million years for the origin of the metazoan, sig-
nificant probabilistic and other difficulties remain with
the neo-Darwinian explanation of the origin of form
and information.

216

“form,” therefore, represents a highly spe-
cific and constrained arrangement of mate-
rial components (among a much larger set
of possible arrangements).

Understanding form in this way suggests
a connection to the notion of information in
its most theoretically general sense. When
Shannon (1948) first developed a mathe-
matical theory of information he equated
the amount of information transmitted with
the amount of uncertainty reduced or elim-
inated in a series of symbols or characters.
Information, in Shannon’s theory, is thus
imparted as some options are excluded and
others are actualized. The greater the num-
ber of options excluded, the greater the
amount of information conveyed. Further,
constraining a set of possible material ar-
rangements by whatever process or means
involves excluding some options and actu-
alizing others. Thus, to constrain a set of
possible material states is to generate infor-
mation in Shannon’s sense. It follows that
the constraints that produce biological form
also impart information. Or conversely, one
might say that producing organismal form
by definition requires the generation of in-
formation.

In classical Shannon information theory,
the amount of information in a system is
also inversely related to the probability of
the arrangement of constituents in a system
or the characters along a communication
channel (Shannon 1948). The more improb-
able (or complex) the arrangement, the
more Shannon information, or information-
carrying capacity, a string or system pos-
sesses.

Since the 1960s, mathematical biologists
have realized that Shannon’s theory could
be applied to the analysis of DNA and pro-
teins to measure the information-carrying
capacity of these macromolecules. Since
DNA contains the assembly instructions for
building proteins, the information-process-
ing system in the cell represents a kind of
communication channel (Yockey 1992:
110). Further, DNA conveys information
via specifically arranged sequences of nu-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

cleotide bases. Since each of the four bases
has a roughly equal chance of occurring at
each site along the spine of the DNA mol-
ecule, biologists can calculate the probabil-
ity, and thus the information-carrying ca-
pacity, of any particular sequence n bases
long.

The ease with which information theory
applies to molecular biology has created
confusion about the type of information that
DNA and proteins possess. Sequences of
nucleotide bases in DNA, or amino acids in
a protein, are highly improbable and thus
have large information-carrying capacities.
But, like meaningful sentences or lines of
computer code, genes and proteins are also
specified with respect to function. Just as
the meaning of a sentence depends upon the
specific arrangement of the letters in a sen-
tence, so too does the function of a gene
sequence depend upon the specific arrange-
ment of the nucleotide bases in a gene.
Thus, molecular biologists beginning with
Crick equated information not only with
complexity but also with “‘specificity,”
where ‘‘specificity’’ or “‘specified’’ has
meant “necessary to function” (Crick
1958:144, 153; Sarkar, 1996:191).> Molec-
ular biologists such as Monod and Crick
understood biological information—the in-
formation stored in DNA and proteins—as
something more than mere complexity (or
improbability). Their notion of information
associated both biochemical contingency
and combinatorial complexity with DNA
sequences (allowing DNA’s carrying capac-
ity to be calculated), but it also affirmed
that sequences of nucleotides and amino ac-
ids in functioning macromolecules pos-
sessed a high degree of specificity relative
to the maintenance of cellular function.

The ease with which information theory
applies to molecular biology has also cre-
ated confusion about the location of infor-

3 As Crick put it, “‘information means here the pre-
cise determination of sequence, either of bases in the
nucleic acid or on amino acid residues in the protein”
(Crick 1958:144, 153).

VOLUME 117, NUMBER 2

mation in organisms. Perhaps because the
information carrying capacity of the gene
could be so easily measured, it has been
easy to treat DNA, RNA and proteins as the
sole repositories of biological information.
Neo-Darwinists in particular have assumed
that the origination of biological form could
be explained by recourse to processes of ge-
netic variation and mutation alone (Levin-
ton 1988:485). Yet if one understands or-
ganismal form as resulting from constraints
on the possible arrangements of matter at
many levels in the biological hierarchy—
from genes and proteins to cell types and
tissues to organs and body plans—then
clearly biological organisms exhibit many
levels of information-rich structure.

Thus, we can pose a question, not only
about the origin of genetic information, but
also about the origin of the information nec-
essary to generate form and structure at lev-
els higher than that present in individual
proteins. We must also ask about the origin
of the “specified complexity,” as opposed
to mere complexity, that characterizes the
new genes, proteins, cell types and body
plans that arose in the Cambrian explosion.
Dembski (2002) has used the term “‘com-
plex specified information”’ (CSI) as a syn-
onym for “specified complexity” to help
distinguish functional biological informa-
tion from mere Shannon information—that
is, specified complexity from mere com-
plexity. This review will use this term as
well.

The Cambrian Information Explosion

The Cambrian explosion represents a re-
markable jump in the specified complexity
or “complex specified information” (CSI)
of the biological world. For over three bil-
lion years, the biological realm included lit-
tle more than bacteria and algae (Brocks et
al. 1999). Then, beginning about 570—565
million years ago (mya), the first complex
multicellular organisms appeared in the
rock strata, including sponges, cnidarians,
and the peculiar Ediacaran biota (Grotzin-

217

ger et al. 1995). Forty million years later,
the Cambrian explosion occurred (Bowring
et al. 1993). The emergence of the Edi-
acaran biota (570 mya), and then to a much
greater extent the Cambrian explosion (530
mya), represented steep climbs up the bio-
logical complexity gradient.

One way to estimate the amount of new
CSI that appeared with the Cambrian ani-
mals is to count the number of new cell
types that emerged with them (Valentine
1995:91—93). Studies of modern animals
suggest that the sponges that appeared in
the late Precambrian, for example, would
have required five cell types, whereas the
more complex animals that appeared in the
Cambrian (e.g., arthropods) would have re-
quired fifty or more cell types. Functionally
more complex animals require more cell
types to perform their more diverse func-
tions. New cell types require many new and
specialized proteins. New proteins, in turn,
require new genetic information. Thus an
increase in the number of cell types implies
(at a minimum) a considerable increase in
the amount of specified genetic informa-
tion. Molecular biologists have recently es-
timated that a minimally complex single-
celled organism would require between 318
and 562 kilobase pairs of DNA to produce
the proteins necessary to maintain life
(Koonin 2000). More complex single cells
might require upward of a million base
pairs. Yet to build the proteins necessary to
sustain a complex arthropod such as a tri-
lobite would require orders of magnitude
more coding instructions. The genome size
of a modern arthropod, the fruitfly Dro-
sophila melanogaster, 1s approximately 180
million base pairs (Gerhart & Kirschner
1997:121, Adams et al. 2000). Transitions
from a single cell to colonies of cells to
complex animals represent significant (and,
in principle, measurable) increases in CSI.

Building a new animal from a single-
celled organism requires a vast amount of
new genetic information. It also requires a
way of arranging gene products—pro-
teins—into higher levels of organization.

218

New proteins are required to service new
cell types. But new proteins must be orga-
nized into new systems within the cell; new
cell types must be organized into new tis-
sues, organs, and body parts. These, in turn,
must be organized to form body plans. New
animals, therefore, embody hierarchically
organized systems of lower-level parts
within a functional whole. Such hierarchi-
cal organization itself represents a type of
information, since body plans comprise
both highly improbable and functionally
specified arrangements of lower-level parts.
The specified complexity of new body
plans requires explanation in any account of
the Cambrian explosion.

Can neo-Darwinism explain the discon-
tinuous increase in CSI that appears in the
Cambrian explosion—either in the form of
new genetic information or in the form of
hierarchically organized systems of parts?
We will now examine the two parts of this
question.

Novel Genes and Proteins

Many scientists and mathematicians have
questioned the ability of mutation and se-
lection to generate information in the form
of novel genes and proteins. Such skepti-
cism often derives from consideration of
the extreme improbability (and specificity)
of functional genes and proteins.

A typical gene contains over one thousand
precisely arranged bases. For any specific ar-
rangement of four nucleotide bases of length
n, there is a corresponding number of pos-
sible arrangements of bases, 4”. For any pro-
tein, there are 20" possible arrangements of
protein-forming amino acids. A gene 999
bases in length represents one of 4°”? possi-
ble nucleotide sequences; a protein of 333
amino acids is one of 20%*? possibilities.

Since the 1960s, some biologists have
thought functional proteins to be rare among
the set of possible amino acid sequences.
Some have used an analogy with human lan-
guage to illustrate why this should be the
case. Denton (1986, 309-311), for example,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

has shown that meaningful words and sen-
tences are extremely rare among the set of
possible combinations of English letters, es-
pecially as sequence length grows. (The ra-
tio of meaningful 12-letter words to 12-letter
sequences is 1/10'4; the ratio of 100-letter
sentences to possible 100-letter strings is
1/10!°°.) Further, Denton shows that most
meaningful sentences are highly isolated
from one another in the space of possible
combinations, so that random substitutions
of letters will, after a very few changes, in-
evitably degrade meaning. Apart from a few
closely clustered sentences accessible by
random substitution, the overwhelming ma-
jority of meaningful sentences lie, probabi-
listically speaking, beyond the reach of ran-
dom search.

Denton (1986:301—324) and others have
argued that similar constraints apply to
genes and proteins. They have questioned
whether an undirected search via mutation
and selection would have a reasonable
chance of locating new islands of func-
tion—representing fundamentally new
genes or proteins—within the time avail-
able (Eden 1967, Shtitzenberger 1967,
Lgvtrup 1979). Some have also argued that
alterations in sequencing would likely result
in loss of protein function before funda-
mentally new function could arise (Eden
1967, Denton 1986). Nevertheless, neither
the extent to which genes and proteins are
sensitive to functional loss as a result of
sequence change, nor the extent to which
functional proteins are isolated within se-
quence space, has been fully known.

Recently, experiments in molecular biol-
ogy have shed light on these questions. A
variety of mutagenesis techniques have
shown that proteins (and thus the genes that
produce them) are indeed highly specified
relative to biological function (Bowie &
Sauer 1989, Reidhaar-Olson & Sauer 1990,
Taylor et al. 2001). Mutagenesis research
tests the sensitivity of proteins (and, by im-
plication, DNA) to functional loss as a result
of alterations in sequencing. Studies of pro-
teins have long shown that amino acid res-

VOLUME 117, NUMBER 2

idues at many active positions cannot vary
without functional loss (Perutz & Lehmann
1968). More recent protein studies (often us-
ing mutagenesis experiments) have shown
that functional requirements place significant
constraints on sequencing even at non-active
site positions (Bowie & Sauer 1989, Reid-
haar-Olson & Sauer 1990, Chothia et al.
1998, Axe 2000, Taylor et al. 2001). In par-
ticular, Axe (2000) has shown that multiple
as opposed to single position amino acid
substitutions inevitably result in loss of pro-
tein function, even when these changes oc-
cur at sites that allow variation when altered
in isolation. Cumulatively, these constraints
imply that proteins are highly sensitive to
functional loss as a result of alterations in
sequencing, and that functional proteins rep-
resent highly isolated and improbable ar-
rangements of amino acids—arrangements
that are far more improbable, in fact, than
would be likely to arise by chance alone in
the time available (Reidhaar-Olson & Sauer
1990; Behe 1992; Kauffman 1995:44;
Dembski 1998:175—223; Axe 2000, 2004).
(See below the discussion of the neutral the-
ory of evolution for a precise quantitative
assessment.)

Of course, neo-Darwinists do not envi-
sion a completely random search through
the set of all possible nucleotide sequenc-
es—so-called ““sequence space.’’ They en-
vision natural selection acting to preserve
small advantageous variations in genetic se-
quences and their corresponding protein
products. Dawkins (1996), for example, lik-
ens an organism to a high mountain peak.
He compares climbing the sheer precipice
up the front side of the mountain to build-
ing a new organism by chance. He ac-
knowledges that this approach up ““Mount
Improbable” will not succeed. Neverthe-
less, he suggests that there is a gradual
slope up the backside of the mountain that
could be climbed in small incremental
steps. In his analogy, the backside climb up
“Mount Improbable” corresponds to the
process of natural selection acting on ran-
dom changes in the genetic text. What

219

chance alone cannot accomplish blindly or
in one leap, selection (acting on mutations)
can accomplish through the cumulative ef-
fect of many slight successive steps.

Yet the extreme specificity and complex-
ity of proteins presents a difficulty, not only
for the chance origin of specified biological
information (i.e., for random mutations act-
ing alone), but also for selection and muta-
tion acting in concert. Indeed, mutagenesis
experiments cast doubt on each of the two
scenarios by which neo-Darwinists envision
new information arising from the mutation/
selection mechanism (for review, see LOnnig
2001). For neo-Darwinism, new functional
genes either arise from non-coding sections
in the genome or from preexisting genes.
Both scenarios are problematic.

In the first scenario, neo-Darwinists en-
vision new genetic information arising from
those sections of the genetic text that can
presumably vary freely without conse-
quence to the organism. According to this
scenario, non-coding sections of the ge-
nome, or duplicated sections of coding re-
gions, can experience a protracted period of
“neutral evolution”’ (Kimura 1983) during
which alterations in nucleotide sequences
have no discernible effect on the function
of the organism. Eventually, however, a
new gene sequence will arise that can code
for a novel protein. At that point, natural
selection can favor the new gene and its
functional protein product, thus securing
the preservation and heritability of both.

This scenario has the advantage of allow-
ing the genome to vary through many gen-
erations, aS mutations “‘search”’ the space
of possible base sequences. The scenario
has an overriding problem, however: the
size of the combinatorial space (i.e., the
number of possible amino acid sequences)
and the extreme rarity and isolation of the
functional sequences within that space of
possibilities. Since natural selection can do
nothing to help generate new functional se-
quences, but rather can only preserve such
sequences once they have arisen, chance
alone—random variation—must do_ the

220

work of information generation—that is, of
finding the exceedingly rare functional se-
quences within the set of combinatorial
possibilities. Yet the probability of random-
ly assembling (or “‘finding,”’ in the previous
sense) a functional sequence is extremely
small.

Cassette mutagenesis experiments per-
formed during the early 1990s suggest that
the probability of attaining (at random) the
correct sequencing for a short protein 100
amino acids long is about 1 in 10° (Reid-
haar-Olson & Sauer 1990, Behe 1992:65—
69). This result agreed closely with earlier
calculations that Yockey (1978) had per-
formed based upon the known sequence
variability of cytochrome c in different spe-
cies and other theoretical considerations.
More recent mutagenesis research has pro-
vided additional support for the conclusion
that functional proteins are exceedingly rare
among possible amino acid sequences (Axe
2000, 2004). Axe (2004) has performed site
directed mutagenesis experiments on a 150-
residue protein-folding domain within a B-
lactamase enzyme. His experimental meth-
od improves upon earlier mutagenesis tech-
niques and corrects for several sources of
possible estimation error inherent in them.
On the basis of these experiments, Axe has
estimated the ratio of (a) proteins of typical
size (150 residues) that perform a specified
function via any folded structure to (b) the
whole set of possible amino acids sequenc-
es of that size. Based on his experiments,
Axe has estimated this ratio to be | to 1077.
Thus, the probability of finding a functional
protein among the possible amino acid se-
quences corresponding to a 150-residue
protein is similarly 1 in 1077.

Other considerations imply additional
improbabilities. First, new Cambrian ani-
mals would require proteins much longer
than 100 residues to perform many neces-
sary specialized functions. Ohno (1996) has
noted that Cambrian animals would have
required complex proteins such as lysyl ox-
idase in order to support their stout body
structures. Lysyl oxidase molecules in ex-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

tant organisms comprise over 400 amino
acids. These molecules are both highly
complex (non-repetitive) and functionally
specified. Reasonable extrapolation from
mutagenesis experiments done on shorter
protein molecules suggests that the proba-
bility of producing functionally sequenced
proteins of this length at random is so small
as to make appeals to chance absurd, even
granting the duration of the entire universe.
(See Dembski 1998:175—223 for a rigorous
calculation of this “Universal Probability
Bound’’; See also Axe 2004.) Yet, second,
fossil data (Bowring et al. 1993, 1998a:1,
1998b:40; Kerr 1993; Monastersky 1993),
and even molecular analyses supporting
deep divergence (Wray et al. 1996), suggest
that the duration of the Cambrian explosion
(between 5—10 X 10° and, at most, 7 X 107
years) is far smaller than that of the entire
universe (1.3—2 X 10!° years). Third, DNA
mutation rates are far too low to generate
the novel genes and proteins necessary to
building the Cambrian animals, given the
most probable duration of the explosion as
determined by fossil studies (Conway Mor-
ris 1998b). As Ohno (1996:8475) notes,
even a mutation rate of 10~° per base pair
per year results in only a 1% change in the
sequence of a given section of DNA in 10
million years. Thus, he argues that muta-
tional divergence of pre-existing genes can-
not explain the origin of the Cambrian
forms in that time.*

*To solve this problem Ohno himself proposes the
existence of a hypothetical ancestral form that pos-
sessed virtually all the genetic information necessary
to produce the new body plans of the Cambrian ani-
mals. He asserts that this ancestor and its “‘panani-
malian genome” might have arisen several hundred
million years before the Cambrian explosion. On this
view, each of the different Cambrian animals would
have possessed virtually identical genomes, albeit with
considerable latent and unexpressed capacity in the
case of each individual form (Ohno 1996:8475—8478).
While this proposal might help explain the origin of
the Cambrian animal forms by reference to pre-exist-
ing genetic information, it does not solve, but instead
merely displaces, the problem of the origin of the ge-
netic information necessary to produce these new
forms.

VOLUME 117, NUMBER 2

The selection/mutation mechanism faces
another probabilistic obstacle. The animals
that arise in the Cambrian exhibit struc-
tures that would have required many new
types of cells, each of which would have
required many novel proteins to perform
their specialized functions. Further, new
cell types require systems of proteins that
must, as a condition of functioning, act in
close coordination with one another. The
unit of selection in such systems ascends
to the system as a whole. Natural selection
selects for functional advantage. But new
cell types require whole systems of pro-
teins to perform their distinctive functions.
In such cases, natural selection cannot con-
tribute to the process of information gen-
eration until after the information neces-
sary to build the requisite system of pro-
teins has arisen. Thus random variations
must, again, do the work of information
generation—and now not simply for one
protein, but for many proteins arising at
nearly the same time. Yet the odds of this
occurring by chance alone are, of course,
far smaller than the odds of the chance or-
igin of a single gene or protein—so small
in fact as to render the chance origin of the
genetic information necessary to build a
new cell type (a necessary but not suffi-
cient condition of building a new body
plan) problematic given even the most op-
timistic estimates for the duration of the
Cambrian explosion.

Dawkins (1986:139) has noted that sci-
entific theories can rely on only so much
“‘luck”’ before they cease to be credible.
The neutral theory of evolution, which, by
its own logic, prevents natural selection
from playing a role in generating genetic
information until after the fact, relies on
entirely too much luck. The sensitivity of
proteins to functional loss, the need for
long proteins to build new cell types and
animals, the need for whole new systems
of proteins to service new cell types, the
probable brevity of the Cambrian explo-
sion relative to mutation rates—all suggest
the immense improbability (and implausi-

221

bility) of any scenario for the origination
of Cambrian genetic information that relies
upon random variation alone unassisted by
natural selection.

Yet the neutral theory requires novel
genes and proteins to arise—essentially—
by random mutation alone. Adaptive advan-
tage accrues after the generation of new
functional genes and proteins. Thus, natural
selection cannot play a role until new in-
formation-bearing molecules have indepen-
dently arisen. Thus neutral theorists envi-
sion the need to scale the steep face of a
Dawkins-style precipice of which there is
no gradually sloping backside—a situation
that, by Dawkins’ own logic, is probabilis-
tically untenable.

In the second scenario, neo-Darwinists
envision novel genes and proteins arising
by numerous successive mutations in the
preexisting genetic text that codes for pro-
teins. To adapt Dawkins’s metaphor, this
scenario envisions gradually climbing
down one functional peak and then as-
cending another. Yet mutagenesis experi-
ments again suggest a difficulty. Recent
experiments show that, even when explor-
ing a region of sequence space populated
by proteins of a single fold and function,
most multiple-position changes quickly
lead to loss of function (Axe 2000). Yet to
turn one protein into another with a com-
pletely novel structure and function re-
quires specified changes at many sites. In-
deed, the number of changes necessary to
produce a new protein greatly exceeds the
number of changes that will typically pro-
duce functional losses. Given this, the
probability of escaping total functional
loss during a random search for the chang-
es needed to produce a new function is ex-
tremely small—and this probability dimin-
ishes exponentially with each additional
requisite change (Axe 2000). Thus, Axe’s
results imply that, in all probability, ran-
dom searches for novel proteins (through
sequence space) will result in functional
loss long before any novel functional pro-
tein will emerge.

222

Blanco et al. have come to a similar con-
clusion. Using directed mutagenesis, they
have determined that residues in both the
hydrophobic core and on the surface of the
protein play essential roles in determining
protein structure. By sampling intermediate
sequences between two naturally occurring
sequences that adopt different folds, they
found that the intermediate sequences “‘lack
a well defined three-dimensional structure.”
Thus, they conclude that it is unlikely that
a new protein fold would evolve from a
pre-existing fold via a series of folded in-
termediates sequences (Blanco et al. 1999:
741).

Thus, although this second neo-Darwin-
ian scenario has the advantage of starting
with functional genes and proteins, it also
has a lethal disadvantage: any process of
random mutation or rearrangement in the
genome would in all probability generate
nonfunctional intermediate sequences be-
fore fundamentally new functional genes or
proteins would arise. Clearly, nonfunctional
intermediate sequences confer no survival
advantage on their host organisms. Natural
selection favors only functional advantage.
It cannot select or favor nucleotide se-
quences or polypeptide chains that do not
yet perform biological functions, and still
less will it favor sequences that efface or
destroy preexisting function.

Evolving genes and proteins will range
through a series of nonfunctional interme-
diate sequences that natural selection will
not favor or preserve but will, in all prob-
ability, eliminate (Blanco et al. 1999, Axe
2000). When this happens, selection-driven
evolution will cease. At this point, neutral
evolution of the genome (unhinged from se-
lective pressure) may ensue, but, as we
have seen, such a process must overcome
immense probabilistic hurdles, even grant-
ing cosmic time.

Thus, whether one envisions the evolu-
tionary process beginning with a noncoding
region of the genome or a preexisting func-
tional gene, the functional specificity and
complexity of proteins impose very strin-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

gent limitations on the efficacy of mutation
and selection. In the first case, function
must arise first, before natural selection can
act to favor a novel variation. In the second
case, function must be continuously main-
tained in order to prevent deleterious (or le-
thal) consequences to the organism and to
allow further evolution. Yet the complexity
and functional specificity of proteins im-
plies that both these conditions will be ex-
tremely difficult to meet. Therefore, the
neo-Darwinian mechanism appears to be
inadequate to generate the new information
present in the novel genes and proteins that
arise with the Cambrian animals.

Novel Body Plans

The problems with the neo-Darwinian
mechanism run deeper still. In order to ex-
plain the origin of the Cambrian animals,
one must account not only for new proteins
and cell types, but also for the origin of new
body plans. Within the past decade, devel-
opmental biology has dramatically ad-
vanced our understanding of how body
plans are built during ontogeny. In the pro-
cess, it has also uncovered a profound dif-
ficulty for neo-Darwinism.

Significant morphological change in or-
ganisms requires attention to timing. Mu-
tations in genes that are expressed late in
the development of an organism will not
affect the body plan. Mutations expressed
early in development, however, could con-
ceivably produce significant morphological
change (Arthur 1997:21). Thus, events ex-
pressed early in the development of organ-
isms have the only realistic chance of pro-
ducing large-scale macroevolutionary
change (Thomson 1992). As John and Mik-
los (1988:309) explain, macroevolutionary
change requires alterations in the very early
stages of ontogenesis.

Yet recent studies in developmental bi-
ology make clear that mutations expressed
early in development typically have dele-
terious effects (Arthur 1997:21). For ex-
ample, when early-acting body plan mol-

VOLUME 117, NUMBER 2

ecules, or morphogens such as_ bicoid
(which helps to set up the anterior-poste-
rior head-to-tail axis in Drosophila), are
perturbed, development shuts down (Niis-
slein-Volhard & Wieschaus 1980, Lawr-
ence & Struhl 1996, Miiller & Newman
2003).° The resulting embryos die. More-
over, there is a good reason for this. If an
engineer modifies the length of the piston
rods in an internal combustion engine
without modifying the crankshaft accord-
ingly, the engine won’t start. Similarly,
processes of development are tightly inte-
grated spatially and temporally such that
changes early in development will require
a host of other coordinated changes in sep-
arate but functionally interrelated devel-
opmental processes downstream. For this
reason, mutations will be much more likely
to be deadly if they disrupt a functionally
deeply-embedded structure such as a spinal
column than if they affect more isolated
anatomical features such as fingers (Kauff-
man 1995:200).

This problem has led to what McDonald
(1983) has called ‘“‘a great Darwinian par-
adox”’ (p. 93). McDonald notes that genes
that are observed to vary within natural
populations do not lead to major adaptive
changes, while genes that could cause ma-
jor changes—the very stuff of macroevo-
lution—apparently do not vary. In other
words, mutations of the kind that macro-
evolution doesn’t need (namely, viable ge-
netic mutations in DNA expressed late in
development) do occur, but those that it
does need (namely, beneficial body plan
mutations expressed early in development)

>Some have suggested that mutations in “master
regulator’ Hox genes might provide the raw material
for body plan morphogenesis. Yet there are two prob-
lems with this proposal. First, Hox gene expression
begins only after the foundation of the body plan has
been established in early embryogenesis (Davidson
2001:66). Second, Hox genes are highly conserved
across many disparate phyla and so cannot account for
the morphological differences that exist between the
phyla (Valentine 2004:88).

223

apparently don’t occur.° According to Dar-
win (1859:108) natural selection cannot act
until favorable variations arise in a popu-
lation. Yet there is no evidence from de-
velopmental genetics that the kind of vari-
ations required by neo-Darwinism—name-
ly, favorable body plan mutations—ever
occur.

Developmental biology has raised anoth-
er formidable problem for the mutation/se-
lection mechanism. Embryological evi-
dence has long shown that DNA does not
wholly determine morphological form
(Goodwin 1985, Niyhout 1990, Sapp 1987,
Miller & Newman 2003), suggesting that
mutations in DNA alone cannot account for
the morphological changes required to build
a new body plan.

DNA helps directs protein synthesis.’ It
also helps to regulate the timing and ex-
pression of the synthesis of various proteins
within cells. Yet, DNA alone does not de-
termine how individual proteins assemble
themselves into larger systems of proteins;
still less does it solely determine how cell
types, tissue types, and organs arrange
themselves into body plans (Harold 1995:

© Notable differences in the developmental pathways
of similar organisms have been observed. For exam-
ple, congeneric species of sea urchins (from genus He-
liocidaris) exhibit striking differences in their devel-
opmental pathways (Raff 1999:110—121). Thus, it
might be argued that such differences show that early
developmental programs can in fact be mutated to pro-
duce new forms. Nevertheless, there are two problems
with this claim. First, there is no direct evidence that
existing differences in sea urchin development arose
by mutation. Second, the observed differences in the
developmental programs of different species of sea ur-
chins do not result in new body plans, but instead in
highly conserved structures. Despite differences in de-
velopmental patterns, the endpoints are the same.
Thus, even if it can be assumed that mutations pro-
duced the differences in developmental pathways, it
must be acknowledged that such changes did not result
in novel form.

7Of course, many post-translation processes of
modification also play a role in producing a functional
protein. Such processes make it impossible to predict
a protein’s final sequencing from its corresponding
gene sequence alone (Sarkar 1996:199—202).

224

2774, Moss 2004). Instead, other factors—
such as the three-dimensional structure and
organization of the cell membrane and cy-
toskeleton and the spatial architecture of the
fertilized egg—play important roles in de-
termining body plan formation during em-
bryogenesis.

For example, the structure and location
of the cytoskeleton influence the patterning
of embryos. Arrays of microtubules help to
distribute the essential proteins used during
development to their correct locations in the
cell. Of course, microtubules themselves
are made of many protein subunits. Nev-
ertheless, like bricks that can be used to as-
semble many different structures, the tu-
bulin subunits in the cell’s microtubules are
identical to one another. Thus, neither the
tubulin subunits nor the genes that produce
them account for the different shape of mi-
crotubule arrays that distinguish different
kinds of embryos and developmental path-
ways. Instead, the structure of the micro-
tubule array itself is determined by the lo-
cation and arrangement of its subunits, not
the properties of the subunits themselves.
For this reason, it is not possible to predict
the structure of the cytoskeleton of the cell
from the characteristics of the protein con-
stituents that form that structure (Harold
2001:125).

Two analogies may help further clarify
the point. At a building site, builders will
make use of many materials: lumber, wires,
nails, drywall, piping, and windows. Yet
building materials do not determine the
floor plan of the house, or the arrangement
of houses in a neighborhood. Similarly,
electronic circuits are composed of many
components, such as resistors, capacitors,
and transistors. But such lower-level com-
ponents do not determine their own ar-
rangement in an integrated circuit. Biolog-
ical systems also depend on hierarchical ar-
rangements of parts. Genes and proteins are
made from simple building blocks—nucle-
otide bases and amino acids—arranged in
specific ways. Cell types are made of,
among other things, systems of specialized

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

proteins. Organs are made of specialized ar-
rangements of cell types and tissues. And
body plans comprise specific arrangements
of specialized organs. Yet, clearly, the prop-
erties of individual proteins (or, indeed, the
lower-level parts in the hierarchy generally)
do not fully determine the organization of
the higher-level structures and organization-
al patterns (Harold 2001:125). It follows
that the genetic information that codes for
proteins does not determine these higher-
level structures either.

These considerations pose another chal-
lenge to the sufficiency of the neo-Darwin-
ian mechanism. Neo-Darwinism seeks to
explain the origin of new information,
form, and structure as a result of selection
acting on randomly arising variation at a
very low level within the biological hier-
archy, namely, within the genetic text. Yet
major morphological innovations depend
on a specificity of arrangement at a much
higher level of the organizational hierarchy,
a level that DNA alone does not determine.
Yet if DNA is not wholly responsible for
body plan morphogenesis, then DNA se-
quences can mutate indefinitely, without re-
gard to realistic probabilistic limits, and still
not produce a new body plan. Thus, the
mechanism of natural selection acting on
random mutations in DNA cannot in prin-
ciple generate novel body plans, including
those that first arose in the Cambrian ex-
plosion.

Of course, it could be argued that, while
many single proteins do not by themselves
determine cellular structures and/or body
plans, proteins acting in concert with other
proteins or suites of proteins could deter-
mine such higher-level form. For example,
it might be pointed out that the tubulin sub-
units (cited above) are assembled by other
helper proteins—gene products—called Mi-
crotubule Associated Proteins (MAPS).
This might seem to suggest that genes and
gene products alone do suffice to determine
the development of the three-dimensional
structure of the cytoskeleton.

Yet, MAPS, and indeed many other nec-

VOLUME 117, NUMBER 2

essary proteins, are only part of the story.
The location of specified target sites on the
interior of the cell membrane also helps to
determine the shape of the cytoskeleton.
Similarly, so does the position and structure
of the centrosome which nucleates the mi-
crotubules that form the cytoskeleton.
While both the membrane targets and the
centrosomes are made of proteins, the lo-
cation and form of these structures is not
wholly determined by the proteins that form
them. Indeed, centrosome structure and
membrane patterns as a whole convey
three-dimensional structural information
that helps determine the structure of the cy-
toskeleton and the location of its subunits
(McNiven & Porter 1992:313—329). More-
over, the centrioles that compose the cen-
trosomes replicate independently of DNA
replication (Lange et al. 2000:235—249,
Marshall & Rosenbaum 2000:187—205).
The daughter centriole receives its form
from the overall structure of the mother
centriole, not from the individual gene
products that constitute it (Lange et al.
2000). In ciliates, microsurgery on cell
membranes can produce heritable changes
in membrane patterns, even though the
DNA of the ciliates has not been altered
(Sonneborn 1970:1—13, Frankel 1980:607—
623; Nanney 1983:163—170). This suggests
that membrane patterns (as opposed to
membrane constituents) are impressed di-
rectly on daughter cells. In both cases, form
is transmitted from parent three-dimension-
al structures to daughter three-dimensional
structures directly and is not wholly con-
tained in constituent proteins or genetic in-
formation (Moss 2004).

Thus, in each new generation, the form
and structure of the cell arises as the result
of both gene products and pre-existing
three-dimensional structure and organiza-
tion. Cellular structures are built from pro-
teins, but proteins find their way to correct
locations in part because of pre-existing
three-dimensional patterns and organization
inherent in cellular structures. Pre-existing
three-dimensional form present in the pre-

225

ceding generation (whether inherent in the
cell membrane, the centrosomes, the cyto-
skeleton or other features of the fertilized
egg) contributes to the production of form
in the next generation. Neither structural
proteins alone, nor the genes that code for
them, are sufficient to determine the three-
dimensional shape and structure of the en-
tities they form. Gene products provide nec-
essary, but not sufficient conditions, for the
development of three-dimensional structure
within cells, organs and body plans (Harold
1995:2767). But if this is so, then natural
selection acting on genetic variation alone
cannot produce the new forms that arise in
history of life.

Self-Organizational Models

Of course, neo-Darwinism is not the only
evolutionary theory for explaining the ori-
gin of novel biological form. Kauffman
(1995) doubts the efficacy of the mutation/
selection mechanism. Nevertheless, he has
advanced a self-organizational theory to ac-
count for the emergence of new form, and
presumably the information necessary to
generate it. Whereas neo-Darwinism at-
tempts to explain new form as the conse-
quence of selection acting on random mu-
tation, Kauffman suggests that selection
acts, not mainly on random variations, but
on emergent patterns of order that self-
organize via the laws of nature.

Kauffman (1995:47—92) illustrates how
this might work with various model sys-
tems in a computer environment. In one, he
conceives a system of buttons connected by
strings. Buttons represent novel genes or
gene products; strings represent the law-like
forces of interaction that obtain between
gene products—i.e., proteins. Kauffman
suggests that when the complexity of the
system (as represented by the number of
buttons and strings) reaches a critical
threshold, new modes of organization can
arise in the system “‘for free’? —that is, nat-
urally and spontaneously—after the manner
of a phase transition in chemistry.

226

Another model that Kauffman develops
is a system of interconnected lights. Each
light can flash in a variety of states—on,
off, twinkling, etc. Since there is more than
one possible state for each light, and many
lights, there are a vast number of possible
states that the system can adopt. Further, in
his system, rules determine how past states
will influence future states. Kauffman as-
serts that, as a result of these rules, the sys-
tem will, if properly tuned, eventually pro-
duce a kind of order in which a few basic
patterns of light activity recur with greater-
than-random frequency. Since these actual
patterns of light activity represent a small
portion of the total number of possible
states in which the system can reside, Kauf-
man seems to imply that self-organizational
laws might similarly result in highly im-
probable biological outcomes—perhaps
even sequences (of bases or amino acids)
within a much larger sequence space of
possibilities.

Do these simulations of self-organiza-
tional processes accurately model the origin
of novel genetic information? It is hard to
think so.

First, in both examples, Kaufmann pre-
supposes but does not explain significant
sources of preexisting information. In his
buttons-and-strings system, the buttons rep-
resent proteins, themselves packets of CSI,
and the result of pre-existing genetic infor-
mation. Where does this information come
from? Kauffman (1995) doesn’t say, but the
origin of such information is an essential
part of what needs to be explained in the
history of life. Similarly, in his light sys-
tem, the order that allegedly arises for “‘for
free’’ actually arises only if the programmer
of the model system “tunes” it in such a
way as to keep it from either (a) generating
an excessively rigid order or (b) devolving
into chaos (pp. 86-88). Yet this necessary
tuning involves an intelligent programmer
selecting certain parameters and excluding
others—that is, inputting information.

Second, Kauffman’s model systems are
not constrained by functional consider-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ations and thus are not analogous to biolog-
ical systems. A system of interconnected
lights governed by pre-programmed rules
may well settle into a small number of pat-
terns within a much larger space of possi-
bilities. But because these patterns have no
function, and need not meet any functional
requirements, they have no specificity anal-
ogous to that present in actual organisms.
Instead, examination of Kauffman’s (1995)
model systems shows that they do not pro-
duce sequences or systems characterized by
specified complexity, but instead by large
amounts of symmetrical order or internal
redundancy interspersed with aperiodicity
or (mere) complexity (pp. 53, 89, 102). Get-
ting a law-governed system to generate re-
petitive patterns of flashing lights, even
with a certain amount of variation, is clearly
interesting, but not biologically relevant.
On the other hand, a system of lights flash-
ing the tithe of a Broadway play would
model a biologically relevant self-organi-
zational process, at least if such a meaning-
ful or functionally specified sequence arose
without intelligent agents previously pro-
gramming the system with equivalent
amounts of CSI. In any case, Kauffman’s
systems do not produce specified complex-
ity, and thus do not offer promising models
for explaining the new genes and proteins
that arose in the Cambrian.

Even so, Kauffman suggests that his self-
organizational models can specifically elu-
cidate aspects of the Cambrian explosion.
According to Kauffman (1995:199—201),
new Cambrian animals emerged as the re-
sult of “long jump”? mutations that estab-
lished new body plans in a discrete rather
than gradual fashion. He also recognizes
that mutations affecting early development
are almost inevitably harmful. Thus, he
concludes that body plans, once established,
will not change, and that any subsequent
evolution must occur within an established
body plan (Kauffman 1995:201). And in-
deed, the fossil record does show a curious
(from a neo-Darwinian point of view) top-
down pattern of appearance, in which high-

VOLUME 117, NUMBER 2

er taxa (and the body plans they represent)
appear first, only later to be followed by the
multiplication of lower taxa representing
variations within those original body de-
signs (Erwin et al. 1987, Lewin 1988, Val-
entine & Jablonski 2003:518). Further, as
Kauffman expects, body plans appear sud-
denly and persist without significant modi-
fication over time.

But here, again, Kauffman begs the most
important question, which is: what produc-
es the new Cambrian body plans in the first
place? Granted, he invokes “long jump mu-
tations” to explain this, but he identifies no
specific self-organizational process that can
produce such mutations. Moreover, he con-
cedes a principle that undermines the plau-
sibility of his own proposal. Kauffman ac-
knowledges that mutations that occur early
in development are almost inevitably dele-
terious. Yet developmental biologists know
that these are the only kind of mutations
that have a realistic chance of producing
large-scale evolutionary change—1i.e., the
big jumps that Kauffman invokes. Though
Kauffman repudiates the neo-Darwinian re-
liance upon random mutations in favor of
self-organizing order, in the end, he must
invoke the most implausible kind of ran-
dom mutation in order to provide a self-
organizational account of the new Cambri-
an body plans. Clearly, his model is not suf-
ficient.

Punctuated Equilibrium

Of course, still other causal explanations
have been proposed. During the 1970s, the
paleontologists Eldredge and Gould (1972)
proposed the theory of evolution by punc-
tuated equilibrium in order to account for a
pervasive pattern of ““sudden appearance”’
and “‘stasis’”’ in the fossil record. Though
advocates of punctuated equilibrium were
mainly seeking to describe the fossil record
more accurately than earlier gradualist neo-
Darwinian models had done, they did also
propose a mechanism—known as species
selection—by which the large morphologi-

227

cal jumps evident in fossil record might
have been produced. According to punctua-
tionalists, natural selection functions more
as a mechanism for selecting the fittest spe-
cies rather than the most-fit individual
among a species. Accordingly, on this mod-
el, morphological change should occur in
larger, more discrete intervals than it would
given a traditional neo-Darwinian under-
standing.

Despite its virtues as a descriptive model
of the history of life, punctuated equilibri-
um has been widely criticized for failing to
provide a mechanism sufficient to produce
the novel form characteristic of higher tax-
onomic groups. For one thing, critics have
noted that the proposed mechanism of
punctuated evolutionary change simply
lacked the raw material upon which to
work. As Valentine and Erwin (1987) note,
the fossil record fails to document a large
pool of species prior to the Cambrian. Yet
the proposed mechanism of species selec-
tion requires just such a pool of species
upon which to act. Thus, they conclude that
the mechanism of species selection proba-
bly does not resolve the problem of the or-
igin of the higher taxonomic groups (p.
96).° Further, punctuated equilibrium has
not addressed the more specific and fun-
damental problem of explaining the origin
of the new biological information (whether
genetic or epigenetic) necessary to produce
novel biological form. Advocates of punc-
tuated equilibrium might assume that the
new species (upon which natural selection
acts) arise by known micro-evolutionary
processes of speciation (such as founder ef-

8 Erwin (2004:21), although friendly to the possibil-
ity of species selection, argues that Gould provides lit-
tle evidence for its existence. ““The difficulty” writes
Erwin of species selection, “... is that we must rely
on Gould’s arguments for theoretical plausibility and
sufficient relative frequency. Rarely is a mass of data
presented to justify and support Gould’s conclusion.”
Indeed, Gould (2002) himself admitted that species se-
lection remains largely a hypothetical construct: “I
freely admit that well-documented cases of species se-
lection do not permeate the literature” (p. 710).

228

fect, genetic drift or bottleneck effect) that
do not necessarily depend upon mutations
to produce adaptive changes. But, in that
case, the theory lacks an account of how
the specifically higher taxa arise. Species
selection will only produce more fit species.
On the other hand, if punctuationalists as-
sume that processes of genetic mutation can
produce more fundamental morphological
changes and variations, then their model be-
comes subject to the same problems as neo-
Darwinism (see above). This dilemma is
evident in Gould (2002:710) insofar as his
attempts to explain adaptive complexity in-
evitably employ classical neo-Darwinian
modes of explanation.’

Structuralism

Another attempt to explain the origin of
form has been proposed by the structuralists
such as Gerry Webster and Brian Goodwin
(1984, 1996). These biologists, drawing on
the earlier work of D’Arcy Thompson
(1942), view biological form as the result
of structural constraints imposed upon mat-
ter by morphogenetic rules or laws. For rea-
sons similar to those discussed above, the
structuralists have insisted that these gen-
erative or morphogenetic rules do not reside
in the lower level building materials of or-

° “T do not deny either the wonder, or the powerful
importance, of organized adaptive complexity. I rec-
ognize that we know no mechanism for the origin of
such organismal features other than conventional nat-
ural selection at the organismic level—for the sheer
intricacy and elaboration of good biomechanical de-
sign surely precludes either random production, or in-
cidental origin as a side consequence of active pro-
cesses at other levels’’ (Gould 2002:710). “Thus, we
do not challenge the efficacy or the cardinal impor-
tance of organismal selection. As previously discussed,
I fully agree with Dawkins (1986) and others that one
cannot invoke a higher-level force like species selec-
tion to explain ‘things that organisms do’—in partic-
ular, the stunning panoply of organismic adaptations
that has always motivated our sense of wonder about
the natural world, and that Darwin (1859) described,
in one of his most famous lines (3), as ‘that perfection
of structure and coadaptation which most justly excites
our admiration’ (Gould 2002:886).

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ganisms, whether in genes or proteins.
Webster and Goodwin (1984:510—511) fur-
ther envision morphogenetic rules or laws
operating ahistorically, similar to the way
in which gravitational or electro-magnetic
laws operate. For this reason, structuralists
see phylogeny as of secondary importance
in understanding the origin of the higher
taxa, though they think that transformations
of form can occur. For structuralists, con-
straints on the arrangement of matter arise
not mainly as the result of historical contin-
gencies—such as environmental changes or
genetic mutations—but instead because of
the continuous ahistorical operation of fun-
damental laws of form—laws that organize
or inform matter.

While this approach avoids many of the
difficulties currently afflicting neo-Darwin-
ism (in particular those associated with its
““genocentricity’’), critics (such as Maynard
Smith 1986) of structuralism have argued
that the structuralist explanation of form
lacks specificity. They note that structural-
ists have been unable to say just where laws
of form reside—whether in the universe, or
in every possible world, or in organisms as
a whole, or in just some part of organisms.
Further, according to structuralists, morpho-
genetic laws are mathematical in character.
Yet, structuralists have yet to specify the
mathematical formulae that determine bio-
logical forms.

Others (Yockey 1992; Polanyi 1967,
1968; Meyer 2003) have questioned wheth-
er physical laws could in principle generate
the kind of complexity that characterizes bi-
ological systems. Structuralists envision the
existence of biological laws that produce
form in much the same way that physical
laws produce form. Yet the forms that phys-
icists regard as manifestations of underlying
laws are characterized by large amounts of
symmetric or redundant order, by relatively
simple patterns such as vortices or gravi-
tational fields or magnetic lines of force. In-
deed, physical laws are typically expressed
as differential equations (or algorithms) that
almost by definition describe recurring phe-

VOLUME 117, NUMBER 2

nomena—patterns of compressible “order”
not “complexity” as defined by algorithmic
information theory (Yockey 1992:77-—83).
Biological forms, by contrast, manifest
greater complexity and derive in ontogeny
from highly complex initial conditions—
i.e., non-redundant sequences of nucleotide
bases in the genome and other forms of in-
formation expressed in the complex and ir-
regular three-dimensional topography of the
organism or the fertilized egg. Thus, the
kind of form that physical laws produce is
not analogous to biological form—at least
not when compared from the standpoint of
(algorithmic) complexity. Further, physical
laws lack the information content to specify
biology systems. As Polanyi (1967, 1968)
and Yockey (1992:290) have shown, the
laws of physics and chemistry allow, but do
not determine, distinctively biological
modes of organization. In other words, liv-
ing systems are consistent with, but not de-
ducible, from physical-chemical laws
(1992:290).

Of course, biological systems do manifest
some reoccurring patterns, processes and be-
haviors. The same type of organism devel-
ops repeatedly from similar ontogenetic pro-
cesses in the same species. Similar processes
of cell division re-occur in many organisms.
Thus, one might describe certain biological
processes as law-governed. Even so, the ex-
istence of such biological regularities does
not solve the problem of the origin of form
and information, since the recurring process-
es described by such biological laws (if there
be such laws) only occur as the result of pre-
existing stores of (genetic and/or epigenetic)
information and these information-rich ini-
tial conditions impose the constraints that
produce the recurring behavior in biological
systems. (For example, processes of cell di-
vision recur with great frequency in organ-
isms, but depend upon information-rich
DNA and proteins molecules.) In other
words, distinctively biological regularities
depend upon pre-existing biological infor-
mation. Thus, appeals to higher-level biolog-
ical laws presuppose, but do not explain, the

229

origination of the information necessary to
morphogenesis.

Thus, structuralism faces a difficult in
principle dilemma. On the one hand, phys-
ical laws produce very simple redundant
patterns that lack the complexity character-
istic of biological systems. On the other
hand, distinctively biological laws—if there
are such laws—depend upon pre-existing
information-rich structures. In either case,
laws are not good candidates for explaining
the origination of biological form or the in-
formation necessary to produce it.

Cladism: An Artifact of Classification?

Some cladists have advanced another ap-
proach to the problem of the origin of form,
specifically as it arises in the Cambrian.
They have argued that the problem of the
origin of the phyla is an artifact of the clas-
sification system, and therefore, does not
require explanation. Budd and Jensen
(2000), for example, argue that the problem
of the Cambrian explosion resolves itself if
one keeps in mind the cladistic distinction
between “‘stem’’ and “‘crown’”’ groups.
Since crown groups arise whenever new
characters are added to simpler more an-
cestral stem groups during the evolutionary
process, new phyla will inevitably arise
once a new stem group has arisen. Thus,
for Budd and Jensen what requires expla-
nation is not the crown groups correspond-
ing to the new Cambrian phyla, but the ear-
lier more primitive stem groups that pre-
sumably arose deep in the Proterozoic. Yet
since these earlier stem groups are by def-
inition less derived, explaining them will be
considerably easier than explaining the or-
igin of the Cambrian animals de novo. In
any case, for Budd and Jensen the explo-
sion of new phyla in the Cambrian does not
require explanation. As they put it, “given
that the early branching points of major
clades is an inevitable result of clade di-
versification, the alleged phenomenon of
the phyla appearing early and remaining
morphologically static is not seen to require

230

particular explanation” (Budd & Jensen
2000:253).

While superficially plausible, perhaps,
Budd and Jensen’s attempt to explain away
the Cambrian explosion begs crucial ques-
tions. Granted, as new characters are added
to existing forms, novel morphology and
greater morphological disparity will likely
result. But what causes new characters to
arise? And how does the information nec-
essary to produce new characters originate?
Budd and Jensen do not specify. Nor can
they say how derived the ancestral forms
are likely to have been, and what processes,
might have been sufficient to produce them.
Instead, they simply assume the sufficiency
of known neo-Darwinian mechanisms
(Budd & Jensen 2000:288). Yet, as shown
above, this assumption is now problematic.
In any case, Budd and Jensen do not ex-
plain what causes the origination of biolog-
ical form and information.

Convergence and Teleological Evolution

More recently, Conway Morris (2000,
2003c) has suggested another possible ex-
planation based on the tendency for evolu-
tion to converge on the same structural
forms during the history of life. Conway
Morris cites numerous examples of organ-
isms that possess very similar forms and
structures, even though such structures are
often built from different material sub-
strates and arise (in ontogeny) by the ex-
pression of very different genes. Given the
extreme improbability of the same struc-
tures arising by random mutation and se-
lection in disparate phylogenies, Conway
Morris argues that the pervasiveness of
convergent structures suggests that evolu-
tion may be in some way “channeled”’ to-
ward similar functional and/or structural
endpoints. Such an end-directed under-
standing of evolution, he admits, raises the
controversial prospect of a teleological or
purposive element in the history of life. For
this reason, he argues that the phenomenon
of convergence has received less attention

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

than it might have otherwise. Nevertheless,
he argues that just as physicists have re-
opened the question of design in their dis-
cussions of anthropic fine-tuning, the ubiq-
uity of convergent structures in the history
of life has led some biologists (Denton
1998) to consider extending teleological
thinking to biology. And, indeed, Conway
Morris himself intimates that the evolution-
ary process might be “underpinned by a
purpose”’ (2000:8, 2003b:511).

Conway Morris, of course, considers this
possibility in relation to a very specific as-
pect of the problem of organismal form,
namely, the problem of explaining why the
same forms arise repeatedly in so many dis-
parate lines of decent. But this raises a
question. Could a similar approach shed ex-
planatory light on the more general causal
question that has been addressed in this re-
view? Could the notion of purposive design
help provide a more adequate explanation
for the origin of organismal form generally?
Are there reasons to consider design as an
explanation for the origin of the biological
information necessary to produce the higher
taxa and their corresponding morphological
novelty?

The remainder of this review will suggest
that there are such reasons. In so doing, it
may also help explain why the issue of tel-
eology or design has re-emerged within the
scientific discussion of biological origins
(Denton 1986, 1998: Thaxton et al. 1992;
Kenyon & Mills 1996; Behe 1996, 2004;
Dembski 1998, 2002, 2004; Conway Mor-
ris 2000, 2003a, 2003b; Lonnig 2001; Lon-
nig & Saedler 2002; Nelson & Wells 2003;
Meyer 2003, 2004; Bradley 2004) and why
some scientists and philosophers of science
have considered teleological explanations
for the origin of form and information de-
spite strong methodological prohibitions
against design as a scientific hypothesis
(Gillespie 1979, Lenior 1982:4).

First, the possibility of design as an ex-
planation follows logically from a consid-
eration of the deficiencies of neo-Darwin-
ism and other current theories as explana-

VOLUME 117, NUMBER 2

tions for some of the more striking “‘ap-
pearances of design” in biological systems.
Neo-Darwinists such as Ayala (1994:5),
Dawkins (1986:1), Mayr (1982:xi—x11) and
Lewontin (1978) have long acknowledged
that organisms appear to have been de-
signed. Of course, neo-Darwinists assert
that what Ayala (1994:5) calls the “‘obvious
design” of living things is only apparent
since the selection/mutation mechanism can
explain the origin of complex form and or-
ganization in living systems without an ap-
peal to a designing agent. Indeed, neo-Dar-
winists affirm that mutation and selection—
and perhaps other similarly undirected
mechanisms—are fully sufficient to explain
the appearance of design in biology. Self-
organizational theorists and punctuational-
ists modify this claim, but affirm its essen-
tial tenet. Self-organization theorists argue
that natural selection acting on self-organiz-
ing order can explain the complexity of liv-
ing things—again, without any appeal to
design. Punctuationalists similarly envision
natural selection acting on newly arising
species with no actual design involved.

And clearly, the neo-Darwinian mecha-
nism does explain many appearances of de-
sign, such as the adaptation of organisms to
specialized environments that attracted the
interest of 19th century biologists. More
specifically, known micro-evolutionary pro-
cesses appear quite sufficient to account for
changes in the size of Galapagos finch
beaks that have occurred in response to var-
iations in annual rainfall and available food
supplies (Weiner 1994, Grant 1999).

But does neo-Darwinism, or any other
fully materialistic model, explain all ap-
pearances of design in biology, including
the body plans and information that char-
acterize living systems? Arguably, biologi-
cal forms—such as the structure of a cham-
bered nautilus, the organization of a trilo-
bite, the functional integration of parts in
an eye or molecular machine—attract our
attention in part because the organized
complexity of such systems seems reminis-
cent of our own designs. Yet, this review

231

has argued that neo-Darwinism does not
adequately account for the origin of all ap-
pearances of design, especially if one con-
siders animal body plans, and the informa-
tion necessary to construct them, as espe-
cially striking examples of the appearance
of design in living systems. Indeed, Dawk-
ins (1995:11) and Gates (1996:228) have
noted that genetic information bears an un-
canny resemblance to computer software or
machine code. For this reason, the presence
of CSI in living organisms, and the discon-
tinuous increases of CSI that occurred dur-
ing events such as the Cambrian explosion,
appears at least suggestive of design.

Does neo-Darwinism or any other purely
materialistic model of morphogenesis ac-
count for the origin of the genetic and other
forms of CSI necessary to produce novel
organismal form? If not, as this review has
argued, could the emergence of novel in-
formation-rich genes, proteins, cell types
and body plans have resulted from actual
design, rather than a purposeless process
that merely mimics the powers of a design-
ing intelligence? The logic of neo-Darwin-
ism, with its specific claim to have account-
ed for the appearance of design, would it-
self seem to open the door to this possibil-
ity. Indeed, the historical formulation of
Darwinism in dialectical opposition to the
design hypothesis (Gillespie 1979), coupled
with neo-Darwinism’s inability to account
for many salient appearances of design in-
cluding the emergence of form and infor-
mation, would seem logically to re-open the
possibility of actual (as opposed to appar-
ent) design in the history of life.

A second reason for considering design
as an explanation for these phenomena fol-
lows from the importance of explanatory
power to scientific theory evaluation and
from a consideration of the potential ex-
planatory power of the design hypothesis.
Studies in the methodology and philosophy
of science have shown that many scientific
theories, particularly in the historical sci-
ences, are formulated and justified as infer-
ences to the best explanation (Lipton 1991:

232

32-88, Brush 1989:1124-1129, Sober
2000:44). Historical scientists, in particular,
assess or test competing hypotheses by
evaluating which hypothesis would, if true,
provide the best explanation for some set of
relevant data (Meyer 1991, 2002; Cleland
2001:987-989, 2002:474—496).'° Those

'0 Theories in the historical sciences typically make
claims about what happened in the past, or what hap-
pened in the past to cause particular events to occur
(Meyer 1991:57—72). For this reason, historical sci-
entific theories are rarely tested by making predictions
about what will occur under controlled laboratory con-
ditions (Cleland 2001:987, 2002:474—496). Instead,
such theories are usually tested by comparing their ex-
planatory power against that of their competitors with
respect to already known facts. Even in the case in
which historical theories make claims about past caus-
es they usually do so on the basis of pre-existing
knowledge of cause and effect relationships. Neverthe-
less, prediction may play a limited role in testing his-
torical scientific theories since such theories may have
implications as to what kind of evidence is likely to
emerge in the future. For example, neo-Darwinism af-
firms that new functional sections of the genome arise
by trial and error process of mutation and subsequent
selection. For this reason, historically many neo-Dar-
winists expected or predicted that the large non-coding
regions of the genome—so-called “junk DNA—would
lack function altogether (Orgel & Crick 1980). On this
line of thinking, the non-functional sections of the ge-
nome represent nature’s failed experiments that remain
in the genome as a kind of artifact of the past activity
of the mutation and selection process. Advocates of
the design hypotheses on the other hand, would have
predicted that non-coding regions of the genome might
well reveal hidden functions, not only because design
theorists do not think that new genetic information
arises by a trial and error process of mutation and se-
lection, but also because designed systems are often
functionally polyvalent. Even so, as new studies reveal
more about the functions performed by the non-coding
regions of the genome (Gibbs 2003), the design hy-
pothesis can no longer be said to make this claim in
the form of a specifically future-oriented prediction.
Instead, the design hypothesis might be said to gain
confirmation or support from its ability to explain this
now known evidence, albeit after the fact. Of course,
neo-Darwinists might also amend their original pre-
diction using various auxiliary hypotheses to explain
away the presence of newly discovered functions in
the non-coding regions of DNA. In both cases, consid-
erations of ex post facto explanatory power re-emerge
as central to assessing and testing competing historical
theories.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

with greater explanatory power are typical-
ly judged to be better, more probably true,
theories. Darwin (1896:437) used this
method of reasoning in defending his the-
ory of universal common descent. More-
over, contemporary studies on the method
of “inference to the best explanation” have
shown that determining which among a set
of competing possible explanations consti-
tutes the best depends upon judgments
about the causal adequacy, or “causal pow-
ers,” of competing explanatory entities
(Lipton 1991:32—88). In the historical sci-
ences, uniformitarian and/or actualistic
(Gould 1965, Simpson 1970, Rutten 1971,
Hooykaas 1975) canons of method suggest
that judgments about causal adequacy
should derive from our present knowledge
of cause and effect relationships. For his-
torical scientists, “‘the present is the key to
the past’ means that present experience-
based knowledge of cause and effect rela-
tionships typically guides the assessment of
the plausibility of proposed causes of past
events.

Yet it is precisely for this reason that cur-
rent advocates of the design hypothesis
want to reconsider design as an explanation
for the origin of biological form and infor-
mation. This review, and much of the lit-
erature it has surveyed, suggests that four
of the most prominent models for explain-
ing the origin of biological form fail to pro-
vide adequate causal explanations for the
discontinuous increases of CSI that are re-
quired to produce novel morphologies. Yet,
we have repeated experience of rational and
conscious agents—in particular ourselves—
generating or causing increases in complex
specified information, both in the form of
sequence-specific lines of code and in the
form of hierarchically arranged systems of
parts.

In the first place, intelligent human
agents—in virtue of their rationality and
consciousness—have demonstrated the
power to produce information in the form
of linear sequence-specific arrangements of
characters. Indeed, experience affirms that

VOLUME 117, NUMBER 2

information of this type routinely arises
from the activity of intelligent agents. A
computer user who traces the information
on a screen back to its source invariably
comes to a mind—that of a software engi-
neer or programmer. The information in a
book or inscription ultimately derives from
a writer or scribe—from a mental, rather
than a strictly material, cause. Our experi-
ence-based knowledge of information-flow
confirms that systems with large amounts of
specified complexity (especially codes and
languages) invariably originate from an in-
telligent source—from a mind or personal
agent. As Quastler (1964) put it, the “‘cre-
ation of new information is habitually as-
sociated with conscious activity” (p. 16).
Experience teaches this obvious truth.
Further, the highly specified hierarchical
arrangements of parts in animal body plans
also suggest design, again because of our
experience of the kinds of features and sys-
tems that designers can and do produce. At
every level of the biological hierarchy, or-
ganisms require specified and highly im-
probable arrangements of lower-level con-
stituents in order to maintain their form and
function. Genes require specified arrange-
ments of nucleotide bases; proteins require
specified arrangements of amino acids; new
cell types require specified arrangements of
systems of proteins; body plans require spe-
cialized arrangements of cell types and or-
gans. Organisms not only contain informa-
tion-rich components (such as proteins and
genes), but they comprise information-rich
arrangements of those components and the
systems that comprise them. Yet we know,
based on our present experience of cause
and effect relationships, that design engi-
neers—possessing purposive intelligence
and rationality—have the ability to produce
information-rich hierarchies in which both
individual modules and the arrangements of
those modules exhibit complexity and spec-
ificity—information so defined. Individual
transistors, resistors, and capacitors exhibit
considerable complexity and specificity of
design; at a higher level of organization,

233

their specific arrangement within an inte-
grated circuit represents additional infor-
mation and reflects further design. Con-
scious and rational agents have, as part of
their powers of purposive intelligence, the
capacity to design information-rich parts
and to organize those parts into functional
information-rich systems and _ hierarchies.
Further, we know of no other causal entity
or process that has this capacity. Clearly,
we have good reason to doubt that mutation
and selection, self-organizational processes
or laws of nature, can produce the infor-
mation-rich components, systems, and body
plans necessary to explain the origination
of morphological novelty such as that
which arises in the Cambrian period.

There is a third reason to consider pur-
pose or design as an explanation for the or-
igin of biological form and information:
purposive agents have just those necessary
powers that natural selection lacks as a con-
dition of its causal adequacy. At several
points in the previous analysis, we saw that
natural selection lacked the ability to gen-
erate novel information precisely because it
can only act after new functional CSI has
arisen. Natural selection can favor new pro-
teins, and genes, but only after they per-
form some function. The job of generating
new functional genes, proteins and systems
of proteins therefore falls entirely to ran-
dom mutations. Yet without functional cri-
teria to guide a search through the space of
possible sequences, random variation is
probabilistically doomed. What is needed is
not just a source of variation (i.e., the free-
dom to search a space of possibilities) or a
mode of selection that can operate after the
fact of a successful search, but instead a
means of selection that (a) operates during
a search—before success—and that (b) is
guided by information about, or knowledge
of, a functional target.

Demonstration of this requirement has
come from an unlikely quarter: genetic al-
gorithms. Genetic algorithms are programs
that allegedly simulate the creative power
of mutation and selection. Dawkins and

234

Ktippers, for example, have developed
computer programs that putatively simulate
the production of genetic information by
mutation and natural selection (Dawkins
1986:47—49, Ktippers 1987:355—369). Nev-
ertheless, as shown elsewhere (Meyer 1998:
127-128, 2003:247—248), these programs
only succeed by the illicit expedient of pro-
viding the computer with a “target se-
quence”’ and then treating relatively greater
proximity to future function (i.e., the target
sequence), not actual present function, as a
selection criterion. As Berlinski (2000) has
argued, genetic algorithms need something
akin to a “forward looking memory” in or-
der to succeed. Yet such foresighted selec-
tion has no analogue in nature. In biology,
where differential survival depends upon
maintaining function, selection cannot oc-
cur before new functional sequences arise.
Natural selection lacks foresight.

What natural selection lacks, intelligent
selection—purposive or goal-directed de-
sign—provides. Rational agents can arrange
both matter and symbols with distant goals
in mind. In using language, the human
mind routinely “‘finds’” or generates highly
improbable linguistic sequences to convey
an intended or preconceived idea. In the
process of thought, functional objectives
precede and constrain the selection of
words, sounds and symbols to generate
functional (and indeed meaningful) se-
quences from among a vast ensemble of
meaningless alternative combinations of
sound or symbol (Denton 1986:309-311).
Similarly, the construction of complex tech-
nological objects and products, such as
bridges, circuit boards, engines and soft-
ware, result from the application of goal-
directed constraints (Polanyi 1967, 1968).
Indeed, in all functionally integrated com-
plex systems where the cause is known by
experience or observation, design engineers
or other intelligent agents applied boundary
constraints to limit possibilities in order to
produce improbable forms, sequences or
structures. Rational agents have repeatedly
demonstrated the capacity to constrain the

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

possible to actualize improbable but initial-
ly unrealized future functions. Repeated ex-
perience affirms that intelligent agents
(minds) uniquely possess such causal pow-
ers.

Analysis of the problem of the origin of
biological information, therefore, exposes a
deficiency in the causal powers of natural
selection that corresponds precisely to pow-
ers that agents are uniquely known to pos-
sess. Intelligent agents have foresight. Such
agents can select functional goals before
they exist. They can devise or select mate-
rial means to accomplish those ends from
among an array of possibilities and then ac-
tualize those goals in accord with a precon-
ceived design plan or set of functional re-
quirements. Rational agents can constrain
combinatorial space with distant outcomes
in mind. The causal powers that natural se-
lection lacks—almost by definition—are as-
sociated with the attributes of conscious-
ness and rationality—with purposive intel-
ligence. Thus, by invoking design to ex-
plain the origin of new _ biological
information, contemporary design theorists
are not positing an arbitrary explanatory el-
ement unmotivated by a consideration of
the evidence. Instead, they are positing an
entity possessing precisely the attributes
and causal powers that the phenomenon in
question requires as a condition of its pro-
duction and explanation.

Conclusion

An experience-based analysis of the
causal powers of various explanatory hy-
potheses suggests purposive or intelligent
design as a causally adequate—and perhaps
the most causally adequate—explanation
for the origin of the complex specified in-
formation required to build the Cambrian
animals and the novel forms they represent.
For this reason, recent scientific interest in
the design hypothesis is unlikely to abate as
biologists continue to wrestle with the prob-
lem of the origination of biological form
and the higher taxa.

VOLUME 117, NUMBER 2

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opmental and evolutionary biology. The M.I.T.
Press, Cambridge, Massachusetts.

Woese, C. 1998. The universal ancestor.—Proceedings
of the National Academy of Sciences, U.S.A.
95:6854—6859.

Wray, G. A., J. S. Levinton, & L. H. Shapiro. 1996.
Molecular evidence for deep Precambrian di-
vergences among metazoan phyla. Science
274:568-573.

Yockey, H. P. 1978. A calculation of the probability of
spontaneous biogenesis by information theo-
ry.—Journal of Theoretical Biology 67:377—
398.

. 1992. Information theory and molecular bi-

ology. Cambridge University Press, Cambridge,

United Kingdom.

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INFORMATION FOR CONTRIBUTORS

Submission of manuscripts.—Submit three copies of each manuscript in the style of the
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CONTENTS

Pseudopaguristes shidarai, a new species of hermit crab (Crustacea: Decapoda: Diogenidae) from
Japan, the fourth species of the genus Akira Asakura

A new species of Procambarus (Crustacea: Decapoda: Cambaridae) from Veracruz, Mexico
Marilu Lopez-Mejia, Fernando Alvarez, and Luis M. Mejia-Ortiz
Brackenridgia ashleyi, a new species of terrestrial isopod from Tumbling Creek Cave, Missouri
(Isopoda: Oniscidea: Trichoniscidae) Julian J. Lewis
New species and records of Bopyridae (Crustacea: Isopoda) infesting species of the genus Upogebia
(Crustacea: Decapoda: Upogebiidae): the genera Orthione Markham, 1988, and Gyge Cornalia &
Panceri, 1861 John C. Markham

Three new species and a new genus of Farreidae (Porifera: Hexatinellida: Hexactinosida)

Kirk Duplessis and Henry M. Reiswig
The origin of biological information and the higher taxonomic categories Stephen C. Meyer

TITUTION LIBRARIES

iN WN

153

169

176

186

199
213

a ~ a t | yw ear 324X a
BUX
\) 4 PROCEEDINGS of THE

BIOLOGICAL SOCIETY
or WASHINGTON

7 DECEMBER 2004

VOLUME 117
NUMBER 3

THE BIOLOGICAL SOCIETY OF WASHINGTON
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117(3):241. 2004.

STATEMENT FROM THE COUNCIL OF THE BIOLOGICAL
SOCIETY OF WASHINGTON

The paper by Stephen C. Meyer, “The
origin of biological information and the
higher taxonomic categories,” in vol. 117,
no. 2, pp. 213-239 of the Proceedings of
the Biological Society of Washington, was
published at the discretion of the former ed-
itor, Richard v. Sternberg. Contrary to typ-
ical editorial practices, the paper was pub-
lished without review by any associate ed-
itor; Sternberg handled the entire review
process. The Council, which includes offi-
cers, elected councilors, and past presidents,
and the associate editors would have
deemed the paper inappropriate for the pag-
es of the Proceedings because the subject
matter represents such a significant depar-
ture from the nearly purely systematic con-
tent for which this journal has been known
throughout its 122-year history. For the
same reason, the journal will not publish a
rebuttal to the thesis of the paper, the su-

periority of intelligent design (ID) over
evolution as an explanation of the emer-
gence of Cambrian body-plan diversity. The
Council endorses a resolution on ID pub-
lished by the American Association for the
Advancement of Science (www.aaas.org/
news/releases/2002/1 106id2.shtml), which
observes that there is no credible scientific
evidence supporting ID as a testable hy-
pothesis to explain the origin of organic di-
versity. Accordingly, the Meyer paper does
not meet the scientific standards of the Pro-
ceedings.

We have reviewed and revised editorial
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(3):242—250. 2004.

A review of the North American subspecies of the Great Blue Heron

(Ardea herodias)

Robert W. Dickerman

Museum of Southwestern Biology, University of New Mexico, Albuquerque, New Mexico 87131,

U.S.A., e-mail: bobdickm @unm.edu

Abstract.—Geographic variation in the great Blue Heron (Ardea herodias)
was comprehensively reviewed by H. C. Oberholser (1912), who recognized
nine North American subspecies—excluding the so-called Great White Heron
(A. occidentalis = A. h. occidentalis). Oberholser’s revision provided the frame-
work generally followed in subsequent subspecific treatments of this species.
However, Payne’s (1979) brief general summary of this species’ geographic
variation rejects most of these North American taxa, recognizing as valid only
the nominate subspecies and those of the Pacific northwest [A. h. fannini] and
Florida [A. h. occidentalis]. My studies verify that A. h. herodias and A. h.
fannini are taxonomically distinct, along with A. h. wardi in which I include
A. h. treganzai, A. h. hyperonca, and A. h. sanctilucae. In addition, I regard
A. h. lessoni, A. h. adoxa, and A. h. olgista as synonymous with A. h. herodias,
as all are based on migrant specimens of this form. In addition, I suspect Payne
is justified as recognizing the Caribbean A. h. occidentalis as valid, based on
its white plumage and shorter head plumes.

The Great Blue Heron (Ardea herodias)
nests in North America from southeastern
Alaska, southern British Columbia, north-
ern Alberta, central Saskatchewan, northern
Manitoba, northern Ontario, southern Que-
bec, New Brunswick, and Nova Scotia
southward to the Gulf states, southern Flor-
ida; on the coastal lowlands of Mexico
south to Tabasco, Nayarit, and Baja Cali-
fornia; and locally in the Caribbean Basin
(A.O.U. 1998). There are no nesting season
specimens of Great Blue Herons taken be-
tween the Yucatan Peninsula of Mexico and
Venezuela. Oberholser (1912) recognized
nine subspecies over this extensive area,
these being A. h. herodias L., 1758 (type
locality: America [= Hudson Bay, Cana-
da]); A. h. wardi Ridgway, 1882 (Oyster [=
Estero] Bay, Florida; A. h. treganzai Court,
1908 (Egg Island, Great Salt Lake, Utah);
A. h. fannini Chapman, 1901 (Skidegate
[Graham Island], Queen Charlotte Islands,
British Columbia); A. h. hyperonca Ober-

holser, 1912 (Baird [Shasta Co.], Califor-
nia); A. h. sanctilucae Thayer and Bangs,
1912 (Espiritu Santo Island, Baja Califor-
nia); A. h. lessoni Wagler, 1831 (Mexico);
A. h. adoxa Oberholser, 1912 (Curacao);
and A. h. olgista Oberholser, 1912 (San
Clemente Island, California). Not included
in Oberholser’s revision was the so-called
Great White Heron (A. occidentalis), which
is now widely regarded as a white morph
of A. herodias (e.g., A.O.U. 1998). In ad-
dition, he did include the endemic A. h.
cognatus of the Galapago Islands, which is
the only nesting population of this species
outside North America.

Oberholser’s (1912) subspecies were
based on differences in plumage coloration
and measurements among populations,
which in some cases included migrants
from other areas. In fact, although Ober-
holser was aware of both migration and oth-
er forms of dispersal in this species, he ap-
pears to have underestimated the extent of

VOLUME 117, NUMBER 3

this phenomenon. For example, Bond
(1935) found that Oberholser’s A. h. adoxa
from Curacao is based on a series of eight
specimens, all of which are southward mi-
grants of A. h. herodias. In addition, the
adult female holotype (examined) for Ob-
erholser’s (abid.) A. h. olgista from San Cle-
mente Island is also an example of the nom-
inate form, based on its dark coloration and
a wing chord of 433 mm, even though pre-
viously synonomized with the locally nest-
ing A. h. “‘hyperonca” (= A. h. wardi) by
Grinnell and Miller (1944) and Hellmayr
and Conover (1948). Hellmayr and Con-
over (1948) also listed A. lessoni Wagler as
a synomym of A. h. herodias, simply noting
“type in Munich Museum examined.”

Oberholser’s revision provided the
framework generally followed in subse-
quent subspecific treatments such as A. O.
U. (1931, 1957), Peters (1931), Friedmann
et al. (1950), Palmer (1962) and Hancock
and Elliot (1978).

More recently, Payne (1979) has treated
overall geographic variation in the A. her-
odias complex (including A. occidentalis),
although he did so only briefly, generally,
and without measurements or references to
subspecific names. He recognized only
three taxa in North America: the wide-
spread A. h. herodias, A. h. fannini of the
north Pacific Coast, and the white-plum-
aged A. h. occidentalis of the Caribbean Ba-
sin. My revisionary work on the Great Blue
Heron began in an attempt to identify then
recently collected Mexican specimens in
the 1960’s. Since then I have examined
most of the available adult specimens in
North American collections. My findings
generally agree with those of Payne, except
that I also recognize the populations of pal-
er and larger birds of southern and western
North America as A. h. wardi. I did not
examine plumage variation in nesting pop-
ulations of the Caribbean Basin, but these
may constitute a valid subspecies (A. h. oc-
cidentalis) based on the dominance of the
white morph (rare elsewhere). If not rec-
ognizable, then these populations and those

243

of A. h. wardi should be merged under the
older name of A. h. occidentalis.

Methods

Several caveats apply to the museum
specimens used in this study, the first being
the dearth of properly labeled and prepared
adult nesting season skins for studies of
geographic variation among Great Blue
Herons in North America. For example, I
found no nesting season adult males from
Delaware, Virginia, West Virginia, and
Kentucky; only single males from Mary-
land, Tennessee, South Carolina, and Ala-
bama; and only two from North Carolina!
Secondly, many specimens lack informa-
tion on gonad size, weight, and fat condi-
tion, making it difficult to ascertain whether
such birds are likely nesting or are mi-
grants. As a result, one must often assume
that birds are nesting on the basis of col-
lection localities and dates, which can be
complicated by (a) regional differences in
the timing of breeding activities and (b) the
migration and other forms of dispersal in
this species. For example, we know post-
nesting southern populations (A. h. wardi)
can be dispersing northward in the north-
eastern U.S. while northern birds (A. h. her-
odias) are in the process of nesting (Dick-
erman 2002). Whereas coloration and mea-
surements do distinguish these subspecies,
some specimens overlap or intergrade be-
tween the two. As a result, these may be
either included in or excluded from nesting
samples, thus introducing some degree of
bias into the data. In any case, I have ar-
bitrarily set the nesting season for most
North American populations of this species
as April to July, subject to modification
based on specimens’ gonadal condition,
weight, fat levels, coloration, and measure-
ments.

A second caveat with Great Blue Heron
specimens is that the plumage coloration
can be altered by a variety of factors, in-
cluding wear, bleaching, molt stage, chem-
icals used to preserve or protect skins, mu-

244

seum age, and especially staining due to the
leakage and oxidation of body fat. In ad-
dition, winter-taken specimens in the north
may be under greater nutritional stress, so
that they may produce less powder down to
coat the feathers. This in turn would greatly
affect feather color, as the powder-down
coating produces a pale bloom that makes
the plumage appear lighter. In fact, this
same effect can be extreme when the plum-
age is washed and the powder-down is re-
moved (Dickerman 2004).

For my final comparisons of plumage
coloration in Great Blue Heron populations,
I borrowed 26 adult skins taken throughout
North America, representing all of the
mainland forms. All were clean but un-
washed specimens, taken as early in the
nesting season and chronologically recently
as possible. As I found no differences in
plumage color between males and females,
I combined the sexes for these comparisons.
In addition, I measured 214 males and 189
females for the following characters: wing
chord, tail length, exposed culmen, and tar-
sus length (all in mm). After a preliminary
analysis, I have variously grouped these
measurements by subspecies, area, and
sometimes type specimens (Table 1). I then
calculated the sample sizes, ranges, means,
and standard deviations for the four men-
sural characters, as well as performing two-
sample t-tests to determine the significances
(P = 0.05) of differences. I did not analyze
either plumage or mensural variation in oth-
er age classes, because sample sizes were
too small for juveniles and nestlings. Im-
matures were not analyzed.

Results

As did Oberholser (1912), I find Great
Blue Herons can be aggregated into three
distinct North American nesting popula-
tions on the basis of plumage coloration,
exclusive of the white-phased birds of the
Caribbean Basin (A. h. occidentalis). More
specifically, this variation involves the col-
oration of the upper-parts, neck, and wing

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

feathers in adult birds, which ranges from
pale to darker gray. The first of these ag-
gregates consists of the moderately gray
populations to which the name A. h. hero-
dias can be applied. These nest in southern
Canada west to interior southern British
Columbia, then southward in the United
States to eastern Washington, North Dako-
ta, Wisconsin, Indiana, Maryland, and
South Carolina. The second aggregate of
paler populations in the southeastern, cen-
tral, and western U.S. and Mexico that Ob-
erholser (ibid.) assigned to four subspecies,
of which the oldest name is A. h. wardi with
A. h. treganzai, A. h. hyperonca and A. h.
sanctilucae here considered synonyms. And
the third is the darker gray A. h. fannini,
whose range I have recommended be re-
stricted to the coastal region of northwest-
ern British Columbia and adjacent Alaska,
specifically the Queen Charlotte Islands
north to Prince William Sound (Dickerman
2004). However, as noted earlier, the slaty-
black coloration of the holotype (Chapman
1901) is abnormally dark, apparently due to
washing that removed the powder down
coating and thus the paler bloom of the
plumage (Dickerman 2004).

Oberholser (1912) further characterized
nesting Great Blue Herons on the basis of
measurements, which he particularly em-
phasized in allotting pale populations to
four subspecies. Given this, I also assessed
measurements in this species, in which
males generally average larger than females
in nesting populations (Table 1). For ex-
ample, my overall samples reveal that
males are 4.1% larger in wind chord, 3.5%
in tail length, 6.5% in exposed culmen, and
6.5% in tarsus length (including only “‘typ-
ical populations of named forms, excluding
fannini because of small sample size).
However, the sexes overlap in all of these
mensural characters, and t-tests often show
the differences are not significant at the P
= 0.05 level. Nonetheless, it is important to
segregate the sexes when using measure-
ments to allocate specimens to subspecies

VOLUME 117, NUMBER 3

and populations. As for the mensural char-
acters themselves, I found the following:

Wing chord.—Nesting populations with
the longest wings are 4.8% and 5.2% great-
er than those with the shortest in males and
females, respectively. Means are smallest in
A. h. herodias, generally becoming pro-
gressively larger through the populations of
the interior western U.S., the Pacific Coast
region, and Mexico to the southeastern U.S.
(Table 1). However, a notable departure
from this is that A. h. occidentalis has the
wing chord intermediate, as opposed to be-
ing among the largest in the species. T-tests
reveal that A. h. herodias averages signifi-
cantly shorter in wing length than all but
two other North American populations, the
exceptions being males of A. h. fannini and
A. h. “‘treganzai.”” By contrast, the latter is
significantly shorter-winged than all but one
of the A. h. wardi populations, that being
the small Texas sample. All other popula-
tional differences in this character are insig-
nificant, with clinal intergradation being
smoother among females than males.

Tail length.—Nesting populations with
the longest tails are 7.7% and 8.8% greater
than those with the shortest in males and
females, respectively. Means are smallest in
A. h. herodias and become progressively
and significantly larger in Texas/Florida
populations of A. h. wardi, A. h. “‘hyperon-
ca” X A. h. fannini, and A. h. fannini (Table
1). All other populational differences in this
character are insignificant, with rather mo-
saic intergradation occurring among both
males and females.

Exposed culmen.—Nesting populations
with the longest culmens are 29.4% and
31.9% greater than those with the shortest
in males and females, respectively. Means
are smallest in A. h. fannini and then A. h.
“hyperonca”’ X A. h. fannini, each of
which has a significantly shorter culmen
than all other populations of the species
(Table 1). Elsewhere, males average small-
est in A. h. herodias, which differ signifi-
cantly from those with the longest culmens
in Florida, Texas, and eastern Mexican A.

245

h. wardi and A. h. occidentalis. However,
these extremes intergrade circuitously
through A. h. “‘treganzai,”’ A. h. “‘hyperon-
ca”’ and A. h. “‘sanctilucae’’ with a similar
pattern of geographic variation, except that
A. h. occidentalis has a significantly longer
culmen than all but one A. h. wardi (sensu
latu) population—that in eastern Mexico,
which has a sample size of only one!

Tarsus length.—Nesting populations
with the longest tarsi are 33.2% and 27.3%
greater than those with the smallest in
males and females, respectively. Means in
males are shortest in A. h. fannini and then
A. h. “hyperonca”’ X A. h. fannini, each of
which has a significantly shorter tarsus than
all other populations of the species (Table
1). The same is true with females, except
that the means of those two populations are
essentially identical. Elsewhere, males and
females average smallest in A. h. herodias
and A. h.“‘hyperonca”’ which differ signif-
icantly from those with the longest tarsi in
Florida A. h. wardi and A. h. occidentalis.
However, these extremes intergrade circui-
tously through A. h. “treganzai,”’ A. h.
“sanctilucae”’ and Texas/eastern Mexican
populations of A. h. wardi.

Discussion and Conclusions

Based on these findings, I recommend
recognizing three subspecies among North
American nesting populations of the Great
Blue Heron, excluding the white-plumaged
A. h. occidentalis of the Carribbean Basin.

The first is Ardea herodias herodias L,
with its moderately gray plumage and a
nesting range as outlined above (see Results
section). This the most highly migratory of
the subspecies, with birds regularly moving
southward into Central America and the
Caribbean and as far as Belize, Panama,
Colombia, Venezuela, Curacao, and the Do-
minican Republic (also eastward to Ber-
muda). In addition, lesser numbers move
elsewhere, including northward to Hudson
Bay, northern Quebec, Anticosti Island, and
Newfoundland (plus as a vagrant to Green-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

246

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248

land), and west to California, Arizona, New
Mexico, and Colorado. A. h. adoxa and A.
h. olgista of Oberholser (1912) and A. h.
lessoni Wagler were based on migrant A. h.
herodias. Oberholser restricted the type lo-
cality of A. lessoni to the Valley of Mexico.
However, there are no records of the species
ever having nested in the Valley, and Payne
(1979) was correct in just citing Mexico as
the type locality. Hellmayr and Conover
(1948) erroneously placed A. h. olgista in
the synonomy of A. h. hyperonca. However,
the wing chord of the type (433 mm) is far
too short for that population and is even
short for A. h. herodias! It is also darker, as
in the nominate population.

The second subspecies recognized here is
the pale A. h. wardi, which includes A. h.
“terganzai,’’ A. h.“‘hyperonca,”’ and A. h.
“santilucae.’’ Payne (1979) wrote in a foot-
note (p.198) ““The type of wardi was taken
on 5 January 1881. It is not known whether
this was a local breeding bird or a wintering
bird from a more northern population.”
There is no doubt that it was from the local
population. The type of A. h. wardi is a
very large bird (Oberholser 1912, Table 1),
larger in all measurements than any male A.
h. herodias, and it has longer tail, culmen,
and tarsus measurements than any other
male A. h. wardi. Size is largest in the
southeast (Table 1) and smallest in the
Great Basin region (A. h. “‘treganzai’’), and
only extremes can be identified based on
measurements (Dickerman 1992, 2002).
Oberholser described A. h. “hyperonca”’ as
the color of A. h. herodias, but larger. The
type is from northern California and is
somewhat intermediate towards A. h. fan-
nini and indeed inseparable from A. h. her-
odias in color, but it is larger than the larg-
est male of A. h. fannini or of A. h. herodias
(Table 1). However, specimens from central
California in the California Academy of
Science and the Museum of Vertebrate Zo-
ology labeled A. h. hyperonca are insepa-
rable in color from a topotype of A. h. tre-
ganzai, from early nesting season speci-
mens from southern New Mexico and south

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Texas, or from a midwinter specimen of A.
h. wardi from Florida.

Contra Payne’s comment on clinal vari-
ation in size in the east (1979), there are
not enough nesting colony or even nesting
season specimens yet available to fully doc-
ument a cline. As mentioned earlier, there
are no nesting season adult males from Del-
aware, Virginia, West Virginia or Ken-
tucky; only single males from Maryland,
Tennessee, South Carolina, and Alabama:
and only two from North Carolina. Indeed
the best clinal variation in size is on the
west coast, with an increase in culmen and
tarsal length from A. h. fannini in the north,
through an intermediate population in
southern British Colombia and Washington,
to the long-billed, long- legged A. h. “‘hy-
peronca”’ population of California (Dick-
erman 2004).

A. h. wardi may be separated from A. h.
herodias as follows:

1. Neck of A. h. herodias darker, colder
““vinaceous” gray vs. ““warmer,” paler
neck of A. h. wardi..

2. Chestnut of mid-ventral neck stripe is
more extensive and darker in A. h. her-
odias, and usually extends to the area
behind and below the eye, thus faintly
outlining the white of the throat; in A. h.
wardi the area behind and below the eye
is always white; chestnut of mid-neck is
reduced and paler.

3. Dorsum and wings are darker in A. h.
herodias and paler in A. h. wardi.

The third subspecies recognized here is
A. h. fannini, which differs from A. h. her-
odias in being darker gray in color and in
having the exposed culmen and tarsus sig-
nificantly shorter and tail longer (plus wing
in males, Dickerman 2004). A. h. fannini
differs in being notably much darker gray
than A. h. wardi (sensu latu), and in having
the exposed culmen and tarsus significantly
shorter. In addition, males have significant-
ly shorter wings than all A. h. wardi pop-
ulations except A. h. “‘treganzai”’ on the in-
terior western U.S. A. h. fannini seems to

VOLUME 117, NUMBER 3

differ from other Great Blue Herons in that
it perforce fishes much of the time from
rocks rather than wading, as do the other
longer-legged subspecies. It appears to be
largely resident within its nesting range
(contra A.O.U. 1957), with the only extra-
limital specimen being an adult taken at
Wainwright on the Arctic coast of Alaska
(Brock 1959). I know of no specimens of
A. h. fannini (as here defined) from south
of the Queen Charlotte Islands. A. h. fannini
intergrades southward with A. h. “‘hyperon-
ca’ and perhaps A. h. “‘treganzai”’ (both
here = A. h. wardi) in southwestern British
Columbia (including Vancouver Island) and
western Washington (Dickerman 2004). For
example, that population is paler gray as in
A. h. wardi (sensu latu), but it is closer to
A. h. fannini in the shorter exposed culmen,
tarsus, and male wing chord.

Acknowledgments

The author has measured or compared
specimens of Great Blue Herons in over 30
museums, sometimes more than once! He
wishes to express his appreciation for the
many courtesies he has received at the fol-
lowing institutions: American Museum of
Natural History, New York; Academy of
Natural Sciences, Philadelphia; California
Academy of Sciences, San Francisco; Car-
negie Museum of Natural History, Pitts-
burgh; Coleccion Ornitologico Phelps, Ca-
racas; Colorado State University Coopera-
tive Wildlife Research Vertebrate Collec-
tion, Fort Collins; Cornell University
Museum of Vertebrates, Ithaca; Cowan Ver-
tebrate Museum, University of British Co-
lumbia; Delaware Museum of Natural His-
tory, Greenville; Denver Museum of Natu-
ral History, Denver; Donald R. Dickey Col-
lection, University California, Los Angeles;
James Ford Bell Museum of Natural His-
tory, University of Minnesota; James R.
Slater Museum of Natural History, Univer-
sity of Puget Sound; Museum of Natural
History of Los Angeles County, Los An-
geles; Museum of Comparative Zoology,

249

Harvard; Museum of Natural Science, Lou-
isiana State University; Museum of South-
western Biology, University of New Mex-
ico; Museum of Vertebrate Zoology, Uni-
versity of California, Berkeley; National
Museum of Canada, Ottawa; National Mu-
seum Natural History, Washington, D.C;
North Carolina State Museum of Natural
Sciences, Raleigh; Peabody Museum of
Natural History, Yale; Royal British Co-
lumbia Museum, Victoria; Sam Noble Mu-
seum of Natural History, University of
Oklahoma; San Diego Museum of Naturai
History, San Diego; Texas Cooperative
Wildlife Collection, Texas A&M; The Field
Museum, Chicago; Thomas Burke Memo-
rial Washington State Museum, University
of Washington; University of Alaska Mu-
seum, Fairbanks; University of Kansas Mu-
seum of Natural History, Lawrence; Uni-
versity of Nebraska State Museum, Lin-
coln; Western Foundation Vertebrate Zool-
ogy; Camarillo; Virginia Tech Museum
Natural History, Blacksburg; Zoology Mu-
seum, University of Wisconsin; and the pri-
vate collection of the late Allan R. Phillips.
He especially wishes to thank the cura-
tors of collections who kindly shipped to
New York specimens that permitted final
color comparisons at the American Muse-
um of Natural History. These include: Cal-
ifornia Academy of Sciences, San Francis-
co; Carnegie Museum of Natural History,
Pittsburgh; Denver Museum of Natural His-
tory; Los Angeles County Museum of Nat-
ural History; Museum of Southwestern Bi-
ology, Albuquerque; Museum of Vertebrate
Zoology, Berkeley; National Museum of
Natural History, Washington, D.C.; Utah
Museum of Natural History, Salt Lake City;
and the Western Foundation of Vertebrate
Zoology, [then in Los Angeles]. Christine
Blake of the AMNH graciously received
and repacked all specimens. John P. Hub-
bard suffered through several revisions of
this manuscript and improved it greatly.

Literature Cited

American Ornithologists’ Union. 1931. Check-list of
North American birds, 4th edition. Lancaster,
Pennsylvania, 526 pp.

250

. 1957. Check-list of North American birds, 5th
edition. Lord Baltimore Press Inc., Baltimore,
Maryland, 691 pp.

Bond, J. 1935. The status of the Great Blue Heron in
the West Indies—Auk 52:76-77.

Dickerman, R. W. 1992. Northeastern records of Ardea
herodias wardi from the southeastern United
States —Kingbird 42:10-13.

. 2002. An adult Ardea herodias wardi from

the northeast.—Kingbird 52:35-37.

. 2004. Characteristics and distribution of Ar-
dea herodias fannini with comments on the ef-
fects of washing on the holotype. Northwestern
Naturalist 85:130—-133.

Friedmann, H., L. Griscom, & R. T. Moore. 1950. Dis-
tributional check-list of the birds of Mexico,
part 1. Pacific Coast Avifauna 29.

Hancock, J., & H. Elliott. 1978. The herons of the
world. Harper and Row Publ. New York.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Hellmayr, C. E. and H. Conover. 1948. Catalogue of
birds of the Americas. part 1. No. 2. Zool. Se-
ries, Field Mus. Nat. Hist. 8:434 pp.

Grinnell, J.. & A. H. Miller. 1944. The distribution of
the birds of California. Pacific Coast Avifauna
27: 608 pp.

Palmer, R. S. 1962. Handbook of North American
birds, vol. 1. Loons through Flamingos. Yale
University Press, New Haven, Connecticut. 567
Pp.

Payne, R. B. 1979. Ardeidae. In E. Mayr and G. W.
Cottrell, eds., Check-list of birds of the world,
vol. 1, 2nd edition. Mus. Comp. Zool., Cam-
bridge, Massachusetts.

Peters, J. L. 1931. Check-list of birds of the world,
vol. 1. Harvard Press, Cambridge. Massachu-
setts. 345 pp.

Oberholser, H. C. 1912. A revision of the forms of the
Great Blue Heron (A. herodias Linnaeus).—
Proc. U.S. Nat. Mus. 43:531—559.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(3):251—265. 2004.

A new species of Microgale (Lipotyphla: Tenrecidae: Oryzorictinae)
from the Forét des Mikea of southwestern Madagascar

Steven M. Goodman and Voahangy Soarimalala

(SMG) Department of Zoology, Field Museum of Natural History, 1400 Roosevelt Road,
Chicago, Illinois 60605, U.S.A., e-mail: goodman@fmnh.org and WWF-Madagascar, BP 738,
Antananarivo (101), Madagascar, e-mail: sgoodman@wwf.mg;

(VS) Département de Biologie Animale, Université d’ Antananarivo, BP 906, Antananarivo (101),
Madagascar and Ecology Training Program, WWE-Madagascar, BP 738, Antananarivo (101),

Madagascar, e-mail: etp@wwf.mg

Abstract.—A new species of Microgale, M. jenkinsae (Lipotyphla: Tenre-
cidae), is described based on two specimens taken during an early 2003 bio-
logical survey of the Forét des Mikea in southwestern Madagascar. It is distin-
guished from other congeners by numerous pelage, cranial, and dental char-
acters. M. jenkinsae is the fourth known species in this genus confirmed to
occur in the dry western and southern forests of the island. The Forét des
Mikea, the only site M. jenkinsae is known from, is the last remaining block
of a distinctive forest habitat and is under considerable threat from human
habitat degradation. Action needs to be taken to protect this unique region.

Résumé.—Une nouvelle espéce de Microgale, M. jenkinsae (Lipotyphla:
Tenrecidae), est décrite a partir de deux spécimens récoltés au cours d’un in-
ventaire biologique mené au début de l’année 2003 dans la forét des Mikea
située au sud-ouest de Madagascar. On le distingue de ses autres congénéres
par divers caractéres de pelage, craniens et dentaires. M. jenkinsae est la qua-
trieme espéce connue de ce genre dont la présence est confirmée dans les foréts
seches de l’ouest et du sud de Vile. La forét des Mikea, le seul site d’ou M.
jenkinsae a été rapporté, est le dernier bloc d’un habitat forestier distinctif qui
est cependant extrémement menacé par la dégradation de |’habitat perpétrée
par ’ homme. Des actions doivent étre prises pour protéger cette région unique.

On the island of Madagascar there is an
endemic family of Lipotyphla, known as
the Tenrecidae, that represents one of the
most remarkable adaptive radiations found
in living mammals (Olson & Goodman
2003). As currently circumscribed, Micro-
gale (shrew tenrecs) a tenrecid genus in the
subfamily Oryzorictinae, comprises 18 spe-
cies (Jenkins 2003). On the basis of biolog-
ical inventories and associated museum
studies conducted over the past few de-
cades, seven species of Microgale new to
science have been named (Jenkins 1988,
1992, 1993; Jenkins et al. 1996, 1997;
Goodman & Jenkins 1998: Jenkins &

Goodman 1999), although one of these, M.
pulla, has since been synonymized (Jenkins
et al. 1997). Subsequent to the publication
of MacPhee’s (1987) taxonomic revision of
the genus Microgale, there has been a re-
newed interest in the small mammals of
Madagascar. With the advent of pit-fall de-
vices to trap these animals, there has been
a massive increase in available shrew tenrec
specimens. This has lead to a series of pub-
lications refining some of MacPhee’s taxo-
nomic conclusions and a greater under-
standing of intra-specific, particularly age
related, and inter-specific variation amongst
these animals. As witnessed by the recent

45:00

252
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Morombe |, angcky |
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22:00 + “ Befandriana Atsimo
Vorehy Ankazoabo Atsimo
it
2:30 | Collection site Vohibasiaf” |
Bes
Ankiloaka Praesens
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Manombo sa ae ~~ ng omSakarahe
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fe &
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23:30 St. Augustin LS Oni = a
= Betioky
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24:30 =
0) 50 100 150
EE _ ee _
Fig. 1.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Antsiranana

i) Vohemar

Meena
ve )

\ Antananarivo

Morondava
Belo-sur-Mer ‘g

Morombe|

200 kilometers

Map showing the collection locality of Microgale jenkinsae in the Forét des Mikea, the forested

zone between Morombe and Manombo, in southwestern Madagascar (expanded inset).

description of six new valid shrew tenrec
taxa (an increase of 33%), Microgale tax-
onomy is in flux as a result of ongoing bi-
ological inventories and molecular and
morphological studies.

Of the currently recognized 18 species of
Microgale, 15 are restricted to the eastern
and northern moister portions of Madagas-
car where they occur in either forests or
marshes. Of the remaining three species,
two have been collected over the past few
decades in the dry western forests. These
include M. brevicaudata, which occurs
from the northern foothills of the Marojejy
Massif in the northeast, a zone of humid
forest but probably with a marked dry sea-
son, north to Vohemar and the region of
Antsiranana at the north end of Madagascar,
and then south along the west portion of the

island to at least the Onilahy River near To-
liara (Goodman et al. unpublished; Fig. 1).
The second species, M. nasoloi, is known
from two inland isolated forests in south-
western Madagascar in the vicinity of Sa-
karaha—the Analavelona Massif and the
Forét de Vohibasia (Jenkins & Goodman
1999; Fig. 1). M. longicaudata is the third
species falling into this group and has been
collected from both eastern humid forests
and western dry forests. However, M. lon-
gicaudata, as currently defined, includes
several cryptic species and will soon be re-
vised (Olson et al. in press).

MacPhee’s (1987, Fig. 13) map of col-
lecting localities for Microgale included 31
sites in the eastern humid forest where a
total of nine species were trapped, two sites
in the western dry deciduous each with sin-

VOLUME 117, NUMBER 3

gle species (one based on owl pellets), and
three sites in the southern spiny-bush each
with two species (all based on owl pellets).
Although considerable advances have been
made concerning the species richness and
distribution of shrew tenrecs since Mac-
Phee’s important revision, these data indi-
cate a greater diversity of this group in the
more mesic portions of the island. Recent
biological inventories of the western and
southern forests of Madagascar have largely
upheld this view. However, during a 2003
survey of the Forét des Mikea, the region
between Morombe and Manombo (Fig. 1),
we captured a Microgale that represents a
previously undescribed species of shrew
tenrec.

Materials and Methods

Our small mammal collection made in
the Forét des Mikea contains two speci-
mens of Microgale, and in order to deter-
mine their taxonomic identity, we have con-
sulted material housed in several natural
history museums, which include: BMNH—
The Natural History Museum, London (for-
merly British Museum of Natural History);
FMNH—Field Museum of Natural Histo-
ry, Chicago; MNHN—Muséum National
d’ Histoire Naturelle, Paris; and UADBA—
Université d’ Antananarivo, Département de
Biologie Animale.

Five external measurements in millime-
ters were taken from our two specimens be-
fore preparation and included: total length,
head and body length, tail length, hind foot
length (not including claw), and ear length.
Mass was measured with the use of a spring
balance and recorded in grams.

An additional six cranial and two dental
measurements were taken using a digital
calipers accurate to the nearest 0.1 mm.
These measurements, and their definitions,
are: breadth of braincase: the greatest dis-
tance measured across the hamular process-
es of the squamosals to the mastoid bullae;
greatest length of skull: the distance be-
tween the tips of the nasals and the poste-

253

rior most portion of the cranium; interor-
bital breadth: the minimum distance
across the frontal bones between the orbital
fossae; length of mandibular tooth row:
the maximum distance from distal surface
of the third molar to anterior surface of the
first incisor; length of nasal: the maximum
distance from the posterior extension of the
nasals to their anterior tip; length of palate:
the shortest distance between the tip of the
postpalatal spine and anterior surface of the
first upper incisor; length of maxillary
tooth row: the maximum distance from dis-
tal surface of the third molar to anterior sur-
face of the first incisor; and zygomatic
breadth: the maximum span between the
zygomatic processes of the maxillae.

Tooth abbreviations include: I = incisor,
d = deciduous, C = canine, PM = pre-
molar, and M = molar. Upper case tooth
abbreviations with superscript are used for
upper teeth and lower case abbreviations
with subscript for lower teeth. Cranial and
dental nomenclature follows Hershkovitz
(1977) and MacPhee (1987).

After comparison of the two specimens
collected in the Forét des Mikea to all de-
scribed forms of Microgale, these individ-
uals could not be allocated to any known
form and are therefore described as a new
species.

Microgale jenkinsae, new species
Fig. 2, 3, Tables 1, 2

Holotype.—FMNH 176215, sub-adult
male, collected on 18 February 2003 by
Steven M. Goodman and Voahangy Soari-
malala, field number SMG 13489. The
specimen was preserved as a round study
skin, with associated skull and partial post-
cranial skeleton. Tissue samples were pre-
served in EDTA. The skin is in good con-
dition with a small hole in the left thigh.
The skull and partial postcranial skeleton
are intact. Dental age is sub-adult with P
still erupting and matches MacPhee’s
(1987) eruption pattern stage 1. The basis-
penoid-basioccipital sutures are unfused.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Photograph of the holotype of Microgale jenkinsae (FMNH 176215), a sub-adult male collected on
18 February 2003 in the Forét des Mikea, 9.5 km west Ankiloaka, 22°46.7’S, 43°31.4’E. (Photograph taken by
S. M. Goodman.)

VOLUME 117, NUMBER 3

Table 1.—External measurements (in millimeters) and weight (in grams) of Microgale jenkinsae and other species of small Microgale. Measurements presented as

mean + standard deviation (minimum—maximum, n). For samples of two or fewer specimens only the measurements are presented.

Head and body length Tail length Hind foot length Ear length Weight

Total length

Species

62 719 15 18 4.9

143

M. jenkinsae

(Holotype FMNH 176215)

M. jenkinsae

5.3

147

(FMNH 176154)

M. nasoloi

14.0

141

(Holotype FMNH 156187)

M. pusilla

3.5 = 0.40

3.14.2,n =7

11.4 + 0.98

10-13, n =7

11.4 + 0.53

11-12, n

69.9 = 4.18

65-77, n = 7

51.4 = 2.70

127.4 + 6.16
119-136, n = 7

= 7

SID 2 ANY

51-64, n = 11

47-56, n

3.2 + 0.56

2.14.1, n = 11

8.6 = 0.45

9.6 + 0.50

9-10, n = 11

58.4 + 4.39
53-66, n = 11

117.7 + 5.83
110-128, n = 11

M. parvula

WD 2= Bw

12-16, n = 13

8-9,

8.9 = 1.59
6.3-12, n= 11

12.4 + 0.96

41.9 + 5.92

115.7 + 6.68 68.9 = 0.35
63-74, n = 13 35-41, n = 13

107-129, n = 13

M. brevicaudata

11-14, n = 13

9.4 + 2.97
6.8-15, n = 10

16.1 + 0.67

15-17, n = 12

15.9 = 0.90

85.8 + 3.98
80-93, n = 12

159.3 + 6.34 70.8 + 6.71
64-85, n = 10

150-169, n = 11

M. fotsifotsy

15-17, n = 12

255

External measurements are: total length
143 mm, head and body length 62 mm, tail
length 79 mm, hind foot length (without
claw) 15 mm, and ear length 18 mm. The
animal weighed 4.9 gm (Table 1).

Type locality.—Madagascar: Province de
Toliara, Forét des Mikea, 9.5 km west An-
kiloaka, 22°46.7'S, 43°31.4’E, elevation
about 80 m above sea level (Fig. 1). The
site is about 17 km inland from the Moz-
ambique Channel.

Habitat.—The holotype was obtained in
partially disturbed dry transitional decidu-
ous forest growing on red sands. It was cap-
tured in a pitfall trap placed in relatively
dense understory composed primarily of
Xerophyta (Velloziaceae), small bushes,
and succulent Euphorbiaceae.

Diagnosis.—A relatively small member
of the genus Microgale with a head and
body length of 59-62 mm, tail length of
79-81 mm, and greatest skull length of
18.7-18.8 mm. Deciduous PM? is simple,
caniniform, and single-rooted. The color of
the dorsal pelage is mixed agouti and the
venter is gray with white-tipped fur. The
ears are notably long (18 mm) for a shrew
tenrec of this size.

Paratype.—FMNH 176154 (SMG
13492), sub-adult female from the same lo-
cality as the holotype, collected 19 Febru-
ary 2003, and prepared as fluid preserved
specimen with extracted skull. The dental
eruption pattern fits MacPhee’s (1987) stage
1. Tissues saved in EDTA.

Distribution.—Microgale jenkinsae is
known only from the type locality in the
Forét des Mikea, southwestern Madagascar.
Description.—A small species of Micro-
gale having a tail longer than the head and
body (Fig. 2). The dorsal fur is relatively
dense and soft. Pelage from the level of the
ears to the base of the tail (including the
flanks), is a mixture of completely black
and tannish-brown hairs, or those that are
tannish-brown along most of their length
and black-tipped, imparting an agouti ap-
pearance. The agouti pattern runs anteriorly
from the level of the ears to the eyes. An-

256 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Cranial and dental measurements (in millimeters) and weight (in grams) of Microgale jenkinsae
and other species of small Microgale. Measurements presented as mean =~ standard deviation (minimum —
maximum, n). For samples of two or less specimens only the measurements are presented.

Greatest length

Species of skull
M. jenkinsae 18.8
(Holotype FMNH 176215)
M. jenkinsae 18.7
(FMNH 176154)
M. nasoloi DBw)
(Holotype FMNH 156187)
M. pusilla 16.6 = 0.71
15.7-17.5, n = 7
M. parvula 16.5 + 0.47
15.5-17.0, = 12
M. brevicaudata 20.7 + 0.79

19.0—21.9, n = 13

M. fotsifotsy 21.1 + 0.76

20-22, n= 12

terior and lateral to this band, the pelage is
distinctly paler in coloration, with the ma-
jority of hairs being pale tan to silvery-
white. Individual hairs along the dorsum
measure 4—5 mm. Guard hairs are medium
gray in color. The ventral pelage, with the
exception of the portion surrounding the
gular to mental regions, is gray based with
off-white tips. The difference between the
ventral and dorsal color pattern is pro-
nounced, but grade into each other laterally
instead of forming a well-demarcated line.
Upper surfaces of fore feet and hind feet
are covered with short silver-white fur,
which on the hind feet extends slightly be-
yond the claws as ungual tufts. The color
of mystacial and rhinarial vibrissae vary
from either completely beige-white or
black, to black at the base and gradually
becoming beige-white at the tips. Mystacial
vibrissae reaching up to 20 mm and rhinar-
ial vibrissae about 5 mm in length. Pinnae
are notably long (18 mm) for a small Mi-
crogale, dark brown in color, and covered
internally and externally with fine, silvery-
gray fur.

The hind foot is relatively long (14-15
mm) for a small species of Microgale (Ta-
ble 1). The first digit of the hind foot is less
than one-third the length of the second dig-

Zy gomatic Interorbital
breadth minimum
6.9 4.0
6.9 4.0
8.3 >. I
6.1 + 0.24 3.4 + 0.17
5.6-6.3, n = 8 3.1-3.7, n = 9
Spall 22 O23 3.7 = 0:19
4.7-5.4, n = 12 3.3-4.0, n = 12
8.0 = 0.51 4.8 = 0.27
7.2-8.8, n = 13 4.3-5.2,n = 14
8.2 = 0.32 5.0 + 0.18
7.7-8.7, n = 11 4.6-5.2, n = 12

it. The second and third digits are subequal
in length, with the fourth digit slightly lon-
ger. The fifth digit is about two-thirds the
length of the fourth. There are five plantar
tubercles and, based on FMNH 176154
(SMG 13492), these are located at the base
of digit 1 and digit 5, in intermediate po-
sitions between the base of digits 2 and 3
and digits 3 and 4, and notably reduced as
distal hypothenar and proximal thenar pads.

The skin of the tail is dark brown dor-
sally and tannish-brown ventrally, and
forming a relatively well-demarcated line
laterally separating these two surfaces. The
tail is clothed with very fine silvery-white
fur, which becomes slightly denser at the
tip. In FMNH 176154 (SMG 13492) the
last 10 mm of the tail is mottled dark-white.

The skull is relatively short (Table 2),
slightly flattened dorsolaterally, with a con-
stricted interorbital region. The rostrum is
relatively short and tapers anteriorly. The
anterior portion of frontals consist of two
slightly concave plates divided at the mid-
dorsal line and the posterior portion is
slightly-domed. The braincase has a slightly
bulbous parietal and interparietal, a rounded
supraoccipital and occipital, and a weakly
defined occipital crest. Dentally the holo-
type is a sub-adult with the erupting crown

VOLUME 117, NUMBER 3

Table 2.—Extended.

Braincase Length Length Length of Length of
width of nasal of palate maxillary toothrow mandibular toothrow
8.4 8.9 8.1 8.2 8.0
8.2 8.0 Wall 8.2 8.1
9.2 10.3 9.8 10.1 9.7
6.9 = 0.11 7.0 + 0.37 1.3) = O34! 7.4 + 0.40 7.0 + 0.36
6.87.1, n = 7 6.4-7.5, n= 8 6.8-7.8, = 8 6.7-7.7, n=8 O54, 0 = 7/
6.8 = 0.18 Wed) = O22 Wo3) 22 O23 TA + 0.28 7.2 + 0.19
6.5—7.0, n = 12 6.9-7.7, n = 12 6.9-7.6 ,n = 11 6.7-7.6, n= 12 2O= Os S12
8.7 = 0.27 9.3 + 0.62 9.3} 2£ O.S7/ 9.3) 22 0.5 8.8 + 0.35
8.2-9.1, n = 13 8.4-10.5, n = 14 8.2-10.2, n = 14 8.3-10.2, n = 14 8.1-9.4, n= 14
9.3 + 0.26 9.4 + 0.45 OFS 0139) 10.0 + 0.43 9.6 + 0.47
8.9-9.7, n = 12 8.9-10.3, n = 12 9.1-10.2, n= 12 9.3-10.7, n = 12 9.0-10.5, n = 12

of I present and all antemolars are decid-
uous, fitting stage 1 in MacPhee’s (1987)
tooth eruption pattern. The upper toothrows
from dI' to dPM? slightly converge anteri-
orly. The lingual margins of dPM? and M!
to M2 are roughly parallel. Palatal foramina
are present. Pterygoids are relatively short
and broad, and the pterygoid processes
winged-shape and curved mid-ventrally.
The glenoid fossa is shallow and narrowly
curved. The mandibles are slender, the cor-
onoid processes are relatively narrow at
their bases and pointed dorsally, and the an-
gular processes are short and narrow, and
the dorsal surface is not expanded (Fig. 3).

The dentition is not markedly robust
(Fig. 3). There is a gap between the dI' and
dI* and between the dpm, and dpm,. The
first upper incisor (dI') is small, bicuspid
(bidentate), and the distostyle moderately
well developed; the dl? with approximately
the same crown height as dl’, the tricuspid
(tridentate) has the anterior accessory cusp
more developed than distostyle; the dl? is
one-half crown the height of dI*? and reach-
ing just beyond the level of the distostyle
of the dI’, the bicuspid with small distos-
tyle; the dC! robust with crown height
reaching that of the dI’, with small acces-
sory anterior cusp and pronounced distos-

tyle; the dP? small, equal in crown height
to distostyle of the dC'; the dP® is large,
slightly greater in crown height than the
dC', lingual ledge with well-developed pro-
tocone, and the parastyle, mesiostyle, an-
terior ectostyle, and distostyle present; the
dP* is large, longer in crown length than M!
to M?, with elongated paracone, the lingual
ledge with a protocone more developed
than M' to M?, anterior ectostyle approxi-
mately same length as paracone, and the
parastyle, mesiostyle, and distostyle pre-
sent; the M! and M? large, parastyle, me-
siostyle, anterior ectostyle, and distostyle
present, and centro-buccal cleft slightly
more prominent in M? than M!; the M? is
reduced in size and compressed anteriopos-
teriorly. The first lower incisor (di,) is large,
slightly shorter in crown length to the di,,
the posterior accessory cusp well-devel-
oped; the di, is large, the posterior acces-
sory cuspid well-developed; the di, is small,
about one-half the crown length of lower
(deciduous) canine; the dc is large, poste-
rior accessory cuspid present, no anterior
accessory cuspid; the dpm, is small, slightly
shorter that posterior accessory cuspid of
lower canine, poorly developed anterior ac-
cessory cuspid and posterior accessory cus-
pid, and single-rooted; the dpm, is moder-

258 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Views of the cranium and mandible of the holotype of Microgale jenkinsae (FMNH 176215): upper
left, dorsal view; upper right, ventral view; lower center lateral view of cranium and mandible. (Photograph
taken by J. Weinstein, image number Z94379_05d.)

VOLUME 117, NUMBER 3

ate in size, slightly longer in crown height
than the pm,, moderately developed ante-
rior accessory cuspid and posterior acces-
sory cuspid; the dpm, is large, equal in
crown height (formed by prominent proto-
conid) to m,, the anterior accessory cuspid
and posterior accessory cuspid present; the
m,and m, are large, the m, slightly subequal
in crown height to the m,, both with well-
developed protoconid, anterior accessory
cuspid, and posterior accessory cuspid, and
slightly elongated anterobuccal cingulum;
the m, is large and equal to m, in crown
height, and with a well-developed proto-
conid, anterior accessory cuspid, and prom-
inent posterior accessory cuspid, and slight-
ly elongated anterobuccal cingulum. Given
that the individuals of M. jenkinsae are
stage 1 sub-adults, no information can be
provided on the adult dentition or antemolar
replacement pattern of this species.

Comparisons.—The fact that our two
specimens of Microgale jenkinsae are stage
1 sub-adults complicates comparisons to a
certain degree. However, sub-adult mem-
bers of this genus, at this stage of dental
eruption, exhibit the pelage coloration of
adults and, in general are similar to adults
in external measurements (MacPhee 1987;
Jenkins et al. 1996, 1997).

M. jenkinsae is readily distinguished ex-
ternally from other relatively small mem-
bers of this genus by pelage coloration and
measurements. The contrasting agouti dor-
sum and grizzled-gray venter is unique
among small shrew-tenrecs. The pelage
pigmentation in M. nasoloi is a relatively
uniform gray; M. fotsifotsy has less gray in
the dorsum than M. jenkinsae and is notably
darker; M. parvula has a dark brown dor-
sum and dark grayish-brown ventrum; M.
brevicaudata is medium-brown dorsally
and dull grayish-brown ventrally; and in M.
longicaudata and M. pusilla the dorsum is
a mixed light brown and medium brown
and ventrum gray broadly edged with dark
tan-brown. Further, there is no overlap in
tail measurements between M. jenkinsae
and any of these taxa, with the exception of

259

M. fotsifotsy, but M. jenkinsae can be dif-
ferentiated from it based on pelage charac-
teristics, a non-white-tipped tail, upper sur-
faces of the feet clothed with short silver-
white fur, and several external measure-
ments (Table 1).

The presence of a single-rooted second
lower premolar separates M. jenkinsae from
all other named small members of the genus
Microgale, with the exception of M. pusilla.
In the latter species the root form is iden-
tical in individuals with deciduous and per-
manent antemolar dentitions. M. pusilla is
notably smaller than M. jenkinsae in all ex-
ternal, cranial, and dental measurements
(Tables 1 and 2), and is separable based on
pelage characters.

A generic revision of all members of Mi-
crogale is currently in preparation and in-
cludes molecular characters (Olson and
Goodman, in prep.). The results of this
study will be presented elsewhere and will
address aspects of the phylogenetic position
and sister-taxa relationships of M. jenkin-
sae.

Etymology.—This new species of Micro-
gale is named after Paulina D. Jenkins of
The Natural History Museum, London, for
her important contributions to Tenrecidae
systematics.

Discussion

Ecology.—Microgale jenkinsae is cur-
rently known only from the Forét des Mi-
kea, between Morombe and Manombo (Fig.
1), in the southwestern portion of Mada-
gascar, a zone of transitional dry deciduous
and spiny bush habitat (Seddon et al. 2000;
Goodman and Soarimalala in press). This
region receives, on average, about 400—500
mm of rainfall per year (Chaperon et al.
1993), with probably more rainfall in the
inland higher ground than along the coastal
plain. Differences in forest types within the
Forét des Mikea tend to follow this pattern,
with more deciduous forest on the slightly
higher ground away from the coast and
spiny bush along the coastal plain.

260

The climax vegetation of the dry decid-
uous forest has been characterized as being
dominated by the genera Dalbergia, Com-
miphora, and Hildegardia (Humbert 1965).
The formation has a canopy 10 to 15 m
high, sometimes reaching 20 m, with a open
medium stratum and diffuse undergrowth.
All trees and most of the shrubs shed their
leaves in the dry season.

The vertebrate communities inhabiting
the Forét des Mikea are similar to other arid
portions of the island, although there are
apparently several strict and regional en-
demic vertebrates (e.g., the reptiles Furcifer
belalandaensis and Paroedura vahiny, the
birds Uratelornis chimaera and Monias
benschi). The known small mammal com-
munity consists of six Lipotyphla (Tenrec
ecaudatus, Setifer setosus, Echinops telfai-
ri, Geogale aurita, Microgale jenkinsae,
and Suncus madagascariensis), one intro-
duced murine rodent (Rattus rattus), and
two endemic Nesomyinae (Macrotarsomys
sp. and Eliurus myoxinus) (Carleton and
Schmidt 1990; Soarimalala and Goodman
2004; Goodman and Soarimalala in press).

Trapping.—During the survey of the
Forét des Mikea, six sites were visited and
systematically trapped using standard live
and pit-fall traps (for more details on tech-
niques see Goodman & Carleton 1996;
Goodman et al. 1996). Pit-fall devices have
been particularly useful for sampling Li-
potyphla difficult to trap by other methods.
For example, the only known specimens of
another western Microgale, M. nasoloi,
were captured with this technique (Jenkins
and Goodman 1999). Three pit-fall lines,
each composed of 11 12-liter buckets
placed 10 m apart, were installed at each of
the six survey sites. There was considerable
variation between sites in trap success and
species diversity. At site 1, 20 individuals
(Tenrec, Setifer, Echinops, M. jenkinsae,
and Geogale) were caught in 198 bucket
nights between 14 and 19 February 2003.
At site 2, 10 individuals (Echinops and
Geogale) were obtained in 198 bucket
nights between 21 and 27 February 2003.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

At site 3, 5 individuals (Tenrec, Echinops,
Rattus rattus, and Geogale) were taken in
132 bucket nights between 2 and 5 March
2003. At site 4, only 4 individuals (Tenrec,
Geogale, and Suncus madagascariensis)
were trapped in 165 bucket nights between
8 and 12 March 2003. At site 5, nothing
was captured in 198 bucket nights between
14 and 19 March 2003. At site 6, 7 indi-
viduals (Geogale) were captured in 132
bucket nights between 22 and 25 March
2003. This level of faunal heterogeneity (as
reflected by trapping-success) may be relat-
ed to microhabitat differences between the
sites, but further research is needed to test
this hypothesis.

Natural history.—Little definitive infor-
mation can be gleaned on the natural his-
tory of Microgale jenkinsae on the basis of
two specimens, and the following extrapo-
lations are tentative. Using foot structure
and the context in which this species was
trapped, it is terrestrial. Both specimens
were taken in a portion of the Forét des
Mikea dominated by more dry deciduous
forest than by spiny bush habitat. Several
of the other sites inventoried during the
Forét des Mikea survey tended to be dom-
inated by spiny bush habitat. The two M.
Jenkinsae were obtained in different pitfall
lines installed in portions of the forest hav-
ing a relatively dense understory—one line
was dominated by Gramineae associated
with the regeneration of an old forest ex-
ploitation track and the other line in a mi-
crohabitat with a dense growth of Xerophy-
ta (Velloziaceae) mixed with other low-
growing plants and some succulent Eu-
phorbiaceae.

An analysis of soil samples taken at each
of the 18 pitfall lines installed during the
Forét des Mikea survey indicate that at site
1, where the two specimens of M. jenkinsae
were collected, the average percent carbon
in the soil was higher (1.9%, range 0.98—
2.3%) than four of the other five sites (all
less than 0.9% carbon). The outlier is site
2 which had a slightly higher soil carbon
content (2.4%, range 0.08—4.5%) than site

VOLUME 117, NUMBER 3

1. Given that shrew tenrecs are believed to
be primarily insectivorous, one may expect
that their distribution would be correlated
with soils relatively rich in organic materi-
al, which in turn would support a higher
density and diversity of invertebrates.

Both specimens are dentally sub-adults,
and, thus, it is not unexpected that they did
not show any signs of reproductive activity.
However, shrew tenrecs with deciduous an-
temolar dentitions can be reproductive (e.g.,
Jenkins et al. 1996). Our survey was con-
ducted during the rainy season, a period of
the year that normally coincides with breed-
ing activity in small mammals in this area
of the island (Ganzhorn et al. 1996; Ran-
drianjafy 2003).

Paleoecological implications.—Remains
of Microgale pusilla have been reported in
disintegrated owl pellets of unknown age,
but by extrapolation almost certainly Hol-
ocene, from the sites of Lelia and Anjo-
himpaty in southwestern Madagascar
(MacPhee 1986, 1987), a zone of xero-
phytic spiny bush habitat on an exposed
limestone substrate. M. pusilla is consid-
ered to be an inhabitant of the more mesic
portions of the island, including the eastern
humid forest and central highlands. More
recently bone remains of M. pusilla have
been identified in owl pellets collected in
the capital city of Antananarivo, at least 80
km from the nearest intact natural forest
block, and it is assumed that this species
may live in surrounding marshlands and
rice fields (Goodman et al. 1997a). At sev-
eral sites in the central highlands it has been
captured in marshlands within close vicinity
to natural forest (Goodman et al. 2000a,
Soarimalala et al. 2001). Thus, its occur-
rence in Owl pellets in southwestern Mad-
agascar can be interpreted in at least two
ways: this species is a generalist and is able
to live in a variety of ecological conditions
from humid forests to marshlands to xero-
phytic bush—however, on the basis of re-
cent inventories of the drier portions of the
island there is no evidence of its occurrence
in this latter habitat—or the undated owl

261

pellets collected in the southwest are from
a past geological period when this region of
Madagascar was distinctly more mesic.
The specimens of M. pusilla described by
MacPhee (1986, 1987) from Lelia and An-
johimpaty were deposited in the Service de
Paléontologie collection at the Université
d’Antananarivo. A detailed search of that
collection, however, did not uncover these
specimens. Nevertheless, a comparison of
our material of M. jenkinsae to the illustra-
tions and description of these specimens
(MacPhee 1986) indicates considerable
similarity in size, morphology, and dental
structure. Most important in this regard is
that dpm, (in M. jenkinsae and M. pusilla)
and pm, (in M. pusilla) are simple in cor-
onal structure and single-rooted, characters
used to separate M. pusilla from all other
small members of this genus before our rec-
ognition of M. jenkinsae. Further, on the ba-
sis of a scale provided with the line drawing
of the Anjohimpaty mandible (MacPhee
1986, Fig. 5), the approximate lower tooth-
row length is 18.2 mm, which is within the
range of M. jenkinsae, but notably larger
than M. pusilla (Table 2). We strongly sus-
pect that these specimens, reported as M.
pusilla, may be referable to M. jenkinsae.
Recent biological surveys of the Parc
National de Tsimanampetsotsa (Fig. 1), for-
merly under the statute of a Réserve Na-
turelle Intégrale, did not find any species of
Microgale living in this protected area
(Goodman et al. 2002), which is relatively
close to Lelia and Anjohimpaty. Our small
mammal surveys in the Forét des Mikea at
six different sites, with a minimum of 132
pit-fall nights per site, yielded a total of
1023 pit-fall nights, yet only two individ-
uals of M. jenkinsae were captured, both at
the first site. This would indicate that this
species is either rare or difficult to capture
and presumably occupies specific micro-
habitats. The important point here is that
before significant paleoenvironmental infer-
ences can be made associated with the pres-
ence of certain taxa known only as subfos-
sils, it is critical that detailed biological in-

262

ventories be conducted in the general re-
gion of the paleontological site to
thoroughly document the extant fauna.

Conservation.—Historically, most field
efforts associated with the exploration and
documentation of Madagascar’s unique fau-
na have been in the humid forests on the
eastern, central, and northern portions of
the island. Further, there is a preponderance
of reserves and parks protecting this biome
as compared to the drier western and south-
ern regions of the island (ANGAP 2001).
On the basis of several recent biogeograph-
ic analyses of small mammals and birds,
species turnover along the nearly 1200 km
long eastern humid forests of the island is
relatively low (Goodman et al. 1997b,
2000b). A number of endemic species in
this biome have broad distributions, many
extending the complete length of this hab-
itat. More recent biological surveys of
Madagascar’s western deciduous forests
and southern spiny bush lands have re-
vealed a previously unrecognized biota, in-
cluding numerous terrestrial vertebrates.
The growing realization is that levels of
plant and animal species turnover along a
latitudinal transect of western Madagascar
is notably higher than the east, and this is
probably related to a greater geological
complexity and associated botanical com-
munities in the west (Du Puy & Moat
2003).

The recent surveys of the Forét des Mi-
kea, which has no official protection, and
forested regions to the north and south of
this zone, are a case in point. Two unde-
scribed species of mammals (Microgale
jJenkinsae and Macrotarsomys nov. sp.)
have been discovered in the Forét des Mi-
kea that are unknown from any other region
in the west. Further to the north, in the vi-
cinity of the Bemaraha Plateau, there are at
least two species of rodents that appear to
be endemic to the region (Carleton et al.
2001). The recent discovery and description
of Microgale nasoloi from unique forest
formations in southwestern central Mada-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

gascar seems to indicate another regional
endemic with a very limited distribution.

The drier western and southern forests of
Madagascar have been subjected to consid-
erable anthropogenic degradation, perhaps
greater than in the humid east (see Smith
1997, Dufils 2003). In areas such as the
Forét des Mikea, which was estimated in
1999 to contain forest cover in excess of
3700 km?, habitat loss rates have increased
over the past few decades associated with
pressures in the form of selective logging,
cattle pasture, hunting, and clearing for ag-
ricultural crops (Seddon et al. 2000). Given
the levels of habitat heterogeneity and mi-
croendemism in the west, action needs to
be taken to protect the remaining large
blocks of natural habitat in this region. On
the basis of recent exploration of the Forét
des Mikea this area should be given priority
amongst the zones in need of rapid protec-
tion.

Acknowledgments

We are grateful to the Direction des Eaux
et Foréts for issuing permits to conduct fau-
nal surveys in the Forét des Mikea. For ac-
cess to specimens in their care we are in-
debted to Géraldine Veron, Muséum Na-
tional d’Histoire Naturelle, Paris; Paulina
D. Jenkins, The Natural History Museum,
London; and Prof. Daniel Rakotondravony,
Université d’ Antananarivo, Département de
Biologie Animale, Antananarivo. This field
project was supported by WWF-Madagas-
car, Fonds Frangaise pour |’ Environnement
Mondiale (l’Agence Frangaise de Dével-
oppement), and the Volkswagen Founda-
tion. We are grateful to Link Olson for
comments on an earlier draft of this paper
and his important aid in numerous ways.
Two anonymous reviewers also helped to
improve this paper.

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VOLUME 117, NUMBER 3

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Appendix |

List of specimens of Microgale spp. examined during
the course of this study.

Microgale parvula.—Province d’ Antsiranana, Parc
National de Marojejy [formerly Réserve Naturelle In-
tégrale de Marojejy], along tributary of Manantenina
River, 8 km NW Manantenina, 14°26.2'S, 49°46.5’E,
450 m (FMNH 159681); Parc National de Marojejy
[formerly Réserve Naturelle Intégrale de Marojejy],
along tributary of Manantenina River, 10 km NW
Manantenina, 14°26.0'S, 49°45.7’E, 775 m (FMNH
159682, 159683, 159684); Parc National de Marojejy
[formerly Réserve Naturelle Intégrale de Marojejy], 11
km NW Manantenina, Antranohofa, 14°26.2’S,
49°44.5'E, 1225 m (FMNH 159685); Pare National de
Marojejy [formerly Réserve Naturelle Intégrale de Ma-
rojejy],10.5 km NW Manantenina, along tributary at
head of Andranomifototra River, 14°26.4’S, 49°44.5’E,
1625 m (FMNH 159686). Province de Fianarantsoa,
approx. 45 km S. Ambalavao, east bank Iantara River,
along Ambalamanenjana-Ambatoboay trail, edge of
Pare National d’Andringitra [formerly Réserve Natu-
relle Intégrale], 22°13'20"S, 47°01'29"E, 720 m
(FMNH 151621); Pare National d’Andringitra [for-
merly Réserve Naturelle Intégrale d’Andringitra],
approx. 43 km S. Ambalavao, junction of Sahanivo-
raky and Sahavatoy Rivers, 22°13’40"S, 47°00'13”E,
810 m (FMNH 151622); Pare National d’Andringitra
[formerly Réserve Naturelle Intégrale d’ Andringitra],
approx. 38 km S. Ambalavao, on ridge east of Volot-
sangana River, 22°11'39”"S, 46°58'16"E, 1625 m
(FMNH 151623, 151723, 151794, 151801).

Microgale nasoloi.—Province de Toliara, Forét de

Vohibasia, 59 km northeast Sakaraha, 780 m,

VOLUME 117, NUMBER 3

22°27.5'S, 44°50.5'E (FMNH 156187); Forét
d’ Analavelona, Antanimena, 12 .5 km NW Andranoh-
eza, 22°40.7'S, 44°11.5'E, 1050 m (FMNH 161576).
Microgale fotsifotsy.—Province de Fianarantsoa,
Pare National d’Andringitra, 8.5 km SE Antanifotsy,
Campement Andohan’ Ambola, PP ANOB2TBaS:
46°56.758'E, 1960 m (FMNH 165694, 165778,
165779) ; 2 km W. Andrambovato, along Tatamaly
River, 21°30.7'S, 47°24.6'E, 1075 m (FMNH 170749);
Forét de Vinantelo, at foot of Mt. Ambodivohitra, 15.5
km SE Vohitrafeno, 21°46.6'S, 47°20.8’E, 1100 m
(FMNH 170750); approx. 40 km S. Ambalavao, along
Volotsangana River, 22°13’22"S, 46°58'18”E, 1210 m
(FMNH 151646, 151647); Province de Mahajanga,
western side of Anjanaharibe-Sud, 13.5 km SW Befin-
gotra, 14°47.0'S, 49°26.5’E, 1200 m (FMNH 167428).
Province de Toliara, Pare National d’ Andohahela [for-
merly Réserve Naturelle Intégrale d’ Andohahela], par-

265

cel I, 8 km NW Eminiminy, 24°37.55'S, 46°45.92’E,
440 m (FMNH 156569); Parc National d’ Andohahela
[formerly Réserve Naturelle Intégrale d’ Andohahela],
parcel I, 13.5 km NW Eminiminy, 24°35.04’S,
46°44.08'E, 1200 m (FMNH 156424); Pare National
d’Andohahela [formerly Réserve Naturelle Intégrale
d’Andohahela], parcel I, 15.0 km NW Eminiminy,
24°35.15’S, 46°43.85’E, 1500 m (FMNH 156570).

Microgale pusilla.—Province d’ Antananarivo, 13
km NE Antananarivo, in Tyto alba pellets (FMNH
151606, 151607); 10 km SE Tsinjoarivo, Forét de Ma-
hatsinjo, Andasivodihazo, 19°40.7’S, 47°46.2’E, 1550
m (FMNH 166123, 166124, 166125) ; Réserve Spé-
ciale d’Ambohitantely, 24 km NE Ankazobe,
18°10.1'S, 47°16.6’E, 1450 m (FMNH 165489). Prov-
ince de Fianarantsoa, Manambolo Forest, Ambavafa-
tra, along Andohabatotany River, 17.5 km SE Sendri-
soa, 22°8'58"S, 47°1'25”, 1300 m (FMNH 167612,
167619, 167621).

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(3):266—270. 2004.

Designation of the type species of Musaraneus Pomel, 1848
(Mammalia: Soricomorpha: Soricidae)

Neal Woodman

USGS Patuxent Wildlife Research Center, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20013

Abstract.—The genus name Musaraneus often is attributed to Brisson
(1762), however, most of Brisson’s names are unavailable. Pomel (1848) sub-
sequently made the name Musaraneus available, but did not designate a type
species. The 18 species that Pomel listed under Musaraneus currently are dis-
tributed among five modern genera, two of which (Cryptotis Pomel, 1848 and
Diplomesodon Brandt, 1852) are predated by Musaraneus. Because Cryptotis
and Diplomesodon potentially could be considered junior synonyms of Musar-
aneus, I propose Sorex leucodon Hermann, 1780 (= Crocidura leucodon) as
the type species for Musaraneus, thereby establishing Musaraneus as a junior

synonym of Crocidura Wagler, 1832.

The generic name Musaraneus Pomel,
1848 derives from mus araneus (“‘spider
mouse’’), one of the terms commonly used
alongside sorex and mus caecus by classical
Latin writers (e.g., Plinius n.d.; Columella
n.d.; Serenus n.d.) to refer to small mam-
mals now generally interpreted as shrews
(family Soricidae). The classical name mus
araneus has a long history of use in early
zoological literature. It was adopted and
used widely by Renaissance natural histo-
rians and made the transition from a Latin
common name to being incorporated into
more formal taxonomies. The vernacular
mus araneus generally was applied to the
small mammal called locally by a variety
of names that included ‘“‘muzeraigne,”’
“spitzmus, shrew, erd shrew,’ or
“shrew-mouse’”’ (Gesner 1551, 1560, 1602:
Marggraf 1648; Jonston 1657; Topsell
1658; Ray 1693). A number of early tax-
onomists attempted to establish the name as
Mus Araneus or Musaraneus within heir-
archical classifications (Charleton 1668;
Klein 1751; Brisson 1756, 1762). It is of
interest that Gesner (1551, 1560, 1602) and
subsequent writers (e.g., Topsell 1658,
Charleton 1668) interpreted sorex as dis-

99 ce 29 ce

tinct from mus araneus, in some cases as a
broader category that might include mus ar-
aneus (e.g., Klein 1751), or as a separate
set of animals, typified by mus avellana-
rum, the “haselmus” or “‘hasel-mouse”
(Gesner 1560, Topsell 1658), or by the
“rat”? (Charleton 1668). Gesner’s (1551,
1560, 1602) print of mus araneus is an il-
lustration of a soricid (Fig. 1A), possibly a
white-toothed shrew of the genus Croci-
dura, whereas his picture of a sorex is iden-
tifiable as a garden dormouse (Eliomys
quercinus—Fig. 1B). His illustrations were
copied and republished by subsequent writ-
ers (e.g., Topsell 1658) and likely influ-
enced later interpretations of the names. In
contrast, Linné (1746, 1748, 1758) explic-
itly and consistently applied Sorex to those
mammals that previous authors had called
mus araneus or Musaraneus, and Sorex
Linné, 1758 is the name that survived in
the taxonomic literature. Musaraneus con-
tinues to be reflected in modern words for
shrew in a number of romance languages,
e.g., musarana (Spanish), musaraigne
(French), musaranho (Portuguese), musar-
agno (Italian). It also survives, in part, in

VOLUME 117, NUMBER 3

Fig. 1. Gesner’s (1602) illustrations of (A) mus araneus and (B) mus avellanarum from Historiae Animalium.
Photographs courtesy of the Smithsonian Institution Libraries, Joseph KR Cullman 3" Library of Natural History,

Washington D.C. Reproduced with permission.

the scientific name for the European com-
mon shrew, Sorex araneus Linné, 1758.

As a genus-level name, Musaraneus is
often attributed to Brisson (1762; see Pomel
1848, Sherborn 1902, Palmer 1904, Mc-
Kenna and Bell 1997, Kretzoi and Kretzoi
2000). Because Brisson (1762) did not con-
sistently apply binomial nomenclature in
his work, however, most of his names are
unavailable in accordance with Article 11.4
of the International Code of Zoological No-
menclature (ICZN 1999; but see Hopwood
1947; ICZN 1998). In a subsequent classi-
fication of insectivores, Pomel (1848) re-
described Brisson’s Musaraneus as one of
four genera (with Talposorex Pomel, Sorex
Linneus, and Galemys Pomel) within the
tribe Soriciens in his family Spalacogalae.
Pomel (1848) made the name Musaraneus
available, and therefore, he is the author of
this name, as noted by Sherborn (1928).
Hopwood (1947) recorded a number of oth-
er generic names used by Brisson (1762)
that similarly were made available by later
authors (see also ICZN 1998).

Pomel’s (1848) Musaraneus comprised

18 species distributed among three “‘sec-
tions”’ (subgenera): Cryptotis, a new taxon
with a single North American species; Myo-
sorex Gray, 1838, an existing taxon com-
prising three African species; and Croci-
dura Wagler, 1832, an existing taxon con-
taining 14 Old World species. Based on the
list of included species, the genus Musar-
aneus included representatives from five
modern genera. In addition to those genera
representing Pomel’s (1848) three sections
(Cryptotis, Myosorex, Crocidura), one spe-
cies (Musaraneus puchellus) represents Di-
plomesodon Brandt, 1852, and three others
(M. crassicaudatus, M. vulgaris, M. Bach-
mani) represent Sorex Linné, 1758.

Pomel (1848) was uneven in designating
species that he considered to be typical of
the genera he described. In his classification
of insectivores, Pomel noted a “typical spe-
cies” for his newly-described Talposorex,
but not for his names Galemys or Musara-
neus. For these latter genera, he provided
lists of species divided among several sec-
tions (subgenera). Pomel (1848) wrote the
latter name as, ““Genre Musaraneus Briss,

268

Pom.,” clearly indicating the influence of
Brisson (1762). It is curious, however, that
his version of Musaraneus did not explic-
itly include any of the three “‘species”’ (Mu-
saraneus, Musaraneus aquaticus, M. bras-
iliensis) that Brisson (1762) had allocated
to his genus Musaraneus. Pomel (1848)
may have considered Brisson’s (1762) uni-
nomial “‘species’” Musaraneus to be repre-
sented by what Pomel called Musaraneus
(Crocidura) vulgaris, which he equated
with araneus of authors.

The type species of Musaraneus Pomel,
1848 is important to modern taxonomists
because Musaraneus Pomel, 1848 predates
Cryptotis Pomel, 1848 and Diplomesodon
Brandt, 1852, and Musaraneus could be in-
terpreted as a senior synonym of one of
these genera. Kretzoi and Kretzoi (2000:
241) indicated that the type species for Mu-
saraneus Pomel is ““Crocidura (M.) priscus
Pomel”’ (sic). There are a number of im-
portant and confusing errors in their ac-
count for this name, however. Both the
original description of the genus and the
designation of Crocidura priscus as the
type species are credited by them to “Po-
mel 1853,” which is referenced as “Arch.
Sci. Phys. Nat., Bibl. Univ. Genéve, 9:
249,’ but this reference is a conflation of
several different publications. Pomel
(1853a, 1853b) are parts of his “Catalogue
des Vertébrés Fossiles,” which was pub-
lished in at least three sections in Annales
Scientifiques, Littéraires et Industrielles de
L’Auvergne: the first in October and No-
vember of 1852 (Pomel 1852), the second
in March and April of 1853 (Pomel 1853a),
and the last in May and June of 1853 (Po-
mel 1853b). Insectivores, including the de-
scription of the fossil species Musaraneus
(Crocidura) priscus, appear in the first part
of this work, and the correct citation for that
name is Pomel (1852:351). Pomel’s original
description of Musaraneus is in an earlier
publication (Pomel 1848) that was pub-
lished in Volume 9 of Archives des Sciences
Physiques et Naturelles, Genéve. In order
for a species to be designated a type species

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

by a subsequent author, it must have been
included in the genus by the original author
(ICZN 1999: art. 69.2). However, Pomel
(1848) did not include the name priscus
among the species he listed under Musar-
aneus, as he had not yet described the spe-
cies. Therefore, the selection by Kretzoi and
Kretzoi (2000) is invalid.

Pomel’s (1848:249) original description
of Musaraneus reads:

Trois intermédiaires en haut, deux en bas, estomac
oblong avec poche bien marquée sous le boyau py-
lorique.

My translation of this description is:

Three upper intermediary teeth, two lowers, stom-
ach oblong with well-marked pouch below the py-
loric constriction.

I interpret Pomel’s upper ““intermédiaires”
to represent the simplified upper dentition
between the large, hooked first incisor and
the roughly molariform fourth premolar
(P4) that commonly are referred to as “‘un-
icuspids”’ (Choate 1970). The lower “inter-
médiaires”’ are the teeth designated the low-
er unicuspid and lower fourth premolar
(p4). Among the five modern genera Pomel
(1848) included within Musaraneus, all but
Myosorex have a single lower unicuspid
and p4; Myosorex typically has two lower
unicuspids in addition to p4. In the upper
dentition, Sorex has five unicuspids, Cryp-
totis and Myosorex each have four, and Di-
plomesodon has two. Only Crocidura has
three upper unicuspids in accord with Po-
mel’s (1848) description. Although it is not
required that the type species match the
original description for a genus, it is highly
desirable.

Among the modern species of Crocidura
that Pomel (1848) included in Musaraneus,
the majority are African, one is from Japan,
and one, Crocidura leucodon, is widespread
in continental Europe, including France,
where Pomel lived. Among the recommen-
dations for selecting a type species for sub-
sequent designation are that the species be
common and that it be well known to the

VOLUME 117, NUMBER 3

original author (ICZN 1999: Recommen-
dations 69.A1, 69.A7). Therefore, I select
Sorex leucodon Hermann, 1780, as used in
the name combination Musaraneus (Croci-
dura) leucodon by Pomel (= Crocidura
leucodon), as the type species of Musara-
neus Pomel, 1848. By designating this tax-
on as the type species, Musaraneus Pomel,
1848 becomes a junior synonym of Croci-
dura Wagler, 1832, thereby stabilizing the
generic names Cryptotis and Diplomesodon
in accordance with their long-established
usage.

Acknowledgments

My thanks to Alfred L. Gardner for orig-
inally pointing out the potential problems
engendered in the choice of a type species
for Musaraneus. The Department of Special
Collections, University of Kansas Libraries;
Dale Miller, Leslie Overstreet and Daria A.
Wingreen in the Joseph EF Cullman 3” Li-
brary of Natural History, National Museum
of Natural History; and Kirsten van der
Veen in the Dibner Library of the History
of Science and Technology, National Mu-
seum of American History graciously pro-
vided access to, and/or photocopies of, im-
portant pre-Linnean manuscripts. Sandy
Feinstein, Alfred L. Gardner, Robert M.
Timm, and an anonymous reviewer provid-
ed valuable comments on previous versions
of my manuscript.

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
117(3):271-302. 2004.

The mammals of Palawan Island, Philippines

Jacob A. Esselstyn, Peter Widmann, and Lawrence R. Heaney

(JAE) Palawan Council for Sustainable Development, P.O. Box 45, Puerto Princesa City,
Palawan, Philippines (present address: Natural History Museum, 1345 Jayhawk Blvd.,
Lawrence, KS 66045, U.S.A.)

(PW) Katala Foundation, PRO. Box 390, Puerto Princesa City, Palawan, Philippines;

(LRH) Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago, IL 60605 U.S.A.

Abstract.—The mammal fauna of Palawan Island, Philippines is here doc-
umented to include 58 native species plus four non-native species, with native
species in the families Soricidae (2 species), Tupaiidae (1), Pteropodidae (6),
Emballonuridae (2), Megadermatidae (1), Rhinolophidae (8), Vespertilionidae
(15), Molossidae (2), Cercopithecidae (1), Manidae (1), Sciuridae (4), Muridae
(6), Hystricidae (1), Felidae (1), Mustelidae (2), Herpestidae (1), Viverridae
(3), and Suidae (1). Eight of these species, all microchiropteran bats, are here
reported from Palawan Island for the first time (Rhinolophus arcuatus, R. ma-
crotis, Miniopterus australis, M. schreibersi, and M. tristis), and three (Rhin-
olophus cf. borneensis, R. creaghi, and Murina cf. tubinaris) are also the first
reports from the Philippine Islands. One species previously reported from Pa-
lawan (Hipposideros bicolor) is removed from the list of species based on re-
identificaiton as H. ater, and one subspecies (Rhinolophus anderseni aequalis
Allen 1922) is placed as a junior synonym of R. acuminatus. Thirteen species
(22% of the total, and 54% of the 24 native non-flying species) are endemic
to the Palawan faunal region; 12 of these are non-flying species most closely
related to species on the Sunda Shelf of Southeast Asia, and only one, the only
bat among them (Acerodon leucotis), is most closely related to a species en-
demic to the oceanic portion of the Philippines. Of the 28 insectivorous bats,
18 species are somewhat to highly widespread in Indo-Australia, 2 are shared
only with the Sunda Shelf and Indochina, 1 with the Sunda Shelf alone, 3
occur on the Sunda Shelf and the oceanic Philippines, 1 occurs in Palawan,
Sulawesi, and the oceanic Philippines, 2 occur only on Palawan and in the
oceanic Philippines, and | occurs on Borneo, Sulawesi, and throughout the
Philippines. Though the insectivorous bats tend to be widely distributed, these
data, particularly the distributions of the non-volant species, strongly reinforce
the perception of Palawan Island (and associated smaller islands) as a biogeo-
graphic unit of the Sunda Shelf, with only limited similarity to other portions
of the Philippine Islands.

The Philippine archipelago is remarkable
for the large number of indigenous land
mammal species (ca. 175), and especially
for the number of endemic species (ca.
112). Given its relatively small land area,
the Philippines has perhaps the greatest
concentration of endemic mammals in the

world (Heaney et al. 1998, Heaney & Re-
galado 1998, Mittermeier et al. 1997).
These species, especially the endemics, are
not distributed homogeneously over the
country; rather, there is a large number of
discrete biogeographic units, and these cor-
respond to the limits of the islands that ex-

272

isted during periods of low sea level during
the late Pleistocene (Heaney 1986, 1991a,
1991b, 2000). With a single exception, cur-
rent geological evidence indicates that none
of these ““Pleistocene islands” has had dry-
land connections to the Asian mainland or
to other areas. Rather, each arose as a de
novo oceanic island, some from a combi-
nation of oceanic crust and volcanic mate-
rials, and some as uplifted areas of conti-
nental rock that had been submerged for
long periods, and all of these have remained
isolated by sea channels (Hall 1998, 2002;
Heaney 1985, 1986, 1991a). The sole ex-
ception is the Palawan faunal region, which
generally has been considered to be a por-
tion of the Sunda Shelf, both geologically
and biogeographically, with many species
shared with Borneo (Dickerson 1928, Ev-
erett 1889, Heaney 1986). Although Pala-
wan was initially also a de novo oceanic
island, its biogeographic affinity to the Sun-
da Shelf has been thought to be due to the
presence of a shallow shelf between Borneo
and Palawan with an intervening depth of
ca. 145 m (Heaney 1986, 1991a). Previous-
ly, evidence indicated that sea levels
dropped to about 165 m below present lev-
els during the penultimate glacial episode
(Gascoyne et al. 1979), which would have
resulted in a dry-land connection of Pala-
wan to Borneo at about 165,000 BP (Hea-
ney 1985, 1986, 1991a). However, recent
evidence suggests that sea level dropped
only to about 135 m (Rohling et al. 1998)
or perhaps as little as 115 m below present
levels (Siddall et al. 2003, Voris 2000) dur-
ing that glacial episode, leaving open the
question of when Palawan was connected
to Borneo, or if the gap simply became very
narrow.

The mammals of Palawan Island, the
largest part of the faunal region at 11,785
km/?, and the associated smaller islands have
been documented over the course of more
than a century (Allen 1910, Allen 1922, Ev-
erett 1889, Heaney et al. 1998, Hoogstraal
1951, Kuntz 1969, Reis & Garong 2001,
Sanborn 1952, Taylor 1934, Timm & Bir-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ney 1980), but the fauna is still poorly
known in many respects. Little information
has been available for most species on ecol-
ogy and distribution, including habitat re-
quirements, and only a few studies have
considered phylogenetic relationships (e.g.,
shrews: Heaney & Ruedi 1994; bats: Mus-
ser et al. 1982; squirrels: Heaney 1979; mu-
rid rodents: Musser 1979, Musser & New-
comb 1983, Musser & Heaney 1992; pan-
golin: Feiler 1998; leopard cats: Groves
1997). In particular, microchiropteran bats
have been only superficially documented.
The limitations of available data have thus
limited understanding of the Southeast
Asian fauna, and the Philippine fauna in
particular, from both biogeographic and
ecological perspectives, and hence limited
conservation planning in a nation that is of-
ten cited as one of the most in need of ef-
fective conservation action (Mittermeier et
al. 1999, Ong et al. 2002, Wildlife Conser-
vation Society of the Philippines 1997).
Two of us (Esselstyn and Widmann) re-
cently conducted extensive surveys of the
mammals of Palawan Island, focusing on
the 15 sites described below. Esselstyn
worked from December 1999 to November
2000 at Sites 1-11, emphasizing (though
not exclusively) insectivorous bats, which
are the mammals most poorly known in the
Philippines (Heaney et al. 1998, Heaney &
Mallari 2002), and Widmann conducted
studies of bats, rodents, and larger mam-
mals from 1997 to 2002 at Sites 12—15;
Heaney visited briefly in April 2000. In this
paper, we report information collected dur-
ing these studies, emphasizing new data on
bats, and we include additional unpublished
records of mammals. We summarize infor-
mation on all additional species that were
not taken during this study but have been
documented on the island, and re-examined
some key specimens from prior studies. We
include descriptions of the habitats where
we conducted our surveys because of a gen-
eral paucity of such information, and use all
available information to evaluate conser-
vation status of the mammals. We include

VOLUME 117, NUMBER 3

measurements of the skulls of selected spe-
cies of insectivorous bats that have been es-
pecially poorly known.

Methods

At Sites 1-11, small non-volant mam-
mals were captured using locally made live
(cage) traps and Victor (snap) rat traps. Ap-
proximately 90% of live traps used mea-
sured 11 X 11 X 24 cm, and 10% measured
13 X 16 X 13 cm. Trap lines consisting of
approximately 70 traps (50 live traps and
20 snap traps) were placed in areas of tra-
versable terrain. Individual traps were
placed in locations of likely capture (e.g.,
near holes, along fallen logs, near root but-
tresses, etc.) along the line spaced at 5—15
m intervals. Most traps were set at ground
level, but in forest habitats we placed 5—
15% of the traps in elevated locations up to
2 m above ground level on fallen logs, hor-
izontal vines, etc. All but two trap lines
were set for three nights; the exceptions at
Sites 3 and 7 were set for five and two
nights, respectively. At Sites 12—14, a mix-
ture of live traps were used (see Site de-
scriptions). Most trap lines were baited with
fresh grilled coconut coated with peanut
butter. Other trap lines were baited with live
earthworms or bananas. All traps were
checked in the early morning and late af-
ternoon. Baits were changed at least once
daily, usually in the afternoon, and as nec-
essary in the early morning. Most live an-
imals were released at the site of capture.

We captured bats using a harp trap (ca. 2
x 2 m, 4 bank), mist nets (2 X 6 m, 16
mm mesh), butterfly nets, and hand capture
at Sites 1-11; only nets were used at Sites
12—15. Harp trap and net locations were se-
lected to be locations of likely capture (e.g.,
natural canopy breaks, over streams and
trails, around fruiting trees, and near poten-
tial roosting locations). The harp trap and
mist nets were usually set in a location for
three nights, and occasionally for only one
or two nights. During surveys of caves, we
primarily used the harp trap, and its loca-

273

tion was frequently changed. Mist nets were
continuously monitored during peak activ-
ity periods from 1800 to 2000 h (at Sites
12 and 13, until 2400 h) and then checked
again in the early morning. The harp trap
was checked periodically between 1800 and
2100 h and again in the early morning. In
forested areas, we searched for bats in po-
tential roosting locations (e.g., hollow trees,
rock formations, new banana leaves, etc.).
Most bats were released at the site of cap-
ture. For many of the bats, we report the
proportion of adult females that were preg-
nant on certain dates. These were palpated
externally to determine the presence or ab-
sence of an embryo before being released.

Forearm and cranial measurements were
taken by L. R. Heaney and D. S. Balete at
the Field Museum of Natural History
(FMNH). All voucher specimens from Es-
selstyn, which are cited below as specimens
examined, were preserved in fluid (with
skulls later removed and cleaned) and cat-
aloged at the FMNH; half of the vouchers
have been deposited at the National Muse-
um of the Philippines (NMP). Additionally,
data on several previously unreported spec-
imens housed in the University of Michigan
Museum of Zoology (UMMZ) and United
States National Museum of Natural History
(USNM) are included here.

Site Descriptions

See Fig. | for approximate locations of
study sites. The province of Palawan in-
cludes the main island called Palawan and
many smaller, nearby islands. The island is
politically divided into 13 municipalities,
one of which is Puerto Princesa City; the
municipalities and the city are subdivided
into barangays, and barangays into sitios.

Site I (10°03’00"N, 119°00'44"E) was in
lowland primary forest located along the
Tarabanan River in northern Puerto Prin-
cesa Municipality, at elevations ranging be-
tween ca. 100 and 200 m. Slope in the area
was generally moderate, rising from the riv-
er to the ridge-tops. Forest in the area was

274 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Jose Rizal
@

Sofronio Espanola

Brookes Point
Bataraza

Fig. 1. Map of Palawan Island, showing the locations of the primary towns (solid circles) and research sites
(solid triangles), and the position of Palawan in the Philippines (inset).

nearly undisturbed, which is uncommon at lection of minor forest products and hunting
this low elevation. We were aware of only of Sus barbatus and Macaca fascicularis.
one small human-made clearing (ca. 0.5 ha) Canopy ranged from 20 to 30 m in height
in the area; other disturbances included col- and was multi-layered. Canopy trees ranged

VOLUME 117, NUMBER 3

in diameter from 40—80 cm and had light
buttress development. Leaf litter was thin.
We surveyed small volant and non-volant
mammals for 13 days in January 2000.
Four trap lines were run for three nights,
yielding 3 Tupaia palawanensis, 21 Max-
omys panglima, and 1 juvenile Viverra tar-
galunga in 792 trap-nights. Three of the
lines were baited with coconut and peanut
butter, while one line was baited with live
earthworms. Forty-eight net-nights pro-
duced 1 Cynopterus brachyotis and 12
harp-nights produced 3 Rhinolophus acu-
minatus, 3 R. arcuatus, and 1 R. creaghi,
plus other means produced 11 Megaderma
spasma and 1 R. acuminatus.

Site 2 (9°42'14"N, 118°32’01”"E) was in
lowland primary forest located mid-way up
Mt. Salakot, between 300 and 700 m ele-
vation. Slope was rolling to moderately
steep. Several small streams dissected the
area. The only major human-caused distur-
bance in the area was an unused helicopter
landing pad and an abandoned road; the
road had a dense regrowth of ferns and
small trees. Sus barbatus was hunted, and
some almaciga trees (Agathis sp.) near the
upper reaches of the site had fallen due to
over-collection of resin. Agoho trees (Ca-
suarina sp.) gradually became more com-
mon as elevation increased. Canopy ranged
from 15 to 30 m in height and was multi-
layered. Canopy trees ranged in diameter
from 35—60 cm with the largest emergents
reaching 80—90 cm. Buttress systems were
only slightly developed and stilt root sys-
tems were present above 600 m, but rare.
Leaf litter was slightly deeper than at lower
elevations. We surveyed small volant and
non-volant mammals at this site for 20 days
during March and July 2000. Six trap lines
(three coconut and peanut butter-baited
lines, one banana-baited line, and two
earthworm-baited lines) yielded 16 Tupaia
palawanensis, 55 Maxomys panglima, and
1 Sundamys muelleri from 1272 trap nights.
Fifty-six net-nights yielded 1 Cynopterus
brachyotis, 1 Hipposideros diadema, 8
Rhinolophus arcuatus, and 1 R. creaghi.

AHS

Ten harp-nights yielded 1 Hipposideros
diadema, 16 Rhinolophus arcuatus, 10 R.
creaghi, 2 R. virgo, and 2 Kerivoula hard-
wickil.

Site 3 (10°07'28"N, 118°59'36"E) was in
primary montane and mossy forest located
near the peak of Cleopatra’s Needle (max-
imum elevation 1603 m), at elevations
ranging from 1300 to 1600 m. Slope was
moderate to extreme. Other than a trail to
the peak occasionally traveled by tourists,
there was little human-caused disturbance
in the area. No above-ground sources of
water were found near the site, but mist was
frequently present: during our stay of two
weeks during dry season, the area was al-
most continuously shrouded by a dense fog.
Vegetation type was montane forest up to
ca. 1500 m. At this elevation, the vegetation
began a transition to mossy forest. In mon-
tane forest, the canopy reached a height of
approximately 10 m. Trees, rocks, fallen
logs, and other stable surfaces were covered
with a thin layer of moss; epiphytic ferns
and orchids were abundant. Many trees had
an adventitious root system, but maintained
straight boles. The canopy was more open
here than at lower elevations. Above 1500
m, trees were shorter (2—4 m in height) and
took on a shrub form above 1550 m. Moss
growth was heavier at this elevation, and
vegetation became extremely dense at the
upper reaches. Pitcher plants (Nepenthes)
began to appear at about 1500 m and were
abundant by 1550 m. We surveyed small
volant and non-volant mammals at this site
for 12 days during February and March
2000. Three trap lines yielded 1 Tupaia pa-
lawanensis, 33 Maxomys panglima, and 3
Rattus tiomanicus in 740 trap-nights. Two
lines were run for three nights each and one
line was run for five nights; all three lines
were baited with coconut and peanut butter.
Forty-eight net-nights produced 2 Rhinolo-
phus arcuatus, and 12 harp-nights produced
no captures; 2 Pipistrellus javanicus were
captured by hand.

Site 4 (9°33'45"N, 118°27'54’"B) is a cave
known locally as ““Ma-ngit’’. It is located

276

along the Iraan River near Sitio Pamolkoan,
Barake, Aborlan, at ca. 430 m elevation.
The cave is in a small valley, restricted by
mountains to the east and west. A small,
seasonal stream flows through the cave. At
least six entrances to the cave were evident,
and many small tunnels connected medium
to small caverns, which ranged from well-
lit to completely dark. Very little distur-
bance was evident in or around the cave
due to its isolated location (ca. 10 hour hike
from the nearest road). The cave was sur-
rounded by a large expanse of primary low-
land forest, but some agricultural areas
were present within ca. 5 km. Disturbances
to local vegetation included collection of
rattan (Calamus spp.). We surveyed bats in
Ma-ngit Cave for six days during December
1999. We captured 819 bats belonging to
seven species (9 Eonycteris spelaea, 43
Hipposideros diadema, 367 Rhinolophus
creaghi, 4 R. virgo, 9 Miniopterus australis,
386 M. schreibersi, and 1 M. tristis) inside
the cave.

Site 5 (10°05'00"N, 118°51’06"E) is a
complex area of limestone karst containing
probably greater than 100 caves in Baran-
gays Tagabinet and Cabayugan, Puerto
Princesa; elevation is ca. 50 m. Caves in the
area ranged from tiny cracks too small to
enter, to large complexes of multiple cav-
erns with multiple entrances. Local terrain
was generally flat except for the sometimes-
massive limestone outcrops, which form
high-rising cliffs throughout the area. Caves
probably are present all over these complex
formations, but only the very few found
near ground level were accessible. We cap-
tured bats in and around five different
caves/cave complexes. These represented
some of the most accessible caves in the
area. Disturbance at the caves was moder-
ate, with vandalism and guano excavation
evident at most caves. Most of the caves
were surrounded immediately by agricul-
tural development. Both primary and sec-
ondary lowland forests were present in the
surrounding hills. We surveyed bats at these
caves for 14 days between March and May

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

2000. A total of 575 bats belonging to 10
species (18 Cynopterus brachyotis, 1 Me-
gaderma spasma, 100 Hipposideros ater,
239 H. diadema, 14 Rhinolophus arcuatus,
33 R. creaghi, 10 R. macrotis, 86 R. virgo,
15 Miniopterus australis, and 59 M. schrei-
bersi) were captured.

Site 6 (9°28'25"N, 118°30'21"E) was a
mostly abandoned agricultural area located
in Barake, Aborlan Municipality, at eleva-
tion ranging from 40—80 m. A small stream
flowed through the area and topography
was flat to rolling. Vegetation was a mosaic
of grassland (primarily Imperata cylindri-
ca) with sparse trees (mostly Vitex sp.),
cashew plantations, dense brush, and very
small (<1 ha) areas of secondary growth.
Frequent fires appeared to maintain this
area as a grassland. We surveyed small vo-
lant and non-volant mammals for four days
during June 2000. A single trap-line baited
with live earthworms yielded 17 Rattus ex-
ulans in 186 trap-nights. We captured 7 Cy-
nopterus brachyotis in 6 net nights, and 1
Kerivoula whiteheadi in 3 harp-nights.

Site 7 (9°29'15"N, 118°29'24”E) was in a
narrow band of secondary forest in Barake,
Aborlan Municipality, located between dis-
turbed habitat at lower elevation (Site 6)
and primary and good secondary forest at
higher elevation. Elevation ranged from
80—140 m; slope was rolling to moderately
steep, and two small streams dissected the
area. Canopy height varied from 5—20 m.
Woody vines and lianas were common and
vegetation was quite dense in areas. Wild
bananas (Musa spp.) were abundant, but
patchy in distribution; leaf litter depth was
highly variable. We surveyed this site for
small volant and non-volant mammals for
four days during June 2000. A single trap
line baited with live earthworms yielding
120 trap-nights produced 1 Rattus exulans.
Six net-nights produced 27 Cynopterus
brachyotis and 1 Macroglossus minimus,
and 3 harp-nights yielded 1 Hipposideros
diadema and | Kerivoula pellucida.

Site 8 (9°59'47"N, 118°56'43"E) is a
large cave complex located in a limestone

VOLUME 117, NUMBER 3

karst formation on top of the first ridge up
from San Rafael, Puerto Princess, at an el-
evation of ca. 250 m. The cave complex is
known locally as ““Taraw’’. The cave sys-
tem appeared to be quite large; we were
unable to explore much of it due to a lack
of climbing equipment and expertise. Some
caverns exceeded 20 m in height, while oth-
ers were quite small. We found no perma-
nent water in or around the cave, but evi-
dence of storm flow was present. Evidence
of vandalism and guano collection was pre-
sent. A mixture of habitats surrounded the
cave complex: between the cave and the
community of San Rafael, the vegetation
was dominated by brushland and agricul-
tural developments, while on the other side
of the cave secondary and primary forest
dominated, with mixed areas of slash-and-
burn fields. We surveyed bats at this cave
for five days during July 2000. We captured
2775 bats representing 10 species (7 Hip-
posideros ater, 43 H. diadema, 90 Rhino-
lophus acuminatus, 240 R. arcuatus, 239 R.
creaghi, 25 R. macrotis, 151 R. virgo, 1257
Miniopterus australis, 711 M. schreibersi,
4 immature M. sp., and 8 Myotis macrotar-
SUS).

Site 9 (9°39'40"N, 118°27'48"F) is a
small cave located in Sitio Labtay, Napsan,
Puerto Princesa City. The cave, which con-
sisted of a single chamber 1—2 m wide, 3—
6 m high, and ca. 30 m long, was in a nar-
row canyon along the Panagurian River at
ca. 280 m. Vegetation in the area consisted
of good-quality secondary forest, second-
growth forest, and agricultural develop-
ments. We trapped bats with a harp trap,
mist nets, and a butterfly net in the cave
and surrounding forest for five days during
August 2000. We captured 73 bats (4 Cy-
nopterus brachyotis, 68 Hipposideros dia-
dema, and 1 Rhinolophus arcuatus).

Site 10 (10°44'00”, 119°34’23”) is a cave
located near sea level in Sitio Sader, Ban-
tulan, Taytay Municipality. The cave con-
sisted of a single chamber (ca. 3—8 m wide,
3-7 m high, and 40 m long) with a large
(>2.5 m diameter) entrance at each end.

277

Minor damage had been done by both trea-
sure hunters and guano collectors. The sur-
rounding vegetation was dominated by ag-
ricultural areas with some strips and patches
of residual forest. Large expanses of sec-
ondary and logged-over forest were found
nearby in the vicinity of Lake Manguao. We
captured 115 bats belonging to 4 species
(48 Eonycteris spelaea, 1 Rousettus am-
plexicaudatus, 64 Hipposideros diadema,
and 2 Miniopterus tristis) using a harp trap
and butterfly net at one of the entrances to
the cave during three days in October 2000.

Site 11 (10°46'34"N, 119°31'52”E) was
located around the perimeter of Lake Man-
guao, in Barangays Poblacion and Bantu-
lan, Taytay Municipality. The area was
dominated by secondary and logged-over
forest, with disturbance from slash and burn
agriculture being found throughout the area.
The area retained ca. 60% forest cover.
Slope was generally moderate to steep, and
elevation ranged from ca. 40-250 m. We
trapped for small volant and non-volant
mammals in forest, agricultural habitats,
and two small caves near the lake for 14
days during October and November 2000.
We totalled 471 trap-nights in three lines
baited with coconut and peanut butter, and
captured 3 Tupaia palawanensis, 23 Max-
omys panglima, and 1 Rattus exulans. For-
ty-two net-nights yielded 53 Cynopterus
brachyotis, 2 Macroglossus minimus, and 1
Rousettus amplexicaudatus, and 13 harp-
nights produced 1 Megaderma spasma, |
Rhinolophus acuminatus, 2 Kerivoula hard-
wickii, and 5 Tylonycteris pachypus. Addi-
tionally, we captured 4 Megaderma spasma
and 4 R. acuminatus by hand.

Site 12 (9°27'48'N, 118°32'16"E) was lo-
cated at the “rainforestation”’ site in Sitio
Kandis, Aborlan Municipality, in forest/
grassland mosaic at ca. 40 m above sea lev-
el, about seven km away from the next
good secondary forest in the foothills of the
Victoria Range. Charcoal making, logging,
rattan collection, grazing and burning were
common until 1994 when such activities
were made illegal. The terrain was flat to

278

rolling, dissected by two creeks. The site
consisted of 5.5 ha Imperata cylindrica
grassland interspersed with single shrubs
and trees, predominantly Antidesma ghae-
sembilla, which forms the fire climax in
more open situations, with Vitex pubescens,
Guioa pleuropteris, Tarenna stenantha, Fa-
graea fragrans, Lantana camara, and Mus-
saenda philippica in protected areas not af-
fected by fire in the last ten to fifteen years.
About 4.5 ha consisted of regenerating for-
est, close to two seasonal creeks, dominated
by Garcinia benthami, G. parviflora, Can-
arium asperum, Polyscias nodosa, and Bar-
ringtonia curranii. Undergrowth was mod-
erate to very dense. Canopy height was on
average eight meters, with some taller
emergents such as Nephelium sp. and Dip-
terocarpus gracilis. The most conspicuous
vine was Gnetum latifolium. Macrophytic
epiphytes were virtually absent. Leaf litter
layer was usually not closed, except in very
dry years. Surveys were conducted regular-
ly from 1997-2000 with 10 medium-sized
Sherman live traps, 10 commercial live rat
traps, and 55 wire mesh traps (measure-
ments equal to medium Shermans). Baits
were roasted coconut with peanut butter,
but mostly fruits available in the area. A
total of 3514 trap-nights yielded 169 small
mammals (28 Tupaia palawanensis, 8 Sun-
dasciurus juvencus, 11 Rattus exulans, 16
Rattus tiomanicus, 104 Maxomys panglima,
and 2 Sundamys muelleri). Mist nets (2 X
6 m) were set along trails in forest, in grass-
land, gaps in the shrub cover, and rarely in
the canopy (ca. 8 m high). Capture sites
were often near or in fruiting shrubs or
trees, since the main focus of the study was
on frugivores. From 1997 to 2000, a total
of 482 net-nights yielded 1257 bats (1 Ac-
erodon leucotis, 829 Cynopterus brachy-
otis, 394 Macroglossus minimus, 5 Eonyc-
teris spelaea, 4 Rousettus amplexicaudatus,
18 Megaderma spasma, \ Hipposideros
diadema, 4 Scotophilus kuhlii, and 1 Mu-
rina cf. tubinaris); all but the M. cf. tubi-
naris were released.

Site 13 (09°13'N, 118°26'E) was on Rasa

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Island, Narra Municipality, a small (8.3
km?) shallow coral island, 1.8 km offshore
in the Sulu Sea. Approximately two-thirds
of the island was covered with mangrove
and one-third with coastal forest over lime-
stone. About five percent of the latter had
been converted into coconut plantation. Se-
lective logging was done until the early
1990s, resulting in the complete loss of ma-
ture Intsia bijuga. The mangrove consisted
of nine species of the genera Rhizophora,
Sonneratia, Avicennia, Bruguiera, Aegicer-
as, and Ceriops. Canopy height was vari-
able, usually between 8 and 15 m. Emer
gent trees (e.g., Garuga floribunda and
Pterocymbium taluto) ranged up to 42 m.
Leaf litter layer was not closed, except un-
der very dry conditions. Barren coral rocks
and crevices were ubiquitous. Buttresses
were a common feature of all emergent for-
est trees. Under open conditions, an herbal
layer consisting of /mpatiens sp. was pre-
sent. Vines were abundant, including climb-
ing bamboo Dinochloa sp., often forming
dense tangles. Macrophytic orchids were
present, but relatively scarce. Traps and nets
were set along a trail within the coastal for-
est. Traps were baited with roasted coconut
with peanut butter, and a few with crickets.
Most traps caught hermit crabs. Nets were
set in the understory, which was very open
from January to April 2002 and only pro-
vided very few fruits due to an extended
dry spell. Trapping totaled 104 trap-nights,
and produced 3 Rattus tanezumi and 2 R.
tiomanicus. Netting totaled 28 net nights,
and yielded 2 Cynopterus brachyotis, 5 Ma-
croglossus minimus, and 5 Megaderma
spasma; all bats were released, and the rats
preserved as vouchers.

Site 14 (9°17'N, 118°27’E) was in fresh-
water swamp forest in Narra Municipality,
in about 5 ha of remnant forest along Tar-
itien River. The habitat was dominated by
two woody species, Nauclea orientalis and
Pandanus sp., at lower elevations, which
are flooded for at least six months. The herb
layer was not extensive and was dominated
by Acrostichum sp. The higher portions

VOLUME 117, NUMBER 3

were dominated by pioneering species of
early to medium successional stages, such
as Trema orientalis, Vitex pubescens, and
Commersonia bartramia. Even during ex-
treme dry spells like that in the first half of
2002, there were isolated open water bodies
left, which connected to several creeks dur-
ing the rainy season. The swamp forest is
bordered by ricefields and grassland. Twen-
ty trap-nights yielded 1 Rattus exulans and
1 R. tiomanicus. Twelve net-nights yielded
27 Cynopterus brachyotis, 5 Macroglossus
minimus, and 1 Megaderma spasma; all
bats were released and the rats preserved as
vouchers.

Site 15 (10°12'N, 118°55’E) was along
the “‘jungle trail” near the Central Park Sta-
tion in Puerto Princesa (formerly St. Paul)
Subterranean River National Park
(PPSRNP). Primary lowland forest on steep
slopes ascended from sea level to about 40
m. Five net-nights on 15 September 1996
yielded 1 Cynopterus brachyotis, 2 Rhino-
lophus arcuatus, 3 Rhinolophus virgo, and
2 Rhinolophus sp.; all were released. Ad-
ditionally, all three authors made visual ob-
servations at various times.

Accounts of Species

Order Insectivora
Family Soricidae—Shrews

Crocidura palawanensis.—We never en-
countered this poorly known species. It is
endemic to the Palawan faunal region and
has been taken in old-growth rain forest and
shrubby second growth (Heaney & Ruedi
1994); the holotype came from “‘deep forest
near the sea at ... Brooke’s Point’”’ (Taylor
1934), a second from near sea level in Ba-
buyan, Puerto Princesa (Hoogstraal 1951,
Sanborn 1952), and a third from 3600—
4350 ft (ca. 1100—1300 m) on Mt. Mantal-
ingajan (USNM); two additional specimens
are from Balabac (Heaney & Ruedi 1994).
IUCN (2002) lists this species as Vulnera-
ble, but current definitions suggest that Data
Deficient would be more appropriate.

Crocidura sp.—Reis & Garong (2001)

279

reported a single humerus of a shrew, sub-
stantially smaller in size than C. palawa-
nensis, from undated sediments in a small
rock-shelter cave near Tabon Cave on Lip-
uun Point, near Malunut Bay, in Quezon
Municipality (near the location of the town
of Quezon as shown in Fig. 1). They de-
scribed the specimen as being similar in
size to C. monticola from Borneo. We ten-
tatively include it in our tallies of native
species of Palawan, but we recommend that
it be sought by trapping with small snap-
traps baited with live earthworms and pit-
fall traps.

Suncus murinus.—This introduced com-
mensal is abundant in urban and agricultur-
al areas (Rabor 1986); in forest, it is rarely
present, but occasionally is common (Hea-
ney et al. 1989, Heaney & Tabaranza 1997).
It is found throughout Asia and Indo-Aus-
tralia, including the Philippines (Heaney et
al. 1998). We observed this species fre-
quently in houses in Puerto Princesa City
and the State Polytechnic College of Pala-
wan in Aborlan Municipality.

Order Scandentia
Family Tupaiidae—Tree Shrews

Tupaia palawanensis.—TVhis common
species is endemic to the Palawan faunal
region (Wilson 1993); it is related to T. glis,
which is widespread on the Sunda Shelf
(Corbet & Hill 1992). It is widespread on
Palawan (Taylor 1934), and is usually com-
mon in secondary and primary lowland for-
est, though local densities may be highly
variable between apparently similar habitats
(Dans 1993, Hoogstraal 1951, Sanborn
1952). It is rare in montane forest, and com-
mon but patchy in agricultural areas. We
captured and/or observed this species in co-
conut and cashew plantations, brushy areas
with a few small trees (Sites 6, 11, and 12),
secondary and logged-over forest (Sites 7
and 11), and primary forest (Sites 1, 2, 3,
and 15) from near sea level to 1400 m.
IUCN (2002) lists this species as Vulnera-
ble, but we concur with Heaney et al.

280

(1998) that the species should be delisted
due to the variety of habitats used and its
apparent abundance. Specimens examined:
Deasitew lan) SiteyZn Gy):

Order Chiroptera
Family Pteropodidae—Fruit Bats

We follow Ingle & Heaney (1992) and
Heaney et al. (1998) in regarding reports of
Haplonycteris fischeri (Kock 1969) and
Ptenochirus minor (Yoshiyuki 1979) from
Palawan as erroneous, probably originating
in mislabeled specimens. Pteropus hypo-
melanus is known from Cuyo Island, at the
northeast edge of the Palawan faunal region
(Heaney et al. 1998), as well as in the oce-
anic Philippines and on islets around Bor-
neo, and should be sought on Palawan.

Acerodon leucotis.—This poorly known
species is endemic to the Palawan faunal
region. Hoogstraal (1951) found the species
in an area with patches of ““much disturbed
remnants of original forest and dense sec-
ond growth forest’? on Busuanga Island,
and two specimens were taken at Santiago,
Iwahig (in Puerto Princesa) on Palawan
(Sanborn 1952). A specimen from Bat Is-
land (Barangay Tagburos, in Honda Bay),
taken by P. O. Glass in 1978, is housed in
the UMMZ. Heaney sighted large numbers
of medium-sized, pale-furred flying foxes at
Site 15 in April 2000, in the clearing of the
old park headquarters near the center of the
park (“Central Park’’), that were probably
this species. Widmann captured one at Site
12 at a height of 5 m, and saw others feed-
ing in the canopy at ca. 8 m height. The
IUCN (2002) lists this species as Vulnera-
ble; we regard it as Data Deficient.

Cynopterus brachyotis.—Found through-
out Southeast Asia; in the Philippines, it is
common to abundant in secondary forest
and agricultural areas, and rare in primary
forest (Heaney et al. 1998); Sanborn (1952)
reported many from Palawan. We netted
this species frequently in secondary forest
and agricultural areas at Sites 6, 7, 11, and
12, in freshwater swamp forest at Site 14,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

and in coastal forest on Rasa Island (Site
13). We also captured this species in pri-
mary forest in a tree fall gap (Site 1), over
a stream (Site 2), and at a place with no
visible disturbance (Site 15). We found
them roosting in various-sized groups in
three caves at Sites 5 (there appeared be
less than 50 in each of two caves) and 9
(ca. 300 individuals); although this species
occasionally roosts in caves on Borneo
(Payne et al. 1985), there are no previous
records of such roosts in the Philippines.
On several occasions we captured them car-
rying whole green figs (Ficus sp.) during
flight; two of these individuals were return-
ing to a cave at Site 5 between 1900—
2000 h.

Out of 20 adult females caught at Site 14
on 20 March 2002, 15 were pregnant and 5
were carrying a single suckling young. On
1 and 2 April 2000, we captured three adult
females at Site 5; all were carrying a single
suckling infant during flight. On 17 May
2000 we captured eight adult females at
Site 5; one was pregnant, one was carrying
a suckling infant during flight, and two
were emaciated and may have recently
weaned their young. On this date, we also
captured a male with enlarged mammaries
(see Francis et al. 1994). On 30 October
2000, among 25 adult females, none were
pregnant but one was lactating. Specimens
examined: 4, Site 1 (1), Site 5 (3).

Eonycteris spelaea.—This widespread
Southeast Asian species 1s common in ag-
ricultural areas in the Philippines, where all
known roosts are in caves (Heaney et al.
1998, Rickart et al. 1993). Sanborn (1952)
reported a large series from a cave above
Tanabog, Palawan. We netted five individ-
uals at Site 12, but most of our records
came from caves in lowland forest. We
found this species roosting 1n caves at Sites
4 and 10; at Site 4, the roosting population
appeared to exceed 2000. At Site 10, there
was an extremely large population (proba-
bly >50,000) of small pteropodids roosting
inside the cave. We captured 49 pteropodids
at the entrance to the cave, 48 of which

VOLUME 117, NUMBER 3

were E. spelaea and one was a Rousettus
amplexicaudatus. On 19 December 1999,
all three adult females we captured at Site
4 were pregnant. On 21 October 2000 at
Site 10, we captured 11 adult female E. spe-
laea, four of which were carrying an infant
during flight and five of which were preg-
nant. This species is heavily hunted in some
areas of the Philippines (Rickart et al. 1993,
Utzurrum 1992), but we observed no evi-
dence of that being the case on Palawan.
Specimens examined: 5, Site 4 (3), Site 10
(2).

Macroglossus minimus.—In the Philip-
pines, this widespread Australasian species
is common in secondary forest and agri-
cultural areas and uncommon in primary
forest up to more than 2000 m (Heaney et
al. 1998, 1999). We captured this species in
secondary lowland forest (Sites 7 and 12)
and agricultural clearings (Site 11), usually
near wild or domestic banana plants (Musa
spp.), and in freshwater swamp forest in
Narra Municipality and Rasa Island (Sites
13 and 14). Specimens examined: 3, Site 7
GD, Sit i),

Pteropus vampyrus.—In the Philippines,
this widespread Southeast Asian species oc-
curs in primary lowland forest and adjacent
agricultural areas (Heaney et al. 1998; Ra-
bor 1955, 1986; Rickart et al. 1993; San-
born 1953; Taylor 1934). Widmann esti-
mated 400 individuals on Malinau Island,
Aborlan Municipality in 1998, 570 on Rasa
Island on 12 November 1999, and a small
colony (ca. 40 individuals) at Lagan on Du-
maran Island on 27 October 2001, based on
departure counts. Flying foxes commonly
sighted in Puerto Princesa City around
mango and guyabano (= sour sop) trees are
probably this species. In 1998, we found
two individuals of this species that ap-
peared to have been electrocuted on power
lines, one at the Provincial Agriculture Cen-
ter in Irawan, Puerto Princesa, and the other
at the State Polytechnic College, Aborlan.
We believe this species to be common over-
all, but under moderate pressure due to

281

hunting and perhaps to electrocution on
power lines.

Rousettus amplexicaudatus.—Within the
Philippines, this widespread Southeast
Asian species is commonly found in agri-
cultural habitats up to 500 m and rarely in
primary lowland forest (Heaney et al.
1998). All known roosting sites are in caves
(Heaney et al. 1989, 1991, 1998, 1999; Hei-
deman & Heaney 1989; Rickart et al.
1993). According to Payne et al. (1985), R.
amplexicaudatus often roosts in association
with Eonycteris spelaea. We netted one in-
dividual from a cave at Site 10 containing
a large population of E. spelaea, one in an
agricultural clearing in Site 11, and four in
forest-grassland mosaic at Site 12; all of
these sites are in heavily disturbed areas be-
low 60 m. Specimens examined: 2, Site 10
(1), Site 11 (1).

Family Emballonuridae—Sheath-tailed
Bats

There are no known records of Saccolai-
mus saccolaimus from Palawan, but its
widespread distribution from India to New
Guinea, including the oceanic Philippines
(Heaney et al. 1998), suggests that it may
be present and should be sought.

Emballonura alecto.—The Philippine
sheath-tailed bat is known from Borneo, the
Philippines, and Sulawesi (Heaney et al.
1998); we never encountered this species
on Palawan, but Taylor (1934:200) captured
five individuals “under an overhanging
rock along Iwahig River, near the base of
Thumb Peak’.

Taphozous melanopogon.—The bearded
tomb bat is widespread in southern Asia
(Heaney et al. 1998). In the Philippines, it
is common in urban areas and lowland ar-
eas with limestone caves and rare in forest
(Rickart et al. 1993, Sanborn 1952). There
is a previous record from the vicinity of
Puerto Princesa (Allen 1922), and A. C. Al-
cala collected 6 specimens from Sitio Mal-
abusog, Tinitian, Roxas Municipality in

282

1984 which are deposited in the UMMZ.
We never encountered this species.

Family Megadermatidae—False Vampire
and Ghost Bats

Megaderma spasma.—This widespread
southern Asian species is common in pri-
mary lowland forest and disturbed forest in
the Philippines (Heaney et al. 1991, 1998,
1999; Rickart et al. 1993). We captured this
species from sea level to ca. 500 m in sec-
ondary forest (Site 7), primary forest (Sites
1 and 2), in a bamboo thicket (Site 11), and
in or near caves (Sites 5 and 11). It was the
most common insectivorous bat netted in
forest-grassland-mosaic (Site 12), in swamp
forest (Site 14), and in coastal forest (Site
13). At Site 1, we found this species roost-
ing in small groups (<10) in four hollow
trees distributed throughout the area. At
Site 11 we found ca. 12 individuals roosting
in a small cave (ca. 0.5—3 m wide, 0.3—1.5
m high, and 10 m long) along with Rhino-
lophus acuminatus. We also found two in-
dividuals roosting in a small cave (also Site
11) that had been severely disturbed by
treasure hunters three years earlier. Cranial
measurements of three individuals (Table 1)
are slightly smaller than those of specimens
from Leyte and Biliran (Rickart et al. 1993)
and southern Luzon (Heaney et al. 1999).
Specimens examined: 3, Site 1 (3).

Family Rhinolophidae—Horseshoe and
Roundleaf Bats

Several poorly known but apparently
widespread species in this family occur on
the Sunda Shelf and in the oceanic Philip-
pines and should be sought on Palawan;
these include Hipposideros cervinus and H.
lekaguli (Balete et al. 1995, Heaney et al.
1998, Ingle & Heaney 1992).

Hipposideros ater.—Occurs from India
to Australia (Heaney et al. 1998). Known
from lowland and montane forest and caves
(Heaney et al. 1991, 1998; Payne et al.
1985, Rickart et al. 1993). We found this
species to be uncommon to abundant in

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

three caves in disturbed lowland forest at
50 to 250 m elevation at Sites 5 (17% of
575 captures) and 8 (<1% of captures).
During March to April 2000, none of the
26 females we captured at Site 5 were preg-
nant or lactating, but on 19 and 20 May
2000, 25 of 30 adult females were pregnant.
We have re-examined a specimen from Pa-
lawan in the UMMZ identified by Allen
(1922) as H. bicolor, and a series from the
Tigoplan River, Palawan in FMNH reported
by Sanborn (1952), and now consider them
to be H. ater; thus, we now know of no
records of H. bicolor from Palawan. Cranial
measurements of 5 individuals (Table 1) are
smaller than those of H. bicolor (Heaney et
al. 1999, Ingle & Heaney 1992) but match
those of H. ater (Ingle & Heaney 1992,
Rickart et al. 1993). Specimens examined:
5, Site 5 (4), Site 8 (1).

Hipposideros diadema.—The diadem
roundleaf bat is widespread from Myanmar
to the Solomon Islands, with many previous
records from Palawan (Allen 1922, Heaney
et al. 1998). In the Philippines, it is com-
mon in disturbed forest, agricultural areas
(Ingle 1992, Rickart et al. 1993), and pri-
mary forest (Heaney et al. 1998, Rickart et
al. 1993). Reis & Garong (2001) reported
specimens from sediments in a rock-shelter
near Tabon Cave, Quezon Municipality dat-
ed to 11,130 BP. We captured this species
from sea level to 600 m in disturbed grass-
land-forest mosaic (Sites 6 and 12), second-
ary forest (Site 7), primary forest (Site 2),
and at nearly all caves we visited (Sites 4,
5, 8, 9, and 10), and we observed large
numbers (probably thousands) in the un-
derground river cave at PPSRNP. All of the
roosts we identified held groups of H. dia-
dema numbering greater than 200. Thirteen
adult females captured in December 1999
included none that were pregnant or lactat-
ing. At Site 5 in March to April 2000, none
of the 54 adult females were pregnant or
lactating, but between 15 and 20 May 2000
26 of 43 were pregnant and one was car-
rying a suckling infant during flight. One of
21, one of 12, and none of 13 adult females

283

VOLUME 117, NUMBER 3

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284

from July, August, and October were preg-
nant (Sites 8, 9, and 10, respectively). Spec-
imens examined: 5, Site 4 (2), Site 5 (2),
Site 9 (1).

Rhinolophus acuminatus.—This poorly
known species occurs from Thailand to
Lombok and Palawan, but not elsewhere in
the Philippines (Heaney et al. 1998; speci-
mens from Negros reported by Csorba et al.
(2003) as this species were mislabelled). It
is found in lowland forest on Borneo
(Payne et al. 1985) and in secondary low-
land dipterocarp forest on Banggi (Md. Nor
1995). We captured this species in caves
from ca. 60 to 250 m in caves (Sites 8 and
11), a bamboo thicket (Site 11), and pri-
mary forest (Site 1). At Site 8, we captured
90 individuals out of 2775 captures. At Site
11 we found ca. 20 individuals roosting in
a small cave (ca. 0.5—3 m wide, 0.3—1.5 m
high and 10 m long) with ca. 12 Megad-
erma spasma. At Site 1, we took two in-
dividuals over a small stream just below a
rock outcrop containing many fissures suit-
able for roosting bats. We also captured a
single individual in our temporary living
quarters at Site 1 after we observed for sev-
eral days a bat feeding inside our semi-en-
closed tent for several minutes daily be-
tween 0430 and 0600 h. Of 15 adult fe-
males taken in July 2000 at Site 8, one was
pregnant and one was lactating. Cranial
measurements (Table 1) match those pre-
viously available (Ingle & Heaney 1992)
that are based on series from Balabac and
Busuanga reported by Kuntz (1969) and
housed in the USNM. We also refer two
specimens collected by A. C. Alcala on 13
July 1984, at Malabusog, Roxas Munici-
pality housed at UMMZ (162885 and
162886) to this species, and include them
in Table 1.

Sanborn (1952) reported a single speci-
men from Palawan (housed in FMNH) that
he regarded as the first record from Pala-
wan. However, we have determined that a
single specimen (UMMZ 53112) collected
in the late 1800s by the Beal/Steere Expe-
dition and subsequently named R. ander-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

seni aequalis (Allen 1922) was this species.
Heaney compared this specimen, which is
the holotype and only known specimen, to
series of all species currently known from
Palawan. It was unambiguously identified
as R. acuminatus; as noted by Medway
(1977:32), a dorsal connecting process with
a prominent triangular point (as shown by
Medway 1977 fig. 6a and by Ingle and Hea-
ney 1992 fig. 13a) is present, the base of
the sella is not expanded into a cup, the
median groove of the horseshoe is not
broadened, and papillae are not present.
The ears (18 mm) are less than half of the
length of head plus body, and the forearm
is 46.4 mm. The skull (measurements in Ta-
ble 1) is virtually identical to those in the
series from Sites 1 & 8, including overall
size and shape, nasal swellings, braincase
breadth and inflation, toothrows, palatal
bridge, foramina in the roof of the posterior
portion of the nasal passage, and bullae. Be-
cause R. acuminatus was named by Peters
in 1871 (from Gadok, Java), we therefore
recognize R. anderseni aequalis as its ju-
nior synonym. We note that Cabrera (1909)
described Rhinolophus anderseni “proba-
bly from Luzon’’. Aside from the holotype
of R. anderseni aequalis, no specimens
have subsequently been referred to this spe-
cies. Csorba et al. (2003) tentatively as-
signed R. anderseni Cabrera as a junior
synonym of R. arcuatus on the basis of the
original description plus new drawings of
the noseleaf and measurements of the skull,
but without examining the holotype. We
provisionally accept this, but point out the
need for direct examination and compari-
sons. IUCN (2002) lists R. acuminatus as
Data Deficient. Specimens examined: 8,
Site 1 (4), Site 8 (1), Malabusog (2), and
the holotype of R. anderseni aequalis.
Rhinolophus arcuatus.—Widespread
from Sumatra to New Guinea (Heaney et
al. 1998). Specimens from the Philippines
currently identified as R. arcuatus may con-
sist of two or more species (Heaney et al.
1991, 1999; Ingle & Heaney 1992: Rickart
et al. 1993). Individuals referred to this

VOLUME 117, NUMBER 3

“‘species”” have been found in agricultural
areas, secondary forest, and primary low-
land, montane, and mossy forest (Heaney et
al. 1991, 1999; Ingle 1992; Rickart et al.
1993). We regularly captured this species at
elevations from sea level to 1400 m in low-
land primary forest (Sites 1 and 2), mon-
tane forest (Site 3), and caves (Sites 5 and
8). We also captured a single individual in
the understory of mature but disturbed for-
est near a cave at Site 9, and we tentatively
identified two individuals from Site 15
(which were released) as belonging to this
species. Of 8 adult females taken between
15 and 20 May 2000 at Site 5, 5 were preg-
nant and one was lactating. Of 22 adult fe-
males taken in July 2000 at Site 2, one was
lactating, and of 189 captured in July at Site
8, none were pregnant but 14 were lactat-
ing. These are the first specimens of this
species from the Palawan faunal region.
Cranial measurements (Table 1) closely
match those of specimens from Leyte
(Rickart et al. 1993) and southern Luzon
(Heaney et al. 1999). Specimens examined:
5, Site 1 (2), Site 2 (1), Site 3 (2).
Rhinolophus cf. borneensis.—A_ single
specimen taken by P. O. Glass on 31 Jan-
uary 1978 at “Sabang, Buenavista” (in
Barangay Cabayugan, near Ulugan Bay in
Puerto Princesa Municipality, ca. 10°05’N,
118°49'E; UMMZ 161395) appears to be
this species. It was previously known from
Indochina, the Malay Peninsula, Java, and
Borneo, as well as some smaller islands in
the southern South China Sea (Corbet &
Hill 1992, Csorba et al. 2003); this is the
first record from Palawan and from the
Philippines. Cranial measurements and fea-
tures (Table 1) closely match specimens in
FMNH from Sarawak, the Natuna Islands,
and Sabah, and external features are similar,
but because we have only a single speci-
men, the identification is tentative. On Bor-
neo, “the species roosts in caves, some-
times in colonies of several hundred indi-
viduals” (Payne et al. 1985). P. O. Glass (in
litt.) noted that he captured the specimen in
a mist net in a small banana grove in an

285

area of mixed agricultural/second growth
forest within 1 km of mature forest. Spec-
imen examined: 1, from Sabang.
Rhinolophus creaghi.—This species was
previously known from Borneo and Madura
Islands, where it often roosts in caves (Cor-
bet & Hill 1992, Csorba et al. 2003, Koop-
man 1993, Medway 1977, Payne et al.
1985). This is the first record of this species
from Palawan Island and the Philippines.
On Palawan, we found it to be common in
primary lowland forest from near sea level
to at least 700 m. We captured one individ-
ual at Site 1 and 11 individuals at Site 2. It
roosts in caves, often in large numbers; we
captured 368 (45% of captures) at Site 4,
239 (9% of captures) at Site 8, and 33 (6%
of captures) at Site 5. Of 135 adult females
captured in December 1999 at Site 4, 8
were pregnant. Of 16 captured at Site 5 in
March to April, none were reproductively
active; of 11 at Site 2 and 151 at Site 8
(July 2000), none were pregnant but one
and 12 were lactating, respectively. Cranial
measurements (Table 1) show this to be the
largest member of the genus on Palawan;
our specimens are not distinguishable from
a small series from Borneo (FMNH 4707 1—
47075). Two previously unidentified speci-
mens from Mt. Salicod, 2300 ft. (which
may be the same mountain as Mt. Salakot,
Site 2; P. O. Glass, in lit.), taken by P. O.
Glass in 1978 and housed in the UMMZ,
were taken earlier but were not reported;
cranial measurements from these specimens
are included in Table 1. This species is list-
ed as Near-Threatened by IUCN (2002).
Specimens examined: 7, Site 4 (3), Site 5
(1), Site 8 (1), Mt. Salicod (2).
Rhinolophus macrotis.—This poorly
known species ranges from India to Su-
matra and the Philippines, where it is
known from lowland forest with some re-
cords from caves (Heaney et al. 1998, Ingle
1992). Our specimens represent the first re-
cord from Palawan. We captured this spe-
cies in or near three caves in disturbed low-
land forest at SO—250 m at Sites 5 (10 cap-
tures) and 8 (25 captures). The species ap-

286

pears to be uncommon. At Sites 5 and 8 it
represented less than 5% and 1% of our
captures, respectively. At Site 5 on 20 May
2000, we captured and examined two adult
females; both were pregnant. Of 8 adult fe-
males captured in July at Site 8, none were
pregnant or lactating. Cranial measurements
of 5 individuals (Table 1) fall within or near
the range for the species in Ingle and Hea-
ney (1992). Specimens examined: 5, Site 5
(4), Site 8 (1).

Rhinolophus virgo.—This Philippine en-
demic is widely distributed within the Phil-
ippines (Heaney et al. 1998). It is known
from secondary forest, primary lowland
forest, reaching mossy forest on small, low-
lying islands, and often roosts in caves
(Heaney et al. 1991, Ingle 1992, Rickart et
al. 1993); Sanborn (1952) reported a series
from Tanabog, Palawan. This species ap-
peared to be rare in the cave at Site 4 (0.5%
of captures), common at Site 8 (151 cap-
tures, about 6% of total) and abundant in
some caves at Site 5 (15% of captures). We
also captured this species in primary forest
at Sites 2 and 15. Of 29 adult females cap-
tured between 27 March and 4 April 2000,
none were pregnant or lactating. Of 35 fe-
males captured at Site 5 between 15 and 20
May 2000, 16 were pregnant and 5 were
lactating. Of 78 females taken at Site 8 in
July 2000, 6 were pregnant and 2 were lac-
tating. Cranial measurements (Table 1) fall
within the range (Ingle & Heaney 1992) or
very near the range (Rickart et al. 1993) of
previously available individuals. IUCN
(2002) lists this species as Near-Threatened.
Specimens examined: 6, Site 4 (2), Site 5
(4).

Family Vespertilionidae—Vesper and
Evening Bats

This diverse family of bats is generally
poorly documented, and more species
should be sought on Palawan, including
such widespread taxa as Harpiocephalus
harpia, Murina cyclotis, Myotis ater (which
Hill 1983 and Corbet & Hill 1992 have

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

shown to be distinct from Myotis muricola
and to be present on Culion Island in Pa-
lawan faunal region), Philetor brachypte-
rus, and Pipistrellus tenuis.

Glischropus tylopus.—This poorly
known species is found from Myanmar to
the Molucca Islands and Palawan (Heaney
et al. 1998). In Peninsular Malaysia it roosts
in rock crevices, bamboo, and in new ba-
nana leaves (Payne et al. 1985). We never
encountered this species, but it is repre-
sented by a specimen in the USNM (Hol-
lister 1913).

Kerivoula hardwickii.—This species is
widespread from India and southern China
to the Lesser Sunda Islands and the Phil-
ippines (Heaney et al. 1998). It was previ-
ously known from lowland, montane, and
ridge-top mossy forest from 500 to 1600 m
in the Philippines (Heaney et al. 1999,
Rickart et al. 1993). Everett (1889) includ-
ed mention of this species from Palawan. A
previous record from UMMZ (Heaney et al.
1998) has been re-identified as K. white-
headi, as noted below. Payne et al. (1985)
reported the species to “‘frequent the un-
derstory of tall forest’”’ on Borneo, and Md.
Nor (1995) caught one in primary lowland
dipterocarp forest on Banggi Island “in the
axil of a leaf on a rattan vine | m above
ground’’, and netted them in the understory
of primary forest on Balambangan Island.
We captured two adult females, one of
which was lactating, in a bamboo thicket at
Site 11 (ca. 60 m asl), and two individuals
in the understory (ca. 2—4 m above the
ground) of primary lowland forest at ca.
650 m elevation at Site 2, all in harp-traps.
Specimens examined: 3, Site 2 (1), Site 11
(2).

Kerivoula_ pellucida.—This poorly
known species is known from the Malay
Peninsula, the Sunda Shelf, Jolo, and Pa-
lawan (Heaney et al. 1998). Known only
from lowland forest (Payne et al. 1985).
Taylor (1934) reported two specimens from
Palawan (no locality given) that he obtained
from a group of seven that he found flying
together in daylight: his map (Fig. 13),

VOLUME 117, NUMBER 3

shows the locality in the vicinity of Broo-
ke’s Point. On 21 June 2000, using a harp-
trap we captured an adult female carrying
a suckling infant in secondary lowland for-
est (ca. 80 m) over a small stream at Site
7. Cranial measurements of the adult female
(Table 2) are smaller than the one specimen
from Davao del Norte, Mindanao available
to Ingle and Heaney (1992), but they are
otherwise very similar; additional speci-
mens are badly needed to examine patterns
of variation. Specimens examined: 2, Site 7
(2).

Kerivoula whiteheadi.—This poorly
known species is widely distributed from
southern Thailand to Borneo and the Phil-
ippines (on Luzon, Mindanao, and Pala-
wan; Heaney et al. 1998). In the Philip-
pines, it is known only from near sea level
in disturbed forest and agricultural areas
(Sanborn 1952). A single specimen in the
UMMZ captured by P. O. Glass on 29 Sept.
1978 at Irawan, Puerto Princesa Municipal-
ity (noted as 2 km N Irawan, at the base of
Mt. Beaufort by P. O. Glass, in litt.) was
erroneously reported by Heaney et al.
(1998) under both K. hardwickii and this
species. Additionally, two specimens taken
“under banana fronds” by C. A. Ross on 8
April 1987 at Barangay Binwang, Quezon
Municipality, are housed in the USNM. We
captured a single individual of this species
in a harp-trap near ground level in a cogon
grassland (Umperata cylindrica) at Site 6.
Cranial measurements of these four individ-
uals (Table 2) are similar to those in Ingle
& Heaney (1992) of a specimen from Min-
danao, though some variation in size is pre-
sent; more specimens are needed to assess
geographic variation. Specimens examined:
4, Site 6 (1), Irawan, Puerto Princesa Mu-
nicipality, 60 m (1), and Binwang, Quezon
(2).

Miniopterus australis.—This common
species is found from India to Australia; it
is widespread in the Philippines, but this is
the first record from Palawan (Heaney et al.
1998). It is known to roost in caves in low-
land areas of agriculture or second growth

287

(Heaney et al. 1991, Rickart et al. 1993,
Sanborn 1952). We captured this species in
varying numbers at several caves. In pri-
mary and disturbed lowland forest at Sites
4 and 5 it was scarce, represented by less
than 1% and less than 3% of captures, re-
spectively. At Site 8 it was abundant (45%
of 2775 captures). Cranial measurements of
A individuals (Table 2) are similar to those
reported by Ingle and Heaney (1992) and
Rickart et al. (1993) from the Philippines,
and by Corbet & Hill (1992) from through-
out the species range. Specimens examined:
4, Site 4 (2), Site 5 (2).

Miniopterus schreibersi.—This common
species is found from Europe to the Solo-
mon Islands and is widespread in the Phil-
ippines, but this is the first record from Pa-
lawan (Heaney et al. 1998). It is common
in caves throughout the lowlands in agri-
cultural areas and forest and known from
both lowland and montane forest (Heaney
et al. 1991, 1999; Rickart et al. 1993; San-
born 1952). We captured this species in and
around caves in disturbed and primary low-
land forest at Sites 4, 5, and 8. At Sites 4
and 8 the species was abundant, represented
by 47% and 26% of captures respectively.
At Site 5 it was common at 10% of 575
captures. Of 214 adult females captured in
December 1999 at Site 4, only one was
pregnant. Of 36 captured between 15 and
20 May 2000 at Site 5, 15 were pregnant
and none were lactating. Of 421 captured
at Site 8 in July 2000, none were pregnant
but 4 were lactating. Cranial measurements
(Table 2) are similar to those reported by
Ingle and Heaney (1992) and Rickart et al.
(1993) from the Philippines, and by Corbet
and Hill (1992) from throughout the species
range. IUCN (2002) lists this species as
Near-Threatened, but its abundance in
heavily disturbed habitat in the Philippines
(Heaney et al. 1998, Rickart et al. 1993)
makes this inappropriate. Specimens ex-
amined: 6, Site 4 (4), Site 5 (1), Site 8 (1).

Miniopterus tristis.—This widespread
species is found from the Philippines to the
Solomon Islands; this is the first record

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

288

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VOLUME 117, NUMBER 3

from Palawan (Heaney et al. 1998). The
species is known to roost in caves and for-
age in disturbed forest (Rickart et al. 1993,
Sanborn 1952). We captured one specimen
in a cave surrounded by old-growth forest
at Site 4 and two in a cave surrounded by
disturbed areas and secondary forest at Site
10. Miniopterus tristis appeared to be con-
sistently less common than the other spe-
cies of Miniopterus. Cranial measurements
of 3 individuals (Table 2) are similar to
those reported by Ingle and Heaney (1992)
and Rickart et al. (1993) from the Philip-
pines, and by Corbet and Hill (1992) from
throughout the species range. Specimens
examined: 3, Site 4 (1), Site 10 (2).
Murina cf. tubinaris.—A single speci-
men of a small tube-nosed bat (genus Mu-
rina) was taken in a lowland grassland-for-
est mosaic at Site 12 on 24 March 1997,
and is housed in the Staatliches Museum fur
Naturkunde in _ Stuttgart, Germany
(#49238). The specimen (Table 2) is very
similar to a series from Tonkin, Vietnam
(FMNH 32203-32204, 46626-46627),
though slightly larger. In our specimen, as
in the Vietnamese series, the upper tooth-
rows converge slightly, the anterior pre-
molars are reduced, and the canines are
short but longer than the premolars, as not-
ed by Koopman and Danforth (1989) and
Corbet and Hill (1992). The length of fore-
arm (33 mm) falls at the center of the range
given by Koopman and Danforth for M.
tubinaris (28—35 mm), and at the high end
given for M. suilla (26-33 mm). Koopman
and Danforth (1989) considered M. florium,
M. suilla, and M. tubinaris to be members
of a species group, and perhaps to be con-
specific, noting that few specimens are
available. Corbet and Hill (1992) re-empha-
sized the uncertainty in current taxonomy,
but took a somewhat different view, refer-
ring specimens from Borneo to M. suilla,
rather than to M. tubinaris. While we agree
entirely on the need for more specimens
and further study, we follow Koopman &
Danforth (1989) on referring Bornean spec-
imens to M. tubinaris, and provisionally re-

289

fer the specimen from Palawan to this same
species. Specimen examined: 1, Site 12 (1).

Myotis horsfieldii.i—This common spe-
cies is distributed from southeastern China
to the Malay Peninsula, Sulawesi, and the
Philippines (Heaney et al. 1998). On Bor-
neo, the species “‘roosts in crevices or bell-
holes in caves, usually not far from large
streams or rivers” (Payne et al. 1985). In
the Philippines, it has been recorded in low-
land forest and agricultural areas, up to 800
m (Heaney et al. 1998). We never encoun-
tered this species; two specimens in the
UMMZ taken by A. C. Alcala in Sitio Mal-
abusog, Tinitian, Roxas Municipality in
1984 were reported by Heaney et al. (1998).

Myotis macrotarsus.—This species is
known from Borneo and the Philippines
(Heaney et al. 1998); it roosts in caves near
sea level and forages in agricultural areas
(Heaney and Utzurrum, unpubl. data). Md.
Nor (1995) caught the species over a dry
river bed and in the understory of primary
lowland forest on Balambangan Island. In
a cave in disturbed lowland forest at Site 8,
this species was represented by <0.5% of
2775 captures. We also observed small
numbers in the cave at PPSRNP. Cranial
measurements (Table 2) of one individual
show it to be slightly larger than those re-
ported by Ingle & Heaney (1992). IUCN
(2002) lists this species as Near-Threatened.
Specimens examined: 1, Site 8 (1).

Myotis rufopictus.—This poorly known
Philippine endemic has been recorded from
primary lowland and montane forest (Hea-
ney et al. 1999, Mudar & Allen 1986). We
never encountered this species; on Palawan,
it is known from a single specimen in
UMMZ reported by Allen (1922). We fol-
low Ingle & Heaney (1992) in regarding
this as one of several distinct species within
the subgenus Chrysopteron, rather than rec-
ognizing only a single species, Myotis for-
mosus, within the subgenus (e.g., Corbet &
Hill 1992). This species is not listed by
IUCN (2002); we recommend listing as
Data Deficient. Measurements in Table 2 of
the specimen reported by Allen (1922) were

290

taken by Heaney. Specimen examined: 1,
Puerto Princesa (1).

Pipistrellus javanicus.—This species is
distributed from Korea to Java and the Phil-
ippines (Heaney et al. 1998). Taxonomic
status is uncertain; P. imbricatus has been
reported from Palawan (e.g., Allen 1922,
Corbet & Hill 1992, Sanborn 1952), but In-
gle and Heaney (1992) were unable to dis-
tinguish more than one species of Pipis-
trellus of this size in the Philippines; de-
tailed study is needed. It is common in pri-
mary montane forest and uncommon in
lowland and mossy forest (Heaney et al.
1999, Ingle 1992, Sanborn 1952). We cap-
tured two individuals from a roost in a hol-
low tree in montane forest (ca. 1300 m) at
Site 3. The opening in the tree appeared to
have formed where a branch had been bro-
ken off the tree and was quite small (ca. 1.5
< 5 cm). Cranial measurements (Table 2)
fall within the range of specimens reported
by Ingle and Heaney (1992), and are slight-
ly smaller than a series from southern Lu-
zon (Heaney et al. 1999). Specimens ex-
amined: 2, Site 3 (2).

Scotophilus kuhliiimTYhis common spe-
cies is widespread from Pakistan to Taiwan
and the Philippines (Heaney et al. 1998); it
is abundant in urban and agricultural areas,
and roosts in buildings and “tents”? made
from modified palm leaves (Heaney et al.
1998; Rickart et al. 1989, 1993). Hollister
(1913) and Taylor (1934) reported it from
Puerto Princesa, Sanborn (1952) reported it
from Brooke’s Point, and we found it to be
abundant in buildings at the Provincial Ag-
riculture Center, Irawan, Puerto Princesa,
and in staff houses of the State Polytechnic
College in Puerto Princesa, and in mixed
urban/agricultural areas (Site 12).

Tylonycteris pachypus.—This tiny bat is
widespread from India to the Philippines
(Heaney et al. 1998). In the Phillipines, it
is known from bamboo stands in agricul-
tural areas (Heaney & Alcala 1986); Hol-
lister (1913) reported a specimen from
Puerto Princesa. We captured several indi-
viduals of this species in a bamboo thicket

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

at Site 11. Very near the capture site we
observed what appeared to be more than a
dozen individuals of this species foraging
over a few remnant trees in a cleared area
with houses that is immediately surrounded
by logged-over and secondary forest. Spec-
imens examined: 5, Site 11 (5).

Tylonycteris robustula.—This species is
also widespread from southern China to the
Lesser Sunda Islands and the Philippines
(including records from Calauit, Luzon, and
Palawan); its habitat is apparently similar to
that of 7. pachypus (Heaney & Alcala 1986,
Heaney et al. 1998). We never encountered
this species.

Family Molossidae—Free-Tailed Bats

This family is generally poorly known in
Southeast Asia, partly because they typi-
cally fly high above the canopy and are
therefore rarely netted. At least one species
(Chaerophon plicata) is widespread in the
region and should be sought on Palawan.

Cheiromeles torquatus.—This poorly
known species is found from Sumatra to
Java, Borneo, and Palawan, but not the rest
of the Philippines (Heaney et al. 1998). It
roosts in large caves and hollow trees and
forages in open areas, over streams, and
above forest canopy on Borneo (Payne et
al. 1985). We never encountered this spe-
cies, which was documented on Palawan by
Sanborn (1952) based on a single specimen.
IUCN (2002) lists this species as Near-
Threatened.

Mops sarasinorum.—This very poorly
known species occurs in Sulawesi and the
Philippines; the Palawan record is based on
a single specimen in the Senckenberg Mu-
seum, Frankfurt (Heaney et al. 1998). It prob-
ably occurs in lowland forest (Heaney et al.
1998). We never encountered this species. It
is listed by IUCN (2002) as Near-Threatened,
but we recommend Data Deficient.

Order Primates
Family Cercopithecidae—Monkeys

Macaca fascicularis.—This common
monkey occurs from Myanmar to Timor

VOLUME 117, NUMBER 3

and the Philippines (Fooden 1995, Heaney
et al. 1998). It is known from agricultural
areas near forest, second growth, secondary
forest, and primary lowland and montane
forest (Heaney et al. 1998, 1999; Rickart et
al. 1993); Sanborn (1952) reported speci-
mens from Iwahig, Puerto Princesa, and
Brooke’s Point. Reis & Garong (2001) re-
ported a specimen from sediments in a
rock-shelter near Tabon Cave, Quezon Mu-
nicipality dated to 11,130 BP. We common-
ly observed this species at all of our sites
(except Site 13), in secondary and primary
forest (including mangrove, swamp forest,
beach forest, and lowland forest) from sea
level to 1000 m; at forest edge near agri-
cultural areas and houses they seem to be
less common and more shy. On Palawan,
the species is under moderate hunting pres-
sure for meat and the local pet trade, but
appeared to have stable populations. In
most areas, it was quite wary of humans,
but in areas such as the PPSRNP (Site 15),
the species did not associate humans with
danger, and had become a regular thief of
picnic baskets. It is listed by IUCN (2002)
as Near-Threatened.

Order Pholidota
Family Manidae—Pangolins

Manis culionensis.—This endemic spe-
cies of the Palawan faunal region, with re-
cords from Palawan and Culion Islands
(Heaney et al. 1998), was formerly included
within Manis javanica (Feiler 1998). It is
known from primary and secondary low-
land forest, possibly localized in distribu-
tion (Allen 1910, Hoogstraal 1951, Sanborn
1952, Taylor 1934). We sighted several in
lowland grassland/forest mosaic at Site 12.
It is hunted for its skin, which is used to
treat asthma. We have seen it for sale in
Puerto Princesa and our guide at Site 11
said that it is hunted in logged-over lowland
forest in that area. The species was de-
scribed by local informants as fairly com-
mon, but hunting pressure is moderately
heavy. Manis javanica is listed by IUCN

291

(2002) as Near-Threatened; M. culionensis
probably deserves the same status.

Order Rodentia
Family Sciuridae—Squirrels

Hylopetes nigripes.—This large gliding
squirrel is endemic to the Palawan faunal
region; the number of museum specimens
(Allen 1910, Sanborn 1952) suggests that it
is common. Reis & Garong (2001) reported
two specimens from sediments in a rock-
shelter near Tabon Cave, Quezon Munici-
pality dated to 11,130 BP. Taylor (1934)
found the species in primary and secondary
lowland forest where they nest in cavities
in large trees. We observed an individual
running up the side of a large hollow tree
in primary forest at Site 1, and we frequent-
ly heard and twice spotlighted them in ma-
ture lowland forest at Site 15. We also
heard the distinctive calls several times in
selectively logged but largely intact forest
near Barake, Aborlan Municipality, in the
Victoria Range. According to local resi-
dents, the species is common in mature for-
est and is occasionally hunted as a source
of food. IUCN (2002) lists this species as
Near-Threatened; by current criteria, it
should be listed as Data Deficient.

Sundasciucus juvencus.—This tree squir-
rel is endemic to central and northern Pa-
lawan Island (Heaney et al. 1998). Hoogs-
traal (1951) and Sanborn (1952) reported
this species from primary and secondary
lowland forest. We commonly observed this
species in primary and secondary lowland
forest at Sites 1, 2, 7, 11, 12, and in both
secondary forest and grassland/degraded
forest mosaic at Site 15. We also found it
in a very small (<1 ha.) patch of secondary
lowland forest surrounded by grassland and
agricultural areas at Site 6. We observed the
species on Dumaran Island, but not on Mal-
inau or Rasa. The species is reportedly a
common pest in coconut plantations. It is
occasionally hunted as a source of food and
for the local pet trade. It is listed by IUCN
(2002) as Endangered, but this is strongly

292

contradicted by the available data, and we
recommend de-listing.

Sundasciurus rabori.—This_ poorly-
known species, described from 5 specimens
taken at 3600—4350 ft (ca. 1100—1300 m)
on Mt. Mantalingajan, is endemic to Pala-
wan Island (Heaney 1979). P. C. Gonzales
deposited 2 specimens at the UMMZ that
he collected on Mt. Gorangbato in Brooke’s
Point Municipality in 1984; these are the
only reported specimens aside from the
original type series from Mt. Mantalingajan
(Heaney 1979). Although we worked in
some seemingly suitable habitats on Cleo-
patra’s Needle (Site 3), we did not specifi-
cally seek this species, and we never en-
countered it. The IUCN (2002) lists S. ra-
bori as Vulnerable, but based on current
IUCN criteria, it should be considered Data
Deficient.

Sundasciunis steerii.—This species is en-
demic to Balabac and southern Palawan Is-
land (Heaney et al. 1998); Sanborn (1952)
reported a large series from Brooke’s Point.
Heaney et al. (1998) listed it as common in
lowland forest and coconut and banana
plantations. Because all of our study sites
were in central and northern Palawan, we
never encountered this species. It is listed
by IUCN (2002) as Near-Threatened; since
its habitat use is similar to the closely-re-
lated S. juvencus, it is probably not threat-
ened.

Family Muridae—Mice

Chiropodomys calamianensis.—This
poorly known arboreal mouse is endemic to
the Palawan faunal region; it is closely re-
lated to species on the Sunda Shelf (Musser
1979). Reis & Garong (2001) reported a
specimen from sediments in a rock-shelter
near Tabon Cave, Quezon Municipality dat-
ed to 11,130 BP. It is known from forest
near sea level (Taylor 1934), coconut plan-
tations, bamboo thickets, and buildings
(Sanborn 1952); there are 12 specimens
from Palawan in FMNH and NMP from the
Hoogstraal expedition (Sanborn 1952). The

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

genus is apparently difficult to capture
(Musser 1979); we never encountered this
species. We recommend IUCN listing as
Data Deficient.

Haeromys pusillus.—This species is
known only from Borneo, Palawan, and Ca-
lauit Islands (Musser & Carleton 1993,
Musser & Newcomb 1983). It is cited in
Heaney et al. (1998) as ““Haeromys sp. A’
as potentially endemic to Palawan, but we
follow Musser (pers. comm.) in treating it
as conspecific with H. pusillus. We never
encountered this species, but Musser &
Carleton (1993) cited a specimen from Pa-
lawan. A specimen of H. pusillus was taken
in Sabah, Borneo, in a pit-fall trap near the
edge of tall dipterocarp forest (Payne et al.
1985), and A. C. Alcala stated that he cap-
tured the specimen from Calauit (in
FMNH) by hand in a bamboo thicket (pers.
comm.). IUCN (2002) listed this species as
Vulnerable, but based on current criteria, it
should be considered Data Deficient.

Maxomys panglima.—This common rat
is endemic to the Palawan faunal region;
the genus is common on the Sunda Shelf,
but is absent from oceanic portions of the
Philippines (Musser et al. 1979). Sanborn
(1952) reported large series from several lo-
calities. We found it to be the most com-
monly captured small mammal in agricul-
tural/forest mosaic at Site 12 (62% of 169
captures), and was common to abundant in
secondary forest (Site 11), primary lowland
(Sites 1 and 2), and montane forest (Site 3)
from near sea level to at least 1550 m. We
captured a single juvenile in mossy forest
at 1580 m at Site 3. Because we found this
species to be common, although sometimes
patchy, in all lowland and montane forested
sites where we trapped extensively, and in
mixed agricultural/second growth areas at
Sites 11 and 12, we consider the IUCN
(2002) listing as Near-Threatened to be un-
justified. Specimens examined: 5, Site 1 (3),
Site 3 (2).

Mus musculus.—This introduced com-
mensal has a nearly world-wide distribu-
tion, although Southeast Asian populations

VOLUME 117, NUMBER 3

are sometimes treated as a separate species,
M. castaneus (Musser & Carleton 1993). It
is common in human habitations in urban
and rural areas (Heaney et al. 1998). We
captured several in a residential area at Site
12, and it is most likely common in such
places throughout Palawan.

Palawanomys furvus.—This_ poorly
known monotypic genus is endemic to Pa-
lawan Island. It has been taken from a sin-
gle locality on Mt. Mantalingajan and prob-
ably occurs in high mountain forest (Mus-
ser & Newcomb 1983). Our survey efforts
on Cleopatra’s Needle (Site 3) failed to find
this species; perhaps it is restricted to the
more extensive mountain ranges of south-
ern Palawan. The IUCN (2002) lists this
species as Endangered; the lack of data and
lack of damage to its presumed habitat
(montane and mossy forest) suggest that it
should be listed as Data Deficient.

Rattus exulans.—This introduced com-
mensal species is widespread from Bang-
ladesh to Easter Island (Heaney et al. 1998).
The first records from Palawan were named
as a distinct species (luteiventris) by Allen
(1910), but it is currently treated as a junior
synonym of R. exulans (Musser & Carleton
1993). It is common in agricultural areas
(Barbehenn et al. 1973, Rabor 1986) and
sometimes present in disturbed forest and
rare in primary forest (Barbehenn et al.
1973; Heaney et al. 1991, 1998). We found
this species in grassland (Site 6), agricul-
tural areas (Sites 6, 11, 12, and 14), and in
secondary lowland forest (Site 7). The spe-
cies appears to be absent from primary
(e.g., Sites 1, 2, and 3) and logged-over for-
est (e.g., Site 11) on Palawan. Specimens
examined: 4, Site 6 (3), Site 11 (1).

Rattus tanezumi.—This introduced com-
mensal, formerly included within Rattus
rattus (Musser & Carleton 1993), is wide-
spread from Afghanistan to New Guinea
and Micronesia (Heaney et al. 1998). It is
often abundant in urban and agricultural ar-
eas and common in disturbed forest up to
1800 m (Danielsen et al. 1994; Heaney et
al. 1989, 1999; Rabor 1986; Sanborn 1952).

293

Hoogstraal (1951) and Sanborn (1952)
found them to be common on Palawan in
some agricultural and residential areas. We
captured three at Site 13, and three in a res-
idential area in Puerto Princesa; vouchers
were deposited in the NMP and the collec-
tion of the Palawan Council for Sustainable
Development.

Rattus tiomanicus.—This indigenous rat
is found on the Malay Peninsula and the
islands of the Sunda Shelf, including Pala-
wan (Heaney et al. 1998). Payne et al.
(1985) reported the species from secondary
forest, agricultural areas and gardens, scrub,
and grassland. We captured this species in
grassland/forest mosaic at Site 12 (9% of
captures), two in selectively logged forest
at Site 13, one in a ricefield at Site 14, and
three individuals from mossy forest and the
transition zone between mossy and montane
forest at Site 3. At Site 3, two individuals
were taken at ca. 1580 m during the night
and one at ca. 1540 m during the day. Spec-
imens examined: 3, Site 3 (3).

Sundamys muelleri.—This moderately
large rat is found from southern Myanmar
to the Sunda Shelf, including Palawan
(Heaney et al. 1998); the genus is absent
from the oceanic Philippines. Sanborn
(1952) described a subspecies endemic to
Palawan, S. m. balabagensis, from a single
specimen taken at 3000 ft (ca. 900 m) “‘in
thick forest near the top of Mt. Balabag”’;
two additional specimens in the USNM are
from Pinigisan, on the lower slopes of Mt.
Mantalingajan at 2100—2500 ft (ca. 640—
760 m). Additional specimens from the Pa-
lawan region are from Culion Island (San-
born 1952), Balabac, and Busuanga
(USNM; Heaney et al. 1998). On Borneo,
the species occurs in forest, often near
streams (Payne et al. 1985) at elevations
usually below 3500 ft (ca. 1070 m; Med-
way 1977). Md. Nor (1995) caught the spe-
cies on Banggi, Balambangan, and Mol-
leangan Islands “‘mostly in primary forest
on low ground and near streams’’. We cap-
tured one individual from a riparian zone in
primary forest at ca. 700 m at Site 2, and

294

two in lowland grassland/forest mosaic at
Site 12. Specimen examined: 1, Site 2 (1).

Family Hystricidae—Porcupines

Hystrix pumila.—The only porcupine
found in the Philippines is endemic to the
Palawan faunal region; other species occur
widely on the Sunda Shelf and continental
Asia. Reis & Garong (2001) reported 3
specimens from sediments in a rock-shelter
near Tabon Cave, Quezon Municipality dat-
ed to 11,130 BP. It is known to occur in
secondary and primary lowland forest and
to den in abandoned mine shafts (Hoogs-
traal 1951, Sanborn 1952). We observed
this species at dusk along the edge of sec-
ondary forest at Site 11, once at night at
Site 15 (where it was feeding on fruit of
Terminalia catappa), and several times at
night in grassland/forest mosaic at Site 12.
Local guides reported them at Site 14, at
the Iwahig Penal Colony in Puerto Princesa,
in Rizal Municipality, and in Dumaran Mu-
nicipality (on the mainland). This was re-
ported as the most important game species
for the Tagbanua ethnic community in Bar-
ake, Aborlan Municipality (Lacema & Wid-
mann 1999); they are often dug out of their
subterranean dens. Not listed by IUCN
(2002) but listing as Data Deficient or Near-
Threatened seems justified.

Order Carnivora
Family Felidae—Cats

Prionailurus bengalensis.—This small
cat is widespread from Siberia to Pakistan
and Bali, with reports from the Philippines
on Busuanga, Cebu, Negros, Palawan, and
Panay Islands only (Heaney et al. 1998,
Taylor 1934). The population from the Pa-
lawan faunal region was described recently
as a distinct subspecies, P. b. heaneyi, by
Groves (1997); it is well represented by
museum specimens (Allen 1910, Sanborn
1952). Rabor (1986) reported the species
from agricultural areas and forest from sea
level to ca. 1500 m. We spotlighted one

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

along a river trail in Barake, Aborlan Mu-
nicipality.

Family Mustelidae—Weasels,
Otters, and Badgers

Amblonyx cinereus.—This otter occurs
from India to Taiwan and the Sunda Shelf
(Heaney et al. 1998). On Palawan, it is
found in coastal rivers and bays (Hoogstraal
1951, Rabor 1986, Sanborn 1952). Payne et
al. (1985) and Sanborn (1952) reported the
species feeds on crustaceans, mollusks, and
fish where there is permanent water and
some tree cover. Rangers at PPSRNP re-
ported to Heaney and Widmann that otters
frequently visit along the beach and small
streams, and local people reported them
from the Iwahig River (Puerto Princesa),
Aborlan River (Aborlan Municipality),
Malatgao and Taritien Rivers (Narra Mu-
nicipality), and adjacent mangrove and
freshwater swamp forest. We received one
report of an otter raiding a prawn pond.
IUCN (2002) lists this species as Near-
Threatened.

Mydaus marchei.—This badger is en-
demic to the Palawan faunal region; it is
related to a species that occurs on the Sunda
Shelf. It has been documented in mixed
grassland and secondary forest (Hoogstraal
1951, Kruuk 2000, Rabor 1986, Taylor
1934), and Sanborn (1952) reported series
from several localities. We occasionally
smelled its strong odor in areas of mixed
agriculture and secondary forest throughout
Palawan; we sighted it often in residential
and cultivated areas, grassland, and grass-
land/forest mosaic at Site 12, and rarely in
ricefields and freshwater swamp forest at
Site 14. One individual living in a den on
campus at the State Polytechnic College
was easily followed and observed. Because
it is widespread and moderately common
on Palawan, and is rarely hunted (Grim-
wood 1976, Kruuk 2000), we agree with
Kruuk (2000) that the IUCN listing of this
species as Vulnerable is not justified.

VOLUME 117, NUMBER 3

Family Herpestidae—Mongooses

Herpestes brachyurus.—The only mon-
goose found in the Philippines is distributed
from the Malaysian Peninsula to Borneo
and Palawan (Heaney et al. 1998). The Pa-
lawan population was named as a distinct
species (H. palawanus Allen 1910), but
currently is treated as a subspecies (Corbet
& Hill 1992). Allen (1910) described them
based on one specimen from Iwahig; San-
born (1952) reported one specimen from
Puerto Princesa and one from Brooke’s
Point, and Rabor (1986) found the species
most often near rivers. On Borneo, Payne
et al. (1985) found the species to occur in
primary and secondary lowland forest,
plantations, and gardens. We never encoun-
tered this species; but we received reports
of them at Site 14.

Family Viverridae—Civets

Arctictis binturong.—The binturong is
known from northern Myanmar to the Sun-
da Shelf (Heaney et al. 1998). On Borneo,
the species is arboreal and terrestrial, most-
ly nocturnal, and occurs in old-growth and
secondary forests, sometimes entering ag-
ricultural areas near forest (Payne et al.
1985). The Palawan population, which ini-
tially was named as a distinct species (A.
whitei Allen 1910) from four specimens, is
still represented by few specimens (Heaney
et al. 1998). Rabor (1986) reported obser-
vations from primary and secondary low-
land forest up to 200 m. Our guide at Site
11 reported that a juvenile repeatedly en-
tered remnant trees that were fruiting in a
clearing surrounded by secondary forest. At
Site 2, we observed A. binturong drinking
water from a stream at ca. 400 m during
mid-day. We spotlighted one in a fruiting
Ficus tree at Site 15, and twice saw one in
grassland/forest mosaic at Site 12 feeding
on fruits of Guioa pleuropteris. Local peo-
ple reported hunting them for food, and also
catching them and selling them as pets. The
IUCN (2002) listing of A. binturong whitei
as Vulnerable seems justified.

295

Paradoxurus hermaphroditus.—This
common species is found from Sri Lanka
to the Lesser Sunda Islands and the Phil-
ippines (Heaney et al. 1998). Recorded in
agricultural areas and forest over a wide
elevational range (Allen 1910; Heaney et al.
1991, 1999; Hoogstraal 1951; Rabor 1986);
Sanborn (1952) reported large series from
several localities. We often saw them feed-
ing in fruiting trees and shrubs in grassland/
forest mosaic at Site 12, and we saw road-
kills along the coastal highway. They are
hunted, but the large number of museum
specimens and sightings indicate that they
traditionally have been and probably remain
the most common carnivore on Palawan
(e.g., Allen 1910, Sanborn 1952).

Viverra tangalunga.—tThis civet is found
from the Malay Peninsula to Sulawesi and
the Philippines (Heaney et al. 1998).
Known from primary and secondary low-
land, montane, and mossy forest (Allen
1910, Heaney et al. 1999, Rickart et al.
1993). We captured and released a juvenile
of this species in a cage trap in lowland
primary forest at Site 1, and we observed
two in forest-grassland mosaic at Site 12.

Order Artiodactyla

Tragulus napu and Axis calamianensis
both occur in the Palawan faunal region,
but we found no evidence of either species
on Palawan Island.

Family Suidae—Pigs

Sus barbatus.—The bearded pig is found
from the Malay Peninsula to Borneo and
Palawan (Heaney et al. 1998). Rabor (1986)
and Payne et al. (1985) reported the species
from primary and secondary forest from sea
level to the highest peaks; Sanborn (1952)
reported a series from Iwahig. Groves
(2001) has tentatively suggested that the
population of this species from the Palawan
faunal region, which has been recognized
as a distinct subspecies, may warrant rec-
ognition as a distinct species, Sus ahoeno-
barbus. We regularly observed this species

296

or evidence of its occurrence in forest hab-
itats (including fragmented forest) from sea
level to montane forest at ca. 1500 m (Sites
1, 2, 3, 4, 7, and 11). We also observed
evidence of the species entering cultivated
areas near forest and damaging crops. Wild
pigs are heavily hunted on Palawan with
snares, low caliber rifles, and small, baited
explosive devices known as “pig bombs”.
The species appears to be locally common,
but is in decline due to heavy hunting pres-
sure (Caldecott et al. 1993, Oliver 1992).
The IUCN (2002) lists S. barbatus ahoe-
nobarbus as Vulnerable.

Discussion

Adequacy of sampling.—Small fruit bats
on Palawan (Cynopterus, Eonycteris, Ma-
croglossus, and Rousettus) appear to have
been fairly completely sampled; no species
have been added in over 50 years (exclud-
ing the apparently erroneous reports of
Haplonycteris fischeri and Ptenochirus mi-
nor), despite extensive netting. It is inter-
esting that our mist netting in primary for-
est produced very few captures; for exam-
ple, in primary lowland forest at 150 m el-
evation (Site 1), we captured | fruit bat in
42 net-nights; in primary lowland forest at
ca. 500 m (Site 2), we captured 1 fruit bat
in 56 net-nights, and in primary montane
forest at ca. 1400 m (Site 3), we captured
no fruit bats in 48 net-nights. Although our
sample size is limited, all of these values
fall well below what would be typical on
islands in the oceanic portion of the Phil-
ippines (Heaney et al. 1989, 1999), sug-
gesting that small fruit bats are not as abun-
dant in primary forest on Palawan (e.g.,
Heideman & Heaney 1989, Heaney et al.
1989). Indeed, all of the small fruit bats
currently known from Palawan predomi-
nately occur in disturbed habitats, in con-
trast to the oceanic Philippines, where sev-
eral endemic genera (Alionycteris, Haplon-
ycteris, Otopteropus, and Ptenochirus) are
most common in old-growth forest.

The ecology of large fruit bats (Acerodon

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

and Pteropus) has been very poorly studied
on Palawan and elsewhere in the Philip-
pines. This is the direct result of the diffi-
culty in capturing these species by any
means other than shooting. Despite the pau-
city of ecological information on these spe-
cies, their distribution among major island
groups appears to be moderately well un-
derstood because their activities and roosts
are highly conspicuous, and no additional
species have been found on Palawan in over
50 years. It seems unlikely that additional
species will be found on Palawan, with the
possible exception of P. hypomelanus.

Insectivorous bats are clearly the least
known of all Palawan mammals. Distribu-
tions remain poorly documented, ecological
information is scanty for most species, and
the taxonomy is often uncertain. Our survey
efforts and examination of previously col-
lected insectivorous bats documented eight
species (Rhinolophus arcuatus, R. ct. bor-
neensis, R. creaghi, R. macrotis, Miniopte-
rus australis, M. schreibersi, M. tristis, and
Murina cf. tubinaris) on Palawan for the
first time, and several more are noted in the
text as very likely to be present.

Our knowledge of small non-volant
mammals (including Soricidae, Tupaiidae,
Sciuridae, and Muridae) on Palawan is un-
even. Some lowland species (e.g., Tupaia
palawanensis, Sundasciurus juvencus,
Maxomys panglima, Rattus tiomanicus,
Sundamys muelleri, and the non-native mu-
rines) are common and well known. Very
little is known of several other species (e.g.,
Crocidura palawanensis, Crocidura sp.,
Sundasciurus rabori, Chiropodomys cala-
mianensis, Haeromys pusillus, and Pala-
wanomys furvus). Perhaps we failed to lo-
cate these poorly known species because
we did little trapping in trees or other places
above the ground surface (C. calamianensis
and H. pusillus), sampled only one site
above 1000 m (S. rabori and P. furvus), and
our trapping techniques were limited, i.e.,
we did not use pitfall traps (Crocidura
spp.). The presence of so many poorly
known species suggests that other species

VOLUME 117, NUMBER 3

may await discovery, especially in high
mountain habitats and high in the canopy.
Musser & Newcomb (1983) suggested that
unknown species are yet to be discovered
on Palawan, citing the report of Hoogstraal
(1951), which narrated their attempt to cap-
ture a “very large rat with a white tail”
described to them by native Palaw’an. Oth-
er large islands in the Philippines have been
shown to support diverse communities of
endemic small mammals which are restrict-
ed to montane areas (e.g., Heaney 2001,
Heaney & Rickart 1990, Rickart 1993), and
perhaps the same awaits discovery on Pa-
lawan.

Medium to large mammals (Cercopithe-
cidae, Manidae, Hystricidae, the carnivores,
and Suidae) are possibly the most thor-
oughly inventoried subset of Palawan’s
mammalian fauna. Because of their large
size, they are easily observed compared to
other mammals. Many of these species are
also commonly hunted, so obtaining speci-
mens is often easier than for non-game spe-
cies. It is unlikely that other medium to
large mammals await discovery on Pala-
wan, though the ecology of all requires
much additional study.

Biogeography.—As noted above, the Pa-
lawan faunal region is part of the Sunda
Shelf and may have been connected to
mainland Asia via Borneo (Everett 1889)
during a Pleistocene episode of glacially-
induced sea-level lowering (Heaney 1985,
1986, 1991a), though current data leave this
uncertain. All other islands in the Philip-
pines are oceanic and have probably never
had a dry-land connection to any mainland
area (Heaney 1985, 1986, 2001). Of the 58
native species currently known from Pala-
wan Island (tentatively including the small
Crocidura sp.), 13 species arc endemic to
Palawan (and usually to some of the smaller
islands that were included in Pleistocene
Greater Palawan; Heaney 1986); 12 of
these are non-volant, all of which have their
closest relatives on the Sunda Shelf. Eight
of Palawan’s 11 native rodents (73%) are
endemic; all three non-endemics are mu-

297

rids. Only one endemic species is a bat (Ac-
erodon leucotis), and only it has its closest
relatives in the oceanic Philippines. This
pattern of endemism is clearly consistent
with the geological history of the Philip-
pines and also highlights the importance of
the greater vagility of bats over non-flying
mammals. Of the 28 insectivorous bats, 18
species are somewhat to highly widespread
in Indo-Australia (and some beyond), 2 are
shared only with the Sunda Shelf and In-
dochina (Rhinolophus acuminatus and
Rhinolophus cf. borneensis), 1 with the
Sunda Shelf only (Cheiromeles torquatus),
3 occur on the Sunda Shelf and the oceanic
Philippines (Kerivoula pellucida, K. white-
headi, Myotis macrotarsus), 1 occurs on
Palawan, Sulawesi and the oceanic Philip-
pines (Mops sarasinorum), 2 occur only on
Palawan and in the oceanic Philippines
(Rhinolophus virgo and Myotis rufopictus),
and one occurs on Borneo, Sulawesi, and
throughout the Philippines (Emballonura
alecto). These data again demonstrate that
the bats are more widely distributed and do
not clearly reflect the geological history, as
do the non-flying mammal species, which
in the Philippines are usually restricted to a
single area that was united by dry land dur-
ing the late Pleistocene. From the highly
diverse fruit bat fauna of Borneo (17 spe-
cies; Payne et al. 1985), only five species
extend to the northern continental land-
bridge islands of Sabah (Md. Nor 1995).
These are the same five non-endemic spe-
cies that can be found just to the north on
Palawan Island.

We note that the combined totals of na-
tive non-volant mammal species on Pala-
wan that either are shared with Borneo and
other portions of the Sunda Shelf or that are
endemic to Palawan and have their closest
relatives on Borneo or adjacent areas is 22
out of 24 (92%). The apparent exceptions
are Hylopetes nigripes (related to H. albon-
iger of Indochina) and Palawanomys furvus
(an enigmatic genus of unclear phylogenet-
ic position). If this analysis were extended
to the entire Palawan faunal region, Axis

298

calamianensis (related to A. porcinus in In-
dochina) would also be included here. Four
species are widespread in Southeast Asia
(including parts of Wallacea), and no spe-
cies is shared with the oceanic Philippines
except for those 4 species (Macaca fasci-
cularis, Prionailurus bengalensis, Paradox-
urus hermaphroditus, and Viverra tanga-
lunga). If Sus barbatus, which occurs on
Palawan, the Sunda Shelf, and in the mar-
ginal islands of the Sulu Archipelago is
counted as a widespread Southeast Asian
species, the total rises to five species. These
data, in sum, strongly reinforce the conclu-
sion of Everett (1889), based on his anal-
ysis of bathymetric features of the ocean
floor and the pattern of relationships of the
18 species then known from Palawan and
associated smaller islands, that the Palawan
faunal region is an extension of the Sunda
Shelf, probably due to a fairly recent dry-
land connection, with only a small portion
of its fauna shared with the oceanic Phil-
ippines.

Patterns of species richness relative to is-
land area are also of interest. Insectivorous
bats are so poorly known in the Philippines
that it is possible only to say that the 28
species documented here compare favor-
ably with most islands in the Philippines;
that is, there is no evidence that Palawan is
species-poor (Heaney et al. 2002). Fruit
bats, on the other hand, are represented on
Palawan by only 6 species, and this places
their diversity far below that on much
smaller islands in the oceanic Philippines;
Maripipi, for example, has 10 species, but
is only 22 km/?. It seems certain that Pala-
wan has a depauperate fruit bat fauna, as
well as probably having lower fruit bat den-
sity, as noted above, compared to the oce-
anic Philippines (Heaney 1991b). Species
richness of non-flying mammals, on the
other hand, at 24 is much above the species/
area curve documented in the oceanic Phil-
ippines, though well below that of islands
on the primary portion of the Sunda Shelf
(Heaney 1984, 1986; Heaney et al. 2002).
It is interesting that carnivores are notably

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

more diverse on Palawan than in the oce-
anic Philippines, and murid rodents notably
less diverse, for an island the size of Pala-
wan.

Conservation issues.—The most pressing
issue facing terrestrial wildlife in Palawan
is the rapid loss of forest cover, especially
in the primary lowland forests that are tar-
geted for logging. Palawan’s forests are less
commercially valuable than the diptero-
carp-dominated forests of the other islands,
and, consequently, deforestation occurred
later on Palawan. However, once forests
were exhausted on Luzon, Mindanao, Ne-
gros, etc., commercial logging operations
began working in lowland forest on Pala-
wan (Environmental Science for Social
Change 1999) during the 1970’s and 1980’s
at unsustainable levels (Quinnell & Balm-
ford 1988, Kummer 1992). Since a logging
ban was imposed in the early 1990s
throughout the Province of Palawan, log-
ging has declined, but the large commercial
operations appear to have been replaced by
small-scale, illegal commercial logging. We
have seen that forest continues to disappear
from the most accessible areas, and forest
edges are gradually creeping higher and
higher up the contours, in a manner similar
to that experienced on Leyte (Rickart et al.
1993) and southern Luzon (Heaney et al.
1999). Lowland primary forest has been
eliminated from many parts of Palawan and
the destruction shows few signs of easing.
Due to almost complete conversion of the
coastal plain into ricefields, coconut, or oth-
er plantations, distinctive ecosystems such
as freshwater swamp forest and beach forest
have virtually disappeared (Widmann
1998). Slash and burn agricultural practices
have also been very damaging to forests,
and Palawan has experienced a population
explosion due to high birth and immigration
rates.

It should be noted that caves are crucial
to maintaining the fauna, since approxi-
mately 18 (32%) of Palawan’s mammals are
bats that roost in caves. Caves have been
the focus of much destruction in the Phil-

VOLUME 117, NUMBER 3

ippines; common activities in caves on Pa-
lawan include guano mining, general van-
dalism, recreational exploration, and trea-
sure hunting. Unlike many other parts of
the Philippines (e.g., Heaney et al. 1991,
1999: Rickart et al. 1993), we never en-
countered any evidence of cave-roosting
bats being hunted on Palawan. We found
guano mining to be common, usually near
the mouth of caves. Recreational explora-
tion of caves is steadily increasing; many
caves are currently being developed or ad-
vertised for this purpose, while many more
proposals are in the planning stages. At
PPSRNP (Site 15), hundreds of visitors
may enter a one kilometer stretch of the
cave daily. The bats are clearly disturbed
by the activity, but the ultimate result of
such disturbance is unknown.

Many medium to large sized mammals
are under significant hunting pressure on
Palawan for their meat, the live animal
trade, and medicinal use, as noted above,
but few data are available on the impact.
The recent and on-going shift from subsis-
tence to market economies among members
of the Tagbanua and other ethnic groups
may contribute to the decline of some spe-
cies (Lacerna & Widmann 1999), such as
Sus barbatus, Pteropus vampyrus, and Hys-
trix pumila for meat, Macaca fascicularis
and Arctictis binturong as pets, and Manis
culionensis for traditional Chinese medi-
cine.

Acknowledgments

Funding for these studies was provided
by the Palawan Council for Sustainable De-
velopment, United States Peace Corps,
Philippine Cockatoo Conservation Program
through the Loro Parque Fundacion, Ten-
eriffe, Spain, and the Barbara Brown and
Ellen Thorne Smith Funds of the Field Mu-
seum. Jun Saldajeno, Apollo Regalo, Ben-
igno Maca, Erlito Porka, and Manual Lar-
dizabal made significant contributions to
field work, often under difficult living con-
ditions. Adelwisa Sandalo, Lualhati Tabu-

299

gon, Ariel Carino, Leilani Berino, Dr. Ter-
esita Salva, Dr. Edgardo Castillo, Dr. Pa-
cencia Milan, Dr. Josef Margraf, Indira Lac-
erna-Widmann, Siegfred Diaz, Deborah
Villafuerte, and the wildlife wardens of
Rasa Island provided valuable logistical and
administrative support. JAE thanks the Ta-
bugon family for their extraordinary hos-
pitality during his stay on Palawan. We
thank A. C. Alcala, P C. Gonzales, and P.
O. Glass for depositing important speci-
mens at UMMZ, and P. Myers for access to
those specimens. H. Kafka, J. Mead, and R.
Thorington provided access to specimens at
the USNM, and E Dieterlen kindly loaned
specimens from SMNH. We are grateful to
P. O. Glass for additional details about spec-
imens he collected. Ding Padilla, Clara
Simpson, and Lisa Kanellos produced the
map in Fig. 1. Eric Rickart gave sound ad-
vice during the early stages of the project
and Danilo Balete helped with the transport
and identification of voucher specimens;
Eric Rickart and Robert Timm provided
constructive reviews of the manuscript. The
Department of Environmental and Natural
Resources, through the Protected Areas and
Wildlife Bureau and the Provincial Envi-
ronment and Natural Resources Office, pro-
vided permits and encouragement. LRH
thanks the Geological Sciences Department,
Northwestern University, for use of office
space during a sabbatical leave.

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(3):303-310. 2004.

A new species of Tropidonophis (Serpentes: Colubridae: Natricinae)
from the D’Entrecasteaux Islands, Papua New Guinea

Fred Kraus and Allen Allison

Bernice P. Bishop Museum, Honolulu, Hawaii 96817, U.S.A., e-mail: (FK) fkraus@hawaii.edu,
(AA) allison@hawaii.edu

Abstract.—We describe a new species of natricine snake of the genus T7ro-
pidonophis from the D’Entrecasteaux Islands, off the southeastern peninsula of
New Guinea. The new species is large, with 15 unreduced scale rows, a high
ventral and low subcaudal scale count, and a distinctive color pattern of dark
mid-dorsal bands and offset lateral blotches on a yellow or brown ground color.
The species is known from two specimens collected at 900—1090 m in primary
lowland hill forest. Close relationships with other members of the genus are

not apparent.

The natricine genus Tropidonophis con-
sists of 18 species of small to medium-sized
snakes distributed from the Philippines (two
endemic species) to the Bismarck Archi-
pelago (two endemic species), with 12 spe-
cies found on New Guinea and its offshore
islands, four in the Moluccas, and one in
Australia (Malnate and Underwood 1988).
The genus is thought to be most closely re-
lated to the Southeast Asian Xenochrophis
on the basis of shared scalation, hemipenial,
and osteological features (Malnate and Un-
derwood 1988). Most Tropidonophis spe-
cies are nondescript, frequently with a uni-
formly dark dorsal ground color of brown
or gray, sometimes with darker spots, short
lines, or narrow bands. Some specimens of
a few species have more conspicuous pat-
terns of dark bands on a lighter ground col-
or (cf. O'Shea 1996: 95). Tropidonophis
species typically inhabit rainforest, occur
from sea level to 2200 m (Malnate and Un-
derwood 1988), and are reported to dwell
frequently near permanent water sources
(O’Shea 1996).

During the course of conducting biolog-
ical surveys in the D’Entrecasteaux Islands
in 2002 we collected a strikingly colored
specimen of Tropidonophis that is unas-
signable to any currently recognized spe-

cies. A search of museum collections re-
vealed another specimen belonging to the
same taxon. We take this opportunity to
provide this species with a name.

Materials and Methods

Specimens were collected under appli-
cable national and provincial permits, fixed
in 10% buffered formalin, and transferred
to 70% ethanol for storage. Measurements
were made to the nearest mm in the field
with a fiberglass tape; mass was measured
to the nearest gram in the field with a Pe-
sola scale. Diagnostic features and compar-
isons to other species were based on data
provided in the comprehensive study of
Tropidonophis by Malnate and Underwood
(1988) and by reference to specimens
housed in the Bernice P. Bishop Museum,
Honolulu (BPBM).

Specimens are deposited in the BPBM
and American Museum of Natural History
(AMNH).

Tropidonophis dolasii, new species
Figs. 1, 2

Holotype.—BPBM 16539 (field tag FK
6118), adult female, collected by D. Sale-
puna on E slope of Oya Tabu (Mt. Kilker-

304

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1.
bar equals one cm.

ran), 9.4555°S, 150.7857°E, 1090 m, Fer-
gusson Island, Milne Bay Province, Papua
New Guinea, on 23 August 2002.
Paratype.—AMNH 73979, adult female,
collected by L. Brass on E slope of Good-
enough Island, 900 m, Milne Bay Province,
Papua New Guinea, on 27 October 1953.
Diagnosis.—A large species of Tropi-
donophis with 15 dorsal scale rows at mid-
body and one head length anterior to the
vent, 161—162 ventrals, 63 subcaudals, 2
preoculars, 3 or 4 postoculars, 8 supralabi-
als, 8 or 9 infralabials, no postocular dark
stripe, and yellow or brown ground color

(A) Lateral, and (B) dorsal view of head of holotype (BPBM 16539) of Tropidonophis dolasti. Scale

with vaguely defined mid-dorsal black
bands offset by lateral black blotches on
scale rows 1—4. These dark bands and
blotches are not solid, rather they are
formed by a network of darkened scale
margins.

Description of holotype.—Adult female.
Dorsal scale rows 15 (reduction to 15 rows
occurs at the level of the 15" ventral); all
rows except first keeled; first row weakly
keeled on those scales posterior to approx-
imately 15 ventrals anterior to vent; keels
on dorsal scales more weakly developed an-
teriorly and laterally and more strongly de-

VOLUME 117, NUMBER 3

veloped posteriorly and dorsally; paired
apical pits obvious on those dorsal scales
retaining the horny epidermal layer; all dor-
sal scales, except on first row, notched at
the posterior tip. Rostral twice as wide as
high; internasals longer than wide; prefron-
tals wider than long, as are frontal, supra-
oculars, and parietals; lateral extension of
parietal contacts middle postocular, exclud-
ing upper postocular from contact with an-
terior temporals on each side. Nasals divid-
ed by large nares; loreal higher than long;
preoculars 2; postoculars 3 (right) and 4
(left); anterior temporals 2, upper a narrow
sliver approximately 20% the size of lower,
lower excluded from contact with posto-
culars on right side, with point contact to
second and third postoculars on left; pos-
terior temporals 3, the most anterior lying
on posterior slope of lower anterior tem-
poral and in contact posteriorly with only
the middle posterior temporal (Fig. 1). Su-
pralabials 8, 4 and 5 contact eye; infra-
labials 9 (right) and 8 (left), four contact
anterior chin shields. Posterior chinshields
separated along their entire length by 1 +
1 + 2 intergenials; lateral gulars separated
from posterior chinshields. Pits present in
the loreal, preoculars, postoculars, anterior
temporals, posterior temporals, parietals,
and supralabials; absent from the rostral,
nasals, internasals, prefrontals, frontal, su-
praoculars, infralabials, and chin shields;
many small tubercles present on all head
shields.

Ventrals 161; anal divided; subcaudals
63, excluding tip; subcaudal pits unobserv-
able because horny epidermal layer missing
for all subcaudals; subcaudals/(ventrals +
subcaudals) = 0.28. Dorsal scales on tail
reduced to six rows at level of subcaudal
18, reduced to four rows at level of sub-
caudal 41, and reduced to two rows at level
of subcaudal 61.

Total length 1145 mm; snout-vent length
905 mm; tail length 240 mm (21% of total
length); mass 285 g.

Maxillary teeth on left side 29, the last
three enlarged.

305

Dorsal ground color in preservative yel-
low, varying from deep orange-yellow an-
teriorly to pale straw-yellow posteriorly. In-
terstitial skin bright orange anteriorly, be-
coming gray posteriorly. Head mustard yel-
low, darker than neck and anterior body,
without dark postocular stripe; supralabials
and infralabials with black posterior mar-
gins (Fig. 1A). Dorsum bears pattern of ~
48 bands; each band spans middle 7—9 scale
rows and is 1-2 scales long; mid-dorsal
bands staggered against equal number of
lateral blotches, each 3—4 scales high and
1—2 scales long. All bands and blotches
formed by black outlining of affected
scales; scale centers (and usually the pos-
terior margins) retain ground color, impart-
ing vague and indefinite appearance to
bands and blotches (Fig. 2). First (reduced)
dorsal band appears at level of ventral 15,
first nearly complete dorsal band at level of
ventral 28, and first trace of lateral blotch
at level of ventral 30. Series of black dashes
on first dorsal scale row up to level of ven-
tral 25, each dash extending for 1—5 scales
(Figs. 1A, 2). Tail with ~ 20 dark bands;
bands increasingly reduced and poorly de-
fined posteriorly. Venter yellow, fading
from deep orange-yellow on chin to light
straw yellow on tail, with gray flecks lat-
erally at origin of dorsal banding and grad-
ually filling in mid-ventrally; the last third
of venter evenly, though not heavily, freck-
led with gray.

Color in life (from field notes).— “‘Dor-
sum mustard yellow with vague mid-dorsal
and lateral blotches created by black outlin-
ing along the margins of affected scales.
Dorsum becoming more orange anteriorly
and top of head orange-brown. Sides turn-
ing to yellow. Venter bright yellow with a
tendency to orange-yellow on chin and
throat. Black flecks scattered on venter be-
ginning ca. 4-way down body and increas-
ing in frequency posteriorly.”

Variation.—The paratype is smaller (to-
tal length ~810 mm, snout—vent length
~756 mm, tail length 54+ mm) than the
holotype, has a broken tail, and is eviscer-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Holotype (BPBM 16539) of Tropidonophis dolasii in life.

ated anteriorly. It differs from the holotype
in having prefrontals longer than wide;
postoculars 4 on right side, 3 on left; an-
terior temporals two on each side, the upper
(anteriormost) % as large as the lower; post-
erior temporals two on each side; four in-
fralabials in contact with anterior chin
shields on left side, five on right; posterior
chinshields meeting anteriorly and separat-
ed posteriorly by 1 + 2 intergenials; pits on
head scales not observable because of loss
of horny epidermal layers; ventrals 162;
subcaudals 15 before tail broken; maxillary
teeth on left side 32, the last 4 enlarged.
Dorsal ground color brown, no darker
posteriorly than anteriorly; dorsum with
~51 bands, all bands and lateral blotches
formed by dark brown, not black, margin-
ing and more solidly filled in than for ho-
lotype. Venter pale yellow anteriorly,
changing to brown posteriorly. Barring on

lips dark brown, less distinct than in holo-
type due to general suffusion of brown pig-
ment on the head.

Comparisons to other species.— Tropi-
donophis dolasii is distinguished from T.
negrosensis in lacking a posterior reduction
in dorsal scale rows; from T. dahlii, T. den-
drophiops, T. doriae, and T. hypomelas in
having 15 (vs. 17) dorsal scale rows; from
T. mairii in having 2 (vs. 1) preocular; from
T. truncatus in having 3 or 4 (vs. 2, rarely
3) postoculars; from T. halmahericus, T.
mcdowelli, and T. punctiventris in having 8
(vs. 9) supralabials; from 7. aenigmaticus,
T. novaeguineae, and T. picturatus in hay-
ing a larger number of ventral scales (161—
162 vs. 140-152, 128-143, and 117—140,
respectively); from 7. elongatus, T. mon-
tanus, T. multiscutellatus, and T. parkeri in
having fewer subcaudal scales (63 vs. 85—
108, 71-89, 74-103, and 80-100, respec-

VOLUME 117, NUMBER 3

tively); and from T. statisticus in its larger
size (~810—1145 mm vs. maximum of 870
mm), dorsal pattern of dark bands, offset
with lateral blotches, on a yellow or brown
ground (vs. uniform gray or brown with
series of dorsal spots), and strongly barred
labials (vs. unbarred).

Only five other species of Tropidonophis
attain a size greater than one meter: 7. dah-
lii, T. doriae, T. elongatus, T. halmahericus,
and T. montanus. The first is restricted to
New Britain and the last three to the west-
ern half of New Guinea or the Moluccas.
Only 7. doriae approaches the geographic
range of 7. dolasii, being found on the ad-
jacent mainland of Milne Bay Province
(Malnate and Underwood, 1988; O’Shea,
1996), but this species has 17 dorsal scale
rows, no more than 153 ventrals in females,
and no fewer than 71 subcaudals in fe-
males.

The conspicuous yellow dorsal and ven-
tral coloring of 7. dolasii (brown dorsally
in the long-preserved paratype) and its pat-
tern of lateral blotches combined with mid-
dorsal bands are apparently unique among
Tropidonophis. In other dorsally banded
Papuan species (e.g., 7. doriae, T. hypo-
melas), the bands are typically solid, in-
stead of being formed by a network of dark-
ened scale margins, and extend across the
entire dorsum, instead of lying just on the
mid-dorsal scale rows.

Ecological notes.—The holotype was
collected in small-crowned lowland hill for-
est (Paijymans, 1975, 1976) on steep terrain
at 1090 m. The collection site faces east but
receives little direct sunlight because sur-
rounding ridges and frequent clouds block
the sun throughout much of the day. Near-
est water source was a small (~30-cm
wide) trickle among rocks in a narrow ra-
vine approximately 50—100 m elevation be-
low the collecting site. At the time of col-
lection, the region had been in a month-
long drought, although moisture was still
present under logs and some rocks. Tem-
perature varied from 15.8—21.0°C during
the two weeks of our stay. The specimen

307

was active in mid-morning and attempted
to escape. Other snakes occurring in the
same area were Aspidomorphus lineaticol-
lis, Boiga irregularis, and Tropidonophis
aenigmaticus.

The paratype was noted to come from
“transition oak-rain forest”’ and had an un-
identified Rana in its stomach.

Etymology.—The name is a patronym
honoring Dolasi Salepuna of Ulua, Fergus-
son Island, who was an invaluable field as-
sistant and captured the holotype.

Distribution.—The species is known
only from the uplands of eastern Fergusson
Island and eastern Goodenough Island (Fig.
3). It likely occurs throughout the higher
elevations of the D’Entrecasteaux Islands.

Remarks

In their revision of Tropidonophis, Mal-
nate and Underwood (1988) placed consid-
erable importance in their key on the num-
bers of anterior and posterior temporal
scales, even while documenting that these
characters show considerable intraspecific
variation. We have not emphasized these
scales for diagnosing 7. dolasii because we
are uncertain of their modal distribution,
given our few specimens and the consid-
erable variation these characters exhibit in
the genus. In considering the holotype, it is
especially uncertain |) whether the third,
small scale in the anterior-posterior series
should properly be considered an anterior
or posterior temporal, and 2) whether the
first and the third small scales in the series
are normally present or are aberrant divi-
sions unique to that specimen. We have re-
ferred to the third, small scale as a posterior
temporal based on the definition provided
by Malnate and Underwood (1988: 75) that
those scales meeting either the posterior
slope of the supralabial apex or the anterior
temporals constitute the posterior tempo-
rals. However, comparison with the para-
type, whose temporals appear more normal
in size and placement than those of the ho-
lotype, shows the region occupied by this

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Elevations above 1000 m are not demarcated.

=
a

Magnetic Nonth

D’Entrecasteaux |
Goodenough Islands

Tis |
a %o

Fergusson
a

Normanby

Coral Sea

Fig. 3.

Milne Bay

Map of southeastern New Guinea showing type locality for Tropidonophis dolasii (star) on Oya Tabu

(Mt. Kilkerran), Fergusson Island, and approximate locality for paratype (dot) on Goodenough Island,

D’Entrecasteaux Islands.

small scale in the holotype to be part of the
parietal in the paratype, suggesting this
scale is not homologous with the other tem-
poral scales. Further comparison of the two
specimens shows the small anterior tem-
poral of the holotype to be of similar place-
ment but much smaller size than the cor-
responding scale in the paratype (~20% the
size of the larger anterior temporal in the
holotype vs. ~66% in the paratype). Given
these observations, it seems likely that the
temporal scalation seen in the holotype is
aberrant.

If one assumes that the temporal scala-
tion seen in the paratype is normal for the

species, then, among New Guinean Tropi-
donophis, having two anterior temporals
would serve as a further character helping
to distinguish 7. dolasii from T. statisticus,
T. m. mairii, T. mcdowelli, and T. truncatus.
Similarly, two posterior temporals would be
a further character diagnosing our species
from 7. dahlii, T. hypomelas, and T. pic-
turatus.

The nearest relatives of Tropidonophis
dolasii are not immediately evident. It
shares with eight other species (7. aenig-
maticus, T. elongatus, T. montanus, T. mul-
tiscutellatus, T. novaeguineae, T. parkeri,
T. picturatus, and T. statisticus) the com-

VOLUME 117, NUMBER 3

mon scale conditions of 15 unreduced dor-
sal scale rows, two preoculars, three or
more postoculars, and eight supralabials. Of
these eight, only 7. elongatus and T. mon-
tanus attain an equivalent size (the remain-
der never exceed 950 mm and individuals
usually are much smaller). Only T. elon-
gatus, T. multiscutellatus, and T. novaegut-
neae sometimes have dorsal bands, al-
though the bands are solid and unlike the
margined construction seen in 7. dolasii
and are superimposed on a brown or gray,
instead of yellow, ground color. Of the eight
species, all except T. novaeguineae typical-
ly bear a postocular dark stripe, not seen in
T. dolasii. Ventral scale counts of Tropi-
donophis dolasii overlap only with those in
T. elongatus, T. montanus, T. parkeri, and
T. statisticus; subcaudal counts overlap
only with 7. aenigmaticus and T. pictura-
tus; and ventrals + subcaudals overlap only
with 7. aenigmaticus, T. multiscutellatus,
and T. statisticus. Given this chaotic pattern
of character-state similarities and our small
sample size, attempts to identify the sister
taxon of 7. dolasii would be premature.

Acknowledgments

We thank E Malesa, D. Salepuna, and J.
Tekwae for field assistance; D. Mitchell and
Conservation International for logistical as-
sistance in Milne Bay Province; D. Libai,
E Malesa, and B. Salepuna for logistical as-
sistance on Fergusson Island; C. Kishinami
for specimen curation; L. Ford and C. Rax-
worthy for loan of the paratype; B. Evans
for preparing the map; A. Kodani for pre-
paring the line drawings; the PNG National
Museum and Art Gallery for providing in-
country collaborative assistance; and the
PNG Department of Environment and Con-
servation, PNG National Research Institute,
and Milne Bay Provincial Government for
permission to work in Milne Bay Province.
This research was supported by National
Science Foundation grant DEB 0103794.

309

Literature Cited

Malnate, E. V., & G. Underwood. 1988. Australasian
natricine snakes of the genus Tropidonophis.—
Proceedings of the Academy of Natural Scienc-
es of Philadelphia 140:59—201.

O’Shea, M. 1996. A guide to the snakes of Papua New
Guinea. Independent Group Pty. Ltd., Singa-
pore, 239 pp.

Paijmans, K. 1975. Vegetation of Papua New Guin-
ea.—CSIRO Land Research Series 35:1—25 +
4 maps.

. 1976. New Guinea Vegetation. Australian Na-

tional University Press, Canberra, 212 pp.

Appendix
Specimens examined

Tropidonophis aenigmaticus: BPBM 16534, E slope
Oya Tabu, Fergusson Island, 9.4556°S, 150.7896°E,
1050 m, Milne Bay Prov., Papua New Guinea; BPBM
16535, Ulua, Fergusson Island, 9.4520°S, 150.8251°E,
0-10 m, Milne Bay Proy., Papua New Guinea; BPBM
16536, S slope Oya Waka, Fergusson Island, 9.4562°S,
150.5596°E, 980 m, Milne Bay Prov., Papua New
Guinea; BPBM 16537, 17241, Saidowai, Normanby
Island, 9.9637°S, 150.9546°E, 0-10 m, Milne Bay
Proy., Papua New Guinea; BPBM 16538, 1.4 km NE
Saidowai, Normanby Island, 9.9530°S, 150.9607°E,
40-80 m, Milne Bay Proy., Papua New Guinea;
BPBM 17243, Sibonai, Normanby Island, 10.13578°S,
150.9708°E, 0-40 m, Milne Bay Prov., Papua New
Guinea; BPBM 17242, 17244, S end Sewa Bay, Nor-
manby Island, 10.0340°S, 150.9822°E, 60 m, Milne
Bay Prov., Papua New Guinea.

Tropidonophis doriae: BPBM 13135, E branch Avi
Avi River, 5.5km S, 5.6km W of Tekadu Airstrip,
7.735°S, 146.496°E, 120 m, Gulf Prov., Papua New
Guinea.

Tropidonophis hypomelas: BPBM 12022, Weitin
River Valley, 10 km N, 8.5 km W of river mouth, New
Ireland, 4.533°S, 152.95°E, 250 m, New Ireland Prov.,
Papua New Guinea; BPBM 12163, Weitin River Val-
ley, 8 km N, 7 km W of river mouth, New Ireland,
4.554°S, 152.964°E, 150 m, New Ireland Prov., Papua
New Guinea.

Tropidonophis mairii: BPBM 3104, 3299-3300,
Balimo, 8.00°S, 142.55°E, 10 m, Western Prov., Papua
New Guinea.

Tropidonophis multiscutellatus: BPBM 5030, Biak
Island, 1.02°S, 136.27°E, Papua, Indonesia; BPBM
3783, May River, 400 m, East Sepik Prov., Papua New
Guinea; BPBM 17232-33, W Alotau, 10.3092°S,
150.3471°E, 5-10 m, Milne Bay Prov., Papua New
Guinea; BPBM 5506, Wau, 7.343°S, 146.713°E, Mo-
robe Prov., Papua New Guinea.

Tropidonophis picturatus: BPBM 4133, Garaina,
7.883°S, 147.142°E, 750 m, Morobe Prov., Papua New
Guinea.

310

Tropidonophis statisticus: BPBM_ 17293-95,
17299300, vic. Bunisi Village, 10.0171°S,
149.6002°E, 1420-1490 m, Milne Bay Proy., Papua
New Guinea; BPBM 17296—98, Siyomu Village,
10.0145°S, 149.5970°E, 1300 m, Milne Bay Prov.,
Papua New Guinea; BPBM 4146, 5125, vic. Wau,
7.343°S, 146.713°E, 1600-1650 m, Morobe Prov.,
Papua New Guinea; BPBM 4128, 5459, 6239—40, Mt.
Kaindi, 7.348°S, 146.667°E, 1800-2250 m, Morobe
Prov., Papua New Guinea; BPBM 5458, 6337, Edie
Creek, 7.358°S, 146.658°E, 2000-2200 m, Morobe

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Prov., Papua New Guinea; BPBM 6484, Bulldog Rd.,
9 km SE Wau, 2200 m, Morobe Prov., Papua New
Guinea; BPBM 3734, Sarawaget Range, 1920 m, Mo-
robe Proy., Papua New Guinea; BPBM 5497, Kililo,
Sarawaget Range, 2100 m, Morobe Prov., Papua New
Guinea; BPBM 5498, SW Kabwum, Sarawaget Range,
2300 m, Morobe Prov., Papua New Guinea; BPBM
2272, Banz, 5.50°S, 144.35°E, 1680 m, Western High-
lands Prov., Papua New Guinea; BPBM 2895, 16 km
NW Banz, Western Highlands Prov., Papua New Guin-
ea.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
117(3):311-316. 2004.

A new species of snake of the genus Omoadiphas
(Reptilia: Squamata: Colubridae) from the Cordillera
Nombre de Dios in northern Honduras

James R. McCranie and Franklin E. Castaneda

(JRM) 10770 SW 164" Street, Miami, Florida 33157-2933, U.S.A.,
e-mail: jmccrani @bellsouth.net;
(FEC) Posgrado en Biologia, Universidad de Costa Rica, San José, Costa Rica,
e-mail: castanek @ yahoo.com

Abstract.—A new species of Omoadiphas is described from the Cerro Tex-
iguat Wildlife Refuge in the Cordillera Nombre de Dios of northern Honduras.
The new species differs from the congeneric O. aurula in number of subcaudal,
supralabial, infralabial, and postocular scales, in color and pattern, and in hay-
ing the posterior nasal scale in contact with the prefrontal scale. Even though
the type-locality is declared a wildlife refuge by the Honduran government,
rapid deforestation of the area does not bode well for the continued existence
of the species at its type (and only known) locality.

Resumen.—Se describe una nueva especie de Omoadiphas del Refugio de
Vida Silvestre Texiguat, ubicado en la Cordillera Nombre de Dios en el norte
de Honduras. La nueva especie difiere de su congenerico O. aurula en el
numero de escamas subcaudales, supralabiales, infralabiales y postoculares, en
color y patron y en que tiene la escama nasal posterior en contacto con la
escama prefrontal. Aunque la localidad tipo ha sido declarada como un Refugio
de Vida Silvestre por el gobierno de Honduras, la rapida deforestaci6n que se

observa en el area es una amenaza para la nueva especie.

The Cordillera Nombre de Dios of north-
ern Honduras is an area of extremely high
endemism among amphibians and reptiles.
The Cerro Texiguat Wildlife Refuge, in the
western portion of this mountain range, is
known to harbor 18 Honduran endemic spe-
cies of amphibians and reptiles, eight of
which have their type-locality within the re-
serve (McCranie, pers. observ.). In Septem-
ber 2003, we collected a specimen of snake
in this reserve that represents an unde-
scribed species of the recently described ge-
nus Omoadiphas Kohler, Wilson, & Mc-
Cranie and another endemic for the refuge.
Herein we describe this species.

Methods

We follow the format of the description
of the holotype in Kohler et al. (2001) in

describing this new taxon. The Dowling
(1951) method was used in counting ventral
scales. Head and scale measurements were
made to the nearest 0.1 mm with dial cali-
pers held under a dissecting microscope.
Snout-vent length and tail length measure-
ments were made to the nearest mm along-
side a ruler. Measurements are abbreviated
to: snout-vent length (SVL); total length
(TL); head length (HL); and head width
(HW). Scale dimensions were made at the
longest or widest points along the longitu-
dinal or breathwise dimensions of the body,
respectively. Color (capitalized) and codes
(in parentheses) in life follow those of Smi-
the (1975-1981). The term “‘goo-eaters”’ is
used in the sense given it by Cadle &
Greene (1993) and Fernandes (1995). Com-
parative statements about other snake gen-

312

era are taken from KGhler et al. (2001) and
references cited therein.

Systematics

Omoadiphas texiguatensis, new species
Figs. 1-3

Holotype.—USNM 559599 (National
Museum of Natural History), an apparently
subadult female from approximately 2.5
airline km NNE of La Fortuna, 15°25’49’N,
87°18'32"W, 1690 m elev., Cerro Texiguat
Wildlife Refuge, Departamento de Yoro,
Honduras, collected 3 September 2003 by
Franklin E. Castaneda & James R. Mc-
Cranie. Original number LDW 13565.

Diagnosis.—Omoadiphas texiguatensis
can be distinguished from the holotype of
O. aurula (SMF 78865; subadult female),
the only known specimen of the only other
known species in the genus, in having 47
subcaudal scales (24 in O. aurula), six su-
pralabials (seven), seven infralabials
(eight), one postocular (two), the posterior
nasal contacting the prefrontal (posterior
nasal separated from prefrontal by loreal),
a dorsal pattern of a dark stripe on scale
row three on each side (dark stripe only on
vertebral row), and dark brown to nearly
black ventral surfaces in preservative (pale
yellow). The affinities of the two species of
Omoadiphas appear to lie with a group of
six other genera of snakes (see Kohler et al.
2001) that are part of a larger group re-
ferred to as “‘goo-eaters.’’ Omoadiphas tex-
iguatensis differs from the species of these
six other genera in the following ways:
from Adelphicos in having 17 dorsal scale
rows (15), 172 ventral scales (120—147),
and no anterior temporal (anterior temporal
present); from all Atractus in having a di-
vided cloacal scute (undivided) and from
select species of Atractus in lacking an an-
terior temporal (anterior temporal present in
some Atractus); from Chapinophis in hav-
ing 172 ventral scales (178-196), 47 sub-
caudal scales (29—40), no anterior temporal
(anterior temporal present), no scale row re-
duction anteriorly on body (scale row re-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

duction present), and a striped body pattern
(stripes absent); from Chersodromus in
having 172 ventral scales (124-142), 47
subcaudal scales (32—43), a divided cloacal
scute (undivided), and a striped body pat-
tern (stripes absent); from all Geophis in
having a divided cloacal scute (undivided)
and a striped dorsal pattern (stripes absent)
and from select species of Geophis in lack-
ing an anterior temporal (anterior temporal
present in some Geophis); and from Ninia
in having 172 ventral scales (122-157), no
anterior temporal (anterior temporal pre-
sent), smooth dorsal scales (keeled), a di-
vided cloacal scute (undivided), and a
striped body pattern (stripes absent).

Description of holotype.—An apparently
subadult female; TL 169 mm; SVL 143
mm; tail length 26 mm (15.4% of TL); HL
8.0 mm from front face of rostral to pos-
terior end of mandible; HW 3.9 mm at
broadest point (level of angle of mouth);
head barely distinct from neck; snout
broadly rounded in dorsal view; eye length
0.8 mm; snout length 1.9 mm, about 2.4
times as long as eye length; pupil circular;
rostral about 2.0 times wider than high (0.6
mm X 0.3 mm); internasals about 2.0 times
wider than long (0.4 mm X 0.2 mm); pre-
frontals much larger than internasals, about
as wide as long (0.9 mm X 0.9 mm), bor-
dering orbit above loreal and anterior to su-
praocular; median prefrontal suture (1.0
mm) 0.4 times as long as frontal; frontal
broadly rounded anteriorly, strongly V-
shaped posteriorly, about 1.6 times longer
than wide (2.3 mm X 1.4 mm), much long-
er than distance from its anterior edge to tip
of snout (1.6 mm); parietals about 2.1 times
longer than wide (3.4 mm X 1.6 mm), me-
dian suture (1.9 mm) shorter than frontal
length; supraoculars longer than wide (0.6
mm X 0.4 mm), bordering orbit, contacting
postocular, separated from loreal by pre-
frontal.

Nasal divided, anterior nasal contacting
rostral, internasal, and first supralabial,
posterior nasal contacting internasal, pre-
frontal, loreal, and first and second supra-

VOLUME 117, NUMBER 3

313

Fig. 1.
Omoadiphas texiguatensis.

labials, nostril located in posterior portion
of anterior nasal; loreal single, about 3.0
times longer than high (0.9 mm X 0.3 mm),
lower edge contacting second and third su-
pralabials, upper edge contacting prefrontal,
loreal bordering orbit (no preocular): post-

Drawing of dorsal (A) and lateral (B) surfaces of the head of the holotype (USNM 559599) of

ocular single, about 2.0 times higher than
long (0.6 mm X 0.3 mm); no anterior tem-
poral, posterior temporal single, about 1.7
times longer than high (1.0 mm X 0.6 mm);
supralabials 6—6, third and fourth bordering
orbit, fifth contacting postocular, parietal,

314

Fig. 2. Schematic drawing of the midbody dorsal
pattern of the holotype (USNM 559599) of Omoadi-
phas texiguatensis.

and posterior temporal, sixth contacting
posterior temporal; mental about 3.0 times
wider than long (0.6 mm X 0.2 mm), sep-
arated from chinshields by first pair of in-
fralabials, which contact each other along
ventral midline; chinshields about 1.3 times
longer than wide (1.5 mm X 1.2 mm), not
extending to border of lip, separated from
first ventral by two gular scales and four
preventral scales; infralabials 7—7, first four
contacting single pair of enlarged chin-
shields (their suture length 1.2 mm); a few
tiny scale organs (tubercles) present dorsal-
ly and ventrally on head; dorsal scales dis-
posed in 17-17-17 longitudinal rows,
smooth throughout, lacking apical pits and
supra-anal tubercles; dorsal scales in 10
rows at level of tenth subcaudal; ventrals
172; cloacal scute divided; subcaudals 47,
paired; tail spine pointed.

Color in life: Dorsal surfaces of head and
neck Chestnut (32) with Sepia (119) spots;
dorsal surface of body Prout’s Brown
(121A) with Sepia (119) spots; Sepia (119)
dorsolateral stripe present on scale row
three on each side, lateral area below stripe
Vandyke Brown (121); dorsal surface of tail

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Prout’s Brown (121A); ventral and subcau-
dal surfaces Vandyke Brown (121); iris
Vandyke Brown (121).

Color in alcohol (about two weeks after
preservation): Dorsal surface of head me-
dium brown; dorsal surface of body dark
brown with indistinct darker brown spots
present on anterior one-third; darker brown
longitudinal stripe present on scale row
three on each side of body; a vague, slightly
darker brown vertebral stripe present; dor-
sal surface of tail darker brown than that of
body; ventral surface of head pale brown,
that of body dark brown anteriorly, becom-
ing even darker brown posteriorly; subcau-
dal surface very dark brown, almost black.

Distribution and natural history notes.—
Omoadiphas texiguatensis is known only
from within the limits of the Cerro Texiguat
Wildlife Refuge (Refugio de Vida Silvestre
Texiguat). The holotype was crawling in
leaf litter next to a rotten log. Only the
snake’s tail was exposed when first sighted;
its body was under the leaves. It was found
at 1000 h in moderately disturbed cloud
forest (Lower Montane Wet Forest forma-
tion of Holdridge 1967) at 1690 m elev. A
hard rain occurred from about 1850 to 1875
h the previous day, but the weather was
clear and sunny when the snake was cap-
tured.

Etymology.—tThe specific name texiguat-
ensis is formed from Texiguat and the Latin
suffix —ensis (denoting place, locality, or
country). The name refers to the Cerro Tex-
iguat Wildlife Refuge where the holotype
was collected. We use this specific name in
an effort to stress the importance of this
wildlife refuge to the conservation status of
many Honduran endemic species of am-
phibians and reptiles (but see Discussion).

Discussion

The genus Omoadiphas is now known
from two apparently subadult females
placed in two species, making it one of the
most poorly known snake genera in the
Neotropics. Kohler et al. (2001) concluded

VOLUME 117, NUMBER 3 315

Fig. 3. Dorsal (A) and ventral (B) views of the holotype (USNM 559599) of Omoadiphas texiguatensis,
total length 169 mm.

316

that its relationships appear to lie with six
other Neotropical genera that are part of a
larger group called “‘goo-eaters” by Cadle
and Greene (1993) and Fernandes (1995).
The discovery of O. texiguatensis appears
to support this relationship as well as sup-
porting the distinctiveness of the genus.

Omoadiphas texiguatensis is truly a dif-
ficult snake to find. After collecting the ho-
lotype, we spent much of the following
three days in the area raking through leaves,
overturning and ripping apart rotten logs,
and overturning rocks in an unsuccessful at-
tempt to find more specimens. We also
walked through the area for several hours
on two nights searching for active snakes.
In addition, this was McCranie’s fourth col-
lecting trip to the area.

As noted by Wilson et al. (2001). and
McCranie & Wilson (2002), most of the
protected areas in Honduras exist on paper
only. Such is the case for the Cerro Texi-
guat Wildlife Refuge. There are no facilities
or personnel of any sort or even signage to
denote the presence of a protected area. In-
deed, people living in San Francisco (the
closest village to the type-locality of O. tex-
iguatensis) and in the area between that vil-
lage and the type-locality that we queried
are unaware that the area is a wildlife ref-
uge. In addition, crop fields and cleared ar-
eas now dominate the area around the type-
locality. We did not encounter any pristine
forest in September 2003 within an hour or
two walk in any direction from where the
holotype of O. texiguatensis was collected.
This is in sharp contrast to the condition of
the area during McCranie’s first visit in
1991 when pristine cloud forest dominated
the region. Clearly, the rapid rate of defor-
estation in the area does not bode well for
the continued existence of O. texiguatensis
or any of the other species of amphibians
and reptiles found in this region of unusu-
ally high endemism.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Acknowledgments

We thank C. Gonzalez, M. Moreno, and
H. Portillo of COHDEFOR, Tegucigalpa,
for issuing collecting and exportation per-
mits. Sefor G. Enamorado, his son Bairon,
and daughters Marina and Norma of San
Francisco, Yoro, provided us with a ride
and good company up the tortuous “road”
to the type-locality and then back to San
Francisco. We also thank EF D. Castaneda
for loaning us the vehicle in which we
reached San Francisco. An earlier draft of
the manuscript was improved upon by L.
D. Wilson. Figs. 1 and 2 were drawn by S.
Mohammadi.

Literature Cited

Cadle, J. E., & H. W. Greene. 1993. Phylogenetic pat-
terns, biogeography, and the ecological struc-
ture of Neotropical snake assemblages. Pp.
281-293 in R. E. Ricklefs and D. Schluter, eds.,
Species diversity in ecological communities:
historical and geographical perspectives. Uni-
versity of Chicago Press, Chicago, 414 pp.

Dowling, H. G. 1951. A proposed standard system of
counting ventrals in snakes.—British Journal of
Herpetology 1:97—99.

Fernandes, R. 1995. Phylogeny of the dipsadine
snakes. Unpublished Ph.D. dissertation, Univer-
sity of Texas at Arlington, 115 pp.

Holdridge, L. R. 1967. Life zone ecology, Revised edi-
tion. Tropical Science Center, San José, Costa
Rica, 206 pp.

Kohler, G., L. D. Wilson, & J. R. McCranie. 2001. A
new genus and species of colubrid snake from
the Sierra de Omoa of northwestern Honduras
(Reptilia, Squamata, Colubridae).—Sencken-
bergiana Biologica 81:269-276.

McCranie, J. R. & L. D. Wilson. 2002. The amphibi-
ans of Honduras.—Society for the Study of
Amphibians and Reptiles, Contributions to Her-
petology 19:1—625 + pls. 1—20.

Smithe, EF B. 1975-1981. Naturalist’s color guide. Part
I. Color guide. The American Museum of Nat-
ural History, New York, 182 color swatches.

Wilson, L. D., J. R. McCranie, & M. R. Espinal. 2001.
The ecogeography of the Honduran herpetofau-
na and the design of biotic reserves. Pp. 109—
158 in J. D. Johnson, R. G. Webb andy O. Flo-
res-Villela, eds., Mesoamerican herpetology:
systematics, zoogeography, and conservation.
Centennial Museum, University of Texas at El
Paso, Special Publication 1:1—200.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(3):317-329. 2004.

A new species of Kolpotocheirodon (Teleostei: Characidae:
Cheirodontinae: Compsurini) from Bahia, northeastern Brazil, with a
new diagnosis of the genus

Luiz R. Malabarba, Flavio C. T. Lima, and Stanley H. Weitzman*

(LRM) Museu de Ciéncias e Tecnologia, PUCRS, Av. Ipiranga 6681, 90619-900, Porto Alegre,
RS, Brazil. and Depto. Zoologia, IB, Universidade Federal do Rio Grande do Sul,
Av. Bento Gongalves 9500, 91501-970, Porto Alegre, RS, Brazil, e-mail: malabarb@pucrs.br;
(FCTL) Museu de Zoologia da USP, Caixa Postal 42594, 04299-970 Sao Paulo, SP, Brazil,

e-mail: fctlima@usp.br;
(SHW) Division of Fishes, WG 12, Department of Systematic Biology, MRC-0159,
Smithsonian Institution, RO. Box 37012, Washington D.C. 20013-7012, U.S.A,
e-mail: weitzman.stan@nmnh.si.edu

Abstract.—Kolpotocheirodon figueiredoi, a new species of the characid sub-
family Cheirodontinae, tribe Compsurini, is described from the upper rio Pa-
raguacu basin, Bahia, Brazil. A new diagnosis for the genus is proposed, based
mostly on scanning electron microscope (SEM) analyses of the caudal organ
of the new species and that of the single previously known species, Kolpoto-
cheirodon theloura. The genus is diagnosed now in part by the presence of a
previously undescribed, sexually dimorphic and apparently glandular, structure
found in the lower caudal-fin lobe of males. The basal relative position of
Kolpotocheirodon within the Compsurini, in which all species are inseminating,
is further supported by the presence of aquasperm in both species rather than
the apomorphic elongate sperm nuclei present in the remaining members of
the tribe.

Resumo.—Kolpotocheirodon figueiredoi é descrito para a por¢ao superior
da bacia do rio Paraguacgu, Bahia, Brasil. Propoem-se uma nova diagnose para
© género, baseada principalmente na analise de Microscopia Eletr6nica de Var-
redura do 6rgao caudal da nova espécie e de Kolpotocheirodon theloura, a
unica espécie conhecida anteriormente. O género é diagnosticado pela presenga
de uma estrutura aparentemente glandular e previamente nao descrita do lobo
ventral da nadadeira caudal dos machos. A posigao relativamente basal de
Kolpotocheirodon em Compsurini, uma tribo de peixes com inseminagao de
Cheirodontinae, é corroborada pela presen¢a de espermatozoides aproximada-
mente esféricos (aquasperm) nas duas espécies, ao invés da presenga de es-
permatozoides de nticleo alongado, como observado nos demais membros da
tribo.

The genus Kolpotocheirodon was recent-
ly described by Malabarba & Weitzman
(2000), from a single species, K. theloura,
from the uppermost tributaries of the rio
Sao Francisco and rio Parana central Brazil.
The genus is a member of the tribe Comp-
surini, subfamily Cheirodontinae (see Mal-

abarba et al., 1998) and was diagnosed pri-
marily by the presence of a unique special-
ized caudal organ at the proximal region of
the ventral caudal-fin lobe of males. This
organ consists of hypertrophied elongate
dermal flaps attached along the fin rays and
a series of relatively flat tabs and papillae

318

attached along the exposed border of those
flaps. These structures were unknown in
other inseminating or externally fertilizing
species of characids.

At the time the research of Malabarba &
Weitzman (2000) was conducted, FE C. T.
Lima and colleagues were collecting in the
rio Pratinha, a tributary of the rio Paragua-
cu, Iraquara, Bahia, Brazil and there they
discovered a new cheirodontine species that
has a caudal organ similar to that present in
K. theloura. This new compsurin species is
herein described and ecological data and
field observations from the type locality are
presented.

Data from the description of the new spe-
cies, examination of a new collection of
better-preserved specimens of K. theloura
than originally available, and scanning elec-
tron microscopy (SEM) observations of the
caudal-fin structures of these two species al-
low a reanalysis of the characters diagnos-
ing Kolpotocheirodon and redescription of
the autapomorphies that distinguish its type
species.

Methods and Materials

The systematic methods for making
counts and measurements for all specimens
studied here are the same as those described
and used by Malabarba & Weitzman (1999)
and are not re-described here. However, un-
like the convention for fin rays wherein the
count for the rays for the holotype is given
first followed by the range and mean sep-
arately for the unbranched and the branched
rays, the counts of jaw teeth do not report
a single value followed by an indication of
variation. Instead, only the range of the
counts, for example, maxilla with 2 or 3
teeth, is provided. This is because we are
confident only in counts taken from cleared
and stained specimens. SEM photographs
were taken from specimens fixed in for-
malin and preserved in 70% ethanol. Before
metalization with gold, the fins were passed
through 99% ethanol, then acetone, and
treated with a critical point dryer.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Institutional abbreviations are as listed in
Leviton et al. (1985). Character polarity for
the diagnoses of the two Kolpotocheirodon
species and a revised analysis of Kolpoto-
cheirodon monophyly is here established
by use of parsimony through a re-analysis
of the cheirodontine clade Compsurini that
was first diagnosed by Malabarba et al.
(1998). This new analysis also includes spe-
cies of the genera Saccoderma, Compsura,
Macropsobrycon, and the species Acinoch-
eirodon melanogramma (“identified”’ as
““New Genus and Species B” in Malabarba
et al, 1998), and Kolpotocheirodon theloura
(then “identified” as “‘New Genus and Spe-
cies A’).

Kolpotocheirodon Malabarba & Weitzman

Kolpotocheirodon Malabarba & Weitzman,
2000:270 (type species: Kolpotocheiro-
don theloura Malabarba & Weitzman,
2000:271 by monotypy and original des-
ignation).

Comments preliminary to the diagnosis.—
The genus Kolpotocheirodon was diag-
nosed in Malabarba et al. (1998) (as New
Genus and Species A) and in Malabarba &
Weitzman (2000) by the presence of three
apomorphic features that occur in its type
species. These characters, as described by
Malabarba and Weitzman (2000), are a spe-
cialized part of a caudal organ located at
the proximal region of the ventral caudal-
fin lobe of mature males and consist of hy-
pertrophied elongate dermal flaps attached
along the fin rays together with a series of
relatively flat tabs and papillae attached
along the exposed border of these flaps (=
character 36 in Malabarba 1998); hooks on
the anal-fin rays of mature males distributed
along the most posterior unbranched and
five anterior branched anal-fin rays (= char-
acter 30 in Malabarba, 1998); and the
twelfth and thirteenth caudal-fin rays are
dorsally concave along their basal halves
and have ventrally expanded segments (=
character 34, state 2 in Malabarba 1998).

VOLUME 117, NUMBER 3

Fig. 1.

Diagnosis.—By using SEM the special-
ized caudal-fin organ described in the pre-
vious diagnosis of Kolpotocheirodon 1s
now found to be more complex than for-
merly known. A new caudal organ, previ-
ously undescribed, corresponds to a second-
ary sexually dimorphic organ found exclu-
sively in the ventral lobe of the caudal fin
of males of both Kolpotocheirodon species.
This “‘pineapple-like’’ organ is easily rec-
ognized by its peculiar shape, somewhat
cone shaped or papilla-like, but completely
covered by smaller papillae-like bodies or
knobs (see Figs. 1, 2). These are distributed
among the large papillae of the caudal fin
of males of K. theloura (see Fig. 3), but
form the entire caudal-fin organ in K. fi-
gueiredoi (see Fig. 1). This organ is found
only in adult males of both species, sug-
gesting that it may have a reproductive

SEM of caudal organ in Kolpotocheirodon figueiredoi, male (MZUSP 55219, 25.5 mm SL), from
rio Pratinha, Iraquara, Bahia, Brazil. (A) lower caudal-fin lobe; (B) detailed image showing the smooth border
of the flap attached along the basal portion of the nineteenth caudal-fin; (C) and (D) detailed images of the
pineapple organs of the ventral lobe of the caudal fin.

function, possibly pheromone in nature.
This pineapple organ has not been found in
other cheirodontines or other characids, and
its presence supports a hypothesis of close
relationship between the two Kolpotochei-
rodon species.

Both Kolpotocheirodon species have a
conspicuous small black spot at the mid-
length of the first branched anal-fin ray of
males (Figs. 5, 7 and 8), absent in females
(Fig. 6). Such a spot is absent in all other
known cheirodontines. It is here considered
derived and a synapomorphy for the genus.

Males of Kolpotocheirodon figueiredoi
and K. theloura have the ventral body sur-
face in the area covering the pelvic bone
with a dark brown mark, nearly in the shape
of an isosceles triangle. This pigment ap-
pears to externally delineate an area corre-
sponding to the muscles inserted on the pel-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2.
caudal-fin ray of males in Kolpotocheirodon theloura. MNRJ 18081, SL 26.2 mm, from lagoa Perta Pé, rio Sao
Francisco drainage, Palmital, Minas Gerais, Brazil. (A) bar = 50 wm; (B) bar = 20 pm.

vic bone (Fig. 8). Such a spot is absent in
all other cheirodontines, and constitutes a
synapomorphy for the two species.
Malabarba & Weitzman (2000) described
the presence of well-developed hooks along
the last unbranched and five anterior
branched anal-fin rays of males as derived,
and diagnostic for Kolpotocheirodon (=
character 30 in Malabarba 1998). The new
specimens available of K. theloura, MNRJ
18081, have the last unbranched and five to
seven anterior branched anal-fin rays of
males bearing hooks (5 branched rays in 7
specimens, 6 in 23 specimens, and 7 in 3
specimens). Males of K. figueiredoi have
the last unbranched and five to six anterior
branched anal-fin rays of males bearing
hooks (5 branched rays in 6 specimens, 6
in 6 specimens; 8 in one specimen). The
anal-fin region bearing hooks also contains
modified soft tissues, absent in the remain-
ing portion of the fin. Although showing
more variability than previously described,
the condition found in both Kolpotochei-
rodon species is different from that found
in other compsurins, which have hooks
along a larger number of anal-fin rays. We
found that only in Saccoderma species
among compsurins are anal-fin hooks re-
stricted to the anterior anal-fin rays, in the
last unbranched and four anterior branched
rays. By parsimony both conditions are
considered derived and autapomorphic for

Detailed SEM images of the pineapple organs found between the papillae of the ventral lobe of the

each genus. Note: Menezes et al. (in press)
and Weitzman et al. (in press) have de-
scribed and discussed glandular soft tissue
in the anal fins of sexually active male char-
acids of many kinds including glandulocau-
dines, and some non-glandulocaudines.
This tissue is most often associated with
anal-fin hooks, but in one species a glan-
dular organ was found.

Kolpotocheirodon theloura
Malabarba & Weitzman
Fig. 7

Kolpotocheirodon theloura Malabarba &
Weitzman, 2000:271—281 (description;
relationships); 272, fig. 1 (holotype);
273-4, fig. 2-3 (paratypes); 275, fig. 4
(caudal-fin hooks); 276, fig. 5 (ventral
caudal-fin lobe); 277, fig. 6 (anal-fin
hooks); 278, fig. 7 (premaxillary and
maxillary teeth), fig. 8 (pelvic-fin hooks).

Material examined: All specimens listed in
Malabarba & Weitzman (1999), plus
MNRJ 18081, 135 spms. (10 examined,
SL 24.3—27.4 mm), Brazil, Minas Gerais,
Palmital, lagoa Perta Pé, rio Sao Francis-
co drainage.

Diagnosis.—Kolpotocheirodon theloura
is diagnosed from the new Kolpotocheiro-
don species and other characid fishes by the
following autapomorphies: As described in
Malabarba & Weitzman (2000), K. theloura

VOLUME 117, NUMBER 3

Fig. 3.

SEM images of caudal organ in Kolpotocheirodon theloura, male, MNRJ 18081, SL 26.2 mm, from

lagoa Perta Pé, rio Sao Francisco drainage, Palmital, Minas Gerais, Brazil. (A) lower caudal-fin lobe, bar = 600
zm ; (B) detailed image of the flap attached along the basal portion of the nineteenth caudal-fin ray with a series
of relatively flat tabs along its exposed border, bar = 100 pm; (C) detailed image of the flaps attached to the
eighteenth through thirteenth or fourteenth fin-rays with a single series of papillae along its exposed border, bar

= 200 pm.

has hypertrophied elongate dermal flaps at-
tached along the fin rays of the ventral cau-
dal-fin lobe of males (= character 36 in
Malabarba 1998). The flap attached along
the basal portion of the nineteenth caudal-
fin ray has a series of relatively flat tabs
along its exposed border (Fig. 3). The flaps
attached to the eighteenth through thirteenth

or fourteenth fin-rays are relatively short,
narrow and bear papillae in a single series
along its exposed border (Fig. 3A, C).
These modified flaps of the caudal organ
are not exclusive to males in K. theloura,
being also found in females, although less
developed (Fig. 4). Modified flaps are also
observed in the dorsal fin of males of K.

322

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4.

theloura (Fig. 9). These modified flaps are
independent of the sexually dimorphic
pineapple-like organs described as a syna-
pomorphy for Kolpotocheirodon and are
absent in K. figueiredoi. The modified flaps
constitute an autapomorphy of K. theloura.

As described in Malabarba & Weitzman
(2000), the twelfth and thirteenth caudal-fin
rays of K. theloura are curved, dorsally
concave along their basal halves, and with
ventrally expanded segments (= character
34, state 2 in Malabarba, 1998). This fea-
ture was not observed in K. figueiredoi and
is considered autapomorphic for the type
species, K. theloura.

Kolpotocheirodon theloura has 3—5 very
small vertical bars crossing lateral body

Detailed SEM images of the flaps bearing papillae along the basal portion of the ventral lobe of the
caudal-fin ray in Kolpotocheirodon theloura, female, MNRJ 18081, SL 26.5 mm, from lagoa Perta Pé, rio Sao
Francisco drainage, Palmital, Minas Gerais, Brazil. (A) bar = 500 pm; (B)bar = 200 pm.

stripe between pseudotympanum and area
ventral to dorsal fin (Fig. 7). Such bars are
absent in the new Kolpotocheirodon species
and in other compsurins and represent an
autapomorphy for K. theloura.

Kolpotocheirodon figueiredoi
new species
Figs. 5, 6

Holotype.-MZUSP 70037, 1 male, 30.5
mm SL, Brazil, Bahia, Iraquara, rio Pratin-
ha, Fazenda Pratinha (12°21'13’S;
41°32'57”"W), 17-21 Dec 1998; P. Gerhard,
E C. T. Lima, FE Di Dario and L. S. Rocha.

Paratypes.—All specimens collected
with the holotype: MCP 22345, 3 males,

Fig. 5.

Kolpotocheirodon figueiredoi, new species, holotype, male, MZUSP 70037, SL 30.5 mm; rio Pratin-
ha, Iraquara, Bahia, Brazil.

VOLUME 117, NUMBER 3

323

Fig. 6.
tinha, Iraquara, Bahia, Brazil.

25.1—-30.5 mm SL, 2 females, 24.0—24.8
mm SL. MZUSP 55219, (6) 14 males,
24.2-28.2 mm SL, 8 females, 24.0—31.0
mm SL; (1 male 28.2 mm SL and 1 female
26.9 mm SL Alizarin red s and Alcian blue
stained specimens cleared with trypsin; 1
male 26.2 mm SL and 1 female 26.4 mm
SL sectioned for histology; 1 male 25.5 mm
SL sectioned for TEM study).
Diagnosis.—Kolpotocheirodon figueire-
doi lacks all autapomorphies described
above for K. theloura, but has no unambig-
uous autapomorphies for its diagnosis. The

Kolpotocheirodon figueiredoi, new species, paratype, female, MZUSP 55219, SL 30.0 mm; rio Pra-

following characters have alternative states
between K. figueiredoi and K. theloura, but
these also occur alternatively among other
compsurin species. Nevertheless they are
most parsimoniously accepted either as au-
tapomorphic for K. figueiredoi or apo-
morphic for K. theloura.

Whereas males of K. figueiredoi have no
hooks on the caudal-fin rays, while males
of K. theloura have the twelfth to the four-
teenth or fifteenth principal caudal-fin rays
bearing 4—6 retrorse hooks on each side in
a row along their dorsal divisions (Mala-

Fig. 7.
Gerais, Brazil.

Kolpotocheirodon theloura, male, MNRJ 18081, SL 25.0 mm; lagoa Perta Pé, rio Palmital, Minas

324

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 8.
covering the pelvic bone showing a dark brown mark, nearly isosceles triangle shape, apparently externally
delineating the area corresponding to the muscles inserted in the pelvic bone. (B) and (C) Left lateral view of
the dorsal (B) and anal fins (C), showing the dark spots of those fins.

barba & Weitzman, 2000: fig. 4). The pres-
ence of hooks on the caudal fin is known
for several compsurins, including Acinoch-
eirodon melanogramma (hooks on caudal-
fin rays 13-14, rarely on ray 15), Saccod-
erma hastata (hooks on caudal-fin rays 13—
18), “Odontostilbe’ dialeptura (hooks on
caudal-fin rays 12-16), and Macropsobry-
con uruguayanae (hooks on caudal-fin rays
12-14, plus several spinelets along the
proximal half of caudal-fin rays 14 to 18).
However, hooks are absent in Compsura
heterura, Compsura gorgonae, and ”’ Od-
ontostilbe”’ mitoptera. Malabarba & Weitz-
man (1999, 2000) pointed out that although
these hooks are present on the ventral lobe
of the caudal fin in all these species, they
do not all occur on the same caudal-fin rays
in all species and are of different shapes. A
previous analysis of character distribution

Kolpotocheirodon. figueiredoi male, MCP23455, SL 30.5 mm. (A) Ventral body surface in the area

(Malabarba et al., 1998) indicated the pres-
ence of caudal-fin hooks as a synapomor-
phy for the compsurin cheirodontines, and
its absence a secondary reversal in some of
its species. The inclusion of a new species
bearing no hooks in the most basal genus
of the tribe allows either the recognition of
the presence of hooks as a synapomorphy
for the tribe Compsurini with a reversal in
K. figueiredoi, or the recognition of inde-
pendent acquisitions of hooks in K. thel-
oura and in the clade including the remain-
ing compsurins. The first hypothesis is pre-
ferred, since it better conforms with the pu-
tative homology of caudal-fin hooks among
compsurins (de Pinna 1991).

Males of K. figueiredoi have a conspic-
uous small black spot in the soft tissue be-
tween midlength of first and second, and
second and third branched dorsal-fin rays

VOLUME 117, NUMBER 3

Fig. 9.
rodon theloura, male, MNRJ 18081, SL 26.2 mm, from lagoa Perta Pé, rio Sao Francisco drainage, Palmital,
Minas Gerais, Brazil. (A) bar = 500 wm; and (B) bar = 100 pm.

Detailed SEM images of the flaps bearing papillae along the anterior dorsal-fin ray in Kolpotochei-

(Figs. 5, 8). This is absent (Fig. 7) in K.
theloura (= character 65 in Malabarba
1998). Among compsurins, a similar spot is
observed in species of Compsura, Macrop-
sobrycon and Acinocheirodon, but is absent
in species of Saccoderma, This spot was
previously proposed as a synapomorphy for
a clade including the last four genera cited
above. Again, the inclusion of a new spe-
cies in the most basal genus of the tribe
allows both the recognition of the presence

of the dorsal black spot as a synapomorphy
for the tribe Compsurini with a reversal in
K. theloura, or the recognition of indepen-
dent acquisitions of the dorsal black spot in
K. figueiredoi and in the clade including re-
maining compsurins. The first hypothesis is
preferred because it better conforms to the
putative homology of the dorsal black spot
among compsurins.

A further character distinguishing K. fi-
gueiredoi is its caudal-peduncle/caudal-fin

Table 1—Morphometrics of Kolpotocheirodon figueiredoi, new species. Standard length is expressed in mm;
measurements through head length are percentages of standard length; the last four entries are percentages of
head length. Range includes the holotype, MZUSP 70037, and paratypes MCP 22345, MZUSP 55219.

Halewee Males Females

male n Low High x SD n Low High xX SD
Standard length (mm) 30.5 13 25.3 303 270 10 23.8 31.0 26.4
Snout to anal-fin origin 9.7) 13 57.3 63.0 60.0 1.30 1©@ © @5,3 ©3.7/ I35
Snout to dorsal-fin origin 50.5 13 45.7 52.9 495 1.80 1@ 4B S24 ayy 135)
Snout to pelvic-fin origin 42.0 13 42.0 46.7 445 1.51 I@ = 4S5).3} E38} E33} INS IN/
Dorsal-fin base length 13}, Ih 13 Wi IS Id i120) 10 Wi 143 132 0.72
Anal-fin base length 25.9 13 24.5 29.0 26.8 1.27 10 235 26.8 25.4 1.08
Caudal peduncle length IS). 13 25 id. 4 O20) 10 Hiei 54 3s 25)
Caudal peduncle depth 13.8 13 13.6 16.1 14.8 0.76 10 Wi 133 125 O93
Depth at dorsal-fin origin Slail 13 Sil B5./ B35) iis 10) 336) 39:4 35:4 Ese
Dorsal-fin height 29.8 12 26.1 29.8 28.0 1.36 Q WZ WMS RA O83
Pelvic-fin length 19.0 13 167 19.3 IS2 O71 10 IS4b 16.3 146 O79)
Pectoral-fin length 19.3 13 175 20.1 18.8 0.77 8 IS. IO2 WA tao
Bony head length 23.3 13 Was) AP) DANS (O59) 1@ 23.0 250 Aj O72
Snout length 21.1 13 20323) 8 Oa 10 IQAE WZ.3) Biles 1335)
Upper jaw length 31.0 13 M54 Zl@ 27.7 Way 10 2a SO2 ZI 2,13)
Horizontal eye diameter 38.0 13 33.3 385 362 i139 I@ 34h} BV Bs 1s
Least interorbital width 29.6 13 ADM Bill ASQ Weil I@ 254 2S Wei 1.32

326

spot (Figs. 5, 6) that is more or less rect-
angular in shape and horizontally elongate.
It never reaches the dorsal and ventral bor-
ders of the caudal peduncle. The same spot
is vertically elongate, lozenge-shaped and
reaches the dorsal and ventral borders of the
caudal peduncle in K. theloura (Fig. 7).

Description.—Morphometric data sum-
marized in Table |. Body elongate and
compressed, greatest depth at dorsal-fin or-
igin. Predorsal profile slightly convex. Pro-
file of body along dorsal-fin base poster-
oventrally inclined, nearly straight from
base of posterior dorsal-fin ray to adipose
fin. Ventral body profile convex from tip of
lower jaw to pelvic-fin origin. Muscles cov-
ering pelvic bone strongly prominent in
ventral body profile, especially in males.
Area between pelvic- and anal-fin origins
slightly concave in females and notably
concave in males, with a pair of concavi-
ties, separated from each other by a small
median keel visible only when pelvic fins
moved out of way. These concavities lodge
pelvic fins when later retracted. Ventral pro-
file along anal-fin base slightly concave in
females. In males same profile, slightly
convex in region of anterior lobe and slight-
ly convex along remaining posterior fin
portion. Dorsal and ventral profiles of cau-
dal peduncle nearly straight in females.
Dorsal and especially ventral surfaces of
caudal peduncle of males convex, with an
internal translucent cavity, covered by cau-
dal peduncle scales.

Head small. Snout shorter than eye di-
ameter. Mouth terminal. Maxilla short, po-
sitioned at an angle of approximately 45 de-
grees relative to long body axis. Posterior
tip of maxilla reaching vertical that passes
through anterior border of eye.

Premaxilla with 4 teeth, each having 9
small evenly spaced cusps all about equal
in size. Cutting edge arched. Maxilla with
2 or 3 teeth similar in form to those of pre-
maxilla, with 7—9 cusps. Cutting edge
slightly arched to almost straight. Dentary
bone with 4 or 5 large teeth each with 7
cusps; followed by 2 or 3 smaller teeth with

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

7 or fewer cusps. Teeth posterior to second
tooth asymmetrical with most lateral cusps
situated towards tooth base and most me-
dial cusp more distally located. Cusps small
and regular and approximately equal in
size. Cutting edge slightly arched to almost
straight.

Dorsal-fin rays, 11, 9, m = 22 (ii, 8 in one
specimen). First unbranched ray less than
half-length of second. Dorsal-fin origin ap-
proximately at midlength of body. Adipose-
fin origin nearly at vertical through inser-
tion of posteriormost anal-fin ray.

Anal-fin rays, ii, 18, (iii-iv X = 3.5, 16—
19, X = 17.5, n = 22). Anal-fin origin
slightly posterior to vertical passing through
base of posteriormost dorsal-fin ray in fe-
males and at a vertical passing through base
of two last dorsal-fin rays in males. Distal
border of anal fin concave in females, with
anterior 5—6 branched rays very long, form-
ing prominent anterior lobe. Distal border
of anal fin of males convex in the anterior
lobe, decreasing in length gradually and
forming a posterior concave border. Anal-
fin rays of males with slender, elongate re-
trorse hooks on longest unbranched ray, and
anterior first 5 or 6 branched rays (scattered
hooks present in branched rays 7 and 8 in
one specimen). Hooks inserted at postero-
lateral border of fin rays, bent over lateral
surface of fin ray and anteriorly directed.
Hooks located on posterior ray branches,
less numerous on anterior ray branches.
One, rarely two, bilateral pair of bony
hooks per ray segment.

Pectoral-fin rays, 1, 9 G, 8-9, X = 8.6, n
= 22). Distal ends of longest rays not
reaching pelvic-fin origin in females; reach-
ing or not in males. Pelvic-fin rays, 1, 7 (1,7
in all specimens, n = 22). Pelvic-fin origin
anterior to vertical passing through dorsal-
fin origin. Distal tip of pelvic fin passing
anal-fin origin in males, but not in females.
Male pelvic fins bearing elongate ventro-
medial retrorse hooks along branched rays
DW ts,

Principal caudal-fin rays 10/10 (10/9, but
10/10 and 9/9, in one specimen each, n =

VOLUME 117, NUMBER 3

21). Lower caudal-fin lobe of males cov-
ered with series of papillae from 12" or 13"
ray to 18" or 19" principal caudal-fin rays.
Papillae most numerous near caudal-fin
base, extending in some specimens to near
tip of lower caudal-fin lobe. Hooks or hy-
pertrophied dorsal fin-ray flaps absent. Dor-
sal fin-rays 9-10, and ventral procurrent
caudal-fin rays 8-10, in two cleared and
stained specimens.

Scales cycloid, moderately large. Lateral-
line pores incomplete, perforated scales 7,
(5-9, X = 7.4, n = 20). Scales in lateral
series 34, (332-36, X = 33.7, n = 19). Scale
rows between dorsal-fin origin and lateral
line 5, (5-6, X = 5.2, n = 20). Scale rows
between lateral line and pelvic-fin origin 4,
(4-5, X = 4.1, n = 20). Predorsal scales,
when in regular series 11 (10-12, X = 10.8,
n = 18). :

Supraneurals, 4; precaudal vertebrae, 16;
caudal vertebrae, 17—18 Gn two cleared and
stained specimens).

Color in alcohol.—(See Figs. 5, 6, 8).
Head dark brownish dorsally with a silvery
color in opercle and infraorbital area, where
guanine not destroyed by fixative. Body
pale brownish yellow; dorsolateral scales
delineated in their borders with dark chro-
matophores. Black lateral body stripe evi-
dent, pale anterior to dorsal-fin origin, pro-
gressively wider and conspicuous posteri-
orly in larger specimens. Humeral spot ab-
sent. A conspicuous caudal spot centered at
posterior termination of caudal peduncle,
rectangle-shaped and extending to base of
middle caudal-fin rays; caudal spot never
reaching ventral and dorsal borders of cau-
dal peduncle. Dorsal fin of males with a
conspicuous small black spot in soft tissue
between approximately mid length of first
and second, and second and third branched
dorsal-fin rays; dorsal fin of females with-
out distinct marks. Anal fin of males with
a concentration of dark chromatophores
along midlength of first branched anal-fin
ray, forming a small and conspicuous spot
in adult male specimens; absent in females.
An inconspicuous dark line present along

327

anal-fin base in both sexes, plus a small
dark line on body nearly parallel to longi-
tudinal lateral body stripe in males and par-
allel to anal-fin base in females. Pectoral
and pelvic fins hyaline. Ventral body sur-
face in area covering pelvic bone of males
with a dark brown mark nearly shaped like
an isosceles triangle, apparently delineating
external area corresponding to muscles
originating from pelvic bone. Ventral mid-
line between pelvic-fin insertion and anal-
fin origin of males with a pair of thin lateral
black lines, seen only when pelvic fin ex-
tended. A dark mark present on lower in-
ternal border of orbits, visible only when
eyes depressed.

Color in life.—Described from color
slides of a male taken in the field by Pedro
Gerhard. Head dark brownish dorsally; op-
ercle and infraorbital area silvery. Body
light brownish yellow; dorsolateral scales
slightly delineated with dark chromato-
phores; belly silvery. Lateral body stripe
evident, silvery, pale anterior to dorsal-fin
origin. Humeral area unpigmented, but a
dark area visible due to presence of a pseu-
dotympanum. As described in preserved
specimens, a conspicuous caudal spot cen-
tered at posterior termination of caudal pe-
duncle, rectangle-shaped, and extending to
base of middle caudal-fin rays; never reach-
ing ventral and dorsal borders of caudal pe-
duncle. Caudal spot in colorful specimens
bordered dorsally and ventrally by two yel-
low spots. Small black spot on dorsal fin of
males, located approximately at mid length
of first and second, and second and third
branched dorsal-fin rays, bordered dorsally
by yellow pigmentation. Anal-fin spot of
males located along mid length of first
branched anal-fin ray, anteriorly bordered
with yellow pigmentation. A small dark line
along anal-fin base and a small dark line on
body nearly parallel to longitudinal lateral
body stripe visible above anterior lobe of
anal fin. Pectoral and pelvic fins hyaline.
Presence of marks on ventral body surface
not visible in available photos.

Sexual dimorphism.—Males are easily

328

recognized by their color pattern, display-
ing two conspicuous small black spots on
the dorsal and anal fins (see Fig. 5), and a
triangular dark brown mark on the ventral
body surface of the pelvic bones (see Fig.
8), absent in females (See Fig. 6). Sexes are
also differentiated by the relative position
of the pelvic and anal fins, both located
more anteriorly in males than in females;
by the larger caudal peduncle depth in
males, having an expanded portion in its
ventral and dorsal profiles; and by the larger
pelvic-fin lengths of males (see Table 1 for
all these measurements).

Distribution.—Known only from the
type locality, the rio Pratinha, Bahia, Brazil.
The rio Pratinha is a tributary of the rio
Santo Antonio, itself a tributary of the rio
Paraguacu, a coastal drainage of eastern
Brazil.

Habitat and natural history notes.—For
a complete description of the site of collec-
tion of K. figueiredoi, the rio Pratinha, see
Lima & Gerhard (2001: 112-113). In the
rio Pratinha, K. figueiredoi was observed
and collected only in those portions with a
moderate water current. The species was
most commonly collected in a riffle at a
narrow stretch of rio Pratinha. Specimens
were observed at midwater, swimming
against the current, probably feeding on
food items drifting downstream. During one
occasion, one individual of this species was
seen picking with its mouth on a large boul-
der in a cave entrance. The ecological pref-
erences of K. figueiredoi may be remark-
able, given the fact that at least some other
species of the Cheirodontinae, for example
Cheirodon interruptus (Jenyns) and Chei-
rodon tbicuhiensis Eigenmann, prefer lentic
waters such as lagoons or pools in slow-
moving water courses in coastal streams of
Rio Grande do Sul, Brazil, personal obser-
vation.

Etymology.—We take great pleasure in
naming this species in honor of José Lima
de Figueiredo, a Brazilian ichthyologist at
the Museu de Zoologia da Universidade de
Sao Paulo.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Discussion.—The two Kolpotocheirodon
species are included in the tribe Compsurini
(Malabarba et al. 1998) by sharing two un-
ambiguous synapomorphies with the mem-
bers of that tribe: they are inseminating
(Character 70 in Malabarba et al. 1998),
and the anal-fin hooks are positioned along
the posterolateral border of the anal-fin rays
and bent more or less anteriorly over the
lateral surface of the anal-fin ray to which
each 1s attached (Character 26 in Malabarba
et al. 1998). The presence of hooks and
their distribution in the caudal fin were also
previously employed for the diagnosis of
the tribe, but are absent in K. figueiredoi.
Alternative hypotheses explaining this are
discussed above under the diagnosis of this
species. Kolpotocheirodon theloura was the
only species of the Compsurini known to
have aquasperm (a nearly spherical or
spherical sperm nucleus), a condition also
found in the new Kolpotocheirodon figuei-
redoi. All other species of the Compsurini
so far investigated have elongate sperm nu-
clei (see Burns et al. 1997:434, fig. 1B—H
& 1998:242, fig. 11).

Acknowledgments

SEM pictures were made at Centro de
Microscopia e Microanalises—PUCRS. We
thank Marco Aurélio Azevedo and John
Burns for the histological analyses of the
gonads and caudal organ that provide im-
portant data concerning the generic and
species diagnoses presented herein. We
thank P. Gerhard for the photos of live spec-
imens. We also thank Pedro Gerhard, Fabio
Di Dario and L. S. Rocha for their help dur-
ing field work and Silvio Arruda and Rai-
mundo Oliveira for logistic support. Finan-
cial support was provided by CNPq (proc.
464545/00-5) and FAPESP (proc. O1/
14449-2),

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, S. H. Weitzmann, and L. R. Malabarba. 1997.
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, & . 2000. A new genus and species of
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rodontinae: Compsurini) from South America
with uniquely derived dermal papillae on caudal
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(3):330—338. 2004.

Astyanax biotae, a new species of stream fish from the Rio
Paranapanema basin, upper Rio Parana system, southeastern Brazil
(Ostariophysi: Characiformes: Characidae)

Ricardo M. C. Castro and Richard P. Vari

(RMMC) Laborat6rio de Ictiologia de Ribeirao Preto,

Departamento de Biologia da Faculdade de Filosofia, Ciéncias e Letras de Ribeirao Preto,
Universidade de Sao Paulo, Avenida Bandeirantes 3900, 14040-901, Ribeirao Preto, SP. Brazil,
e-mail: rmcastro @ffclrp.usp.br; (RPV) Vertebrate Zoology Section, Division of Fishes,
National Museum of Natural History, Smithsonian Institution,

Washington, D.C. 20560-0159, U.S.A., e-mail: vari.richard@nmnh.si.edu

Abstract.—Astyanax biotae, a new species of characid, is described from a
first-order stream in the Rio Paranapanema basin, upper Rio Parana system, in
the interior of the state of Parana, southeastern Brazil. The species differs from
its congeners in that region in a combination of morphometric and pigmentary

features.

Resumo.—Astyanax biotae, uma nova éspecie de caracideo € descrita de um
riacho de primeira ordem da bacia do Rio Paranapanema, sistema do Alto Rio
Parana, interior do Estado do Parana, sudeste do Brasil. A espécie descrita
difere das demais espécies do género Astyanax ocorrentes na mesma regiao
por uma combinagao de caracteres morfométricos e pigmentares.

Astyanax Baird & Girard includes nearly
90 nominal species of neotropical characid
fishes distributed from the southwestern
United States to Argentina (Lima et al.
2003:106). The numerous nominal species
assigned to Astyanax, in conjunction with
the lack of a comprehensive treatment of
the genus subsequent to Eigenmann (1921,
1927), often makes the identification of spe-
cies problematic. Furthermore, Astyanax as
now delimited is likely non-monophyletic,
and various species encompassed in the ge-
nus as traditionally defined (.e., characids
with two rows of teeth in the upper jaw and
with the inner tooth row consisting of five
teeth) have been generically reassigned in
recent years (e.g., Zanata 1997).

This uncertainty applies even in regions
such as the upper Rio Parana that until re-
cently had been thought to be well known
ichthyologically. Evidence from a series of
fish groups (Britski & Langeani 1988; Me-
nezes 1988; Vari 1988; Weitzman et al.

1988; Langeani 1990; Menezes 1996a,
1996b; Castro & Casatti 1997) demon-
strates that the Rio Parana system upstream
from the now submerged Sete Quedas Falls
is an area of endemism (see Castro et al.
2003), a phenomenon likely correlated with
the formidable barrier to fish migration pre-
sented, until recently, by those falls. The
numerous streams and headwaters that con-
tribute to the large rivers of this system are
inhabited primarily by fish species of small
body sizes (mostly less than 12 cm in stan-
dard length). Such small-sized species con-
stitute at least 50% of the described fresh-
water fish species of South America and
typically demonstrate a high degree of geo-
graphic endemism (Castro 1999). Such spe-
cies are highly dependent on riparian veg-
etation for food, shelter, and reproduction
(see Bohlke et al. 1978; Lowe-McConnell
1987), but those habitats are threatened by
a number of anthropogenic activities, most
notably deforestation and the extensive use

VOLUME 117, NUMBER 3

of fertilizers and pesticides in intensive ag-
ricultural practices (see Lowe-McConnell
1975, 1987; Menezes et al. 1990; Sabino &
Castro 1990; Aratyo Lima et al. 1995; Cas-
tro & Menezes 1998).

The lacunae in our understanding of the
fish diversity within the upper Rio Parana
basin and the possibility of extirpation of
as-yet unrecognized species is clearly dem-
onstrated by the species of Astyanax in that
basin. In their comprehensive overview of
the then-known species of Astyanax in the
upper Rio Parana basin, Garuti and Britski
(2000) recognized seven species of the ge-
nus within that river system. Nonetheless,
recent collecting efforts in that basin re-
vealed at least two undescribed species of
Astyanax, one of which is known only from
a narrow first-order stream running through
a narrow gallery forest that is a remnant of
the originally widespread subtropical me-
sophytic forest of that region. This species,
which may be in danger of extinction, is
described herein.

Material and Methods

Measurements are given as proportions
of standard length (SL) except for subunits
of the head that are presented as proportions
of head length. Lateral-line scale counts in-
clude all pored scales along that series, in-
cluding scales posterior to the hypural joint.
In fin-ray counts, lower-case Roman nu-
merals indicate unbranched rays, and Ara-
bic numerals indicate branched rays. The
last anal-fin rays that are joined at the base
were counted as one element. Counts for
the holotype are indicated in square brack-
ets in the text. Measurements were made
following the methods outlined in Fink &
Weitzman (1974:1—2) with the addition of
head height measured at the vertical at the
base of the supraoccipital spine. Cleared
and stained specimens were prepared fol-
lowing a modification of the method out-
lined by Taylor & Van Dyke (1985). Ver-
tebral counts include the four vertebrae as-
sociated with the Weberian apparatus.

33)

Stomach contents were analyzed on eight
specimens (37.5 to 52.5 mm SL) using the
methods of frequency of occurrence and
percent composition described by Bowen
(1992) and Hynes (1950), respectively. The
food items were grouped in broad taxonom-
ic or ecological categories reflecting their
origins, with aquatic insects and algae con-
sidered autochthonous and terrestrial in-
sects, arachnids, and vascular plants allo-
chthonous.

The following institutional abbreviations
are used: LIRP—Laborat6rio de Ictiologia
de Ribeirao Preto, Departamento de Biolo-
gia da Faculdade de Filosofia, Ciéncias e
Letras de Ribeirao Preto, Universidade de
Sao Paulo, Ribeirado Preto, Brazil;
MZUSP—Museu de Zoologia da Univer-
sidade de Sao Paulo, Sao Paulo, Brazil; and
USNM—National Museum of Natural His-
tory, Smithsonian Institution, Washington,
D.C., U.S.A.

Astyanax biotae, new species
Fig. 1, Table 1

Astyanax sp. 2. Castro et al., 2003:13, 20,
21, fig. 6.6 [Brazil, Parana, Rio Parana-
panema basin; ecology].

Holotype.—LIRP 4009, 49.8 mm SL;
Brazil, Parana State, upper Rio Parana sys-
tem, Rio Paranapanema basin, Municipio
de Diamante do Norte, Fazenda Agua
Mole, Cérrego Agua Mole (22°38’31.7'S,
52°48'59.0"W); collected by Ricardo M. C.
Castro, Hertz EK Santos, Ricardo C. Benine,
Katiane M. Ferreira, and Flavio C. T. Lima,
7 August 2000 (station PPAO29).

Paratypes.—LIRP 2734, 15 specimens,
27.5—52.3 mm SL; LIRP 4021, 2 cleared
and stained specimens, 51.3—52.5 mm SL;
USNM 373492, 15 specimens, 31.2—52.2
mm SL; MZUSP 79807, 10 specimens,
32.4—45.6 mm SL; LIRP 4276, 34 speci-
mens, 33.0—47.4 mm SL; collected with
holotype.

Diagnosis.—Astyanax biotae is readily
distinguished from all congeners in the up-
per Rio Parana basin in having the terminus

332 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1.
system, Rio Paranapanema basin, Municipio de Diamante do Norte, Fazenda Agua Mole, Corrego Agua Mole
(22°38'31.7"S, 2°48'59.0"W).

of the base of the dorsal fin situated along
the vertical through the base of the first or
second branched anal-fin ray, versus
through the origin of the anal fin (A. fas-
clatus, A. trierythropterus) or in the area of
the vent (A. altiparanae, A. cf. eigeman-

Astyanax biotae, new species, holotype, LIRP 4009, 49.8 mm SL. Brazil, Parana, upper Rio Paranda —

niorum, A. paranahybae, A. scabripinnis,
and A. schubarti). Furthermore, A. biotae
has a distinct overall reticulate pattern
formed by dark pigmentation on the ex-
posed portion of the scales versus the lack
of such a pigmentation pattern in all of the

Table 1.—Morphometric values for holotype and 30 paratypes of Astyanax biota. Standard length is expressed
in millimeters; measurements 1-15 as percentages of standard length; 16—21 as percentages of head length.

Standard length
1. Greatest body depth
2. Snout to dorsal-fin origin
3. Length of base of dorsal fin
4. Posterior terminus of dorsal fin to adipose fin
5. Posterior terminus of dorsal fin to caudal-fin base
6. Snout to origin of pelvic fin
7. Snout to anus
8. Snout to origin of anal fin
9. Length of base of anal fin
10. Length of caudal peduncle
11. Length of longest dorsal-fin ray
12. Length of first pectoral-fin ray
13. Length of first pelvic-fin ray
14. Least depth of caudal peduncle
15. Head length
16. Head height
17. Snout length
18. Gape width
19. Orbital diameter
20. Postorbital head length
21. Interorbital width

Holotype Paratypes Mean SD
49.8 27.5—52.3 44.20 6.11
34.7 34.7-41.8 38.68 1.83
54.5 50.4—56.9 53.86 32
13.3 12.3-15.1 IBS) 0.69
24.5 19.3—24.5 2290) 1.30
38.4 35.6-41.9 37.39 1.30
49.6 45.7-49.8 48.33 0.98
60.0 54.6-61.3 58.35 1.63
65.1 61.4-66.8 64.07 1.44
31.6 29.1-39.6 32.09 1.95
10.4 9.4-12.8 11.31 1128)
AY <3) 26.8—30.8 28.21 1.19
21.3 19.2—24.4 22.01 1.25
16.5 16.0-19.2 17.44 0.82
12.0 10.9-13.7 12.40 0.55
Ql 25.4-28.7 27.23 0.86
94.2 94.2—113.5 102.15 4.40
26.8 23.5-29.3 25.86 1.53
29.0 26.3-34.8 30.59 1.88
31.9 31.9-40.0 34.85 2.17
42.8 35.143.6 37/2 2.13
37.0 34.8-40.9 38.12 1.70

VOLUME 117, NUMBER 3

other species of Astyanax that occur in the
upper Rio Parana basin. Astyanax biotae
and A. paranahybae can also be distin-
guished by the difference in their relative
body heights (approximately 35—42% of SL
versus 25%, respectively).

Description.—Morphometrics of holo-
type and paratypes presented in Table 1.
Body relatively deep, less so in individuals
of less than 30 mm SL; greatest body depth
located along vertical through insertion of
pelvic fin. Dorsal profile of head distinctly
convex from margin of upper lip to vertical
through posterior nostril, straight to very
slightly convex from that point to tip of su-
praoccipital spine. Dorsal profile of body
slightly to moderately convex from rear of
head to origin of dorsal fin, straight and
posteroventrally slanted along base of dor-
sal fin, straight to slightly convex from pos-
terior terminus of base of dorsal fin to ad-
ipose fin, and slightly concave along caudal
peduncle. Slight middorsal ridge present
along predorsal region of body. Body trans-
versely rounded overall dorsally, but some-
what flattened middorsally between poste-
rior terminus of base of dorsal fin and adi-
pose fin. Ventral profile of head strongly
convex anteriorly and then slightly convex
as far as vertical through posterior margin
of eye. Ventral profile of body convex to
insertion of pelvic fin, nearly straight but
slightly posteroventrally aligned from that
point to origin of anal fin, straight to slight-
ly convex and posterodorsally slanted along
base of anal fin, straight to slightly concave
along caudal peduncle.

Head obtusely rounded anteriorly in lat-
eral profile; mouth terminal, albeit very
slightly upturned. Upper jaw with maxilla
distinctly posteroventrally angled and ex-
tending under orbit as far as vertical
through anterior margin of pupil. Nostrils
of each side very close together; anterior
opening circular, posterior crescent-shaped.
Eye relatively large and without distinct ad-
ipose eyelid. Median fronto-parietal fonta-
nel extending from mesethmoid to supra-
occipital spine; width of fontanel approxi-

685)

mately one-fourth of interorbital distance.
Infraorbital series complete with third infra-
orbital by far the largest. All infraorbitals
carrying laterosensory canal segments
proximate to inner margin of orbital rim.
Supraorbital absent. Branchiostegal rays
four. Gill-rakers long and _ setiform;
6+1+11 rakers on outermost gill-arch of
52.5 mm SL cleared and stained specimen.

Description of dentition based on two
cleared and stained specimens. Teeth on
premaxilla in two rows, with teeth of inner
row larger. Inner row with five teeth. Sym-
physeal tooth of inner series quadricuspid
and more elongate than other teeth. Second
tooth more massive and pentacuspid. Re-
maining teeth pentacuspid, with third and
fourth teeth somewhat smaller than second
tooth, and fifth tooth distinctly smaller than
all other teeth in series. Outer row of teeth
on premaxilla consisting of four tricuspid
teeth arranged in regular series with size of
teeth gradually decreasing laterally. Fourth
tooth of outer tooth row separated from
third tooth by distance twice that separating
other teeth of series. Maxilla with single tri-
cuspid or pentacuspid tooth. Dentary with
eight to 10 teeth. Anterior five dentary teeth
pentacuspid and arranged in single row.
First four dentary teeth massive and fol-
lowed by much smaller fifth tooth. Anterior
five dentary teeth followed by gap and then
three to five very small, elongate, conical
teeth.

Scales cycloid, relatively large, and firm-
ly implanted. Lateral line decurved anteri-
orly and then nearly straight along midla-
teral line, completely pored from supra-
cleithrum to base of caudal fin and followed
by apparently unossified tubular extension
running along membrane between middle
rays of caudal fin. Lateral line scales 32 to
35 [34]; scales in transverse series from or-
igin of dorsal fin to lateral line 6 or 7 [6];
scales in transverse series from insertion of
pelvic fin to lateral line 4 or 5 [4]; scales
in transverse series from origin of anal fin
to lateral line 4 or 5 [5]; scales along mid-
dorsal line between tip of supraoccipital

334

process and origin of dorsal fin 10 to 14
[11]; scales along mid-dorsal line between
posterior termination of base of dorsal fin
and adipose fin 8 to 11 [9]; horizontal scale
rows around caudal peduncle 13 to 15 [14].

Vertebrae 32(3), 33 (17), or 34 (7) [33].

Dorsal-fin rays 11,9 [11,9]; anal-fin rays 11
to iv,22 to 26 [111,24]; total number of anal-
fin rays 24 to 30 [28]; pectoral-fin rays 1,10
to 12 [1,12]; pelvic-fin rays typically 1,7,
with i,6 in three specimens, and 1,4 in both
fins in one apparently anomalous individual
[1,7]; some specimens with anteriorly di-
rected hooks on dorsal surface of pelvic-fin
rays in adpressed fin; principal caudal-fin
rays 10/9 [10/9].

Dorsal-fin margin distally rounded to
slightly truncate; first unbranched ray ap-
proximately 40% length of second un-
branched ray. Dorsal-fin origin situated at
vertical approximately at middle of SL. Or-
igin of adipose fin located slightly anterior
of vertical through posterior terminus of
base of anal fin. Pectoral fin relatively well
developed, profile distinctly acute in ad-
pressed fin. Tip of pectoral fin extending to,
or falling slightly short of, vertical through
insertion of pelvic fin. Profile of expanded
pelvic fin pointed, with lateral rays longest.
Insertion of pelvic fin located distinctly an-
terior to vertical through origin of dorsal
fin. Tip of adpressed pelvic fin extending to
origin of anal fin. Distal margin of anal fin
ranging from somewhat concave to straight,
with third unbranched and first and second
branched rays longest and subequal or first
through third branched rays longest; sub-
sequent branched rays gradually decreasing
in length. Caudal fin forked with lobes
rounded.

Color in life.—Description based on col-
or transparencies of live holotype (see also
Castro et al., 2003:fig 6.6). Overall colora-
tion silvery-brownish with silvery high-
lights on scales, particularly in abdominal
region. Basal region of exposed portions of
scales darker, particularly along regions
slightly dorsal of midlateral line. Iris, an-
teroventral portions of infraorbital region,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

lower jaw, and ventral regions of head sil-
very. Iris with green highlights. Dark pig-
mentation as in preserved specimens.

Coloration in alcohol.—Overall ground
color of specimens fixed in formalin yel-
lowish-brown on body, with guanine still
present on ventral portion of head and on
abdomen. Snout and dorsal portion of head
relatively dark. Middorsal and immediately
adjoining portions of body dark. Distinct,
ventrally attenuated humeral spot extending
from approximately two scales ventral of
dorsal midline to about one scale dorsal of
horizontal through insertion of pectoral fin.
Scales of lateral surface of body posterior
of humeral mark with dark pigmentation
field on exposed portion of each scale. Dark
spots forming irregular, discontinuous dark
stripe along midlateral surface of body.
Caudal peduncle with distinct, anteriorly-at-
tenuating, dark mark.

Dorsal, anal, and caudal fins with inter-
radial membranes covered with small dark
chromatophores. Dorsal fin with dark pig-
mentation on interradials more prominent
on distal one-half of central rays of fins,
particularly in larger individuals. Dark pig-
mentation on caudal fin particularly well
developed along middle rays of fin. Anal
fin with dark pigmentation distinctly more
developed on distal half of fin in some in-
dividuals; otherwise pigmentation of uni-
form intensity across fin. Adipose fin often
freckled with small dark spots. Pectoral and
pelvic fins with small dark spots along fin-
ray margins and on membranes.

Etymology.—The species name, biotae, 1s
in recognition of the important pioneering
role of the ““-BIOTA/FAPESP—The Virtual
Biodiversity Institute Program” (www.
biota.org.br/) in the inventory, conserva-
tion, and sustainable use of the biodiversity
resources of the State of Sao Paulo, Brazil.
This special research program of the Fun-
dacgao de Amparo a Pesquisa do Estado de
Sao Paulo (FAPESP) supported the collect-
ing efforts that yielded all known speci-
mens of the species.

VOLUME 117, NUMBER 3

Fig. 2.

Common name.—Brazil, Parana, Dia-
mante do Norte: ““Lambari’” a name also
generically applied to all other species of
Astyanax and other small characids in
southeastern Brazil.

Distribution.—Known only from the type
locality in the region called the Pontal do
Paranapanema.

Ecology.-The sample of Astyanax biota
was collected during the winter dry season
in the Corrego Agua Mole (see Castro et al.
2003:fig. 5.6), a first-order stream running
through a narrow, not very dense gallery
forest within an extensive cattle grazing
area, at an elevation of approximately 300
m above sea level. This location lies within
what was originally an extensive subtropi-
cal mesophytic forest in southern and
southeastern Brazil (Huek & Seibert 1981).
The width of the stream varied between
0.7—1.0 m and the depth between 0.17—0.40
m, with a current speed of approximately
0.2 m.s''. The marginal vegetation was
dominated by grasses of the family Cyper-
acea (Fimbristylis sp.) and ferns (Pterydo-
phyta) of the family Polypodiaceae. Water
temperature was 18.6°C; pH 8.7; dissolved
oxygen 10.6 mg.l-'; conductivity 17

335

Son, ae SR ae
Map of the upper Rio Parana basin showing type locality for Astyanax biotae (star) and major river
systems in the basin; A = Rio Paranapanema; B = Rio Parana, C = Rio Uruguay, D = Rio Tieté.

S.cm~!; and horizontal water transparency
0.4 m.

Collecting efforts along an 100 m long
stretch of the stream yielded seven fish spe-
cies in addition to Astyanax biotae: Calli-
chthys callichthys, Corydoras aeneus,
Crenicichla britskii, Gymnotus cf. inaequil-
abiatus, Gymnotus cf. sylvius, Rhamdia
quelen, and Phalloceros caudimaculatus.
Astyanax biotae was the most abundant
species in the sample (approximately 70%
of the 110 specimens in the sample) and the
second largest contributor to the fish bio-
mass (approximately 31% of the total col-
lected fish biomass) after Rhamdia quelen
(approximately 53%). These values clearly
indicate the ecological importance of Asty-
anax biotae at this site.

Although our food analysis results are
derived from a single collecting event, the
stomach content analysis of eight individ-
uals (37.5 to 52.5 mm SL; one with an
empty stomach) clearly demonstrates that
Astyanax biotae feeds primarily on arthro-
pods (approximately 80% of the diet com-
position), with debris and seeds of vascular
plants (approximately 15%) and filamen-
tous algae (approximately 6%) significantly

336

less important in the diet. Aquatic insects
(mostly aquatic larvae of the Chironomidae
followed in order by aquatic larvae of the
aquatic Coleoptera, aquatic larvae of the
Plecoptera and Trichoptera (equal amounts
of each), nymphs of the Ephemeroptera, na-
iads of the Odonata, and a single adult of
the aquatic Hemiptera) and terrestrial in-
sects (primarily worker ants, Formicidae;
followed by worker termites, Isoptera, and
adult terrestrial Coleoptera) account for ap-
proximately 30% of the ingested arthro-
pods, followed by distinctly lower numbers
of arachnids (mostly spiders, Aranae, and a
pseudoscorpion). Overall, approximately
55% of the items in the stomachs of A. bio-
tae were allochthonous and 45% were au-
tochthonous, a clear indication of the im-
portance of the riparian vegetation as a food
source for this species of Astyanax. One of
the examined specimens, a 52.5 mm SL fe-
male (USNM 373492) with a greatly dis-
tended abdomen was found to contain ap-
proximately 350 roundish, well-developed,
deep yellow oocytes 0.7—0.8 mm in diameter.

Comparative material examined.—Asty-
anax altiparanae, LIRP 35, 126 specimens,
43.0-80.1 mm SL; USNM 373491, 10
specimens, 41.1—79.9 mm SL. Astyanax cf.
eigenmanniorum, LIRP 3401, 10. speci-
mens, 55.0—70.8 mm SL; USNM 373495,
10 specimens, 48.5—68.3 mm SL. Astyanax
fasciatus, LIRP 32, 28 specimens, 42.0—
93.5 mm SL; USNM 373493, 10 speci-
mens, 45.7—83.8 mm SL. Astyanax schu-
barti, MZUSP 4263, holotype, 82.9 mm
SL; MZUSP 4264, 1 paratype, 90.4 mm
SL. Astyanax scabripinnis, LIRP 124, 562
specimens, 19.1-75.0 mm SL; USNM
373494, 10 specimens, 36.5—74.8 mm SL.
Astyanax trierythriopterus, LIRP 2017, 138
specimens, 26.3—41.2 mm SL; USNM
373496, 10 specimens, 27.8—41.1 mm SL.

Acknowledgments

The specimens of Astyanax biotae that
served as the basis for this description were
collected during a collaborative LIRP-
MZUSP expedition supported by FAPESP

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(Fundagao de Amparo a Pesquisa do Estado
de Sao Paulo) within the “BIOTA/FA-
PESP—The Virtual Biodiversity Institute
Program” (www.biota.org.br/) through the
Thematic Project “Fish diversity of the
headwaters and streams of the upper Parana
River system in the State of Sao Paulo, Bra-
zil’”’ (FAPESP grant No. 98/05072-8, Ri-
cardo M. C. Castro (LIRP) principal inves-
tigator). Research associated with this pro-
ject was supported by that grant, the Neo-
tropical Lowland Research Program of the
International Environmental Sciences Pro-
gram of the Smithsonian Institution, and the
PRONEX Project ““Conhecimento, conser-
vacao e utilizag¢ao racional da diversidade
da fauna de peixes do Brasil” (FINEP/
CNPq grant No. 661058/1997-2). The first
author is a Conselho Nacional de Desen-
volvimento Cientifico e Tecnoldgico do
Brasil researcher (grant No. 301309/91-4).
The success of the collecting effort was as-
sured by the assistance of H. FE Santos, K.
M. Ferreira and R. C. Benine (all of LIRP),
and FE C. T. Lima (MZUSP). H. E Santos
(LIRP) assisted with the preparation of the
clear and stained specimens, the stomach
extractions for diet analysis and the prepa-
ration of the photograph of the holotype. A.
L. A. Melo (LIRP) processed the speci-
mens. A. C. Ribeiro (LIRP) produced the
distribution map and K. M. Ferreira (LIRP)
helped with the identification of the stom-
ach contents. This paper was greatly im-
proved by the suggestions and criticisms of
S. H. Weitzman and C. J. Ferraris, Jr.

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neotropical distribution patterns. Academia and W. R. Heyer, eds., Proceedings of a work-
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eastern Brazil. Pp. 379—427 in P. E. Vanzolini gia, Série Zoologia, 83:88—106.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(3):339-345. 2004.

Tetragonopterus lemniscatus (Characiformes: Characidae), a new
species from the Corantijn River basin in Suriname

Ricardo C. Benine, Gabriela Zanon Pelicao, and Richard P. Vari

(RCB) Laboratorio de Ictiologia de Ribeirao Preto, Departamento de Biologia,
FFCLRP- Universidade de Sao Paulo, Av. Bandeirantes, 3900, 14040-901, Ribeirao Preto,
Sao Paulo, Brazil, e-mail: rbenine@hotmail.com;

(GZPD) Laboratério de Ictiologia de Ribeirao Preto, Departamento de Biologia,
FFCLRP- Universidade de Sao Paulo, Av. Bandeirantes, 3900, 14040-901, Ribeirao Preto,
Sao Paulo, Brazil, e-mail: gzpd@usp.br;

(RPV) Division of Fishes, Smithsonian Institution, RO. Box 37012,

National Museum of Natural History, WG-14, MRC 159, Washington, D.C. 20013-7012, U.S.A.,
e-mail: vari.richard@nmnh.si.edu

Abstract.—Tetragonopterus lemniscatus, a new species of characid characi-
form, is described from the Corantijn River basin in western Suriname. The
species is readily distinguished from its congeners (7. argenteus, T. chalceus)
by the presence of dark, longitudinal stripes positioned between adjacent scale

rows of the lateral surface of the body.

Resumo.—Tetragonopterus lemniscatus, uma nova espécie de caraciforme
caracideo, é descrita de bacia do Rio Corantijn, oeste de Suriname. Esta espécie
é€ prontamente distinguida de seus congéneres pela presenga de um padrao
estriado de coloragao ao longo do corpo, formado por faixas escuras presentes

entre as fileiras de escamas adjacentes.

The Neotropical characid characiform
genus Tetragonopterus is characterized ex-
ternally by a relatively deep body with a
transversely-flattened prepelvic region that
is bordered laterally, particularly proximate
to the pelvic-fin insertion, by distinctly-an-
gled scales, a pronounced ventral curvature
of the anterior portion of the lateral line, an
anal fin with a long base, and a complete
outer row of teeth on the premaxilla. Recent
authors (e.g., Géry 1977:450; Reis 2003:
212) have recognized only two species of
Tetragonopterus, T. argenteus and T. chal-
ceus, but the examination of samples of the
genus that originated in the Corantijn River
basin of western Suriname revealed a third
species of the genus, which we describe
herein.

Material and Methods

Measurements are given in terms of stan-
dard length (SL). Lateral-line scale counts

include all pored scales along that series, in-
cluding scales posterior to the hypural joint.
In fin-ray counts, lower-case Roman numer-
als indicate unbranched rays, and Arabic nu-
merals indicate branched rays. The last anal-
fin rays that are joined at the base were
counted as one element. Morphometric and
meristic data were taken following the pro-
cedures outlined in Fink & Weitzman
(1974). Individual meristic values in the de-
scription are followed by their frequency in
parentheses, with values for the holotype in-
dicated in square brackets. Gill rakers counts
were taken from specimens that were cleared
and counterstained following the method of
Taylor & Van Dyke (1985). Vertebral counts
were taken via radiographs and include the
four vertebra of the Weberian apparatus and
the terminal centrum.

Institutional abbreviations follow Leviton
et al. (1985) with the addition of LIRP, La-

340 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Tetragonopterus lemniscatus, new species, holotype, USNM 225366, 47.5 mm SL; Suriname, Nick-
erie District, tributary to Sisa Creek.

boratorio de Ictiologia de Ribeirao Preto, Tetragonopterus lemniscatus,
Departamento de Biologia da RPRFCLRP, new species
Universidade de Sao Paulo, Ribeirao Preto, Fig. 1, Table 1

SP, Brazil; and NZCS, National Zoological
Collection of Suriname, Paramaribo, Suri- Holotype.—USNM 225366, adult male,
name. 47.5 mm SL, Suriname, Nickerie District,

Table 1.—Morphometric data for holotype and 11 paratypes of Tetragonopterus lemniscatus.

Paratypes
Holotype Range Mean
Standard length (mm) 47.5 41.8-81.4
Percentages of standard length
Greatest body depth Silat 46.2—53.9 50.0
Snout to dorsal-fin origin 52.1 51.3-53.6 52.4
Snout to pectoral-fin origin 28.9 27.6-29.9 29.1
Snout to pelvic-fin insertion 50.9 47.5-52.2 49.6
Snout to anal-fin insertion 67.9 64.6-69.4 67.2
Caudal peduncle depth 12.0 10.3—12.0 11.0
Caudal peduncle length 9.3 7.1-9.5 8.3
Pectoral-fin length 22.8 21.7-24.3 23.0
Pelvic-fin length 19.6 17.6—21.1 IS), 7/
Dorsal-fin length 32.8 32.6-44.3 37.2
Orbit to dorsal-fin origin 38.5 37.1-42.1 38.8
Head length 26.0 25.6-29.4 28.1
Head depth DOPED, 20.4—22.2 21.4
Percentages of head length
Snout length 27.0 24.4—28.2 26.2
Upper jaw length 41.3 41.3-44.9 43.3
Horizontal orbital diameter 41.8 38. 147.0 42.9

Least interorbital width 38.6 29.6-38.6 32.9

VOLUME 117, NUMBER 3

tributary to Sisa Creek, north side of stream
approximately 700 m downstream of cross-
ing of road from Amotopo to Camp Geo-
logie, approximately 3°42’'N, 57°42'W, R. P.
Vari et al., 20 Sep 1980.

Paratypes.—All collected in Suriname,
Nickerie District. USNM 374750, 4 speci-
mens, 42.0—46.6 mm SL. LIRP 4928, 2
specimens, 47.5—47.9 mm SL, cleared and
counterstained, collected with holotype.
USNM 225523, 2 specimens, 74.0—81.4
mm SL. LIRP 4929, 1 specimen, 79.8 mm
SL, stream at km 212 of Amotopo to Camp
Geology road, at Machine Park-Camp 212,
approximately 3°50’N, 57°34’W, R. P. Vari
et al., 15 Sep 1980. NZCS F7062, 1 spec-
imen, 62.1 mm SL, formerly USNM
225320, stream entering Corantijn River, at
approximately km 385, slightly N of Tiger
Falls, approximately 4°00'N, 58°02’W, R. P.
Vari et al., 16 Sep 1980. USNM 224367, 2
specimens, 48.4-60.1 mm SL, Kamp
Kreek, 100 m N of turnoff to Camp Geol-
ogy, approximately 4°49’N, 57°28’W, R. P.
Vari et al., 13 Sep 1980.

Diagnosis.—Tetragonopterus lemnisca-
tus 1s distinguished from its two recognized
congeners, 7. argenteus and T. chalceus, by
the dark pigmentation on the lateral surface
of the body (presence of dark, longitudinal
stripes formed by pigmentation fields along
the margins of the adjoining scale rows ver-
sus the absence of dark stripes, respective-
ly). Tetragonopterus lemniscatus further
differs from T. argenteus in the number of
median scales between the tip of the supra-
occipital spine and the base of the first dor-
sal-fin ray (8 versus 12—16, respectively).

Description.—Morphometric data are
summarized in Table 1. Overall body size
moderate (41.8—81.4 mm in SL). Body pro-
portionally deep. Greatest depth of body at
origin of dorsal fin. Dorsal profile of head
slightly concave above orbit. Each nostril
located closer to anterior margin of orbit
than to each other. Supraoccipital spine
elongate, but tip of spine not extending be-
yond vertical through posterior margin of
opercle.

341

Dorsal profile of body convex from tip
of supraoccipital spine to posterior terminus
of base of dorsal fin; slightly convex from
that point to end of base of adipose fin.
Caudal peduncle profile concave both dor-
sally and ventrally. Ventral profile of body
convex from tip of lower jaw to beginning
of caudal peduncle. Prepelvic region of
body transversely flattened, with flattening
more pronounced proximate to pelvic-fin
insertion. Scales along lateral margins of
flattened region immediately anterior to in-
sertion of pelvic fin with distinct angle. Ob-
tuse median keel extending from immedi-
ately posterior of insertion of pelvic fin to
urogenital opening.

Mouth terminal. Premaxillary teeth in
two rows. Outer premaxillary tooth row
with 4 (5) or 5 (7) [5] tricuspid teeth with
median cusps most developed. Inner row
with 5 teeth with tetracuspid symphyseal
tooth followed by two pentacuspid, and
then two, rarely one, tricuspid teeth. Max-
illa with 3 tricuspid teeth along anterodorsal
portion of free anterior margin. Dentary
with 4 (4) or 5(8) [5] pentacuspid teeth fol-
lowed by series of small tricuspid teeth
(Fig. 2).

Dorsal-fin rays 11,9 (12) [41,9]. Distal mar-
gin of dorsal fin straight. Adipose fin well-
developed. Anal-fin rays iv,29 (3), iv,30 (5),
or iv,31 (4) [iv,30]. Posterior unbranched
and anterior branched anal-fin rays longest,
with distal margin of remainder of fin mod-
erately concave. Principal caudal-fin rays
i,17,1 (12) [1,17,1]. Pectoral-fins rays 1,11
(2), 1,13 (7), or 1,14 (3) [1,13]. Tip of pec-
toral fin extending beyond vertical through
insertion of pelvic fin. Pelvic-fin rays 1,7
(12) [1,7]. Tip of pelvic fin reaching to base
of first or second unbranched anal-fin ray
in smaller individuals, barely falling short
of base of first unbranched ray in larger
specimens.

Scales cycloid. Median scales anterior to
origin of dorsal fin 8 (12) [8]. Lateral line
distinctly ventrally curved anteriorly, with
33(3), 34(5), or 35(4) [33] pored scales.
Rows of scales above lateral line to origin
of dorsal fin 6 (11) or 7 (1) [6]. Rows of

342 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2.

Tetragonopterus lemniscatus, paratype, LIRP 4928, 47.5 mm SL. Premaxilla, maxilla, and lower jaw

showing form of dentition; left side, lateral view. Scale bar = 1 mm.

scales below lateral line to origin of anal fin
5 (11) or 6 (1) [5]. Scales around caudal
peduncle 14 (11) [14]. Scale sheath formed
of one row of scales overlaps basal portions
of all but three or four posterior most anal-
fin rays. Field of small scales covering base
of caudal fin; scale field extending further
distally on fins along its dorsal and ventral
margins.

Two cleared and stained specimens with
9 gill-rakers on upper limb and 13 gill-rak-
ers on lower limb of first gill arch.

Vertebrae 30 in all specimens including
holotype.

Coloration in life-—(Based on photo-
graph of recently captured specimen from
the Corantijn basin by third author). Overall
coloration silvery, but somewhat purplish
on portion of body dorsal to horizontal run-
ning approximately through dorsal margin
of orbit. Humeral spot faintly apparent.
Dark stripes on lateral surface of body ap-
parent, but slightly masked by overlying
guanine. Infraorbital series, opercle, ventral

VOLUME 117, NUMBER 3

Atlantic Ocean

Silas

Fig. 3.

56°

Map of Suriname showing collecting sites of Tetragonopterus lemniscatus. Star = holotype locality

French
Guiana

and dots = paratype localities (some symbols represent more than one locality or lot).

portion of head, and most of body bright
silver. Iris yellowish with indications of red
dorsally. Fins dusky with yellowish cast.
Color in alcohol.—Overall ground col-
oration yellowish tan. Dorsal portion of
head, jaws, nape, and portion of middorsal
region of body anterior and posterior to
base of dorsal fin distinctly darker. Posterior
margins of scales with band of dark chro-
matophores. Dark pigmentation particularly
well-developed on dorsal and ventral por-
tions of exposed regions of scales and form-
ing undulating, narrow, horizontal stripes
along regions of overlap of scale rows on
lateral surface of body. Stripes extending on
anterior portion of body from horizontal
through base of insertion of pectoral fin to
region about two scales ventral of origin of
dorsal fin. Stripes ventral of horizontal
through dorsal margin of orbit decurved

ventrally anteriorly, with posterior portion
of ventralmost stripes posterodorsally-an-
gled in region over base of anal fin. Smaller
individuals with 9 or 10 dark stripes appar-
ent. Dorsalmost stripes becoming variably
masked by overall darker pigmentation on
dorsolateral region of body in larger spec-
imens. Humeral region with indistinct,
slightly posterodorsally-aligned bar in area
above second and third scales of lateral line.
Humeral spot becoming progressively less
apparent in larger specimens. Caudal pe-
duncle with large, rounded, dark spot con-
tinuing posteriorly onto basal portions of
middle caudal-fin rays. Short, irregular, hor-
izontal stripes extending anteriorly from an-
terior margin of spot in some larger indi-
viduals.

Median fins with small, dark chromato-
phores overlying both membranes and rays

344

of rayed fins and lateral surface of adipose
fin. Distal margin of caudal fin somewhat
darker in some large specimens. Pectoral
and pelvic fins hyaline or with few, small,
dark chromatophores.

Distribution.—Tetragonopterus lemnis-
catus is only known from localities in the
Corantijn River basin in western Suriname
(hig33)):

Habitat.—The holotype locality of Tetra-
gonopterus lemniscatus was a black water
rainforest stream with a limited amount of
emergent vegetation and shadowed by
overhanging trees. The stream had a mod-
erate rate of water flow over a sand bottom
with areas of detritus. Although all other
population samples of the species were also
collected in black water, some of the loca-
tions were in full sun and at other collecting
sites the current was swift. Some locations
at which the species was collected had areas
of clay, rock, or mud bottom.

Etymology.—The species name, lemnis-
catus, from the Latin for beribboned, is in
reference to the series of dark stripes along
the lateral surface of the body in this spe-
cies.

Remarks.—Tetragonopterus was first re-
ported from Suriname by Kner (1859:38)
who cited T. chalceus for that country. That
citation may have been the basis for the in-
clusion of the species in the Surinamese
ichthyofauna by Eigenmann (1912:68;
1917:58) and for the report of the occur-
rence of the species throughout the Guianas
by Géry (1977:450). Ouboter and Mol
(1993:146) reported T. chalceus from both
the upper portion of the Corantijn River and
from the Kabalebo River, the major right
bank tributary to the Corantijn River. It is
likely that the above citations, in particular
that of Ouboter and Mol (1993), were
based, at least in part, on 7. lemniscatus.
Tetragonopterus has not been reported from
elsewhere in Suriname, although 7. chal-
ceus has been reported from a series of lo-
calities across French Guiana including the
Fleuve Maroni along the Surinamese-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

French Guiana border (Planquette et al.,
1996:320).

Comparative material.—Tetragonopterus
chalceus: MNHN A9812 (holotype); MCP
15145 (4, 1 C&S); USNM 66293 (1);
MZUSP 29820 (3) (1 C&S); MCP 14015
(1, C&S); MZUSP 40819 (2, 1 C&S). Te-
tragonopterus argenteus: MNHN A-9807
(1); MZUSP 15570 (4, 2 C&S); MZUSP
5091 (2, 1 C&S); USNM 224789 (4).

Acknowledgments

Financial support was provided by FA-
PESP (Proc. 00/1920-6, 98/05072-6, and
98/10337-0), PRONEX (Proc. 059/97), and
the Neotropical Lowland Research Program
of the Smithsonian Institution. We thank
Oswaldo T. Oyakawa (MZUSP), Carlos A.
Lucena and José Pezzi da Silva (MCP) for
the loan of specimens. Sandra Raredon
(USNM) prepared Fig. 1 and Alexandre C.
Ribeiro (LIRP) Figs. 2 and 3. Patrice Pru-
vost (MNHN) provided radiographs of
specimens of Tetragonopterus deposited at
that institution. Tatiana X. Abreu (LIRP)
and Angela M. Zanata (MZUSP) examined
and provided photographs of the holotype
of 7. chalceus. Marcelo R. de Carvalho,
Flavio A. Bockmann, and Ricardo M. C.
Castro (LIRP) and Thomas B. Vari provid-
ed valuable comments on earlier drafts of
the manuscript.

Literature Cited

Eigenmann, C. H. 1912. The freshwater fishes of Brit-
ish Guiana, including a study of the ecological
grouping of species and the relation of the fauna
of the plateau to that of the lowlands—Mem-
oirs of the Carnegie Museum 5:xii + 578, 103
plates.

1917. The American Characidae.—Memories
of the Museum of Comparative Zoology 53(1):
1-102.

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
172:1-42.

Géry, J. 1977. Characoids of the World. TFH Publi-
cations, Neptune City, New Jersey, U.S.A., 672
PP.

VOLUME 117, NUMBER 3 34

Kner, R. 1859. Zur Familie der Characinen, III. Folge Planquette, P, P. Keith, & P-Y. Le Bail. 1996. Atlas

der ichthyologischen Beitrage——Denkschriften des poissons d’eau douce de Guyane, vol. 1.
der Akademie der Wissenschaften, Wien 17: Muséum National d’histoire Naturelle and In-
137-182. stitut National de la Recherche Agronomique,
Leviton, A. E., R. H. Gibbs, Jr, E. Heal, & C. E. Paris, 429 pp.
Dawson. 1985. Standards in herpetology and’ Reis, R. E. 2003. Subfamily Tetragonopterinae. Pp.
ichthyology, part I. Standard symbolic codes for 212 in R. E. Reis, S. O. Kullander and C. J.
institutional resource collections in herpetology Ferraris, Jr., orgs., Check list of the freshwater
and ichthyology.—Copeia 1985(3):802—832. fishes of south and central America. Edipucrs,
Ouboter, P. E., & J. H. A. Mol. 1993. The fish fauna Porto Alegre, Brazil, 729 pp.
of Suriname. Chapter 8. Pp. 133-154 in P. E. Taylor, W. R. & G. C. Van Dyke. 1985. Revised pro-
Ouboter, ed., Freshwater ecosystems of Surina- cedures for staining and clearing small fishes
me. Kluwer Academic Publishers, The Nether- and other vertebrates for bone and cartilage.—

lands. Cybium 9:107-119.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(3):346-362. 2004.

Longipalpa saltatrix, a new genus and species of the meiofaunal
family Nerillidae (Annelida: Polychaeta) from an anchihaline cave
in Bermuda

Katrine Worsaae, Wolfgang Sterrer, and Thomas M. Iliffe

(KW) Zoological Museum, University of Copenhagen, Denmark, e-mail: kworsaae @zmuc.ku.dk;
(WS) Bermuda Natural History Museum, Flatts FLBX, Bermuda, e-mail: museum.bamz @ibl.bm;
(TMI) Texas A&M University at Galveston, Texas 77553-1675, U.S.A., e-mail: iliffet@tamug.edu

Abstract.—A new genus and species of the meiofaunal family Nerillidae is
described from an anchihaline cave in Bermuda. The description is based on
studies of live animals with dissecting and light microscopes, as well as studies
of fixed material with light and scanning electron microscopy. Longipalpa sal-
tatrix, new species, differs from all other nerillids by possessing a pair of
extremely long latero-ventral palps on the prostomium and a pair of ciliated
pygidial lobes. It is further characterized by the combination of the following
characters: three very short dorsal prostomial antennae, eight chaetigerous body
segments, single parapodial cirri from segment three to eight, compound chae-

tae, and hermaphroditism.

With 48 species in 17 genera (300 wm—
2 mm in length), the Nerillidae is the largest
meiofaunal family in the Polychaeta. The
family has been a member of the now re-
jected group “Archiannelida’ (e.g., Beau-
champ 1910, Goodrich 1912). The Nerilli-
dae are now believed to be more closely
related to a macrofauna family among the
Aciculata and possibly have evolved by
progenesis (Westheide 1990, Westheide &
Purschke 1996, Rouse & Fauchald 1997,
Rouse & Pleijel 2001).

Nerillids are nearly all marine and dis-
tributed worlwide, from the intertidal to
abyssal depths (3660 m—see Worsaae &
Kristensen 2003). While most nerillids are
members of the interstitial sand fauna, some
have been found in mud, fine silt, organic
debris, bacterial mats, green algae and mac-
rophytes (Jouin & Swedmark 1965, Gelder
1974, Sterrer & Iliffe 1982, Saphonov &
Tzetlin 1997, Miiller et al. 2001, Worsaae
& Kristensen 2003). Several nerillids are
known from caves: Leptonerilla prospera
(Sterrer & Iliffe, 1982) was described from
caves in Bermuda with fine silt; Mesone-

rilla diatomeophaga Nunez, 1997 in Nunez
et al. (1997) was described from a cave in
Lanzarote with diatom carpets on lapilli;
Nerilla marginalis TYilzer, 1970 was de-
scribed from a marginal cave in Istra; and
Troglochaetus beranecki Delachaux, 1921
has been reported from various freshwater
caves, groundwater reservoirs and moun-
tain rivers in Europe and Colorado, U.S.A.
[see Morselli et al. (1995) for review]. Ne-
rillids are known from all continents, except
Antarctica, and the wide geographical dis-
tribution as well as the diversity in habitats
may well reflect an old evolutionary origin
of the family.

The anchihaline Bermudian caves are in-
habited by a rich and diverse fauna, con-
sisting primarily of crustaceans (Sket & Il-
iffe 1980; Iliffe et al. 1983; Manning et al.
1986; Iliffe 1993, 1994, 2000). The most
abundant stygobiont taxa are copepods and
ostracods with 18 species each. Non-crus-
taceans include two ciliates, two gastro-
pods, and two annelids—the nerillid Lep-
tonerilla prospera and the tubificid oligo-
chaete Phallodriloides macmasterae (Er-

VOLUME 117, NUMBER 3

séus, 1986). Although most of these species
are endemic to Bermuda, many of them
have cave-adapted congeners from the Ca-
ribbean, Mediterranean and the Pacific. Sty-
gobiont taxa with such highly anomalous
distributions are believed to be Tethyan rel-
icts.

Materials and Methods

The geology of Bermuda is particularly
unusual in that the island consists of a mid-
ocean volcanic seamount, capped with ma-
rine and eolian limestone of Pleistocene
age. The numerous inland caves of Ber-
muda are totally within this limestone and
often contain tidal, anchihaline pools that
extend below sea level to a maximum depth
of about 25 m. Surface waters in these
pools are brackish, with salinity increasing
with depth to approach fully marine levels
at 3—5 m depths. The island and its caves
have been profoundly affected by changes
in sea level associated with Pleistocene gla-
ciation. During the Ice Ages, sea level was
as much as 100 m lower and the caves of
Bermuda were all dry and air filled. Large
speleothems (stalactites and stalagmites)
formed at this time by rainwater percolating
through the ground and dripping into the
caves. As glacial periods ended, sea level
rose and flooded a substantial portion of the
caves such that they are only accessible
with the use of specialized cave diving
techniques (Iliffe 1993, 1994).

The material was collected in Roadside
Cave, a small anchihaline cave located in
the Walsingham Tract of Bermuda
(32°21'N, 64°43'W) on 15, 20 and 21 Jan
2002. A low entrance crawlway opens to a
small dark chamber containing a narrow
marine lake, which extends underneath a
rock ledge and has a maximum depth of no
more than 10 m. Surface salinity and tem-
perature recorded on 28 Oct 1981 were
30%0 and 23°C, respectively. Tidal magni-
tude in the pool is 57% of that in the open
ocean, with a lag of 80 minutes. A number
of other anchihaline stygobionts inhabit this

347

small pool, including the platycopioid co-
pepods Antrisocopia prehensilis Fosshagen,
1985 in Fosshagen & Iliffe (1985) and Nan-
ocopia minuta Fosshagen, 1988 in Fosshag-
en & Iliffe (1988); the calanoid copepod
Paracyclopia naessi Fosshagen, 1985 in
Fosshagen & Iliffe (1985); the misophrioid
copepod Speleophria bivexilla Boxshall &
Iliffe, 1986; the bogidiellid amphipod Ber-
mudagidiella bermudensis (Stock, Sket &
Iliffe, 1987); the pseudoniphargid amphi-
pod Pseudoniphargus grandimanus Stock,
Holsinger, Sket & Iliffe, 1986; the halocy-
prid ostracod Spelaeoecia bermudensis An-
gel & Iliffe, 1987; the mictacean Mictocaris
halope Bowman & Iliffe, 1985; and the
gastropod Caecum troglodyta Moolenbeek
& Faber, 1987 in Moolenbeek et al. (1987).
Similar to the new species of polychaete
here described, the copepods Antriscopia
prehensile, Nanocopia minuta and Speleo-
phria bivexilla are known only from this
cave.

Samples were collected with a conical
plankton net with a diameter of 30 cm and
a mesh size of 40 wm. Rocks and projec-
tions below the water surface were covered
with a thin layer of fine silt. Before the
samples were taken, the surface layer was
whirled up with hands, fins or loose stones
from 0.5—6.5 meter’s depth and thereafter
the net was dropped and dragged through
the suspended material.

More than 70 specimens were sorted out
alive from the collected samples. Several of
these were observed and video recorded
alive with a Hitachi VK C-350 video cam-
era mounted on a Wild M 420 Makroskop
dissecting microscope. Fourteen animals
were studied and photographed alive with
an Olympus BX51 light microscope mount-
ed with a digital camera (Olympus c-3030).
Twelve of these were afterwards prepared
as permanent whole mounts. Unless other-
wise mentioned, measurements were made
on live animals. Before fixation, the ani-
mals were anesthetized in an isotonic so-
lution of MgCl,, which was added under the
cover slip for the whole mounts. The MgCl,

348

of the whole mounts was replaced by a fix-
ative (2% formaldehyde or a trialdehyde so-
lution) and then by a glycerol series from
2—100% (diluted in distilled water). When
the glycerol was fully dehydrated after two
days, the cover slip was sealed with Gly-
ceel®:

Twenty-six specimens were fixed for
scanning electron microscopy (SEM) in a
modified trialdehyde solution (Lake 1973)
and postfixed in 1% OsO,, or fixed directly
in 1% OsO,. The specimens were trans-
ferred to distilled water, dehydrated through
an acetone series, critical point dried,
mounted on stubs, sputter coated with pal-
ladium, and examined with a JEOL JSM-
6335F Field Emission scanning electron
microscope.

The study of live animals was carried out
at the Bermuda Aquarium and. Zoo
(BAMZ) and the study of fixed material
was carried out at the Zoological Museum,
University of Copenhagen (ZMUC). Types
are deposited in the Zoological Museum,
University of Copenhagen (ZMUC), Den-
mark, and in the Smithsonian Institution,
National Museum of Natural History
(USNM), Washington D.C., U.S.A.

Family Nerillidae Levinsen, 1883
Longipalpa, new genus

Diagnosis.—Longipalpa is unique
among nerillids by having two extremely
long palps on the prostomium and two
densely ciliated lobes on the dorsal side of
the pygidium. It is further characterized by
the combination of the following charac-
ters: eight chaetigerous segments between
prostomium and pygidium; prostomium
with three very short simple dorsal anten-
nae; compound serrated chaetae; single
parapodial cirri from segment three to
eight; two pygidial lobes; one anterior and
one posterior group of non-motile cilia ar-
ranged in distinct patterns and a pair of
short bands of motile cilia on prostomium;
tranverse discontinuous rows of ciliary tufts
on dorsal and ventral surface; cuticular

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

plates in pharyngeal apparatus; two pairs of
segmented nephridia from segment II-III
and IJI-IV; hermaphroditic reproduction
with one pair of spermioducts from seg-
ment VI-VII and one pair of oviducts from
segment VII-VIII.

Type species.—Longipalpa saltatrix,
new species, by present designation.

Gender.—Feminine.

Etymology.—From the Latin longus
(=long) + English palp (=prostomial ap-
pendage), in reference to the greater length
of these appendages when compared to oth-
er genera in the family.

Similarity.—Longipalpa differs from the
seventeen described nerillid genera by the
two extremely long prostomial palps and
two ciliated pygidial lobes. It furthermore
differs from most genera by the very short
length of the prostomial antennae, lack of
parapodial cirri in segment 2 and possible
lack of pygidial cirri.

Four characters have been important in
defining nerillid genera in recent years:
number of body segments (7—9), compound
or capillary chaetae, number of antennae
(O—3), and number of cirri per parapodium
(1-2) (Tzetlin & Larionov 1988, Tzetlin &
Saphonoy 1992, Westheide & Purschke
1996, Miiller et al. 2001, Miiller 2002).
Longipalpa resembles eight genera (Afro-
nerilla, Akessoniella, Micronerilla, Nerilli-
dium, Nerillidopsis, Thalassochaetus, Tro-
chonerilla, Troglochaetus) by having eight
segments. It thereby differs from four gen-
era with seven segments (Aristonerilla,
Bathychaetus, Paranerilla, Psammoriedlia)
and five genera with nine segments (Lep-
tonerilla, Meganerilla, Mesonerilla, Neril-
la, Xenonerilla). It resembles seven genera
with compound chaetae (Aristonerilla, Lep-
tonerilla, Mesonerilla, Micronerilla, Neril-
lidopsis, Paranerilla, Thalassochaetus) and
six genera with three antennae (Aristoneril-
la, Leptonerilla, Mesonerilla, Micronerilla,
Nerilla, Trochonerilla), although only Tro-
chonerilla possesses antennae of similar
short length. Two genera (Leptonerilla, Mi-
cronerilla) differ from Longipalpa by the

VOLUME 117, NUMBER 3

presence of two cirri per parapodium (ver-
sus One cirrus per parapodium in Longipal-
pa).

Six genera show resemblance to Longi-
palpa in three out of the four “generic”
characters mentioned above: Aristonerilla,
Mesonerilla, Micronerilla, Nerillidopsis,
Thalassochaetus, and Trochonerilla (see
Table 1). Micronerilla may show the great-
est resemblance with Longipalpa, but dif-
fers by having two cirri per parapodium,
pygidial cirri and two eyes. It furthermore
diverges by the much longer antennae;
many ciliary tufts on antennae, parapodial
and pygidial cirri; parapodial cirri present
on segment 2 (and sometimes on segment
1 as well) and absent on the last segment;
gonochoristic reproduction and two pairs of
spermioducts (Swedmark 1959, Jouin 1970,
Saphonov & Tzetlin 1997, Miiller 2002).
The other five genera likewise differ from
Longipalpa in several important characters
mentioned in Table 1.

Leptonerilla prospera has previously
been described from the caves of Bermuda
(Sterrer & Iliffe 1982). The two Bermudian
cave species have not been found in the
same cave or Ccave-systems, although Road-
side Cave and Walsingham Cave (type lo-
cality for L. prospera) are separated by only
290 m. Their morphology is very different,
and there is no reason to suspect that these
two species should be closely related.

Longipalpa saltatrix, new species
Figs. 1-6, Table 2

Gen sp. A in Worsaae & Miiller (2004).
Type material.—Holotype: ZMUC-POL
1675 (whole mount), 763 wm long, Road-
side Cave, Bermuda (32°21'N, 64°43’W),
0.5—6.5 m depth, 20 Jan 2002. Paratypes:
All paratypes with same locality as for ho-
lotype, 0.5—6.5 m depth, collected 15, 20
and 21 Jan 2002. Nine specimens as whole
mounts (ZMUC-POL 1676-1684) and 26
specimens on nine SEM-stubs (ZMUC-
POL 1685-1693) are deposited in the Zoo-
logical Museum, University of Copenhagen

Table 1.—Comparison with relevant genera. Abbreviations: ( ), exceptions from remaining species of a genus; —, not applicable; segm, segment.

Spermioducts

Cirri per Pygidial

parapodium

Cirri

Antennae

Others

segm

Sex

cirri

segm

Chaetae Eyes

dimensions*

Antennae

Segm

Genus

VI-VII long palps, py-

compound 3-8 hermaphroditic

short

Longipalpa

gidial lobes
ciliary tufts on

VI-VII

gonochoristic

2-7

compound

long

Aristonerilla

antennae

IV-V, V-VI

gonochoristic,

1-9

compound (O)

3 (2) long

Mesonerilla

+ VI-VII
VI-VII +

hermaphroditic
gonochoristic

ciliary tufts on

1-7

compound

long

Micronerilla

antennae

VIL-VIl
V-VI

hermaphroditic

2-7

capillary +

medium

Nerillidopsis

compound
compound

2-7
2-7

8
8

Thalassochaetus

VI-VII + palps present

gonochoristic

capillary

short

Trochonerilla

VII-VIII

4 Short = antennae shorter than prostomium and palps, medium = antennae longer than prostomium but shorter than palps, long = antennae longer than prostomium

and palps.

349

+ Unpublished observations of Trochonerilla mobilis with palps (Worsaae, personal observation).

350 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Wf j
Wi4

sn
yj
en eg
sd hg
od
4 100 um

Fig. 1. Reconstruction from light micrograph of live holotype of Longipalpa saltatrix, new species, dorsal
view. Not all chaetae are drawn. Detailed information on nephridia, gonoducts and external dorsal ciliation is
included from confocal laser scanning microscopy and scanning electron microscopy. Abbreviations: as, anterior
field of sensory cilia; be, band of cilia; bm, bulbus muscle; ct, ciliary tuft; cp, cuticular plates; dg, dorsal glands;
eg, eggs; en, enteronephridium; hg, hindgut; la, lateral antennae; mg, midgut; mo, mouth; no, nuchal organ; od,
oviduct; pa, palp; pc, parapodial cirrus; pl, pygidial lobe; ps, posterior field of sensory cilia; sd, spermioduct;
sg, salivary glands; sn, segmented nephridium; tb, tranverse ciliary band.

VOLUME 117, NUMBER 3

ies)
Nn
—

Table 2.—Meristics and morphometric characters of holotype and total type material (measurements on ju-
veniles in parentheses). Abbreviations: excl., exclusive; incl., inclusive; L, length; min., minimum; max., max-

imum; segm., segment; W, width.

Holotype
Total
L excl. appendages, chaetae 763
max. W incl. parapodia 224
max. W excl. parapodia 192
prostomium
L 72
W 81
max. L palps 696
L median antenna 43
max. L lateral antennae 56
trunk
L segm. | 80
L segm. 2 123
L segm. 3 113
L segm. 4 109
L segm. 5 103
L segm. 6 67
L segm. 7 56
L segm. 8 35
L pygidium 8
max. L parapodia segm. 1 63
max. L other parapodia 48
max. L parapodial cirri 67
max. L pygidial lobes 42
chaetae
max. no. chaetae segm. 1?

max. no. chaetae notopodia*
max. no. chaetae neuropodia?
max. total L chaetae

max. L shaft>

L distal extension shaft?

L blade?

Min Max Average n
624 (471) 985 788 14
127 (116) 285 202 13
108 (99) 268 1 13

59 77 68 12
67 (66) 90 80 12
680 (660) 718 696 6
24 (17) 43 33 7
41 (31) 65 53 10
55 (46) 97 74 2
98 (77) 150 126 12
76 (71) 152 108 12
73 121 104 12
71 (69) 103 93 12
54 106 80 12
54 89 68 11
25 58 42 11
8 37 20 12
35 63 53 13
30 50 40 12
4] 73 60 13
30 50 4] 9

7 (6) 13 10 8

7 10 8 6

6 10 8 6
135 145 139 8

86 109 103 5

0) 2 1 5)

33 41 37 5)

4 Measured on fixed material by SEM.

> Measured alive by LM and on fixed material by SEM.

(ZMUC), Denmark. Two paratypes as
whole mounts (USNM 1022181-1022182)
are deposited in the Smithsonian Institution,
National Museum of Natural History,
Washington, D.C., U.S.A.

Diagnosis.—Characters of the genus.

Etymology.—From the Latin saltator
(=dancer), in reference to the swimming
skills of the species, which may swim in
loops while waving and twisting the long
palps.

Description (see Table 2 for principle
counts and measurements).—A relatively

hyaline nerillid with brown pigmentation,
especially along intestinal wall. The body
consists of prostomium, eight chaetigerous
segments and pygidium (Figs. 1, 2A, 4).
Adults with eight segments and a total
length of 624—985 wm (only 454-825 wm
when fixed); juveniles with six to seven
segments and a total length of about 500
wm. Maximum body width generally at
segment five, up to 268/285 wm (excl./incl.
parapodia); narrow restriction between seg-
ment one and two, posterior to the esoph-
agus. Prostomium up to 77 pm long, 90 ~m

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

a re & F dee
22) BOOT
—

Fig. 2. Light micrographs of live holotype of Longipalpa saltatrix, new species. A, Whole specimen with
two palps. B, Closer view of ciliation on palp and prostomium. C, Closer view of prostomium and segment one.
Abbreviations: see Fig. 1; pp,, parapodium of segment one; pr, prostomium; I-VI, segments one to eight.

VOLUME 117, NUMBER 3

Fig. 3. Light micrographs of live specimens of Longipalpa saltatrix, new species. A, Dorsal view of segment
three to six showing midgut lining of glandular cells with vesicles. B, Ventral view of middle segments showing
diffuse glandular pattern. C, Posterior part of animal. D, Parapodium. E, Segments seven-eight and pygidium.
Abbreviations: see Fig. 1; arrowhead, extension of shaft; cb, chaetal blade; cs, chaetal shaft; dg, diffuse glandular
pattern; fa, fascicle; gc, glandular cells; ov, ovoids; py, pygidium; VI-VIII, segments six to eight; ve, vesicles.

354

wide; first four segments longest, decreas-
ing in length posteriorly, pygidium even
shorter.

Prostomium short, with two ventro-lat-
eral palps and three dorsal antennae. Palps
filiform and long, up to 718 wm (up to
about 90% of body length in adults, and up
to about 130% in juveniles), and with com-
plex ciliation (see below) (Figs. 1, 2, 5A).
Antennae short, filiform, with few distal cil-
ia. Medium antenna up to 43 pm long, lat-
eral antennae up to 65 pm long (Figs. 1,
4A, B, 5A). Nuchal organs paired, situated
between palps and parapodia of segment
one on a round elevated bulge on each lat-
eral. side of the prostomium (Figs. 1, 5A,
C). Parapodia of segment one very large
(up to 63 pm long), up to twice the length
of the following parapodia (Figs. 1, 2A, 4A,
C).

Parapodial cirri (with few distal cilia) be-
tween dorsal and ventral chaetal bundles of
parapodia of segment three to eight; length
up to 73 wm, increasing towards the pos-
terior segments (Fig. 4A). No trace of at-
tachment of parapodial cirri on segment one
and two, neither of scars from detached cir-
ri, or rudimentary cirri. Appendages like
cirri and palps, and even chaetae, were eas-
ily lost during handling and fixing of the
animals. Of the more than 70 specimens ob-
served alive, none possessed parapodial cir-
ri On segment one and two and scars were
not found with SEM. Pygidial cirri were
never observed, but it was difficult to ex-
amine the pygidium thoroughly for scars of
cirri with SEM. On one specimen, a pair of
scar-like structures was found at the pygid-
ium, which could be scars from lost pygid-
ial cirri or just an artifact (Fig. 6G). All
adult (but no juvenile) animals possessed
two very special structures, here named py-
gidial lobes due to their location on the dor-
sal side of the pygidium. Each lobe is up to
50 wm long, with two projections and a
dense ciliation (Figs. 1, 3E, 4A, 6G, H).

All chaetae compound and relatively
straight, shaft with minor pointed distal ex-
tension, less than 2 wm long (Figs. 3D, 6B).

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Chaetae very slightly serrated and generally
with a hairy appearance (Fig. 6A—C). Seg-
ment one uniramous, with up to thirteen
chaetae in one chaetal fascicle; segments
two to eight biramous, with dorsal and ven-
tral fascicles comprising up to ten chaetae
each. Similar numbers of chaetae in seg-
ments two to six, somewhat fewer chaetae
in the last two segments. No noticeable dif-
ferences in number or length between dor-
sal and ventral chaetae. Similar lengths in
all segments of shaft, blade and total length
of chaeta. Shaft up to 109 um, blade up to
41 pm, total length up to 145 pm (Figs. 3D,
4, 0A).

Dorsal surface of prostomium with very
specific ciliation characterized by three dif-
ferent groups of cilia: a pair of short bands
with motile cilia (>20 cilia, up to 20 pm
long), one on each dorso-lateral surface
next to the lateral antennae (Figs. 1, 5A);
two transverse rows of non-motile cilia in
front of antennae on the anterior most part
of the prostomium (Fig. 5A, B), the last of
which arranged in a distinct pattern; and a
posterior group of twenty non-motile cilia
(Fig. 5D, E), arranged in complex pattern
near the origin of median antenna on pos-
terior part of the prostomium.

The patterns of the anterior field of cilia
(probably sensory in function, see Discus-
sion) and the posterior fields of cilia (prob-
ably sensory) are characteristic of the spe-
cies and are here given in detail: anterior
field of cilia (S-15 pm long) with two
transverse rows of cilia (Fig. 5B). Posterior
row contains about 5 cilia, spaced 2—5 wm
apart (cilia no. 1-5 in Fig. 5B). Anterior
row with about 20 cilia (no. 6—25 in Fig.
5B). Cilia arranged in distinct pattern mir-
rored halfway along the row. Moving to-
wards the middle from the lateral sides, the
first cilia are two groups of four cilia (no.
6-9 and 10-13), a single cilium next to
them (no. 14 and 15), two groups of three
cilia (no. 16—18 and 19—21), three cilia in
the middle (no. 22—24), and one cilium (no.
25) in front of the middle cilium.

ie)
Nn
Nn

VOLUME 117, NUMBER 3

Fig. 4. Scanning electron micrographs of Longipalpa saltatrix, new species. A, Dorsal view of whole spec-
imen with both palps lost. B, Lateral view of specimen with one palp lost. C, Ventral view of specimen with
two palps lost. Abbreviations: see Figs. 1, 2; arrowhead, connection of chaetal shaft and blade; db;, dorsal ciliary
band of segment three; ma, median antennae; mv, midventral ciliary band; vb,-vb,, ventral ciliary band on
segments one to seven.

356 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Scanning electron micrographs of Longipalpa saltatrix, new species. A, Dorsal view of prostomium,
right palp, and segment one. B, Closer view of anterior field of twenty-five cilia. C, Closer lateral view of nuchal
organ and extra ventral ciliary band. D, E, Close dorsal view of posterior field of twenty cilia from two speci-
mens. Abbreviations: see Figs. 1, 2, 4; bc, band of cilia; sc, scar from lost palp; vc, ventral ciliation around
mouth; xb, extra ventral ciliary band.

VOLUME 117, NUMBER 3 357

Fig. 6. Scanning electron micrographs of Longipalpa saltatrix, new species. A, Chaetal bundle and para-
podial cirrus. B, Closer view of chaetae with microvillar hairy appearance. C, Two hairy chaetae showing
serration pattern (indicated by arrowheads). D, Left ventral side of prostomium and segment one. E, Right side
of segment six with half dorsal ciliary band. E Left ventral side of segment three with half ventral ciliary band.
G, Left dorsal side of posteriormost segments. H, Closer view of left pygidial lobe. Abbreviations: see Figs. 1-
5; ch, holes after lost chaetae; db, dorsal ciliary band of segment six; ex, extension of shaft; gr, dense ciliary
groups; sc, scars from lost pygidial cirri or an artifact.

358

Posterior field of twenty cilia (3-15 pm
long) covers an area about 12 wm wide and
8 wm long (Fig. 5D, E). Four cilia in a close
square (no. 1—4 in Fig. 5D) surrounded by
a common elevation of the cuticle, are
found in the center of the field, posterior to
the basal part of the antenna. Right next to
these four cilia one cilium is found on each
lateral side (no. 5—6), which as all the single
situated cilia in the pattern, is surrounded
by a cuticular collar. On each lateral side of
the antenna is one cilium (no. 7—8). About
3 wm posterior of these two clusters are
found, each with three cilia in a transverse
line, surrounded by a common elevation of
the cuticle (no. 9-14). Next to these, on the
level of the 4 central cilia, are found 2 pairs
of cilia next to each other on each lateral
side (no. 15-18). The last pair of cilia (no.
19—20) is located a few micrometers poste-
riorly with about 5 pm in between the cilia.

Palps with complex ciliation containing
transverse ciliary bandlets in a longitudinal
row on the inner and outer lateral surfaces
of the palp, respectively. More than 20 cilia,
up to 20 wm long in each ciliary bandlet,
positioned 5—20 pm apart in a row extend-
ing to the tip. Farthest distance between
bandlets on outer lateral surface of the palp.
Longitudinal row of single ciliary tufts on
both dorsal and ventral surface between the
longitudinal rows of bandlets (Figs. 1, 5A).
Less than 5 cilia, up to 7 wm long in each
ciliary tuft, located in a row extending to
the tip. Ciliary bandlets beat in metacronal
waves, creating a water current leading par-
ticles towards the base of the palp. Motility
of ciliary tufts not clearly distinguishable
due to intense beating of ciliary bandlets.
However, we suspect these cilia to be non-
motile due to their small number and short
length.

Dorsal surface of body segments not cil-
iated, except for few ciliary tufts. Two to
four tufts of motile cilia are situated in a
transverse line across each segment between
the parapodia on each side (Figs. 4A, B, 6E).
Each tuft contains 20—200 motile cilia, up to
ca. 25 4m long. Each pygidial lobe possess-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

es two large groups of cilia, one on each of
the two projections of the lobe (Figs. 1, 4A,
6G, H). Each group contains more than 100
cilia, up to ca. 25 wm long.

Ventral surface with dense ciliation
around mouth on ventral side of prostomi-
um, continuous with relatively narrow mid-
ventral ciliary band extending to the anus
on the dorsal side of the pygidium. Tran-
verse rows of ventral ciliary tufts on each
segments at the level of the parapodia: four
pairs on segments one to three, three pairs
on segments four to seven, two pairs on
segment 8. Three additional pairs of ciliary
tufts: one pair between prostomium and
segment one, almost connecting ciliation
around mouth with that of the nuchal or-
gans; and two pairs between segment one
and two (Figs. 4C, 6D).

Pharynx with ventral opening between
prostomium and segment one, and muscular
bulb in segment one (Figs. 1, 2A, C). About
six pairs of ventral brown glands (may have
salivary function) open into buccal cavity
on ventral side of pharynx (Figs. 1, 2A, C).
Two additional dark brown, round cell-
groups (probably glandular in function)
dorsally of pharynx in prostomium. All
groups contain several cells with relatively
large round vesicles. A pair of triangular
cuticular plates on ventral side of pharyn-
geal bulb in anterior part of pharyngeal or-
gan (Figs. 1, 2C). Large round glandular
cells with many small vesicles and brown
pigmentation line stomach wall (Figs. 1,
3A); large ciliated cells line hindgut (Figs.
1, 3C). Diffuse superficial glands create a
unique pattern in the ventro-caudal epithe-
lium (Fig. 3B).

Studies by confocal scanning microscopy
showed the distribution of nephridia and
gonoducts in this and other species (Wor-
saae & Miiller 2004). Longipalpa saltatrix
is hermaphroditic with one pair of sper-
mioducts in segments six to seven and one
pair of oviducts in segments seven to eight
(see Fig. 1 and Worsaae & Miiller 2004, fig.
2J). Two pairs of segmented nephridia are
present, from segments two to three, and

VOLUME 117, NUMBER 3

from segments three to four, respectively
(see Fig. 1, and Worsaae & Miiller 2004,
fig. 2G—I). Several enteronephridia line the
hindgut (see Fig. 1, and Worsaae & Miiller
2004, fig. 2G, J). Fertile animals contain a
maximum of two large eggs with diameters
up to 170 wm, and an additional large num-
ber (up to ca. 40 has been counted) of
smaller ovoids with a diameter about 10—
20 wm.

Distribution.—Presently known only
from a certain anchihaline cave pool in Ber-
muda.

Motility.—The animals swim beautifully
in the water column, describing loops and
turns. Less frequently, they glide over the
surface and if provoked make an escape re-
action or quick turn by undulation of the
body and fast curling up of the palps in a
narrow spiral. When swimming, they are
capable of bending the prostomium and
body as well as waving, bending and curl-
ing the long palps. The pygidial lobes are
flapped between positions flat along the
body to an almost right angle to the dorsal
surface, thereby using the densely ciliated
lobes as helms. The forward drift seems to
be created mainly by the ventral ciliation,
possibly with additional force from the cilia
on palps and pygidial lobes.

Remarks.—The description of Longipal-
pa saltatrix not only adds a new genus to
the family Nerillidae, but also expands the
definition of the family. The extremely long
palps of this species are not only unusual
in their length but probably also in their
function. The longitudinal rows of ciliary
bandlets create a water current propelling
particles towards the mouth opening, which
has never been observed in other nerillids.
It seems possible that the animals collect
food particles by help of the palps, thereby
increasing their feeding radius extensively.
Foraging could happen when gliding over
or through the substrate as well as when
swimming through and above the sediment
collecting particles in suspension. In several
species of nerillids, the ciliation of the
much shorter palps creates a water current

359

transporting particles away from the mouth
(Worsaae, personal observations). This
transport may indicate that other nerillid
palps may also be functional in feeding be-
havior, however, by transporting rejected
particles away from the mouth and not by
gathering them. The previous understand-
ing of the nerrillid palps as being mainly
sensory in function should probably be ex-
panded to include a function in feeding be-
havior. This view stands in contrast to the
general comprehension of the ventral palps
of the major taxon group Aciculata (see
Rouse & Fauchald 1997) with which the
family has the most apparent resemblance
(see e.g., Schmidt 1848, Quatrefages 1866,
Westheide 1990, Westheide & Purschke
1996, Rouse & Fauchald 1997, Rouse &
Pleijel 2001). Aciculates are generally char-
acterized by short sensory palps with no di-
rect function in food collecting. If the neril-
lids truly belong to the Aciculata, then the
long palps of Longipalpa saltatrix may also
influence the conception of the conserva-
tiveness of the palps in this taxon group.
The extremely long palps would proba-
bly be disadvantageous in the interstitial
habitat from which many nerillids are de-
scribed. This disadvantage may be one of
the reasons why these long palps have not
been found in other nerillids. In the Ber-
mudian caves with only a sparse layer of
very fine silt on top of bare rocks, it may
even be an advantage to be able to actively
swim and perhaps also feed on suspended
particles with the aid of the palps. However,
Leptonerilla prospera which lives under
rather similar conditions, except from more
light in the Walsingham Caves in Bermuda,
does not possess long feeding palps. Dif-
ferences in habitat characteristics between
cave and interstitial habitats include more
space in caves, thus allowing for swimming
as opposed to crawling, and different types
of food, hence different feeding mecha-
nisms. Many other anchihaline species are
most commonly found within the water col-
umn rather than on the sediments, implying
that this is where food is primarily located.

360

The motile ciliary bands and the anterior
field of cilia on the prostomium could easily
be detected on live animals with light mi-
croscopy (LM), whereas the posterior field
of cilia could only be detected with SEM.
The motile ciliary bands (be in Figs. 1, 5A)
are most likely not mechanoreceptors be-
cause of their long length, motility, and
dense grouping. However, the anterior and
posterior fields of cilia are probably sensory
in function because of their non-motility,
shorter length, and single appearance of cil-
ia—each with a cuticular collar. Two fields
of cilia (suggested to be sensory in func-
tion) have also been described for the neril-
lid Paranerilla limicola Jouin & Swed-
mark, 1965 (Worsaae & Kristensen 2003).
The anterior field of cilia in P. limicola con-
sists of a little group of cilia and, except for
the anterior position; it is very different
from the two transverse rows of cilia ar-
ranged in a pattern found in L. saltatrix.
The posterior field in P. limicola is more
similar with 14 cilia arranged in a distinct
pattern. However, this pattern differs some
from the pattern of 20 cilia found in L. sal-
tatrix. It seems very possible that the two
systematically significant prostomial fields
of cilia (probably sensory in function) are
a common feature of nerillids, which just
demands SEM techniques to be described.

A few of the unusual characteristics of L.
saltatrix have previously been found in sin-
gle occasions in otherwise very different
nerillid species. Structures remarkably sim-
ilar to the special triangular cuticular plates
on the ventral part of the pharynx have
been described for Thalassochaetus palpi-
foliaceus Ax, 1954. A different, although
also distinct pattern of diffuse superficial
ventral glands are described for Nerillidium
renaudae Jouin, 1970. Pygidial lobes have
been described for the aberrant Nerillidium
simplex Levi, 1953 (see also Jouin 1966,
Swedmark 1959). These lobes are appar-
ently not double-lobed or cilicated as in L.
saltatrix; however, their position and
square, non-cirriform appearance is very
similar to L. saltatrix. The pygidial lobes

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

found in N. simplex have been interpreted
as modified pygidial cirri (Levi 1953,
Swedmark 1959), which may also count for
the lobes of L. saltatrix. The superficial re-
semblance with the lobes of N. simplex is
probably a matter of convergence; however,
the lobes may be functionally comparable.
The two groups of cilia on each pygidial
lobe of L. saltatrix show great resemblance
in number and length with the two ciliary
tufts found on each side of the body seg-
ment dorsal to the parapodia. Furthermore,
examined juveniles did not possess pygidial
lobes, which would be expected if the lobes
were modified pygidial cirri. These obser-
vations could mean that the pygidial lobes
are modified rudiments of a strongly re-
duced ninth body segment. However, stud-
ies with LM and SEM of L. saltatrix show
that there are no remains of chaetae or para-
podial muscles and cLSM studies show that
there are no segmental nerves posterior to
the eighth body segment. Further exami-
nation of the development of L. saltatrix is
needed to clarify the origin of the pygidial
lobes.

Acknowledgments

We thank Dr. Martin V. Sgrensen for as-
sistance with collecting of the material. The
study was financially supported by the Ber-
muda Zoological Society. This paper is
contribution # 63 from the Bermuda Bio-
diversity Project (BBP), Bermuda Aquari-
um, Natural History Museum and Zoo.

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(3):363-—367. 2004.

Neostrengeria lemaitrei, a new species of freshwater crab from
Colombia (Crustacea: Decapoda: Pseudothelphusidae), and the
vertical distribution of the genus

Martha R. Campos

Universidad Nacional de Colombia, Instituto de Ciencias Naturales, Apartado Aéreo 103698,
Bogota, Colombia, S. A. e-mail: mhrocha@ciencias.unal.edu.co

Abstract.—A new species of the genus Neostrengeria Pretzmann, 1965, N.
lemaitrei from Magdalena Valley, Cundinamarca Department, is described. The
genus is endemic to the Eastern Andes of Colombia, at altitudes ranging from
300 to 300 m above sea level. With the addition of N. lemaitrei the total number
of species rises to 21. This new species, like all others in Neostrengeria, is
distinguished primarily by the morphology of the first male gonopod, partic-
ularly by the form of lateral and accessory lobes, and the shape of the apex.

The genus Neostrengeria Pretzmann,
1965, comprises 21 species of freshwater
crabs that inhabit mountain springs and
streams on the slopes and high plain of the
Eastern Andes in Colombia (2° to 9°40'N,
73° to 74°50'W), at altitudes ranging from
300 to 3000 m above sea level (Campos
1994).

The taxonomy of Neostrengeria was re-
viewed by Rodriguez (1982), with follow
up studies by Campos (1992, 1994, 2000).
Campos & Lemaitre (1998) presented a key
for the identification of the species based
on the morphology of the male first gono-
pod. The distribution of the genus has been
discussed by Campos & Rodriguez (1985),
and Campos (1992, 1994). The present new
species was found in the Magdalena Valley,
at altitude of 720 m above sea level.

The general carapace morphology of
Neostrengeria species is very similar. The
species are characterized primarily by the
shape of the first male gonopod which has
a distinct lateral lobe generally divided in
two halves forming an accessory lobe. The
form of the gonopod’s apex is also variable
according to the species, and can be oval,
oblong, or expanded into a projection.

The terminology used for the different

processes of the gonopod is that established
by Smalley (1964), Rodriguez (1982) and
Campos & Lemaitre (1998). The material is
deposited in Museo de Historia Natural, In-
stituto de Ciencias Naturales, Universidad
Nacional de Colombia, Bogota (ICN-
MHN). The abbreviations cb and cl, re-
ported as cl X cb, indicate carapace breadth
and carapace length, respectively. Color no-
menclature follows Smithe (1975).

Family Pseudothelphusidae Rathbun, 1893
Tribe Hypolobocerini Pretzmann, 1971
Genus Neostrengeria Pretzmann, 1965
Neostrengeria lemaitrei, new species

Fig. |

Holotype.—Agua Blanca stream, Vereda
Lamal, Inspecci6n Guadualito, Municipio
Yacopi, Cundinamarca Department, Colom-
bia, 720 m alt., 4 Nov 1995, leg. M. R.
Campos: | male, 13.9 X 23.6 mm, ICN-
MHN-CR 1991.

Paratypes.—Same locality data as holo-
type: 5 males, size range 8.1 X 12.9 mm,
to 12.5 X 20.0 mm, 4 females, size range
9.4 X 14.7 mm, to 7.4 X 11.2 mm, ICN-
MHN-CR 1533.

Type locality.—Agua Blanca stream,
Vereda Lamal, Inspeccion Guadualito, Mu-

364 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Neostrengeria lemaitrei, new species, male holotype, ICN-MHN-CR 1991. A, left first gonopod,
caudal view; B, same, lateral view; C, same, cephalic view; D, same, mesial view; E, same, apex, distal view;
E right carapace half, dorsal view; G, left opening of efferent branchial channel, external view; H, left third
maxilliped, external view. 1, lateral lobe; 2, accessory lobe; 3, cephalic expansion; 4, mesocaudal projection of
spermatic channel.

VOLUME 117, NUMBER 3

nicipio Yacopi, Cundinamarca Depatment,
Colombia, 720 m alt.
Diagnosis.—Carapace without median
groove; front lacking distinct upper border.
Third maxilliped with exognath 0.67 times
length of ischium. First male gonopod with
lateral lobe semicircular distally, proximally
narrow, with external margin concave; ac-
cessory lobe elongated, semi-acute distally,
forming excavated ridge on caudal surface;
accessory lobe as long as lateral lobe. Apex
outline oval with expansion projected ce-
phalically into prominent, acute spine.
Description of holotype.—Carapace (Fig.
1F) with cervical groove straight, shallow,
ending some distance from lateral margin.
Anterolateral margin lacking depression be-
hind external orbital angle. Lateral margin
with series of approximately 15 papilliform
teeth. Postfrontal lobes oval, high, indicated
anteriorly by 2 transverse depressions. Me-
dian groove lacking. Front without distinct
upper border, frontal area sloping down-
wards, slightly bilobed in dorsal view, low-
er margin visible in dorsal view, strongly
sinuous in frontal view. Dorsal surface of
carapace smooth, covered by small papillae,
regions well demarcated. Third maxilliped
with distal half of external margin of merus
rounded, exognath 0.67 times length of is-
chium (Fig. 1H). Orifice of efferent bran-
chial channel open, irregularly ovate (Fig.
1G). First pereiopods heterochelous; palm
of larger chela strongly swollen, fingers
slight gaping when closed, smaller chela
slight swollen, fingers not gaping when
closed. Walking legs (pereiopods 2-5) slen-
der, but not prominently elongated (total
length 1.10 times the breath of carapace).
First male gonopod wide in caudal view;
mesial side forming convex expansion with
deep subdistal notch; caudal margin wide
with excavated surface, festooned (Fig. 1A,
D); lateral lobe wide, semicircular distally,
proximally narrow with external side con-
cave, separated from accessory lobe by
deep notch (Fig. 1A—D); accessory lobe
elongated, semi-acute distally, forming ex-
cavated ridge, covered with diminute papil-

365

lae and row of spinules on external border
on caudal surface; accessory lobe as long
as lateral lobe (Fig. 1A, C); apex outline
oval in distal view with expansion projected
cephalically into prominent, acute spine;
mesial lobe subtriangular; mesocaudal pro-
jection of spermatic channel with bifid tip;
spermatic channel with conspicuous rows
of spinules; proximal cephalic border with
two setae (Fig. 1C, D, E); conspicuous se-
tae along outline of prominent basal round-
ed lobe, and a patch of setae on caudal sur-
face (Fig. 1A).

Color.—The holotype, preserved in al-
cohol, is brown-olive (near 129, Dark
Brownish Olive) on the dorsal side of the
carapace. The dorsal and ventral surfaces of
the chelae and the walking legs are brown
(near 223, Raw Umber). The ventral sur-
face of the carapace is beige (near 92, Pale
Horn Color).

Habitat.—The specimens were collected
in shaded, moist banks of springs and
streams. They were found in soft mud, un-
der rocks.

Etymology.—The species is named in
honor of Colombian scientist Dr. Rafael Le-
maitre, who has dedicated his life to study-
ing Crustaceans. This species is not only a
recognition of Rafael’s contributions to sci-
ence, but to the stimulus he has provided to
a new generation of up and coming Col-
ombian scientists.

Remarks.—A comparison of both de-
scriptions and material of other species of
the genus with that of this new species re-
vealed that it is most similar to Neostren-
geria gilberti Campos, 1992. The main dis-
tinguishing feature between both species is
the form of the first gonopod. The male first
gonopod of N. gilberti has been described
and illustrated by Campos (1992: 542, fig.
2). In this new species, the mesial side of
the gonopod is convex expanded with deep
subdistal notch, whereas in N. gilberti it is
rounded basally, straight tapering distally
without subdistal notch. The lateral lobe in
N. gilberti is rounded distally with the prox-
imal external side straight, whereas in UN.

366

Table 1.—Vertical distribution of the Neostrengeria
species.

Meters
above sea
Species level
Neostrengeria appressa Campos, 1992 1125-1900
N. aspera Campos, 1992 1600
N. binderi Campos, 2000 470
N. botti Rodriguez & Tiirkay, 1978 1350-2600
N. boyacensis Rodriguez, 1980 2350-3000
N. charalensis Campos & Rodriguez, 1450-2150
1985
N. gilberti Campos, 1992 950-1250
N. guenteri (Pretzmann, 1965) 500-1575
N. lasallei Rodriguez, 1980 1110-2150
N. lemaitrei, new species 720
N. libradensis Rodriguez, 1980 1200
N. lindigiana (Rathbun, 1897) 1800-2350
N. lobulata Campos, 1992 1700-2350
N. macarenae Campos, 1992 300-500
N. macropa (H. Milne Edwards 1853) 2200-2900
N. monterrodendoensis Bott, 1967 1320-1500
N. niceforoi (Schmitt, 1969) 1000-1750
N. perijaensis Campos & Lemaitre, 1998 1270-1800
N. sketi Rodriguez, 1985 1800
N. tencalanensis Campos, 1992 1600-2400
N. tonensis Campos, 1992 1600-2400

lemaitrei it 1s distally semicircular, and
proximally narrow with the external side
concave. The apex outline in N. gilberti is
oblong in distal view with a mesially di-
rected semi-acute spine; the mesocaudal
projection of spermatic channel is awl-
shaped with a distal spinule on the inner
side. In contrast, in N. lemaitrei, the apex
outline is oval in distal view with the ex-
pansion projected cephalically into a prom-
inent, acute spine, and the mesocaudal pro-
jection of spermatic channel has the tip bi-
fid.

Distribution of Neostrengeria species

The distribution of the species of Neos-
trengeria comprises both slopes and the
high plain of the Eastern Cordillera of Co-
lombia that encompasses the Magdalena,
Orinoco and Catatumbo basins. It is limited
to the north by Serrania de Perija, and to
the south by Serrania de La Macarena (2°
to 9°40'N, 73° to 74°50'W), (H. Milne Ed-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

wards 1853; Rathbun 1897; Pretzmann
1965; Bott 1967; Schmitt 1969; Rodriguez
& Tiirkay 1978; Rodriguez 1980, 1982,
1985; Campos & Rodriguez 1985; Campos
1992, 1994, 2000; Campos & Lemaitre
1998).

Based on the collected material, the ver-
tical distribution of the species of the genus
Neostrengeria (Table 1) ranges from 300 m
to 3000 m. Neostrengeria botti has the
greatest altitude range of between 1350 and
2600 m. The species that exhibit a range of
between 300 and 1000 m are N. binderi, N.
macarenae and N. lemaitrei, new species.
Most of the species are distributed between
1000 and 2400 m. The highest altitude,
3000 m, is reached by N. boyacensis.

Acknowledgments

I am especially grateful to the referees
for providing useful comments of the paper.
I also indebted to David H. Campos for crit-
ically reading the manuscript. The illustra-
tion was prepared by Juan C. Pinzon.

Literature Cited

Bott, R. 1967. Fluss-krabben aus dem westlichen Sii-
damerika.—Senckenbergiana Biologie 48(5/6):
365-372.

Campos, M. R. 1992. New species of fresh-water crabs
of the genus Neostrengeria Pretzmann, 1965
(Crustacea: Decapoda: Pseudothelphusidae)
from Colombia.—Proceedings of the Biological
Society of Washington 105:540—554.

. 1994. Diversidad en Colombia de los cangre-

jos del género Neostrengeria.—Academica Co-

lombiana de Ciencias Exactas Fisicas y Natur

ales. Col. Jorge Alvarez Lleras No. 5:1—143.

. 2000. Neostrengeria binderi, a new species of
pseudothelphusid crab from the eastern Andes
of Colombia (Crustacea: Decapoda: Brachyu-
ra).—Proceedings of the Biological Society of
Washington 113:401—405.

Campos, M. R., & R. Lemaitre. 1998. A new fresh-
water crab of the genus Neostrengeria Pretz-
mann, 1965, from Colombia (Crustacea: Deca-
poda: Brachyura: Pseudothelphusidae) with a
key to the species of the genus.—Proceedings
of the Biological Society of Washington 111:
899-907.

Campos, M. R., & G. Rodriguez. 1985. A new species
of Neostrengeria (Crustacea: Decapoda: Pseu-

VOLUME 117, NUMBER 3

dothelphusidae) with notes on geographical dis-
tribution of the genus.—Proceedings of the Bi-
ological Society of Washington 98:718—727.

Milne-Edwards, H. 1853. Observations sur les affini-
tiés zoologiques et la classification naturelle des
crustacés.—Annales des Sciences Naturelles,
Zoologie 20:163—228.

Pretzmann, G. 1965. Vorlaufiger Bericht tiber die Fam-
ilie Pseudothelphusidae.—Anzeiger der Oster-
reichischen Akademie der Wissenschaften
Mathematische Naturwissenschaftliche Klasse
(1),1:1-10.

. 1971. Fortschritte in der Klassifizierung der
Pseudothelphusidae.—Anzeiger der Mathema-
tisch Naturwissenschaftliche der Osterreichisch-
en Akademie der Wissenschaften (1)179:14—24.

Rathbun, M. 1893. Descriptions of new species of
American freshwater crabs.—Proceedings of
the United States Naitonal Museum 16(959):
6459-661, pl. 73-77.

. 1897. Descriptions de nouvelles espéces de

crabes d’eau douce appartenant aux collections

du Muséum d’ Histoire naturelle de Paris.—Bul-
letin du Muséum National d’ Histoire Naturelle,

Paris 3(2):58-61.

367

Rodriguez, G. 1980. Description préliminaire de qu-
elque espéces et genres nouveaux de crabes
d’eau douce de 1’Amérique tropicale (Crusta-
cea, Decapoda, Pseudothelphusidae).—Bulletin
du Muséum National d’ Histoire Naturelle, Paris
4(3):889—-894.

. 1982. Les crabes d’eau douce d’ Amérique.

Famille des Pseudothelphusidae.—Faune Trop-

icale 22:1—223.

. 1985. A new cavernicolous crab (Crustacea,
Decapoda, Pseudothelphusidae) from Colom-
bia.—Bioloski vestnik, Ljubljana 33(2):73—80

Rodriguez, G., & M. Tiirkay. 1978. Der generische
Status einiger Kolombianischer Stisswasser-
krabben.—Senckenbergiana Biologica 59:297—
306.

Schmitt, W. 1969. Colombian freshwater crab notes.—
Proceedings of the Biological Society of Wash-
ington 82:93-112.

Smalley, A. 1964. A terminology for the gonopods of
the American river crabs.—Systematic Zoology
13:28-31.

Smithe, E B. 1975. Naturalist’s color guide: The
American Museum of Natural History. New
York, part 1: unnumbered pages.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(3):368—376. 2004.

A new species of Agostocaris (Caridea: Agostocarididae) from
Acklins Island, Bahamas

Fernando Alvarez, José Luis Villalobos, and Thomas M. Iliffe

(FDA, JLV) Colleccion Nacional de Crustaceos, Instituto de Biologia,
Universidad Nacional Aut6noma de México, Apartado Postal 70-153,
México 04510 D.E, México, e-mail: falvarez@servidor.unam.mix;
(TMI) Department of Marine Biology, Texas A&M University at Galveston, Galveston,
Texas 77553-1675, U.S.A.

Abstract.—The new bresilioid shrimp Agostocaris acklinsensis is described
from an anchialine cave in Acklins Island, Bahamas. This is the third species
described in the genus. The new species is characterized by having small ex-
opods on the third and fourth pereiopods, one spine on the ischium of the fifth
pereiopod, and an outer ramus of the uropods with one distolateral spine. A
key to the species of Agostocaris is provided.

The family Agostocarididae Hart &
Manning, 1986, was created to accommo-
date Agostocaris williamsi, from Grand Ba-
hama and Turks and Caicos, a species that
appeared to be morphologically similar to
some species in the Atyidae De Haan, 1849,
and the Bresiliidae Calman, 1896, but had
a distinct morphology of the propodus and
dactylus of the first two pereiopods. Ken-
sley (1988) described a second species from
Cozumel, Mexico, Agostocaris bozanici,
which exhibits the same unique pereiopodal
morphology, placing it also in the Agosto-
carididae. Holthuis (1993) synonymized the
Agostocariidae with the Bresiliidae. How-
ever, Martin & Davis (2001) have proposed
to recognize the family Agostocarididae
within the superfamily Bresilioidea Cal-
man, 1896, where a hetereogeneous assem-
blage of forms are included in five families.

At best, as pointed out by Kensley
(1988), the relationships of Agostocaris are
unclear. The particular articulation of the
proprodus of the first pair of legs, and the
morphology of the chela of the second pair
of legs, are unique characters not shared by
any other genus in the Bresilioidea. With
respect to the diagnosis of Agostocaris,
with the new species described herein, the

range of variation in taxonomically impor-
tant characters increases, making it neces-
sary to provide a new diagnosis for the ge-
nus.

Materials and Methods

Specimens of the new Agostocaris de-
scribed herein were collected during an ex-
pedition to Crooked and Acklins Islands,
Bahamas, in January 1999. The new spe-
cies was captured in Jumby Hole Cave
(22°29.275'N, 73°53.501’W), Snug Corner,
Acklins Island, Bahamas, 11 January 1999
(Fig. 1). This cave is located about 250 m
inland from the west side of the island fac-
ing the shallow water Bight of Acklins. It
is actually a complex of closely associated
caves that were mined for guano in the past.
More than 3 m of soil and guano were re-
moved from pits within dry portions of the
cave. One of these caves contains a 20 m
diameter, shallow (30 to 50 cm deep) pool.
Sediments in the pool consist of a thick lay-
er of guano from a bat roost located directly
above. Tidal range in the pool appeared to
be about 30 cm. This pool is in total dark-
ness but is close to 4 or more entrances on
all sides. Salinity was measured at 32.5%o

VOLUME 117, NUMBER 3 369

GULF OF MEXICO

Crooked
Island

Bight of
Acklins

Acklins
Island

Liza Bay Cave

Fig. 1. Map showing the location of the type locality of Agostocaris acklinsensis, Acklins Island in the
Bahamas.

370

with a refractometer and water temperature
was 25.5°C. Specimens of Agostocaris were
observed walking across the surface of
rocks and the guano bottom in 50 cm depth.
They were collected by hand using glass
vials. Other invertebrates collected from the
cave pools included copepods, archiannelid
and other polychaetes, mites and the shrimp
Barbouria cubensis (von Martens, 1872)
(Hippolytidae).

The specimens representing the new spe-
cies are deposited in the Coleccion Nacion-
al de Crustaceos (CNCR), Instituto de Biol-
ogia, Universidad Nacional Autonoma de
México. Other abbreviations used are: cl,
postorbital carapace length, and tl, total
length.

Results
Agostocaris Hart & Manning, 1986

Diagnosis.—Rostrum well developed,
with or without dorsal teeth. Carapace lack-
ing spines and grooves. Eyes reduced,
fused, without pigment or weakly pigment-
ed. Antennal scale with lateral spine. First
maxilliped with lash on exopod. Second
maxilliped with terminal segments serial.
Pleurobranchs on all pereiopods or on pe-
reiopods 2—5. First and second pereiopods
chelate, first pair heavier than second one.
First pereiopod with propodus articulating
with carpus at one third of its length. Sec-
ond pereiopod with carpus undivided; dac-
tylus digitiform, heavier and longer than
propodus, both fingers without teeth or
spines. Telson with 4—5 pairs of dorsal
spines, posterior margin with variable num-
ber of spines.

Agostocaris acklinsensis, new species
Figs. 2—4

Material examined.—Holotype, female,
cl 7.3 mm, tl 21.5 mm; 11 January 1999;
Jumby Hole Cave, Snug Corner, Acklins Is-
land, Bahamas; collected by T. M. Iliffe;
CNCR 19601. Paratypes, 8 females, cl 4.0—
8.0 mm, tl 13.6—21.7 mm; same locality,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

date and collector as holotype; CNCR
19602.

Description.—Carapace globose, smooth,
devoid of spines. Rostrum laterally com-
pressed, triangular, ending in sharp tip,
reaching distal end of first antennular seg-
ment; without teeth in mature individuals,
with three dorsal teeth with alternating setae
in juveniles (Fig. 2a, b). Carapace without
grooves, inferior margin of orbit and pter-
ygostomian angle slightly produced (Fig.
2a), pterygostomian regions produced lat-
erally (Fig. 2b).

Abdomen smooth, somites 1—2 with
rounded pleura, somites 3—5 with posterior
angle of pleura subacute, sixth somite with
posterior margin sinuous at insertions of
telson and uropods. Telson 2.5 times as
long as its basal width, tapering distally,
distal width less than half of basal width;
bearing four pairs of movable spines on
dorsal surface, spines located on distal two
thirds of dorsal surface; posterior margin
rounded, bearing 9 spines, second pair from
external one longest (Fig. 4g).

Eyes pigmented, fused, forming part of a
single plate, peduncle and cornea not dis-
cernible, projected dorsally (Fig. 2c). An-
tennule with first segment as long as seg-
ments 2 and 3 combined; stylocerite acute,
reaching distal margin of first segment (Fig.
4e). Antennal scale 1.8 times as long as
wide, laterodistal tooth short not exceeding
distal margin of blade (Fig. 4f), flagellum
1.25 times total length (Fig. 2a).

Mandible with stout 2-segmented palp,
incisor process with six distal teeth, molar
process conical, sharp distal end (Fig. 2d).
Both mandibles approximately symmetri-
cal. First maxilla with distal lacinia oval
shaped, bearing three rows of short, thick
setae on mesial surface; proximal lacinia
with single row of short, thick setae on dis-
tomesial margin; palp bearing one distal,
long setae and two subdistal short ones on
internal margin (Fig. 2e). Second maxilla
with scaphognathite approximately rectan-
gular distally, subtriangular proximally; dis-
tal margin with long, plumose setae; lateral

VOLUME 117, NUMBER 3 37

Fig. 2. Agostocaris acklinsensis, new species, a female holotype, b-f female paratype: a, total lateral view:
b, carapace, dorsal view; c, dorsal view of eyes, carapace removed; d, mandible: e, first maxilla; f, first maxil-
liped. Scale bar represent: a—c, f, 1 mm; d—e, 0.5 mm.

372 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Agostocaris acklinsensis, new species, female paratype: a, second maxilla; b, second maxilliped; c,
third maxilliped; d, first pereiopod; e, detail of propodus and dactylus of first pereiopod; f, second pereiopod.
Scale bars represent: a—d, f, 1 mm; e, 0.5 mm.

VOLUME 117, NUMBER 3 373

Fig. 4. Agostocaris acklinsensis, new species, female paratype: a, third pereiopod; b, detail of proximal
segments of third pereiopod; c, fourth pereiopod; d, fifth pereiopod; e, antennule; f, antenna; g; telson and
uropods, left side omitted; h, first pleopod; i, second pleopod. Scale bars represent 1 mm.

374

margin with short plumose setae; internal
margin with long simple setae, increasing
in length distally, almost as long as sca-
phognathite; palp digitiform, devoid of se-
tae; distal endite trapezoidal, middle and
proximal endites approximately rectangular,
all three bearing simple setae on distal mar-
gins (Fig. 3a).

First maxilliped with triangular endite
bearing marginal setae; palp digitiform,
with apical tuft of setae; exopod elongated,
bearing long, simple setae distally; caridean
lobe broadly rounded, with submarginal
row of short setae and long plumose setae
along margin; epipod bilobed, both lobes
trapezoidal, distal one smaller, devoid of se-
tae (Fig. 2f). Second maxilliped with en-
dopod pediform, 4-segmented, with contin-
uous row of setae along margin; exopod
slender, bearing long simple setae distally;
epipod simple, flat, rounded (Fig. 3b). Third
maxilliped with endopod 4-segmented,
bearing setae on mesial margin; exopod as
long as first segment of endopod, with dis-
tal tuft of long setae; epipod digitiform, less
than half the length of exopod; arthro-
branches present (Fig. 3c).

First pereiopod with ischium and merus
of about same length and width, carpus
wider proximally, propodus articulating
with carpus at one third of its length, palm
as long as fingers, cutting edges of both fin-
gers with minute sharp teeth, dactylus with
long setae arising from proximal half teeth
(Fig. 3d); exopod as long as ischium and
merus combined, with apical tuft of long
setae; arthrobranch and pleurobranch pre-
sent (Fig. 3d). Second pereiopod longer
than first one, with merus slightly shorter
than ischium, carpus becoming wider dis-
tally and as long as merus, propodus with
palm shorter than fixed finger, dactylus
heavier and longer than fixed finger; exo-
pod shorter than ischium and merus com-
bined, bearing apical tuft of long setae; ar-
throbranch and pleurobranch present (Fig.
3f). Third pereiopod with ischium with two
spines, merus the longest segment, carpus
and propodus of about the same length,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

dactylus with corneous sharp tip and four
smaller teeth on internal surface, arthro-
branch and pleurobranch present, finger-
like exopod arising from basis (Fig. 4a, b).
Fourth pereiopod with ischium with two
spines, merus the longest segment, carpus
and propodus of about the same length,
dactylus with corneous sharp tip and three
smaller teeth on internal surface, arthro-
branch and pleurobranch present, finger-
like exopod arising from basis (Fig. 4c).
Fifth pereiopod with ischium with one
spine, propodus the longest segment, dac-
tylus with corneous sharp tip and eight
smaller teeth on internal surface, arthro-
branch and pleurobranch present (Fig. 4d).

First pleopod with exopod setose, endo-
pod devoid of setae, one third the length of
exopod (Fig. 4h). Second pleopod with en-
dopod and exopod setose, appendix interna
slender more than half the length of endo-
pod (Fig. 41).

Uropods with external ramus bearing one
distolateral movable spine, distal margin
broadly rounded, with long plumose setae
on distal and internal margins. Internal ra-
mus bearing marginal long plumose setae
except on proximal third, distal margin sub-
acute (Fig. 4g).

Etymology.—The specific name is de-
rived from “‘Acklins”, the name of the Ba-
hamian island where the new species was
captured.

Key to the species of Agostocaris

1. First maxilliped with palp 2-segmented,
ischium of fifth pereiopod devoid of
spines, outer ramus of uropods devoid of
distolateral spines, .. Agostocaris williamsi

— First maxilliped with palp unsegmented,

ischium of fifth pereiopod with spines,

outer ramus of uropods with spines, out-

er ramus of uropods with distolateral

SPINES ce Paes see Sime aoe eee eee et 2
. Ischium of fifth pereiopod with two

spines, outer ramus of uropod with two

distolateral spines, telson with five pairs

of dorsal spines

VOLUME 117, NUMBER 3

— Ischium of fifth pereiopod with one
spine, outer ramus of uropod with one
distolateral spine, telson with four pairs
of dorsal spines Agostocaris acklinsensis

Remarks.—Agostocaris acklinsensis can
be easily distinguished from the other two
known species in the genus by the presence
of: exopods on the third and fourth pereio-
pods, a fifth pereiopod with one spine on
the ischium and one distolateral movable
spine on the outer ramus of the uropods.
Other taxonomically important characters
vary among the three species. A second
maxilla with a palp devoid of setae and an
unsegmented palp of the first maxilliped
distinguish A. acklinsensis from A. william-
si, whereas the number of dorsal spines on
the telson, unpigmented eyes and two dis-
tolateral spines on the outer ramus of the
uropods seprate A. bozanici (Table 1).

Noteworthy are the eyes of Agostocaris,
which are composed of one single plate not
differentiated into peduncle and cornea.
This plate is projected outside the orbits
creating the eye-like structures, which in
the three species are pointed distally. Since
all the species of Agostocaris are cave
dwellers it is reasonable to suppose that the
cornea was lost and later the peduncle was
reduced, in such a way that the “‘eyes”’ we
see now are part of the basal plate. This
singular morphology merits further studies
on its ontogeny and functionality.

The placement of the genus Agostocaris
is a matter of controversy. Holthuis (1993),
by synonymizing Agostocarididae with the
Bresiliidae, gave more weight to characters
that are shared by many taxa in the Caridea
(mandible with palp, carpus of second legs
undivided, first two pairs of legs chelate,
first pair of legs more robust than second
one, Williams, 1984) with little resolution
among families, than to exceptional auta-
pomorphic characters such as the fused
eyes and the particular morphology of the
first two pereiopods of Agostocaris.

We agree with Martin & Davis’ (2001)
proposal of recognizing a superfamily Bre-

Table 1.—Comparison of selected characters of the three species of Agostocaris.

A. acklinsensis

A. bozanici

A. williamsi

Weakly pigmented

Without pigment

Weakly pigmented
Palp with setae

Eyes

Palp without setae

Second maxilla

Palp unsegmented Palp unsegmented

Palp 2-segmented

First maxilliped

Basis with finger-like exopod

Basis without exopod
Basis without exopod

Third pereiopod

Basis with finger-like exopod
Ischium with one spine

Fourth pereiopod
Fifth pereiopod

Ischium with two spines
Appendix interna less than half

Ischium devoid of spines

Appendix interna less than half

Appendix interna two thirds length of

Second pleopod

length of endopod
With four pairs of dorsal spines

length of endopod
With five pairs of dorsal spines

endopod
With four pairs of dorsal spines

Telson

Outer ramus without distolateral Outer ramus with two distolateral Outer ramus with one distolateral

Uropods

spine

spines

375

376

silioidea, which includes five families, and
concur with the opinion that this taxon still
represents an artificial grouping. While it is
beyond the scope of this paper to discuss
the relationships among bresilioids, it is
clear that Agostocarididae represents a dis-
tinct family that can be easily separated
from the other four bresilioid families. The
Alvinocarididae Christoffersen, 1986, and
Mirocarididae Vereshchaka, 1997, lack ex-
opods on all pereiopods, whereas the Agos-
tocarididae can have exopods on all five pe-
reiopods. The Diascididae Rathbun, 1902,
have well developed eyes with peduncle
and cornea, a dorsoventrally flattened ros-
trum and a disc-like dactylus of the first pe-
reiopod, contrasting with the fused eyes,
acuminate rostrum and typically shaped
dactylus of pereiopod 1 of the Agostocari-
didae. Finally the Bresiliidae, and the rest
of the bresilioid families, can be separated
from the Agostocarididae based on the car-
pus-propodus articulation of the first pereio-
pod which is normal in the former, being
the distal end of the carpus articulated to
the proximal end of the propodus; while in
the latter the carpus is articulated to an area
close to the middle portion of the propodus.
In addition, the chela of the second pereio-
pod in the Agostocarididae is unique in that
the digitiform dactylus is longer than the
fixed finger and lacks teeth or spines.

Acknowledgments

Collection of shrimp described herein
was part of the January 1999 Anchialine
Caves Expedition to the southern Bahamas
led by Thomas Iliffe. Other members of the
expedition included Texas A&M University
graduate students Brett Dodson and Shelley
Fetterolf. This expedition was funded by
National Science Foundation, Biotic Sur-
veys and Inventories Program award num-
ber 9870219. We thank Neil Sealey (Media
Publishing Ltd, Nassau, Bahamas), Dr.
Nancy Elliott (Sienna College) and Dr. Wil-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

liam Keegan (Florida Museum of Natural
History) for providing invaluable logistical
information on Crooked and Acklins Is-
lands. The drawings were prepared by Ro-
lando Mendoza.

Literature Cited

Calman, W. T. 1896. On deep sea Crustacea from the
south west of Ireland.—Transactions of the
Royal Irish Academy 31:1—22.

Christoffersen, M. L. 1986. Phylogenetic relationships
between Oplophoridae, Atyidae, Pasiphaeidae,
Alvinocarididae fam. n., Bresiliidae, Psalido-
popidae and Disciadidae (Crustacea Caridea
Atyoidea).—Boletim Zoologico, Universidade
do Sao Paulo 10:273-281.

De Haan, W. 1849 (1833-1850). Crustacea. Jn P. EK von
Siebold, ed., Fauna Japonica sive descriptio an-
imalium, quae in itinere per Japonium, Jussu et
auspices superiorum, qui summum in India Ba-
tava imperium tenent, suscepto, annis 1823-
1830 collegit, notis, observationibus et adum-
brationibus illustravit. Lugduni-Batavorum, 243
Pp-

Hart, C. W., Jr, & R. W. Manning. 1986. Two new
shrimps (Procarididae and Agostocarididae,
new family) from marine caves of the western
north Atlantic_—Journal of Crustacean Biology
6:408-416.

Holthuis, L. B. 1993. The Recent Genera of the Car-
idean and Stenopodidean Shrimps (Crustacea,
Decapoda). Nationaal Natuurhistorisch Muse-
um, Leiden, 328 pp.

Kensley, B. 1988. New species and records of cave
shrimps from the Yucatan Peninsula (Decapoda:
Agostocarididae and Hippolytidae).—Journal of
Crustacean Biology 8:688—699.

Martin, J. W., & G. E. Davis. 2001. An updated clas-
sification of the recent Crustacea. Natural His-
tory Museum of Los Angeles County, Science
Series 39, 124 pp.

Rathbun, M. J. 1902. Papers from the Hopkins Stan-
ford Galapagos Expedition 1898-1899. VIII.
Brachyura and Macrura.—Proceedings of the
Washington Academy of Sciences 4:275—292.

Vereshchaka, A. L. 1997. A new family for a deep-sea
caridean shrimp from North Atlantic hydrother-
mal vents.—Journal of the Marine Biological
Association of the United Kingdom 77:425—
438.

Williams, A. B. 1984. Shrimps, lobsters, and crabs of
the Atlantic coast of the eastern United States,
Maine to Florida. Smithsonian Institution Press,
550 pp.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(3):377-384. 2004.

A new species of caridean shrimp of the family Stylodactylidae from
the eastern Pacific Ocean

Mary K. Wicksten and Joel W. Martin

(MKW) Department of Biology, Texas A&M University, College Station, Texas 77843-3258,
U.S.A., e-mail: wicksten@mail.bio.tamu.edu
(JWM) Natural History Museum of Los Angeles County, 900 Exposition Boulevard,
Los Angeles, California 90007, U.S.A., e-mail: jmartin@nhm.org

Abstract.—Four specimens of shrimp of the family Stylodactylidae were
collected at two stations off Baja California, Mexico, and California, U.S.A.
These are the first specimens of the family reported from the eastern Pacific.
The shrimp are described as a new species, Bathystylodactylus echinus. The
species can be recognized by the following features: rostrum straight, much
longer than the carapace, bearing at least 23—27 dorsal and 18—25 ventral
spines; eye small and without pigment, stylocerite slender and not reaching
middle of first segment of antennular peduncle, carapace without prominent
posterior dorsal hump, body set with minute spinules, posterior pereopods con-
siderably longer than anterior two pair, slender and lacking fringe of setae.

Shrimp of the family Stylodactylidae are
recognized by their peculiar first and sec-
ond pereopods, which end in elongate but
nearly equal fingers with setae on the cut-
ting edges. These pereopods and the max-
illipeds are densely setose. Species of the
family are widely distributed from tropical
to temperate regions (e.g., Cleva 1990a), al-
though most of the species described to date
have come from the tropical Indo-Pacific
(Chace 1983; Cleva 1990b, 1994, 1997;
Okuno and Tachikawa 2000). Chace (1983)
and Cleva (1994) reviewed the members of
the family, described new species, and pro-
vided keys. Hanamura and Takeda (1996)
described an additional genus, Bathystylo-
dactylus, for a new species (B. inflatus)
from off Taiwan (and for the former Sty-
lodactylus bathyalis from the Coral Sea),
bringing to 5 the number of recognized
genera in the family (Stylodactylus, Neos-
tylodactylus, Parastylodactylus, Stylodac-
tyloides, and Bathystylodactylus). There
have been no previous reports of the family
in the eastern Pacific Ocean.

While sorting specimens in the Benthic

Invertebrate Collection of Scripps Institu-
tion of Oceanography, we found four spec-
imens of shrimp of this family from three
stations taken off California, U.S.A., and
Baja California, Mexico. The specimens in-
clude both males and females. We com-
pared these specimens with specimens of
Stylodactylus rectirostris in the collections
of Texas A&M University (catalog number
2-7212, Oregon station 5916) and with
published descriptions of other species in
the family. The specimens represent an un-
known species of Bathystylodactylus, de-
scribed herein.

Systematic Account

Bathystylodactylus echinus, new species
Figs. 1-5

Holotype: Male, carapace length (CL)
32.7. Basin off Magdalena Bay, Baja Cali-
fornia, Mexico (24°35’N, 113°25’W),
3563-3621 m, 6-foot Sigsbee trawl, 24
June 1965, ship Horizon sta. MV65-I-38,
Carl Hubbs, collector; Scripps Institution of

378

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1.
Scale bar = 10.0 mm.

Oceanography (SIO) catalog number
C3188.

Paratypes: Male, CL 41.4, same station
as holotype, LACM CR 1965-349.1 (Nat-
ural History Museum of Los Angeles
County). Female, CL 29.7. Basin off Mag-
dalena Bay, Baja California, Mexico
(24°23'N, 113°17’W), 3427-3621 m, 45-
foot otter trawl, 25 June 1965, ship Horizon
2 sta., MV65-I-39, C. Hubbs, SIO cat. no.
3203. Female CL 27.6 Patton Escarpment
(32°25'N, 120°40'W), 3689-3630 m, 40-
foot otter trawl, 7 Feb. 1981, ship New Ho-
rizon sta. 133, collector S. Luke, SIO cat.
no. C10324.

Description: Rostrum (Figs. 1, 2B, C)
nearly straight, nearly 2X length of cara-
pace but broken in all specimens, with 23—
27 movable dorsal and 18—25 ventral
spines; series of 7—9 minute spinules on
carapace just posterior to rostrum proper,
long setae along distal ventrolateral surface.
Carapace (Fig. 2A) with hepatic depression
well delineated. Antennal and branchioste-
gal spines short but obvious, antennal spine

Bathystylodactylus echinus, new species, male holotype, Scripps Institution of Oceanography C3188.

located ventral to suborbital angle. Lateral
surface of carapace inflated over branchial
region, suprabranchial carina curved. Area
posterior to eye and antennal origin slightly
depressed. Anterior regions of carapace set
with small, simple, movable spinules, pos-
terior regions punctate or with few spinules.
Abdomen (Fig. 1) with small spinules on
dorsal and lateral surfaces, somites one and
two rounded dorsally, somite three weakly
carinate dorsally; somite four rounded to
weakly carinate, with or without shallow
depression interrupting dorsal carina; pleura
of somites rounded, those of somites four
and five (Fig. 5B) each with sharp poster-
oventral spine; one specimen with minute
spine on pleuron of somite three. Telson
(Fig. 5C, E) 8X longer than wide, tapering
to apex, with 11—13 pairs of dorsolateral
spines located on weak ridges and numer-
ous small spinules; two mesial spines flank-
ing apex on either side. (Apex of telson pre-
served in only one specimen; observed
asymmetry may be due to injury.)

VOLUME 117, NUMBER 3

37/9)

Fig. 2. Bathystylodactylus echinus, new species, male holotype, carapace and rostrum. A, carapace and eye,
lateral view. B, rostrum (attached at area of dashed lines in A and illustrated at same scale as A). C, higher
magnification of region of rostrum shown in B and denoted by arrows. Scale bar = 10.0 mm A, B; 2.5 mm C.

Eyes (Figs. 1, 2A, 5A) reduced, cornea
without trace of pigment.

Antennular peduncle (Fig. 5A) elongate.
Stylocerite slender, not reaching middle of
first segment. First and second segments
subequal in length, third segment very
short. Antennal scale (scaphocerite) more
than 4X long as broad, outer margin slight-
ly concave, with microscopic spinules, not
reaching end of second segment of anten-

nular peduncle, blade exceeding distolateral
spine. Carpocerite covered by minute spi-
nules, reaching second segment of anten-
nular peduncle. Basicerite bearing strong
lateral spine.

Mandible (Fig. 3A) with molar process
bearing teeth in the following configuration:
2 small, one large, 4 small and large blunt
process; stout, 2-jointed palp present. First
maxilla (Fig. 3B) with distal endite broad

380 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Bathystylodactylus echinus, new species, male paratype (LACM CR 1965-349.1). A, mandible; B,
first maxilla; C, second maxilla; D, inner surface of second maxilla slightly enlarged and showing palp:; E, first
maxilliped. Scale bar = 5.0 mm A, B, D; 10.0 mm C, E.

and with stiff mesial setae; proximal endite 2 more proximal endites; long palp ending
curved inward and ending in brush of setae; in 5 setae, scaphognathite with anterior half
palp ending in long setae and having tufts rounded, posterior half slender and curved
of setae on lateral surface. Second maxilla mesially, bearing long setae. First maxilli-
(Fig. 3C, D) with distal endite larger than ped (Fig. 3E) with long distal and short

VOLUME 117, NUMBER 3

proximal endites; palp reaching 3/4 length
of distal endite and ending in tuft of setae;
exopod with lash, well developed caridean
lobe and deeply bilobed epipod.

Second maxilliped (Fig. 4A) much larger
than inner mouthparts, with exopod having
lash and reaching end of basal segments;
podobranch and epipod present; basal seg-
ments fringed with stiff curved setae; an-
tepenultimate segment short, with long sim-
ple setae on flexor margin at articulation
with basal segments; penultimate segment
with fringe of long setae on flexor margin;
two terminal segments; that on flexor side
longer than one on extensor side, both
fringed with long setae. Third maxilliped
(Fig. 4B) setose, with arthrobranch but
without exopod, exceeding antennular pe-
duncle by about length of distal segment.
Ultimate segment longest, with dense setae
on flexor side. Penultimate segment with
long, pinnately branched setae. Antepenul-
timate segment with both long and short se-
tae.

Pereopods all lacking exopods or epi-
pods. First pereopod (Fig. 4B, C) with en-
tire flexor surface fringed with long setae,
merus longer than carpus, propodus about
equal in length to carpus, ending in elon-
gate chela (Fig. 4C); fingers simple, with
long setae and shorter spine-like setae along
cutting edges. Second pereopod similar to
first. Third to fifth pereopods (Fig. 1) elon-
gate, with few scattered setae; merus of
third pereopod with 8—10 spines on flexor
and lateral surfaces; merus of fourth pereo-
pod with 15, merus of fifth pereopod 14;
carpus shorter than merus; propodus broken
and dactylus missing in all specimens.

All pleopods densely setose. First pleo-
pod shorter than second to fifth pleopods.
Male second pleopod (Fig. 4D, E) with ap-
pendix interna and appendix masculina, ap-
pendix masculina reaching nearly % length
of appendix interna, with apex notched and
bearing small hooks.

Lateral branch of uropod with spinules,
margin nearly straight, two small teeth by

381

suture (Fig. 5D). Uropods shorter than tel-
son.

Etymology.—The specific name is de-
rived from the Greek word for spiny.

Remarks.—The new species can be as-
signed to the genus Bathystylodactylus ac-
cording to the features given by Hanamura
and Takeda (1996). The new species bears
a well-developed and two-jointed mandib-
ular palp. Both sexes bear well-developed
arthrobranchs on the four anterior pereo-
podal somites. There is no supraorbital
spine. The stylocerite falls far short of the
mesiodistal margin of the basal segment.
There are no fringes of setae on pereopods
3—5, as there are in Stylodactylus rectiros-
tris and other species of Stylodactylus. Han-
amura and Takeda (1996) mentioned that
the third to fifth abdominal somites were
““weakly carinate”’ dorsally. In our speci-
mens, only somite three is consistently
weakly carinate. The posterior three pereo-
pods definitely are longer than the anterior
two in the new species, but due to breakage,
their relative lengths to each other cannot
be determined.

Two species of Bathystlyodactylus have
been described previously: B. bathyalis
(Cleva, 1994), from the Coral Sea (as Sty-
lodactylus bathyalis); and B. inflatus Han-
amura and Takeda (1996), from off Taiwan
(Hanamura and Takeda 1996). Bathystylo-
dactylus echinus can be distinguished from
the former by its curved rostrum and char-
acteristic sharp spine on the ventral margin
of abdominal pleuron three. Like Bathys-
tylodactylus inflatus, B. echinus has a
straight rostrum with numerous dorsal and
ventral spines. The pleura of the fourth and
fifth abdominal somites each bear a poster-
oventral spine. However, in B. inflatus the
carapace has a marked wide elevation near
the posterodorsal margin. This is not pre-
sent in B. echinus. The shape of the supra-
branchial carina is more sinuous in B. infla-
tus than in B. echinus. In B. inflatus, there
are 11 spinules on the carapace posterior to
the rostrum; in B. echinus, there are 8—9. In
B. inflatus, there are 9—10 dorsal rostral

382

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

~~
SSS
SSS

5
SSS

e e Pi tiy a Fi bia )
PER miei 1] \ us

SNARK

Fig. 4.

Bathylstylodactylus echinus, new species, male paratype (LACM CR 1965-349.1) (A) and holotype
(SIO C3188) (B-E). A, second maxilliped (paratype). B, right third maxilliped (upper appendage) and first

pereopod (holotype). C, higher magnification of chela of first pereopod (tips of fingers broken). D, second
pleopod (holotype). E, higher magnification of appendix interna and appendix masculina (arrow from D). Scale
bar = 10.0 mm A, E, E; 7.5 mm B, C.

VOLUME 117, NUMBER 3 383

TD;
7

ZS
>

LEE

\ )
y] K

—

Fig. 5. Bathystylodactylus echinus, new species, male paratype (A) and holotype (B—E). An, antennule,
antenna, and eye (e), right side, dorsal view, male paratype (LACM CR 1965-349.1). sc = scaphocerite; st =
stylocerite. B, lateral view of abdominal somite 6 plus portions of the telson, uropods, and pleurae of somites
4 and 5, holotype. C, telson and right uropods, dorsal view, holotype. D, higher magnification of distolateral
area of outer uropod (arrow from C). E, higher magnification of tip of telson (arrow from C). Scale bar = 10.0
mm A, B; 7.5 mm C; 3.75 mm D, E.

384

spines located proximally to the origin of
the first ventral rostral spine; in B. echinus,
there are no more than 4. The integument
of B. inflatus was described as “thin” and
the body consequently “soft.” In B. echi-
nus, the integument appears to us to be typ-
ical of a benthic caridean, and not membra-
nous (as seen in midwater species of the
Oplophoridae, for example).

Cleva (1994) and Hanamura and Takeda
(1966) described the body of Bathystylo-
dactylus species as “‘pubescent.”’ Their il-
lustrations show a very light coating of pile.
In B. echinus, the spinules on the body are
characteristic and easily seen, especially on
the dorsal aspect of the carapace. These spi-
nules conform in shape and structure to tac-
tile or vibrational sensory structures seen in
other crustaceans (Cohen and Dijkgraaf
1961).

Bathystylodactylus echinus is the largest
and deepest species known in its family. It
was collected with the flatback lobster Wil-
lemoesia inornata Faxon at stations MV65-
I-38 and MV65-I-39, and with the galatheid
crab Munidopsis antonii (A. Milne-Ed-
wards) at station MV65-I-39.

Acknowledgments

We thank Larry Lovell, Scripps Institution
of Oceanography, for allowing us to ex-
amine the specimens and offering assis-
tance and hospitality during a visit. We also
thank an anonymous reviewer for alerting
us to a potential synonymy. The study ben-
efited from partial support from NSF grants
DEB 9978193 to J. Martin and D. Jacobs
(from the PEET Initiative of Systematic Bi-
ology), DEB 0120635 to Cliff Cunningham
et al. (from the Biocomplexity Genome-En-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

abled Research program), and DEB
0138674 to J. Martin et al. (for collection
support).

Literature Cited

Chace, FE A. Jr. 1983. The caridean shrimps (Crustacea:
Decapoda) of the Albatross Philippine Expedi-
tion, 1907-1910, part 1. Family Stylodactyli-
dae.—Smithsonian Contributions to Zoology
381: 1-21.

Cleva, R. 1990a. Sur les Stylodactylidae (Crustacea,
Decapoda, Caridea) de Il’ Atlantique —Bulletin
Muséum National d’Histoire Naturelle 4, sér.
12, sect. A, 1: 165-176.

. 1990b. Crustacea Decapoda: les genres et les

espéces indo-ouest pacifiques de Stylodactyli-

dae. Pp. 71-136 in A. Crosnier, Résultats des

Campagnes MUSORSTOM, vol. 6.—Mémoires

du Museum National d’ Histoire Naturelle (A),

145.

. 1994. Some Australian Stylodactylidae (Crus-

tacea: Decapoda) with descriptions of two new

species.—The Beagle, Records of the Museum

and Art Galleries of the Northern Territory 11:

53-64.

. 1997. Crustacea Decapoda: Stylodactylidae
récoltés en Indonése, aux iles Wallis et Futuna
et au Vanuatu (Campagnes KARUBAR, MU-
SORSTOM 7 et 8). Données complémentaires
sure les Stylodactylidae de Nouvelle-Calédonie.
Pp. 385—407 in A. Crosnier & P. Bouchet, eds.,
Résultats des Campagnes MUSORSTOM, 16.
Mémoires du Muséum National d’ Histoire Na-
turelle, Paris 172.

Cohen, M. J., & S. Dijkgraaf. 1961. Chapter 2. Mech-
anoreception. Pp. 65—108 in T. H. Waterman,
ed., The physiology of Crustacea, vol. II. Aca-
demic Press, New York, 681 pp.

Hanamura, Y., & M. Takeda. 1996. Establishment of a
new genus Bathystylodactylus (Crustacea: De-
capoda: Stylodactylidae), with description of a
new species from northwestern Pacific.—Zoo-
logical Science 13:929—934.

Okuno, J., & H. Tachikawa. 2000. A new species of
the genus Neostylodactylus Hayashi & Miyake,
1968 (Crustacea, Decapoda: Stylodactylidae)
from southern Japan.—Proceedings of the Bio-
logical Society of Washington 113:39—47.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(3):385-397. 2004.

A new pedunculate barnacle (Cirripedia: Heteralepadidae) from the
Northwest Atlantic

L. Buhl-Mortensen and W. A. Newman

(LB-M) Benthic Habitat Research Group, Institute of Marine Research,
PB. 1870 Nordnes N-5817 Bergen, Norway, e-mail: lene.buhl-mortensen @imr.no;
(WAN) Marine Biological Research Division, Scripps Institution of Oceanography,
La Jolla, California 92093-0202, U.S.A.

Abstract.—A species of Heteralepas has been discovered attached to a gor-
gonian coral from 500 meters of depth off Nova Scotia (~42° N). A brief
review of the previously described Heteralepas species is presented. Of the 29
previously described species (including 2 in synonymy), the new species is
more similar to some from the Indo- West Pacific than to any of the 8 previously
known species from the Atlantic. While the new species can be distinguished
from Atlantic but not some Pacific species by some characters, it can be dis-
tinguished from all species of the genus by small but marked differences in
the configuration of the apertural region of the capitulum. Therefore it is pro-
posed as a new species, Heteralepas cantelli, the most northern known member

of the family.

Introduction

A specimen of Heteralepas was discov-
ered during a survey of the coral-associated
fauna at ~42°N on the continental shelf and
slope off Nova Scotia, Canada, in 2002.
This is not only several degrees of latitude
farther north than any previously known
species of the genus along the Atlantic sea-
board, but at a higher latitude than any pre-
viously known species of Heteralepas
(Young 1999, Zevina 1982). It was collect-
ed by a benthic trawl from ~500 m of
depth, attached to the gorgonian Primnoa
resedaeformis (Gunnerus, 1763).

Thirty species of Heteralepas have been
described, and 2 of these are presently in
synonymy. Of the 28 recognized species
(Table 1), 8 have been recorded from the
Atlantic; 3 from the Western Atlantic, 4
from the Eastern Atlantic and 1 found in
both areas; H. cornuta (Darwin, 1852), H.
lankesteri (Gruvel, 1901), H. belli (Gruvel,
1901), and A. luridas (Zevina, 1975), and
H. microstoma (Gruvel, 1901), H. meteo-

rensis Carriol, 1998, H. alboplaculus Zev-
ina & Kolbasov, 2000 and H. segonzaci
Young, 2001 plus H. cornuta respectively
(Zevina 1975, Young 2001). All these spe-
cies are from relatively low latitudes and
none compare favorably with the new form.
The closest affinities of the new form are
with species like A. japonica (Aurivillius,
1892) from the Indo-West Pacific. However,
the new form can be distinguished from all
previously described species by character-
istics of the apertural regions, and therefore
it is considered to represent a new species.

Systematics

Subclass Cirripedia Burmeister, 1834
Superorder Thoracica Darwin, 1854
Order Pedunculata Lamarck, 1818
Suborder Heteralepadomorpha
Newman, 1987
Family Heteralepadidae
Nilsson-Cantell, 1921

Pilsbry (1907b) revised Alepas and ex-
tracted two distinct but related taxa from it,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

386

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VOLUME 117, NUMBER 3

Heteralepas and Paralepas, but he left
them in the family Lepadidae. Nilsson-Can-
tell (1921) noted that these two genera, in
addition to lacking calcareous plates, dif-
fered from the remaining Lepadidae in the
nature of their trophi and cirri and therefore
he proposed a new family, the Heteralepa-
didae, for them. Species of Heteralepas are
generally considered to have ctenopod or
lasiopod cirri used for setose feeding, while
those of Paralepas have acanthopod cirri,
generally used to feed on the food or tissues
of their hosts, including the eggs of hosts
such as spiny lobsters. The two genera are
further distinguished by the inner ramus of
the posterior two pairs of cirri (cirri V &
VI) being similar to the outer rami in Par-
alepas, but conspicuously reduced in length
and breadth in Heteralepas. However, as
will be noted in the discussion, there is at
least one species that is somewhat inter-
mediate in these characters and it likely
should be assigned a genus of its own.

Heteralepas Pilsbry 1907b
Heteralepas cantelli sp. nov.
(Figs. 1—4)

Type material.—The sole specimen (ho-
lotype) is deposited in the National Muse-
um of Washington, Washington, D.C.
USNM 1019509.

Etymology.—Named in honor of the
Swedish cirripedologist, Carl August Nils-
son-Cantell (cf. Newman 1990) who erect-
ed the family Heteralepadidae.

Material.—Known from a single speci-
men collected in the Northeast Channel,
south of Nova Scotia, Canada (41°55.9’N,
65°42.5'W), by a commercial bottom trawl-
er on October 9, 2002 from 500 m depth.
It was attached to the exposed skeleton of
the gorgonian Primnoa resedaeformis.

Diagnosis: Capitulum and peduncle rel-
atively smooth, without tubercles, carinal
ridge or indications of the insertions of the
carapace adductor muscle; apertural region
slightly recessed or depressed below gen-
eral surface; aperture ~ % height of the ca-

387

pitulum, with crenulate lips restricted to up-
per %.

Description

The fresh specimen was translucent yel-
lowish pink. The capitulum is 3 cm high
and 2 cm wide, globular or nearly ovoid in
lateral aspect, slightly pointed apically, lat-
erally compressed, frontal margin interrupt-
ed by a depressed apertural region with
slightly protuberant lips in the upper % of
the aperture (Figs. 1A, B, 2A, B). The
slightly recessed apertural region is outlined
by a thin edge and in the region below it,
where the carapace adductor muscle is
found, there is a chin-like thickening. Oth-
erwise the capitulum is smooth, without
carinal crest or ridge, warts, bumps or pro-
tuberances. Aperture % height of capitulum;
crenate lips produced in upper 4%. Peduncle
1.2 cm in diameter, equal in length to ca-
pitulum and marked with several folds and
lines in the otherwise smooth cuticle, basal
portion expanded into attachment disc. La-
brum too damaged to describe; mandible
(Fig. 3B) with 4 teeth including inferior an-
gle, surface covered with numerous fine se-
tae, lower margins of teeth 1—3 with a few
fine pectinations (5 under the first and sec-
ond, and 3 under the third tooth; Fig. 3B,
al-a3). First maxilla (Fig. 3C) with cutting
edge stepped (plane of superior cutting
edge indented relative to plane of inferior
cutting edge) rather than notched, with
three major spines (one large flanked by
two somewhat smaller ones) above and ap-
proximately 14 spines below step, with soft
setae in a group along the superior margin
and spread out along the inferior margin,
lateral surfaces clothed with numerous se-
tae. Second maxilla (Fig. 3A) with a prox-
imal cluster of long spine-like setae and a
similar array of setae separated into two
groups along the cutting edge.

Cirrus I not separated from posterior
pairs but modified as a maxilliped of rela-
tively short, unequal, densely setose rami;
cirri II-VI basically similar in structure, se-

388

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

lang, Il.
ertural region; B, lateral aspect of entire animal;

tation lasiopod (Fig. 4C). However, while
cirri II-IV have long subequal rami nearly
equal in length to the outer rami of cirri V
and VI, the inner rami of V and VI are at-
rophied (Fig. 4D, Table 2). The number of
articles comprising the cirri is as follows:

Cirrus: I Il Ill
Inner ramus: 18 42
Outer ramus: Mal 5)II

IV V VI
30) I) 2D" 2
S71 33 OO

Caudal appendage (Fig. 4D) of 13 articles,
slightly longer than pedicel of cirrus VI. Pe-
nis (Fig. 4A, B) relatively long, slender, an-
nulated, without specialized hooks or grap-

Heteralepas cantelli sp. nov.: A, right-frontal aspect of capitulum, enlarged to show details of ap-

ples but clothed with numerous, long soft
setae distally.

Discussion: The cirri of the new species
are fully lasiopod (Fig. 4C) and the inner
rami of the cirri V & VI are substantially
reduced in length as well as breadth (Fig.
4D; Table 2). Therefore the new species is
a Heteralepas in the strict sense. The num-
ber of articles comprising the rami of the
cirri and the form of the mouthparts, while
sometimes useful in distinguishing species,
are considered somewhat variable (Nilsson-
Cantell 1921), as are elaborations of the ca-
pitulum as well as its length relative to the
peduncle (Young 2001). Therefore keys,

VOLUME 117, NUMBER 3

1 cm

389

Fig. 2.
region; B, lateral aspect:

such as that presented by Zevina (1982) for
the 19 species of Heteralepas recognized at
the time, should be used with caution.
Zevina (1982) did not include complete
synonymies in her monograph, and at least
two species once attributed to Heteralepas
were assigned to Paralepas without amend-
ing either genus. Therefore we review all
species attributed to the genus and, as can
be seen from Table 1, 28 species (including
the new form) are presently recognized. In

Heteralepas cantelli sp. nov.: A, left-frontal aspect of capitulum, enlarged to show details of apertural

the process we encountered some problem-
atic forms, and these are briefly discussed
below before moving on to those that are
strictly relevant to the new species.
Heteralepas quadrata_ (Aurivillius,
1892). This shallow-water species [includ-
ing 1) A. percnonicola as a junior synonym
(Hiro 1937), 2) the forms attributed to the
species by Rosell (1972), and 3) a little-
known form from the Eastern Pacific (Zullo
1991] sat uncomfortably in Heteralepas un-

390

Wi A
WW
Wy

| \\y

Fig. 3.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1mm

Heteralepas cantelli sp. nov., mouth parts: A, second maxilla; B, mandible (al’-a3’, enlarged un-

dersides of teeth al-a3); C, first maxilla. (A—C same scale).

til Foster (1979) transferred it to Paralepas,
a decision accepted by Zevina (1982).
However, the species also sits uncomfort-
ably in Paralepas because in some ways it
is morphologically intermediate between
the two genera. The characters largely in-
volve the cirri, their setation being neither
strictly lasiopod nor acanthopod, the rela-
tively low number of articles of their rami,
and the somewhat reduced inner rami of
cirri V & VI, as well as the somewhat in-
termediate armature of the mandible and
first maxilla. This suggests that proposal of
a new genus is in order, and such a study
would be of interest to evolutionary biolo-
gists as well as cirripedologists in light of
the inferred relative primitiveness of the
Heteralepadidae (Foster 1979), a view re-
cently corroborated genetically and mor-

phologically (Harris et al. 2000; Pérez-Lo-
sada et al. 2004). While there are a number
of samples from the Eastern Pacific attri-
buted to this species in the Benthic Inver-
tebrate Collection at Scripps Institution of
Oceanography, an appropriate review of the
situation would also require studying ma-
terials from the Western Pacific. However,
such a study is beyond the scope of the
present paper.

?Heteralepas malaysiana (Annandale,
1905). From a telegraph cable at approxi-
mately 54 m depth in the Gaspar Straits.
Annandale (1909:84) accepted Pilsbry’s
(1907b) revision of Alepas and transferred
Alepas xenophorae Annandale, 1906 to
Heteralepas (Paralepas) and described
Heteralepas (Heteralepas) nicobarica sp.
nov. In the same paper, in a list of species

VOLUME 117, NUMBER 3

Amm (ww““7~ry,
i en
ji aie )

391

Fig. 4. Heteralepas cantelli sp. nov.,

thoracic appendages: A, penis; B, enlargement of distal portion of

penis; C, intermediate segments of outer ramus of cirrus VI; D, posterior of thorax supporting right caudal
appendage and pedicel of cirrus VI with proximal portions of inner and outer rami in outline (narrow and wide

respectively, boundaries of articles omitted).

contained in the Indian Museum, Annan-
dale (1909:130) included Heteralepas ma-
layana (sic) under the subgenus Heterale-
pas. This was presumably because Annan-
dale (1905:81) had clearly stated that the
posterior (= inner) ramus of cirrus V was
““... reduced to a mere thread, less than
one-third as long as the anterior ramus”
and that cirrus VI was “... in much the
same condition”. However, subsequently,
and without a word of explanation, he (An-
nandale 1916:298) transferred Heteralepas
malaysiana to the subgenus Paralepas.
While Newman (1960) retained malaysiana
in Heteralepas s.s., Zevina (1982) followed

Annandale by returning it to Paralepas.
This is puzzling, considering the habitat as
well as the characteristics of cirrus V & VI
given in the original description. In light of
these considerations, and the fact that the
ornamentation of the capitulum appears
more similar to that of Heteralepas rex
(Pilsbry, 1907a) from Hawaii and ZH. uti-
nomit Newman, 1960 from Tasmania than
it does to any species of Paralepas, we
have tentatively returned the species to Het-
eralepas.

?Heteralepas ovalis (Hoek, 1907): This
species is represented by a single specimen
taken along with Paralepas morula from an

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

392

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VOLUME 117, NUMBER 3

echinoid spine in Malaysian waters. Hiro
(1936:223) noted that nothing is known of
the internal parts, but from the original fig-
ures it is evident that the capitulum to ap-
erture ratio is approximately 3:1. This is
suggestive of Paralepas, but for lack of
more conclusive evidence we have left it in
Heteralepas.

Heteralepas cygnus Pilsbry, 1907b: The
original description was based on a speci-
men acquired from the ‘“Ward’s Natural
Science Establishment, Monterey, Califor-
nia’, and hence the specimen was presum-
ably from California, but it has not been
recorded from this region since. Further-
more, Annandale (1909) indicated that
there is a specimen in the Edinburgh Mu-
seum, questionably from the West Indies.
The description may be adequate to distin-
guish it from similar albeit relatively undis-
tinguished forms, but what ocean it came
from remains uncertain.

Heteralepas cornuta (Darwin, 1852): A
species usually having more-or-less distinc-
tive carinal protuberances on its capitulum,
first reported from the Caribbean (presum-
ably from 90 m or so). It has since turned
up in the Gulf of Mexico (Gittings et al.
1986), off Madeira and other West African
islands (cf. Haroun et al. 2003), and along
the coast of Northwest Africa. Furthermore
it has been found in the Indian Ocean, the
Philippines and the Southeast Pacific, off
Chile (4315 m!) (cf. Young 2001 for re-
view). Young (2001) commented not only
on its wide geographical range and the ex-
traordinary depth of the Chilean record
compared to other populations attributed to
the species, but on differences in cirral se-
tation of the Chilean form compared to the
population he has studied from the eastern
Atlantic. Thus H. cornuta may represent a
number of similar species. In any event,
like the previous species, it is sufficiently
distinct from the new form to no longer
concern us here.

Heteralepas microstoma (Gruvel, 1901):
Known from off Madeira, the Azores and
Meteor Seamount immediately to the south.

393

While known to range from between 269—
623 m, it is most commonly found around
300 m (Young 2001). Zevina & Kolbasov
(2000) illustrated and compared it to anoth-
er recently described species, H. meteoren-
sis Carriol, 1998, as well as to their new
species, H. alboplaculus Zevina & Kolba-
sov, 2000, which was also from Meteor
Seamount. Having such similar forms sym-
patric on Meteor Seamount, and then large-
ly from the same depth, is troubling. Young
(2001) synonymized H. meteorensis with
H. microstoma, but he was apparently un-
aware of the work of Zevina & Kolbasov
(2000) who claimed that all three species
can be distinguished from each other by mi-
nute cuticular structures revealed by SEM.
However, their photographs are not clear in
this regard. As can be seen from our Table
2, there appears to be little in the way of
macro-morphological differences among
them, although the peduncle of H. meteo-
rensis seems to be relatively longer and the
aperture does not appear as tubular as in H.
microstoma. On the other hand, Heterale-
pas alboplaculus is described as having the
capitulum and to some extent the peduncle
covered by well-spaced tubercles contain-
ing calcareous structures. Such calcareous
structures are unprecedented in the family
and could be the work of a pathogen. We
hope that workers in the Atlantic will clar-
ify this situation in the near future. In the
meantime, while the specific status of H.
meteorensis and H. alboplaculus is uncer-
tain (Table 1), we have included the former
as well as H. microstoma in Table 2 for
comparative purposes.

When it comes to determining the affin-
ities of the new species, the logical place to
begin is in the Atlantic. Of the 8 previously
known species of Heteralepas noted in the
introduction, 5 occur in the eastern Atlantic.
These include H. cornuta, alboplaculus and
segonzaci, and taking their capitular fea-
tures at face value, they are distinct from
the new form and therefore no longer con-
cern us here. This leaves H. microstoma
and meteorensis, which are very similar if

394

not synonymous, but as noted above both
have been characterized in Table 2 for com-
parative purposes.

As for the western Atlantic species, H.
cornuta, which ranges as far north as the
Carolinas, was noted above as being dis-
tinct from the new form. This leaves H. lur-
idas, belli and lankesteri. The first, from
300-700 m of depth in the Caribbean, is
known to range between 2 and 9.5 mm in
height and the specimen illustrated in the
original description is less than 6 mm high,
so it is a small species. Its capitulum, with
a somewhat tubular or flaring apertural re-
gion, is otherwise undistinguished, and its
cirral and caudal appendage counts are low-
er than in the new species. So, assuming H.
luridas is not based on juveniles, it too need
no longer concern us. This leaves H. belli
and lankesteri, and since in outward ap-
pearance they are similar to the new form,
they have been included in Table 2. As we
shall see, so far none of the species includ-
ed in Table 2 agree well with the new spe-
cies in numerous detail; but what about spe-
cies from the Indo-Pacific?

Of the Indo-West Pacific species, H. ja-
ponica and similar species such as H.. fulva
from the Southeast Pacific are rather close
to the new form. The former has been re-
ported from between 18 and 1020 m depth
from Japan to Singapore, Australia and
New Zealand, the Nicobars in the Andaman
Sea and Réunion Is. (Foster & Buckeridge
1995). Therefore, while not as wide—rang-
ing as H. quadrata, it is wide-ranging com-
pared to most species of the genus. Part of
this range is due to synonymies, and that of
Nilsson-Cantell’s (1927, 1938) for H.
indica (Gruvel, 1901) has long been ac-
cepted. This extended the range of the spe-
cies to Singapore and into the Indian Ocean
where it was reported from Nicobar Is. on
floating wood. Furthermore, Foster (1979),
in his report on New Zealand cirripeds, syn-
onymized H. dubia Broch, 1922 from 55—
72 m in Disaster Bay, Australia, with H.
japonica. However, Zevina (1982), without
explanation, continued to recognize H. du-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

bia as a distinct species, and subsequent au-
thors have followed suit.

Considerable variability in characters
might be expected in such a wide-ranging
species and for present purposes we are ac-
cepting the opinion of these authors. How-
ever, considering such variation in cirripeds
as geographical rather than indicative of ge-
netically distinct populations has generally
proven wrong (Newman 1993). Thus cau-
tion seems in order because the reported
variations in the mandible of presumed H.
Japonica from different populations, appear
to go beyond the range of variability found
within a species. Although Aurivillius
(1894) did not illustrate the mandible of H.
Japonica, his written description agrees
with Nilsson-Cantell’s (1921:247, fig. 43b)
and Pilsbry’s (1911:71, fig. 4A) illustra-
tions, and also with that of Gruvel (1902:
284, Pl. 24, fig. 24) for H. indica. Thus the
teeth of the mandible appear to be without
pectinations, but on close inspection of
Nilsson-Cantell’s illustration there might
have been low pectinations of the lower
margins of teeth 1—3, especially 2 and 3.
However, since there is no such suggestion
in the other illustrations, the evidence fa-
vors the mandible being simple.

The situation at the southern end of the
range for H. japonica looks quite different
with regard to the mandible. Foster (1979)
synonymized H. dubia Broch from Austra-
lia, and the population he was studying in
New Zealand, with H. japonica. While
Broch (1922:288, fig. 37B) gave no indi-
cation of pectinations on the first tooth, he
clearly illustrated them on the upper sides
of teeth 2—4 as well as the lower sides of 2
and 3. Foster (1979:16, fig. 3J) illustrated
the same for the upper sides, but limited
pectinations on the underside to tooth 1. So,
the populations attributed to this species
from north and south of the equator appear
to differ in the characteristics of the man-
dible, and the new species, with its incon-
spicuous pectinations on the lower sides of
teeth 1-3 (Fig. 3, al-a3), differs from both
of them.

VOLUME 117, NUMBER 3

In view of the foregoing considerations
we include H. dubia in Table 1 as a ques-
tionable species rather than a synonym of
H. japonica. Nonetheless, H. japonica still
includes sufficient variability to make it an
ideal Indo-Pacific representative similar to
the new form from the Atlantic, and there-
fore it is included in Table 2 for compara-
tive purposes.

Summary and Conclusions

The essentially naked heteralepadids pre-
sent a difficult problem to systematists
since, being unarmored, they lack a number
of distinct features customarily utilized in
separating genera and species (Zullo &
Newman 1964). Aside from the work of
Nilsson-Cantell (1921, 1927), and to some
extent, Young (2001), no studies have eval-
uated the usefulness of morphological char-
acters in distinguishing Heteralepas spe-
cies. Thus it is difficult to establish a new
species with a high degree of certainty. But,
in spite of the latitude allowed by synony-
my, the present form could not be assigned
to any known species.

As can be observed in Table 2, the At-
lantic species most similar to the new spe-
cies (H. microstoma, meteorensis, belli and
lankesteri) are readily distinguished from it
as well as from H. japonica from the Indo-
West Pacific, by several characters. How-
ever, the new species, H. cantelli, cannot be
distinguished from H. japonica by the char-
acters presented in the table. This is due in
part to the variability attributed to H. ja-
ponica, but there are notable differences be-
tween these two species, not included in the
table, that distinguish them. These include
1) the lack of any indication of a carinal
thickening, crest, or protuberances along
the carinal margin (but sometimes also
found lacking in individuals of H. japoni-
ca), 2) a marked crenation of the apertural
margin largely restricted to the upper third
rather than along its entire margin, and 3)
a slightly depressed area around the entire
apertural region, setting it off from the gen-

395

eral surface of the capitulum. The last two
differences are sufficient not only to distin-
guish the new form from H. japonica, but
from all known heteralepadids.

Acknowledgments

We thank Paulo S. Young, Museu Na-
cional/UFRJ, Rio de Janeiro, Brazil, who
died tragically before the publication of this
paper, for advice on the Atlantic species
during preparation of the manuscript, and
Pal B. Mortensen, Bedford Institute of
Oceanography, Dartmouth, Canada and
Vladimir E. Kostylev, Natural Resources
Canada, Dartmouth, Canada, for helping
with the translation of publications in Ger-
man and Russian, respectively. While we
would also like to thank two judicious ref-
erees (John S. Buckeridge, EOS, Auckland
University of Technology, as well as Paulo
S. Young) for reviewing the manuscript, we
are solely responsible for any errors that re-
main.

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tional and Applied Science Bulletin 24(4):104—
283.

Young, P. S. 1999. The Cirripedia (Crustacea) collected
by the “Fisheries Steamer Meteor” in the East-
ern Atlantic—Arquivos do Museu Nacional,
Rio de Janeiro 58:1—54 (1998).

. 2001. Deep-sea Cirripedia Thoracica (Crus-
tacea) from the northeast Atlantic collected by
French expeditions.—Zoosystema 23(4):705—
756.

Zevina, G. B. 1975. Cirripedia Thoracica of the Amer-
ican Mediterranean.—Trudy Instituta Okeanol-
ogii 100:233—258 (in Russian).

. 1982. Barnacles of the suborder Lepadomor-

pha of the world ocean. II. Pp. 1-222 in Fauna

U.S.S.R., Zoological Institute, Russian Acade-

my of Science, Leningrad, 133 (in Russian).

, & G. A. Kolbasov. 2000. Barnacles of the

genus Heteralepas (Thecostraca, Cirripedia,

Thoracica) from the Canary Islands and the

VOLUME 117, NUMBER 3 397

Azores. Description of mantle ultrastructure.— jacent regions in the tropical eastern Pacific. Pp.
Zoologicheskii Zhurnal 79(11):1275-1283 (in 173-192 in M. J. James, ed., Galapagos marine
Russian). invertebrates. Taxonomy, biogeography, and
, & M. Y. Schreider. 1992. New species of Cir- evolution in Darwin’s Islands. Plenum Publish-
ripedia (Crustacea) from the Indian Ocean.—Zool- ing Company, New York, New York.

ogicheskii Zhurnal 71(10):39—46 Gn Russian). , & W. A. Newman. 1964. Thoracic Cirripedia
Zullo, V. A. 1991. Zoogeography of the shallow-water from a southeast Pacific guyot—Pacific Sci-
cirriped fauna of the Galapagos Islands and ad- ence 18(4):355-372.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
117(3):398—407. 2004.

Two new species of seven-spined Bathyconchoecia from the
North Atlantic and Indian oceans
(Crustacea: Ostracoda: Halocypridae)

Louis S. Kornicker and J. A. Rudjakov

(LSK) Department of Zoology—IZ, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20013-7012, U.S.A.,
e-mail: kornicker.louis@nmnh.si.edu;
(JAR) Museum of Comparative Zoology, Harvard University, 26 Oxford St., Cambridge,
Massachusetts 02138-2902, U.S.A, e-mail: rudyakov @fas.harvard.edu

Abstract.—A new species of halocyprid ostracode Bathyconchoecia omega
from abyssal depths of the North Atlantic Ocean, off Newfoundland, Canada,
is described and illustrated, and a new species Bathyconchoecia georgei is
proposed for a specimen from the Indian Ocean previously referred to Bathy-
conchoecia deeveyae Kornicker, 1969.

The R/V Chain, operated by the Woods
Hole Oceanographic Institution, collected
in 1972 at a depth of 4400 m in the North
Atlantic Ocean, off Newfoundland, Canada,
a bottom sample containing a single A-1
male of Bathyconchoecia omega, new spe-
cies. The A-1 male from off Newfoundland
is considerably larger than previously de-
scribed seven-spined species of the genus,
and is the northernmost occurrence of the
group. Additional ostracodes in the sample
are mostly bottom-living Podocopida, Cla-
docopida and Myodocopida, which sug-
gests a bottom or near-bottom habitat for B.
omega. However, a specimen of pelagic
species of Conchoecia in the sample sug-
gests that it contains some shallow water
contaminants.

Only three species of Bathyconchecia
having seven spines on the carapace (four
on right valve, three on left), have been de-
scribed previously: B. deeveyae Kornicker,
1969, B. septemspinosa Angel, 1970, and
B. longispinata Ellis, 1987. One of the
specimens previously referred to B. deev-
eyde is proposed as a new species herein.
Thus, the number of 7-spined species of
Bathyconchoecia is now five. Their distri-
bution is shown in Fig. 1.

Correction.—Kornicker (1981:1237) re-
ported that the slide containing the append-
ages of the holotype of B. deeveyae (USNM
123335) had been lost. It has been recov-
ered.

Bathyconchoecia omega, new species
Figures 2—6

Holotype.—Unique specimen, A-1 male
on slide and in alcohol, MCZ Harvard Uni-
versity, MCZ50432.

Type locality.—R/V Chain 106, 30 Aug
1972, Station 334, North Atlantic Ocean,
off Newfoundland, Canada, 40°42.6'’N—
40°44'N, 46°13.8’W—46°14.6'W, epibenthic
sled, depth 4400 m.

Material.—Holotype.

Description of A-I male (Figs. 2—6).—
Carapace with linear dorsal margin except
for slight bulge near middle just posterior
to base of dorsal spine. Posterodorsal corner
of each valve with gland on very slight
bulge. Posterodorsal corner evenly rounded
except for long spine on right valve; spine
parallel with length of valve, but at slight
upward angle (very tip of spine of specimen
broken off; soft matter projects from broken
tip). Base of spine projects slightly medial

VOLUME 117, NUMBER 3

Fig.1.

to slightly overlap posterior edge of left
valve (Fig. 2B). Rostrum of each valve with
anterior spine at slight angle to each other
(Fig. 2B). Spine at midlength of dorsal mar-
gin of each valve at slight outward and up-
ward angle (Fig. 2A, B). Spine near ventral
margin of each valve at about % length of
valve at slight downward and outward an-
gle. Anterior spines on rostra and posterior
spine on right valve with surface ridges par-
allel to lengths of valves; a few of the ridg-
es of the rostral spines bear short stout
spines. Other long spines with minute sur-
face spines. Carapaces completely covered
by distinct punctae and slightly curved ver-
tical frills (not all shown in Fig. 2A). Frills
generally on each side of 2 or 3 rows of
punctae (Fig. 2C). Indistinct reticulations
and ridges on anteroventral surface of valve
ventral to incisure (Fig. 2A).

Pigmentation: No black pigment spots on
either carapace or body.

Central adductor muscle attachments
(Fig 2A): Indistinct, near center of valve

399

B. deeveyae

B. georgei

B. longispinata
B. omega

B. septemspinosa

Distribution of species of seven-spined Bathyconchoecia.

and consisting of 2 individual scars; stria-
tions of muscle ends indistinctly visible
from outside view of valve; scars not cov-
ered by punctae.

Carapace size (mm): Length including
spines 3.79, length excluding spines 2.92,
height excluding spines 1.68, width without
spines 1.52.

First antenna (Fig. 2E): Shaft short with
indistinct segmentation. Brush-like struc-
ture with about 315 filaments in about 9
rows, each with about 35 filaments. Dorsal
bristle on segment following brush-like
structure stout, spinous, about % length of
brush filaments. Terminal segment with 4
bristles: 1 long stout bristle reaching well
past brush filaments and with widely scat-
tered marginal spines (not shown); 3 shorter
than brush filaments. Limb with densely
packed amber-colored cells.

Second antenna (Fig. 3A—D): Protopod
bare. Endopod: Ist article with 2 spinous
dorsal bristles (1 long, 1 short) and few in-
distinct medial spines near ventral margin;

400 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Bathyconchoecia omega holotype, MCZ 50432, A-1 male: A, Complete carapace from right side,
length without spines 2.92 mm; B, Complete carapace, ventral view; C, Left valve, detail of ornamentation on
outer surface; D, Posterodorsal corner of complete specimen (spine on right valve, glandular opening on left
valve); E, Right Ist antenna, lateral view.

VOLUME 117, NUMBER 3 401

Sen Rene

Fig. 3. Bathyconchoecia omega holotype, MCZ 50432, A-1 male: A, Left 2nd antenna, medial view; B,
Endopod right 2nd antenna, lateral view; C, Proximal part exopod left 2nd antenna, medial view; D, Distal part
exopod right 2nd antenna, lateral view; E, Proximal part left Sth limb, lateral view; EF Right 5th limb drawn on
body, lateral view.

402

2nd article with 1 minute bristle medial to
3rd article and 2 stout terminal bristles with
few indistinct marginal spines (inner bristle
stouter, both about same length as exopod
bristles); 3rd article with 3 bristles: middle
bristle longer and stouter than others, more
than 1/2 length of bristles of 2nd article,
with few marginal spines; outer bristle
about % length of middle bristle, with many
marginal spines; inner bristle similar in
length to outer bristle, with marginal spines;
base of 3rd article lateral to distal end of
2nd article; endopods of left and right limbs
similar. Exopod: Ist article with short ven-
tral spines and small medial terminal bris-
tle; articles 2 to 8 with long natatory bristle;
Oth article with 4 bristles (2 short lateral; 1
ventral of medium length and with short
marginal spines; 1 long dorsal, with nata-
tory hairs).

Mandible (Fig. 4): Coxa (Fig. 4B-—E):
Pars incisivus with 5 ventral teeth and slen-
der distal tooth at ventral tip of triangular
posterior section; anterior edge serrate (Fig.
4C). Proximal list with 10 teeth in 2 layers
(Fig. 4D); distal list with 19 teeth in 3 lay-
ers (Fig. 4E). Spined posterior part with 6
lobes with numerous spines and 7th lobe
with short stout spinous bristle and minute
spines along distal posterior edge of lobe
near bristle (Fig. 4C). Anterior margin of
coxa evenly rounded, without triangular
process. Basis (Fig. 4A, E G): 2 long plu-
mose bristles present on or near dorsal mar-
gin and | long spinous medial bristle near
midwidth some distance from dorsal margin
(Fig. 4F); lateral surface with 3 long bare
distal bristles and long spines (Fig. 4F);
posterior margin spinous and with 2 short
distal bristles (proximal bristle sclerotized
and with ventral spines; distal bristle tube-
formed) (Fig. 4G); anterior margin with
long bare distal bristle (Fig. 4G); ventral
margin with 5 short teeth with minute sec-
ondary teeth (Fig. 4G); | short tooth with
minute secondary teeth on posterior margin
proximal to posterior ventral tooth. Poster-
odorsal corner of basis with oval sclerite
(Fig. 4A, B). Endopod (Fig. 4E H): Article

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1 with dorsal, ventral, lateral, and medial
slender spines and 4 bristles (1 long, ter-
minal, dorsal, spinous; 1 long, distal, ven-
tral, bare; 2 medium length, medial with ba-
ses close to ventral bristle, bare). Article 2
with 5 bristles (2 long and 1 shorter, ter-
minal, dorsal, spinous; 2 medium length,
distal ventral, bare) and few distal spines on
ventral margin and medial and lateral sur-
faces near ventral margin. Article 3 with
long spinous terminal claw and 6 bristles (2
long, spinous, terminal (shorter of these
with base on medial side), and 4 short, ven-
tral, bare (1 of these with lateral base)) and
medial spines.

Maxilla (Fig. SA—C): Endite of precoxa
with 2 tube-formed bristles, 3 claws, and 2
long spinous bristles. Coxa: dorsal margin
with long stout dorsal bristle (Fig. 5B);
proximal endite with 3 tube-formed bristles
and total of 4 claws and claw-like bristles;
distal endite with 2 tube-formed bristles and
4 claws. Basis with 2 long stout plumose
bristles near dorsal margin, and short bare
ventral bristle. Endopod: article 1 spinous
with 4 dorsal bristles (3 proximal, | distal);
medial surface with 4 distal bristles (3 long,
1 short); article 2 with 2 stout claws of un-
equal length and 4 slender bristles.

Fifth limb (Fig. 3E, F): Epipod with 3
groups of 4 stout plumose bristles; dorsal
group with additional small 5th bristle (Fig.
3F). Precoxa with 3 ventral bristles (Fig.
3E). Coxa with 11 or 12 ventral bristles (not
all shown). Basis with 6 bristles plus long
terminal dorsal exopod bristle with minute
widely separated marginal spines (not all
shown). Endopod: article 1 with dorsal and
medial spines and 3 bristles (not all shown).
Article 2 with dorsal and medial spines and
4 bristles (3 near ventral margin and | lon-
ger dorsal). Article 3 with 2 long terminal
slender claws and 1 long ringed, terminal,
slender ventral bristle. A muscle terminates
at base of exopod bristle.

Sixth limb (Fig. 6A): Epipod with 3
groups of 5, 5, and 6 (dorsal) long plumose
bristles; dorsal group with additional short
7th bristle (Fig. 6A). Coxa with 1 spinous,

VOLUME 117, NUMBER 3 403

Fig. 4. Bathyconchoecia omega holotype, MCZ 50432, A-1 male: A, Left mandible, junction of coxa and
basis, lateral view. B—H, Right mandible, lateral views: B, Proximal part of coxa; C, Distal end of coxa; D,
Detail of proximal tooth of coxa (detail from C); E, Detail of distal tooth of coxa (detail from C): E Basis and
endopod; G, Distal end of basis; H, Endopod.

404 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Bathyconchoecia omega holotype, MCZ 50432, A-1 male: A, Right maxilla, medial view (arrow
indicates tube-formed bristles); B, Left maxilla, lateral view; C, Right maxilla, oblique dorsal view (not all
bristles shown); D, Anterior of body from right side showing upper and lower lips (esophagus dashed); E, F
Lower lip from left side, anterior of body to left; G, Upper lip, dorsal view.

VOLUME 117, NUMBER 3

~
LZ
4222 =
LEE
a
M1
ANS
\
A
eV
Ul] \

CRAP SesSone
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405

Fig. 6. Bathyconchoecia omega holotype, MCZ 50432, A-1 male: A, Right 6th limb, medial view; B, 7th
limb; C, Left lamella of furca and unpaired bristle, lateral view; D, Right lamella of furca, lateral view; E,
Posterior view of ventral end of body showing unpaired bristle and furca; E Posterior view of body showing
copulatory organ on left side; G, Posterior of body from right side showing furca and copulatory organ; H,

Copulatory organ from left side, anterior to upper left.

406

ventral, terminal bristle. Basis with spines,
4 spinous bristles and 1 long, terminal, dor-
sal, exopod bristle with widely scattered
minute spines (basis may consist of medial
and lateral parts). Endopod: article 1 with 4
bristles; article 2 with 2 bristles; article 3
with 3 long terminal bristles (dorsal 2 claw-
like). A muscle terminates at base of exo-
pod bristle.

Seventh limb (Fig. 6B): Broad thumb-
like process with 2 long unequal bare bris-
tles.

Furea (Fig. 6C—G): Each lamella with 7
claws with teeth along posterior margins; |
unpaired spinous bristle following claws on
lamellae (Fig. 6E).

Bellonci organ: Not developed.

Lips (Fig. 5D—G): Upper lip with spinous
posterior edge (Fig. 5D, G). Lower lip spi-
nous (Fig. 5E, F).

Copulatory organ (Fig. 6F—H): Organ
with 2 separate branches on left side of
body. Broad anterior branch with minute
terminal teeth; narrow posterior branch with
small tapered tip.

Comparisons.—The length of the unique
A-1 male from off Newfoundland (exclud-
ing spines) is 2.92 mm, whereas A-1 instars
of B. deeveyae and B. septemspinosa are
shorter than 1.8 mm (Kornicker and Angel,
1975: table 1; Kornicker, 1981:1240). A
length of 0.66 mm was reported for an A-
4 instar of B. deeveyae by Kornicker (1991:
30). The adult male of B. longispinata has
a range of lengths of 1.95—2.11 mm (Ellis,
1987: Table Il), much shorter than the 2.92
mm length of the A-1 male referred herein
to B. omega. The 2 mid-dorsal spines on
the carapace of the later specimen are short-
er than those of B. longispinata. Also, the
fossae and frills of B. omega are on all parts
of the valve, whereas, they cover only cer-
tain areas on B. longispinata. The length of
the adult male of B. georgei, new species,
is 1.28 mm, much smaller than the length
(2.92 mm) of the A-1 male of B. omega.
The carapace of the former species is with-
out the frills present on the carapace of B.
omega.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

The 2nd endopod articles of both man-
dibles of the A-1 instar of B. omega bear 5
bristles compared to 4 on the A-1 mandi-
bles of B. septemspinosa and B. deeveyae
and the adult male mandible of B. longis-
pinata, and 3 on the adult male mandible
of B. georgei. Mandibles of a total of six
A-1 and A-2 instars of B. septemspinosa
examined by Kornicker and Angel (1975:
Table 1) indicate that the number of bristles
on the 2nd endopod article of the mandible
of those instars do not vary from 4 bristles
and, therefore, may be a reliable character
to use to discriminate specimens of B. ome-
ga, but reliability of the character in the lat-
ter species is unknown.

Bathyconchoecia georgei, new species

Bathyconchoecia deeveyae Kornicker.—
George, 1971: 141, figs. 1-9.

Not Bathyconchoecia deeveyae Kornicker,
1969: 403, pl. 1, figs. 1-2.

Etymology.—Species named in honor of
Jacob George, National Institute of Ocean-
ography, Cochin-18, India, who described
the specimen upon which the new species
is based.

Holotype.—Unique specimen, adult
male. Specimen is in a vial labeled 07.43,
with serial number 0130, deposited in the
archive room at the Indian Ocean Regional
Centre, National Institute of Oceanography,
Cochin — 14, India (there are no mounted
slides). (Information about specimen sup-
plied by Dr. Rosamma Stephen, Scientist,
National Institute of Oceanography Region-
al Center, Cochin, in correspondence with
the junior author. Dr. Stephen did not ex-
amine specimen in vial, but stated that she
“could make out that there is a white spec-
imen inside.’’)

Type locality.—International Indian
Ocean Expedition station Co. 62 (I. O. B.
C.1969), in vertical haul from 200 to O m,
off SW coast of India, 10°39'N, 75°22’E.

Material.—None examined.

Discussion of B. deeveyae Kornicker,
1969.—This species was described from an

VOLUME 117, NUMBER 3

A-1 juvenile collected at a depth of 508—
523 m in a benthic trawl in the Peru-Chile
Trench System, Pacific Ocean (Kornicker,
1969:403). A second specimen, an adult
male, was collected in a vertical plankton
haul from 200 to O m in the Indian Ocean
off the SW coast of India (George, 1971:
141). A third specimen, an adult or A-1 fe-
male, was collected at a depth of 520 m in
an epibenthic sled from off Surinam, Atlan-
tic Ocean (Kornicker, 1981:118). Ellis
(1987:83) observed, “It is possible that
these three specimens are not conspecific.”
That observation prompted the present au-
thors to reconsider the three specimens that
had been referred to B. deeveyae, and led
to our conclusion that the Indian Ocean
specimen is not conspecific with the other
two specimens of B. deeveyae from the At-
lantic and Pacific Oceans. The Indian
Ocean specimen was adequately described
by George (1971: 141), so that only a brief
diagnosis based on the adult male is pre-
sented here.

Diagnosis (adult male).—Carapace 1.28
mm long, excluding spines. Second endo-
pod article of mandible with 3 bristles. Fur-
ca with 8 claws on each lamella.

Comparisons.—The carapace of the new
species, B. georgei is much smaller than
equivalent stages of B. septemspinosa, B.
deeveyae, and B. longispinata (because
only the adult male of B. georgei is known,
the relative sizes of its instars is an extrap-
olation). The 2nd endopod article of the
mandible of the adult male B. georgei bears
3 bristles compared to 5 on the adult male
of B. longispinata and 4 on the A-1 instars
of both B. septemspinosa and B. deeveyae.
The adult male B. georgei bears 8 claws on
each lamella compared to 7 on the adult
male B. longispinata.

Acknowledgments

We thank Elizabeth Harrison-Nelson for
preparing the illustrations and text for pub-

407

lication, Molly Ryan for producing the spe-
cies distribution map (Fig. 1), and Megan
Bluhm for inking the illustrations from pen-
ciled Camera Lucida drawings by the first
author. We are greatly indebted to Dr. Ro-
samma Stephen, National Institute of
Oceanography, Cochin, India, for providing
information about the present location of
the type specimen of B. georgei. The junior
author would like to thank Dr. Gonzalo Gi-
ribet and Mrs. Ardis B. Johnston for their
help and encouragement.

Literature Cited

Angel, M. V. 1970. Bathyconchoecia subrufa n. sp.
and B. septemspinosa n. sp., two new halocy-
prids (Ostracoda, Myodocopida) from the trop-
ical North Atlantic and the description of the
larval development of B. subrufa. —Crusta-
ceana 19:181—199.

Deevey, G. B. 1968. Pelagic ostracods of the Sargasso
Sea off Bermuda: Descriptions of species, sea-
sonal and vertical distribution. —Peabody Mu-
seum of Natural History (Yale University) 26:
1-125.

Ellis, C. J. 1987. Bathyconchoecia longispinata n. sp.,
a new species of halocyprid Ostracod with sey-
en carapace spines.—Crustaceana, 53:83—93.

George, J. 1971. On the occurrence of Bathyconchoe-
cia deeveyae Kornicker (Ostracoda, Halocypri-
didae) in the Indian Ocean. —Crustaceana 21:
141-144.

Kornicker, L. S. 1969. Bathyconchoecia deeveyae, a
highly ornamented new species of Ostracoda
(Halocyprididae) from the Peru-Chile Trench
system.—Proceedings of the Biological Society
of Washington 82:403—408.

. 1981. Range extension and supplementary de-

scription of Bathyconchoecia deeveyae (Ostra-

coda: Halocyprididae).—Proceedings of the Bi-

ological Society of Washington 94:1237—1243.

. 1991. Myodocopid Ostracoda of Hydrother-

mal Vents in the Eastern Pacific Ocean. Smith-

sonian Contributions to Zoology, 516, 46 pages,

25 figures, 2 tables.

, & M. V. Angel. 1975. Morphology and on-

togeny of Bathyconchoecia septemspinosa An-

gel, 1970 (Ostracoda: Halocyprididae). —

Smithsonian Contributions to Zoology 195: 1—

21.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(3):408—422. 2004.

The hermaphroditic sea anemone Anthopleura atodai n. sp.
(Anthozoa: Actiniaria: Actiniidae) from Japan, with a redescription
of A. hermaphroditica

Kensuke Yanagi and Marymegan Daly

(KY) Costal Branch of Natural History Museum and Institute, Chiba, 123 Yoshio, Katsuura,
Chiba Pref., 299-5242 Japan, e-mail: yanagi@chiba-muse.or.jp;
(MD) Department of Ecology and Evolutionary Biology, University of Kansas,
Lawrence KS 66045 U.S.A., Current address: Dept. of Evolution, Ecology & Organismal
Biology, Ohio State University, Columbus, Ohio 43210 U.S.A. e-mail: daly.66@osu.edu

Abstract.—A new species of internally brooding sea anemone, Anthopleura
atodai, is described from the middle to northern Pacific coasts of Honshu,
Japan. This species attaches to mussels or in rock crevices of the higher tidal
zone. This is the second hermaphroditic species and fourth internally brooding
species of Anthopleura to be reported; it is distinguished from other members
of Anthopleura by a combination of the following features: brooding its young,
synchronously hermaphroditic, S-shaped basitrichs in filaments, 40 to 68 ten-
tacles, verrucae in the proximal part of the column larger than those in the
distal part, cobalt-blue spot at the distal end of each siphonoglyph. Anthopleura
hermaphroditica, the species that most closely resembles A. atodai, is rede-
scribed to clearly differentiate it from A. atodai and to resolve questions about

its taxonomy and identity.

Anthopleura Duchassaing and Michelot-
ti, 1861, one of the largest genera in the
Actiniaria, includes about 50 species (Carl-
gren 1949; Dunn 1974, 1978, 1982a; Fautin
2003). In Japanese waters, six species of
Anthopleura are known: Anthopleura asia-
tica Uchida & Muramatsu, 1958; A. fusco-
viridis Carlgren, 1949; A. kurogane Uchida,
1938; A. mcmurrichi Wassilieff, 1908; A.
pacifica Uchida, 1938; A. uchidai England,
1992. Additionally, Atoda (1954) reported
the post-larval development of an uniden-
tified species of Anthopleura, which broods
its young in the colenteron. Although Atoda
(1954) mentioned that the species could be
distinguished from other species of Antho-
pleura by its coloration, it has never named;
we formally describe it here as a new spe-
cies, A. atodai.

Internal brooding is widely known in the
Actiniaria: e.g. Actinia spp. (Chia & Ros-
tron 1960; Rossi 1971; Black & Johnson

1979; Ayre 1983; Manuel 1988; Russo et
al. 1994; Yanagi et al. 1996, 1999), Aulac-
tinia sp. (Dunn et al. 1980), Cereus pedun-
culatus (Rossi 1971), Cnidopus japonicus
(T. Uchida 1934, T. Uchida & Iwata 1954),
Epiactis spp. (Dunn 1975, Fautin & Chia
1986, Edmands 1995), and Bunodactis her-
maphroditica (McMurrich 1904). Aside
from A. atodai, three species of Anthopleu-
ra are reported to brood internally: A. handi
Dunn, 1978, from the Philippines, Hong
Kong, and Malaysia (Dunn 1978, England
1987); A. aureoradiata (Stuckey, 1909a)
from New Zealand (Stuckey 1909a, 1909b;
Carlgren 1949, 1954; Parry 1951); and A.
hermaphroditica (Carlgren, 1899) from
Chile (Carlgren 1899, 1927, 1949, 1959).
Anthopleura atodai most closely resem-
bles A. hermaphroditica. Because the anat-
omy and cnidom of A. hermaphroditica is
incompletely known, and its taxonomic sta-
tus is unclear, we redescribe it to clearly

VOLUME 117, NUMBER 3

Asamushi
Hachinohe

Katsuura

Tateyama

Pacific Ocean

Fig. 1.
species. Stars indicate records of Anthopleura sp. giv-
en by Atoda (1958); circles indicate sites visited in this
study.

Distribution of Anthopleura atodai, new

distinguish A. hermaphroditica from A. ato-
dai and to evaluate the proposed synonymy
between A. hermaphroditica and A. handi.
We find that A. hermaphroditica and A. ato-
dai can be distinguished based on color,
number of tentacles, cnidom, and geograph-
ic range, and that A. hermaphroditica 1s dis-
tinct from A. handi.

Materials and Methods

Specimens of Anthopleura atodai were
collected from high intertidal rocky shore
around Asamushi (40°54’N, 140°51’E), Ot-
suchi (39°22’N, 141°58’E), Katsuura
(35°07'’N, 140°16’ E), and Tateyama
(34°58'N, 139°46’E) (Fig. 1). Anatomical
observations were made on 17 specimens
of A. atodai; histological sections were
made from 11 specimens. Anatomical ob-
servations were made on 10 preserved
specimens of A. hermaphroditica; histolog-
ical sections were made from 5 animals. For
specimens of both A. atodai and A. her-
maphroditica, histological sections 6—8 4m
thick were stained with hematoxylin and
eosin or with Haidenhain’s Azan (Presnell
and Schreibman, 1997).

409

Cnidae data were gathered following the
method of England (1987) and Williams
(1996). Cnidae were measured from both
live and preserved specimens of A. atodai,
and from preserved specimens of A. her-
maphroditica. Cnidae were measured in
smash preparations at 1000 X using differ-
ential interference light microscopy. The
terminology for cnidae follows Weill
(1934), Mariscal (1974), and England
(1991).

The material examined was deposited in
the Costal Branch of Natural History Mu-
seum and Institute, Chiba (CMNH), Nation-
al Science Museum, Tokyo (NSMT), Swed-
ish Museum Natural History, Stockholm
(SMNH), State Zoological Museum, Mu-
nich (ZSM), and The University of Kansas
Natural History Museum and Biodiversity
Research Center (KUMNH).

Systematic Account

Family Actiniidae Rafinesque, 1815
Genus Anthopleura Duchassaing and
Michelotti, 1860
Anthopleura atodai, new species
Figs. 2—5

Anthopleura sp.—Atoda, 1954: 274, figs.
1-29, pls. 6—7.—Isomura et al., 2003:
M3), 103, Ik.

Holotype.—Kenashi-jima, Otsuchi, Iwate
Pref., Honshu, Japan (39°21'30’N,
141°57’S50"E), 14 July 1997, collected by
KY, | specimen, with histological sections
and cnidae preparations (CMNH-ZG 64).

Paratypes.—All from Honshu, Japan and
collected by KY: Kenashi-jima, Otsuchi,
Iwate Pref., 14 Jul 1997, 1 specimen with
cnidae preparations (CMNH-ZG 65), 1
specimen with histological sections
(NSMT-Co 1373), 1 specimen (NSMT-Co
1374), 1 specimen (KUMNH 1808), 1 spec-
imen entirely sectioned longitudinally
(CMNH-ZG 3692), 1 specimen entirely
sectioned transversely (CMNH-ZG 3693);
Banda, Tateyama, Chiba Pref., 28 Oct 1996,
1 specimen with cnidae preparations

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. A-C, Photographs of Anthopleura atodai, new species (A, collected at Banda, 24 Feb 1997; B, C,
collected at type locality, Kenashi-jima, 14 Jul 1997): A, 2 specimens expanded and 1 specimen contracted; B,
semi-expanded specimen; C, fully contracted specimen. D, E, Photographs of A. hermaphroditica (collected
from Chiloe Island, Chile): D, Two specimens expanded; E, Typical oral disc patterning. Photographs in D, E
courtesy of V. Haussermann. Scale Bars: A, D = 10 mm; B, C = 5 mm, E = 20 mm.

VOLUME 117, NUMBER 3

(CMNH-ZG 115), 1 specimen (CMNH-ZG
200), 11 Dec 1996, 24 Feb 1997, 1 speci-
men with cnidae preparations (CMNH-ZG
44), 1 specimen (NSMT-Co 1372), 5 Dec
1997, 1 specimen entirely sectioned longi-
tudinally (CMNH-ZG 3695), 1 specimen
entirely sectioned transversely (CMNH-ZG
3696); Hadaka-jima, Asamushi, Aomori
Pref., 22 Jun 1998, 1 specimen with histo-
logical sections and cnidae preparations
(CMNH-ZG 209), 1 specimen with cnidae
preparations (CMNH-ZG 210), 1 specimen
(KUMNH 1809), 1 specimen entirely sec-
tioned transversely (CMNH-ZG 3697).
Non-type material examined.—All spec-
imens collected from Honshu, Japan by
KY: Kedo-ura, Katsuura, Chiba Pref., 1
May 1999, 4 specimens (CMNH-ZG 253);
Banda, Tateyama, Chiba Pref., 11 Dec
1996, 4 specimens (CMNH-ZG 201), 24
Feb 1997, 6 specimens (CMNH-ZG 3694),
15 May 2000, 2 specimens (CMNH-ZG
906); Nojima, Otsuchi, Iwate Pref., 8 Aug
2001, 4 specimens (CMNH-ZG1060), 4
specimens (CMNH-ZG 1061); Hadaka-
jima, Asamushi, Aomori Pref., 22 Jun 1998,
60 specimens (CMNH-ZG 3698), 5 speci-
mens (KUMNH 1809-1810), 3 specimens
(NSMT-Co 1375-1378).
Description.—Column and pedal disc:
Freshly collected specimens brown or blu-
ish-green, proximal verrucae whitish (Fig.
2A-C). In living, expanded animals, col-
umn width 6-12 mm, almost equal to
height (Fig. 2A—C); oral and pedal disc of
almost equal width. Column of contracted
animals dome-like (Fig. 2A, C). Adhesive
endocoelic verrucae in regular vertical rows
from margin to limbus; in some individuals,
becoming dense and irregular distally (Fig.
2B); number of rows 24—39 (37 in holo-
type) distally, 24 proximally. Diameter of
verrucae increases proximally: 0.4 mm at
margin, 0.6—1.2 mm at limbus. In life, ver-
rucae hold bits of gravel and broken shells.
Marginal endocoels bear 9-32 pale,
opaque, spherical acrorhagi that curve into
fosse (Table 1). Pedal disc weakly adherent,

»’ indicates that an attribute was not measured

Table 1.—Morphological variability of 11 specimens of Anthopleura atodai, n. sp. collected from three localities. **

or counted for that specimen.

Number of
siphonoglyphs

Number of pairs
of directive
mesenteries

Number of pairs of mesenteries
Number of
(in mm) acrorhabi Distal column Proximal column

Height of
column

Diameter of
pedal disc
(in mm)

Specimen

32
23

10.7

12.0

ype (CMNH-ZG 64)
ype (CMNH-ZG 65)

28
34
34

9.7

12.4

34
34

14.4

10.0

11.9

11.7

39
24

12.1

9.3

7.9
9.8

6.1

9.5

38
34
35

9.7

11.8
7.8

11.9

ype (NSMT-Co 1373)

ype (KUMNH 1808)

CMNH-ZG 3693)
CMNH-ZG 209)

KUMNH 1808)

CMNH-ZG210)

CMNH-ZG44)

CMNH-ZG115)

ype (CMNH-ZG 3696)

ere ere ve eevee ie

holo

para

para’

para

para
para

para

411

412

circular in outline, paler in color than col-
umn.

Oral disc and tentacles: Diameter of oral
disc of slightly contracted, fixed anemone
approximately equal to that of pedal disc
and column. Center of oral disc somewhat
elevated into oral cone that bears mouth;
mouth elongate along directive axis. Each
siphonoglyph marked with a bright cobalt-
blue spot in life (Fig. 2A, B); color fades
in preservation. Tentacles marginal, slender,
shorter than oral disc diameter, number 40
to 62 (59 in holotype). Each tentacle trans-
lucent whitish to gray, with parallel longi-
tudinal grayish streaks and/or white flecks
on oral surface (Fig. 2A, B). Circular mus-
cles of tentacles endodermal, longitudinal
muscles of tentacles ectodermal (Fig. 3B).
Numerous zooxanthellae in endoderm.

Marginal sphincter muscle: Endodermal,
circumscribed-pinnate to circumscribed-dif-
fuse, with highly branched mesogleal pro-
cesses (Fig. 4B, C).

Mesenteries and internal anatomy: Actin-
opharynx whitish, half to two-thirds length
of column, with two siphonoglyphs each at-
tached to a pair of directive mesenteries.
Distinct marginal stomata; oral stomata not
seen. Mesenteries in 24—39 pairs, arranged
hexamerously in three to four cycles, same
number proximally and distally (Table 1).
Mesentery arrangement irregular in speci-
mens that have regeneration scars. All older
mesenteries, including directives, fertile; all
specimens hermaphroditic, with gametes of
both sexes on same mesenteries or not (Fig.
3C). Zooxanthellae more numerous in en-
doderm of column than in endoderm of
mesenteries. Each specimen may contain as
many as 22 brooded young, early embryos
through young adults with two cycles of
mesenteries and tentacles (Fig. 3D—F);
brooded young posses zooxanthellae.

Mesenterial retractor muscles strong, dif-
fuse to restricted (Fig. 4A). Parietobasilar
muscles well developed, extend half to en-
tire distance between column wall and re-
tractor muscle, with small free pennon dis-
tally (Fig. 4A). Basilar muscles distinct

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(Fig. 3A). Cnidom: Spirocysts, basitrichs,
holotrichs, heterotrichs, microbasic p-mas-
tigophores, microbasic p-amastigophores
(Fig. 5). Sizes and distribution of cnidae
given in Table 2.

Distribution and habitat.—Known from
the middle to northern Pacific coasts of
Honshu, Japan (Fig. 1). Found in high in-
tertidal, attached to Mytilus or in crevices
of rock. Typically forms dense populations.

Etymology.—The species is named after
Dr. K. Atoda, who first identified this as a
new species.

Anthopleura hermaphroditica
(Carlgren, 1899)
lags, Z, SD, ©

Bunodes hermaphroditicus Carlgren, 1899:
23).

Anthopleura hermaphroditica Carlgren,
1899.—Carlgren 1927: 32.—England
1987: 245.

Anthopleura hermafroditica Carlgr. Carl-
gren 1949: 54.—1959: 22.

non Cribrina hermaphroditica Carlgren,
1899.—MceMurrich, 1904: 287.—Daw-
son, 1992: 38.

Material examined.—SMNH 1177 (syn-
type), SMNH 40829, 40830; ZSM (un-
numbered)

Description.—Column and pedal disc:
Freshly collected specimens olive green to
rosy pink, proximal verrucae paler (Fig.
2D). In living, expanded specimens, col-
umn width 15—20 mm, height 17—25 mm.
In contraction, column dome-like, width 4—
10 mm, height 3—12.5 mm. Adhesive, en-
docoelic verrucae (Fig. 6A) in regular ver-
tical rows from margin to limbus; number
of rows 23—42. Verrucae larger and more
prominent distally than proximally; maxi-
mum diameter of distal verrucae 0.5 mm in
preserved specimens. In life, verrucae hold
small stones and pieces of shells. Margin
denticulate, with endocoelic conical projec-
tions that bear 1—3 verrucae on the outer
surface; projection may bear a swollen ac-
rorhagus on the inner surface. Acrorhagi

VOLUME 117, NUMBER 3 413

Fig. 3. Anthopleura atodai, new species (A, B, holotype CMNH-ZG 64; C, paratype CMNH-ZG NSMT-Co
1373; D, paratype CMNH-ZG 3692; E, paratype CMNH-ZG 3695): A, cross section of proximal column showing
directive mesenteries flanked by those of the second (II), third (IID) and fourth (IV) cycles; B—E, cross sections
thorough circumscribed marginal sphincter. Scale Bars: A = 1 mm; B—E = 200 wm. Abbreviations.—4d, directive
mesentery; p, parietobasilar muscle; 1, retractor muscle. Arrow indicating brooded young.

414 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

SRE:

_—

Fig. 4. Anthopleura atodai, new species (A, E, paratype CMNH-ZG 3692; B-C, holotype CMNH-ZG 64;
D, paratype CMNH-ZG 3969; G, paratype CMNH-ZG 3696; H, paratype CMNH-ZG 209): A, longitudinal
section through pedal disc showing basilar muscles; B, longitudinal section through a tentacle; C—D, cross
sections through a mesentery showing both spermatocysts and oocytes; E-H, internally brooded young in the
enteron (E, EK H), and tentacles (G). Scale Bars: A, C, D = 200 pm; B, G, H = 500 pm; E, F = 100 pm (E-
FE Abbreviations.—o, oocyte; s, spermatocysts.

VOLUME 117, NUMBER 3

Fig. 5. Cnidae of Anthopleura atodai, new species
(paratype CMNH-ZG65); see Table 2 for size ranges.
The cnidae of A. hermaphroditica are identical in mor-
phology and in distribution in the body, but differ in
size; see Table 3 for sizes. A-C from tentacles, D-G
from acrorhagus, H-J from column, K-M from actin-
opharynx, N-R from filaments. A, spirocyst; B, basi-
trich-1; C, basitrich-2; D, spirocyst; E, basitrichs; FE
Holotrich; G, Holotrich; H, basitrich-1; I, S-basitrich;
J, heterotrich; K, basitrich-1; L, basitrich-2; M, micro-
basic p-mastigophore; N, basitrich-1; O, basitrich-2; P,
S-basitrich; Q, microbasic p-mastigophore; R, micro-
basic p-mastigophore. Scale Bar = 20 um.

endocoelic, opaque, tan to white, approxi-
mately 0.5 mm tall. Fosse deep. Pedal disc
adherent, roughly circular in outline, paler
in color than distal column.

Oral disc and tentacles: Oral disc diam-
eter of expanded individuals slightly greater
than pedal disc diameter. center of dic ele-
vated into an oral cone that bears mouth;
mouth elongate along directive axis, pale
gray to rosy pink in life. Oral disc with
Opaque marks; marks grouped into six
wedge-shaped zones or forming a stellate
pattern of concentric, lighter and darker
stripes (Fig. 2D, E); pattern fades in pres-
ervation. Tentacles slender, marginal, coni-
cal, shorter than oral disc diameter: approx-
imately 4 mm long in an expanded pre-
served individual; innermost tentacles

415

slightly longer than outermost tentacles.
Tentacles number 34—80, in three to five
cycles. In life, tentacles translucent, typi-
cally with opaque white base and cross-bars
on oral surface (Fig. 2D, E). Circular mus-
cles of tentacles endodermal, longitudinal
muscles ectodermal. Zooxanthellae in en-
doderm.

Marginal sphincter muscle: Endodermal,
circumscribed-pinnate, pedunculate, asym-
metrical, with closely spaced, highly
branched mesogleal processes (Fig. 6C).

Mesenteries and internal anatomy: Actin-
opharynx one-half to two-thirds length of
column, with two aborally prolonged si-
phonoglyphs each attached to a pair of di-
rective mesenteries. Marginal stoma slight-
ly larger than oral stoma. Mesenteries in
24—48 pairs, arranged hexamerously into
three to five cycles, same number proxi-
mally and distally. Mesenteries of first three
cycles typically perfect, those of fourth cy-
cle imperfect. All perfect mesenteries, in-
cluding directives, fertile, each typically
bears both male and female gametes (Fig.
6D). Mesenteries of specimens that contain
many brooded young typically lack gametic
tissue. Zooxanthellae more numerous in en-
doderm of column than in that of mesen-
teries. A specimen may contain as many as
nine brooded young; brooded young up to
2 mm long, with an oral disc diameter of 1
mm, and as many as 20 tentacles. Largest
brooded young zooxanthellate, with small
endocoelic verrucae and marginal projec-
tions.

Mesenterial retractor muscles diffuse-re-
stricted; retractor typically abuts parietal
muscle pennon (Fig. 6E). Parietobasilar
muscles strong, each with a broad pennon
and many short, thick, lateral processes. Pa-
rietal muscle may span as much as half the
distance between the column and the free
edge of the mesentery. Basilar muscles
strong (Fig. 6B).

Cnidom: Spirocysts, basitrichs, hetero-
trichs, holotrichs, microbasic p-amastigop-
hores, microbasic p-mastigophores. Sizes

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

416

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VOLUME 117, NUMBER 3 417

pe 4 “iy Uy
Aa,

»,
See

Fig. 6. Internal anatomy and histology of Anthopleura hermaphroditica. A, longitudinal section through a
verruca; B, longitudinal section through pedal disc showing basilar muscles; C, cross section through sphincter
muscle; D, cross section through a mesentery showing both spermatocysts and oocytes; E, cross section through
proximal to actinopharynx showing mesenteries of first (1), second (II), third (III), and fourth (IV) cycles. Scale
Bars: A = 150 um; B, C = 100 um; D = 200 pm; E = 600m. Abbreviations.—p, parietobasilar muscle; r,

retractor muscle; s, spermatocyst. Arrow indicating oocyte.

418 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

and distribution of cnidae given in Table 3;

cnidae illustrated in Fig. 5. =
ne ee : @ +t
Distribution and habitat.—Known only bs a ¥
from high intertidal zone of Chile. il ii S
Ri Seow 9 oO < as
i és Ean onse) Rim sk CK WEA
Discussion = Sent ae }+
< ere OE Seg ee ae nan
‘ . . xx WOKE <x
Differential diagnosis. “Anthopleura ais Ae x x ore ne x Sa
atodai and A. hermaphroditica belong to a FFs S a = a A 3 x a =
the genus Anthopleura by virtue of pos- NANA lL | Aon lo oo
: : satQoTTFaHHnoIM +9
sessing verrucae, acrorhagi, and columnar SSINOAD ASSO AN SS
heterotrichs. The hermaphroditism and
brooding habit of A. atodai distinguishes it es
from other species of Anthopleura from wa- mS
L . . Im .
ters around Japan, viz. A. mcmurrichi Was- Sy fe on 2 >
silieff, 1908; A. pacifica Uchida, 1938; A. J 2 Al was ©
iridi. : iati a|\| ATA = TT SPO AT eer
Juscoviridis Carlgren, 1949; A. asiatica E| ax Gn ex ma cx qa
Uchida & Muramatsu, 1958; A. kurogane ~l ono dg Pet todamdtes
. aa S| aanit CNW dn a Cada
Uchida & Muramatsu, 1958; A. uchidai BN Fe ay eRe Cay
England, 1992, and from most other nom- QRom~ Boqgnrannms
: é : ae jawortoaetrrnHsoarka
inal species of Anthopleura. It is distin- WAT YAAVTIAITP IDG
ished fi A di by the ch aon Tenaee ttn ae
guished from A. aureoradiata by the char- BOR HOOGR eC aaace
Sn 0 ON OOO) al mea OS I Th a Th |
acter of verrucae at the lower column and
the coloration of the column: in A. aureo-
radiata, the verrucae diminish in size prox-
“ “1)5RSSSSeAnRSAoHA
imally (Parry 1951), and “near the bottom oD Se ne

of the column these become mere mark-
ings” (Stuckey 1909a: 369); in A. atodai,
the verrucae increase in size proximally. In
A. aureoradiata, the coloration of the col-
umn differs distally and proximally (Stuck-
ey 1909a, Parry 1951), whereas in A. ato-
dai, the coloration of the column is uni-
form. Anthopleura atodai is distinguished
from A. handi in its hermaphroditism, pos-

Table 3.—Cnidae of Anthopleura hermaphroditica. Letters refer to Fig. 5. Sizes are given as a length by width range, in 1m; measurements of exceptionally large
or small capsules in parentheses. ““N’’ is the number of specimens examined containing that type of cnidae, ‘‘n’’ is the number of capsules measured. The size range

of cnidae for A. handi, combined from Dunn (1974) and England (1987) is given for comparison.

eee
g 62
£48
session of zooxanthellae, and circumscribed z & = &
marginal sphincter muscle. Al Big ee ee. : 3
Anthopleura atodai and A. hermaphrod- a eleguzc¥d KEQODAR
itica are both hermaphroditic, and brood = ase a 3 ao 2 BOS 2 2
young internally. However, they are distin- oe a s SEOEE ro Ele Ss 6
guished by number of tentacles, coloration, Baaee ae SEZI2a2So8
size of cnidae, and geographical distribu- wes Fe I
tion. The maximum number of the tentacles
observed in members of A. atodai is 62, al
whereas Carlgren (1899) reported a maxi- aS
mum of 90 in specimens of A. hermaphrod- 5 om 3 8
itica. The column of living specimens of A. Q x 3 cs ES 5
atodai is bluish-green or brown; in speci- gs ER: 4 5
mens of A. hermaphroditica, the column is eee At, ra

VOLUME 117, NUMBER 3

419

Table 4.—Summary of differences in size and distribution of cnidae between Anthopleura atodai and A.

hermaphroditica.

Cnidae type and location

Tentacle basitrichs
Acrorhagus holotrichs
Proximal column heterotrichs
Proximal column basitrichs
Filament basitrich-2

Shorter in A. atodai

gray or pink. The nematocysts of the ten-
tacles, acrorhagi, column, and filaments fur-
ther distinguish the two (Table 4).

Taxonomy of Anthopleura hermaphrodi-
tica—The taxonomy of Anthopleura her-
maphroditica has been confused because of
a series of misidentifications and because of
a proposed synonymy between A. herma-
phroditica and A. handi. In the original de-
scription of the species, as Bunodes her-
maphroditicus, Carlgren (1899) mentioned
two notable features: hermaphroditism and
acrorhagi. McMurrich (1904) found speci-
mens of a hermaphroditic actiniid from
Chile that had pseudoacrorhagi, rather than
true acrorhagi and identified these as Cri-
brina hermaphroditica, changing the gener-
ic assignment of Carlgren’s species and
contesting Carlgren’s (1899) assertion that
the species had acrorhagi. Carlgren (1927)
transferred the species to Anthopleura, a ge-
nus characterized as having acrorhagi, but
maintained that the species he had origi-
nally called Bunodes hermaphroditica and
the specimens described by McMurrich
(1904) as C. hermaphroditica were the
same species. However, after examining ad-
ditional material from Chile, Carlgren
(1959) reversed this opinion, and erected a
new species, B. hermaphroditica, which he
attributed to McMurrich.

Carlgren’s (1959) description constitutes
a new combination for C. hermaphroditica
McMurrich 1904, rather than an original
description. According to the International
Code of Zoological Nomenclature (ICZN
1999: Art. 11.6), the name C. hermaphrod-
itica was made available by its subsequent
use as valid (e.g., Clubb, 1908), and its au-

Difference

One size class in A. hermaphroditica; two distinct classes in A. atodai
Narrower in A. atodai

One size class in A. hermaphroditica; two distinct classes in A. atodai

thorship dates from its publication by
McMurrich (1904) as a synonym of Buno-
des hermaphroditica (International Code of
Zoological Nomenclature: Art. 50.7; ICZN
1999). Therefore, the specimens Mc-
Murrich examined constitute the type series
for C. hermaphroditica McMurrich, 1904;
the type specimens of Bunodes herma-
phroditicus Carlgren, 1899 (SMNH 1177)
belong to Anthopleura as they have true ac-
rorhagi with holotrichous nematocysts.

The surviving material from the Lund
University Chile Expedition includes two
recognizable species: A. hermaphroditica
(Carlgren, 1899) and Bunodactis herma-
Dhroditica (McMurrich, 1904). There are
many more specimens belonging to Buno-
dactis hermaphroditica than to A. herma-
phroditica; the difference in number of
specimens collected reflects their abun-
dance in the field (V. Haussermann, pers.
comm.). Specimens belonging to Bunodac-
tis hermaphroditica \ack holotrichous nem-
atocysts in the distal column and in the
proximal column; both of these features are
diagnostic at the level of genus (e.g., Eng-
land, 1987). Specimens of Bunodactis her-
maphroditica have more prominent verru-
cae than specimens of A. hermaphroditica,
especially proximally.

England (1987) suggested that A. her-
maphroditica might be synonymous with A.
handi. We disagree with this proposition of
synonymy because A. hermaphroditica and
A. handi differ in several important re-
spects. Most importantly, members these
two species differ in key life history fea-
tures: members of A. hermaphroditica are
hermaphroditic and zooxanthellate, mem-

420

bers of A. handi are gonochoric and azoox-
anthellate. Furthermore, the basitrichs in
both the distal and proximal column are
larger in members of A. handi than in mem-
bers of A. hermaphroditica. Finally, there is
a considerable disparity in the geographic
range and habitat of the two species: A.
handi is found in the tropical Indo-Pacific
around Malaysia, Singapore, and New
Guinea (Dunn 1978, 1982b; England 1987;
Fautin 1988); A. hermaphroditica is re-
stricted to cold waters of the western Pacific
(Carlgren 1899, 1959).

Biology of Anthopleura atodai.—Antho-
pleura atodai clearly corresponds to Ato-
da’s Anthopleura sp.: the two have identical
distributions, life history, and coloration.
All specimens examined, regardless of size,
were simultaneously hermaphroditic. In ac-
tiniarians, hermaphroditism is unexpectedly
rare (Shick 1991) in view of the “low den-
sity model” of Ghiselin (1969). Among
hermaphrodites, simultaneous hermaphrod-
itsm is the most common mode; known ex-
ceptions include the protandrous hermaph-
rodite Sicyopis (= Kadosactis) commensal-
is (Gravier, 1918) and the gynodioecious
species Epiactis prolifera (Verrill, 1869)
and Cereus pedunculata (Pennant, 1777)
(see Bronsdon et al. 1993, Dunn 1975, Ros-
si 1971).

The reproductive biology of A. atodai re-
mains ambiguous. Isomura et al (2003)
were unable to find gametogenic tissue in
any specimens that they identified as An-
thopleura sp. sensu Atoda, although they
regularly found brooded young. The mes-
enteries of some specimens bore spherical
protuberances proximally that were inter-
preted to be early stages of the brooded
young; from this they inferred that the
brooded young were asexually produced
(Isomura et al. 2003). None of our results
refute an asexual origin for the brooded
young. However, our finding of fertile spec-
imens from the study site of Isomura et al.
(2003), including those that contained both
gametes and brooded young (e.g., Fig. 4A),
indicates that the species is not exclusively

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

asexual, and lends support to the contention
by Isomura et al. (2003) that the Mutsu Bay
population is remarkable in lacking fertile
individuals. In general, the gametes and the
gametogenic region are small in A. atodai,
making it possible that Isomura et al.
(2003) overlooked them in the specimens
they examined. The presence of gametes
does not rule out an asexual origin for the
brooded young; some species of Actinia
have both gametes and asexually produced
young in their enteron (Yanagi et al., 1999).
Therefore, further investigation is necessary
to definitively demonstrate the asexual ori-
gin of the brooded young and to clarify re-
productive ecology of A. atodai.

Acknowledgments

MD supported by NSF- DEB 9978106
(to D.G. Fautin). A part of this work also
supported by Fujiwara Natural History
Foundation (to the first author, KY). We
thank D.G. Fautin (Department of Ecology
and Evolutionary Biology, University of
Kansas, Lawrence KS, U.S.A.) and E. A.
Robson (School of Animal and Microbial
Sciences, The University of Reading, Read-
ing, U.K.) for comments that improved this
manuscript. We especially thank V. Haus-
sermann (ZSM) for permission to use her
photographs, for her generous loan of ma-
terial, and for her insight into the ecology
and biology of A. hermaphroditica. S.D.
Cairns (USNM) and K. Sindemark
(SMNH) also provided specimens. We
thank the staff of Otsuchi Marine Research
Center, Ocean Research Institute, Univer-
sity of Tokyo; Asamushi Marine Biological
Station, Biological Institute of the Faculty
of Science of Tohoku University; Banda
Marine Laboratory of Tokyo University of
Fisheries for help in sampling. We are deep-
ly grateful to the late E. Tsuchida (formerly
Ocean Research Institute, University of To-
kyo) for his kind help in sampling around
Otsuchi and for the opportunity to under-
take this work.

VOLUME 117, NUMBER 3

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(3):423—446. 2004.

New species and new combinations in Rhysolepis
(Heliantheae: Asteraceae)

Harold Robinson and Abigail J. Moore

(HR) Department of Botany, National Museum of Natural History, MRC-166,
Smithsonian Institution, PO. Box 166, Washington, DC 20013-7012;
(AJM) Department of Biology, University of Utah, Salt Lake City, UT 84112

Abstract.—A narrow circumscription of the genus Viguiera Kunth results in
transfer of 58 species of Helianthinae with glabrous stamen filaments, exap-
pendiculate style appendages, and a persistent pappus into Rhysolepis
S.EBlake. Rhysolepis dillonorum from Peru, R. emaciata from Bolivia, and R.
goyasensis, R. hatschbachii, R. laxicymosa, R. santacatarinensis, and R. sub-
truncata from Brazil are new species. Viguiera pazensis and V. procumbens
are placed in synonymy under Rhysolepis helianthoides, and V. misionensis 1s

combined with R. pilosa.

The present study began with a project
by the junior author to clarify the limits of
two Andean species of Viguiera Kunth in
H.B.K. and join in the description of a spe-
cies from Brazil known to be undescribed.
The project was undertaken with the knowl-
edge that none of the species involved were
truly congeneric with the type species of
Viguiera, V. helianthoides Kunth in H.B.K.
= V. dentata (Cav.) Spreng. The arrival of
additional material from Gert Hatschbach
of the Museo Botanico Municipal de Curi-
tiba, led to review of other species prob-
lems and discovery of additional species
needing description. In view of the number
of species involved and because of the ge-
neric redelimitations of Schilling and Pa-
nero (2002), the decision has been made to
abandon the long misapplied name Vigui-
era and use a more phyletically appropriate
generic concept for the species in this study.

Viguiera traditionally has contained spe-
cies related to Helianthus L., but differing
by a more persistent pappus with squamel-
lae. The most recent treatment of Viguiera
in the broad sense was that of Blake (1918).
Blake’s treatment excluded some genera
such as Tithonia Desf. with broadened, fis-
tulose peduncles (La Duke 1982); Syncre-

tocarpus S. F Blake (1916) with a glabrous
strip just inside the lateral margins of its
achenes that was misinterpreted as a wing;
and Rhysolepis S. E Blake (1917) with
transverse corrugations on its paleae. The
broad Blake concept of Viguiera included
some elements now placed in Hymenoste-
phium Benth. in Benth. & Hook.f., but ex-
cluded others (Schilling and Panero 2002).
Some single species once placed in Vigui-
era have been moved to other genera, in
example a Peruvian species named by
Blake in 1918, Viguiera acutifolia, has been
transferred to Pappobolus (Panero 1992)
and a Mexican species included in Viguiera
by Blake (1924) was subsequently trans-
ferred to Stuessya (Turner & Davies 1980).

In a brief review of members of the Hy-
menostephium group, Robinson (1977) re-
tained the broad concept of Viguiera in
spite of the realization that the type species
of Viguiera was individually distinctive
with pubescent anther filaments and a small
apical appendage on the branches of the
style. Hymenostephium was retained in Vi-
guiera because it had an apical appendage
on the style branches and was technically
closer to the type of Viguiera than most
other species placed in the latter genus. The

424

needed generic revisions of the concept of
Viguiera were fully initiated by Schilling
and Panero (2002); but the South American
species and their relatives in Mexico with
exappendiculate styles have not yet been
treated.

The South American species are in need
of transfer to some genus other than Vigui-
era. The problem has been that none of the
synonyms given by Blake (1918) seems to
be applicable. Leighia Cass. belongs to the
group, but that name is a later homonym of
Leighia Scop. As noted by Blake, the type
of Harpalium Cass., H. rigidum (Desf.)
Cass. ( = Helianthus rigidus Desf.), is not
a Viguiera. Other Blake synonyms, Heliom-
eris Nutt. and Bahiopsis Kellogg, are con-
sidered separate genera (Schilling & Panero
2002). The type of Gymnolomia Kunth in
H.B.K., after some confusion, proved to be-
long to Eleutheranthera Poit. ex Bosc.
(Robinson 1992). Thus, none of the syno-
nyms from Blake (1918) can be used. A
name is found, however, outside the syn-
onymy of Viguiera as circumscribed by
Blake in 1918. His genus Rhysolepis, in
spite of its sometimes weakly transversely
corrugated paleae, is not distinct from the
group treated here, and so the name can be
applied.

Rhysolepis S. F Blake, Contr. Gray Herb.
52: 36 (1917).—Type: Viguiera pal-
merit A. Gray
Leighia Cass. in E Cuvier, Dict. Sci.
Nat. ed. 2. 25: 435 (1822).—Type: He-
lianthus linearis Cav. Not Leighia
Scop. (1777). = Ethulia L.f.

Annual to perennial herbs or shrubs; of-
ten with tubers or with fusiform nodes on
roots. Stems and leaves usually strigose, pi-
lose, or hispid. Leaves alternate or opposite,
sessile or petiolate, filiform to ovate, lan-
ceolate, oblong, or broadly rounded; blade
often trinervate with secondary veins near
and subparallel to lower margin; margins
entire to serrate. Inflorescences usually with
1—6 heads, sometimes heads over 50; pe-
duncles usually elongate, 3-30 cm long, of-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ten stout, not enlarged and fistulose distally;
involucre broadly campanulate; bracts in 2—
5 series, gradate to subequal, oblong to
ovate or lanceolate, at base usually with in-
durated ribs, at tips herbaceous, appressed
or reflexed, rounded to acute; receptacle
convex to conical; pales persistent, partially
enclosing achenes, mostly ribbed and in-
durate, sometimes transversely corrugated,
usually with blunt apex. Ray florets usually
8—24, sterile, sometimes lacking; corollas
yellow, with yellow or orange resin ducts
along veins. Disk florets usually 40—200,
tightly packed, bisexual; corollas yellow or
greenish-yellow, 5-lobed, with basal tube
usually 0.5—1.0 mm long and narrow, usu-
ally scabrid on abruptly broadened base of
throat and on backs of lobes, with yellow
or orange resin ducts along 5 veins of
throat; anther filaments without hairs or pa-
pillae; thecae blackish, shortly hastate at
base; endothecial cells with nodes on trans-
verse walls; apical appendage usually yel-
low, blackish in some annual species, ovate,
concave abaxially often with cluster of
glands in concavity; style with resin ducts
outside of veins not restricted to branches;
style branches spreading radially, with tuft
of hairs or papillae at tip, without apical
appendage, with stigmatic papillae covering
whole inner surface. Achenes compressed,
with or without setulae, without differenti-
ated intramarginal bare strip; walls with
phytomelanin interrupted by striations of
pale cells; pappus mostly persistent, with
pair of awns usually longer than squamellae
on margins between awns, but awns some-
times not longer than squamellae. Chro-
mosome numbers n = 17, 34.
Rhysolepis was described from Mexico
and has previously been credited with only
three Mexican species as recognized by
Robinson (1972):
Rhysolepis kingii H. Rob., Phytologia 24:
210 (1972).

Rhysolepis morelensis (Greenm.) S. E
Blake, Contr ‘Gray silesba 523550
(1917).

VOLUME 117, NUMBER 3

Viguiera morelensis Greenm., Proc.
Amer. Acad. 40: 40 (1904).
Rhysolepis palmeri (A. Gray) S. FE Blake,
Contr. Gray Herb. n.s. 52: 37 (1917).
Viguiera palmeri A. Gray in S.Watson,
Proc. Amer. Acad. 22: 427 (1887).

The broadened concept of Rhysolepis
recognized here includes the rather overlap-
ping Blake (1918) sections and series, Ten-
uifoliae consisting of perennial herbs with
linear leaves, solitary heads and involucral
bracts 2-seriate and subequal; Revolutae
with perennial herbs or subshrubs of the
Chilean and Argentine Andes with large
solitary heads and involucral bracts 2—5-se-
riate, gradate and lanceolate; Grandiflorae
with perennial herbs having one or few
large, long-pedunculate heads and having
few leaves with the lowest opposite and
scale-like; Aureae, primarily Andean, in-
cluding annuals to shrubby perennials with
broad leaves and involucral bracts mostly
3—5-seriate, usually gradate, lanceolate, and
with herbaceous tips not strongly differen-
tiated; Bracteatae, mostly of Brazil and
Paraguay, including herbaceous perennials
similar to the Aureae but with involucral
bract tips shortly and abruptly herbaceous
and blunt; Leighia, mostly Mexican, but
similar to the Aureae and Bracteatae with
involucral bracts strongly gradate, oblong
and usually with an abrupt herbaceous tip;
Trichophylla consisting of slender virgate
perennials with linear to filiform leaves,
revolute leaf margins and involucral bracts
lanceolate to linear-lanceolate; and subge-
nus Verbalesia containing perennial herbs
with pappus awns equalled in length by and
partially fused to the squamellae. The fol-
lowing new combinations agree, to a con-
siderable extent, with species concepts of
Blake (1918), although that work left ques-
tions about the real distinctions of many
species. As a result, more recent synony-
mies are taken into account, and other syn-
onymies are to be expected. Many poorly
known species are omitted.

425

Rhysolepis anchusifolia (DC.). H. Rob.
& A. J. Moore, comb. nov.

Leighia anchusaefolia DC., Prodr. 5: 580
(1836).

L. dissitifolia DC., Prodr. 5: 581 (1836).

L. immarginata DC., Prodr. 5: 581 (1836).

L. lomatoneura DC., Prodr. 5: 581 (1836).

L. stenophylla Hook. & Arn., J. Bot. 3: 313
(1841).

L. baldwiniana Nutt., Trans. Amer. Philos.
Soc. ser. 2, 7:365 (1841).

Viguiera stenophylla (Hook. & Arn.) Gri-
seb., Goett. Abh. 24: 193 (1879).

V. anchusaefolia (DC.) Baker in Mart., FI.
bras. 6(3): 222 (1884). Argentina, Brazil,
Uruguay.

Rhysolepis arenaria (Baker) H. Rob. &
A. J. Moore, comb. nov.

Viguiera arenaria Baker in Mart., Fl. bras.
6(3): 228 (1884). Brazil, north central
Sao Paulo.

Rhysolepis aspilioides (Baker) H. Rob. &
A. J. Moore, comb. nov.

Viguiera aspilioides Baker in Mart., FI.
bras. 6(3): 228 (1884). Brazil, Matto
Grosso.

Rhysolepis atacamensis (Phil.) H. Rob.
& A. J. Moore, comb. nov.

Viguiera atacamensis Phil., Anales Mus.
Nac. Chile, Segunda Secc., Bot. 1891: 48
(1891). Chile.

Rhysolepis australis (S.EBlake) H. Rob.
& A. J. Moore, comb. nov.

Viguiera australis S. F Blake, Contr. Gray
Herb. n.s. 54: 148 (1918). Chile.

Rhysolepis bakeriana (S. FE Blake) H.
Rob. & A. J. Moore, comb. nov.

Viguiera bakeriana S. F Blake, Contr. Gray
Herb. n.s. 54: 130 (1918). Brazil, Minas
Gerais.

426

Rhysolepis bishopii (H. Rob.) H.Rob. &
A. J. Moore, comb. nov.

Viguiera bishopii H. Rob., Phytologia 45:
458 (1980). Bolivia.

Rhysolepis bracteata (Gardn.) H. Rob. &
A. J. Moore, comb. nov.

Viguiera bracteata Gardn., London J. Bot.
7: 404 (1848). Brazil. Distrito Federal,
Goias, Minas Gerais.

Rhysolepis breviflosculosa (S. FE Blake)
H. Rob. & A. J. Moore, comb. nov.

Viguiera breviflosculosa S. F Blake, Contr.
Gray Herb. n.s. 54: 158 (1918). Uruguay.

Rhysolepis brittonii (Hochr.) H. Rob. &
A. J. Moore, comb. nov.

Viguiera brittonii Hochr., Bull. New York
Bot. Gard. 6: 294 (1910). Peru.

Rhysolepis discolor (Baker) H. Rob. &
A. J. Moore, comb. nov.

Viguiera discolor Baker in Mart., Fl. bras.
6(3): 228 (1884). Brazil, Minas Gerais.

Rhysolepis ellenbergii (Cuatrec.) H. Rob.
& A. J. Moore, comb. nov.

Viguiera ellenbergii Cuatrec., Proc. Biol.
Soc. Wash. 77: 146 (1964).

Peru. A second specimen from the type
locality is as follows: Peru. Cuzco: Prov.
Urubamba, ruinas de Machu Picchu, high
above Rio Urubamba, 80 km WNW of
Cuzco, rock walls, rock piles, terraces &
cliffs, Intyhuatana (Solar Observatory);
2500-2600 m, 27 May 1963, Ugent 5376
(US).

Rhysolepis fabrisii (Saenz) H. Rob. & A.
J. Moore, comb. nov.

Viguiera fabrisii Saenz, Darwiniana 22: 50
(1979). Argentina.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Rhysolepis fusiformis (S. E Blake) H.
Rob. & A. J. Moore, comb. nov.

Viguiera fusiformis S. F Blake, Contr. Gray
Herb. n.s. 54: 145 (1918). Bolivia.

Rhysolepis gardneri (Baker) H. Rob. &
A. J. Moore, comb. nov.

Viguiera gardneri Baker in Martt., Fl. bras.
6(3): 224 (1884).

Originally described from Brazil, Goias.
Two more recent collections matching the
type photograph are: Brazil. Goidas, Piren-
Opolis (Morro da Caixa Dagua); cerrado
seco, arborizado, com pedras no solo, su-
jeito ao fogo periddico; planta com 80 cm,
ramificada inflorescéncia terminais, flores
roxas, 23 Apr 1976, Heringer 15560 (UB,
US); Municipio de Niquelandia, entrada no
km 8 da Rodovia Niquelandia/Uruagu; Fa-
zenda Trairas. Morro. Relévo ondulado;
14°29'19"S, 48°33'19"W. Cerrado com mui-
tas pedras de cor branca; arbusto, ca. 70 cm
de altura; flores com corola amarela e an-
teras alaranjadas. Nome comum: margarida;
13 Apr 1996; Mendonca, Marquete, Fon-
seca & Oliveira 2453 (UB, US).

Rhysolepis gilliesii (Hook. & Arn.) H.
Rob. & A. J. Moore, comb. nov.

Leighia gilliesii Hook. & Arn., J. Bot.
(Hooker) 3: 313 (1841).

Helianthus heteropappus Gill. ex Hook. &
Arn., J. Bot. (Hooker) 3: 314 (1841),
nom. nud.

Viguiera gilliesii (Hook. & Arn.) Hieron.,
Actas Acad. Nac. Ci. Cordoba 4: 39
(1882).

Flourensia hispida Phil., Anales Univ.
Chile 36: 186 (1870). Argentina, Chile.

Rhysolepis grandiflora (Gardn.) H. Rob.
& A. J. Moore, comb. nov.

Leighia grandiflora Gardn. in Field &
Gardn., Sert. Pl. t. 54-55 (1844).

Viguiera grandiflora (Gardn.) Gardn., Lon-
don J. Bot. 7: 404 (1848).

VOLUME 117, NUMBER 3

Rhysolepis guaranitica (Chod.) H. Rob.
& A. J. Moore, comb. nov.

Viguiera guaranitica Chod., Bull. Herb.
Boiss. ser. 2, 3: 724 (1903). Argentina,
Brazil, Paraguay.

Rhysolepis helianthoides (L. Rich.) A. J.
Moore & H. Rob., comb. nov.
Fig. |

Sanvitalia helianthoides L. Rich. in Willd.,
Sp. Pl. 3: 2190 (1803).

Helianthus procumbens Pers., Syn. Pl. (Per-
soon) 2: 475 (1807).

Viguiera pazensis Rusby, Mem. Torrey Bot.
Club 3(3): 59 (1893).

Viguiera pflanzii Perkins, Bot. Jahrb. Syst.
49: 226 (1913)..

Viguiera punensis S. FE Blake, Bot. Jahrb.
Syst. 54, Beibl. 119: 48 (1916). Argen-
tina, Bolivia, Peru.

The present complex was maintained as
two separate species by Blake (1918) based
on longer, relatively narrower leaf shape
and more prominent leaf venation in V. pa-
zensis. Separation was maintained by Saenz
as recently as 1979 based on larger, ovate-
lanceolate rather than ovate to oblong
leaves, multiple rather than single heads per
stem, and smaller involucres in V. pazensis.
We could not separate the species using
these characters, nor pubescence type or
shape of the involucral bracts. Tips of the
involucral bracts were sometimes reflexed
and thus looked different from bracts with-
out reflexed tips, but the lengths and shapes
were the same.

The broadened concept of Rhysolepis he-
lianthoides is characterized by leaves tu-
berculate-pilose adaxially, pilose abaxially
with hairs denser on veins; stems ribbed
and villous; and involucral bracts oblanceo-
late, subequal, often recurved, and with an
indurate base and herbaceous apex. In ad-
dition, the achenes tend to have rather read-
ily deciduous awns and squamellae, a char-
acter reportedly shared with R. lanceolata
(Blake 1918).

427

The concept of Viguiera pazensis in this
study includes two isotypes, Bang 44 (US).
Some more southern material might prove
distinct, and the name Helianthus ataca-
mensis Phil. (not Viguiera atacamensis
Phil.) is for the present omitted from the
synonymy. For an additional specimen that
was determined as V. pazensis, but is not
this species, see Rhysolepis dillonorum A.
J. Moore & H. Rob. below.

Rhysolepis hilairei (S. EF Blake) H. Rob.
& A. J. Moore, comb. nov.

Viguiera hilairei S. FE Blake, Contr. Gray
Herb. 54: 153 (1918). Brazil, Minas Ger-
ais.

Rhysolepis hypoleuca (S. F Blake) H.
Rob. & A. J. Moore, comb. nov.

Viguiera hypoleuca S. F Blake, Contr. Gray
Herb. n.s. 54: 165 (1918). Brazil, Matto
Grosso.

Rhysolepis incana (Pers.) H. Rob. & A.
J. Moore, comb. nov.

Helianthus incanus Pers., Syn, Pl. 2: 475
(1807).

Helianthus aureus Kunth in H.B.K., Nov.
Gen. Sp., ed fol. 4: 176 (1818).

Harpalium aureum (Kunth) Cass., Dict.
Sci. Nat. 25: 438 (1822).

Viguiera chimboensis Hieron., Bot. Jahrb.
Syst. 29: 38 (1900).

Viguiera lehmannii Hieron., Bot. Jahrb.
Syst, 29: 38 (1900).

Viguiera aurea (Kunth.) Hieron., Bot.
Jahrb. Syst. 28: 608 (1901).

Viguiera incana (Pers.) S. E Blake, Contr.
U.S. Nat. Herb. 26: 252 (1930). Ecuador.

Rhysolepis kunthiana (Gardn.) H. Rob.
& A. J. Moore, comb. nov.

Viguiera kunthiana Gardn., London J. Bot.
7: 399 (1848). Brazil, Goias.

428 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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i

AR THGERIN( 2004

Fig. 1. Rhysolepis helianthoides (L. Rich.) A. J. Moore & H. Rob.., A. Habit. B. Head with ray florets
removed and involucral bracts not recurved. C. Head showing ray florets. D. Receptacular pale. E. Ray corolla
showing lack of style. F Disk floret showing striated achene with pappus of awns and squamellae. G. Disk
corolla in section, showing filaments and anthers with small glands on outer surfaces of anther appendages. H.
Disk style showing branches with continuous stigmatic area on inner surfaces and apex with hairs but no
appendage. Drawn mostly from Bang 44 (US, isotype of Viguiera pazensis Rusby); C. from Buchtien 8579
(US).

VOLUME 117, NUMBER 3

Rhysolepis lanceolata (Britton) H. Rob.
& A. J. Moore, comb. nov.

Viguiera lanceolata Britton, Bull. Torrey
Bot. Club 19: 149 (1892).

V. mandonii Sch.Bip. ex Rusby, Mem. Tor-
rey Bot. Club 3(3): 60 (1893).

Helianthus szyszylowiczii Hieron., Bot.
Jahrb. Syst. 36: 491 (1905). Bolivia,
Peru.

Rhysolepis linearifolia (Chod.) H. Rob.
& A. J. Moore, comb. nov.

Viguiera linearifolia Chod., Bull. Herb.
Boiss. ser. 2, 2: 392 (1902).

Viguiera trichophylla Dusén, Ark. Bot.
9(15): 30, f. 12 & t. 7. f. 4 (1910). Brazil,
Goids, Matto Grosso, Parana; Paraguay.

Rhysolepis linearis (Cav.) H. Rob. & A.
J. Moore, comb. nov.

Helianthus linearia Cav., Icon., 3: 9, t. 218
(1794)[1795].

Helianthus squarrosus Kunth in H.B.K..,
Nov. Gen. & Sp., ed. fol. 4: 174, t. 377
(1818).

Leighia elegans Cass., Dict. Sci. Nat. 25:
A435 (1822).

Leighia linearis (Cav.) DC., Prodr. 5: 581
(1836).

Viguiera linearis (Cav.) Sch.Bip. ex
Hemsl., Biol. Centr-Amer., Bot. 2:178
(1881). Mexico.

Rhysolepis macbridei (S. F Blake) H.
Rob. & A. J. Moore, comb. nov.

Viguiera macbridei S. EF Blake, J. Wash.
Acad. Sci. 16: 218 (1926). Peru.

Rhysolepis macrocalyx (S. E Blake) H.
Rob. & A. J. Moore, comb. nov.

Viguiera macrocalyx S. E Blake, Contr.
Gray Herb. 54: 171 (1918). Brazil, Minas
Gerais.

429

Rhysolepis macropoda (S. F Blake) H.
Rob. & A. J. Moore, comb. nov.

Viguiera macropoda S. F Blake, Contr.
Gray Herb. 54: 128 (1918). Brazil, Minas
Gerais.

Rhysolepis macrorhiza (Baker) H. Rob.
& A. J. Moore, comb. nov.

Viguiera macrorhiza Baker in Mart., FI.
bras. 6(3): 225 (1884). Paraguay.

Rhysolepis media (S. E Blake) H. Rob.
& A. J. Moore, comb. nov.

Viguiera media S. FE Blake, Contr. Gray
Herb. 54: 138 (1918). Ecuador.

Rhysolepis mollis (Griseb.) H. Rob. & A.
J. Moore, comb. nov.

Viguiera mollis Griseb., Abh. K6nigl. Ges.
Wiss. Gottingen 19: 183 (1874).

Helianthus argentinus Saenz, Darwiniana
22: 64 (1979) Argentina.

We do not know why Saenz (1979) ex-
cluded the species from Viguiera in his
treatment, creating the new name Helian-
thus argentinus. Panero (1992) was correct
in returning the species to Viguiera as then
delimited.

Rhysolepis nervosa (Gardn.) H. Rob. &
A. J. Moore, comb. nov.

Viguiera nervosa Gardn., London J. Bot. 7:
403 (1848). Brazil, Goias.

Rhysolepis nudibasilaris (S. E Blake) H.
Rob. & A. J. Moore, comb. nov.

Viguiera nudibasilaris S. FE Blake, Contr.
Gray Herb. 54: 149 (1918). Brazil, Minas
Gerais.

Rhysolepis nudicaulis (Baker) H. Rob. &
A. J. Moore, comb. nov.

Viguiera nudicaulis Baker in Mart., FI.
bras. 6(3): 228 (1884). Uruguay.

430

Rhysolepis oblongifolia (Gardn.) H. Rob.
& A. J. Moore, comb. nov.

Viguiera oblongifolia Gardn., London J.
Bot. 7: 402 (1848).

Rhysolepis oblongifolia was described
from Brazil, Goias. Some more recent col-
lections include: Brazil. Matto Grosso: Ser-
ra do Roncador, Mun. de Barra do Gargas,
230 km along new road NNE of village of
Xavantina, 6.0 km S of Coérrego dos Porcos,
30 km due S of 12°51’S, 51°45’W. ca. 450
m, 26 Nov 1969, Eiten & Eiten 9547 (SP,
US); 209 km NNE of Xavantina; 9 Dec
1969; Eiten & Eiten 9818 (SP, US); Minas
Gerais. 56 km along road NE of Barrocao,
towards Porteirinha, 2400 ft.; 21 Jan 1981,
King & Bishop 8585 (MO, US); Brasilandia
de Minas, 1 Jun 2001, Soares 32] (BHCB,
US); Maranhao, Balsas, approx. 25 km
along road west from Balsas to fazenda of
Sr. Damiao; 7°40’S, 46°10’W; 4 Dec 1981;
Jangoux et al. 1783 (US).

Rhysolepis obtusifolia (Baker) H. Rob. &
A. J. Moore, comb. nov.

Viguiera obtusifolia Baker in Mart., FI.
bras. 6(3): 226 (1884). Brazil, Goias?

Rhysolepis ovatifolia (DC.) H. Rob. & A.
J. Moore, comb. nov.

Leighia ovatifolia DC., Prodr. 5: 583
(1836).

Viguiera ovatifolia (DC.) Baker in Mart.,
Fl. bras. 6(3): 226 (1884).

The type is from Brazil, Sao Paulo. Ad-
ditional specimens seen from Parana match
the type photograph: Jaguariahyva, ad mar-
ginem silvulae, 19 Apr 1910; Dusén 9723
(US)(det. Dusén as Viguiera robusta). Ja-
guariahyva opp., in campo, 740 m.s.m, 5
May 1914, G. Jonsson 262a (US)(det. Mal-
me as V. robusta).

Rhysolepis peruviana (A. Gray) H. Rob.
& A. J. Moore, comb. nov.

Viguiera peruviana A. Gray, Proc. Amer.
Acad. Arts 5: 124 (1861-62).

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Viguiera weberbaueri S. FE Blake, Bot.
Jahrb. Syst. 54, Beibl. 119: 49 (1916).
Peru.

Rhysolepis pilicaulis (S. FE Blake) H.
Rob. & A. J. Moore, comb. nov.

Viguiera pilicaulis S. F Blake, Contr. Gray
Herb. 54: 164 (1918).

Rhysolepis pilicaulis was described from
Paraguay. A recent collection has been seen
from Brazil: Matto Grosso do Sul. Rod.
BR-267, proximo do km 447, descida da
chapada (Mun. Guia Lopes de Laguna, 9
Mar 2003, G. & H. Hatschbach & Barbosa
74393 (MBM, US). The Brasilian specimen
is most like the Field Museum type photo-
graph of the now destroyed, broad-leaved
Berlin specimen. The inflorescence is char-
acteristically rather profusely branched with
short peduncles, and there are only 8 or 9
short, slender rays while Blake cited 10 to
11. The species has antrorse prorulosity in-
side the disk corolla throat.

Rhysolepis pilosa (Baker) H. Rob. & A.
J. Moore, comb. nov.

Viguiera pilosa Baker in Mart., Fl. bras.
6(3): 223 (1884).

Viguiera malmei S. F Blake, Contr. Gray
Herb. 54: 151 (1918).

Viguiera misionensis Saenz, Darwiniana
2 O2 (D7).

Viguiera misionensis of northern Argen-
tina shows no obvious differences from R.
pilosa from southern Brazil in Parana, Rio
Grande do Sul, Santa Catarina.

Rhysolepis pusilla (A. Gray) H. Rob. &
A. J. Moore, comb. nov.

Tithonia pusilla A. Gray, Proc. Amer. Acad.
Arts 5: 124 (1861-62).

Viguiera pusilla (A. Gray) S. E Blake,
Contr. Gray Herb. 54: 160 (1918). Peru.

VOLUME 117, NUMBER 3

Rhysolepis radula (Baker) H. Rob. & A.
J. Moore, comb. nov.

Viguiera radula Baker in Mart., Fl. bras.
6(3): 223 (1884). Brazil, Minas Gerais.

Rhysolepis retroflexa (S. E Blake) H.
Rob. & A. J. Moore, comb. nov.

Viguiera retroflexa S. F Blake, Contr. Gray
Herb. 54: 146 (1918). Bolivia.

Rhysolepis revoluta (Meyen) H. Rob. &
A. J. Moore, comb. nov.

Helianthus revolutus Meyen, Reise Erde 1:
311 (1834).

Helianthus lanceolatus Meyen, Reise Erde
1: 311 (1834), not V. lanceolata Britton

Flourensia corymbosa DC., Prodr. 5: 592
(1836).

Viguiera poeppigii A. Gray, Proc. Amer.
Acad. Arts 19: 6 (1883).

Viguiera corymbosa (DC.) S. E Blake,
Proc. Amer. Acad. Arts 49: 349 (1913).

Viguiera revoluta (Meyen) S. E Blake,
Contr. Gray Herb. 54: 121 (1918). Ar-
gentina, Chile.

Rhysolepis robusta (Gardn.) H. Rob. &
A. J. Moore, comb. nov.

Viguiera robusta Gardn., London J. Bot. 7:
403 (1848). Brazil, Goias.

Rhysolepis rojasii (S. E Blake) H. Rob.
& A. J. Moore, comb. nov.

Viguiera rojasii S. KF Blake, Contr. Gray
Herb. 54: 179 (1918). Paraguay.

Rhysolepis salicifolia (Hassl.) H. Rob. &
A. J. Moore, comb. nov.

Viguiera salicifolia Hassl., Repert. Spec.
Nov. Regni Veg. 14: 274 (1916).

Viguiera villaricensis S. FE Blake, Contr.
Gray Herb. 54: 152 (1918). Argentina,
Paraguay.

431

Rhysolepis simsioides (S. EF Blake) H.
Rob. & A. J. Moore, comb. nov.

Viguiera simsioides S. F Blake, Bot. Jahrb.
Syst. 54. Beibl. 119: 48 (1916). Peru.

Rhysolepis sodiroi (Hieron.) H. Rob. &
A. J. Moore. comb. nov.

Helianthus sodiroi Hieron., Bot. Jahrb.
Syst. 29: 41 (1900).

Viguiera sodiroi (Hieron.) S. FE Blake,
Contr. Gray Herb. 54: 139 (1918). Ec-
uador.

Rhysolepis speciosa (Hassl.) H. Rob. &
A. J. Moore, comb. nov.

Viguiera speciosa Hassl., Repert. Spec.
Nov. Regni Veg. 14: 272 (1916).

Viguiera simulans S. F Blake, Contr. Gray
Herb. 54: 127 (1918).

Rhysolepis speciosa has been known
from Paraguay; Brazil, Matto Grosso. It
also occurs in the Distrito Federal with
specimens previously identified as Viguiera
squalida as follows: Peunsula Norto, 1000
m, s. d., Valério de Carvalho dos grupos
II (UB, US); Reserva Ecolégia do IBGE,
7 Nov 1977, Heringer et al. 249 (IBGE,
US); Area do Cristo Redentor: 15°57'07"S,
47°53'37'"W, 19 Oct 1988, Azevedo 180
(IBGE, US); Reserva Ecologica do IBGE,
Campo Limpo; 21 Aug 1990, Silva et al.
1009 (IBGE, US); Cristo Redentor, 10 Oct
1990, Brochado 70 (IBGE, US); Tampao
das parcelas de campo sujo do Projeto
Fogo—IBGE, 9 Dec 1991, Landim de Sou-
za 83 (IBGE, US); Ecolégica do IBGE,
15°56’41"S, 47°53'07’W, 7 Nov 1994, Apa-
recida da Silva 2457 (IBGE, US).

Rhysolepis squalida (S. Moore) H. Rob.
& A. J. Moore, comb. nov.

Viguiera squalida S. Moore, J. Bot. 42: 37
(1904). Brazil, Goids, Matto Grosso,
Matto Grosso do Sul.

432

Rhysolepis subdentata (S. FE Blake) H.
Rob. & A. J. Moore, comb. nov.

Viguiera subdentata S. FE Blake, Contr.
Gray Herb. 54: 131 (1918). Brazil, Minas
Gerais.

Rhysolepis tenuifolia (Gardn.) H. Rob. &
A. J. Moore, comb. nov.

Viguiera tenuifolia Gardn., London J. Bot.
7: 400 (1848). Brazil, Minas Gerais.

Rhysolepis tuberculata (S. FE Blake) H.
Rob. & A. J. Moore, comb. nov.

Viguiera tuberculata S. F Blake, Contr.
Gray Herb. 54: 151 (1918). Brazil, Minas
Gerais.

Rhysolepis tuberosa (Griseb.) H. Rob. &
A. J. Moore, comb. nov.

Viguiera tuberosa Griseb., Abh. KO6nigl.
Ges. Wiss. Gottingen 24: 192 (1879). Ar-
gentina; Brazil, Rio Grande do Sul, Uru-

guay.

Rhysolepis tucumanensis (Hook. & Arn.)
H. Rob. & A. J. Moore, comb. nov.

Leighia tucumanensis Hook. & Arn., J. Bot.
(Hooker) 3: 314 (1841)

Viguiera stenophylla (Hook. & Arn.) Gri-
seb. var. discoidea Griseb., Abh. Konig].
Ges. Wiss. Gottingen 24: 193 (1879).

Viguiera discoidea (Griseb.) S. E Blake,
Contr. Gray Herb. 54: 157 (1918).

Viguiera oligodonta S. FE Blake, Contr.
Gray Herb. 54: 146 (1918). Argentina.

Rhysolepis weddellii (Sch.Bip. ex S. FE
Blake) H. Rob. & A. J. Moore,
comb. nov.

Viguiera weddellii Sch.Bip. ex S. FE Blake,
Contr. Gray Herb. 54: 126 (1918). Boliv-
ia; Brazil, Goias-Matto Grosso.

In addition to the species listed above we
include the following seven previously un-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

described species. Of the new species, the
ones from Bolivia and Santa Catarina, Bra-
zil, seem to fit Blake’s series Aureae;
whereas the others fit his series Bracteatae.
One character seen in the new species
seems to partially reenforce the distinction.
All of the new members of the Bracteatae
except R. subtruncata have bands of pro-
rulose cells on the inner surface of the disk
corolla throats, midway between the veins
and often also along the veins. Prorulosity
is the condition where elongate cells have
the upper ends projecting as papillae. The
two new species in the Aureae lack such
prorulose bands. The specimens were de-
scribed partially from dissections of florets
mounted in Hoyer’s solution (Anderson
1954).

Rhysolepis dillonorum A. J. Moore & H.
Rob., sp. nov.
Fig. 2

Type: Peru. Arequipa: Prov. Caraveli,
Lomas of Atiquipa, ca. 10.5 km N of turn-
off to Atiquipa, 584 km S of Lima; ca.
150—200 m, 1 Nov 1983, M. O. Dillon &
D. Dillon 3775 (holotype US; isotype F).

E speciebus omniibus in habitis fruticosis
et in indumento appresse strigulosis et in
bracteis involucri plerumque obtusis dis-
tincta.

Shrub to 1 m high, moderately and alter-
nately branched at 30—45° angles; roots not
seen; stem tan to dark brown, closely ap-
pressed-strigulose, glabrescent with age.
Leaves usually opposite in middle of
branches, alternate at base of branches and
distally, not decrescent except near heads;
petioles none to 0.2 cm long, bases some-
times continuous across node; blades ovate
to oblong-ovate, 1.5—4.2 cm long, 0.7—2.2
cm wide, base broadly acute to rounded,
margins entire, apex acute, both surfaces
densely appressed-strigulose, abaxially with
scattered glandular dots, triplinervate from
near base, secondary veins reaching distal
¥Y%. Heads borne singly on long branches or
with 2 or 3 heads on short branches; brac-

VOLUME 117, NUMBER 3

UNITED STATES

3026289

NATIONAL HERBARIUM

Fig. 2. Rhysolepis dillonorum A. J.

Rhyselepis Aillenorum AdMoorer jit

Holotype £

QAO

0073069;

PLANTS OF PERU
Field Museum of Natural History

DEPTO: AREQUIPA PROV: CARAVELI

ASTERACEAE
Viguiera pazensis [Rusby

Lomas of Atiquipa, ca. 10.5 km N of turn-off to
Atiquipa. KM 584 S of Lima. ca. 150-200 m.

%
Erect shrub to 1 m.; ray & disc florets yellow.

1 Nov 1983

M. 0. Dillon & D, Dillon 3779

Planis collected under the sponsorship of the National Geographic Society (Grant No. 2706-83]
Distributed by Field Museum of Natural History

Moore & H. Rob., holotype, Dillon & Dillon 3775 (US).

434

teoles decrescent, oblong, 1.1—0.4 cm long;
peduncles 3.5—17 cm long, 0.5—5.0 cm from
last bracteole, appressed-strigulose. Invo-
lucre 0.4—0.6 cm high, 1.2—1.5 cm diam.;
bracts 2—3-seriate, obovate, gradate, 4—8
mm long, 2—4 mm wide, 3—5-nerved, tips
obtuse to short-acuminate, indurate in prox-
imal % to %, distally herbaceous, abaxially
and adaxially appressed-strigulose especial-
ly on tips; paleae obovate, indurate, ca. 7.5
mm long, ca. 1.0—1.5 mm wide, scabridu-
lous on tip, apex short-acute. Ray florets
13-14; corollas yellow, tube ca. 1.2 mm
long, sparsely scabridulous; limb oblong-el-
liptical, 1.5 cm long, 0.5—0.8 cm wide,
sparsely scabridulous abaxially, apex 3-
lobed. Disk florets at least 50; corolla yel-
low, ca. 5 mm long, tube | mm long, sca-
bridulous; throat 3 mm long, slightly cam-
panulate at base, scabridulous proximally,
glabrous distally, with vertical bands of an-
trorsely prorulose cells inside, lobes 1 mm
long, glabrous outside, papillose inside es-
pecially near margins; anther thecae 2 mm
long; appendages yellow, 0.6—0.75 mm
long, ca. 0.45 mm wide. Ray achenes ca.
3.5 mm long, ca. 0.7 mm wide, sericeous
on margins, with pappus crown ca. 0.1 mm
high. Disk achenes 3.5 mm long, 0.9 mm
wide, sericeous with setulae over whole
surface; pappus awns 2.0—2.2 mm long,
fimbriate-margined, squamellae ca. 4, 1.0—
1.2 mm long, ca. 1 mm wide, margins fim-
briate. Pollen 30—33 ym in diam. in Hoy-
er’s solution.

Rhysolepis dillonorum is_ presently
known only from the type collection. The
specimen was earlier identified as Viguiera
Pazensis; it was seen as distinct in a recent
review of the latter by the senior author of
the new species. The low elevation at 150—
200 m near the coast, the shrubby habit, the
appressed minute hairs on the stems, leaves,
and involucral bracts, and the blunt tips of
the involucral bracts are all distinctive. The
supposed Andean relatives are found at
2000 m or above, are more herbaceous,
have longer, mostly spreading hairs, and

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

have more lanceolate, subequal involucral
bracts.

The blunt involucral bracts and the ver-
tical bands of prorulose cells inside the disk
corolla throat seem to relate the new species
to members of Blake’s section Bracteatae
that are most common in Brazil and distin-
guish the species from the section Aureae
to which V. pazensis ( = R. helianthoides)
belongs. Opposite leaves are common on
the specimen, but the branching is alternate,
and the basal nodes of the branches have
alternate leaves.

Rhysolepis emaciata H. Rob. & A. J.
Moore, sp. nov.
Fig. 3

Type: Bolivia. Cochabamba: 10 NE;
2465 m; Campero, pajonal de Elyonurus
tripsacoides, 2 May 1999, Antezana 1276
(holotype US, isotype MO).

E speciebus aliis boliviensis in seriebus
Aureis in ramis nullis in foliis dense spir-
aliter insertis et in bracteis involucri ca. tris-
eriatis gradatis differt.

Slender subshrub or shrub 0.4—0.6 m tall,
apparently unbranched above base; roots
not seen; stems reddish-brown, densely his-
pid with long hairs. Leaves rather densely
spirally inserted, sessile; laminae herba-
ceous, lanceolate, 1.5—2.8 cm long, 0.5—0.8
cm wide, base rounded, margins often with
single blunt tooth near basal 4, subentire to
remotely undulate distally, apex acute, mu-
cronulate, adaxial surface densely scabrous
with slender hairs, abaxially densely villous
with white hairs and densely gland-dotted;
triplinerved from near base, reaching to %
leaf length. Inflorescence example seen
with single terminal head, with leaves on 7
cm below head becoming smaller, upper-
most bractlike; peduncle 2.5 cm long from
last foliiform bract, densely villous. Head
with involucre 1 cm high, 2 cm wide; bracts
ca. 3-seriate, oblong-lanceolate, gradate, 6—
10 mm long, 1.5—2.0 mm wide, appearing
herbaceous throughout, villous with white
hairs abaxially, without distinct cilia on

VOLUME 117, NUMBER 3

Fig. 3.

UNITED STATES

3376064

NATIONAL HERBARIUM

I@-"7 ays

aoe ya

HOM

00730693

BRS

Rhysalepis emaciata HRbsAd. Hooxe-
Heletype ;

BOLIVIA
ASTERACEAE
Viguiera
Cochabamba: Campero
Pajonal de Elyonurus tripsacoides, 10

NE. Arbusto lenoso, inflorescencia
amarilla.

2465 m

02-05-1999

Antezana, C. 1276
MISSOURI BOTANICAL GARDEN HERBARIUM (MO)

Rhysolepis emaciata H. Rob. & A. J. Moore, holotype, Antezana 1276 (US).

435

436

margins distally, tips acute to slightly mu-
cronulate, erect on inner bracts, shortly re-
curved on other bracts, scabridulous on
both surfaces; paleae yellowish-tan, indu-
rate, oblong, 7.5 mm long, ca. 2.5 mm
wide, tip minutely hispidulous, acute to,
sometimes, trifid. Ray florets 17—18; corol-
las yellow, tube 1.5 mm long, hispidulous,
limb oblong, ca. 1.0—1.1 cm long, 0.4 cm
wide, abaxial surface strigose and gland-
dotted, apex 2- or 3-lobed. Disk florets 50
or more; corollas darker yellow, ca. 6 mm
long, tube ca. 1 mm long, scabridulous,
throat ca. 4 mm long, campanulate and sca-
bridulous at base, smooth inside, lobes 1.0—
1.2 mm long, strigulose outside, papillose
inside; anther thecae 2.5—3.0 mm long, with
slender basal hastation much longer than
collar, essentially short-tailed; appendage
0.58—0.70 mm long, 0.35—0.41 mm wide.
Achenes ca. 2.5—3.0 mm long, 0.8 mm
wide, sparsely strigulose with stiff setulae;
pappus color a very light tan, awns 3.0—3.5
mm long, with fimbriate margins, squamel-
lae ca. 5, 1.0-1.5 mm long, 0.2—0.5 mm
wide, deeply fimbriate. Pollen grains 25—28
jum in diam. in Hoyer’s solution.

Rhysolepis emaciata is known only from
the type collection. Relation might be ex-
pected to R. australis, R. fusiformis, R. he-
lianthoides, and R. lanceolata of the Boli-
vian Andes, but the latter all have thicker
stems, more obvious branching, and only
about 2 series of subequal involucral bracts.
The spirally inserted leaves of R. emaciata
characteristically seem to contract slightly
in width near the basal fourth. The bases of
the anthers seem unusually long and tailed
for a member of the Heliantheae.

Rhysolepis goyasensis H. Rob. & A. J.
Moore, sp. nov.
Fig. 4

Type: Brazil. Goids: Serra Geral do Pa-
rana, ca. 3 km S of Sao Joao da Alianga,
near Riacho, ca. 850 m, gallery forest and
adjacent cerrado, 15 Mar 1971, /rwin, Har-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ley & G. L. Smith 31821 (holotype US; iso-
types NY, UB, US).

A R. breviflosculosam in pubescentibus
caulis et involucri similis sed in foliis su-
perioribus decrescentibus et in laminis base
non cordatis differt.

Subshrub to 1 m high; usually un-
branched between base and inflorescence;
part of rhizome seen, roots moderately
stout, spreading, without evident fusiform
enlargements; stem reddish-brown to tan,
pilose to lanulose, denser above, hairs
spreading to retrorse. Leaves alternate, re-
duced to bracteoles distally, often gradually
decrescent; petioles ca. 2 mm long; blades
oblong-elliptical, 0.6—6.5 cm long, 0.3—2.1
cm wide, at base broadly acute, margins en-
tire, apex short-acute to short-acuminate,
adaxially villous with tubercle-based hairs,
abaxially villous, triplinervate from near
base, secondary veins reaching distal 4 of
blade. Inflorescences unbranched or with 1—
3 branches on each side, conic to cylindri-
cal when multibranched, elongate branches
shorter than main axis, spreading at ca. 45°
angles; bracteoles narrowly oblong-ellipti-
cal 0.7—3.0 mm long; peduncles 3.5—5.0 cm
long, 1-2 mm long from last bracteoles,
lanulose as in stems. Heads usually 1—5; in-
volucre 0.9—1.1 cm high, 1.2—1.9 cm wide;
bracts ca. 3-seriate, oblong to oblong-lan-
ceolate, gradate, 10-15 mm long, 3—4 mm
wide, 3-nerved, tips abruptly acute to
slightly acuminate, slightly indurate at base
or herbaceous throughout; paleae rather ob-
long, coriaceous, ca. 8 mm long, ca. 2 mm
wide, apex short-acute to mucronulate. Ray
florets 14-15; corollas yellow; tube 3.5 mm
long, pilosulous; limb 1.5 cm long, 0.4 cm
wide, pilosulous abaxially on veins, apex 2
or 3 lobed. Disk florets ca. 50; corolla yel-
low, ca. 5 mm long, tube 0.7—1.0 mm long,
nearly glabrous; throat 2.5—3.0 mm long,
slightly campanulate at base, glabrous, in-
side with vertical bands of antrorsely pro-
rulose cells, lobes 0.7—1.0 mm long, acute,
sparsely scabrid outside, papillose inside;
anther thecae 2.5 mm long, appendages yel-
low, 0.55—0.60 mm long, 0.3—0.4 mm wide.

VOLUME 117, NUMBER 3 437

Rhy sole "$ goyasensis He p.zAs,Teore.
Neletype:

THES NEW YORK BOTANICAL GARDEN
Plants of the Planalto do Brasil
Extado do Golds

No. 37821 Serra Geral do Parana

Viguiera quinqueremis Blake
(ex descript. of phyllaries)
6 det. John Pruski (NY), 1992
UNITED STATES

Erect simple or few-branched subshrub
Ga. Im tall. Ligules yellow; disc yellow-
brown. Gallery forest and adjacent
3314050 cerrado, ca. 3km S. of Sao Joao da Aliansa,
near riacho, ca. 850m. elev.

NATIONAL HERBARIUM TT H. S, Irwin, R, M. Harley, G, L, Smith 15 March 4971

Field work condocted with the collaboration of tho Univers(dade do Brasilia and
06 tho Instituto de Pesquisas © Experimentaclo Agricola do Norte, supported In
part by funds from the Natlonal Sclence Foundation.

Fig. 4. Rhysolepis goyasensis H. Rob. & A. J. Moore, holotype, Irwin, Harley & Smith 31821 (US).

438

Achenes 3 mm long, 0.8—1.0 mm wide, gla-
brous except few, small, marginal setulae;
pappus awns 2.0—3.1 mm long, minutely
scabrid on margins and keel, squamellae ca.
8, 1.1-1.7 mm long, 0.3-0.5 mm wide,
margins fimbriate. Pollen 27-30 wm in
diam. in Hoyer’s solution.

Paratypes: Brazil. Goids: Sao Joao de Al-
ianga, estrada para Vaozinho, campo cer-
rado, solo rochoso, 9 Feb 1994, G. & M.
Hatschbach 60230 & Silva (MBM, US);
Corrente (Mun. Sao Joao da Alianga), cam-
po cerrado, solo rochoso, 20 Feb 2000, G.
& M. Hatschbach & O. S. Ribas 70471
(MBM, US).

Rhysolepis goyasensis has pubescence of
the stems and involucres reminiscent of that
in R. breviflosculosa far to the south in Uru-
guay. The new species differs by the leaf
blades lacking cordate subamplexicaul ba-
ses and by the decrescent size of the distal
leaves of the stem. The new species seems
related to the R. robusta species group, but
has few or single heads or a narrowly conic
to cylindrical inflorescence borne well be-
yond the larger stem leaves.

Rhysolepis hatschbachii H. Rob. & A. J.
Moore, sp. nov.
Bigs

Type: Brazil. Matto Grosso do Sul: Ro-
dovia Bonito, Campo dos Indios, proximo
de Trés Morros (Mun. Bonito); encosta do
morro; solo calcario, 10 Mar 2003, G. &
M. Hatschbach & E. Barbosa 74469 (ho-
lotype MBM, isotype US).

A R. gardneri in formibus capituli similis
sed in foliis abaxialiter dense pilosulis et in
pedunculis ebracteatis longioribus et in lim-
bis radii abaxialiter glabris distincta.

Perennial herb or subshrub to 1 m high,
with lateral branches ascending at 35—40°
angles; roots not seen; stems tan to dark
brown, densely hispid to strigose. Leaves of
main stems alternate, 4.5—8.5(11) cm long,
1.5—2.8(4.2) cm wide, on branches often
opposite, 2.0-5.5 cm long, 0.5—1.5 cm
wide; petioles 1-2 mm long; laminae her-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

baceous, oblong-elliptical, base obtuse,
margins entire to remotely serrulate, apex
short-acute and apiculate, adaxial surface
densely pilosulous with bases of hairs often
enlarged, abaxial surface densely scabridu-
lous on veins, less densely pilosulous be-
tween veins, gland-dotted; triplinervate
with strongly ascending secondary veins
reaching middle or distal % of blade. Inflo-
rescence of few heads terminal on stems
and branches; peduncles 9-30 cm long
without leaves or bracts, strigulose to hispid
with white hairs, hairs denser below heads;
involucre 10-13 mm high, 13-20 mm
wide; bracts broadly 3—4-seriate, slightly
unequal, oblong with obtuse to acute tips,
7-11 mm long, ca. 4 mm wide, bases of
inner bracts indurate and strongly ribbed,
abruptly shortly herbaceous and sometimes
recurved at tips, outer bracts canescent with
white, densely strigulose pubescence, mar-
gins densely fimbriate with short cilia, inner
surface usually glabrous, rarely strigulose
near tip; paleae oblanceolate, ca. 9 mm
long, 1.5 mm wide, acute, essentially gla-
brous. Ray florets 9-14; corollas yellow,
tube ca. 1 mm long, sparsely pilosulous;
limb narrowly elliptical, ca. 1.7—2.4 cm
long, 0.5—0.6 cm wide, abaxially glabrous,
apex minutely bilobed. Disk florets 35—45
or more; corollas yellow, ca. 5 mm long,
tube ca. 1 mm long, scabridulous, throat ca.
3 mm long, sparsely scabridulous on nar-
rowly campanulate base, with vertical
bands of antrorsely prorulose cells inside,
lobes ca. | mm long, nearly glabous abax-
ially, papillose inside; anther thecae ca. 2.3
mm long; appendage yellow, 0.6—0.7 mm
long, ca. 0.4 mm wide. Sterile ray ovaries
with pair of squamellae 0.5—0.9 mm long;
disk achenes ca. 4 mm long, ca. 1.2 mm
wide; awns 3.0—3.5 mm long, squamellae
narrow, 0.5—0.8 mm long, fimbriate. Pollen
grains ca. 32 pm in diam. in Hoyer’s so-
lution.

Paratype: Brazil. Matto Grosso do Sul:
Serra de Bodoquena, Fazenda Bodoquena,
Reserva da Tercola (Mun. Miranda); Mata,
5-8 m, Sopé de morro, solo argiloso raso,

VOLUME 117, NUMBER 3 439

Your ad,

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EG rys-

A ‘a
PREFEITURA MUNICIPAL DE CURITIBA | ERBARIO N:

MUSEU BOTANICO MUNICIPAL

Asteraceae

Rodovia Bonito—Campo dos Indios, proximo de Trés Morros
(Mun. Bonito) Mato Grosso do Sul

UNITED STATES Rhysolepis htchbochi He 6

Haw. G. Hatschbach, M. Hatschbach & E. Barbosa 74469, 10.111.2003

Isofypes Ereta, Im, capitulo amarelo. Encosta do morro; solo calcario.

3441382

era keen WAC

00730695

PMC -DPP 004

Fig. 5. Rhysolepis hatschbachii H. Rob. & A. J. Moore, isotype, G. & M. Hatschbach & Barbosa 74469
(US).

440

encharcado, 17 Mar 1995, A. Pott & al.
7026 (US, MBM).

Rhysolepis hatschbachii is known from
only the two cited collections. The paratype
was previously determined as near R. gard-
neri of Goias, and the latter is possibly the
closest relative. Differences include the
long peduncles of the latter having many
foliiform bracts, its involucral bracts being
distinctly narrower and pale at the base, and
its rays being shorter and puberulous abax-
ially. The general habit of the new species
is closer to R. ovatifolia of Sao Paulo and
Parana, but that species has distinctive nar-
rower involucral bracts that are essentially
glabrous except for the densely ciliate mar-
gins.

Rhysolepis laxicymosa H. Rob. & A. J.
Moore, sp. nov.
Fig. 6

Type: Brazil. Minas Gerais: Serra do Ca-
bral, estradad para Francisco Dumont
(Mun. Joaquim Felicio); campo rupestre,
950 m, 16 May 2001, G. & M. Hatschbach
& Barbosa 72088 (holotype MBM, isotype
US).

A speciebus novis R. goyasensem similis
sed in caulibus non lanulatis et in inflores-
centiis laxe cymiformibus et in limbis radii
brevibus distinctis.

Erect perennial herb or subshrub 50—90
cm high, apparently unbranched between
base and inflorescence; roots not seen; stem
reddish-tan, pilose to strigose or thinly vil-
lous. Leaves alternate; petioles ca. 1 mm
long; laminae coriaceous, oblong-elliptical,
3.7—1.5 cm long, 1.6—0.6 cm wide, decres-
cent toward inflorescence, base rounded to
broadly acute, margins entire or remotely
1—3-subserrulate, apex short-acute and
slightly mucronulate, adaxial surface
sparsely strigose and densely scabridulous,
abaxial surface with prominent veins and
prominulous veinlets, densely strigose on
veins, strigulous to subsericeous between
veins, gland-dotted; triplinerved from near
base, secondary veins reaching distal % or

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

more of blade. Inflorescences are sparingly
branched, cymiform, branches long, as-
cending at ca. 30° angles; with bracteoles
mostly at branch bases 1.5—0.7 cm long,
0.6—0.3 cm wide; peduncles 6—23 cm long,
strigose, more densely villous near heads,
with bracteoles 7-3 mm long, 3-1 mm
wide. Heads ca. 5; involucre 0.5—0.6 cm
high, 1.0—1.2 cm wide; bracts ca. 3-seriate,
oblong, somewhat gradate, 3.0—6.5 mm
long, 0.8-1.2 mm wide, tips obtuse to
short-acute, outer bracts indurate in basal %,
herbaceous in distal 4%, inner bracts almost
completely indurate with broad sclerified
bands between veins, exposed surfaces
densely pilosulous; paleae pale tan, papery,
lanceolate to oblong, ca. 7 mm long, ca. 1.5
mm wide, scaberulous at base and tip,
gland-dotted at tip. Ray florets ca. 18; co-
rollas yellow, tube ca. 1.2 mm long, sca-
bridulous; limb broadly oblong, 6.5 mm
long, 1.8—3.0 mm wide, puberulous abaxi-
ally on veins, apex trilobed. Disk florets
30-35 or more; corollas yellow, 4 mm long,
basal tube | mm long, scabridulous, throat
ca. 2.5 mm long, base slightly campanulate
and scabridulous, with vertical bands of an-
trorsely prorulose cells inside, especially
midway between veins, lobes ca. 0.7 mm
long, pilosulous distally outside, papillose
inside; anther thecae ca. 1.8 mm long; ap-
pendages yellow, 0.4—0.5 mm long, 0.33—
0.38 mm wide. Achenes ca. 3.5 mm long,
ca. 1.1 mm wide, sericeous with slender se-
tulae; pappus whitish, awns mostly 2.0—2.5
mm long, fimbriate on margins and midrib;
squamellae 5 or 6, ca. 1 mm long, 0.2—0.5
mm wide, margins fimbriate. Pollen grains
22-28 wm in diam. in Hoyer’s solution.
Rhysolepis laxicymosa seems mostly
closely related to R. goyasensis, but it 1s
smaller in all parts. The pubescence of the
stem is shorter, the inflorescence is more
slender with fewer bracts, the involucre is
smaller with narrowly oblong bracts, and
the rays are scarcely twice as long as the
involucre. In the length of its rays, R. lax-
icymosa is closer to R. subtruncata, also of
Goias, which has distinctive subtruncate

VOLUME 117, NUMBER 3 44]

nm
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MUSEU BOTANICO MUNICIPAL
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Asteraceae

Vequiers oblong ele Crate,

re Serra a Cabral, estrada para Francisco Dumont (Mun, Joaquim Felicio)
Q\e ayy Minas Gerais

UNITED STATES G. Hatschbach, M. Hatschbach & E. Barbosa 72088, 16.V.2001
.

Ereta, 50cm, capitulo amarelo. Campo mupestre. Alt.: 950m.

3407290

NATIONAL HERBARIUM

PMC - OPP 004

Fig. 6. Rhysolepis laxicymosa H. Rob. & A. J. Moore, isotype, G. & M. Hatschbach & Barbosa 72088
(US).

442

leaves that are not decrescent below the in-
florescence.

Rhysolepis santacatarinensis H. Rob. &
A. J. Moore, sp. nov.
Fig. 7

Type: Brazil. Santa Catarina: Serra do
Faxinal (Mun. Praia Grande), paredOes ro-
chosos, 1200 m, 15 Apr 1993, G. & M.
Hatschbach 59135 & J. M. Silva (holotype
MBM, 2 isotypes US).

A R. pilosam in foliis lanceolatis et brac-
teis involucris lanceolatis similis sed in fo-
liis distincte petiolatis et in nervis pinnatis
et in caulibus densius lanulatis differt.

Subshrub or shrub to 1 m high, moder-
ately branched; roots not seen; stems tan to
reddish-brown, villous, hairs denser near
heads. Leaves alternate; petioles 0.2—1.7 cm
long, sometimes slightly winged, villous;
laminae herbaceous, lanceolate, 4-17 cm
long, 0.4—3.5 cm wide, base and apex at-
tenuate to acuminate, margins remotely cre-
nate-serrulate, adaxially tuberculate-sca-
brous, abaxially densely canescent, pilose
to subvillous, denser on veins, with glan-
dular dots; venation pinnate or essentially
pinnate. Inflorescence with | or 2 heads per
branch, often overtopped by leaves; pedun-
cles 0.2—2.0 cm long. Heads 4—8; involucre
0.75—1.25 cm high, 2—3 cm wide, 3.5 cm
wide in fruit; bracts 2—3-seriate, narrowly
lanceolate to narrowly oblanceolate, 12—22
mm long, 2-3 mm wide, apices acuminate
to mucronulate, tips strongly recurved, bas-
al % to % indurate, 5-ribbed, distally her-
baceous, abaxially villosulous, adaxially at
tip pilosulous to subglabrous, sparsely
gland-dotted, margins finely ciliate; paleae
oblong, ca. 9-11 mm long, ca. 2 mm wide,
indurate, to 7-ribbed, apex acute and mu-
cronulate, sometimes with teeth, glabrous
with scabridulous midvein. Ray florets ca.
23; corollas yellow, tube ca. 1 mm long,
sparsely puberulous; limbs narrowly ellip-
tical, 1.5-3.5 cm long, 0.3-0.4 cm wide,
apex 1- or 2 -(3-) lobed, abaxially puberu-
lous, gland-dotted. Disk florets to 120 or

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

more; corollas yellow, 5—6 mm long, tube
1.5 mm long, glabrous, throat 3.5 mm long,
base moderately campanulate, scabridulous
on base and veins, smooth inside, lobes
0.5—1.0 mm long, acute, sometimes sparse-
ly scabridulous outside, papillose on distal
Y% inside; anther thecae 2.5—3.0 mm long;
appendage yellow, 0.7—0.8 mm long, 0.3—
0.4 mm wide. Achenes 4 mm long, ca. |
mm wide, glabrous except for marginal se-
tulae near pappus; awns 2-3 mm long,
squamellae separated into broad segments,
ca. 0.5 mm long, fimbriate. Pollen 25—28
ym in diam. in Hoyer’s solution.

Paratypes: Brazil. Santa Catarina: Mun.
Lauro Miller; 20 km west of Lauro Miller,
lower and middle slopes of serra by Rio do
Rastro, 700-1000 m, 3 Apr 1957, L. B.
Smith & R. Klein 12339 (FLOR, US); Rod.
SC-438, Serra do Rio do Rastro (Mun. Lau-
ro Miiller); Pareddes rochosos; 1000 m, 7
Apr 1991, G. & M. Hatschbach & E. Bar-
bosa 55311 (MBM, US).

Rhysolepis santacatarinensis would be-
long to the series Aureae of Blake (1918)
on the basis of its lanceolate involucral
bracts, and it would key to various species
in the Blake key depending on the emphasis
given to the dense canescent pubescence of
the abaxial faces of its leaves. Its distribu-
tion in southern Brazil and shape of its
leaves suggest closest relation to R. pilosa,
which has much sparser pilose pubescence,
usually no petiole, and much smaller heads.
The large heads with 120 or more disk flo-
rets distinguish the new species from most
other members of the genus in Brazil and
elsewhere. The venation of the leaves is
also distinctive, lacking strongly ascending
lateral veins at the base. The basal second-
ary veins are either strictly pinnate or only
slightly more ascending.

Rhysolepis subtruncata H. Rob. & A. J.
Moore, sp. nov.
Fig. 8

Type: Brazil. Goias: Chapada dos Vead-
eiros, ca. 42 km N of Alto do Paraiso, ca.

VOLUME 117, NUMBER 3 443

PREFEITURA MUNICIPAL DE CURITIBA
MUSEU BOTANICO MUNICIPAL

M.
Asteraceae

ISOTYPE OF: NC. Viguiers

~ inensis Rb. a AN. Moore
Rhysolepis Sanlacaferinensis HRA» 4 \0C: Serra do Faxinal (mune Praia Grande) Santa
Catarina
L6G. G. & M- Hatschbach 59135 & JeMe Silva, 15-IV-1993

3422582 “Alte: 1200 me

. UNITED STATES NATIONAL HERBARIUM IANO

PMC - OPP 004

Fig. 7. Rhysolepis santacatarinensis H. Rob. & A. J. Moore, isotype, G. & M. Hatschbach 59135 & Silva
(US).

444 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

THB NEW YORK BOTANICAL GARDEN
Plants of the Planalto do Brasil
Estado do Golds

Rhysolepi's subtrun cia HReb, 2 No. 33151A Chapada dos Veadeiros

Li A,d.Aoore Viguiera sp.

Hololype ‘

UNITED STATES

Subshrub to ca. 2.5m tall. Ligules yellow;
disc yellow-brown. Riacho margin in
cerrado. Cerrado on rocky Slopes and

3313597 adjacent campo, ca. k2km N. of Alto do
Paraiso, ca. 1250m. elev.

NATIONAL HERBARIUM

(ho Toatiluto de Derquisas 0 Experimentacio Awrfeola do Norte, supported In

H, S, Irwin, R, M. Harley, G, L, Smith 25 March 1971
7 ,

HON NILALA Field work condveted with tho collaboration of tho Universidade do Drasilia and
4 davon part by funds from tho Natlonal Scteoce Foundation.

Fig. 8. Rhysolepis subtruncata H. Rob. & A. J. Moore, holotype, Jrwin, Harley & Smith 33151A (US).

VOLUME 117, NUMBER 3

125 m elev., riacho margin in cerrado, on
rocky slopes and adjacent campo, 25 Mar
1971, Irwin, Harley & G. L. Smith 33151A
(holotype US, isotypes NY, UB).

E speciebus aliis in foliis coriaceis saepe
subtruncatis et in ramis inflorescentibus
longis valde ascendentibus et in floribus ra-
diis brevibus differt.

Subshrub to 2.5 m high, with few or no
branches between base and inflorescence;
roots not seen; stems tan to reddish-brown,
strigose to stiffly pilose. Leaves alternate,
petioles O-1 mm long, 1—2 mm broad,
densely villosulous abaxially; laminae co-
riaceous, obovate to cuneate, 1.8—4.5 cm
long, 0.8—2.4 cm wide, scarcely smaller but
more remote up to inflorescence, base cu-
neate, margins slightly crenulate-serrulate
above, apex subtruncate to scarcely retuse,
adaxial surface nearly smooth, hairs stri-
gose with enlarged bases, abaxial surface
with prominulous veinlets, pilose to thinly
sericeous, triplinervate from near base, lat-
eral veins reaching distal 4. Inflorescence
loosely corymbiform, with 2 or 3 long
branches on each side, ascending at ca. 30°
angles, pilose; bracts foliiform, mostly % to
Y% as large as leaves, mostly at bases of
branches, with few bracteoles on distal
branches; peduncles 0.4—2.0 cm long be-
yond bracteoles. Heads ca. 9; involucre 0.8
cm high, ca. 1.5 cm wide; bracts ca. 2-se-
riate, lanceolate, 5-8 mm long, 1-2 mm
wide, acute to slightly acuminate, basal 4%
to % indurate, apices herbaceous, appressed
to slightly spreading, abaxially puberulous,
adaxially at tip pilosulous; paleae rather ob-
long, obtuse, ca. 5.5 mm long, ca. 1.5 mm
wide, indurate, glabrous or with midvein
strigulose. Ray florets ca. 20; corollas yel-
low, tube ca. 1.2 mm long, pilosulous; limb
broadly oblong, 5—6 mm long, 3.5—4.0 mm
wide, apex unlobed or 2-lobed, abaxially
pilosulous mostly on veins, Disk florets ca.
50?; corollas yellow-brown, 4 mm long;
tube 0.8 mm long, sparsely scabrid, throat
2.5 mm long, base scabrid, narrowly cam-
panulate, glabrous distally, smooth inside,
lobes deltate, ca. 1 mm long, scabrid out-

445

side; anther thecae 1.8—2.0 mm long; ap-
pendage yellow, 0.35—0.40 mm long, 0.45—
0.55 mm wide. Achenes (immature) 2.5
mm long, 0.8—1.0 mm wide, setulae over
whole surface, sericeous; pappus awns Ca.
1.5 mm long, squamellae ca. 0.5 mm long,
deeply fimbriate. Pollen grains 22—26 wm
in diam. in Hoyer’s solution.

Rhysolepis subtruncata has distinctive
cuneate, coriaceous leaves and ascending
branches of the inflorescence reaching the
level of the terminal central head. The rays
are very short compared to many other spe-
cies of the genus. The leaves below the in-
florescence are not or are scarcely decres-
cent. The throats of the disk corollas lack
the vertical bands of prorulose cells found
in other species of sect. Bracteatae. Any
additional collections should be readily
identifiable by the leaf shape and by the
overall habit of the leafy plants and inflo-
rescence.

Acknowledgments

A. J. Moore was supported in this study
by the National Science Foundation Re-
search Experience for Undergraduates pro-
gram Award Number DBI-0243512. The
extensive technical help of Marjorie Know-
les is also acknowledged. The drawing of
Rhysolepis helianthoides was prepared by
Alice Tangerini, staff illustrator, Depart-
ment of Botany.

Literature Cited

Anderson, L. E. 1954. Hoyer’s solution as a rapid
mounting medium for bryophytes.—Bryologist
57:242-247.

Blake, S. F 1916. Compositae novae imprimis andinae
Weberbauerianae.—Bot. Jahrb. Syst. 54:47—51.

. 1917. New and noteworthy Compositae, chief-

ly Mexican.—Contr. Gray Herb. 52:16—59.

. 1918. A revision of the genus Viguiera.—

Contr. Gray Herb. n.s. 54:1—205, pl. 1-3.

1924. New American Asteraceae.—Contr.
U.S. Nat. Herb. 22(8):—viii, 587-661, pl. 54—
63, 1X=x1.

La Duke, J. C. 1982. Revision of Tithonia.—Rhodora
84:453—522.

Panero, J. L. 1992. Systematics of Pappobolus (Aster-

446 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

aceae-Heliantheae).—Syst. Bot. Monogr. 36:1—
195.

Robinson, H. 1972. Studies in the Heliantheae I. A
new species of Rhysolepis.—Phytologia 24:
209-210.

. 1977. Studies in the Heliantheae (Asteraceae).

VIII. Notes on genus and species limits in the

genus Viguiera.—Phytologia 36:201—215.

1992. New combinations in Elaphandra

Strother (Ecliptinae-Heliantheae-Asteraceae).—

Phytologia 72:144—151.

Saenz, A. A. 1979. El género Viguiera (Compositae)
en la Argentina.—Darwiniana 22:45—66.
Schilling, E. E., & J. L. Panero. 2002. A revised clas-
sification of subtribe Helianthinae (Asteraceae:
Heliantheae). I. Basal lineages.—J. Linn. Soc.,
Bot. 140:65—76.

Turner, B. L., & EF Davies. 1980. Stuessya (Asteraceae:
Heliantheae), a new genus from southcentral
Mexico.—Brittonia 32:209—212.

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CONTENTS

A review of the North American subspecies of the Great Blue Heron (Ardea herodias)
Robert W. Dickerman
A new species of Microgale (Lipotyphla: Tenrecidae: Oryzorictinae) from the Forét des Mikea of
southwestern Madagascar Steven M. Goodman and Voahangy Soarimalala
Designation of the type species of Musaraneus Pomel, 1848 (Mammalia: Soricomorpha: Soricidae)
Neal Woodman
The mammals of Palawan Island, Philippines
Jacob A. Esselstyn, Peter Widmann, and Lawrence R. Heaney
A new species of Tropidonophis (Serpentes: Colubridae: Natricinae) from the D’ Entrecasteaux Islands,

Papua New Guinea Fred Kraus and Allen Allison
A new species of snake of the genus Omoadiphas (Reptilia: Squamata: Colubridae) from the Cordillera
Nombre de Dios in northern Honduras James R. McCranie and Franklin E. Castafieda

A new species of Kolpotocheirodon (Teleostei: Characidae: Cheirodontinae: Compsurini) from Bahia,
northeastern Brazil, with a new diagnosis of the genus
Luiz R. Malabarba, Flavio C. T. Lima, and Stanley H. Weitzman
Astyanax biotae, a new species of stream fish from the Rio Paranapanema basin, upper Rio Parana sys-
tem, southeastern Brazil (Ostariophysi: Characiformes: Characidae)
Ricardo M. C. Castro and Richard P. Vari
Tetragonopterus lemniscatus (Characiformes: Characidae), a new species from the Corantijn River
basin in Suriname Ricardo C. Benine, Gabriela Zanon Pelicao, and Richard P. Vari
Longipalpa saltatrix, anew genus and species of the meiofaunal family Nerillidae (Annelida: Polychaeta)
from an anchihaline cave in Bermuda
Katrine Worsaae, Wolfgang Sterrer, and Thomas M. Iliffe
Neostrengeria lemaitrei, a new species of freshwater crab from Colombia (Crustacea: Decapoda:
Pseudothelphusidae), and the vertical distribution of the genus Martha R. Campos
A new species of Agostocaris (Caridea: Agostocarididae) from Acklins Island, Bahamas
Fernando Alvarez, José Luis Villalobos, and Thomas M. Iliffe
A new species of caridean shrimp of the family Stylodactylidae from the eastern Pacific Ocean
Mary K. Wicksten and Joel W. Martin
A new pedunculate barnacle (Cirripedia: Heteralepadidae) from the Northwest Atlantic
L. Buhl-Mortensen and W. A. Newman
Two new species of seven-spined Bathyconchoecia from the North Atlantic and Indian oceans

(Crustacea: Ostracoda: Halocypridae) Louis S. Kornicker and J. A. Rudjakov
The hermaphroditic sea anemone Anthopleura atodai n. sp. (Anthozoa: Actiniaria: Actiniidae) from
Japan, with a redescription of A. hermaphroditica Kensuke Yanagi and Marymegan Daly

New species and new combinations in Rhysolepis (Heliantheae: Asteraceae)
Harold Robinson and Abigail J. Moore

SMITHSONIAN INSTITUTION LIB

01118 3

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251

266

271

303

311

37)

330

339

346

363

368

377

385

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423

G2 H- ISSN 0006-324X

BH X
NH PROCEEDINGS oF THE

BIOLOGICAL SOCIETY
or WASHINGTON

JAN 1 4 20d

20 DECEMBER 2004

VOLUME 117
NUMBER 4

THE BIOLOGICAL SOCIETY OF WASHINGTON
2003-2004
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PROCEEDINGS

Editor: Richard C. Banks

Associate Editors

Plants: Carol Hotton Invertebrates: Stephen L. Gardiner
Insects: Wayne N. Mathis Christopher B. Boyko
Vertebrates: Gary R. Graves Janet W. Reid

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(4):447—487. 2004.

Studies on western Atlantic Octocorallia (Coelenterata: Anthozoa).
Part 5: The genera Plumarella Gray, 1870; Acanthoprimnoa, n. gen.;

and Candidella Bayer, 1954

Stephen D. Cairns and Frederick M. Bayer

Department of Zoology, National Museum of Natural History, Smithsonian Institution, PO. Box

37012, Washington, D.C. 20013-7012, U.S.A., e-mail: cairns.stephen@nmnh.si.edu

Abstract.—The nine western Atlantic species belonging to three genera, Plu-
marella, Acanthoprimnoa, and Candidella, are described and illustrated. Four
new species of Plumarella are described, as well as one new species of Acan-
thoprimnoa; the genus Acanthoprimnoa is also described as new, differentiated
from Plumarella by lacking tubercles on the undersurfaces of its sclerites. Two
western Pacific species are transferred to Acanthoprimnoa: A. serta and A.
cristata. Three varieties are recognized of the common Plumarella pourtalesii,
one previously described as a variety (P. p. robusta) and another proposed
herein (P. p. var. obtusa). A dichotomous key and table of comparisons is
provided for the species and forms of Plumarella, as are a table of comparisons
for the two Atlantic species of Acanthoprimnoa, and an indented key to the
eleven genera of western Atlantic Primnoidae. Specimens of these genera were
found to be extremely common at lower shelf and upper slope depths primarily
in the temperate western Atlantic; over 1500 specimens were examined in this

study, including types of all included species.

This is the fifth in a series of revisions
(Cairns 2001; Cairns & Bayer 2002, 2003,
2004) of the western Atlantic deep-water
octocorals, and the fourth dealing with the
Primnoidae, a family consisting of about
205 species and 32 genera worldwide, of
which approximately 33 species and 11
genera occur in the western Atlantic. Bay-
er’s revision of western Atlantic Calyptro-
Phora (2001) should also be considered as
the first unnumbered part of this series,
which also deals with primnoids. In order
to facilitate identification at the generic lev-
el within this family a key is provided be-
low for those 11 genera that occur in the
western Atlantic. Two genera occur twice
in the key since they have both dichoto-
mous and pinnate branching. In this part we
review the genera Plumarella and Candi-
della, as well as describe a new genus,
Acanthoprimnoa, separated from Plumar-
ella on the basis of its lacking tubercles on

the undersurfaces of its sclerites. Specimens
of Plumarella and Acanthoprimnoa are ex-
tremely common at shelf and upper slope
depths (137—1160 m) in the western Atlan-
tic, occurring there opportunistically as
weeds do on dry land. Ironically, species of
these two genera were previously known
from only 11 stations and as many speci-
mens from the western Atlantic; this report
lists approximately 1425 specimens from
145 localities. The third genus, Candidella,
is known from deeper water (514—2139 m),
and occurs farther north (New England Sea-
mounts) as well as in the eastern Atlantic.

Indented Key to the 11 Western Atlantic
Genera of Primnoidae (fraction indicates
number of western Atlantic species/total
number of species; genera in bold face
treated in this part)

I. Colonies unbranched or extremely
sparsely branched: Primnoella (4/14)

448 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

II. Branching in the form of a bottle-
brush: Thouarella (1+/25)
Ill. Colonies pinnately branched
A. Polyps arranged in whorls: Callo-
gorgia (3/28)
B. Polyps arranged biserially and al-
ternately
1. Tubercles not present on un-
dersurfaces of sclerites: Acan-

thoprimnoa (2/4)

2. Tubercles present on undersur-
faces of sclerites

a. Undersurface of opercular
scales with a prominent
keel: Amphilaphis (1/6) _

b. Undersurface of opercular
scales without a longitu-
dinal keel: Plumarella (6/
20)

IV. Colonies dichotomously branched
A. Polyps arranged in whorls
1. Polyps face outward from
branch; 4 marginal scales:

Candidella (1/4)

2. Polyps face up or down; 2
marginal scales

a. Polyps encased by only
two pairs of large abaxial
body wall scales
i. Members of two pairs

of body wall scales
inseparably fused to
form a complete ring
surrounding polyp;
polyps face up or
down: Calyptrophora
(4/13)

ii. Body wall scales not
fused; polyps face
downward: Paraca-
lyptrophora (3/6)

b. Polyps encased by 3 or 4
pairs of large abaxial
body wall scales: Narella
(7/25)

B. Polyps arranged biserially and al-
ternately

1. Tubercles not present on un-

dersurfaces of sclerites: Acan-

thoprimnoa (2/4)

2. Tubercles present on under-
surfaces of sclerites: Plumar-

ella (6/20)

C. Polyps irregularly arranged: Prim-
noa (1/3)

Material and Methods

This study was based on the examination
of approximately 1505 specimens, collected
at 161 deep-water stations by 18 research
vessels (Appendix: Station data). Except for
those reported from the Bibb and Atlantis,
which were borrowed from the MCZ, the
specimens are deposited at the National
Museum of Natural History (USNM). Syn-
onymies for all species are purported to be
complete for all previously published re-
cords. Unprefaced SEM stub numbers per-
tain to the series made by Bayer; those pref-
aced with C, to the series made by Cairns.

The following abbreviations are used:
Alb—U.S.EC.S. Albatross; Atl—R/V_ At-
lantis; BM—British Museum (now The
Natural History Museum, London); C/—R/
V Colombus Iselin; G—R/V Gerda; Gos—
R/V Gosnold; H:W—height to maximum
width of an opercular or marginal scale;
JS—Johnson-Smithsonian Deep-Sea Expe-
dition (Caroline); MCZ—Museum of Com-
parative Zoology, Harvard, Cambridge;
O—M/V and R/V Oregon and Oregon IT;
P—R/V Pillsbury; SB—R/V_ Silver Bay;
USNM—United States National Museum
(now the National Museum of Natural His-
tory, Smithsonian, Washington, D.C.).

Subclass Octocorallia
Order Gorgonacea
Suborder Calcaxonia
Family Primnoidae Gray, 1858
Genus Plumarella Gray, 1870

Cricogorgia Milne Edwards, 1857:6, pl.
B2, fig. 6 (nom. nud.).—Gray, 1870:36—
Bik

Plumarella Gray, 1870:36.—Studer, 1887:
51.—Wright & Studer, 1889:73-74,
281.—Versluys, 1906:13—14.—Kinoshi-
ta, 1908:6—8.—Kiikenthal, 1915:144;
1919:340—342; 1924:255.—Deichmann,
1936:155-156.—Bayer, 1956:F220.—
Fabricius & Alderslade, 2001:244—245.

VOLUME 117, NUMBER 4

Type species.—Gorgonia penna La-
marck, 1815, by subsequent designation
(Kiikenthal 1915:144).

Diagnosis.—Primnoidae with a well-de-
fined operculum; polyps usually inclined
apically, each polyp completely surrounded
by 8 rows of body wall scales; polyps ar-
ranged biserially or irregularly, but never in
whorls; 8 marginal scales, often pointed or
spinose; undersurfaces of all sclerites tu-
berculate, opercular scales not keeled; col-
onies uniplanar, usually pinnately (plu-
mose) branched but sometimes dichoto-
mous.

Distribution.—Western Pacific; Patagonia;
western Atlantic; 10—1914 m.

Remarks.—The only revision of the genus
Plumarella was that of Kiikenthal (1919),
reiterated in 1924: (Kiikenthal, 1924), which
included the description and synonymy of
all 17 species as well as a key to their iden-
tification. He used the following characters
to distinguish species, as emphasized in his
key: shape of distal edge of marginal scales,
presence of a longitudinal keel on the body
wall scales, number of scales in the ab- and
adaxial body wall rows, polyp size, and tex-
ture of surface of body wall scales. These
characters have also been used in this re-
view (Table 1), along with the additional
characters such as branching mode, termi-
nal branchlet length and flexibility, number
of polyps/em branch length, shape of the
operculars, presence of tubercles on the un-
dersides of the sclerites, ornamentation on
the edges of the opercular scales, and
coarseness of coenenchymal granulation,
the last three characters being used to dis-
tinguish a closely related new genus once
confused with Plumarella.

Key to the Species and Forms of the Six
Species of Plumarella known from the
Western Atlantic

1. Distal edges of marginal scales straight,
gently rounded or only slightly angular

1’. Distal edges of marginal scales promi-
nently spined

449

2. Branching alternate pinnate; colonies
often large (up to 33 cm) ............ 3)
2’. Branching dichotomous; colonies fairly
small (less than 11 cm)
3. Body wall scales smooth
3’. Body wall scales granular
se An ems Mee a Bn err 5 (P. pourtalesii)
4. Closely-pinnate branching; opercular
scales elongate and granular; 10—12
polyps/cm P. pellucida
4’. Loosely-pinnate branching; opercular
scales shorter and smooth; 14—21 pol-
yps/cm
5. Distal edges of marginal scales of some
polyps in a colony slightly angled (but
not spinose) ....P. pourtalesii var. obtusa
5’. Distal edges of marginal scales straight
to slightly rounded
6. 11-13 polyps/cm; distance between pol-
yps on one side of branch 1.0—1.2 mm
aati tidtgce atengs. coeceeneea P. pourtalesti var. typical
6’. 14-16 polyps/cm; distance between pol-
yps 0.5—0.8 mm
Ne iv avec @ P. pourtalesii var. robusta
7. Hach marginal scale with a prominent
spine
7’. Only 4—7 marginal scales with an elon-
gate needle-shaped spine, those scales
corresponding to operculars with at
least one uncovered edge ..... P. aculeata

... P. dichotoma

RE CEE te ke re es P. laxiramosa

PE th Ue oho a ent a ene aed P. aurea

Plumarella pourtalesii (Verrill, 1883)
Figs. 1[A-B, 2A—C, 3A—C

Primnoa Pourtalesii Verrill, 1883:28—29,
pl. 2, figs. 2, 2a—e (S. Carolina).—Not
Hargitt & Rogers, 1901:281, fig. D (prob-
ably A. goesi).

Plumarella pourtalesi.—Wright & Studer,
1889:73, 74, 280 (new comb.).—Ver-
sluys, 1906:15—16 (comments only).

Plumarella pourtalesii.—Versluys, 1906:
342 (key), 345-346 (German translation
of Verrill); Ktikenthal, 1919:345—346;
1924:257-258 (same as 1919).—Deich-
mann, 1936:156, pl. 25, figs. 17-18, pl.
26, figs. 10, 10a (two new records: Bibb
22, MCZ 4822 and Bibb 135, MCZ
4821).— Bayer, 1954a:281 (listed); 1956:
F220, fig. 159-7; 1957:388 (two records,
forma obtusa).—Bayer, Grasshoff & Ver-
seveldt, 1983: fig. 53.—Bayer, 1973: fig.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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VOLUME 117, NUMBER 4

Table 1.—Continued.

A. goesi
(Aurivillius, 1931)

P. aurea
(Deichmann, 1936)

A. pectinata, n. sp.

P. aculeata, n. sp:

P. dichotoma, n. sp.

1.6—2.2
Ridged; granular; granu-

lar

1.5—2.0 1.6—2.2 1.8-3.9

1.7-2.3

H: W of operculars

Ridged; spinose; spinose

Smooth; smooth; smooth All low granular (body

Granular; smooth; granu-

lar

Ornamentation of opercu-
lars; bw; coenenchymal

scales

wall sclerites may also be

smooth)

Yes

Tubercles present on un-
dersides of sclerites

Distribution

Off northeastern Yucatan

Bahamas (northern Straits Straits of Florida to Yu-

off S. Carolina to Florida; off S. Carolina to Cuba;

494-1065 m

Peninsula (164476 m);
Lesser Antilles, Straits of

Florida (614-686 m)

catan, Bahamas, Puerto
Rico, Virgin Islands;

137-595 m

of Florida); 400-900 m

310-878 m

451

16.—Bayer & Cairns (Verrill), 2004: pl.
26, fig. 1.

Plumarella pourtalesii var. robusta Deich-
mann, 1936:156—157, pl. 25, figs. 14—16,
pl. 26, fig. 9.

Material examined.—Typical form: Alb-
2416, 10 branches, USNM 10531; Alb-
2662, 4 branches, USNM 14609; Alb-2663,
10 branches, USNM 14479; Alb-2667, 4
branches, USNM 49425; Alb-2668, 11 col-
onies, USNM 1019275; Alb-2669, over 50
colonies, USNM 14475; Atl-266-2, 6 colo-
nies, USNM 1019276; Atl-266-7, 3 colo-
nies, USNM 1021650; Ati-266-41, 7 colo-
nies, USNM 1019277; Bibb 22, 2 branches,
MCZ 4822 (reported by Deichmann 1936);
Bibb 135, 2 branches, MCZ 4821 (reported
by Deichmann 1936); Cape Hatteras SA6-
5, 13 colonies, USNM 79782 (topotypic);
CI-123, 9 colonies, USNM 1019278; CI-
246, 26 branches, USNM 59491; Clelia 78,
1 colony, USNM 93910; Clelia 79A, 1 col-
ony, USNM 93909; Combat 174, 1 colony,
USNM 50799; Eastward 26017, 14 branch-
es, USNM 59490; Eastward 26022, 2
branches, USNM 1019279; Eastward
26023, 10 branches (dry), USNM 1019280;
Eastward 26028, 2 branches, USNM
1019281; Eastward 26052, 2 branches,
USNM 1021652; G-170, 1 colony, USNM
1019305; G-177, 12 colonies, USNM
1019282; G-235, 1 branch, USNM
1019283; G-386, 3 colonies and SEM stub
250, USNM 53010; G-598, 1 colony,
USNM 1019284; G-672, 1 colony, USNM
59498; G-785, 3 colonies, USNM 52994;
Gos-2344, 10 branches, USNM 58448;
Gos-2385, 10 colonies, USNM 56895; Gos-
2387, 1 colony, USNM 57308; Gos-2413,
8 colonies, USNM 1019285; Gos-2414, 17
colonies, USNM 1019286; Gos-2461, 2
branches, USNM 1019287; O-1343, 3
branches, USNM 50183; O-1349, 3 branch-
es, USNM 50443 (Bayer 1958); O-11703,
2 colonies, USNM 59501; O-11717, 1
branch, USNM 59507; O-11725, 5 colo-
nies, USNM 1019288; P-105, 2 branches,
USNM 52999; SB-453, 2 branches, USNM

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

2416: C, P. aurea,

2662; B, P. pourtalesii var. robusta, Alb

Alb-

Alb

1 cm.

2354. All scale bars

Acanthoprimnoa pectinata,

452

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Fig. 1.
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Vee

Fig. 3. A-—C, Plumarella pourtalesii typical (A-B, G-386; C, syntype, MCZ 5749): A, upper and undersur-
faces of 5 opercular scales; B, upper surfaces of 3 body wall scales; C, upper and undersurfaces of 4 coenen-
chymal scales. D—E Plumarella pellucida (first opercular scale is from G-859, all other scales from the holotype):
D, upper and undersurfaces of 5 opercular scales; E, upper and undersurfaces of 4 body wall scales; K upper
and undersurfaces of 4 coenenchymal scales. All scale bars = 0.10 mm.

VOLUME 117, NUMBER 4

51260; syntypes (MCZ, USNM, see be-
low).

Forma robusta: Alb-2416, over 25 colo-
nies and | unnumbered SEM stub, USNM
49430, 49433: Alb-2666, 2 colonies,
USNM 49422; Alb-2668, 10 branches,
USNM 14474; Alb-2669, 8 branches,
USNM 1019289; Anton Dohrn 6392, 3 col-
onies, USNM 1019290; Atl-266-4, 1 colo-
ny, 1 branch, USNM 1019291; Azi-266-40,
15 branches, USNM 1019293; Atl-266-41,
1 branch, USNM 1019292; Cape Hatteras
SA6-1, 5 colonies, USNM 79783; CI-140,
1 branch, USNM 59497; Combat 368, 14
colonies and SEM stub 253, USNM 50800;
Eastward 26023, 1 dry branch, 4 alcohol
branches, USNM 1019294; G-170, 1
branch, USNM 1019295; G-177, over 20
colonies and 1 unnumbered SEM stub,
USNM 52995; G-247, 1 colony, USNM
53012; G-261, 3 colonies, USNM 1019296;
G-598, 7 colonies, USNM 52997; G-835, 3
branches, USNM 52998; Gos-2413, 5
branches, USNM 1019297; Gos-2461, 8
colonies, USNM 1019298; O-11726, 1
branch, USNM 73756; P-197, 1 colony,
USNM 53013; holotype (see below).

Forma obtusa: Alb-2416, 1 branch,
USNM 1019299; Alb-2668, 8 branches,
USNM 1019300; Alvin 77-761, 2 colonies,
USNM 1019301; Alvin 1335, 2 colonies,
USNM 73741; Atl-3780, 10 dry branches,
MCZ 54321; Cape Hatteras SA6, 8 branch-
es, USNM 79781; CI-140, 3 branches,
USNM 59495; CI-246, 9 branches, USNM
1019302; Eastward 26022, 1 branch,
USNM 76986; Eastward 26023, 1 branch,
USNM 1019303; G-56, 1 colony and 1 un-
numbered SEM stub, USNM 53005; G-
169, 7 colonies, USNM 53008; G-235, 8
colonies and SEM stub 260, USNM 53007;
G-241, 9 colonies, USNM 53004; G-246, 2
branches, USNM 53003; G-261, 10 colo-
nies, USNM 53009; G-386, 3 branches,
USNM 1019304; G-391, 6 colonies and
SEM 252, USNM 1019305; G-598, 1 col-
ony, USNM 1019306; G-664, 3 branches,
USNM 53001; G-679, 4 colonies, USNM
53002; G-1012, 1 colony, USNM 53011;

455

G-1314, 9 colonies and SEM stub 251,
USNM = 53006; — Gilliss, 2a OUNE
79°24'36"W, 603 m, 25 May 1973, 1
branch, USNM 79515; Gos-2387, 5 colo-
nies, USNM 57306; O-1328, 1 colony,
USNM 50528; P-209, 4 branches, USNM
53000.

Types and type localities.—TYwo colonies
and several fragments of the typical form
were mentioned by Verrill (1883) collected
from Blake 318, all of which must be con-
sidered as syntypes. Deichmann (1936) at-
tributed catalog number MCZ 4821 to the
syntype series, but this number was preoc-
cupied by Deichmann by specimens col-
lected from “off Florida’’, thus the syntype
series was later re-cataloged as MCZ
42887, and consists of three small branches.
Four fragments and SEM stubs 249 and
C1084 of one of these syntypes are also de-
posited at the USNM (5749). All syntypes
are preserved in alcohol. Type Locality:
Blake 318: 31°48'50"N, 77°51'50"W (Blake
Plateau off South Carolina), 616 m.

The holotype of forma robusta, two
small branches in alcohol, the largest 9 cm
high, is deposited at the MCZ (4823). It
also bears Verrill’s personal number of
8032. It is also represented by an unnum-
bered SEM stub at the USNM. Type Lo-
cality: The type locality was stated to be
“18 fms. off Alligator Reef, Florida”’
(Deichmann, 1936:157), but Deichmann
correctly queried the extremely shallow
depth of the collection. A newer label with
this type indicates that it was collected at
Bibb station 192 (24°48'05"N, 80°34'45’W:
off Alligator Reef), at 216 m, this depth be-
ing more consistent with the known bathy-
metric range.

Diagnosis.—Distal edge of marginal
scales rounded (scalloped) or straight;
branching close pinnate, branchlets of mod-
erate length and stiff; body wall scales
granular, surface of operculars ridged; 11—
13 polyps/cm (14—16 for forma robusta and
obtusa).

Description.—Colonies are flabellate and
pinnately branched (plumose), consisting of

456

a main branch, which gives rise to 1—4 pri-
mary branches (depending on the size of
the colony), each main and primary branch
giving rise to numerous stiff branchlets up
to 55 mm in length, in an alternating se-
quence (alternate pinnate). Although flabel-
late, colonies may be uniplanar or slightly
convex, such that the polyps are directed
toward the convex side. A branchlet occurs
on the main branch within 5 mm of the base
of the colony; additional branchlets occur
at regular intervals of 1.8—3.2 mm on the
main and primary branches, producing a
characteristic zig-zag pattern for the larger
branches, the branchlets being straight and
parallel to one another. The main stem is
usually anchored by a dense, white, calcar-
eous holdfast, which is often attached to the
dead corallum of a deep-sea scleractinian.
The axis is yellow-brown or gold and lon-
gitudinally striate; overall the colony is
white in alcohol. The largest colony (Com-
bat 174) is 33 cm in height, 13 cm in width,
and 2.7 mm in basal diameter, but most col-
onies examined were considerably smaller.

Polyps are arranged on all branches (1.e.,
main, primary, and branchlets) biserially in
the plane of the flabellum in an alternating
fashion, but angled upward such as to pro-
duce a 30°—-45° angle with the branch, as
well as being slightly curved toward the an-
terior (convex) side of the flabellum. Two
to six polyps occur on the internodes of the
main and primary branches. Polyps are
rarely more than | mm in height, slightly
wider at the apex (0.5—0.6 mm) than the
base, and fairly well spaced: polyps on the
same side of a branchlet are separated by
1.0—1.2 mm. In general, 11—13 polyps oc-
cur/em of branchlet length, this number
sometimes lower in small colonies.

Each polyp is encased by 8 opercular and
8 rows of body wall scales. The abaxial
rows of body wall scales consist of 5 or 6
scales, the adaxial rows usually one less (4—
5 scales), the latter rows always shorter due
to having less scales and being located on
the shorter, concave, side of the upturned
polyp. The marginal scales are up to 0.37

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

mm in width, bearing sparse, low granules
(10 wm in diameter) on their exterior sur-
face and complex tubercles on undersurface
(up to 13 pm in diameter). Body wall scales
proximal to the marginals are progressively
smaller. Body wall scales overlap one an-
other within each row as well as overlap-
ping the edges of the scales in both adjacent
rows. Body wall scales, including margin-
als, are somewhat crescent-shaped, wider
than tall (H:W = 0.6-0.7), and slightly
curved to accommodate the curvature of the
polyp, the distalmost scales in each row
(the marginals) having a straight or slightly
rounded, finely serrate distal margin, pro-
jecting only about 10—20 pm beyond the
articulation with the base of the operculars,
and, if rounded, producing a slightly scal-
loped calyx margin. The 8 opercular scales
are similarly-shaped (H:W = 1.5—1.9),
symmetrical, isosceles triangles, having an
apical angle of 41°-51° but with rounded
tips. The abaxial operculars are up to 0.42
mm in height, the adaxials about 0.30 mm
in height. The upper surface of the each
opercular is covered with small spines on
the lower half and 3 or 4 longitudinal ridges
apically. The lower two-thirds of the un-
dersurface is covered with complex tuber-
cles of the same size as those of the body
wall scales; the distal region is smooth, not
keeled. The lateral edges of each opercular
is finely serrate like the distal edges of the
body wall scales. The 8 opercular scales
fold together forming a moderately tall,
conical operculum.

Coenenchymal scales occur in one layer,
are flat, and elliptical to elongate in shape,
the largest scales being 0.40 mm in length.
Like the other scales, they are sparsely
granular above, tuberculate below, and bear
finely serrate edges. Tentacular sclerites
were not noted.

Comparisons.—See Table 1.

Distribution.—Typical form: Blake Pla-
teau from North Carolina (34°45'N)
through Straits of Florida (insular side) and
north central part of Cuba; 196-882 m. For-
ma robusta: Blake Plateau from off N. Car-

VOLUME 117, NUMBER 4

olina (34°15'N) through Straits of Florida
to Florida Keys; 183-877 m. Forma obtusa:
Blake Plateau from off Georgia (31°26’N)
through Straits of Florida and Northwest
Providence Channel; 183—743 m.

Remarks.—Deichmann (1936) described
a variety of P. pourtalesii called robusta,
which differs from the typical form in hay-
ing a stouter corallum, thicker sclerites, a
flatter operculum, and longitudinal ridges
on the opercular scales. This form is well
represented in our collection (listed sepa-
rately above), and differs from the typical
form (Table 1) in having a stouter, stiffer
colony, and slightly larger and thicker pol-
yps (1.0—1.2 mm tall, 0.65 mm in diameter)
that are more closely spaced on the branch
(distance between adjacent polyps on one
side of a branch 0.5—0.8 mm), such that
even though the polyps are slightly larger,
there are more/cm, i.e., 14—16, the latter
number achieved when polyps also bud
from the anterior face of the branch. As
Deichmann stated, in general, the opercu-
lum is flatter, even concave in some highly
contracted polyps, but both forms have lon-
gitudinal ridges on the opercular scales.
These several differences are fairty consis-
tent but do not include any characters rou-
tinely used to differentiate species of Plu-
marella (see Ktikenthal 1919). Further-
more, the distribution and bathymetric
range of both taxa are virtually the same, 7
of the 22 records of robusta being from
common stations. We thus concur with
Deichmann in considering this form as just
an environmental variation of the typical
form.

Another common form of P. pourtalesii,
represented in our collection by 27 lots, is
otherwise similar to forma robusta but dif-
fers in that the marginal scales of at least
some polyps of every colony have an an-
gled or evei pointed distal edge (Fig. 2D).
The angle o the distal edge is often a sharp
right angle extending about 0.1 mm, or less
commonly may be a discrete spine up to
0.25 mm in length. There is great variation
in the expression of this character. In some

457

colonies all marginal scales of all polyps
will have a short angled distal edge, where-
as in other colonies only those polyps to-
ward the distal branch tips will be so mod-
ified, the marginal scales of the remaining
polyps having a typically straight or round-
ed edge. Furthermore, individual polyps
may have one or all eight marginal scales
with an angled edge. The long-spined mar-
ginals are infrequent and when present only
occur one to a polyp. All three forms some-
times occur at the same station, and their
bathymetric and geographic ranges are
quite similar. Because of the great variation
of this single character that separates this
taxon from forma robusta and the typical
form, it is considered to be an environmen-
tal or genetic variation without taxonomic
validity, but, in order to easily refer to this
variation, the form name obtusa is applied
to it, an allusion to the often obtuse angle
formed by the distal edges of its marginal
spines.

Plumarella pellucida, n. sp.
Figs. 3D—-E 4A-—B, 10A

Material examined/types and type locali-
ty.—Holot pe: G-647, 1 colony and SEM
stubs C1079—1080, 1085-1086, USNM
52992. Paratypes: Alb-III 9-19, 1 colony,
USNM 50573; Cape Hatteras SA6-5, 2 col-
onies, USNM 1019404; CI-266, 3 colonies,
USNM 59506; G-647, 2 colonies, 3 branch-
es, USNM 1019403; G-808, | colony,
USNM 59499; G-859, 3 branches and 1 un-
numbered SEM stub, USNM 52993; O-
11705, 5 branches, USNM 1019405. Type
Locality: 26°16'N, 79°43’W (Straits of Flor-
ida off Fort Lauderdale), 520—549 m.

Diagnosis.—Distal edge of marginal
scales rounded or straight; branching close-
ly pinnate, branchlets of moderate length
and stiff; opercular scales granular but not
ridged, other scales faintly granular, ap-
pearing almost smooth and translucent; ab-
axial and outer-lateral opercular scales quite
long and curved; 10—12 polyps/cm.

Description.—Colonies are flabellate and

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

458

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VOLUME 117, NUMBER 4

closely pinnately branched, as in P. pour-
talesii, the distance between successive
branchlets 2.5—5.0 mm, and the entire fla-
bellum is usually convexly shaped, the pol-
yps curving toward the convex (anterior)
face. Branchlets are up to 60 mm long and
are fairly stiff. The main stem is anchored
by a dense calcareous holdfast. The axis is
yellow and faintly longitudinally striate; in
alcohol the colony is light brown. The larg-
est colony (holotype) is 17 cm in height, 16
cm in width, and 2.2 mm in basal stem di-
ameter.

Polyps are quite regularly arranged in a
biserial, alternating fashion on all branch
edges, and are inclined distally. Polyps on
the same side of a branchlet are well sep-
arated by 1.0—1.2 mm, resulting in 10—12
polyps/cm. Polyps are 1.1—1.2 mm in
height and slightly flared distally (0.65 mm
in diameter).

Each polyp is covered with 8 opercular
and 8 rows of body wall scales, the abaxial
body wall scales numbering about 6, the ad-
axial usually consisting of 5. The body wall
scales, including the marginals, are similar
in size and shape to those of P. pourtalesii;
however, their exterior granulation is much
reduced and the scales appear to be thinner,
producing a smooth, almost translucent as-
pect. Like P. pourtalesii, the distal edges of
the marginals are straight to slightly round-
ed, never spinose or angled. The 8 oper-
cular scales are isosceles triangles (H:W =
1.7—2.4), having an apical angle of 20°-35°,
ranging from sharply pointed to slightly
rounded. The abaxial and outer-lateral oper-
culars measure up to 0.54 mm in height, the
adaxial and inner-lateral operculars only
0.37 mm in height. The distal third of the
abaxial and outer-lateral operculars are at-
tenuate, with slightly serrate edges; these
scales are curved downward to follow the
curvature of the polyp and almost reach the
opposite side of the polyp, considerably
overlapping the shorter adaxial and inner-
lateral opercular scales. The opercular
scales bear low sparse granules, lack distal
ridges, and are tuberculate on the undersur-

459

face with no trace of a keel. In general, the
opercular scales fold together in a conical
operculum in a manner similar to that
shown in Fig. 9f, the abaxial opercular hav-
ing two exposed edges, the adaxial having
none.

Coenenchymal scales occur in one im-
bricating layer, are flat, and elliptical to ir-
regular in shape, the largest being about
0.41 mm in width. Their granulation is re-
duced similar to that on the opercular
scales. Tentacular scales were not noted. All
sclerite types, body wall, opercular and coe-
nenchymal bear complex tubercles on their
undersurfaces, the largest of which mea-
sures about 13 pm in diameter.

Etymology.—The species name pellucida
(Latin: pellucidus, transparent, translucent,
clear) refers to the translucent nature of the
body wall and coenenchymal scales when
viewed in liquid, probably due to their thin-
ness and sparse granulation, which allows
a view of the outline of the branch axis and
of 8 faint white longitudinal lines in the
polyps corresponding to the mesenteries.

Comparisons.—Plumarella pellucida be-
longs to a closely related species complex
characterized by having large, pinnately
branched colonies and smooth to straight-
edged (not spinose or pointed) marginal
scales, this complex consisting of: P. pour-
talesii, P. laxiramosa, and P. pellucida. It
is probably most closely related to P. pour-
talesii, but is distinguished by having non-
ridged opercular scales, the adaxial and out-
er-laterals of which are quite long and
curved; and smooth, almost translucent
body wall and coenenchymal scales. It dif-
fers from P. laxiramosa in having closer
pinnate branching; longer, more attenuate
opercular scales that are granular; and fewer
polyps/cm. (Table 1). All three species oc-
cur in roughly the same geographic and
bathymetric range and often occur at the
same stations.

Distribution.—North Carolina through
Straits of Florida, Bahamas; 549-1160 m.

460

Plumarella laxiramosa, 0. sp.
Figs. 4C—E, 5A—E, 10B

Material examined/types and type local-
ity.—Holotype: Cape Hatteras SA6-1, 1
colony and SEM stubs C1081-1083,
USNM 1019406. Paratypes: Alb-2416, 7
colonies, over 50 branches and 1 unnum-
bered SEM stub, USNM 50594 and 79458;
Atl-266-41, 1 branch, USNM_ 1019407;
Cape Hatteras SA6-1, 19 branches, USNM
79778; Cape Hatteras SA6-5, 8 colonies,
USNM 79779: Combat 368, 1 branch,
USNM 1019408; Gos-2387, 1 colony,
USNM 1019409. Type Locality:
31°17'18"N, 79°00'39"W (Charleston Bump
region, South Carolina), 572—575 m.

Diagnosis.—Distal edge of marginal
scales rounded or straight; branching loose-
ly pinnate, branchlets long and flabby; body
wall, opercular, and coenenchymal scales
smooth; 14—21 polyps/cm, polyps often oc-
curring on anterior face.

Description.—Colonies are flabellate and
pinnately branched, as in P. pourtalesii, but
differ from that species in having more
widely-spaced (loose pinnate) and thus few-
er branchlets, each branchlet separated form
their adjacent by 6-11 mm. Also, unlike P.
pourtalesii, this species has flat, not curved,
colonies, and its branchlets are longer (up
to 80 mm) and less stiff, altogether produc-
ing a limp or languid colony tension. The
main stem is anchored by a dense, white
calcareous holdfast, which often attaches to
the corallum of a dead scleractinian or sty-
lasterid coral with an encrustation up to 1
cm in diameter. The axis is golden-yellow
and faintly longitudinally striate; overall the
colony is light brown in alcohol. The ho-
lotype is 18 cm in height, 17 cm in width,
and has a main stem diameter of 2.7 mm,
although the stem is broken from the sub-
strate. The largest colony, which has an in-
tact base (A/b-2416), is 24 cm in height.

Polyps are closely arranged on all
branches biserially in the plane of the fla-
bellum and in an alternating fashion; how-
ever, most larger colonies often have a third

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

row of polyps on the anterior side, which
produces a very crowded arrangement of
polyps that may number up to 21/cm. Up
to 20 polyps may occur on the rather
lengthy internodes of the main and primary
branches. Polyps are 1.1—1.2 mm in height
and are slightly flared distally (0.65 mm).

Each polyp is covered with 8 opercular
and 8 orderly rows of body wall scales, the
abaxial body wall scales numbering 6 or 7/
row, the adaxial consisting of usually only
5. Body wall scales are roughly rectangular
and slightly curved to fit around a segment
of the polyp; the distalmost body wall
scales (the marginals) are 0.30—0.34 mm in
width and 0.20—0.25 mm in height, the
scales becoming progressively smaller to-
ward the branch. The distal edges of the
marginals are finely serrate and straight to
slightly rounded, never pointed or spinose.
The undersurfaces of all scales are tuber-
culate (tubercles 12—26 wm in diameter),
whereas the upper surfaces of the body wall
scales, as well as those of the opercular and
coenenchymal scales, are virtually smooth.
The 8 opercular scales are similarly-shaped
(H:W = 1.75—2.20), symmetrical, isosceles
triangles, having a blunt, rounded apical an-
gle of about 30°. The abaxial operculars are
up to 0.45 mm in height, the adaxials only
slightly less tall (e.g., 0.37 mm). As men-
tioned above, the surface of the operculars
is smooth and without ridges, whereas their
undersurface is tuberculate. When contract-
ed, the operculars form a closely fitted,
overlapping, low operculum.

Coenenchymal scales occur in one im-
bricating layer, are flat, and elliptical to ir
regular in shape, the largest scales being
about 0.25 mm in length. Tentacular scales
were not noted.

Etymology.—The species name laxira-
mosa (Latin: laxus, loose, slack, and ra-
mosus, branching) refers to the loose
branching mode of the colonies as well as
the limp tension of the branchlets.

Comparisons.—Among those western
Atlantic species of Plumarella having
smooth-edged marginal scales (Table 1), P.

VOLUME 117, NUMBER 4

Fig. 5. A—E, Plumarella laxiramosa, holotype: A, upper and undersurfaces of 4 opercular scales; B, tip of
undersurface of an opercular showing fine serration of edge; C, upper and undersurfaces of 3 body wall scales;
D, tubercles on underside of a body wall scale; E, upper and undersurfaces of 4 coenenchymal scales. F-I,
Plumarella dichotoma, holotype: FE upper and undersurfaces of 4 opercular scales; G, tubercles on underside of
an opercular scale; H, upper and undersurfaces of 5 body wall scales; I, upper and undersurfaces of 5 coenen-
chymal scales. Scale bars for A, C, E-K H-I = 0.10 mm; B = 25 pm; D, G = 10 pm.

462

laxiramosa is most easily distinguished by
its growth form (loose pinnate) and by hav-
ing a third, anterior row of branchlet pol-
yps, which leads to a high number of pol-
yps/cm. It is also distinguished by having
no surface granulation on any scales.
Distribution.—Off North Carolina (to
34°15'N) and South Carolina; 348—625 m.

Plumarella dichotoma, n. sp.
Figs. 5F—I, 6A—-C, 10C

Material examined/types and type local-
ity.—Holotype, Gos-2387, 1 colony and
SEM stubs C1087—1089, USNM 57307.
Paratypes: Alb-2666, 10 colonies, USNM
49423; Alb-2667, 9 colonies, USNM
49431; Alvin 77-762, 1 colony, USNM
1019427; Alvin 1335, 1 dry colony, USNM
73739; Anton Dohrn 65-32, 3 dry branches,
USNM 1019429; Cape Hatteras SA6-5, 5
colonies, USNM 79780; Eastward 26004,
2 colonies, USNM 1019430: Eastward
26023, 1 colony, USNM 1019431; G-169,
2 colonies, USNM 52990; G-170, 1 colony,
USNM 52996: Gos-2344, 3 colonies,
USNM 58447; Gos-2387, 10 colonies,
USNM_ 1019428; Gos-2413, 7 colonies,
USNM 1019432; Gos-2469, 1 colony,
USNM 59021. Type Locality: 31°14'48”"N,
78°59'W (off South Carolina), 530 m.

Diagnosis.—Distal edge of marginal
scales straight or rounded; branching di-
chotomous, sometimes lyrate, branchlets
relatively short and wiry; opercular and
body wall scales smooth; 8-10 polyps/cm
(polyps widely spaced), standing perpendic-
ular to branches on large-diameter branch-
lets.

Description.—Colonies are flabellate and
usually slightly curved, as in P. pourtalesii,
but consistently dichotomously branched.
The main stem is attached to a substrate by
a thin calcareous encrustation and rises only
12-16 mm before it bifurcates, producing
an axil angle of 65°—85°. Subsequent equal,
dichotomous branching occurs at intervals
of 5—12 mm, although some end branches
are up to 40 mm in length and are the result

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

of 7-11 previous branching nodes. Termi-
nal branchlets are wiry to limp in tension.
Higher order axil angles are slightly small-
er, 1.e., 40°—-45°, undamaged colonies usu-
ally being slightly broader than tall. In
some large colonies the two outermost
branches remain slightly larger than their
inner branchlets (unequal dichotomous
branching), as in the holotype, producing a
lyrate form. The largest colony (the holo-
type) is 11 cm in height, 13 cm in width,
and has a basal main stem diameter of 1.4
mm, although most colonies examined were
considerably smaller. The axis is golden
yellow and the colony appears white in al-
cohol.

As in all species of Plumarella, the pol-
yps are biserially arranged in alternating
fashion on the edges of all branches, angled
slightly toward the anterior side of the fla-
bellum, and standing perpendicular to
large-diameter branches, but inclined dis-
tally in smaller-diameter distal branches.
Polyps are widely spaced, adjacent polyps
on the same side of a branch separated by
as much as 2.0 mm. Polyps are fairly tall
and slender, up to 1.3 mm in height and
0.65 mm in apical diameter.

The polyps are protected by 8 opercular
and 8 rather disorganized rows of body wall
scales, both ab- and adaxial rows containing
5 or 6 scales. Body wall scales are smooth,
quickly decreasing in size from the margin-
al to the more proximal ones. The distal
edges of the marginals are straight to slight-
ly rounded and the scales themselves are
square to slightly rectangular. The opercular
scales are modified isosceles triangles (al-
most pentagonal), the two long sides of the
triangle being parallel for much of the
length, only the distal third having an apical
angle of 45°—50°, culminating in a blunt tip.
Abaxial opercular scales are up to 0.50 mm
in height, adaxial, 0.35 mm; the H:W rang-
es from 1.7—2.3. Operculars have a granular
upper surface and a tuberculate lower sur-
face, devoid of a keel. They infold to form
an operculum as illustrated in Fig 9f.

Coenenchymal scales are mildly granular

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464

above, tuberculate below, and irregularly
elliptical in shape, rarely over 0.25 mm in
greater diameter.

Etymology.—The species name dichoto-
ma (Greek: to be divided into two parts), is
an allusion to the dichotomous branching of
the colonies.

Comparisons.—Among the western At-
lantic species of Plumarella, P. dichotoma
is unique in having both dichotomous
branching and smooth-edged marginal
scales (Table 1). It is also distinctive in hav-
ing such widely spaced polyps that are of-
ten oriented perpendicular to the branches.

Distribution.—Off southeast coast of
United States from South Carolina to off
Dry Tortugas, Florida; 494—1065 m.

Plumarella aurea (Deichmann, 1936)
Figs. 1C, 6D-E, 7A—D

Thouarella aurea Deichmann, 1936:165—
166, pl. 25, figs. 12-13, pl. 26, fig. 11.—
Bayer, 1954a:281 (listed).

Plumarella pourtalesii.—Deichmann,
1936:156 (Gin part: 2 of 4 specimens from
Bibb 22, Bahia Honda).

Plumarella aurea.—Bayer, 1981:934, fig.
70 (new combination).—Bayer & Cairns
(Verrill), 2004: pl. 25, 6a, pl. 83, la.

Material examined.—Alb-2666, 4 colo-
nies, USNM 52984; Alb-2667, 5 colonies
and | unnumbered SEM stub, USNM
52985; Alb-2668, 4 branches, USNM
1019312; Alvin 77-762, 3 colonies, USNM
1019313; Azl-3780, 15 dry colonies, MCZ;
Atl-3782, 1 dry colony, MCZ 54327; Atl-
266-40, 5 colonies and SEM stub 390,
USNM 58443; Atl-266-41, 5 colonies,
USNM 1019314; Cape Hatteras SA 6-5, 2
colonies, USNM 1019315: Discoverer X, 1
colony and SEM stub 391, USNM 58446;
Eastward 26004, 1 colony, USNM
1019316; Eastward 26022, 2 branches,
USNM 1019317; Eastward 26023, 4 dry
branches and | in alcohol, USNM 1019318;
G-672, 1 colony, USNM 52974; G-679, 1
colony, USNM 1019319; G-936, 1 branch,
USNM 1019320; Gos-2385, 4 colonies and

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF V ASHINGTON

SEM stub C1092, USNM 56892; Gos-
2414, 6 colonies, USNM 1019321; O-
11716, 1 colony, USNM 59500; P-105, 3
colonies, USNM 52976; SB-453, 1 colony,
USNM 51265; syntypes (see below); spec-
imens misidentified as P. pourtalesii by
Deichmann (1936) from the type locality,
Bibb 22 (MCZ 59442).

Types and type locality.—Five small
branches (syntypes) preserved in alcohol
are deposited at the MCZ (4801), which
also bear Verrill’s number 8042. An un-
numbered SEM stub of one of these
branches is also deposited at the USNM.
Type Locality: 24°14’20"N, 80°59’40"W
(off Bahia Honda, Straits of Florida off
Florida Keys), 310 fathoms (=567 m). AI-
though not stated in the original description,
a label with the type specimens indicates
they were collected at Bibb 22 (dredge 12),
made on 4 May 1868.

Diagnosis.—Distal edge of most margin-
al scales prominently spinose; branching di-
chotomous; opercular, body wall, and coe-
nenchymal scales smooth; polyps crowded,
sometimes on anterior face, 14—22 polyps/
cm.

Description.—Colonies are flabellate and
dichotomously branched. The main stem is
attached to the substrate by a thin calcare-
ous expansion and rises only 5—8 mm be-
fore it bifurcates, producing an axil angle
of about 55°; subsequent axial angles are
40°—45°. Branching is usually equal and di-
chotomous, occurring at intervals of 5—10
mm, but some terminal branches are up to
10 cm in length. Branches and colonies are
quite flexible in tension, almost limp. The
largest colony examined (Gos-2385) is 13
cm in height, 12 cm in width, and has a
main stem diameter of 1.5 mm. The axis is
golden-yellow and the colony appears white
in alcohol.

Polyps are crowded, occurring biserially
in alternating or opposite fashion on the
branchlets and often with occasional polyps
on the anterior side, resulting in 14—22 pol-
yps/cm. Polyps are oriented perpendicular
to the branches or tilted only slightly ante-

VOLUME 117, NUMBER 4 465

Fig. 7. A—D, Plumarella aurea, Gos-2385: A, upper and undersurfaces of 3 opercular scales; B, upper and
undersurfaces of 2 marginal scales; C, upper and undersurfaces of 4 body wall scales; D, upper and undersurfaces
of 4 coenenchymal scales. E-I, Plumarella aculeata, paratype from G-252: E, upper and undersurfaces of 2
opercular scales; EK upper and undersurfiaces of 2 marginal scales; G, spination on marginal spine; H, upper and
undersurfaces of 5 body wall scales; L. upper and undersurfaces of 5 coenenchymal scales. Scale bars for A—F

H-I = 0:10 mm; G = 25 pm.

466

riorly. They are usually squat, cylindrical,
and robust, 0.8—1.2 mm in height (depend-
ing on contraction) but always 0.75—0.80
mm in apical diameter.

Each polyp is protected by 8 opercular
and 8 well-defined rows of body wall
scales, the abaxial rows having 6 or 7
scales, the adaxial, 5 or 6. The distal edges
of the marginal body wall scales are usually
strongly spinose, the 8 tooth-like spines
forming a small crown encircling the oper-
culum and often rising above it. Occasion-
ally 1 or 2 of the marginals of a polyp lack
spines or have reduced spines, but most
polyps have 8 prominent, equal-sized
spines. The marginal spines are sharp (api-
cal angle 20°—25°), often constituting half
the height of the marginal scale, a large
spine being up to 0.25 mm in length and
0.08 mm in basal diameter, contributing to
a H:W for this kind of scale of up to 1.3—
1.5, the low value due to the wide base of
the marginal scales. The marginal spines
are circular in cross section and have finely
serrate edges where they join the lower
rectangular section of the scale (Fig. 7B).
Opercular scales are fairly flat (not curved)
and isosceles triangular in shape, the distal
point being somewhat rounded, forming an
angle of 33°—45°. Abaxial opercular scales
are up to 0.55 mm in height, adaxial only
0.30 mm; the H:W ranges from 1.5—2.0.
The upper surfaces of the body wall and
opercular scales are smooth, the undersur-
faces covered with complex tubercles that
are up to 15—16 pm in diameter.

Coenenchymal scales are also smooth
above, tuberculate below, and irregularly
elliptical, elongate, or circular in shape; and
up to 0.40 mm in greater diameter.

Comparisons.—Among the western At-
lantic Plumarella having spinose marginal
scales, P. aurea is most similar to P. acu-
leata (see that description and Table 1).

Distribution.—Blake Plateau from off
South Carolina (32°10'N) through Straits of
Florida to off Bahia Honda; Northwest
Providence Channel, Bahamas; 310—878 m.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Plumarella aculeata, n. sp.
Figs. 7E-I, 8A—B, 9a—g, 10E

Material examined/types and type local-
ity.—Holotype: G-707, USNM 52980, 1 di-
chotomous colony and SEM 248. Para-
types: Cape Florida X, 11 pinnate colonies,
USNM 1019533; Eastward 26535, 15 pin-
nate branches, USNM 1019534; Eastward
26547, 2 dichotomous colonies, USNM
1019535; G-241, 1 pinnate branch, USNM
1019536; G-252, 2 pinnate branches and
SEM stubs 255 and C1090, USNM 52979;
G-633, 2 pinnate colonies, USNM 52983;
G-692, 8 pinnate colonies, USNM 52986;
G-695, 1 dichotomous colony, USNM
52978; G-707, 5 dichotomous and 1 lyrate
colonies, USNM 52981-52982; G-1125, 3
pinnate colonies, USNM 52977; G-1312, 1
lyrate colony, USNM 52975; SB-440, 4 dry
pinnate branches, USNM 51292. Type Lo-
cality: 26°27'N, 78°40'W (Northwest Prov-
idence Channel, Bahamas), 514—586 m.

Diagnosis.—Distal edges of 4—7 margin-
al scales prominently spinose, the spines
corresponding to those opercular scales that
have one or both of their edges overlapped
by flanking opercular scales; branching var-
iable, including dichotomous, close-pin-
nate, and lyrate; all scales covered with a
low, often inconspicuous, granulation; 10—
12 perpendicularly oriented polyps/cm.

Description.—Colonies are flabellate,
slightly convex, and occur in three branch-
ing forms: dichotomous, lyrate, and close-
pinnate. The most commonly collected
form is close-pinnate, colonies up to 12 cm
in height and 11 cm in width, with a basal
branch diameter of 1.7 mm, and consisting
of 4 or 5 distinct plumes. Internodes are
only 3—5 mm apart, producing a series of
closely spaced, parallel, wiry branches that
rarely exceed 4 cm in length. Dichotomous
colonies are usually smaller, the holotype
only 6 cm tall, 7.5 cm in width, and having
a basal stem diameter of 0.8 mm. The first
bifurcation occurs 7—11 mm above the sub-
strate; subsequent branching occurs every
3-10 mm, distal branchlet rarely more than

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(46 ‘BIJ 99S) JuOWIASUBIIe ONSTIO}OvIeYS SUIMOYS WIn;NdIedoO uv JO MIA OdI9}s puv dAjod vB JO MAIA [RID}e[ ODIO}s :odAjO[OY ‘VIvajnov D]jJa/DUIN] g ‘G-—Y 8 BIA

+
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jaa
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2)
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468

Fig. 9.

3 cm. The lyrate form is believed to be a
variation of the dichotomous form. The axis
is yellow-gold; polyps (in alcohol) are
white.

Polyps are arranged biserially in the
plane of the flabellum in an alternating
fashion and are well spaced (1 mm apart),
resulting in 10—12 polyps/cm. Polyps are
oriented away from the convex side of the
flabellum (toward the anterior side) and
usually perpendicular to the branchlet. Pol-
yps are distally flared, and including the
elongate marginal spines, measure up to 1.8
mm in height and 0.7—0.8 mm in distal di-
ameter.

Each polyp is protected by 8 operculars
and 8 rows of body wall scales, both ab-
and adaxial rows having the same number
of scales (3 or 4) as the polyp is not curved
toward the branch. Four to seven (usually
5 or 6) of the marginal scales bear extreme-
ly long, slender, sharp (apical angle 8°—10°)
spines, that are cylindrical in cross section.
They are slightly curved over the polyp

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Diagrammatic representation of seven arrangements (a—g) of marginal and opercular scales in Plu-
marella aculeata, as viewed from above the operculum. The eight large triangles of each figure are the 8
operculars; the smaller triangles are the corresponding marginal scale spines. The numbers on the operculars
indicate how many of its two edges overlap an adjacent opercular, the arrows also indicating which edge overlaps
an adjacent opercular. Spines occur only on those marginal scales corresponding to operculars having one or
both of its edges overlapped by an adjacent opercular; marginal corresponding to operculars that are overlapped
by both adjacent operculars do not have a spine.

face and often lack granulation, thus ap-
pearing translucent, or may be covered with
aligned spinules (Fig. 7G). The spine por
tion of the marginal scales constitutes 60—
65% of the length of the scale, resulting in
a H:W of 1.8—2.8. The basal portion of the
spined marginals is massive: rounded or
shield-shaped. The number and position of
marginal spines appears to be directly cor-
related to the corresponding opercular
scales that have one or both of their edges
overlapped by flanking opercular scales.
Opercular scales that overlap both adjacent
operculars do not have a corresponding spi-
nose marginal, their distal margins being
only slightly rounded. Because every polyp
has 8 operculars and thus 16 opercular edg-
es, and every edge must either overlap or
be overlapped by an adjacent opercular, it
is mathematically possible for 4 to 8 oper-
culars to have one or two edges overlapped,
resulting in a polyp with 4—8 marginal
spines (Fig. 9). Polyps having only 4—7
marginal spines have been observed; the

VOLUME 117, NUMBER 4

nent

Preteen

!
4

Fig. 10. A, Plumarella pellucida, holotype; B, P. laxiramosa, holotype; C, P. dichotoma, holotype; D,
Candidella imbricata, Gos-2384; E, Plumarella aculeata, holotype; EK Acanthoprimnoa goesi, Atl-3465, MCZ
3741. Scale bars for A, B, F = 5 cm; C—-E = 2.5 cm.

470

hypothetical 8-spined polyp has not been
seen. Body wall scales of the second and
third tier are large, thick, rectangular, and
have rounded upper edges; they are smooth
or bear only low granules. The fourth tier
of scales consists of small scales indistin-
guishable from the coenenchymal scales.
The opercular scales are isosceles triangular
in shape with a broad base and attenuate
rounded tips that form an apical angle of
20°—25°; their edges are finely serrate and
their upper surface smooth to inconspicu-
ously granular. Operculars are up to 0.62
mm in height and have a narrow range of
H:W of 1.6—2.2.

Coenenchymal scales are flat, irregular in
shape, 0.20—0.40 mm in width or diameter,
and have a conspicuously granular upper
surface. The undersurface of all scales and
the upper proximal sides of most where the
scale is overlapped by an adjacent scale, are
covered with complex tubercles up to 18
zm in diameter.

Etymology.—The species name aculeata
(Latin: aculeatus, sharp-pointed) is an al-
lusion to the extremely long, sharp-pointed
spines of the marginal scales.

Comparisons.—Six species of Plumar-
ella are characterized by having spinose
marginal spines (Kiikenthal 1919); four of
those six endemic to Japan. Plumarella
aculeatus differs from these in having ex-
tremely elongate and sharp marginal spines
that occur only on those marginal scales
that correspond to opercular scales that
have overlapped margins (Figs. 8B, 9).

Distribution.—Insular northern Straits of
Florida; Northwest Providence Channel,
Bahamas; 400—900 m.

Acanthoprimnoa, n. gen.

Type species.—Plumarella goesi Aurivil-
lius, 1931, here designated.

Diagnosis.—Primnoidae with a well-de-
fined operculum; polyps usually inclined
apically, each completely surrounded by 8
rows of body wall scales; polyps arranged
alternately and biserially; 8 marginal scales,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

each with a spinose or finely serrate (pec-
tinate) distal margin; no sclerites bear tu-
bercles on their under surfaces; opercular
scales not keeled; colonies uniplanar, usu-
ally pinnately branched (plumose), dichot-
omous, or lyrate. Brooding polyps are com-
mon.

Distribution.—Straits of Florida, Baha-
mas, Yucatan Peninsula, Lesser Antilles;
Japan; 60-1125 m.

Remarks.—In his unpublished manu-
script on the western Atlantic deep-water
octocorals (Bayer & Cairns 2004), Verrill
referred to the type species of this genus as
Acanthoprimnoa aspera, a species later de-
scribed by Aurivillius as Plumarella goesi.
Because only Verrill’s plates and not the
text survived we do not know what criteria
he used to distinguish his new genus. We
separate this genus from the morphologi-
cally similar Plumarella by three criteria:
the lack of tubercles on the undersurfaces
of the sclerites, the distinctive pectinate dis-
tal edges of the body wall and opercular
scales, and the coarsely granular coenen-
chymal scales. Two other species, previous-
ly placed in Plumarella, also share these
characteristics and are transferred to Acan-
thoprimnoa: A. serta (Kiikenthal & Gor-
zawsky, 1908), n. comb. and A. cristata
(Kiikenthal & Gorzawsky, 1908), n. comb.

Etymology.—The genus name Acantho-
primnoa (Greek: acantha, a thorn + prim-
noa, a common suffix used in this family)
is an allusion to the spiny nature of the pol-
yps of the type species.

Acanthoprimnoa goesi (Aurivillius,
1931), n. comb.
Figs. 8C—D, 10K 11A-—I, 12A—B

?Primnoa Pourtalesii.—Hargitt & Rogers,
1901:281, fig. D.

Plumarella goési Aurivillius, 1931:244—
248, pl. 5, figs. 6a—b, text fig. 47, 3-5. —
Bayer, 1957:388 (Cay Sal Bank).

Thouarella goési Deichmann, 1936:164—
165, pl. 25, figs. 2, 19-23, pl. 26, fig.
8.—Bayer, 1954a:281 (listed).

VOLUME 117, NUMBER 4 471

Fig. 11. A-I, Acanthoprimnoa goesi, G-633: A, upper and undersurfaces of 3 opercular scales; B, pectinate
edge of an opercular; C—D, under and upper surfaces of 2 marginal scales; E, upper and undersurfaces of 3
body wall scales; F—H, upper and undersurfaces of 3 coarsely granular coenenchymal scales, F having a central
spine; I, granules on upper surface of a coenenchymal scale. J-M, Acanthoprimnoa pectinata, holotype: J, upper
and undersurfaces of 5 opercular scales; K, upper and undersurfaces of 4 butterfly-shaped body wall scales; L,
upper and undersurfaces of 3 coarsely granular coenenchymal scales; M, coenenchymal scales on a branch.
Scale bars for A, C-H, J-K, M = 0.10 mm; B, I, L = 25 pm.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

472

“WUT QC'0
= q ‘wu 670 = D-V JJ seq ajeog “déjod v Jo MaIA [eASIe] ODIE}S “[EQ9T PLVMISDY “VIvoLqui vyapipuvD ‘q ‘dAjod & Jo MatA [eIA}v] ODIAIS ‘769-9 ‘YINUNIAad
pouusdoyjupoy “> ‘sourds [eulss1eu SUIpuNnoMs puv winjnosiedo Jo MaIA O919}8 pur dATOd v Jo MoIA [eIO\L] O9I9)S :168-D ‘78208 nouumsdoyjupoy ‘g—yW *Z] “Biq

I UO LT RES

VOLUME 117, NUMBER 4

“‘Acanthoprimnoa aspera” Bayer & Cairns
(Verrill), 2004: pl. 10, fig. 8, pl. 13, fig.
8a, pl. 27, figs. 5a—b, pl. 141, fig. 6.

Material examined.—Alb-2342, 10 dry
pinnate colonies, USNM 10236; Alb-2343,
1 pinnate colony, USNM 10243; A/b-2346,
1 pinnate colony, USNM 10783; Alvin 846,
3 dry pinnate colonies, USNM 79517; Alvin
77-764, 4 pinnate colonies, USNM 96825;
Atl-2999, 10 dry pinnate branches, MCZ
54324 and 3832; Azl-3402, 4 dry pinnate
branches, MCZ 3703; Atl-3403, 7 dry pin-
nate branches, MCZ 54335; Az/-3438, 3 dry
pinnate branches, MCZ 3751; Atl-3463, 7
dry pinnate colonies, MCZ 3604; A7l-3465,
54 dry pinnate colonies, MCZ 3744a, 3741,
and 3737; Atl-3466, 17 dry pinnate colo-
nies, MCZ 3603; .At/-3478, 20 dry pinnate
colonies, MCZ 3605; Atl-3479, 17 dry pin-
nate colonies, MCZ-3608 and 3759; Atl-
3480, 11 dry pinnate colonies, MCZ 3668
and 3762; Aftl-3482, 32 dry pinnate colo-
nies, MCZ 3654 and 3663; Cape Florida
X, 8 pinnate colonies, USNM 73932; JS-
43, 6 pinnate colonies and 1 unnumbered
SEM stub, USNM 43801; JS-102, 1 dry
pinnate colony, USNM 1011364 (topotyp-
ic); JS-103, 3 pinnate colonies, USNM
50951 (topotypic); Eastward 26537, 2 pin-
nate (USNM 98161) and 13 dichotomous
colonies (USNM 98850); Eastward 26538,
10 pinnate branches, USNM 1019537;
Eastward 26549, | pinnate colony (USNM
75064) and 24 dichotomous colonies
(USNM 75065, 76987, and 79485); East-
ward 26550, 53 pinnate colonies, USNM
94500; Eastward 26559, 8 dichotomous
(USNM 98851) and 1 pinnate colony
(USNM 98852); Eastward 31281, 15 pin-
nate colonies, USNM 94522; G-235, 3 pin-
nate colonies, USNM 98853; G-241, 16
pinnate colonies, USNM 52966; G-242, 1
pinnate colony, USNM 52968; G-251, 2
pinnate colonies, USNM 52969; G-252, 1
pinnate branch, USNM 98854; G-254, 1
pinnate colony, USNM 52962; G-387, 12
pinnate colonies, USNM 52970; G-533, 1
pinnate colony, USNM 52971; G-633, 13

473

pinnate colonies and SEM stubs 258 and
C1091, USNM 52973 and 52983; G-679, 8
pinnate colonies and SEM stub 256, USNM
52963; G-680, 4 dichotomous colonies,
USNM 1019538; G-696, 1 pinnate colony,
USNM 52972; G-704, 1 pinnate colony,
USNM 52965; G-706, 25 pinnate colonies,
USNM 52967; G-707, 1 pinnate colony,
USNM 98855; G-879, 2 dichotomous col-
onies, USNM 76988; G-897, 10 dichoto-
mous colonies and SEM stub 257, USNM
52964; P-594, over 50 pinnate colonies,
USNM 52961, and 4 dichotomous colonies,
USNM 98856: P-596, 2 dichotomous col-
onies, USNM 52960; P-598, 1 pinnate col-
ony, USNM 52957 and | dichotomous col-
ony, USNM 98858; specimens reported by
Deichmann (1936) and Bayer (1957); a
syntype (USNM 44192).

Types and type locality.—Three speci-
mens are mentioned in the original descrip-
tion, only one of which was figured; all
three are considered to be syntypes. They
are deposited at the Stockholm Museum
(#28); a fragment of one of the colonies is
also deposited at the USNM (44192). Type
Locality: ““Virgin Islands”’, 457-548 m.

Diagnosis.—Distal edges of 7 marginal
scales (not one of the adaxial marginals)
prominently spinose; branching dichoto-
mous in shallow-water form and close pin-
nate in deeper-water form, branchlets flex-
ible but not flaccid; opercular scales cov-
ered with numerous tiny spines, abaxial and
outer-lateral body wall scales bear single,
robust spines on distal margin; coenenchy-
mal scales highly granular and sometimes
bear a single tall spine; 13—15 polyps/cm;
branch axis bronze; polyp brood chambers
common.

Description.—Colonies are flabellate and
occur in two branching forms. The deeper
water form is larger (up to 30 cm in height
and equally broad), with closely pinnate
branching colonies consisting of 2 or 3 reg-
ular plumes. Pinnate branchlets begins
within 5 mm of the base and are subse-
quently arranged in a regular parallel fash-
ion, the internodes being only 2—3 mm in

474

length; unbranched terminal branchlets are
flexible and rarely exceed 4 cm in length.
These large colonies are attached by a cal-
careous holdfast that may reinforce the
main stem as much as 15 mm above the
base and attain a diameter of 4 mm. As with
most species of Plumarella, the colony fla-
bellum is slight convex, with the polyps di-
rected slightly upward and toward the con-
vex face. The shallow-water form is much
smaller (rarely exceeding 4 cm in height)
and is dichotomously branched, often re-
sulting in a colony broader than tall. Inter-
nodes are 2—4 mm in length; terminal
branchlets are rarely more than 15 mm and
number only about 10—15; the basal axis is
about 0.5 mm in diameter. In both forms
the axis is a rich bronze color, which con-
trasts with the white (in alcohol) color of
the polyps.

Polyps are arranged biserially in the
plane of the flabellum in an alternating
fashion and are well spaced approximately
0.6—0.8 mm apart, resulting in 13—15 pol-
yps/cm. Polyps of the deep-water form are
0.8—1.2 mm in height Gncluding the mar-
ginal spines) and about 0.5 mm in diameter;
polyps of the shallow water form are small-
er, usually less than 0.8 mm in height. As
mentioned in the remarks section, some
polyps of both forms have brood chambers
that greatly swell the base of the polyp.

Each polyp is protected by 8 opercular
and 8 rows of body wall scales, the abaxial
row having 5 or 6 scales, the adaxial, 4 or
5. Seven of the 8 marginal body wall scales
bear a prominent distal spine, the 8th (ad-
axial) marginal having a very reduced
spine, allowing the abaxial opercular scales
to overlap the polyp edge at that point of
the circumference (Fig. 8D). The marginal
scales have a flat, rectangular to ellipsoidal
base up to 0.4 mm wide from which the
elongate, sharp-tipped (apical angle 7°—8°),
often crooked spine emerges. The entire
marginal scale may be up to 0.85 mm in
height, the spinose part constituting 75—
85% of its height and contributing to a rath-
er high H:W of 1.7—3.2. The elongate spi-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

nose part of each marginal scale is spinose
itself, bearing prominent rows of smaller
spines (25-30 wm in length), which are ar-
ranged in rows on both the upper and lower
surface of the spine. The smaller spines also
cover the edges and upper surfaces of the
base of the marginals, but the undersurfaces
of the marginal base, covered by tubercles
in all species of Plumarella, is smooth. The
body wall scales of the abaxial and inner-
lateral rows that lie proximal to the margin-
als also have apical spines, but these are
quite variable in size, some quite large (up
to half the height of the scale), others in-
conspicuous. The body wall scales of the
adaxial and outer-lateral rows that lie prox-
imal to the marginals have greatly reduced
or no apical spines. The opercular scales are
similar to the marginal scales in many ways
but are isosceles triangular in shape, not
having a rectangular base, and often have a
notch on either side near the base. The ab-
axial operculars are quite elongate (up to
0.7 mm) and, when closed, often complete-
ly traverse the polyp. Their tips are pointed
(apical angle 15°—25°), with a H:W ranging
from 1.8—3.9. As with the marginal spines
they are covered with prominent spines on
the upper and undersurfaces, except for the
undersurface of the base, which is smooth.
All three edges of the opercular scales are
serrate, but in the region of the proximal
notches the serrations are developed into
elongate (up to 40 pm long and 9 pm in
diameter), finely granular pillars that often
bi- and trifurcate (Figs. 11A—B).

Coenenchymal scales are rather large (up
to 0.5 mm) and have coarse granular edges
and surfaces, the granules rounded and up
to 12 wm in diameter and often twice as
tall. Some coenenchymal scales also bear a
prominent, centrally located, perpendicular
spine up to 0.3 mm in height and 0.1 mm
in basal diameter. These spines are orna-
mented with smaller spines similar to those
on the opercular and marginal scale spines.
The undersurface of the coenenchymal
scales is smooth.

Comparisons.—See A. pectinata.

VOLUME 117, NUMBER 4

Distribution.—Throughout the Straits of
Florida to Arrowsmith Bank, Yucatan
Channel; Northwest Providence Channel;
Old Bahama Channel; Puerto Rico (Hargitt
& Rogers 1901); Virgin Islands. In general,
the dichotomous form occurs from 137—350
m and the pinnate form deeper, 320-595 m.

Remarks.—The only differences between
the two forms, aside from their different
range of capture depths, are that the deeper-
water form has close pinnate branching, a
larger colony, and larger polyps, whereas
the shallow-water form has dichotomous
branching, a smaller colony, and smaller
polyp size. All other characters are quite
similar, unique characters including the
brooding polyps, spinose body wall and
coenenchymal scales, and long, spiny oper-
culars. Several stations contain both forms
(see material examined), but in general the
forms occur at different depth ranges. Some
colonies are transitional in form, beginning
as dichotomous but with a tendency toward
pinnate branching at least in part of the up-
per colony. Such was the syntype illustrated
by Aurivillius (1931: pl. 5, fig. 6a), al-
though he unequivocally classified that col-
ony as dichotomous.

About one-third of the colonies exam-
ined contained polyps with bulbous brood
chambers in their base, this feature occur-
ring in both the dichotomous and pinnate
forms. Among the colonies containing pol-
yps with brood chambers, approximately
one in 50 polyps would be so modified, but
oftentimes there would be 2 or 3 contiguous
brooding polyps. There appears to be no
seasonality regarding the presence of the
brooding polyps.

Acanthoprimnoa pectinata, n. sp.
Figs. 1D, 11J—M, 12C

Material examined/types and type local-
ity.—Holotype: G-899, 1 colony and SEM
stubs C1093—1095, USNM 1019539. Para-
types: Alb-2354, 20 colonies and 1 unnum-
bered SEM stub, USNM 43026 and 75112;
Alvin 77-760, 5 dichotomous colonies,

475

USNM 1019540; Azl-3303, 3 dry pinnate
colonies, MCZ 3627; G-692, 2 branches
and SEM stub 254, USNM 52954; G-889,
4 colonies, USNM 52952; G-898, 1 pinnate
colony, USNM 52956; G-899, 19 colonies,
USNM 52955; O-4940, 2 colonies, USNM
52953; P-592, 20 colonies, USNM 52958;
P-954, 1 colony, USNM 52959; SB-5190,
6 pinnate colonies, USNM 1019541. Type
Locality: 20°57'N, 86°34'W (off Arrows-
mith Bank, Yucatan, Mexico), 40—164 m.

Diagnosis.—Distal edge of marginal
scales straight or only slightly spinose;
branching loosely pinnate, branchlets long
and flaccid; opercular scales ridged and
covered with numerous tiny spines; lateral
edges of opercular and distal and proximal
edges of body wall scales bear a series of
comb-like spines; coenenchymal scales
coarsely granular, but without a central
boss; 11—13 polyps/cm; branch axis bronze;
polyp brood chambers common.

Description.—Colonies are flabellate and
loosely pinnate, each colony consisting of
2 or 3 plumes, although one colony (P-594)
is lyrate in branching. The first branchlets
occur very near the base of the colony, suc-
ceeding branchlets at a periodicity of every
4—5 mm (internode length), the branchlets
up to 55 mm in length and flaccid in ten-
sion. The main stem is anchored by a dense,
calcareous, white holdfast, although the
holdfasts of only five of the colonies are
intact. Like A. goesi, the axis is bronze in
color, which contrasts with the white pol-
yps. The holotype is 18 cm in height, 7 cm
in width, but lacks a base; the largest spec-
imen (A@/-3303) is 22 cm tall. The largest
main stem of an attached colony (G-899)
has a diameter of only 1.1 mm.

Polyps are arranged biserially on the
branchlets and main stem (6—7 polyps/in-
ternode on main stem) in an alternating
fashion 0.8—1.1 mm apart, resulting in 11—
13 polyps/cm. Polyps are relatively small,
only 0.8—0.9 mm in height and 0.40—0.45
mm in diameter. Colonies from all stations
recorded contain some polyps with brood

476

chambers, which greatly swell the base of
those polyps.

Each polyp is protected by 8 opercular
and 8 rows of body wall scales, the abaxial
row consisting of 8—10 scales, the adaxial,
7-9. All body wall scales, including the
marginals, are slightly curved to accom-
modate the curvature of the polyp, and con-
siderably wider than tall, such that a rela-
tively high number occurs in the wall of a
relatively short polyp. The upper surface of
the body wall scales bears many small
spines, especially toward the center of the
scale, and their distal and proximal margins
bear a series of fine, comb-like (pectinate)
projections measuring up to 32 wm in
length. Only rarely will the marginal body
wall scale have a larger, projecting spine,
the largest up to 0.25 mm in length and
constituting about half the height of the
scale. The opercular scales are isosceles tri-
angular in shape (H:W = 1.6—2.2), and
strongly curved in order to cover the top of
the rounded polyp. Operculars are up to
0.38 mm in height and have an apical angle
of 35°—45°. They are sculptured as in A.
goesi.

Coenenchymal scales are relatively small
(0.09—0.21 mm in width) and circular to ir-
regular in shape. As in A. goesi, they are
densely covered on their upper surface with
prominent, blunt granules measuring up to
15 wm in height and 10—12 pm in diameter,
but smooth on the undersurface.

Etymology.—The species name pectinata
(Latin: pectinatus, comblike) refers to the
comb-like serration of the edges of the
opercular and body wall scales.

Comparisons.—Acanthoprimnoa pectin-
ata resembles A. goesi in the morphology
of its opercular spines, color of the branch
axis, coarsely granular coenenchymal
scales, and the common presence of brood
polyps. Further, A. pectinata differs (Table
1) in lacking distally spinose body wall
scales (instead having pectinate distal and
proximal margins), lacking a central boss
on the coenenchymal scales, having more
scales/body wall row, having much shorter

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

operculars, and in having a looser pinnate
branching mode. A. pectinata is most sim-
ilar to the Japanese A. cristata, but lacks the
longitudinal ridges on the body wall scales.

Distribution.—Off northeastern Yucatan
Peninsula and northwestern Cuba (164—476
m); Straits of Florida; Mona Passage and
off Montserrat, Lesser Antilles (614—686
m).

Remarks.—All but two colonies of A.
pectinata occur in relatively shallow water
(164—476 m) off the Yucatan Peninsula, but
the colonies from P-954 (off Montserrat,
Lesser Antilles) and Alvin 77-760 (Straits
of Florida) occur in deeper water (614—686
m) and are the only colonies to have non-
pinnately (dichotomous, lyrate) branching
colonies.

Genus Candidella Bayer, 1954

Primnoa.—Johnson, 1862:245 (in part).
Stenella Gray, 1870:48 (Gunior primary
homonym of Stenella Gray, 1866, a ce-
tacean).—Wright & Studer, 1889:56 (Gn
part).—Ktkenthal, 1919:443—445 Gn
part); 1924:303 (Qn _ part).—Aurivillius,
1931:289—290 (in part).
Narella.—Studer, 1878:643 (Gn part).
Stenella (Primnoa).—Roule, 1896:304.
Stenella (Stenella).—Versluys, 1906:38—39.
Candidella Bayer, 1954b:296 (nom. nov.);
1981:937.—Tixier-Durivault, 1987:171.
Candidella (Candidella).—Bayer, 1956:
F222.

Type species.—Primnoa imbricata John-
son, 1862, by monotypy.
Diagnosis.—Primnoidae with a well-de-
fined operculum; polyps stand perpendicu-
lar to branch (not bent); polyp body wall
completely surrounded by 2—4 rows of
sclerites; polyps arranged in whorls; only
four marginal scales; undersurfaces of all
sclerites tuberculate, opercular scales
strongly keeled; colonies dichotomously
branched in one plane.
Distribution.—North Atlantic, Ascension,
central and western Pacific; 183-2139 m.
Remarks.—Four species are known in

VOLUME 117, NUMBER 4

this genus: C. imbricata (Johnson, 1862);
C. johnsoni (Wright & Studer, 1889), As-
cension; C. gigantea (Wright & Studer,
1889), Fiji; and C. helminthophora (Nut-
ting, 1908), Hawaiian Islands. After its
original description, the monographers Kiu-
kenthal, Studer, and Aurivillius took a
broad view of the genus Stenella, including
similar species but some differing in having
5 or 8 marginal scales, these species later
being transferred to Parastenella, Pteros-
tenella, and Dasystenella. Versluys (1906)
was the first to relegate what is now known
as Candidella to a monophyletic group, the
nominate subgenus of Stenella. After re-
naming the genus Candidella (Bayer,
1954b), because the name Stenella was a
junior homonym, Bayer (1956) also recog-
nized it as a monophyletic subgenus: Can-
didella (Candidella), subsequently elevat-
ing it to generic rank in 1981. Characters
used to distinguish species include the ar-
rangement of polyps, colony branching, and
polyp size (Kiikenthal 1924).

Candidella imbricata (Johnson, 1862)
Figs. 1OD, 12D, 13A—G, 14A—D

Primnoa imbricata Johnson, 1862:245, pl.
31, figs. 2, 2a (Madeira); 1863:299 (ver-
batim).

Stenella imbricata.—Gray, 1870:48—49, 2
figs. (listed, new comb.).—Wright & Stu-
der, 1889:56, 281 (listed).—Kiikenthal,
1919:448—449 (Blake from Cuba, first re-
cord for western Atlantic); 1924:305—306
(diagnosis, key).—Thomson, 1927:32—
33, fl, 2, m3, GS, lla 3, ins, D5 Toll, DS, ws
5—6 (Azores, Morocco).—Aurivillius,
1931:290 (mentioned).—Deichmann,
1936:167—168, pl. 26, fig. 5 (West In-
dies).—Bayer, 1954a:281 (listed for Gulf
of Mexico); 1964:532 (Straits of Florida).

Narella imbricata.—Studer, 1878:643 (list-
ed, new comb.).

?Stenella (Primnoa) johnsoni.—Roule,
1896:304 (Gulf of Gascogne).

Stenella (Primnoa) imbricata.—Roule,
1896:304 (comparison to C. johnsoni).

477

Stenella (Stenella) imbricata.—Versluys,
1906:42—43, 44, fig. 46 (redescription of
type, key to spp.)

Candidella imbricata.—Bayer, 1954b:296
(new. comb.).—Tixier-Durivault &
d’Hondt, 1974:1412—1413.—Grasshoff,
1981:222, map 1 (mid-Atlantic Ridge sw
of Azores); 1982a:738, maps 4, 20;
1982b:948—949, figs. 20—21.—Grasshoff
& Zibrowius, 1983:119—-120, 122, pl. 2,
fig. 7 (mid-Atlantic Ridge), pl. 3, fig. 14
(Biscay Bay).—Carpine & Grasshoff,
1985:6 (frontispiece), 33 (Musée Océan-
ographique de Monaco catalog num-
bers).—Pasternak, 1985:29 (Rockaway
Seamount).—Pettibone, 1991:705, 707
(polychaete commensal).

Candidella (Candidella) imbricata.—Bay-
er, 1956:F222, fig. 159—4b.

Candidella johnsoni.—Bayer, 1981:934,
fig. 74.

Stenella “‘florida” Bayer & Cairns (Verrill),
2004: pl. 13, figs. 1, la, pl. 25, fig. 13a—
b, pl. 82, figs. 2, 2a, pl. 83, fig. 6, 6a.

Material examined.—Alb-2753, 3 branch
fragments, USNM 44126; Alvin 762, 6
branches, USNM 80939, 80940, and
1017255; Alvin 1335, 2 fragments (one
dry), USNM 73744 and 73745; Alvin 3885-
5, 1 complete colony, USNM 1019238; Al-
vin 3903-101-—2, 1 branch, USNM
1019273; Atl-266-47, branch fragments,
USNM 60337; Atl-280-9, 3 branches and
SEM stub 273, USNM 57552; CI-63, 1 col-
ony, USNM 60223; CI-140, 1 branch,
USNM 60341; Eastward 26019, 6 colonies,
USNM 60338; Eastward 26022, 2 colonies,
USNM 60340; Eastward 26023, dry branch
fragments, USNM 1011365; Eastward
26031, 3 colonies (some dry) and SEM
stubs 274 and C1071—1076, 1078, USNM
57553 and 60339; G-169, 6 colonies,
USNM 52778; G-170, 2 colonies and 1 un-
numbered SEM stub, USNM 52779; G-
177, 2 colonies, USNM 52780; G-386, 14
colonies and numerous branches, USNM
52784; G-660, 1 branch, USNM 52781; G-
661, 1 branch, USNM 52782; G-936, 2

478

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 13.
tuberculate undersurface of an opercular; C, upper surface of 2 basal body wall scales; D, K upper and under-
surfaces of 3 medial body wall scales; E, undersurface of 2 marginal scales; G, upper and undersurfaces of 3
coenenchymal scales. Scale bars for A, C-G = 0.25 mm; B = 25 pm.

branches, USNM 52783; G-965, 1 colony,
USNM 52787; Gos-2383, 1 colony, USNM
57309; Gos-2384, 1 colony, USNM 57310;
Gyre CO4, | colony (dry), USNM 89124;
P-197, 2 colonies, USNM 52785; P-881, 2
colonies, USNM 52786; P-892, 2 colonies,
USNM 52911; P-1146, 1 branch, USNM
52912; off Bermuda, 1200 m, 1 colony,
USNM 75104.

Types and type locality.—The holotype

A-G, Candidella imbricata, Gos-26031: A, upper and undersurfaces of 4 opercular scales; B,

is deposited at the BM (1863.1.31.1). Type
locality: Madeira, depth unknown.
Description.—Colonies consist of a ro-
bust vertical main stem up to 9 mm in basal
diameter, which supports a uniplanar fan
achieved by dichotomous branching. The
main stem is anchored by a dense, white,
encrusting, calcareous holdfast, which often
encrusts other calcareous Coelenterata, such
as the scleractinians Enallopsammia pro-

‘WU SZ’ =
q ‘wur gy, = q ‘wu Qs’9 = D-V JO} seq a[eog ‘sayeos Iejno1odo ouros Jo dn asojo ‘gq ‘dAjod & Jo apis [elxepe oY) JO MOIA OdIOIS ‘OD :[EQ9T PuvMIsD” “q ‘D ‘dn
youvig ‘q ‘se[evos [eUISIeUI INOJ dy) SuIMOYs Ose ‘MIA Ie[NoIedo O919}s “gq ‘dAjod B JO MIA [eIO}V] OOIO}S “WV :6-O8T-4V ‘d ‘A-V :BIpoluqui DjjapipuvD, “p| “314

Se VES a= kee Le ERE

480

funda, Lophelia prolifera, Javania cailetti,
the stylasterid Stylaster erubescens, and
various bryozoans. The calcareous deposits
may reinforce the basal stem as much as 2
cm upwards from the base. The holdfast
and basal reinforcement are composed of
100% aragonite, consistent with the find-
ings of Bayer & Macintyre (2001) for the
congeneric C. helminthophora. The axis is
yellow-gold in color and longitudinally stri-
ate; overall the colony is white. The largest
known colony (the holotype) is reputed to
be 21.6 cm in height and 27.9 cm in width.
Branching is dichotomous at intervals of 3—
12 mm, but unequal, resulting in asymmet-
rical branching; there is little to no branch
anastomosis.

Polyps are arranged in whorls of 3 or 4
polyps (rarely as pairs); if in a whorl of 3,
2 polyps are usually directed in the plane
of the fan in opposite directions, the third
polyp standing perpendicular to the plane
of the fan and thus at 90° to the other 2, the
polyp projecting perpendicular to the fan
defining the anterior face of the fan; few
polyps originate from the posterior face of
the fan. When 4 polyps constitute a whorl,
the angular separation between polyps is
not 90°, but about 60°, polyps avoiding the
posterior face. Polyp whorls are closely
spaced, about every 1.2—2.0 mm, 5—6 oc-
curring per cm, polyps present even on the
calcified region of the basal stem and hold-
fast. Most polyps are 2.1—2.5 mm in height
and slightly clavate (1.3—-1.4 mm in distal
diameter), encased by the distal margin of
the flared marginal scales, but some geo-
graphic outliers have larger polyps (see Re-
marks). Polyps are fairly rigid, projecting
perpendicularly from the branches; howev-
er, those in the plane of the fan are some-
times slightly curved toward the anterior
face.

Polyps are protected by 4 marginal
scales, 2—4 medial scales, 4—8 basal scales,
and 8 operculars. The marginal scales are
dimorphic in size and shape, consisting of
2 adjacent larger (0.9 mm in height, 1.1 mm
in width), highly curved scales that define

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

the abaxial side of the polyp and 2 adjacent
smaller (0.65 mm in height, 0.62 mm in
width), slightly curved adaxial scales,
which overlap with the edges of the larger
marginals. Three opercular scales corre-
spond to each of the larger marginals,
whereas about 1.5 operculars correspond to
the smaller marginals, the number of oper-
culars adding to more than 8 because of the
overlap of marginal scales. The marginals
are flared outward distally, rising about 0.15
mm above the junction with the opercular
scales, but not enclosing the operculum.
Medial body wall scales are roughly rect-
angular and flat, with sides measuring 0.45—
0.65 mm in length; their lateral edges over-
lap one another. Sometimes it appears that
only 2 medial scales are present, these oc-
curring on the adaxial side. Basal scales are
dimorphic in size, consisting of 2 large,
square to rectangular scales up to 0.65—0.70
mm in side length, and outwardly concave,
as though squeezing the base of the polyp
into a narrow opening. When polyps be-
come abraded from the branches, these
large scales often remain to mark the orig-
inal position of the polyp. Between the 2
large basal scales, on the adaxial side, are
1 or 2 pairs of much smaller basal scales
that are overlapped and overshadowed by
the larger basals. The 8 operculars are elon-
gate triangular, having a H:W of 1.6—2.0,
pointed distally, highly convex above, and
prominently keeled below. They form a
tight conical operculum over the polyp, ris-
ing well above the marginal scales. One of
the 8 operculars is slightly larger (e.g., 0.8
mm tall, 0.5 mm wide) than the others and
is positioned opposite the smallest opercu-
lar (e.g., 0.6 mm tall, 0.3 mm wide), these
two operculars defining the sagittal axis of
the polyp. The remaining 6 operculars are
of similar size, constituting 3 pairs mirrored
across the sagittal axis. The 2 sagittal oper-
culars are symmetrical, in that their keels
are in a medial position, whereas the other
6 operculars are asymmetrical, their keels
being offset toward the abaxial side (the
side toward the large sagittal opercular),

VOLUME 117, NUMBER 4

producing a longer and slightly upturned
edge of their adaxial side. Each upturned
adaxial opercular edge overlaps the abaxial
edges of the adjacent operculars, the edges
of the small sagittal opercular being over-
lapped by both adjacent operculars and the
large sagittal opercular overlapping both
adjacent operculars (compare to Fig. 9f).

Coenenchymal scales are large (up to 1.0
mm in length), occur in one layer, are po-
lygonal in shape, and are usually slightly
concave above. As mentioned below, they
sometime orient perpendicular to the branch
in order to contribute to the formation of
the worm tube. The upper surfaces of all
sclerites are finely and uniformly granular,
the granules 11-13 sm in diameter; their
undersurfaces are covered with complex tu-
bercles 15-17 wm in diameter. Tentacular
sclerites were not noted.

Comparisons.—There is only one other
species of Candidella known from the At-
lantic, C. johnsoni (Wright & Studer, 1889),
described from Ascension. As summarized
by Versluys (1906), that species differs in
having a very low operculum, marginal
scales that are equal in size, and polyps that
occur in pairs and singly. Although these
two species are probably distinct, the only
subsequent report of C. johnsoni is by Rou-
le (1896) from the Gulf of Gascogne, which
is probably C. imbricata, as he implied that
his C. johnsoni might be a deep-water va-
riety of C. imbricata.

Candidella imbricata is morphologically
more similar to the central Pacific C. hel-
minthophora (Nutting, 1908), both species
having dimorphic marginal scales and a
similarly shaped polyp. However, C. hel-
minthophora differs in having two rings of
medial body wall scales, a larger colony
with longer internodes (up to 4 cm), larger
polyps, and more flexible branches.

Distribution.—Western Atlantic: New
England Seamounts (San Pablo, Rockaway,
Kelvin, Muir), Bermuda, eastern coast of
Florida, Bahamas, Greater and Lesser An-
tilles, northern Gulf of Mexico; 514—2063
m. A rather large distributional gap exists

481

between the New England Seamounts and
the coast of Florida. Eastern Atlantic: com-
monly collected in the Bay of Biscay, off
Morocco, Canary Islands, Madeira, Azores,
and the mid-Atlantic Ridge southwest of
the Azores; 815—2139 m (see Grasshoff
1982b for map).

Remarks.—Colonies of even small size
will usually host the commensal polynoid
polychaete Gorgoniapolynoe caeciliae
(Fauvel, 1913), larger colonies often host-
ing 5 or 6 worms. The polychaete has es-
sentially the same known distribution as C.
imbricata, despite the fact that it occurs in
at least two other gorgonians (Pettibone
1991). The gorgonian appears to be induced
to form a tube that is slightly elliptical in
cross section, the greater diameter being ap-
proximately 2.3—2.5 mm and the length up
to 25 mm, the tube always occurring on the
anterior side of the fan; the length of the
polychaete is about 11 mm. The tubes are
formed predominantly of greatly enlarged
and outwardly curved basal scales from two
adjacent polyps. These basal scales, nor-
mally only 0.7 mm in height, increase in
size up to 1.6 mm in height and up to 2.9
mm in width. The curvature is such that
basal scales from two adjacent polyps in the
same whorl meet and sometimes fuse along
the dorsal midline of the tube, whereas the
proximal and distal edges of these enlarged
basal scales meet and sometimes fuse with
those of adjacent whorls, altogether form-
ing a somewhat porous tube that is open at
both ends. Occasionally, small coenenchy-
mal scales that project perpendicular to the
branch will fill in the spaces between basal
scales of adjacent whorls. Although an ob-
vious advantage is gained for the worm in
this association, no advantage can be con-
jectured for the gorgonian.

Several specimens, collected at the mar-
gins of the known distribution, show some
variation in morphology. The single speci-
men known from the northern Gulf of Mex-
ico (USNM 89124) has a very low oper-
culum, like that of C. johnsoni, but other-
wise is similar to C. imbricata. The colo-

482

nies from Bermuda (USNM 75104) and
San Pablo Seamount (USNM 57552) have
unusually large polyps, 4.0 and 3.2 mm, re-
spectively, but are otherwise similar to C.
imbricata.

Acknowledgments

We wish to thank Ardis Johnston for the
loan of Plumarella specimens deposited at
the MCZ, and Elly Beglinger (Zoological
Museum, Amsterdam) for the loan of typi-
cal specimens of Plumarella penna. We
thank Ian Macintyre for the mineralogical
determination of the axis of C. imbricata.
Molly Ryan, staff illustrator, produced Fig-
ure 9, and Tim Coffer helped produce the
plates. Specimens of C. imbricata from AI-
vin stations made in 2003 were collected by
the “‘Mountains-in-the-Sea’’ Expedition,
Les Watling, Chief Scientist, funded by the
NOAA Ocean Exploration program.

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Associate Editor: Stephen L. Gardiner

484 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

APPENDIX: Station Data

Station Latitude (°N) Longitude ((W) Depth (m) Date

U.S.F.C.S Albatross

2342 23°10'39" 82°20'21” 384 19 Jan 1885
2343 23°11'35” 82°19'25" 510 19 Jan 1885
2346 23°10'39” 82°20'21” 366 20 Jan 1885
2354 20°59’30" 86°23'45" 238 22 Jan 1885
2416 31°26' 79°07' 505 1 Apr 1885
2662 29°24'30" 79°43' 794 4 May 1886
2663 29°39’ 79°49' 770 4 May 1886
2666 29°47'30" 79°49' 494 5 May 1886
2667 30°53’ 79°42'30" 499 5 May 1886
2668 30°58'30" 79°38'30" 538 5 May 1886
2669 31°09’ 79°33'30" 644 5 May 1886
2753 13°34’ 61°03’ 514 4 Dec 1885
IIIl9-19 off Beaufort, N. Carolina ? 27 May 1949
Alvin (submersible)
77-760 27°04.9' 79°20.1' 613-654 Jun 1977
77-761 27°04’ 79°18.8' 600 Jun 1977
77-762 27°03.3’ 79°20.0’ 600 Jun 1977
771-164 27°55.8' 79°09' 410 Jun 1977
846 26°26' TES. 525 3 Nov 1978
1335 27°05' 79°40' 608 21 Feb 1984
3885-5 33°46.17’ 62°33.9' 1821 4 Jun 2003
3903-101-2 38°47.33’ 64°07.95' 2063 15 Jul 2003
Anton Dohrn
6392 30°49’ 79°49' 400 2
65-32 Tortugas 1064 30 Jul 1932
R/V Atlantis (Atl)
266-02 31°58’ 77°18.5' 813 25 Jun 1961
266-04 31°56’ 77°26' 768 26 Jun 1961
266-07 31°53’ YDS! 750 28 Jun 1961
266-40 30°53’ T8°47' 804 13 Jul 1961
266-41 30°59’ 78°14’ 877 15 Jul 1961
266-47 30°53’ T8°47' 819 19 Jul 1961
280-09 38°51'18” 60°29'00" 1902 17 Jun 1962
2999 23°10’ 81°29’ 265-512 17 Mar 1938
3303 23°05’ 82°33’ 476 23 Mar 1939
3402 22°36’ 78°21' 421 28 Apr 1939
3403 22°36’ 78°22 384 28 Apr 1939
3438 23°05’ 79°37' 485 2 May 1939
3463 23°09’ 81°26 421 9 May 1939
3465 23°09’ 81°27’ 320 9 May 1939
3466 23°09’ 81°27’ 366 9 May 1939
3478 23°09’ 81°27'30" 240 11 May 1939
3479 23°10’ 81°26’ 384 11 May 1939
3480 23°10’ 81°28’ 366 11 May 1939
3482 23°09’ 81°27’ 348 11 May 1939
3780 30°27' USS) 458-485 24 Feb 1940
3782 30°10! 78°44’ 795-804 24 Feb 1940
U.S.C. S. S. Bibb
19 23°03’ 83°10/30" 567 4 May 1868
Dp) 24°14'20" 80°59'40" 567 4 May 1868

135 24°20'30" 81°58'30" 229 17 Feb 1869

VOLUME 117, NUMBER 4 485

APPENDIX: Continued

Station Latitude (°N) Longitude (°W) Depth (m) Date

Johnson-Smithsonian Deep-Sea Expedition (JS)

43 18°04’ 67°48’ 439-549 11 Feb 1933
102 18°51’ 64°33’ 90-500 4 Mar 1933
103 18°51’ 64°33’ 274-732 4 Mar 1933
R/V Cape Florida
x 27°31’ 79°15' 350-400 Jul 1984
R/V Cape Hatteras
SA6 31°18'08”" 79°00'08” 545-549 17 Nov 1985
SA6-1 31°17'18" 79°00'39”" 572-575 17 Nov 1985
SA6-5 31°49'40" 78°19'16" 625 18 Nov 1985
R/V Colombus Iselin (CI)
63 28°06’ 77°08' 1023-1153 21 Sep 1980
123 24°12'06" T7T18' 1435 24 Sep 1973
140 26°24 79°36' 738 28 Sep 1973
246 26°23’ 79°37' 743-761 29 Oct 1974
266 24°18.5' WP iy" ? 3 Nov 1974
Clelia (submersible)
78 32°43'38" 78°05'38” 175-196 5 Jul 1993
T9A 32°43'38" 79°05'50” 210 7 Jul 1993
M/V Combat
174 34°45’ 75°28' 320 14 Nov 1956
368 34°15’ T5231 348 16 Jun 1957
Discoverer
x 32°10’ 78°07'18" 446 5 Oct 1967
R/V Eastward
26004 28°08’ 79°33' 785-830 Nov 1974
26017 26°38'30" 79°32'30" TIS Nov 1974
26019 27°16'48”" 79°25' 655-685 Nov 1974
26022 27°28'30" 79°25'18" 655-685 Nov 1974
26023 DIPS 79°22' 690 Nov 1974
26028 27°09.5’ 79°25' 635-700 Nov 1974
26031 27°00' 79°24'18" 645-690 Nov 1974
26052 25°42.7' 79°47.5' 660-770 Nov 1974
26535 DINGO! 79°15.6' 480 29 Mar 1975
26537 2TimAY 79°15’ 520 29 Mar 1975
26538 DPD! 79°13.7' 420 29 Mar 1975
26547 27°18’ 79°17' 520 Mar 1975
26549 27°17'30" 79°12'30" 370 30 Mar 1975
26550 27°16'24” 79°14'18" 440 30 Mar 1975
26559 26°30.3’ 79°14.7' 2 31 Mar 1975
31281 26°53'54" 79°07'18" 320 1977
R/V Gerda (G)
56 DSB" 79°20' 458 28 Aug 1962
169 27°01’ 79°21.5' 229-275 29 Jun 1963
170 27°06’ 79°32’ 659-677 29 Jun 1963
7 2T°17' 79°34 686 30 Jun 1963

235 25°44’ PDs 531 30 Jun 1963

486 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

APPENDIX: Continued

Station Latitude (°N) Longitude ((W) Depth (m) Date
241 25°26’ 79°18" 494-502 30 Jan 1964
242 25°36’ 79°21' 485-530 30 Jan 1964
246 26°57’ 79°12.5' 512 5 Feb 1964
247 27°07’ 79°21" 567 5 Feb 1964
251 Dif Dy 78°41’ 293-311 5 Feb 1964
252 DDO! 78°37.5' 485-496 5 Feb 1964
254 igs Amo 78°49' 488-516 6 Feb 1964
261 27°20’ TO D24 494-511 7 Feb 1964
386 27°09" 78°18’ 604 19 Sep 1964
387 DP 79°15’ 412 19 Sep 1964
391 27°20’ USD! iy 19 Sep 1964
533 26°27' 78°43’ 383-403 4 Mar 1965
598 24°47’ 80°26’ 183 15 Apr 1965
633 25°59’ I9A9’ 479-458 30 Jun 1965
647 26°16’ 79°43’ 520-549 15 Jul 1965
660 26°59’ 79°21’ 631 17 Jul 1965
661 27°07’ 79°32" 695-718 17 Jul 1965
664 DBS 79°22’ 567 17 Jul 1965
672 DSB) 79°03" 796 18 Jul 1965
679 25°56’ 78°09’ 595-711 20 Jul 1965
680 25°56’ 78°05’ 571-657 20 Jul 1965
692 26°34’ 78°25’ 329-421 21 Jul 1965
695 26°28’ 78°37' 535-575 22 Jul 1965
696 26°28’ 78°43’ 458-467 22 Jul 1965
704 26°29’ 78°40’ 275-366 22 Jul 1965
706 26°27’ 78°43’ 489-522 22 Jul 1965
707 26°27’ 78°40’ 514-586 22 Jul 1965
785 24°39" 80°40’ 205-210 16 Aug 1966
808 26°38’ 79°33’ 751 13 Sep 1966
835 24°22' 81°11’ 187-198 11 Jul 1967
859 23°54’ 81°57’ 1160-1190 21 Aug 1967
879 21°00 86°25’ 210 9 Sep 1967
889 20°55’ 86°28’ 175-220 10 Sep 1967
897 20°59’ 86°24" 210-290 10 Sep 1967
898 21°04’ 86°19" 340-360 10 Sep 1967
899 20°57’ 86°34’ 40-164 10 Sep 1967
936 26°35’ 79°20’ 600 1 Oct 1967
965 23°45’ 81°49" 1394-1399 1 Feb 1968
1012 23°35’ 79°33’ 509-531 14 Jun 1968
1125 26°45’ 79°05’ 900-950 17 Jun 1968
1312 26°38’ 79°02’ 505-527 31 Mar 1971
1314 26°52’ 79°11’ 532 ?

R/V Gosnold (Gos)

2344 30°29’ 77°29.5' 882 ?
2383 30°56'24” 78°34'18" 869 27 Aug 1965
2384 30°54'24" 78°44'00" 820 27 Aug 1965
2385 30°57'12" 78°54'36" 379 27 Aug 1965
2387 31°14'48” 78°59'00" 530 27 Aug 1965
2413 30°14.5' 79°44.7' 585-622 2 Sep 1965
2414 30°16' 79°55.1' 494 3 Sep 1965
2461 28°14.4' 79°30.5' 850 15 Sep 1965

2469 29°43'12" 79°51'48" 640 16 Sep 1965

VOLUME 117, NUMBER 4

APPENDIX: Continued

Station Latitude (°N) Longitude ((W) Depth (m)

M/V, R/V Oregon and Oregon IT (O)

1328 24°33’ 83°34’ 366
1343 22°59' T9°1T' 457
1349 24°03’ 80°30’ 274
4940 20°30' 86°14’ 310-330
11703 30°28’ 79°51’ 494
11705 30°26 79°44! 640
11716 30°52’ 79°39' 576
11717 30°52’ 79°34’ 658
11725 31°44" 79°02' 543
11726 31°42’ 78°53’ 512
R/V Gyre
CO4 27°28'06" 89°43'36" 1358-1518
R/V Pillsbury (P)
105 30°28’ 79°42’ 388-403
197 27°59’ 79°20' 567-586
209 26°59' 79°16' 550
592 215007 86°23’ 180
594 21°00.5' 86°23.0’ 330
596 24°42" 80°32’ 137
598 21°07' 86°21’ 155-205
881 13°20.8’ 61°02.5' 576-823
892 14°17’ 60°45'12” 1236-1313
954 16°55’ 62°43’ 686-1125
1146 20°08’ WG 1110-1189
M/V, R/V Silver Bay (SB)

440 D9 oa 79°15! 439-503
453 29°38’ 78°26! 879
5190 18°24’ 68°05’ 366

487

Date

Jul 1955

18 Jul 1955
12 Jun 1964
19 Jan 1972
19 Jan 1972
21 Jan 1972
21 Jan 1972
22 Jan 1972
12 Jan 1972

13 Apr 1984

27 Jul 1964
11 Aug 1964
12 Aug 1964
15 Mar 1968
15 May 1968
15 May 1968
15 May 1968
6 Jul 1969

6 Jul 1969
16 Jul 1969
14 Jun 1970

8 Jun 1958
12 Jun 1958
17 Oct 1963

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(4):488-504. 2004.

A new species of the sea anemone Megalactis (Cnidaria: Anthozoa:
Actiniaria: Actinodendridae) from Taiwan and designation of a
neotype for the type species of the genus

Adorian Ardelean and Daphne Gail Fautin

(AA) Universitatea de Vest din Timisoara, Facultatea de Chimie-Biologie-Geografie, str.
Pestalozzi, nr 16, Timisoara, 1900, Romania, e-mail: adorian@mynature.net;
(DGF) Department of Ecology and Evolutionary Biology, and Division of Invertebrate Zoology,
University of Kansas Natural History Museum and Biodiversity Research Center,
University of Kansas, Lawrence, Kansas 66045 U.S.A., e-mail:fautin@ku.edu

Abstract.—Megalactis comatus, new species, from Taiwan is the third species
in this genus of sea anemones with highly branched tentacles. The others are M.
hemprichii Ehrenberg, 1834, from the Red Sea, and M. griffithsi Saville-Kent,
1893, from the Great Barrier Reef. Size of nematocysts from acrospheres and
column clearly separate M. comatus from the other species of Megalactis. One
of us (A.A.) observed asexual blastulae in VM. comatus. This is the first record
of asexual reproduction in the genus. Because type specimens of M. hemprichii
have not been found and the original description cannot be used to distinguish
this species from other species of Megalactis, we designate a neotype for the
type species of the genus, M. hemprichii Ehrenberg, 1834. All the specimens of
actinodendrids examined lacked basilar muscles; this calls into question the
placement of family Actinodendridae among thenarian sea anemones.

The family Actinodendridae is a group of
three genera of exclusively tropical Indo-
Pacific sea anemones: Actinodendron
Blainville, 1830, Megalactis Ehrenberg,
1834, and Actinostephanus Kwietniewski,
1897. An actinodendrid has the oral disc
drawn out into a number of branched ten-
tacles that make it resemble a tree (Blain-
ville 1830, 1834; Quoy & Gaimard 1833;
Haddon 1898; Carlgren 1949). The last
branches of tentacles terminate in acro-
spheres that appear as white swellings of
tissue; they are packed with nematocysts
and spirocysts. Because the actinodendrids
have been documented to sting humans
badly (Saville-Kent 1893, Halstead 1970),
knowledge of these animals is significant
not only for taxonomy and phylogeny, but
also for medicine and toxicology.

Actinodendridae was considered by Carl-
gren (1900, 1949) to belong to the supra-
familial group Thenaria. Basilar muscles,

which are structures “running along both
sides of the base of the mesentery, close to
the pedal disc” (Carlgren 1949, p. 8), were
used by Carlgren (1899, 1900, 1942, 1949)
to define two major groups in sea anemo-
nes, Athenaria ““Nyantheae without basilar
muscles” (Carlgren 1949, p. 21) and Then-
aria ‘“‘Nyantheae with basilar muscles”
(Carlgren 1949, p. 41). We did not find bas-
ilar muscles in specimens of actinodendrids
studied, which makes placement of Actin-
odendridae among Thenaria questionable.
The morphology of the tentacles of these
sea anemones varies with environment, be-
havior, and conditions of preservation. Al-
though the number of species described in
Actinodendridae is small, the lack of ter-
minology for describing branched struc-
tures and the enormous variety that can be
found makes identification of species diffi-
cult. In this paper we describe one species
and redescribe two others of Megalactis,

VOLUME 117, NUMBER 4
and standardize terminology for the
branched tentacles of Actinodendridae.

Actinodendrids are found in shallow wa-
ter in sheltered places with sandy or muddy
bottoms. Members of the genus Megalactis
reportedly attach the pedal disc to hard sub-
strata in sand or mud into which the anem-
ones burrow (Saville-Kent 1893, Fishelson
1970). The new species of Megalactis de-
scribed here lives in thickets of the scler-
actinian coral Acropora in Taiwan; this
might be the same species as that reported
by den Hartog (1997) as an unidentified ac-
tinodendrid living attached to coral branch-
es in Indonesia.

The description of Megalactis hemprichii
Ehrenberg, 1834, the type species of the ge-
nus, was diagnostic in the early 19th cen-
tury. Mentioning only that a sea anemone
had bipinnately branched tentacles was suf-
ficient to distinguish M. hemprichii from all
sea anemones known at that time. With the
current state of knowledge, the original de-
scription of M. hemprichii does not distin-
guish it from other species of Megalactis:
bipinnate disposition of the branches is ge-
neric rather than specific. Type specimens
of M. hemprichii Ehrenberg, 1834 have not
been found (Klunzinger 1877, Favtin 2004
Hexacorallians of the World: http://hercules.
kgs.ku.edu/hexacoral/anemone2/index.cfm).
We designate a neotype for M. hemprichii
in accordance with Article 75.3 of the In-
ternational Code of Zoological Nomencla-
ture (International Commission of Zoolog-
ical Nomenclature 1999); no new species
can be described within Megalactis without
having a basis of comparison with the type
species of the genus.

In the course of this research, one of us
(A.A.) found unusual gametogenic struc-
tures in male specimens: nodes filled with
spermatic packets that have a three-dimen-
sional struc ire more voluminous than the
thickened b nd typical for gametogenic tis-
sue in members of Actiniaria. Male and
small individuals of M. comatus had blas-
tulae inferred to be of asexual origin among
the mesenteries. This is the first record of

489

asexual reproduction in a member of Acti-
nodendridae. The only female found con-
tained no gametogenic nodes or blastulae
among its mesenteries.

Materials and Methods

Specimens of the new species of Mega-
lactis were investigated alive by diving and
as preserved material; museum specimens
of other species of Megalactis and Actino-
dendron were investigated for internal mor-
phology and histology (Table 1); results
from this study are based on examination
of more than 400 museum lots and photo-
graphic documents of actinodendrids.

Animals were recorded in situ on Hi8
videotape using a CanonES6000A video
camera in an Amphibico underwater hous-
ing. Live material was collected underwater
by hand using gloves for protection against
stinging. Geographic coordinates were read
with an Eagle 12-channel GPS receiver at
the point of collection. The animals were
kept in aquaria with running seawater for
two days; no food was given. Photographs
were made in the aquarium using a Nikon
Coolpix 950 digital camera. Archived vid-
eotapes an 1 photographs are in the collec-
tion of the Division of Invertebrate Zoolo-
gy, University of Kansas Natural History
Museum (KUNHM). Specimens were re-
laxed with magnesium sulfate in seawater,
then preserved in 10% seawater formalin.
After at least two months, they were trans-
ferred to 10% freshwater formalin.

Undischarged cnidae from preserved an-
imals were examined at 1000 in squash
preparations using a light microscope
equipped with differential interference op-
tics. Squash preparations were made from
acrospheres, the oral face of the main
branches of the tentacles, the proximal,
middle, and distal column, the actinophar-
ynx, and the mesenterial filaments. Sigma
Scan Pro version 4.01.003 measurement
software was used to measure the length
and the width of undischarged capsules pro-
jected onto a Summa Sketch digitizing tab-

490

let (Summagraphics). Sampling nemato-
cysts was done following the recommen-
dations of Williams (1996).

For histology, tissue was embedded in
Paraplast, sectioned at 9 «wm, and stained
with Heidenhain’s Azan or hematoxylin and
eosin (Presnell & Schreibman 1997). Serial
sections for three-dimensional reconstruc-
tion were obtained from mesenterial struc-
tures, column, and two entire juvenile in-
dividuals. Images were obtained using a Ni-
kon Coolpix 995 digital camera connected
to an Olympus microscope through an Op-
tem eyepiece digital coupler. Serial images
were aligned manually using layers in Ado-
be Photoshop. Three-dimensional recon-
struction was done using the software Vay-
tek VoxBlast Version 3.0 Light (http://
www.vaytek.com/).

In the following discussion, as is conven-
tional in sea anemones, the proximal direc-
tion is toward the pedal disc and distal is
the opposite. Tentacles are arranged in four
cycles. Branches of the tentacles are or-
dered by how close they are to the oral disc:
a branch arising from the oral disc is con-
sidered to be of the first order; a branch that
ramifies from a branch of the first order is
of the second order, etc. (Fig. 1).

Abbreviations: CAS, California Acade-
my of Sciences, San Francisco, CA, USA;
KUNHM, University of Kansas Natural
History Museum, Lawrence, KS, USA;
NNM, Nationaal Natuurhistorisch Museum,
Leiden, The Netherlands; NMNS, National
Museum of Natural Sciences, Taichung,
Taiwan; TAUI, Zoological Museum, Tel-
Aviv University, Tel-Aviv, Israel.

Taxonomic Account

Order Actiniaria
Family Actinodendridae Haddon, 1898

Diagnosis (modified from Carlgren 1949;
see remarks below).—Limbus not well de-
fined. No marginal sphincter muscle. Fosse
absent. Up to 48 branched tentacles cycli-
cally arranged. Terminal branches of tenta-
cles with acrospheres. Two or more well de-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Ramifications in a tentacle of the first cy-
cle; view looking down a tentacle, 1.e., proximally, to
the oral disc. The arrow labeled “‘d”’ indicates the dis-
tal direction, that labeled “‘p” indicates the proximal
direction. Abbreviations: enI, endocoel of the first cy-
cle; enlII, endocoel of the second cycle; ex, exocoel;
od, oral disc; TI, first order branch; TIl-a, lateral sec-
ondary order branch; TII-b, oral face secondary order
branch; TIIJ, third order branch; TIV, fourth order
branch.

veloped siphonoglyphs. Twenty-four pairs
of mesenteries, all or almost all perfect and,
apart from the directives, fertile. Retractor
muscles diffuse, broad, band-like. Cnidom:
spirocysts, basitrichs.

Remarks.—Carlgren (1949, p. 67) indi-
cated a “‘well developed disc” and “pairs
of mesenteries up to 48” for Actinodendri-
dae. Pedal disc size varies greatly: that of
some specimens is wide, but that of others
is narrow with a limbus that is hard to rec-
ognize. None of the specimens studied had
more than 24 pairs of mesenteries. Carlgren
(1949) asserted that parietobasilar and bas-
ilar muscles are distinct in actinodendrids,
but we found them to be absent in all gen-
era of the family.

Genera.—Actinodendron Blainville,

VOLUME 117, NUMBER 4

1830, type genus; Megalactis Ehrenberg,
1834; Actinostephanus Kwietniewski,
1897.

Genus Megalactis Ehrenberg, 1834

Diagnosis (modified from Carlgren,
1949; see remarks below).—Actinodendri-
dae with ramified tentacles having second-
order branches arranged bipinnately. Last
order branches with capitate acrospheres.

Remarks.—Carlgren (1949, p. 68) stated
that Megalactis has “the oral face of the
arms [branches of the first order] free from
tentacles.” All specimens of Megalactis we
studied had two to three second-order
branches on the oral face of branches of the
first order. Carlgren (1949, p. 68) stated in
his diagnosis for Megalactis that “the ulti-
mate branches of the tentacles are simple
and pointed.”’ One of us (A.A.) found spec-
imens of Megalactis that have capitate ter-
minal tentacles.

Species.—Megalactis hemprichii Ehren-
berg, 1834, type species by monotypy, Ras
Kafil, Red Sea; Megalactis griffithsi Sa-
ville-Kent, 1893, Warrior Reef, Torres
Strait, Great Barrier Reef, 9°30’S,
143°06’E. Coordinates from Gazetteer of
Australia, 2001 (http://www.ga.gov.au/).

Megalactis hemprichii Ehrenberg, 1834

Megalactis Hemprichii Ehrenberg, 1834:
263 (original description).

Megalactis Hemprichii Ehrenberg: Milne
Edwards & Haime, 1851:11.

Actineria Hemprichii Ehrb.: Klunzinger,
1877:90-91.

Megalactis Hemprichii Ehr.: Andres, 1883:
308-309.

Megalactis Hemprichii E.: Carlgren, 1899:
14.

Megalactis Hemprichii Klunzinger: Delage
& Hérouard, 1901:539.

Megalactis hemprichii Ehrenberg,
Carlgren, 1949:68.

Megalactis hemprichi Ehrenberg: Fishel-
son, 1970:109.

1834:

491

non Megalactis hemprichii Ehrenberg,
1834: Cutress & Arneson, 1987:53—62.

Description.—Dimensions: column di-
ameter 14—26 mm distally and 14-15 mm
in the middle; pedal disc diameter 5—9 mm;
column length 23—41 mm; oral disc diam-
eter 21—23 mm; tentacles of the first cycle
45—51 mm long; tentacles of the fourth cy-
cle 10-11 mm long.

Color: Of live specimens unknown. Pre-
served specimens beige to pale yellow.

Column: Pyramidal to elongate with nar-
row pedal disc; limbus hardly recognizable
(Fig. 2A). Column smooth and mesenterial
insertions clearly visible through column in
relaxed specimens. In contracted speci-
mens, column with circumferential folds
(Fig. 2A).

Oral disc and tentacles: Oral disc narrow.
In preserved specimens, mesenterial inser-
tions on oral disc visible as dark lines; ra-
dial bumps near mouth mainly on exocoelic
intervals (Fig. 2D). Forty-eight tentacles ar-
rayed in four cycles (6 + 6 + 12 + 24).
Tentacles of first, second, and third cycles
ramified in branches of up to three orders.
Proximal secondary branches of first, sec-
ond, and third tentacle cycles short (Fig.
2B).

Branches regularly oriented. Secondary
branches pinnately disposed in one row on
each side of a branch of the first order (Fig.
2E). Up to two long and broad secondary
branches on aboral side of primary branch-
es of tentacles belonging to first, second,
and third cycles (Fig. 2E). Up to 45 sec-
ondary branches on tentacles of first and
second cycle; up to 25 secondary branches
on tentacles of third cycles; up to 11 sec-
ondary branches on tentacles of fourth cy-
cle. Branches of last order relatively long.
Large, round acrospheres.

Internal structure: Actinopharynx short
with two deep siphonoglyphs. Twenty-four
pairs of mesenteries in three cycles (6 + 6
+ 12); first two cycles usually perfect. Oral
stomata large; marginal stomata very small.
Retractor muscles diffuse and strong. Fila-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2.

Megalactis hemprichii, external morphology (TAUI 21560). A, Aboral view of entire animal. B,
Crown of tentacles, oral view of entire animal. C, Regenerated tentacles (TAUI 7812). Arrows indicate tentacles
with missing secondary branches. D, Detail of oral disc and mouth. E. First order branch. Abbreviations: b,
radial bumps on exocoelic intervals; co, column; od, oral disc; pd, pedal disc; s, siphonoglyph; TI, branch of
the first order; TI, short proximal secondary branch; TIl-a, lateral secondary order branch; TII-b, oral face

secondary order branch. Scale bars: A, B = 15 mm; C = 10 mm; D, E = 5 mm.

ments absent on mesenteries proximally.
Parietobasilar and basilar muscles not seen.
Gonochoric. The only specimen sectioned
was female (Fig. 3).

Cnidae: Basitrichs densest in acrospher-
es. Cnidom: spirocysts and basitrichs (Fig.
4). Measurements in Table 2.

Type specimen and _ locality.—Neotype
TAUI 31623, Red Sea, Gulf of Aqaba, Ei-
lat, 29°30'N, 34°55’E. Coordinates from
GEOnet Names Server of National Imagery
and Mapping Agency (http://www.nima.
mil).

Voucher specimens.—Table 1.

Megalactis comatus, new species
Figs. 4—10

Description.—Dimensions: Diameter of
column 2—38 mm distally and 5—21 mm in
the middle, of pedal disc 2—8 mm; column
length 8-26 mm; tentacles of the first cycle
9-11 mm long; tentacles of the fourth cycle
2-3 mm long; oral disc diameter 13—25
mm; tentacle crown diameter 5O—100 mm.

Color: In live specimens, oral disc and
tentacle color ranges from dark brown to
pale orange or pink. Tentacles translucent,
without pattern (Fig. 5). Oral disc with ra-

VOLUME 117, NUMBER 4

Fig. 3. Megalactis hemprichii, histology
(KUNHM 001948). Abbreviations: f, filament; m, me-
soglea; 0, ova; r, retractor. Scale bar = 1 mm.

dial rows of white spots aligned along ex-
ocoelic spaces; radial spots may spread lat-
erally onto adjacent endocoelic spaces (Fig.
5F). Insertions of mesenteries on oral disc
visible as lighter lines (Fig. 5F). Column
beige to white; distal column translucent
tinged with brown or pale orange. Female
gametogenic tissue purple and male game-
togenic tissue white (Oscar Chen, currently
at Institute of Oceanography, National Tai-
wan University, pers. comm.). Preserved

493

specimens beige, column paler than oral
disc or crown.

Column: Pyramidal to elongate with a
narrow pedal disc; limbus hardly recogniz-
able (Fig. 5C). Pedal disc and proximal col-
umn adhesive with strong ripples of ecto-
dermal tissue in preserved specimens. Cir-
cumferential folds resulting from contrac-
tion of the column between pedal region
and distal-most third of column (Fig. 5C).
Distal-most third of column thinner and
smoother than proximal column. Mesenter-
ial insertions clearly visible through col-
umn.

Oral disc and tentacles: Oral disc narrow.
Mesenterial insertions on oral disc visible
as light lines in live specimens. Radial
bumps close to mouth mainly on exocoelic
intervals.

Appearance of tentacle crown shaggy be-
cause of numerous branches not regularly
oriented (Fig. 5A, E). Forty-eight tentacles
arrayed in four cycles (6 + 6 + 12 + 24).
Tentacles of first, second, and third cycles
ramified in branches of up to four orders.
Proximal secondary branches of first, sec-
ond, and third tentacle cycles long.

Secondary branches pinnately disposed
in One row on each side of a primary branch
(Fig. 5D). On contracted tentacles, pinnate
arrangement unclear: secondary branches
appear to be arranged in two or more lateral
rows on each side of a primary branch.
Some large secondary branches occur on
aboral side of primary branches of tentacles
belonging to first, second, and third cycles.
Secondary branches variable in length. Up
to 48 secondary branches on each tentacle
of first and second cycle; up to 40 second-
ary branches on each tentacle of third cycle;
up to 12 on each tentacle of fourth cycle.
Branches of last order relatively long, ter-
minate in small round to pointed acro-
spheres.

Internal structure and histology: Actino-
pharynx short, with two deep siphono-
glyphs (two specimens had three: Fig. 6),
each connected to a pair of directive mes-
enteries. Twenty-four pairs of mesenteries

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496 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF V ASHINGTON

eeet
“See hah hh) Lh ALR ee

AA RAee

tye RS

Fig. 4. Cnidae. Basitrichs of acrospheres (A, B), and middle column (C, D, E). Spirocyst (F). Image of a
squash preparation from an acrosphere showing numerous basitrichs (G). Long basitrich (H) from filaments of
M. hemprichii (TAUI 21560). Scale bars = 10 pm:

Fig. 5. Megalactis comatus, external morphology. A, Crown of tentacles, oral view of entire animal. B,
Tentacles of the fourth cycle oriented toward substrate. C, Column in a preserved specimen. Arrow indicates
deep ripples in the pedal disc region. D, Secondary branches in bipinnate arrangement. E, Long proximal
secondary branches (arrow). EK Oral disc. Scale bars = 10 mm.

VOLUME 117, NUMBER 4

Fig. 6. Megalactis comatus, internal anatomy of a
specimen with three siphonoglyphs (KUNHM 1664):
transverse view. Abbreviations: f, filaments; g, game-
togenic tissue; s, siphonoglyphs. Scale bar = 5 mm.

in three cycles (6 + 6 + 12); first two cy-
cles usually perfect. Stomata not seen. Re-
tractor muscles diffuse and strong (Fig. 7A—
C). Filaments absent on mesenteries proxi-
mally. Parietobasilar and basilar muscles
not seen.

Gonochoric: Mesenteries in male speci-
mens have nodes filled with spermatic
packets. Each spermatic node formed
through plications of mesentery along oral-
aboral axis; node digitiform, closed on one
side of mesentery and open on the other
(Fig. 8). The only female specimen found
had ova in arrangement typical of Actini-
aria.

Cnidae: Largest and densest basitrichs in
acrospheres (Fig. 4G). Cnidom: spirocysts
and basitrichs (Fig. 4). Measurements in Ta-
blew

Type specimens and locality.—Holotype
KUNHM 1663, Pacific Ocean, Taiwan,
Henchun Peninsula, Nanwan, power plant
water intake basin, DNS 2 IN|
120°45.22’E. See Table 1 for paratype and
voucher specimens.

Etymology.—tThe epithet comatus, which
means “with long hair, shaggy” in Latin
(Brown 1978), refers to the hairy and irreg-
ular aspect of the tentacle crown in this spe-
cies.

497

Natural history.—Animals live in sym-
biosis with zooxanthellae. We found speci-
mens of M. comatus in water a few centi-
meters to 4 m deep. Each specimen of M.
comatus attaches to a coral skeleton with
its pedal disc and proximal part of the col-
umn. The color, similar to that of brown and
red algae, and shaggy aspect of the tentacle
crown make specimens difficult to find
even when abundant.

The water intake basin of the nuclear
power plant from which the type specimens
were collected was 18 years old at the time.
It was inhabited by a large number of spec-
imens of M. comatus and other species of
sea anemones tentatively identified as Bol-
oceroides memurrichi (Kwietniewski,
1898), Thalassianthus sp., and a species of
family Actiniidae. The initially large pop-
ulation of M. comatus had decreased in the
previous decade (Dr. Keryea Soong, Na-
tional Sun Yat-sen University, Kaohsiung,
Taiwan, and Oscar Chen, pers. comm.), and
has been replaced by the actinid.

One of us (A.A.) found in nature speci-
mens of M. comatus that appeared to be
undergoing transverse fission; several spec-
imens had their columns strongly constrict-
ed. One specimen, KUNHM 1667, lacks a
pedal disc, having a circular opening into
the gastrovascular cavity (Fig. 9A, B);
specimens KUNHM 1670 and KUNHM
1251 are sacciform, lack tentacles, and have
a small opening rather than an oral disc
(Fig. 9C), or have small undeveloped ten-
tacles (Fig. 9D). Specimens of M. comatus
are easy to collect, so it is not likely that
the pedal or oral disc of a specimen was
torn off during collection as can happen in
other sea anemones that attach or are deeply
buried in the substrate. Further observations
in aquaria should be made to confirm trans-
verse fission.

Some sectioned individuals of M. coma-
tus, including males and infertile individu-
als, had blastulae among their mesenteries
(Fig. 10). These larvae contained syncitial
blastoderm (solid blastula or stereoblastula
in Fautin et al. 1992) and were similar to

498 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 7. Megalactis comatus, histology. A, Retractor muscle. B, Mesenteries. C, Detail of retractor muscle.
D, Detail of mesenterial filament. E, Male gonads with the beginning of spermatic node. Abbreviations: c,
column; cg, cnido-glandular tract; ct, ciliated tract; e, endoderm; ec, ectoderm; f, filament; gl, glandular cells;
m, mesoglea; sn, spermatic node; sp, spermatic packet; zo, zooxanthellae. Scale bars: A, B, D, E = 100 pm; C
= 50 pm.

those depicted in Yanagi et al. (1999). A.A. vals (Fig. 10D, E). Juvenile stages were not
also found larvae in an individual lacking found in histological sections.
tentacles presumably because of transverse

fission. Some larvae showed incipient blas- Discussion
topores, indicating an early gastrula stage
(Fig. 10B, C). In some larvae, the outer lay- Systematics.—Type specimens of Mega-

er contained nematocysts at regular inter- lactis hemprichii have not been found

VOLUME 117, NUMBER 4

Fig. 8. Three-dimensional reconstruction of sper-
matic nodes in mesenteries of M. comatus from 20
serial slices each 9 wm thick. Abbreviations: f, fila-
ment; 1, retractor muscle; sp, spermatic packet; sn,
spermatic node. Scale bar = 0.5 mm.

(Klunzinger 1877, Fautin 2004 Hexacoral-
lians of the World: http://hercules.kgs.ku.
edu/hexacoral/anemone2/index.cfm). To
typify the genus, we designate a neotype for
M. hemprichii. Specimens of M. hemprichii
from the type locality of Ras Kafil in the
Red Sea bordering Sinai (now part of
Egypt) were unavailable and collection in
this region is not feasible. We designate as
neotype specimen TAUI 31623 from the
Gulf of Aqaba in the Red Sea, a locality
“as near as practicable from the original
type locality” (Art. 75.3.6, International
Commission of Zoological Nomenclature
1999).

Because of poor descriptions and com-
plex morphology of the tentacles, species of
Megalactis are difficult to distinguish from
each other. Ehrenberg’s (1834) description
of M. hemprichii includes a very brief Latin

499

Fig. 9.
B, Column without pedal disc KUNHM 1667. C,
Specimen without oral disc (KUNHM 1670). D, Spec-
imen with short tentacles (KUNHM 1668). Scale bars
= 5 mm.

Megalactis comatus, transverse fission. A,

description and no illustration. The only il-
lustration for M. hemprichii in Klunzinger
(1877) is based on drawings left by Ehren-
berg. Subsequent references to M. hempri-
chii are translations of the original descrip-
tion (Milne-Edwards 1857, Andres 1883,
Delage & Hérouard 1901) and a distribu-
tion record (Fishelson 1970). The specimen
identified as M. hemprichii depicted in fig-
ure 2A of Cutress & Arneson (1987) has
secondary branches not bipinnately dis-
posed, and therefore is probably a specimen
of Actinodendron.

Differences and similarities between the
species of Megalactis are presented in Table
3. Type specimens of all the species de-
scribed by Saville-Kent (1893), if they ex-
isted, have not been located (Fautin 2004
Hexacorallians of the World: http://hercules.
kgs.ku.edu/hexacoral/anemone2/index.
cfm). The photograph and description of the
color pattern of the oral disc in M. griffithsi

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

: . on 78 Ne
Lats paths S35) is

Fig. 10. Megalactis comatus, asexual larvae. A. Larva (arrow) among mesenteries. B. Late blastula (arrow).
C. Three-dimensional reconstruction of a larva from 17 serial slides each 9 pm thick. D, E. Larva with nema-
tocysts. Abbreviations: b, blastopore; c, column wall; m, mesentery; n, nematocyst. Scale bars = 0.25 mm.

Saville-Kent, 1893, can be used to identify
specimens and distinguish this species from
M. comatus.

Haddon (1898) used the shape of acros-
pheres to distinguish M. griffithsi from M.
hemprichii: clubbed for M. hemprichii and
pointed for M. griffithsi. The shape of ac-
rospheres cannot be used as a diagnostic
character in either living or preserved spec-
imens of Megalactis because it is influ-
enced by behavior and preservation. It is
common to find a museum specimen that
has acrospheres of both shapes.

Nematocysts from the acrospheres and
middle column differ in size between spec-
imens of M. comatus and M. griffithsi. The
ratio between length and width of nemato-
cysts shows a clear difference between the
two species (Fig. 11). Three specimens of

M. hemprichii from the Red Sea have a
similar gross morphology to specimens of
M. griffithsi but the nematocysts of the ac-
rospheres have size values close to those of
M. comatus. The nematocysts in the middle
column of M. comatus are larger than those
in M. hemprichii.

The number of tentacles for all species
of Megalactis is given as 10+10 for M.
hemprichii by Ehrenberg (1834), Milne-Ed-
wards (1857), Andres (1883), Delage &
Hérouard (1901), and Klunzinger (1877)
and 6+6+12 for M. griffithsi by Saville-
Kent (1893) and Haddon (1898). We agree
with Haddon (1898) that the number of ten-
tacles indicated by Ehrenberg (1834) for M.
hemprichii might be an individual peculiar-
ity. One of the three specimens of M. hem-
prichii studied (TAUI 7812) had only 41

VOLUME 117, NUMBER 4

missing data.

Table 3.—Diagnostic characters of species of Megalactis. ?

M. hemprichii

M. hemprichii
original description

neotype

M. griffithsi

M. comatus

Species/character

Regular Regular

Regular

Irregular, shaggy

Tentacle crown aspect
Secondary branches

Relatively short, usually

?, may be constricted proxi-

Elongated, usually constrict-

Relatively short, constricted

constricted proximally

Up to 45
Short

mally

proximally
Up to 35
Short

ed proximally

Up to 48

Number secondary branches
Proximal secondary branches

Distal secondary branches

Long to very long

Present

No pattern

Complex pattern of radiat-

Rows of white spots

Oral disc pattern of live specimens

ing lines and alternating
dark and white regions

Oral disc and tentacles

Light brown

Oral disc brick red and

Oral disc and tentacles pink

Color of live specimens

gray; tentacles pale pink;

column white

brown or green; column

beige

to brown; column white,

beige

501

width

Fig. 11. Length in pm of basitrichs from acros-
pheres of M. comatus (gray dots) and M. griffithsi
(black dots). In the region delimited by the rectangle
are measurements of nematocysts from the region
where acrosphere (a) meets peduncle (p); open circles
represent basitrichs type 2 (see Fig. 4B).

tentacles, all of which showed signs of re-
generation—lacking secondary branches, or
having branches not bipinnately arranged
(Fig. 2C). It is possible that M. hemprichii
has predators that feed on its tentacles. We
infer that in both previously described spe-
cies of Megalactis, the fourth cycle of ten-
tacles was overlooked, being probably con-
sidered secondary branches on the adjacent
tentacles. In situ, members of Actinoden-
dridae usually orient the tentacles of the
fourth cycle towards the substrate. All spec-
imens of actinodendrids studied, including
those belonging to Megalactis, had a typical
tentacle arrangement in multiples of six (6
ae ©) ar IID ap Dab.

Because we did not find basilar muscles
in specimens of Actinodendron plumosum
Haddon, 1898, A. glomeratum Haddon,
1898, Megalactis griffithsii Saville-Kent,
1893, and M. comatus, the position of fam-
ily Actinodendridae among Thenaria as de-
fined by Carlgren (1899, 1900, 1942, 1949)
is questionable. It is possible that basilar
muscles are reduced in size or have been
lost in the family Actinodendridae; basilar
muscles are reduced or absent in many bur-
rowing sea anemones (Carlgren 1949, Daly

502

et al. 2002). Basilar muscles are absent in
the thenarian family Aliciidae. Another ex-
planation may be that the basilar muscles
were not present in the ancestral lineage of
Actinodendridae and this family does not
belong to Thenaria.

Spermatic nodes.—We report for the first
time spermatic nodes in Actiniaria. Hyman
(1940, p. 583) stated that generally the ga-
metogenic tissues in actiniarians “occur as
thickened bands on the septa behind the
septal filaments.” Atypical organization of
gametogenic tissue is reported in the hex-
acorallian groups Actiniaria (Excoffon &
Zamponi 1999), Zoanthidea (Ryland 2000),
and Scleractinia (Harrison & Wallace
1990). The most similar structure to sper-
matic nodes in M. comatus are the ““gonadal
nodes” reported by Ryland (2000) that are
lens-shaped folds in the perfect mesenteries
of females of the zoanthid Parazoanthus
anguicomus and of a male of P. axinellae.
Spermatophores were described by Excof-
fon & Zamponi (1999) in the sea anemone
Sagartia troglodytes. The spermatic nodes
in M. comatus are not stalked like the sper-
matophores in S. troglodytes but have a
three-dimensional structure more developed
than a simple fold of the mesentery like the
‘““gonadal nodes’’ reported by Ryland
(2000). Excoffon & Zamponi (1999) re-
ported that spermatozoa in S. troglodytes
were released from spermatophores through
the stalk, the region by which the sper-
matophores are attached to the mesenteries,
and the mesogleal wall of the spermato-
phores is continuous with that of adjacent
mesentery. Thus, like spermatic nodes,
spermatophores must develop from folds of
mesenteries through evagination. We agree
with Ryland (2000) that one function of the
“gonadal nodes”’ is to increase the number
of “gonadal packets” with no increase in
length of body.

Asexual larvae.—The origin of larvae
found in the coelenteron of some sea anem-
ones is uncertain (Fautin 2002). Chia &
Rostron (1970) assumed that the larvae in-
side Actinia equina (Linnaeus, 1758) were

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

sexually produced, but Carter & Thorp
(1979) found this to be unlikely because the
phenotypes were identical between a brood
and the adult host. In fungiid corals, any
tissue fragment in the coelenteron is able to
transform into a larva (Kramarsky-Winter
& Loya 1996). Because one of us (A. A.)
found blastulae in immature and male in-
dividuals of M. comatus, they are consid-
ered to be of asexual origin.

Acknowledgments

We especially thank Dr. Keryea Soong,
National Sun Yat-sen University, Kao-
hsiung, Taiwan, for bringing the specimens
to our attention. His graduate student Oscar
Chen was A.A.’s buddy and showed the an-
imals in situ. Dr. M. Daly, and H.-R. Cha
critically read the manuscript and made
suggestions. W. N. Eschmeyer (CAS) pro-
vided advice on designating a neotype.
Thanks also go to Dr. Y. Benayahu and A.
Shlagman for providing specimens from the
collection of Zoological Museum, TAUI. N.
E. Chadwick and G. Ayalon (The Interuni-
versity Institute of Eilat, Israel) and Fan
Tung Yung and Tsai Wan Hsu (National
Taiwan University, Taiwan) collected or
provided specimens used in this study. Sug-
gestions from an anonymous reviewer im-
proved the manuscript. This research was
supported by NSF grants DEB-9521819
and DEB-9978106 in the PEET program to
D.G.E and OCE-0003970 to D.G.EF and R.
W. Buddemeier.

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504 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Saville-Kent, W. 1893. The Great Barrier Reef of Aus- Yanagi, K., S. Segawa, & K. Tsuchia. 1999. Early de-

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parameters and taxonomic relevance.—Scientia
Marina 60:339-351. Associate Editor: Stephen L. Gardiner

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(4):505—513. 2004.

A new genus and new species of crab of the family Xanthidae
MacLeay, 1838 (Crustacea: Decapoda: Brachyura) from the
southwestern Gulf of Mexico

Ana Rosa Vazquez—Bader and Adolfo Gracia

(ARVB), (AG) Instituto de Ciencias del Mar y Limnologia, UNAM, Apdo. Postal 70-305,
México, D.F, 04510, Mexico, e-mail: ana_rosav@yahoo.com.mx; gracia@mar.icmyl.unam.mx

Abstract.—A new genus and new species belonging to the Euxanthinae sub-
family, Batodaeus adanad, are described from the southwestern part of the
Gulf of Mexico. Whereas the new genus is similar to Monodaeus Guinot, 1967
in the carapace ornamentation and shape of pereopods, it differs in the structure
of the abdomen and male telson, sternoabdominal cavity, and shape and or-

namentation of the first gonopod.

Resumen.—Se describe un nuevo género y una nueva especie perteneciente
a la subfamilia Euxanthinae, para el suroeste del Golfo de México. Este género
nuevo es similar a Monodaeus Guinot, 1967, en la ornamentacion del capar-
azon y forma de los pereopodos; sin embargo, difiere de éste en la forma y
tamano del abdomen, telson y cavidad esterno-abdominal, asi como en la es-

tructura de los apéndices sexuales.

In the course of deep-water biodiversity
surveys in the Cayo Arcas and west side of
Triangulos in the southwestern part of the
Gulf of Mexico, specimens of an unusual
species of xanthid crab were obtained. Al-
though superficially similar to species of
Monodaeus Guinot, 1967, they possess sev-
eral atypical features that suggest other-
wise. They are here described as a new ge-
nus and new species.

Material and Methods

Specimens were collected in 1998 during
surveys investigating the marine fauna in
the deep southwestern Gulf of Mexico,
cruise BATO (Biota de los Arrecifes, de la
Plataforma y Talud continental en el no-
roeste del Banco de Campeche), carried out
on board the R/V Justo Sierra by the Insti-
tuto de Ciencias del Mar y Limnologia,
UNAM. The samples were caught using a
semicommercial otter trawl.

The material was deposited in the refer-
ence collection of the Instituto de Biologia,

UNAM (CNCR). Measurements listed are
in millimeters (mm): total carapace length
(CL) and carapace width (CW).

Batodaeus, new genus

Diagnosis.—Carapace subhexagonal,
broader than long; dorsal surface convex
and granulated. Regions in male well de-
marcated especially in the anterior half,
front strongly deflexed; inner orbital teeth
conspicuous; thoracic sternum relatively
narrow. Anterolateral margins armed with 4
teeth (excluding outer orbital tooth), sube-
qual in size to posterolateral margins; pos-
terior margin of epistome triangular, median
part depressed, with distinct median fissure,
and a pair of shallow but clearly visible lat-
eral notches; preorbital and postorbital
lobes conspicuous, granulated. Incomplete
endostomial ridges present. Basal antennal
segment long, subcylindrical, almost touch-
ing front, not filling space between front
and inner orbital teeth; a small gap between
basal antennal segment and suborbital mar-

506

gin. Third maxillipeds not filling buccal
cavity; merus with deep, rounded depres-
sion near mesial margin. A longitudinal tu-
berculated ridge just below suture separat-
ing subhepatic and subterygostomian re-
gions. Pereopods 2-5 long, slender, with
conspicuous spines on upper margin of
merus and carpus. Sternoabdominal cavity
relatively narrow and deep; thoracic sternite
4 with median part slightly raised, with
short longitudinal furrow. Abdomen long,
completely covering sternoabdominal cavi-
ty. Second abdominal segment in each sex
not reaching coxae of fifth pereopod, leav-
ing a small portion of sternite 8 visible.
Sternite 7 covering part of penis groove on
sternite 8. First gonopod slender, slightly in-
curved; a row of spines on mesial margin.
Second gonopod very short, sigmoid, ter-
minal process curved with short setae on
distal end.

Type species.—Batodaeus adanad, new
species, by present designation.

Gender.—Masculine.

Etymology.—The name is a combination
of Bato, the name of the cruise during
which it was collected, and “‘daeus”’ to in-
dicate its similarity with the genus Mono-
daeus.

Remarks.—TYhe taxonomic position of
Batodaeus is uncertain. Although the spec-
imens studied here share some characteris-
tics with the subfamily Actaeinae in having
sternite 4 with a longitudinal furrow and the
male abdomen with locking mechanism on
sternite 5, they also share some features
with the Xanthinae in having the carapace
with four teeth and with thoracic sternite 4
being rather long (see Serene, 1984). In
fact, Batodaeus more closely resembles the
euxanthinid genus Monodaeus Guinot,
1967 in the following characters: regions of
the carapace well demarcated; hepatic re-
gion inflated; presence of four anterolateral
teeth; surface of carapace with several
strong granules; basal antennal segment just
touching the front; anterior border of buccal
cavity with a conspicuous crest; third max-
illipeds not completely closing buccal cav-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ity and merus with prominent distolateral
angle; and thoracic sternite 4 with a median
longitudinal furrow. However, the carapace
of Batodaeus is more convex and the re-
gions are less well demarcated and with
acute spines, particularly on the hepatic re-
gion; the posterolateral margins are sube-
qual in length to the anterolateral; the front
is more advanced, with a deep sulcus; the
preorbital lobe is conspicuous; the sternoab-
dominal cavity is relatively deeper and nar-
rower; thoracic sternite 4 has a shallow but
distinct longitudinal median furrow; the ab-
domen is relatively narrow and longer; and
the first male gonopod is long, slender and
slightly incurved with few spines, not se-
tose or stout as in Monodaeus.

The bathymetric and geographic distri-
bution of these two genera is also different.
Monodaeus (type species Xantho couchii
Bell, 1851) at present contains eight spe-
cies: M. arnaudi Guinot & Macpherson,
1988; M. couchii (Bell, 1851); M. cristu-
latus Guinot & Macpherson, 1988; M. guin-
otae Forest, 1972; M. pettersoni Garth,
1985; M. rectifrons (Crosnier, 1967); M.
rouxi (Capart, 1851); and M. tuberculidens
(Rathbun, 1911). They all occur in the
Western Indian Ocean, the Eastern Atlantic,
the Mediterranean Sea and the Eastern Pa-
cific, from 20 to 500 meters. Batodaeus
adanad, however, was collected in the
Western Atlantic at depths from 160 to
250 m.

In comparison with Medaeops Guinot,
1967 (only known from the Indo-West Pa-
cific), Batodaeus has a carapace which has
the regions less inflated, the merus of third
maxilliped with a prominent distolateral an-
gle; the pereopods are more slender and
longer, the thoracic sternum is not flat; and
the first male gonopod lacks setae. It differs
from the superficially similar Indo-West Pa-
cific genus Alainodaeus Davie, 1992 in
having a carapace which is subhexagonal,
not ovoid; a straight frontal margin; a
rounded telson (triangular in Alainodaeus)
and an almost straight first male gonopod.

Batodaeus, for the moment, is tentatively

VOLUME 117, NUMBER 4

placed in the Euxanthinae, as it seems to fit
there considering the primary character of
the subfamily stated by Seréne (1984): the
form of the anterolateral margin which has
the anterior part gradually sloping down-
wards via subhepatic region to meet the in-
fraorbital margin. The similarity to Mono-
daeus confirms that it can be placed as Eux-
anthinae, even though the subfamily is now
poorly defined.

Batodaeus adanad, new species
Figs. 1-4

Material examined.—Holotype: 1 ¢
CNCR (21023), 16.6 mm X 23.2 mm; Sta.
9, 22°17.19'N, 91°43.08’W (Banco de
Campeche, off Cayo Arenas), 251 m, 23
May 1999. Allotype: 1 2 CNCR (22509),
11.8 mm X 16.6 mm, Sta. 9, 22°17.19'N,
91°43.08’W (Banco de Campeche, off Cayo
Arenas), 251 m, 23 May 1999. Paratype: 1
2 CNCR (21024), 12.1 mm X 17.6 mm,
off Sta. 8, 22°13.72’N, 91°46.64’W (Banco
de Campeche, off Cayo Arenas), 162 m, 23
May 1999.

Description.—Carapace (Fig. 1A) sub-
hexagonal, about 1.4 to 1.5 times broader
than long; dorsal surface convex, granulat-
ed, granules coarse, more abundant on an-
terolateral and posterolateral margins, with
sparse short setae on frontal, hepatic and
protogastric regions. Regions in male well
demarcated, especially in the anterior half;
orbital region with small, acute spines; he-
patic region inflated, with 3—4 rows of
spines; a depression between cardiac and
gastric region; meso and urogastric regions
less granulated; front straight, strongly de-
flexed; margin dentate, at most %4 as long
as CW, separated from protogastric region
by a long transverse furrow. Deep notch be-
tween frontal and preorbital lobes, the latter
strongly granulated. Anterolateral margins
armed with 4 teeth (excluding the outer or-
bital), first small, with margins granulated,
second and third longest, subequal in size,
spinose and directed anteriorly, fourth big-
ger than first, with borders and base gran-

507

ulated. Posterolateral and posterior margins
of carapace almost straight, granulated.
Pterygostomian region (Figs. 2A, 4C)
densely granulated, with a longitudinal, spi-
nose crest just below suture that divides
subhepatic and pterygostomian regions;
pterygostomian ridges present, marked with
small tubercles.

Orbits % as wide as front, separated from
it by a deep, long notch, borders conspic-
uously dentate; 2 large sutures on supraor-
bital region; preorbital and postorbital lobes
dentate. An acute granulated tooth on infra-
orbital angle. Eyestalks completely fitting
in orbits when retracted, with 4 sharp spi-
nules and stiff setae.

Antennules (Figs. 2A, 4C) with basal
segment considerably inflated, and with a
longitudinal crest of small granules; penul-
timate and ultimate segments slender.

Basal antennal segment subcylindrical,
with 4 sharp spinules; second to fourth seg-
ments mobile, longer than broad; flagellum
long.

Ischium of third maxilliped (Fig. 2B)
longer than broad; outer surface with me-
dian longitudinal furrow. Merus subquad-
rate, outer surface coarsely granulated, me-
sial margin dentate, setose; distolateral an-
gle ending in a semiacute lobe, directed an-
teriorly. Palp marginally setose. Exopod
reaching to tip of distolateral angle of mer-
us.

Left cheliped shorter than right and more
spinose (Figs. 4A—B); merus granulose, up-
per and lower margins delimited by a row
of strong, acute, inward spines; outer surface
of carpus spinose, inner margin armed prox-
imally with strong acute spine curving in-
wards, junction between carpus and chelae
fringed with setae, inner surface densely
granulated. Palm of long chela with 3 upper
rows of acute spines directed inwards, di-
minishing in number and size towards lower
margin, inner surface slightly punctate. Dac-
tylus about as long as palm; fingers leaving
small gap when closed, each terminating in
inwardly curved corneous claw; movable

508 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Batodaeus adanad, new genus, new species. A. Holotype male 16.6 mm X 23.2 mm (CNCR 21023):
B, Allotype female 11.8 mm X 16.6 mm (CNCR 22509), dorsal view.

VOLUME 117, NUMBER 4

Fig. 2.
regions; B, third left maxilliped, outer view; C, sternoabdominal cavity. D, allotype female, sternoabdominal
cavity. Abbreviations: g2, gonopod 2; s1—8, sternites 1—8.

part with upper margin armed with small
acute spines, cutting edges with blunt teeth.

Pereopods 2—5, slender, subequal in
length; pereopod 4 slightly long, and pereo-
pod 2 slightly short; all segments with lat-
eral and mesial faces spinose, covered by
dense, thin setae. Dactylus slightly longer
than propodus, terminating in corneous
claw. Propodus with long thin setae on up-
per border, outer surface punctate. Upper
border of carpus with 4—6 small spines.
Merus with 12 spines on upper border,
which diminish in size proximally, directed
anteriorly.

Thoracic sternum in male (Fig. 2C) rel-
atively narrow, densely granulated; sternal
sutures 1—2 indistinct, 2—3 complete, 3—4
incomplete and confined to lateral regions;

509

Batodaeus adanad. A—C, holotype male. A, antennal, pterygostomian, suborbital, and epistomal

4—5 and 5—6 interrupted medially; 6—7 and
7-8 complete. Sternite 4 with slightly raised
median part, with short longitudinal furrow.
Locking mechanism on sternite 5 just be-
low suture 4—5. Sternoabdominal cavity
deep and relatively narrow.

Male abdomen (Fig. 3A) with short mar-
ginal setae on segments 1—6 and telson.
First segment long, slender. Second as long
as first, broadest, not reaching coxa of fifth
pereopod, with small portion of sternite 8
visible (Fig. 3B). Third to fifth segments
fused, punctate, longer than broad. Sixth
segment as long as broad. Posterior margin
of telson rounded. Male sexual openings
coxal.

First gonopod (Fig. 3C, D) long, reach-
ing beyond suture separating sternites 4 and

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3.
5 and sternite 8; C, gonopod 1; D, tip of gonopod 1; E, gonopod 2. E allotype female, abdominal segments.
Abbreviations: al—2, abdominal segments | and 2; cx 5, coxa 5; ep7, episternite 7.

5, when in situ, slender and with distal part
slightly incurved. Second gonopod (Fig.
3E) very short, curved, tip sharp, recurved.

Females with regions of carapace less de-
marcated (Fig. 1B); front straight, less de-
flexed; right cheliped longer than left, palm
with less conspicuous spines, pereopods
more setose, without spines on upper border
of merus, carpus with spines more acute.
Thoracic sternum (Fig. 2D) with longitu-
dinal furrow on sternite 4 less marked; ab-
dominal cavity less deep and broad; in lon-
gest specimen, locking mechanism on ster-
nite 5 not visible. Abdomen (Fig. 3F), with
first and second segments as in male, leav-
ing visible a small portion of sternite 8; seg-
ments 3—6 free and subequal in size; pos-
terior part of telson rounded. Pleopods long,

Batodaeus adanad. A—E, holotype male. A, abdominal segments; B, abdominal segments 1—3, coxa

slender, extending past edge of telson; gon-
opores small, ovate.

Color in life.—Cream, with tip of chelae
fingers dark.

Etymology.—This species name is
formed from an arbitrary combination of '
the two first letters of each of our sons’
names: Adolfo, Andrés, and Adrian, and is
used as a noun in apposition.

Distribution.—Western Atlantic; south-
western Gulf of Mexico; Banco de Cam-
peche.

Remarks.—The shape and ornamentation
of carapace and pereopods of B. adanad are
superficially similar to the species of Mon-
odaeus, notably M. rouxi. Also, the ptery-
gostomial region is densely granulated and
the incomplete endostomial ridges of B.

VOLUME 117, NUMBER 4 511

Fig. 4. Batodaeus adanad, holotype male. A, right chela; B, left chela; C, ventral view of anterior part of
carapace.

SID

adanad are similar to those seen in M. tub-
erculidens and M. couchii. The morphology
of the merus of the third maxilliped of B.
adanad also resembles those of M. guinotae
and M. tuberculidens.

However, Batodaeus adanad is easily
separated from all Monodaeus species in
that the former has the posterolateral mar-
gin almost as long as the anterolateral, the
chelipeds are less stout, spinose, and are
covered by strong tubercles; the sternoab-
dominal cavity is deeper and with a less
marked longitudinal furrow on sternite 4.
The abdomen of Batodaeus is long, com-
pletely covering the sternoabdominal cavi-
ty, whereas in Monodaeus species the ab-
domen does not completely cover the ster-
noabdominal cavity, leaving a longitudinal
furrow on sternite 4 exposed; the pereopods
2—5 without a granulated crest on the su-
perior border of merus; and the telson of
male is rounded. In addition, the morphol-
ogy of the first gonopod in B. adanad dif-
fers from that in any known Monodaeus
species.

The new species differs from Medaeops
edwardsi Guinot, 1967, M. neglectus
(Balss, 1922), and M. granulous (Haswell,
1882) in that all these species have a less
convex carapace, with the regions hardly
projecting; their pereopods are shorter and
broader; and the fingers of their chelipeds
are granulated. The first gonopod, thoracic
sternum, and sternoabdominal cavity, too,
are also different in morphology.

Batodaeus adanad can be easily separat-
ed from Alainodaeus akiaki Davie, 1992
and A. rimatara Davie, 1992 in that those
species have a carapace that is transversally
ovoid; the front is less deflexed; the cheli-
peds are more robust; the first male gono-
pod is stout with slightly twisted tip; and
the second male gonopod is moderately
longer.

Acknowledgments

Dr. Rafael Lemaitre and Dr. Janice Clark
are greatly appreciated for their help and

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

loan of specimens during our visit to the
Smithsonian Institution. Special thanks are
given to Dr. Michel Hendrickx for his kind
review and valuable suggestions on the
manuscript. The comments of Dr. P. Ng
greatly improved the quality of the manu-
script. We thank the crew and scientific
staff of R/V Justo Sierra for field work dur-
ing cruise BATO. We also thank Ana Elena
Viniegra for drawings and Ana Isabel Bie-
ler for taking the photographs.

Literature Cited

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Expédition Océanographique belge dans les
eaux cOtiéres africaines de 1’Atlantique Sud
(1948—1949)—Bruxelles 3(1):11—205.

Couch, R. Q. 1851. Notice of a Crustacean, New to
Cornwall.—Transactions of Penzance Natural
History Society: 13-14.

Crosnier, A. 1967. Remarques sur quelques Crustacés
Décapodes benthiques ouest-africains. Descrip-
tion de Heteropanope acanthocarpus et Me-
daeus rectifrons sp. noy.—Bulletin du Muséum
national d’Histoire naturelle, Paris, (2), 39(2):
320-344.

Davie, P. J. EF 1992. Deepwater Xanthid crabs from
French Polynesia (Crustacea, Decapoda, Xan-
thoidea).—Bulletin du Muséum national
d’Histoire naturelle, Paris, 4°. sér., 14, section
A, n° 2:501—561.

Drach, P., & J. Forest. 1953. Description et répartition
des Xantho des mers d’ Europe.—Archives de
Zoologie expérimentale et générale, 90(1):1—35.

Forest, J. 1972. Une espéce nouvelle de Xanthidae des
eaux bathyales de Méditerrane: Monodaeus
guinotae sp. nov.—Thalassia Jugoslavica 8(1):
63-69.

Garth, J. S. 1985. On a small collection of brachyuran
Crustacea from Easter Island obtained by the
Scripps Institution of Oceanography Downwind
Expedition of 1958.—Occasional Papers of the
Allan Hancock Foundation (3):1—12.

Guinot, D. 1967. Recherches préliminaires sur les
groupements naturels chez les Crustacés Brach-
youres. II. Les anciens genres Micropanope
Stimpson et Medaeus Dana.—Bulletin du Mu-
séum national d’ Histoire naturelle, Paris, 39(2):
354-374.

, & E. Macpherson. 1988. Remarques sur le

genre Monodaeus Guinot, 1967, avec la de-

scription de deux espéces nouvelles (Crustacea

Decapoda Brachyura).—Bulletin du Muséum

national d’ Histotoire naturelle, Paris 4°. sér., 10:

731-757.

VOLUME 117, NUMBER 4

MacLeay, W. S. 1838. On the brachyurous decapod

Crustacea brought from the Cape by Dr.Smith.
Tn Wlustrations of the Annulosa of South Africa:
being a portion of the objects of natural history
chiefly collected during an expedition into the
interior of South Africa, under the direction of
Dr. Andrew Smith, in the years 1834, 1835, and
1836; fitted out by “The Cape of Good Hope
Association for Exploring Central Africa’’.
London: 53-71.

513

Seréne, R. 1984. Crustacés Décapodes Brachyoures de

l’‘Océan Indien Occidental et de la Mer Rouge.
Xanthoidea: Xanthidae et Trapeziidae. Avec un
Addendum par Crosnier (A):Carpillidae et
Menippidae.—Faune Tropicale. Office de la Re-
cherche Scientifique et Technique Outre-Mer.
Paris, 24:1—400.

Associate Editor: Christopher Boyko

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(4):514—522. 2004.

A new anchialine shrimp of the genus Procaris (Crustacea:
Decapoda: Procarididae) from the Yucatan Peninsula

Richard v. Sternberg and Marilyn Schotte

(R.v.S) NCBI--GenBank, Building 45, Room 6An, 18D-30, National Institutes of Health,
Bethesda, Maryland 20892-6510 U.S.A. e-mail: sternber@ncbi.nlm.nih.gov
(MS) Department of Zoology, National Museum of Natural History, Smithsonian Institution,
Washington, D.C., 20013-7012 U.S.A. e-mail: Schottem@si.edu

Abstract.—A fourth species of the anchialine shrimp genus Procaris is de-
scribed from Cozumel Island, Quintana Roo, México. The combination of char-
acter states observed for the abdomen, antennal scale/stylocerite, second an-
tennular segment, carapace, eyes, rostrum, and telson is unique in the genus.
The new species appears to be morphologically most closely related to P.
ascensionis from Ascension Island. Cladistic analysis of differentiating char-
acter states supports a sister group relationship between P. ascensionis and the
Mexican species, in two out of three most parsimonious hypotheses. In addi-
tion, the Bermudan P. chacei and Hawaiian P. hawaiiana are positioned as
sister taxa in all minimal length trees. While the discovery of a new Procaris
species adds to our biogeographical knowledge of the genus, it has pointed to
the possibility that the Atlantic taxa may be a paraphyletic assemblage.

Shrimps of the family Procarididae are
restricted to anchialine habitats, and occupy
an unclear position within the Decapoda
relative to the Caridea (Christoffersen 1988,
1990; Felgenhauer & Abele 1983; Kensley
& Williams 1986; Schram 1986). The Pro-
carididae contains two genera, Procaris and
Vetericaris Kensley & Williams, 1986. Pro-
caris has perhaps the most interesting dis-
tribution of any anchialine decapod: P. as-
censionis Chace & Manning, 1972 is re-
stricted to Ascension Island in the mid-
south Atlantic, P. chacei Hart & Manning,
1986 is endemic to Bermuda, and P. ha-
watiana Holthuis, 1973 is found on the Ha-
waiian archipelago. [A photograph of an
undescribed “*Procarid sp.’ from Christmas
Island in the Indian Ocean has been pub-
lished (Jones & Morgan 2002), although
the habitus of the pictured specimen looks
more atyid than procaridid.]

What is even more remarkable is the con-
servative morphology of Procaris species,
considering the disjunct biogeography of

the taxa, as the three species differ in only
a few characters (Hart & Manning 1986).
Vetericaris is monotypic with the Hawaiian
V. chaceorum Kensley & Williams, 1986
separated from any Procaris species by a
plethora of character states. Despite the dis-
tinctiveness of Procaris and Vetericaris, the
monophyly of the family has not been ques-
tioned. A recently described family for a
genus of abyssal shrimp, the Galatheacari-
didae Vereshchaka, 1997, overlaps with the
Procarididae in several key (albeit plesio-
morphic) character states, indicating that
the hypothesized connection between an-
chialine and abyssal caridean taxa of Hart
et al. (1985) may not be entirely without
merit. Interrelationships aside, an open
question is how many additional anchialine
and submerged caverniculous carideans
await discovery that could, potentially,
complete the known biogeographical gaps.

Here we describe a fourth species of Pro-
caris, from the Yucatan Peninsula. The dis-
covery of this new species adds consider-

VOLUME 117, NUMBER 4

Sea SE IE as era
qa SS

515

Sree

SERIE EES yy

a
,

Fig. 1.
of telson.

ably to our biogeographic knowledge of the
genus. The new Procaris material was col-
lected by Drs. Dennis Williams and Jeff
Bozanic who during the years 1988, 1989,
and 1995 collected them from the cenotes
of Quintana Roo, México. CL numbers re-
fer to carapace length; USNM numbers de-
note catalog numbers in the National Mu-
seum of Natural History, Smithsonian In-
stitution.

WA
Siac
oo

eos S SSeS

Procaris mexicana, n. sp. A, habitus, lateral view; B, anterior region; C, telson and uropods; D, apex

Procarididae Chace & Manning, 1972
Procaris mexicana, new species
Figs. 1-3, Table 1

Procaris sp. Kensley, 1988:688.

Material.—Holotype (USNM_ 1068789):
México, Cueva Quebrada, Chankanaab Park,
Cozumel, Quintana Roo, 25 September
1987, coll. Dennis Williams, CL 8 mm.
Paratypes: USNM 1068790, 1 specimen, CL

516

6.5 mm, same locality as holotype, coll.
Dennis Williams, 23 Sep 1987; USNM
1068791, 3 specimens (1 damaged), CL 5.1
mm, 5.5 mm, and 5.9 mm, Cueva Quebrada,
depth of 25-30 feet, coll. Jeff Bozanic, 5
April 1988; USNM 1068792, 4 specimens,
all CL 6 mm, Cueva Quebrada, coll. Dennis
Williams, Feb. 1993; USNM 1068793, 1
specimen, CL 8 mm, Lagoon Cave, Cozu-
mel, Quintana Roo, México, coll. Jeff Boz-
anic, 3 Apr. 1988.

Description.—Integument fragile and
thin. Rostrum acutely triangular and lacking
teeth, only reaching medial concavity of
eyes. Carapace devoid of spines; anterior
margin distinctly convex and slightly emar-
ginate below distinct cervical sulcus; prom-
inent anteroventral sulcus positioned parallel
to ventral margin, and meeting ventral end
of cervical sulcus; posterodorsal margin
markedly concave.

Eyestalk produced into two lobes, the me-
dial lobe sharply triangular and extending
beyond the more bluntly triangular lateral
lobe; eye lacking facets and with irregular
mass of pigment.

Antennular peduncle does not reach distal
one-third of antennal scale, broad; stylocer-
ite tapering distally to acute apex, almost
reaching distal margin of second antennular
article; segments subequal in length; anterior
margin of basal article with distinct V-
shaped dorsomedial cleft.

Antennal scale lacking distolateral tooth,
distal margin convex, length approximately
2.5 times the width; distal margin of scale
reached by antennal peduncle.

Mandible pronouncedly developed, with
three-segmented palp, molar and incisor pro-
cesses forming one piece; incisor process
subtrapezoidal, lacking distinct marginal
teeth except for the two angular regions,
scooplike. Paragnath sinuous, surrounding
incompletely mandibular bases, distal end
pointed, broadest around midlength. Endites
of first maxilla well-developed, broad; palp
simple. Second maxilla with two endites,
distal endite with deep incision, palp pro-
nounced and broader proximally, tapering

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

slightly distally, scaphognathite small in
comparison to the endites and palp. Maxil-
liped 1 with near tongue-shaped endite,
well-developed palp; long, simple epipod;
caridean lobe prominent. Maxilliped 2 en-
dopod with seven segments of roughly sim-
ilar width throughout; exopod long, strap-
like; epipod simple, reduced. Maxilliped 3
with seven-segmented endopod, distal half
of merus broader than all other parts of the
appendage; exopod long, subequal to endo-
pod length; epipod simple, small.

Pereiopods 1—5 similar in organization,
flexor margins lined with simple setae; dac-
tyli approximately 0.12—0.13 times length of
propodi, with strong, curved spines. All five
pereiopod pairs with straplike exopod; pe-
reiopods 1—4 with distinct simple epipod,
and pleurobranch and setobranch; pereiopod
5 lacking epipod, pleurobranch, and seto-
branch.

Third abdominal somite with dorsal cap
not reaching middle of fourth somite; pos-
teroventral margin of the six anterior somites
broadly rounded. Abdominal sternites 1—5
with median tubercle between coxae of ple-
opods; sternite 6 with bulbous tubercle pos-
teriorly directed between uropod bases. Tel-
son approximately 1.4 times length of so-
mite 6, not including posterior spines, armed
with two pairs of dorsal spines; posterior
margin armed with four pairs of spines, lat-
eral spines shortest, two mesial pairs roughly
half the length of sublateral spines.

All pleopods similar in organization; en-
dopods short and weakly developed; appen-
dices internae and masculinae absent from
all pleopods.

Distribution.—Known only from anchia-
line habitats of Cozumel, Quintana Roo, Yu-
catan Peninsula, México.

Remarks.—All Procaris species are re-
markably similar in morphology, differing
slightly but specifically in a set of characters
(Table 1; Hart & Manning 1986). This is
significant given the immense distances sep-
arating all four taxa, especially P. hawaiiana
vis-a-vis the three Atlantic species. On the
basis of biogeography, one might expect the

VOLUME 117, NUMBER 4 517

Fig. 2. Procaris mexicana: A, pleopod 4; B and C, mandible; D, second maxilliped; E, paragnaths; E first
maxilliped; G, pleopod 1; H, first maxilla; I, second maxilla; J, third maxilliped; K, pleopod 3; L, pleopod 2;
M, pleopod 5.

518 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Procaris mexicana: A, pleopod 1; B, same, dactyl; C, pereopod 2; D, same, dactyl; E, pereopod 3;
F same, dactyl; G, pereopod 4; H, same, dactyl; I, pereopod 5; J, same, dactyl.

519

VOLUME 117, NUMBER 4

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520 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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characters unordered (the other tree identical to A).

VOLUME 117, NUMBER 4

three Atlantic species to form a clade, with
the Indo-Pacific P. hawaiiana as the sister
group of the lineage. However, a comparison
of the character states presented in Table 1
affords no clear-cut separation between At-
lantic and Pacific congeners. Each Procaris
species instead appears to be a mosaic of
character states found in the other taxa; spe-
cies differences are due then to specific char-
acter state combinations as opposed to the
presence of apomorphies. To test the possi-
bility that P. ascensionis, P. chacei, and P.
mexicana may be more closely related to
each other than to P. hawaiiana, a data ma-
trix was prepared for parsimony analysis
(Table 2), and Vetericaris was used as the
outgroup for character state polarization (Ta-
ble 1). The purpose of the cladistic test was
twofold: to identify a parsimonious hierar-
chy of Procaris taxa, and to compare this
hierarchy with biogeography.

When characters 1—7 were treated as or-
dered transformation series, one tree was ob-
tained by Exhaustive Search using PAUP
3.1 software (Swofford 1993), with a length
of 18 steps, consistency index (CI) value of
0.833, and a retention index (RI) .umber of
0.571 (Fig. 4A). This first hypotl esis indi-
cates that P. ascensionis and P. :nexicana
are sister species, with P. chacei and P. ha-
waiiana forming a species pair. Placing the
cladogram into the context of time and
space, the split between Atlantic and Pacific
Procaris species would have occurred after
the emergence of two Atlantic clades: P. as-
censionis and P. mexicana on the one hand,
and the proto-P. chacei/P. hawaiiana ances-
tor.

A second exhaustive search was_per-
formed though this time all characters were
parameterized as unordered series. Two trees
most parsimonious were found with lengths
of 17 steps, CI = 0.882, and RI = 0.667.
The topolo; y of one of the cladograms is
identical in structure to the one in Fig. 4A.
The second hypothesis is also a resolved hi-
erarchy, though with P. ascensionis branch-
ing off first, followed by P. mexicana, and

521

Table 2.—Data matrix used in the parsimony anal-
ysis. See Table 1 for explanation of character states.

Character 12345678
Vetericaris 00000000
P. ascensionis 10012223
P. chacei 21101222
P. hawatiana 21101311
P. mexicana 10112111

with P. chacei and P. hawaiiana positioned
as sister taxa (Fig. 4B).

Cladistic analysis of Procaris interrela-
tionships indicates three things. First, P.
chacei and P. hawaiiana are more closely
related to each other on morphological
grounds than either is to any other Procaris
species. Second, relationships between P.
ascensionis and P. mexicana are ambiguous.
Parsimony searches conducted with ordered
and unordered characters support a sister
group relationship between the two (Fig.
4A). Yet the hypothesis that P. ascensionis
is basal to the remaining Procaris species
(Fig. 4B) cannot be dismissed. Finally, the
Atlantic species appear not to form a clade;
i.e., they are a paraphyletic assemblage mi-
nus the inclusion of P. hawaiiana.

One seri pus caveat of the parsimony study
is the paucity of characters (eight) relative
to the number of taxa (five). This reflects the
extremely conservative morphology of Pro-
caris species. Another caveat is the coding
of character states (Table 1). Character states
were coded to maximize hierarchical reso-
lution given a limited number of characters.
For instance, the rostrum character was di-
vided into three character states: not reach-
ing medial concavity of eyes (plesiomorph-
ic); reaching medial concavity (apomorph-
ic); overreaching medial concavity (also
apomorphic). The way this character was
coded for Procaris species, P. chacei and P.
hawaiiana have the same state. Yet the ros-
trum only reaches the median lobe in P.
chacei although it overreaches the eyes in P.
hawaiiana. The same critique applies to
character 2. Nevertheless, if characters are
recoded to reflect all the differences seen,

522

(SRE DU EO

P. ascensionis

3(1)

P. mexicana P. hawatiana

Fig. 5.
ings. Numbers to the right of a box denotes derived
character states supporting a particular set of taxa.

Venn diagram of apomorphy-based group-

the tree obtained is identical to that shown
in Fig. 4A (unpublished results).

Figure 5 shows a Venn diagram of apo-
morphy-based relationships in Procaris, un-
derscoring the polythetic nature of species
differences.

Hart & Manning (1986) suggested that the
remarkable similarity of Procaris species
may be explained by the reduction of vari-
ability by natural selection. The “reduced
variability’ hypothesis appears rather weak
considering that anchialine caridean taxa oc-
curring with Procaris often exhibit consid-
erable variability, morphs, and species-spe-
cific apomorphies (e.g., Kensley & Williams
1986, Smith & Williams 1981). It may be
that the distribution of Procaris is much
more extensive than currently known, with
gene flow over great distances occurring via
semi-continuous populations distributed
among shallow submerged ‘“‘crevicular”’
habitats (Hart et al. 1985, Maciolek 1983).

Acknowledgments

We are most grateful to Drs. Dennis Wil-
liams and Jeff Bozanic who collected the
new Procaris material, and to Drs. Charles
Fransen and Mark Siddall for their com-
ments on an earlier draft of the manuscript.

Literature Cited

Chace, F A., Jr, and R. B. Manning. 1972. Two new
caridean shrimps, one representing a new family,
from marine pools on Ascension Island (Crusta-
cea: Decapoda: Natantia).—Smithsonian Contri-
butions to Zoology 131:1-18.

Christoffersen, M. L. 1988. Phylogenetic systematics of

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

the Eucarida (Crustacea Malacostraca).—Revista

Brasileira de Zoologia 5(2):325-351.

. 1990. A new superfamily classification of the
Caridea (Crustacea: Pleocyemata) based on phy-
logenetic pattern.—Zeitschrift fiir Zoologische
und Systematik Evolutionsforschung 28(2):94—
106.

Felgenhauer, B. E., and L. G. Abele. 1983. Phylogenetic
relationships among shrimp-like decapods. In: E
R. Schram, ed., Crustacean issues 1. Crustacean
phylogeny, pp. 291-311. A. A. Balkema, Rotter
dam. Pp. 1-372.

Hart, C. W., Jr, and R. B. Manning. 1986. Two new
shrimps (Procarididae and Agostocarididae, new
family) from marine caves of the western north
Atlantic—Journal of Crustacean Biology 6(3):
408-416.

5 , and T: M. Iliffe. 1985. The fauna of
Atlantic marine caves: evidence of dispersal by
sea floor spreading while maintaining ties to deep
waters.—Proceedings of the Biological Society
of Washington 98:288-292.

Holthuis, L. B. 1973. Caridean shrimps found in land-
locked saltwater pools at four Indo-West Pacific
localities (Sinai Peninsula, Funafuti Atoll, Maui,
and Hawaiian Islands), with the description of
one new genus and four new species.—Zoolo-
gische Verhandelingen 128:1- 48.

Jones, D. S., and G. J. Morgan. 2002. A Field Guide to
Crustaceans of Australian Waters. Western Aus-
tralian Museum. Reed New Holland, Sydney,
Australia, 224 pp.

Kensley, B. 1988. New species and records of cave
shrimps from the Yucatan Peninsula (Decapoda:
Agostocarididae and Hippolytidae)—Journal of
Crustacean Biology 8(4):688—699.

, and D. Williams. 1986. New shrimps (families
Procarididae and Atyidae) from a submerged
lava tube on Hawaii.—Journal of Crustacean Bi-
ology 6(3):417—437.

Maciolek, J. A. 1983. Distribution and biology of Indo-
Pacific insular hypogeal shrimps.—Bulletin of
Marine Science 33(3):606—618.

Schram, FE 1986. Crustacea. Oxford University Press,
Oxford.

Smith, M. J., and W. D. Williams. 1981. The occurrence
of Antecaridina lauensis (Edmondosn) (Crusta-
cea, Decapoda, Atyidae) in the Solomon Islands:
an intriguing biogeographical problem.—Hybro-
biologia 85:49—58.

Swofford, D. L. 1993. PAUP: Phylogenetic Analysis
Using Parsimony, Version 3.1. Smithsonian In-
stitution, Washington, D.C.

Vereshchaka, A. L. 1997. New family and superfamily
for a deep-sea caridean shrimp from the Gala-
thea collections.—Journal of Crustacean Biology
17(2):361-373.

Associate Editor: Christopher Boyko

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(4):523-528. 2004.

Macrobrachium patheinense, a new species of freshwater prawn
(Crustacea: Decapoda: Palaemonidae) from Myanmar

Hla Phone and Hiroshi Suzuki

Laboratory of Aquatic Resource Science, Faculty of Fisheries, Kagoshima University,
4-50-20 Shimoarata, Kagoshima 890-0056, Japan, e-mail: suzuki @fish.kagoshima-u.ac.jp

Abstract.—A new species of freshwater palaemonid prawn, Macrobrachium
patheinense, is described from Mayan Creek near Pathein City, Ayeyawaddy
Division, Myanmar. The new species is most closely related to M. mirabile
(Kemp, 1917), M. palaemonoides Holthuis, 1950, M. superbum (Heller, 1862)
and M. inflatum Liang & Yan, 1985, but can be differentiated by the rostrum
shape and dentition, telson shape, and the second pereiopod chela proportions.

Like other South East Asian countries,
Myanmar has a wealth of freshwater
streams, lakes, ponds and rivers, but unlike
its adjacent countries, the freshwater crus-
taceans have been poorly studied. Important
relevant investigations on freshwater deca-
pods have been undertaken in India, Thai-
land, China, Malaysia, Philippines, and In-
donesia (Cai & Dai 1999, Cai & Ng 2001,
Chace & Bruce 1993, Holthuis 1978, Jali-
hal et al. 1988, Liang & Yan 1985, Shokita
& Takeda 1989, Tiwari 1947, Wowor &
Choy 2001, Yeo et al. 1999, and others).

It was not until 1918 that the first signif-
icant carcinological study of Myanmar’s
fauna was conducted, and since then only
12 species of the shrimp genus Macro-
brachium Bate, 1868 have been recorded
(see Jalihal et al. 1988, Jayachandran 2001,
Kemp 1918, Tiwari 1952). A major study
by Cai & Ng (2002) reviewed the taxono-
my of the Myanmar palaemonid freshwater
prawns, reporting one new species and five
new records of Macrobrachium for the
country.

Myanmar’s unique geographic position
means that it has close connections with In-
dia, China, and the rest of the Indo-Malay-
sian region to the east and south. Thus,
there is a strong likelihood that further in-
vestigation will lead to more new taxonom-
ic and zoogeographic discoveries. In Myan-

mar, freshwater shrimps and prawns are im-
portant components of inland fisheries, and
further taxonomic and ecological studies
must be made an urgent priority in order to
ensure sustainable management and conser-
vation of stocks.

Specimens were collected from Mayan
Creek near Thayet Kone village, about five
miles west of Pathein City, Ayeyawaddy
Division, on 6 September 2001. All speci-
mens were preserved in formalin for ship-
ping to Japan and examined at the Labo-
ratory of Aquatic Resource Science, Fac-
ulty of Fisheries, Kagoshima University.
Among the collected specimens, 38 individ-
uals of a Macrobrachium species possessed
similar distinctive characteristics that could
not be attributed to any known species, and
are thus here described as a new species.

The holotype and 33 paratypes are de-
posited in the Laboratory of Aquatic Re-
source Science, Faculty of Fisheries, Ka-
goshima University, Kagoshima, Japan
(KUMB). Additional paratypes are also de-
posited in the Kitakyushu Museum of Nat-
ural History, Kitakyushu, Japan (KMNM),
and the Zoological Reference Collection
(ZRC), Raffles Museum, National Univer-
sity of Singapore.

Numbers in parentheses in “Materials
examined”’ indicate the post-orbital cara-

524

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1.
5 mm.

pace length in millimeters. Abbreviations
used include: M, male; FE female.

Family Palaemonidae Rafinesque, 1815
Genus Macrobrachium Bate, 1868
Macrobrachium patheinense, new species
Figs. 1-2

Materials examined.—Mayan Creek,
Thayet Kone village, Pathein City, Ayeya-
waddy Division, 6 Sep 2001: holotype, M
(8.96), KUMBcr 1101, paratype, 2M (9.16,
8.85), KMNH IvR 400.100, KMNH IvR
400.101, 2M (7.21, 8.75), ZRC 2003.0324,
33M OAS, Vij, B47, SID, 8.50;
8.65, 8.85, 8.61, 8.33, 8.68, 8.48, 9.08,
QM, BAS, BS, S07, BOS, 9.67, SS,
8.64, 8.10, 8.83, 7.97, 8.61, 8.16, 8.24,
7.92, 8.63, 9.10, 9.07, 7.91, 8.55), KUMBcer
1102-1134.

Diagnosis.—Carapace smooth, with an-
tennal and hepatic spine. Rostrum slender,
long; dental formula 2+10/5. Mandible
with 3-segmented palp. Scaphocerite broad,
with slightly concave outer margin. First
pereiopod slender, reaching to end of sca-
phocerite. Second pereiopod equal, ex-
tremely slender; carpus 2 times as long as
merus, finger 1.8 times as long as palm,
without teeth on cutting edge. Telson with
2 pairs of dorsolateral spinules; posterior

Macrobrachium patheinense. Lateral view of holotype, KUMBcr 1101; male (8.9 mm). Scale equals

margin ending in median tooth; 2 spines
and 2 plumose setae on each side, inner
spines well developed, outer spine very
short, plumose setae shorter than inner
spines.

Description.—Rostrum (Figs. 1, 2a)
long, slender, reaching beyond end of an-
tennular peduncle almost to end of scapho-
cerite, tip curving slightly upwards, upper
margin with 12 teeth (mode 13, range 11—
17), of which 2 teeth (mode 2, range 2—3)
are placed behind orbit; first tooth smaller
than second, placed further from second
than third; upper margin of rostrum with
single row of setae between teeth; lower
margin with 5 ventral teeth (mode 4, range
3-8), first tooth level with seventh and
eighth teeth; ventral portion with single row
of setae. Carapace (Fig. 1) has strong an-
tennal spine below lower orbital angle, pro-
duced anteriorly to broadly rounded lobe;
hepatic spine smaller than antennal spine,
placed below and some distance behind an-
tennal spine; branchiostegal groove present.

Abdomen smooth, glabrous, with broadly
rounded first to third pleurites; fourth and
fifth pleurites produced posteriorly, sixth
abdominal somite about 1.5 times as long
as fifth. Telson (Figs. 2b, c) 1.4 times length
of sixth abdominal somite, with 2 pairs of

VOLUME 117, NUMBER 4

S25

3
*
t
3
4
z
e

specter s <*

Fig. 2. Macrobrachium patheinense. Holotype, KUMBcr 1101; male (8.9 mm). a, lateral view of rostrum;
b, dorsal view of telson; c, tip of telson; d, antennule; e, antenna; f, mandible; g, second pereiopod; h, chela of
second pereiopod; i, dactylus and propodus of third pereiopod; j, first pleopod; k, second pleopod; 1, uropodal
diaeresis. Scales equal 1 mm.

526

dorsolateral spinules; posterior margin end-
ing in a median tooth, flanked on each side
by 2 spines and 2 plumose setae; inner
spine well developed, 4 times as long as
median tooth, outer spine very short, 2 plu-
mose setae slightly shorter than inner spine.

Eyes well-developed, with cornea as long
as stalk.

Basal segment of antennular peduncle
(Fig. 2d) broad, stylocerite very short, dis-
tinctly pointed, not reaching middle of basal
segment; anterolateral spine of basal seg-
ment reaching about middle of second seg-
ment; second segment as long as third seg-
ment; anterior margin of basal segment
strongly curved. Scaphocerite (Fig. 2e) 3.2
times as long as broad, not reaching tip of
rostrum; outer margin slightly concave,
ending in a tooth, not reaching end of la-
mella. Mandible (Fig. 2f) with outer, lateral,
3-segmented palp. Other mouth parts typi-
cal for genus.

First pereiopod slender, reaching end of
scaphocerite (Fig. 1); fingers slightly longer
than palm, with numerous setae; carpus
about twice as long as chela, broadest dis-
tally, narrowing proximally; merus shorter
than carpus; ischium about half as long as
merus. Second pereiopods (Figs. 2g, h)
equal in size and shape, extremely slender,
carpus reaching beyond scaphocerite by
half its length; chela gradually narrowing
proximally, finger very long, slender, about
1.8 times as long as palm (mean 1.7, range
1.4-1.9), same width throughout length,
cutting edge entire, tip curves inwards; car-
pus as long as chela, unarmed, with distal
portion broadest; merus about half as long
as carpus (mean 0.7, range 0.5—0.9) but
equal with ischium. Pereiopods 3—5 slender,
subequal in size; third pereiopods over-
reaching scaphocerite by length of entire
dactylus; dactylus (Fig. 21) slender, concave
on ventral, with numerous setae on dorsal
surface, measuring about % of propodus
length; propodus with eight spinnules on
ventral surface, about twice as long as car-
pus; merus nearly as long as propodus; is-
chium about same length as carpus; fourth

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

pereiopods shorter than fifth but longer than
third.

Exopod of first pleopod (Fig. 2j) oval-
shaped, with small endopod, inner margin
concave. Second to sixth pleopods nearly
equal with endopods and exopods; endopod
with a slender appendix interna. Second
pleopod (Fig. 2k) with appendix masculina,
placed between appendix interna and en-
dopod; appendix masculina longer, stronger
than appendix interna, bearing several stiff
setae. Uropods reaching beyond end of tel-
son; exopods ovate, outer margin straight,
inner margin convex, uropodal diaeresis
(Fig. 21) with a spine slightly longer than
outer angle; endopods broadly ovate, small-
er than exopods.

Color.—Grayish white when live.

Etymology.—The specific name is adapt-
ed from the type locality (Pathein) where
the specimens were collected.

Distribution.—Macrobrachium pathei-
nense inhabits freshwater and slightly
brackish water habitats, known so far only
from the type locality.

Remarks.—Macrobrachium patheinense
is similar to the Palaemon-like Macro-
brachium species, that have slender and
delicate pereiopods, especially M. mirabile
(Kemp, 1917), M. palaemonoides Holthuis,
1950, M. superbum (Heller, 1862) and M.
inflatum Liang & Yan, 1985. Macrobrach-
ium patheinense is, however, distinguish-
able from M. mirabile by the shape of the
rostrum and telson. The rostrum of M. path-
einense is slender and longer than the sca-
phocerite, while that of M. mirabile is
shorter than the scaphocerite, and has a
high dorsal crest (Kemp 1917). The telson
of the former has two pairs of plumose se-
tae slightly shorter than the inner spine, but
that of the latter has only one pair of plu-
mose setae longer than the inner spine. The
new species is also distinguished from M.
palaemonoides by shapes of the rostrum
and telson, and the proportions of chelae of
the second pereiopods. The rostrum of M.
patheinense is armed with teeth along the
entire upper margin, but that of M. palae-

VOLUME 117, NUMBER 4

monoides has an unarmed area on its distal
half (Holthuis 1950, Kamita 1974). The tel-
son terminates in a short median tooth in
M. patheinense, but this tooth is longer in
M. palaemonoides (Kamita 1974). The
movable finger of the second pereiopod is
1.4—1.9 (mean 1.7) times as long as the
palm in M. patheinense, but 1.3—1.4 times
as long as the palm in M. palaemonoides
(Chace & Bruce 1993, Holthuis 1950). The
new species can easily be distinguished
from M. superbum by the shape of the ros-
trum and the second pereiopod chela pro-
portions. The rostrum doesn’t reach beyond
the distal end of the scaphocerite in M. su-
perbum (Cai & Dai 1999, Holthuis 1950),
but distinctly further in M. patheinense. In
addition, the upper margin of the rostrum is
generally straight in M. superbum, but dis-
tally upcurved in M. patheinense. The mov-
able finger of second pereiopod is 1.2—1.5
times as long as the palm in M. superbum,
but 1.4—1.9 (mean 1.7) times in M. path-
einense. The rostral shape and formula of
M. patheinense is most similar to those of
M. inflatum, but the second pereiopods and
telson of both species are different. The
movable finger of the second pereiopod of
M. inflatum is subequal to the length of the
palm (0.9-1.0 from the figures of Cai &
Dai (1999) and Liang & Yan (1985)), but
that of M. patheinense is much longer than
the palm (1.4—1.9, mean 1.7). The telson of
M. inflatum bears three pairs of plumose se-
tae, these setae being longer than the inner
spine on the posterior margin, but M. path-
einense has only two pairs of plumose setae
that are slightly shorter than the inner spine.

The unique chela of M. patheinense re-
sembles that of Leandrites stenopus Hol-
thuis, 1950, and Pseudopalaemon bouvieri
Sollaud, 1911, however the presence of a
mandibular palp in M. patheinense confirms
its placement in Macrobrachium and distin-
guishes it from all Leandrites and Pseudo-
palaemon species (Holthuis, 1993).

Thus, the new species appears to occupy
an interesting phylogenetic position and
should be included in future studies inves-

527

tigating generic relationships within the

family Palaemonidae.

Acknowledgments

We are grateful to Peter J. EK Davie of
Queensland Museum, Australia, Peter K. L.
Ng and Yixiong Cai of National University
of Singapore, L. B. Holthuis of National
Museum of Natural History, Leiden, The
Netherlands, and an anonymous reviewer
for their critical readings of the manuscript.

Literature Cited

Cai, Y., & A. Y. Dai. 1999. Freshwater shrimps (Crus-
tacea: Decapoda: Caridea) from the Xishuang-
banna region of Yunnan Province, southern
China.—Hydrobiologia 400:211—241.

, & PK. L. Ng. 2001. The freshwater decapod

crustaceans of Halmahera, Indonesia.—Journal

of Crustacean Biology 21(3):665—695.

, & . 2002. The freshwater palaemonid
prawns (Crustacea: Decapoda: Caridea) of
Myanmar.—Hydrobiologia 487:59—83.

Chace, E A., Jr., & A. J. Bruce. 1993. The Caridean
Shrimps (Crustacea: Decapoda) of the Albatross
Philippine Expedition 1907-1910. Part 6: Su-
perfamily Palaemonoidea——Smithsonian Con-
tributions to Zoology 543:1—52, pls. 1-7.

Heller, C. 1862. Neue Crustaceen, gesammelt wahrend
der Weltumseglung der k. k. Fregatte Novara.
Zweiter vorlauliger Bericht.—Verhandlungen
der kaiserlich-koniglichen zoologisch-botan-
ischen Gesellschaft in Wien 12:519-528.

Holthuis, L. B. 1950. The Decapoda of the Siboga Ex-
pedition. Part 10. The Palaemonidae collected
by the Siboga and Snellius Expeditions with re-
marks on other species. 1. Subfamily Palae-
moninae.—Siboga Expeditie 39(a9):1—268,
figs. 1-52.

. 1978. A collection of decapod Crustacea from

Sumba, Lesser Sunda Islands, Indonesia.—

Zoologische Verhandelingen 162:1—55.

. 1993. The Recent genera of the Caridean and
Stenopodidean Shrimps (Crustacea, Decapoda),
with an Appendix on the Order Amphionidacea.
Nationaal Natuurhistorisch Museum, Leiden,
328 pp.

Jalihal, D. R., S. Shenoy, & K. N. Sankolli. 1988.
Freshwater prawns of the genus Macrobrach-
ium Bate, 1868 (Crustacea, Decapoda, Palae-
monidae) From Karnataka, India.—Records of
the Zoological Survey of India, Miscellaneous
Publication, Occasional Paper No. 112:1—74.

Jayachandran, K. V. 2001. Palaemonid Prawns: Bio-

528

diversity, Taxonomy, Biology and Management.
Science Publishers, Enfield, 624 pp.

Kamita, T. 1974. Four species of the Nepalese
prawns.—Researches on Crustacea 6:1—16, pls.
1-2.

Kemp, S. 1917. Notes on Crustacea Decapoda in the
Indian Museum. IX. Leander styliferus, Milne-
Edwards, and related forms.—Records of the
Indian Museum 13:203—231, pls. 8-10.

. 1918. Crustacean Decapoda of the Inle Lake
Basin.—Records of the Indian Museum. 14:81—
102, pls. 15-16.

Liang, X.-Q., & S.-I. Yan. 1985. New species and new
record of palaemonidae from China (Crustacea
Decapoda) (in Chinese with English summa-
ry).—Acta zootaxonomica Sinica, 10(3):253—
258. figs. 1—4.

Shokita, S., & M. Takeda. 1989. A new freshwater
prawn of the genus Macrobrachium (Decapoda,
Caridea, Palaemonidae) from Thailand.—Bul-
letin of the National Science Museum, Tokyo,
Series A (Zoology), 15(3):147-154.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Sollaud, E. 1911. Pseudopalaemon Bouvieri, nouveau
genre, nouvelle espece, de la Famille des Palae-
monidae.—Bulletin du Museum National
d’ Histoire Naturelle, Paris 17:12—16.

Tiwari, K. K. 1947. Preliminary descriptions of two
new species of Palaemon from Bengal.—Re-
cords of the Indian Museum 45(4):329—331.

. 1952. Diagnosis of new species and subspe-
cies of the genus Palaemon Fabricius (Crusta-
cea: Decapoda).—Annals and Magazine of Nat-
ural History 5:27-32.

Wowor, D., & S. C. Choy. 2001. The freshwater
prawns of the genus Macrobrachium Bate, 1868
(Crustacea: Decapoda: Palaemonidae) from
Brunei Darussalam.—The Raffles Bulletin of
Zoology 49(2):269—289.

Yeo, D. C. J., Y. Cai, & P. K. L. Ng. 1999. The fresh-
water and terrestrial decapod Crustacea of Pulau
Tioman, Peninsular Malaysia.—The Raffles
Bulletin of Zoology, Supplement No. 6:197—
244.

Associate Editor: Christopher Boyko

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(4):529-540. 2004.

A new species of Enhydrosoma Boeck, 1872 (Copepoda:
Harpacticoida: Cletodidae) from the Eastern Tropical Pacific

Samuel Gomez

Instituto de Ciencias del Mar y Limnologia, Unidad Académica Mazatlan. Joel Montes Camarena
s/n, Ap. Postal 811, Mazatlan 82040, Sinaloa, México, e-mail: samuelgomez @ola.icmyl.unam.mx

Abstract.—Some enhydrosomids were found while sorting samples taken
from the Urias system during a short-term study on the effects of organic
enrichment on the abundance and diversity of benthic copepods. Upon careful
examination, these specimens proved to belong to a new species, Enhydrosoma
brevipodum, of the species-group defined by the lack of sexual dimorphism on
the male P3 and can be separated by the reduced exopod of female P5. En-
hydrosoma brevipodum, whose full description is herein provided, constitutes
the fourth record of the genus from the Pacific Mexican coast.

The genus Enhydrosoma Boeck, 1872 is
a group of harpacticoid copepods common-
ly found in shallow brackish and marine
coastal systems worldwide. Some Enhydro-
soma specimens were found in sediment
samples from two shallow brackish systems
in central (Ensenada del Pabellon lagoon)
and southern (Urias system) Sinaloa during
the course of two short-term studies about
the effects of organic enrichment on the dis-
tribution and abundance of meiofauna (see
Gomez-Noguera & Hendrickx 1997) and
on the diversity of benthic harpacticoids.
Some of these specimens belong to three
species recently described by Go6mez
(2003), whereas some specimens constitute
the Pacific counterpart of Enhydrosoma la-
cunae Jakubisiak, 1933 (Gomez 2003),
originally described from Cuba and rede-
scribed by Fiers (1996) from the Yucatan
Peninsula. While sorting samples taken
from Urias system, some specimens of a
different species of Enhydrosoma were
found. These specimens proved to belong
to a new species mainly characterized by
the reduced exopod of female P5. A de-
tailed description of this species is herein
provided.

Materials and Methods

Quantitative sediment cores were taken
for the analysis of the effects of organic en-
richment on benthic copepods along a pol-
luted estuary (Urias system) in southern
Sinaloa (north-western Mexico) during
2001 and 2002. Sediment samples were
taken with an Eckman box corer with a
sampling area of 225 cm?, and subsamples
were taken using plastic corers with a sam-
pling surface of 7 cm?. Sediment cores were
subdivided vertically into separate 1 cm
slices to a depth of 5 cm. Each slice was
fixed with 10% formalin, and sieved
through 500 and 63 pm sieves to separate
macro- and meiofauna. Meiofauna was pre-
served in 70% ethanol and stained with
Bengal Rose until further inspection. Mei-
ofaunal major taxa were quantified and co-
pepods (cyclopoids, poecilostomatoids and
harpacticoids) were separated from the rest
of meiofauna and stored in 70% ethanol for
further investigation. Observation and
drawings of the species described herein
were made from whole and dissected spec-
imens mounted in lactophenol, under 100
oil immersion objective using a Leica com-
pound microscope equipped with drawing

530

tube and phase contrast. The type material
was deposited in the collection of the Insti-
tuto de Ciencias del Mar y Limnologia, Ma-
zatlan Marine Station. The terminology
proposed by Huys & Boxshall (1991) for
the general description and armature for-
mulae was adopted. Abbreviations used in
the text and tables: P1l—P6, first to sixth
swimming leg; EXP, exopod; ENP, endo-
pod.

Family Cletodidae T. Scott, 1904 sensu
Por (1986)
Genus Enhydrosoma Boeck, 1872

Enhydrosoma brevipodum, new species

Type material—One female holotype
preserved in 70% ethanol (EMUCOP-
090301-73), one dissected male allotype
(EMUCOP-090301-62), and one dissected
female paratype (EMUCOP-090301-61);
collected from station 10; 9 Mar 2001; leg.
S. Gomez.

Type _ locality.—Urias system, Sinaloa,
northwestern Mexico (23°09’—23°13'N,
106°20’—106°25'W).

Etymology.—The specific name alludes
to the reduced exopodal lobe of female P5.

Female.—Body (Fig. 1A, 2A) tapering
from posterior margin of cephalothorax,
curved in lateral view; length of holotype,
420 wm from tip of rostrum to posterior
margin of caudal rami. Cephalic shield
about % total length, with strongly folded
lateral and dorsal surface, posterior margin
plain, with sensilla arising from distinct
cones. Rostrum triangular, fused to cephalic
shield, with rounded tip, with two sensilla.
Dorsal surface of free thoracic somites (P2—
P4) smooth, with sensilla arising from dis-
tinct cones along plain posterior margin.
First urosomite (P5-bearing somite) as pre-
ceding somites except for fewer sensilla.
Surface of genital double somite smooth,
with dorsolateral division between first and
second genital somite (second and third
urosomites), posterior margin of both gen-
ital somites plain, first somite with sensilla
arising from distinct cones along posterior

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

margin, second somite as first one except
for two additional tube pores (arrowed in
Fig. 1A), both somites with additional sen-
silla arising from paired bulbous structures
laterally; genital somites completely fused
ventrally, first somite bearing pair of P6 and
genital pore, the former each bearing a
short spinulose spine, and with an associ-
ated tube-pore (arrowed in Fig. 2A), copu-
latory pore covered by integumental fold,
ventral surface of second segment smooth,
except for spinules and fragile setules along
posterior margin between pair of sensillum-
bearing cones. Dorsal surface of fourth and
fifth urosomite as in preceding somite, ex-
cept for lack of central pair of sensilla on
posterior margin of fourth somite, and lack
of sensilla along posterior margin of fifth
urosomite, both somites with pair of tube
pores (arrowed in Fig. 1A); ventral surface
of fourth and fifth urosomite smooth, fourth
urosomite ornamented with spinules and
setules as in second genital somite, fifth
urosomite with only spinules along poste-
rior margin. Anal segment smooth, rounded
anal operculum without ornamentation and
flanked by pair of sensilla. Caudal rami cy-
lindrical and about 8.3 times as long as
wide, with seven setae in all, setae I and II
located in proximal fifth, the former arising
ventrally and about % total length of seta
II, the latter dorsal to seta I, seta III slightly
longer than seta II and arising in the middle
along outer margin of ramus, seta IV and
V fused, seta VI located in distal inner cor-
ner and as long as seta II, seta VII arising
in proximal third at the level between seta
II and III.

Antennule (Fig. 2B). 5-segmented; sur-
face of segments smooth except for spinular
row on first, third and fourth segment. Ar-
mature formula 1-(1), 2-(7), 3-(7+ae), 4-
(1), 5-(11+ae).

Antenna (Fig. 3A, B) with proximal and
distal set of spinules on inner margin of al-
lobasis; with very small distal abexopodal
seta close to distal set of spinules, the latter
difficult to see and can be easily mistaken
for a setule (arrowed in Fig. 3A). Exopod

a gS
NET ea i aS

Diy N Se ie)

ee awe AI Cs Gee ae

ry
; ¥
! \ SS ia a
s,s Se 3 1 3e
We. ms
i i
an Ake, Pc
a \ footy
i “ & SS

532 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

\ |
=
=
|
BB i —
Z | ec
— . >=
=
LZ
LEE

Fig. 2. Enhydrosoma brevipodum, female paratype EMUCOP-090301-61. A, urosome, ventral (P5 bearing-
somite omitted; tube pores in genital field arrowed); B, antennule. Scale bar: A, 100 wm; B, 75 wm.

VOLUME 117, NUMBER 4

Fig. 3.
B, normal exopod of antenna; C, maxilliped; D, Pl. Scale bar, 50 pm.

1-segmented and armed with two bipinnate
elements (Fig. 3B). Endopodal segment or-
namented with two strong spines subdistal-
ly along inner margin; distal margin five se-
tae/spines (outer pectinate spine seemingly
without fused small seta), and ornamented
with two hyaline frills on outer margin.
Mandible (Fig. 4A, B) with slender gna-
thobase; biting edge with uni- and multi-
cuspidate teeth, and one bare seta at distal

533

Enhydrosoma brevipodum, female paratype EMUCOP-090301-61. A, antenna with aberrant exopod;

inner corner. Palp well-developed, 1-seg-
mented and armed with one endopodal and
two basal setae (Fig. 4B).

Maxillule (Fig. 4C). Arthrite with five
distal and two lateral elements, and two sur-
face setae; coxal endite fused to basis and
represented by one seta, basis represented
by two distal setae, endopod and exopod
represented by one seta each.

Maxilla (Fig. 4D) with short spinular row

534

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Enhydrosoma brevipodum, female paratype EMUCOP-090301-61. A, mandible; B, mandibular palp;
C, maxillule; D, maxilla. Scale bar: A, B, 50 wm; C, D, 25 pm.

on distal inner corner of syncoxa; proximal
syncoxal endite with two slender and bare
setae and one bipinnate element; distal syn-
coxal endite with two elements (inner
strongly pinnate, outer anvil shaped, fused
to endite and with only one pinnule). Al-
lobasal endite with non-articulated spine
and two setae. Endopod represented by two
setae fused at base.

Maxilliped (Fig. 3C) prehensile, with
short and unarmed syncoxa; basis with spi-
nules along inner margin; claw slender and
curved distally, with accessory seta.

P1 (Fig. 3D). Coxa and basis ornamented
as depicted, the latter with inner and outer
setae. Exopod three-, endopod 2-segment-
ed, the latter reaching distal third of last ex-
opodal segment.

P2—P4 (Figs. 5A, B, 6A) with coxa and
basis ornamented as shown, the latter with
outer plumose seta. Exopod three-, endopod
2-segmented. Endopod of P2 slightly longer
than, of P3 as long as, of P4 clearly shorter
than first and second exopodal segments
combined. Armature formulae of P1—P4 as
follows:

235)

VOLUME 117, NUMBER 4

Sl,

—— :
he Se aN eS ‘i
— ee EN 3
—< = ;
=> ri ae . J
2S SSS iS : :

SSS A

Ss 5

ig <

a ee oa Sus se 5
Z Se a PZ :
a SX te a S

== AY SO \y \ 8,

3 = Ss NX ( 2
oS NR Sa \ Ny Ay :

3 Ce Gr IN 2

ee. \ . Os E

_ a :

y S < &

Ss Ns :

oe <a > Z . 8

aeitig E/4,

a <e Cig Re By / / §

= t es. ee Se. ~ yyy :

536

Pl p2 P3 P4

EXP 1J-0;1-0;11,2,0 I-0;1-0;11,2,0 I-0;I-0;11,2,1 J-0;I-0;11,2,1
EXP 0-0;1,1,1 0-0;0,2,0 0-0;1,2,0 0-0;1,2,0

P5 (Fig. 6B). Baseoendopod and exopod
fused but with partial suture still visible on
posterior face. Exopod almost square, with
a few setules on outer margin, three setae
(one minutely pinnate) on distal margin and
a large tri-pinnate seta on inner margin.

Baseoendopod with transverse rows of
spinules on endopodal lobe and at base of
exopod; outer seta on long pedicel. Endo-
podal lobe reduced with three elements (a
strong inner pinnate spine and a long pin-
nate seta on distal margin, and a slender
pinnate seta on inner margin).

Male.—Total body length, 450 wm. Gen-
eral dorsal body shape (not shown) as in
female. Urosome (Fig. IB) as in female
dorsally; second and third urosomites dis-
tinct ventrally (Fig. 7A); first urosomite
without ventral spinular ornamentation
along posterior margin; second urosomite
with P6 represented by one fused and one
free plate close to posterior margin ventral-
ly (Fig. 7A), and ornamented with spinules
along posterior margin; third to fifth uro-
somites with spinules and fragile setules
along posterior margin; fifth urosomite
without sensilla. Caudal rami (Figs. 1B,
7A), mouth parts, and P1l—P4 (not shown)
as in female.

Antennule (Fig. 7B) 6-segmented, sub-
chirocer; surface of segments smooth ex-
cept for short spinules of first segment and
longitudinal row of long spinules on fourth
globose segment. Armature formula diffi-
cult to define.

P5 (Fig. 7A). Baseoendopod and exopod
fused, the former with outer extension bear-
ing outer seta and ornamented with spinules
at base of exopod and distally on endopodal
lobe, the latter small and armed with one
inner seta and one outer spine. Exopod
elongate, about 1.5 times as long as wide,
and armed with two apical setae (outermost
slender and about % total length of inner-
most bipinnate seta).

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Remarks.—In his outstanding prelimi-
nary revision of Enhydrosoma, Gee (1994)
accurately stated that the armature comple-
ments of the second endopodal segment of
P1 are not always reliable. This is true, as
shown by Gee (1994), for E. curticauda
Boeck, 1872, and also for E. propinquum
(Brady, 1880) (compare Apostolov & Mar-
inov (1988) and Sars (1909)). Also, Gee
(1994) suggested that the often reduced in-
ner seta on the Pl second endopodal seg-
ment could either be overlooked or mistak-
enly regarded as a spinule or setule in pre-
vious descriptions. On the other hand, it has
to be noted that the males of some species
(e.g., E. intermedia Chislenko, 1978, E. ca-
soae G6mez, 2003 and E. solitarium G6-
mez, 2003) remain unknown and their po-
sition within Enhydrosoma regarding the
male P3 endopod is still pending. Even
when the male of a known species is found,
the description of the P3 endopod is often
omitted, e.g., Arlt (1983) for E. longifur-
catum Sars, 1909 and E. sarsi (TV. Scott,
1904); Bodin (1970) for E. propinquum;
Apostolov & Marinov (1988) for E. gariene
Gurney, 1930; Monchenko (1967), Apos-
tolov & Marinov (1988) and Bodin (1970)
for E. caeni Raibaut, 1965; Marinov &
Apostolov (1985) for E. longicauda Mari-
nov & Apostolov, 1983). Therefore, a thor-
ough revision of the species currently as-
signed to Enhydrosoma is urgently needed
to unravel the phylogenetic relationships
within the genus.

The species herein described belongs to
Gee’s (1994) type 1 (being the male P3 en-
dopod 2-segmented, with the same arma-
ture formula and form of armature elements
as in the female), which seems to be rather
common in Enhydrosoma. In fact, the un-
modified condition of the male P3 endopod
has been reported for 10 species—E. migoti
Monard, 1926: E. tunisensis Monard, 1935
(considered as incertae sedis within Enhy-
drosoma by Wells 1965); E. propinquum,
E. sarsi, E. longifurcatum, E. latipes (A.
Scott, 1909); and E. gariene, E. longicauda,
E. pectinatum Wells & Rao, 1987; E. sor-

VOLUME 117, NUMBER 4

32)

Ss“

<<<

Fig. 6. Enhydrosoma brevipodum, female paratype EMUCOP-090301-61. A, P4; B, PS. Scale bar: 50 pm.

didum Monard, 1926); and E. caeni and E.
rosae Fiers, 1996. Of these, only Sars’
(1909) E. propinquum, E. latipes, E. rosae
and E. migoti have been reported bearing
three setae on the second endopodal seg-
ment of P1 (although the armature formula
of the endopod of Pl needs confirming in
some other species), and E. brevipodum

could well be most closely related to the E.
longiforcatum species-group (E. longifur-
catum, E. gariene, E. sordidum, E. caeni
and E. pectinatum) based on the combina-
tion of the following character states: (a)
the caudal ramus shape and arrangement of
setae I, II, IJ and VII, (b) rostrum, (c) num-
ber of setae on the mandibular palp, (d) fu-

538 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

SFT 77 PR

Fo jimi

iN
|
ai mA

re sa aq (ee

Fig. 7. Enhydrosoma brevipodum, male allotype EMUCOP-090301-62. A, urosome, ventral, showing PS
and P6; B, antennule. Scale bar: A, 100 1m; B, 75 pm.

VOLUME 117, NUMBER 4

sion of coxal endite and basis and number
of setae on the whole palp, (ec) number of
elements on distal syncoxal endite of the
maxilla, (f) lack of sexual dimorphism on
P3 endopod, (g) the fusion of exopod and
baseoendopod on P5 of both sexes, and (h)
number of setae on female P6. The species
described herein differs from the other spe-
cies of the E. longifurcatum species group
in the following characters: (a) presence of
a seta on abexopodal margin of the first en-
dopodal segment of the antenna, (b) number
of setae on the second endopodal segment
of Pl (except for Sars’ (1909) E. propin-
quum), (c) proportions of P5 exopod in both
sexes and arrangement of setae in female
(in all other species in this group the large
inner seta is on distal margin and outer se-
tae on outer margin, but in E. brevipodum
the large seta is located internally and the
others on distal margin. In this regard it has
to be noted that EF. longifurcatum and E.
sordidum were reported to have only three
setae on P5 exopod, but it is highly prob-
able that there are four setae, being that the
middle outer element is very small and
weak (J. M. Gee, in litt.). The inner and
outer elements of female exopod of P5 are
strong and pectinate spines in E. longifur-
catum, E. gariene, E. sordidum, E. caeni
and E. pectinatum, but it is a pinnate seta
in E. brevipodum. On the other hand, the
genital field of E. brevipodum is similar to
that described for E. curticauda by Gee
(1994) in that the single copulatory pore is
covered by an integumental fold, but differs
in the armature formula of P6 (with two and
one seta in E. curticauda and E. brevipod-
um, respectively), spinular ornamentation
(with and without spinules in E. curticauda
and E. brevipodum, respectively), and num-
ber and location of tube pores (with three
tubular extensions arising from two pores
in E. curticauda, and with two tubular ex-
tensions arising from two pores in E. brev-
ipodum). The genital field of E. brevipodum
seems to be similar to that described by Gee
(1994) for E. propinquum in the number
and location of the tube pores, and similar

539

to that described for Strongylacron buch-
holzi (Boeck, 1872) and E. gariene in the
armature formula and lack of spinular or-
namentation of P6 (see Gee, 1994:95, figs.
9C-E). Also, according to Gee (1994) the
position of spinules and/or setules on the
inner margin of the antennal allobasis could
be used to discern which abexopodal seta
has been lost in a given species. The distal
abexopodal seta observed for E. brevipod-
um is very small and can be easily mistaken
for a setule. However, careful examination
revealed the presence of small spinules at
the base of this element which, following
Gee (1994), could indicate either the site
where a seta is possibly attached or the site
where the abexopodal seta must have been
situated in those cases where the proximal
(basal) or the distal seta was lost. All the
above suggests the presence of a very re-
duced distal seta and loss of the proximal
(basal) seta in E. brevipodum, which is also
the case for most species within Enhydro-
soma (Gee, 1994). It is interesting to note
that a female paratype of E. brevipodum
(EMUCOP-090301-61) was found to pos-
sess an aberrant antennal exopod bearing
three setae (two well-developed and one
dwarfed element) (see Fig. 3A). The same
has been observed for a female of FE. cur-
ticauda from East Finnmark (Gee, 1994:
97). To the best of my knowledge, E. brev-
ipodum is unique within the genus by the
reduced exopod of the female P5.
Enhydrosoma brevipodum was found in
sandy sediments taken in the mouth of the
Urias brackish system. The sampling sta-
tion (stn. 10) where the newly found spe-
cies was taken is under the direct effects of
marine water and is characterized by sandy
bottom and low contents of chlorophyl *“‘a”’
(3189.2 mg 1!) and organic mater (2.8%).

Acknowledgments

The author is grateful to Mrs. I. M. Bus-
tos Hernandez, Mr. EK N. Morales Serna and
Dr. J. Salgado Barragan for their support
and help during field work and sample pro-

540

cessing. This is a contribution to project
IN202400 funded by the Research and
Technological Innovation Projects Support
Programme (Programa de Apoyo a Proy-
ectos de Investigaci6n e Innovacion Tec-
noldgica) of the Office for General Affairs
of the Academic Staff (Direccion General
de Asuntos del Personal Académico) of the
National Autonomous University of Mexi-
co (U. N. A. M). The author is grateful to
one anonymous referee and to Dr. J. M. Gee
for their corrections and suggestions to im-
prove the content of the manuscript.

Literature Cited

Apostolov, A. M., & T. M. Marinov. 1988. Copepoda,
Harpacticoida, ((Fauna Bulgarica)) 18, In Ae-
dibus Academiae Scientiarum Bulgaricae.—So-
fia: 1-384.

Arlt, G. 1983. Taxonomy and ecology of some har-
pacticoids (Crustacea, Copepoda) in the Baltic
Sea and Kattegat.—Zoologische Jahrbticher
Systematik 110:45-85.

Bodin, P. 1970. Copépodes harpacticoides marins des
environs de la Rochelle. 1. Espéces de la vase
intertidale de Chatelaillon.—Tethys 2:385—436.

Fiers, EF 1996. Redescription of Enhydrosoma lacunae
Jakubisiak, 1933 (Copepoda, Harpacticoida);
with comments on the Enhydrosoma species re-
ported from West Atlantic localities, and a dis-
cussion of cletodid development.—Sarsia 81:1—
Die

Gee, J. M. 1994. Towards a revision of Enhydrosoma
Boeck, 1872 (Harpacticoida: Cletodidae sensu
Por); a re-examination of the type species, E.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

curticauda Boeck, 1972, and the establishment
of Kollerua gen. nov.—Sarsia 79:83—107.

Gomez, S. 2003. Three new species of Enhydrosoma
and a new record of Enhydrosoma lacunae (Co-
pepoda: Harpacticoida: Cletodidae) from the
Eastern Tropical Pacific—Journal of Crusta-
cean Biology 23:94—118.

Gomez-Noguera, S. E., & M. E. Hendrickx. 1997. Dis-
tribution and abundance of meiofauna in a sub-
tropical coastal lagoon in the south-eastern Gulf
of California, México.—Marine Pollution Bul-
letin 34:582—587.

Huys, R., & G. A. Boxshall. 1991. Copepod evolu-
tion.—The Ray Society, London, 468 pp.
Marinov, T., & A. Apostoloy. 1985. Copépodes har-
pacticoides de l’?Océan Atlantique. 1. Espéces
des cotes du Sahara Espagnol.—Cahiers de

Biologie Marine 26:165—180.

Monchenko, V. I. 1967. Recent data on the harpacti-
ciod (Crustacea, Copepoda) of the Black Sea.—
Doklady Akademii Nauk Ukrajin RSR 29:461—
465.

Por, E D. 1986. A re-evaluation of the family Cleto-
didae Sars, Lang, (Copepoda, Harpacticoida).
Pp. 420—425 in G. Schriever, H. K. Schminke,
& C.-t. Shih, eds., Proceedings of the Second
International Conference on Copepoda.—Syl-
logeus 58.

Sars, G. O. 1909. Copepoda Harpacticoida. Parts XXV
& XXVI. Laophontidae (concluded), Cletodidae
(part.). An account of the Crustacea of Norway,
with short descriptions and figures of all the
species. Bergen Museum, Bergen, 5:277—304.

Wells, J. B. J. 1965. Copepoda (Crustacea) from the
meiobenthos of some Scottish marine sub-lit-
toral muds.—Proceedings of the Royal Society
of Edinburgh 69 B, Part I:1—33.

Associate Editor: Janet Reid

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(4):541—544. 2004.

New record of Ophiosyzygus disacanthus Clark, 1911
(Echinodermata: Ophiuroidea: Ophiomyxidae) in the Caribbean Sea

Giomar Helena Borrero-Pérez and Milena Benavides-Serrato

(GHBP) Museo de Historia Natural Marina de Colombia—MHNMC, Instituto de Investigaciones
Marinas y Costeras—INVEMAR, Apartado Aéreo 1016, Santa Marta, Colombia, E-mail:
ghborrero @invemar.org.co;

(MBS) Facultad de Biologia, Universidad de Puerto Rico, Recinto Universitario de Mayagtiez,
Cédigo Postal 00680, Mayagtiez, Puerto Rico, U.S.A., E-mail: milbese@hotmail.com

Abstract.—Ophiosyzygus disacanthus Clark, 1911 is reported for the first
time in the Caribbean Sea; this is the third record of this species in the liter-
ature. A comparison with two other records from the southwestern coast of
Japan (Clark 1911) and the Gulf of Mexico (Turner & Heyman 1995) is pre-

sented.

Ophiosyzygus disacanthus was described
by Clark (1911) from the southwestern
coast of Japan. This species was further
documented by Turner & Heyman (1995),
who revised the diagnosis of the monotypic
genus Ophiosyzygus and the description of
its type species, O. disacanthus, based on
the type material (two specimens) and on
new material (two specimens) collected re-
cently from the Gulf of Mexico, off the
southwestern coast of Florida. Among other
characters, the genus was diagnosed by
Clark (1911) as lacking radial shields and
dorsal arm plates; but Turner & Heyman
(1995) found these structures, and they
emended the generic diagnosis, completed
the original description of O. disacanthus,
and commented on the family Ophiomyxi-
dae, specifically about its small radial
shields and thin dorsal arm plates, which
have been often overlooked in this family.
In this note, we record O. disacanthus from
the Caribbean Sea, specifically off the Co-
lombian coast.

Materials and Methods

As part of a project developed by the
Marine and Coastal Research Institute (IN-
VEMAR), designed to inventory the ben-

thic macrofauna from the continental shelf
and upper slope region of the Caribbean
coast of Colombia, two specimens of
Ophiosyzygus disacanthus were collected at
9°46'61"N, 76°13'72”W in 155 m depth on
26 Apr 2001. Sampling was conducted on
board the B/I Ancén; a 5 m opening trawl
net was used. The material is deposited in
the collection of the Museo de Historia Nat-
ural Marina de Colombia (MHNMC), cat-
alogue number INV EQU01927. The spec-
imens were measured, and photographs
were taken, after fixing in 70% ethanol. The
plates of the ventral interradius of the disc
were measured after treatment with sodium
hypochlorite, and also the disc granules
were measured.

Family Ophiomyxidae Ljungman, 1867
Ophiosyzygus Clark, 1911
Ophiosyzygus disacanthus Clark, 1911
Fig. 1

Remarks.—The specimens of O. disa-
canthus collected off the Colombian coast
agree with the diagnosis of the genus as
emended by Turner & Heyman (1995), as
well as with the characteristics included in
the species description about the thornier
arm spines, the presence of dorsal arm

542 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1.

Ophiosyzygus disacanthus. A. Ventral surface of disc (disc diameter: 15 mm). Note shape of oral

papillae and white, opaque and irrregular granules. B. Dorsal surface of disc and proximal part of arm. Note
small radial shields (rs). C. Lateral view of arm. Note upper arm spines successively united by broad, thin,
horizontal membrane. D. Four plates of ventral interradii of disc.

plates, and of flat and multiperforate plates
embedded in the skin of the ventral inter-
radii of the disc; these are visible after treat-
ment with sodium hypochlorite (Fig. 1D).
The ventral interradial plates are slightly

longer (115.7 + 35.1 wm, n = 23) than
those of the Gulf of Mexico specimens (111
+ 22 wm, n = 24) measured by Turner &
Heyman (1995). The number of oral papil-
lae is variable; 3—6 in the Colombian spec-

VOLUME 117, NUMBER 4

imens (Fig. 1A), 2—4 in those from Japan,
and 2—5 in Florida specimens. The irregular
granules of the disc are white, opaque, and
are present in the ventral (Fig. 1A) and dor-
sal integument. One of the specimens has
ventral granules larger (155 + 34.6 pm, n
= 16) than the dorsal ones (104 + 15.4 pm,
n = 15); in the other specimen, the dorsal
and ventral granules are similar in size
(111.4 + 21.6 pm, n = 21), but the dorsal
granules are elongate. In general the gran-
ules of the two specimens from Colombia
are smaller than those of the holotype (232
+ 67.8 wm, n = 34) and paratype (221 +
53.8 wm, m = 21) and similar in size to the
specimens from Florida (84 + 21.9 um, n
= 9; 124 + 32.8 pm, nm = 10) (Turner &
Heyman 1995). The arms of both speci-
mens are broken. The discs are damaged,
disc diameters are 10 and 15 mm, 3 mm
more than the maximum size of previously
collected specimens (Clark 1911, Turner &
Heyman 1995). The depth (155 m) where
the Colombian specimens were taken is
within the range recorded for other speci-
mens: 188—278 m for the type material in
the Pacific Ocean and 127—159 m for the
specimens collected in the eastern Gulf of
Mexico (Turner & Heyman 1995).

In accordance with previous findings,
this species is found mostly in rocky and
hard substrata, covered with a veneer of
sand or in deep sand in the case of Gulf of
Mexico samples. The station from which
specimens were collected in the Colombian
Caribbean was one of the most diverse
among all those sampled in the INVEMAR
project and produced a large number of
fishes and invertebrates characteristic of
hard substrata or reef bottoms. Among
these, 39 species of echinoderms were
found, collected from gorgonians (e.g., As-
trocnida isidis, Asteroporpa annulata, As-
teroschema cf. laeve, A. oligactes) and oth-
er substrata (e.g., Nemaster rubiginosus,
Endoxocrinus parrae, Ophioderma appres-
sum, Ophiothrix suensonii). The most di-
verse taxa, in descending order, were echi-
noderms, cnidarians, fishes, decapods, crus-

543

taceans, and mollusks; also many individ-
uals and possibly many species of sponges
were collected, but they are not yet identi-
fied (Reyes et al. 2004). Turner & Heyman
(1995) also reported a diverse habitat, with
cnidarians, echinoderms, sponges, and crus-
taceans at one station and crustaceans, cni-
darians, echinoderms, and sponges at the
other station.

The world distribution of O. disacanthus
is interesting because its presence in the
Gulf of Mexico and the Colombian Carib-
bean might be a relict of a wider distribu-
tion in the Tethys Sea, as in the case of the
genus Quadratus, a myxine fish that was
considered restricted to the Western Pacific,
but that was collected also during this pro-
ject in the Western Atlantic (Mok et al.
2001).

Acknowledgments

We extend our gratitude to Dr. Richard
Turner for his helpful suggestions and thor-
ough revision of a draft of this note. This
study was made possible by the financial
support of the projects INVEMAR FON-
AM, cod. 001065, INVEMAR COLCIEN-
CIAS 2105-09-10401 and INVEMAR-
COLCIENCIAS, cod. 210509-11248. This
is contribution #849 from INVEMAR.

Literature Cited

Clark, H. L. 1911. North Pacific ophiurans in the col-
lection of the United States National Muse-
um.—Bulletin of the United States National
Museum 75:1—302.

Ljungman, A. 1867. Ophiuroidea viventia huc usque
cognita enumerat.—Ofversigt af Kongliga Ve-
tenskaps-Akademiens Forhandlingar 1866 (9):
303-336.

Mok, H., L. M. Saavedra-Diaz, & A. Acero. 2001.
Two new species of Eptatretus and Quadratus
(Myxinidae, Myxiniformes) from the Caribbean
Coast of Colombia.—Copeia 2001(4):1026—
1033.

Reyes, J., N. Santodomingo, A. Gracia, G. Borrero-
Pérez, G. Navas, L. M. Mejia-Ladino, A. Ber-
mudez, & M. Benavides. 2004. Southern Carib-
bean azooxanthellate coral communities off Co-
lombia. In A. Freiwald & M. J. Roberts, eds.,
Cold-water Corals and Ecosystems. Springer

544 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Printing House, Heidelberg, Germany (in specimens from Japanese waters and new ma-
press). terial from the Gulf of Mexico.—Proceedings

Turner, R. L., & R. M. Heyman. 1995. Rediagnosis of of the Biological Society of Washington 108:
the brittlestar genus Ophiosyzygus and notes on 292-297.

its type species O. disacanthus (Echinodermata:
Ophiuroidea: Ophiomyxidae) based on the type Associate Editor: Stephen L. Gardiner

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(4):545—550. 2004.

Sunagocia sainsburyi, a new flathead fish (Scorpaeniformes:
Platycephalidae) from northwestern Australia

Leslie W. Knapp and Hisashi Imamura

(LWK) Department of Zoology, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560-0159, U.S.A.; e-mail: knappl @si.edu;
(HI) Hokkaido University Museum, Faculty of Fisheries, Hokkaido University, 3-1-1 Minato-cho,
Hakodate, Hokkaido 041-8611, Japan, e-mail: imamura@museum.hukudai.ac.jp

Abstract.—Based on two specimens taken by bottom trawl from northwest-
ern Australia, Sunagocia sainsburyi differs from its congeners in having: 4—5
preorbital spines; 5 total gill rakers on first arch; a bony expansion of suborbital
ridge base on cheek bearing 1—2 rows of small spines; and no papillae on upper
surface of eye. It also tends to have more spines on the ethmoid and on the
supraorbital and suborbital ridges. A table compares features of the new species
to the other three species currently included in the genus Sunagocia.

Imamura (1996) erected the genus Eu-
rycephalus for three species formerly
placed in the genus Thysanophrys Ogilby,
1898; E. arenicola (Schultz, 1966), E. car-
bunculus (Valenciennes in Cuvier & Valen-
ciennes, 1833), and E. otaitensis (Cuvier
(ex Parkinson) in Cuvier & Valenciennes,
1829). The primary features distinguishing
the new genus were: suborbital ridge bear-
ing four or more distinct spines; iris lappet
finger-like or branched; lateral-line scale
pores with two openings posteriorly; and
sensory tubules weakly developed or absent
from the cheek region. Recently, Imamura

(2003) learned that the name Eurycephalus
was preoccupied by the cerambycid beetle
genus Eurycephalus Gray in Cuvier & Grif-
fith, 1832 and proposed Sunagocia as a re-
placement name.

During the trawling surveys of north-
western Australia conducted by the F/V
Courageous in 1978 and by the F/V Soela
in 1980, two small specimens of an unde-
scribed species of Sunagocia were taken.
Comparisons of features distinguishing
these specimens from the other three spe-
cies of Sunagocia appear in Table 1. The
two collections of the new species represent

Table 1.—Comparison of features in species of Sunagocia (value for paratype in parentheses).

arenicola carbunculus otaitensis sainsburyi
Character n = 20 n=8 n = 21 n=2
SL (mm) 37-135 70-116 83-159 86 (97)
Total gill-rakers 6 6-7 6-7 5)
Preocular spines 1 1 1 5 (4)
Preorbital spines O, rarely 1—2 1, rarely 2 O, rarely 1 4
Maxilla reaches to just past front of anterior 4 of eye just past front of mid-eye
eye eye
Ocular flaps absent present absent absent
Labial papillae absent absent present absent

Suborbital ridge upper base smooth, scaled

smooth, scaled

smooth, scaled bony expansion with
1—2 rows of small

spines

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

546

Paratype of Sunagocia sainsburyi, CSIRO H 5856-01, Northern Australia, 97 mm SL.

Fig. 1.

> >
2»? 5,
377
RF SO?
ae >
=> ae ‘
= SSD OLY?
> Le Re
—_— >
3 4 “vt
N ay
EIEN
7TS 4
ot =
23 =>??
> pre ° 4%
Ses =
>> dd > f 2 ae
— 34 a 4
>) 2 = iL
= ‘S > rs’
QIN? ? Sy
>)
ee

10mm

Cranial spines of holotype of Sunagocia sainsburyi,

26230-007.

WAM

Fig. 2.

VOLUME 117, NUMBER 4

547

5 mm

Fig. 3.
below suborbital ridge.

the first records of the genus taken with
trawling gear, the other species having typ-
ically been taken with rotenone and SCU-
BA.

Methods

Counts and measurements were taken ac-
cording to Hubbs & Lagler (1949). Mea-
surements were made with calipers and
rounded to the nearest mm. Vertebrae were
counted from radiographs. Terminology of
head spines follows Knapp et al. (2000). In-

1mm

Fig. 4. LL scale, circa 22nd scale from right side
of holotype of Sunagocia sainsburyi.

Head of holotype of Sunagocia sainsburyi, right side, showing lack of sensory tubules on cheek

stitutional acronyms follow Leviton et al.
(1985) except for South African Institute
for Aquatic Biodiversity (SAIAB), formerly
RUSI. Standard length and head length are
abbreviated as SL and HL, and lateral-line
aclele:

Sunagocia sainsburyi, new species
Sainsbury’s flathead
Fig. 1

Holotype.-—WAM _ 26230-007, 86 mm
SL, Western Australia, 125 km NE of Port
Hedlund, 19°07’S, 119°25'’E, EV. Coura-
geous, 28 May 1978, 73-74 m, K. Sains-
bury et al.

Paratype.—CSIRO H 5856-01, 97 mm,
Northern Australia, near Darwin, 11°53’'S
131°15’E, EV. Soela, Cr. 5, Sta. 49, 6 July
1980, trawl, 20—22 m.

Other material examined.—Sunagocia
arenicola, USNM 362804 (12, 37-117
mm), Western Indian Ocean, Amirantes Is.,
D’Arros I., R/V Anton Bruun Cr. 9, 5°24'S,
53°13"E, 8 Dec. 1964, rotenone, 4—8 m,
RS-40, R. D. Suttkus et al.; SAIAB 8219
(8, 46-131), Mozambique, Pinda Reef, Bay
of Bocage, 14°10’S, Sept. 1956, M. M.
Smith. S. carbunculus, USNM 99703 (8.

548

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

CH
TS
CN
ye

3mm

Fig. 5.

70-116), Malaysia, Sabah, North Borneo,
Sandakan Bay, 2 Mar. 1908, U.S. Fisheries
Steamer Albatross, seine. S. otaitensis,
USNM 366402 (6, 106-151), Northern
Philippines, Babuyan Is., Fuga I., circa
18°51’N 121°22’E, coral and tide pools, 11
Mar. 1990, rotenone, A. Ross; USNM
366403 (5, 83-159), Central Philippines,
Negros, Apo I., 9°4.5'N 123°16.4’E, 18
May 1979, LK 79-20, rotenone, 0—2.4 m,
L. W. Knapp et al.; ROM 42303 (10, 37—
80) Indian Ocean, Chagos Arch., Peros
Banhos, Isle du Coin, 5°26’21”S
71°46'52”E’, 6 Feb. 1979, WE 79-06, ro-
tenone, O—7 m, R. Winterbottom et al.

3mm

Fig. 6. Sketch of iris lappet from right eye of ho-

lotype of Sunagocia sainsburyi.

Area around LL scales 19—27, right side of holotype of Sunagocia sainsburyi.

Diagnosis.—A species of Sunagocia
with 4—5 preorbital spines; 5 total gill rak-
ers on the first arch; a bony expansion of
the suborbital ridge upper base on cheek
bearing 1—2 rows of small spines; maxilla
reaching to below middle of eye; no papil-
lae on upper surface of eye; a series of
spines on the ethmoid and several pairs of
nasal spines (Fig. 2); and smaller, more nu-
merous spines on the supraorbital and sub-
orbital ridges. Sensory tubules are absent
from the cheek area below the suborbital
ridge (Fig. 3).

Description.—Data for holotype given,
followed by that of paratype in parentheses
when differing. Dorsal-fin damaged in holo-
type, last 1-2 spines missing, VII(IX), 11;
anal-fin rays 12; pectoral-fin rays 2 un-
branched + 14 branched + 3 unbranched
(2+13+4) = 19; pelvic fin with 1 spine and
5 rays, innermost is unbranched; caudal-fin
branched rays 8; vertebrae 27; total gill rakers
on first arch 5; pored scales in LL 52, ante-
rior 3 scales bearing a small spine; 6 rows
of scales between 2nd dorsal fin origin and
LL. Number of oblique scale rows above
LL about equal to number of LL scales. LL

VOLUME 117, NUMBER 4

9
8
@
td ® e
@
O ®

Snout Length / Interorbital Width

0 20 40 60 80
Standard Length (mm)

Fig. 7.
holotype, solid diamond).

scale pores with two openings to the exte-
rior (Fig. 4). Relationship of LL scales to
adjacent scale rows is shown in Fig. 5. Iris
lappet bears short branches with bifurcate
tips (Fig. 6). Lip margins without papillae.

Body depressed, upper body covered
with ctenoid scales, breast scales largely cy-
cloid. Interopercular flap lacking. HL 2.8
(2.9) in SL; orbit going 1.1 times in snout.
Ratios of least interorbital width into snout
length for the four species of Sunagocia ap-
pear in Fig. 7. Villiform teeth in bands on
jaws and palatines, in two separate patches
on vomer.

Top and sides of head armed with nu-
merous spines (Fig. 2). Preopercular spines
3, uppermost longest, not bearing an acces-
sory spine on base; a pair of stout nasal
spines, with 2—3 smaller spines running an-
teriorly to each; base of opercular spines

549

@ carbunculus
Mi arenicola

100 120 140 160 180

Ratios of least interorbital width into snout length for the four species of Sunagocia (S. sainsburyi

covered by scales, not bearing serrae. Sub-
orbital ridge with about 17—20 serrae.

Color observations were taken on the
paratype after it thawed, prior to preserva-
tion. Dorsum brownish, with about six
darker bands crossing back, venter whitish.
Two brown infraorbital bands and two
brown suborbital bands present. Cheek be-
low suborbital ridge with a series of brown
blotches. A brown band angling back from
anterior ethmoid to front of eye. Dorsal-fin
spines and rays bearing small dark spots;
pectoral fin with several vertical brownish
bands above, clear below; pelvic fin with
four reddish-brown bands; and caudal fin
with about four vertical dark brown bands.

Etymology.—The species is named in
honor of Keith J. Sainsbury, collector of the
holotype and other flatheads later during the
EV. Soela cruises.

550

Acknowledgments

We thank the following individuals for
the loan of specimens or other assistance:
P. R. Last and A. Graham (CSIRO); J. B.
Hutchins (WAM); R. Winterbottom and M.
Rouse (ROM); P C. Heemstra and V.
Mthombeni (SAIAB) and S. Jewett
(USNM). Thanks are also due J. Finan and
S. Raredon, USNM, for considerable tech-
nical assistance. The fine drawing of the
paratype was made by Keiko Hiratsuka
Moore.

Literature Cited

Cuvier, G., & E. Griffith. 1832. The Animal Kingdom,
Arranged in Conformity with its Organization,
Vol. 15, Insecta 2. Whittaker, London, 796 pp.

, & A. Valenciennes. 1829. Histoire Naturelle

des Poissons. EF G. Levrault, Paris, vol. 4: i—

xxvi + 2 pp. + 1-518 pp.

, & . 1833. Histoire Naturelle des Pois-
sons. EF G. Levrault, Paris, vol. 9: i—xxix + 3
pp. + 1-512 pp.

Hubbs, C. L., & K. FE Lagler. 1949. Fishes of the Great
Lakes region.—Bulletin of the Cranbrook Insti-
tute of Science 26: 186 pp.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Imamura, H. 1996. Phylogeny of the family Platyce-
phalidae and related taxa (Pisces: Scorpaenifor-
mes).—Species Diversity 1(2):123—233.

. 2003. Sunagocia, a replacement name for the
platycephalid genus Eurycephalus (Actinopter-
ygii: Percomorpha), with taxonomic comments
on the species of the genus.—Species Diversity
8(3):301—306.

Knapp, L. W., H. Imamura, & M. Sakashita. 2000.
Onigocia bimaculata, a new species of flathead
fish (Scorpaeniformes: Platycephalidae) from
the Indo-Pacific.—J. L. B. Smith Institute of
Ichthyology Special Publication No. 64:1—10.

Leviton, A. E., R. H. Gibbs, Jr, E. Heal, & C. E.
Dawson. 1985. Standards in herpetology and
ichthyology: part 1. Standard symbolic codes
for institutional resource collections in herpe-
tology and ichthyology.—Copeia 1985(3):802—
832.

Ogilby, J. O. 1898. New genera and species of fish-
es.—Proceedings of the Linnean Society of
New South Wales 23:32—41.

Schultz, L. P., E. S. Herald, E. A. Lachner, A. D. We-
lander, & L. P. Woods. 1966. Fishes of the Mar-
shall and Marianas Islands.—Bulletin of the
United States National Museum 202, Vol. 3:
45-62.

Associate Editor: Carole Baldwin

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
117(4):551—563. 2004.

A new species of Nannocharax (Characiformes: Distichodontidae)
from Cameroon, with the description of contact organs and breeding
tubercles in the genus

Richard P. Vari and Carl J. Ferraris, Jr.

(RPV) Department of Zoology, Division of Fishes, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560-0159, U.S.A., e-mail: vari.richard@nmnh.si.edu;
(CJF) Research Associate, Department of Zoology, Division of Fishes, National Museum of
Natural History, Smithsonian Institution, Washington, D.C. 20560-0159, U.S.A.,
e-mail: cferraris@msn.com

Abstract.—Nannocharax reidi, new species, is described from several local-
ities in the upper Cross River basin in Cameroon. The species possesses the
synapomorphies of the clade comprising Nannocharax and Hemigrammochar-
ax. It is assigned to Nannocharax on the basis of its possession of a completely-
pored lateral line, a feature distinguishing that questionably monophyletic ge-
nus within the clade composed of these two genera. Nannocharax reidi is
distinguished from its congeners on the basis of a combination of meristic and
morphometric features and details of pigmentation on the body. Comparative
studies revealed the presence of hook-shaped contact organs on the pectoral
fins of some species of Nannocharax and epidermal breeding tubercles on the
head, body, and fins of at least one species of the genus. These observations
represent the first reports of contact organs and breeding tubercles in African
members of the order Characiformes. Some species of Nannocharax were
found also to possess variably-developed fields of hook-shaped contact organs
on the exposed surfaces of scales of the midlateral portion of the body posterior
of the pectoral girdle. This latter feature has not been previously reported

among fishes.

Species of the African distichodontid ge-
nus Nannocharax are relatively small-sized
fishes inhabiting the Nile River and many
sub-Saharan rivers, with the greatest spe-
cies-level diversity of the genus occurring
in West Africa and the Congo River basin.
Species of Nannocharax share a number of
distinctive modifications relative to phylo-
genetically-proximate taxa including trans-
versely-flattened ventral surfaces of the
head and body, a down-turned mouth, and,
in many species, expanded pelvic and pec-
toral fins; these features apparently corre-
late with their habits of resting on, and
feeding off of, the substrate or vegetation
(e.g., Nannocharax fasciatus, see Géry
1977:89). The most recent comprehensive

treatment of Nannocharax was that of Bou-
lenger (1909:279) who discussed seven
nominal species that recent authors have as-
signed to the genus. Subsequent decades
saw the progressive descriptions of addi-
tional species of Nannocharax, resulting in
24 species being recognized in the compen-
dium of the genus by Daget & Gosse (1984:
200). Two treatments of the West African
species of Nannocharax have been pub-
lished (Daget 1961, Gosse & Coenen
1990), and recent decades have seen the de-
scription of several new species of the ge-
nus from that region (Vari & Géry 1981,
Coenen & Teugels 1989, Van den Bergh et
al. 1995). Numerous uncertainties, nonethe-
less, remain concerning the species-level

552

diversity within Nannocharax. Perhaps the
major question is whether the geographi-
cally widespread N. fasciatus, whose distri-
butional range in West Africa reportedly ex-
tends from Guinea to Gabon (Daget &
Gosse 1984:201), is a single widely-distrib-
uted species or a complex of similar spe-
cies. Reid (1989:24, 56), followed by Teu-
gels et al. (1992:43), noted that population
samples of an N. fasciatus-like form from
the upper Cross River system in Cameroon
differed from the more-typical N. fasciatus
populations from that region, but those au-
thors deferred from pursuing the question
of the identity of these samples.

Studies of the species of Nannocharax in
the lower Guinea region encompassing
Cameroon, Rio Muni, Gabon, and the
coastal portions of the Republic of Congo,
Brazzaville demonstrate that some popula-
tions of an N. fasciatus-like form from the
upper Cross River represent an undescribed
species that we describe herein. We also de-
scribe unusual modifications of the scales,
fins, and epidermis in some species of Nan-
nocharax that were discovered during our
comparative studies. These noteworthy
modifications are either elaborations of
some body scales of a form unique to the
species of Nannocharax within the Chara-
ciformes (and perhaps fishes), or are elab-
orations of the fin rays and epidermis that
were previously thought to be restricted to
New World members within that order.

Materials and Methods

Measurements are given as a percentage
of standard length (SL) except for subunits
of the head that are presented as percent-
ages of head length. Lateral-line scale
counts include all pored scales along that
series, including scales located posterior to
the hypural joint. In fin-ray counts, lower-
case Roman numerals indicate unbranched
rays, and Arabic numerals _ indicate
branched rays. The two posteriormost anal-
fin rays, which are joined at their bases,
were counted as one element. Morphomet-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ric and meristic data were taken following
the procedures outlined in Fink & Weitz-
man (1974). Counts of gill-rakers, teeth,
cteni, and branchiostegal rays were taken
from two specimens that were cleared and
counterstained following the method of
Taylor & Van Dyke (1985). Vertebral
counts were acquired via radiographs and
include the four vertebrae of the Weberian
apparatus and the terminal centrum. Insti-
tution abbreviations are: AMNH, American
Museum of Natural History, New York;
CU, Cornell University, Ithaca, New York;
MRAC, Musée Royal de |’ Afrique Centra-
le, Tervuren, Belgium; and USNM, Nation-
al Museum of Natural History, Smithsonian
Institution, Washington, D.C.

Nannocharax reidi, new species
Fig. 1

Nannocharax sp. 1, Reid, 1989:24, 56
[Cameroon, upper Cross River system].
Nannocharax sp., Teugels et al., 1992:43

[upper Cross River system].

Holotype.—USNM 304046, 62.7 mm
SL; Cameroon, Cross River system, col-
lecting points on southern Munaya River
draining northern Korup, on Basep River at
junction with Munaya River (5°49'30’N,
9°03'30"E); collected by Gordon McG.
Reid, 22 February 1988.

Paratypes.—20 specimens, 34.3—59.0
mm SL. USNM 375193, 16 specimens (2
cleared and counterstained for cartilage and
bone), 34.3-59.0 mm SL; AMNH 233622,
2 specimens, 37.3—41.6 mm SL; MRAC
A3-47-P-1-2, 2 specimens, 37.1—43.1 mm
SL; collected with holotype.

Non-type specimens examined.—19
specimens, 33.3-44.7 mm SL. USNM
375195, 3 specimens, 34.5—38.8 mm SL;
Cameroon, Manya, Cross River system,
collecting points on main Cross River,
downstream of Mamfé, Mam River, junc-
tion with Cross (3°50'30"N, 9°14’50"B).
USNM 375196, 1 specimen, 41.7 mm SL;
Cameroon, Cross River system, collecting
points on main Cross River below Mamfé

VOLUME 117, NUMBER 4

Fig. 1.
points on southern Munaya River draining northern Korup, on Basep River at junction with Munaya River
(05°49'30"N, 09°03’30"E); lateral and ventral views.

SSL 25 N, QIU SO). WSINIM S7SIOT, Z
specimens, 36.4—44.5 mm SL; Cameroon,
Cross River system, collecting points on
southern Munaya River draining northern
Korup, southern Munaya River, junction
with Cross River (5°53'N, 9°00’E). USNM
375194, 6 specimens, 36.3—36.7 mm SL;
Cameroon, Cross River system, collecting
points on southern Munaya River draining
northern Korup, on Basep River at junction
with Munaya River (5°49'30'N, 9°03'30"E);
collected with holotype. MRAC 88-053-P-
0163-0168, 6 specimens, 33.3—44.7 mm
SL; Cameroon, mainstream of Cross River,
5—15 km downstream of Mamfé (approxi-
mately 5°46’N, 9°17’E). MRAC 88-053-P-
0170, 1 specimen, 42.4 mm SL; Cameroon,
mainstream of Cross River, approximately
5 km downstream of Mamfé (approximate-
ly 5°46’N, 9°17’E).
Diagnosis.—Nannocharax reidi is distin-
guished from all congeners by the combi-
nation of: the lack of a large, dark, rounded
spot extending from the posterior portion of
the caudal p: duncle to the basal portions of
the middle c iudal-fin rays; the lack of a dis-
tinct, dark, midlateral stripe extending from
the snout at least to the rear of the caudal
peduncle; the absence of a series of very
narrow, vertical, dark bars positioned along

353)

Nannocharax reidi, holotype, USNM 304046, 62.7 mm SL; Cameroon, Cross River system, collecting

the lateral surface of the body; the location
of the origin of the dorsal fin posterior to
the vertical through the insertion of the pel-
vic fin; the possession of 47 to 49 scales
along the lateral line, 5, rarely 6, scales dor-
sal of the lateral line to the origin of the
dorsal fin, and 4 scales ventral of the lateral
line to the origin of the anal fin; and the
overall body form.

Descript on.—Morphometric values for
holotype and paratypes are presented in Ta-
ble 1. Body elongate, relatively wide trans-
versely in region from rear of head to ver-
tical through posterior terminus of base of
dorsal fin and increasingly transversely-
compressed posterior to latter region. Trans-
verse widening of anterior portion of body
proportionally more pronounced in larger
examined individuals. Ventral region of
head and body anterior to insertion of pel-
vic fin distinctly flattened; degree of flatten-
ing more pronounced in larger examined
specimens. Dorsal profile of head gently
convex from tip of snout to vertical through
posterior margin of orbit, straight or very
slightly convex from that point to posterior
limit of supraoccipital spine. Predorsal pro-
file of body slightly convex in all examined
specimens. Dorsal profile of body slightly
posteroventrally-inclined along base of dor-

554

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Morphometrics and meristics of holotype and paratypes of Nannocharax reidi, new species, n =
21. Standard length is expressed in mm; measurements | to 14 are percentages of standard length; 15 to 18 are

percentages of head length; mean includes holotype.

Holotype Paratypes Mean
Morphometrics

Standard Length 62.7 34.3-59.0
1. Snout to anal-fin origin 75.6 72.1—16.3 74.3
2. Snout to pelvic-fin insertion 40.7 37.2-41.4 39.3
3. Snout to pectoral-fin insertion 23.4 23.2—26.3 24.8
4. Snout to dorsal-fin origin 43.7 42.1-45.9 44.3
5. Dorsal-fin origin to hypural joint 54.7 53.7—-59.6 56.3
6. Dorsal-fin origin to anal-fin origin 35.4 32.3-36.7 34.3
7. Dorsal-fin origin to pelvic-fin insertion 20.2 17.9-20.0 18.8
8. Dorsal-fin origin to pectoral-fin insertion 26.2 23.9-27.1 25.7
9. Caudal-peduncle depth 10.2 9.8-10.4 10.1
10. Pectoral-fin length ADA) 23.9-27.2 25.6
11. Pelvic-fin length 25.4 22.4-25.6 23.8
12. Dorsal-fin length AN) 21.5—25.4 ABI
13. Anal-fin length 17.1 15.9-17.8 16.8
14. Head length 25.8 24.5—27.5 25.8
15. Postorbital head length 37.8 37.8-44.7 42.1
16. Snout length 34.6 31.5-35.3 33.3
17. Bony orbital diameter Die 27.1—31.9 29.9
18. Interorbital width 21.0 16.3—20.9 19.2

Meristics

Lateral-line scales 47 47-49 47.8
Scale rows between dorsal-fin origin and lateral line S) 5-6 5.1
Scale rows between anal-fin origin and lateral line 4 4 4.0
Predorsal median scales 10 10-12 11.2
Branched dorsal-fin rays 9 8-10 9.4
Branched anal-fin rays 6 6-8 7.0
Branched pelvic-fin rays 7 6-8 WM
Pectoral-fin rays 14 13-15 14.1

sal fin and slightly convex from posterior
terminus of base of fin to caudal peduncle.
Ventral profile of head straight and slightly
posteroventrally-inclined. Ventral profile of
body nearly straight along prepelvic region
and slightly convex from insertion of pelvic
fin to caudal peduncle.

Mouth slightly subterminal. Lower jaw
comparatively wide relative to condition in
many congeneric species, with width of
posterior portion of jaw equal to height of
orbit in larger specimens. Jaw teeth elon-
gate, bicuspid, and slightly expanded dis-
tally (see Daget 1961, fig. 4 for shape of
teeth in genus), with single series of func-
tional teeth in each jaw. Dentary with 5 or
6 teeth. Dentary teeth gradually decreasing

in size posteriorly with terminal tooth in se-
ries approximately one-half as long as tooth
proximate to dentary symphysis. Replace-
ment teeth on dentary arranged in single se-
ries within enlarged dentary replacement
tooth trench. Dentary lacking segment of
laterosensory canal system and movably at-
tached to lateral surface of anterodorsal sur-
face of angulo-articular. Contralateral den-
taries immovably attached syndesmotically
along medial surfaces. Premaxilla with 5 or
6 teeth of same morphology as dentary
teeth. Premaxillary teeth gradually decreas-
ing in size posteriorly with terminal tooth
in series approximately one-half as long as
tooth proximate to premaxillary symphysis.
Premaxillary replacement teeth arranged in

VOLUME 117, NUMBER 4

single row embedded in fleshy covering of
inner surface of premaxillae. Contralateral
premaxillae immovably attached syndes-
motically along medial surfaces; but with
premaxillary complex vertically mobile on
mesethmoid. Maxilla edentulous, with pos-
terior portion of bone flat, plate-like, and
extending nearly entirely under first infra-
orbital bone when mouth closed. Pupil
ovoid, with pronounced emargination of an-
terior portion of iris (see Vari & Géry 1981,
fig. 2, for illustration of this condition in
Nannocharax maculicauda). Snout and dor-
sal portions of upper lip and head covered
with small papillae-like processes in holo-
type and to lesser extent in larger paratypes.
Such papillae may represent intermediate
developmental stages of well-developed
breeding tubercles present in those regions
and elsewhere on head, body, and fins in at
least one congeneric species (see comments
on breeding tubercles under “Contact or-
gans and breeding tubercles in species of
Nannocharax’”’ below).

First gill arch with 13 or 14 gill rakers in
2 cleared and stained specimens. Bran-
chiostegal rays 4.

Scales ctenoid (sensu Johnson 1984,
Roberts 1993), with cteni formed by series
of independent ossifications positioned
along posterior margin of scale. Scales of
lateral surface of body with 23 to 28 cteni
along scale margin. Lateral-line scale series
completely pored, with last scale in series
horizontally elongate. Body scales extend-
ing onto base of middle rays of caudal fin
in triangular pattern. Many smaller individ-
uals with scaleless region on median por-
tion of prepelvic region immediately pos-
terior of ventral margin of pectoral girdle.

Largest examined specimens with lateral
surface of scales in region posterior and
posterodorsal to insertion of pectoral fin
bearing some scattered, elongate, contact
organs (sensu Collette 1977). Contact or-
gans of scales most concentrated in region
proximate to posterior margin of pectoral
girdle and best developed in holotype, the
largest examined specimen. Contact organs

555

elongate, laterally-directed, and with ante-
riorly-directed distal tips. Field of contact
organs neither as dense as, nor as extensive
as, pattern of hook-shaped processes pre-
sent on that region of body in at least one
congeneric species (see “Contact organs
and breeding tubercles in Nannocharax”
below).

Dorsal-rays 11,8 to 10. Distal margin of
dorsal fin nearly straight. Anal-fin rays 11,6
to 8 or rarely ii1,7. Distal margin of anal fin
concave. Individual lepidotrichia of un-
branched anal-fin rays anteroposteriorly ex-
panded with overall form of distal portion
of fin rays somewhat club-shaped; such
rays proportionally more expanded in larger
individuals. Anterior and lateral surfaces of
first unbranched dorsal-fin ray and distal
portions of second unbranched ray envel-
oped by overlying, thick, fleshy layer in
many, but not all, specimens; fleshy cov-
ering more developed in holotype and larg-
er paratypes. Caudal fin distinctly forked.

Pectoral and pelvic fins proportionally
longer than in many congeneric species.
Pectoral-fin rays 11,13 to 15. Dorsal surface
of basal portions of second unbranched, and
first through fifth branched rays of pectoral
fin with basally-directed, hook-shaped,
bony contact organs on basal one-half to
two-thirds of rays in holotype, the largest
examined specimen. Larger male paratypes
with such contact organs developed to less-
er degree, but still obvious. Hook-shaped
processes on pectoral-fin rays apparently
limited to mature males. Unbranched pec-
toral-fin rays and distal portions of first
branched pectoral-fin ray with individual
lepidotrichia widened, more so on distal
portions of fin rays; expanded portion of fin
rays consequently somewhat club-shaped.
First unbranched pectoral-fin ray distinctly
shorter than second ray, with second un-
branched pectoral-fin ray somewhat shorter
than first branched ray; latter ray longest of
fin. Ventral and lateral surface of first un-
branched pectoral-fin ray, and distal por-
tions of second unbranched, and first
branched, pectoral-fin rays with thick,

556

fleshy covering. Fleshy layer on fin rays
thicker and extending farther basally along
rays in larger individuals, and particularly
well-developed in holotype. Tip of pectoral
fin extending distinctly beyond vertical
through insertion of pelvic fin in specimens
of all sizes.

Pelvic-fin rays 11,6 to 8. Pelvic fin with
unbranched rays and distal portion of first
branched ray with individual lepidotrichia
thickened, more so on distal portions of fin
rays that consequently have a somewhat
club-shaped form. Ventral and lateral sur-
face of first unbranched, and distal portions
of second unbranched and first branched
pelvic-fin rays with thick, fleshy covering;
fleshy layer also extending over dorsal sur-
face of distal portions of ray. Fleshy layer
on pelvic-fin rays thicker and extending fur-
ther basally along rays in larger examined
individuals. First unbranched pelvic-fin ray
distinctly shorter than second unbranched
ray; second unbranched ray somewhat
shorter than first branched ray with medial
branch of latter ray longest in fin. Tip of
longest pelvic-fin ray reaching vent in
smaller individuals, but falling slightly
short of opening in larger specimens.

Vertebrae 36 to 38 [37 in holotype].

Coloration in alcohol.—Ground colora-
tion light dusky-brown, with scattered dark
chromatophores in holotype and paratypes;
tan with fewer dark chromatophores in
some more lightly-pigmented, non-type
specimens. Lateral and dorsal surfaces of
head with irregular field of small, dark
chromatophores; chromatophore field more
concentrated on upper lip, snout, and dorsal
surface of head. Some larger individuals
with concentration of dark chromatophores
located posterior to orbit and on dorsal two-
thirds of operculum. Ventral surface of head
ranging from unpigmented to having scat-
tered, small, dark chromatophores.

Body with pattern of relatively wide, ir-
regular bars on dorsal and sometimes ven-
tral surfaces; bars often narrowing towards
midlateral region with dorsal and ventral
bars variably in contact in that region. Re-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

gions of contact between bars sometimes
appearing as irregular, darker, midlateral
patches of dark pigmentation. Smaller,
more lightly-pigmented, non-type speci-
mens with deep-lying, diffuse region of
dusky pigmentation positioned along mid-
lateral region, particularly on posterior two-
thirds of body. Ventral surface of body
ranging from pale with few, scattered,
small, dark chromatophores in some small-
er non-type specimens to dusky in holotype
and paratypes. Some darker specimens with
variably-shaped, unpigmented, typically
scaleless area on anteroventral portion of
body immediately posterior of margin of
pectoral girdle.

Dorsal fin with transverse band of dark
pigmentation located slightly dorsal of ba-
ses of fin rays and second, more distally-
positioned, wider band of dark pigmenta-
tion extending across entire fin. Wider band
of pigmentation distinctly separated from
distal margin of fin anteriorly, but angling
toward and reaching margin of fin along
distal portions of first or second branched
dorsal-fin rays. Anal fin with variably-de-
veloped patch of dark pigmentation on bas-
al portions of anterior rays and with dark
band extending from more distal portions
of anterior rays across fin to its posterior
margin. Caudal fin with patch of dark pig-
mentation situated basally and with vari-
ably-shaped and -positioned patches of dark
pigmentation located on both lobes of fin.
Pectoral-fin rays overlain dorsally by small,
dark chromatophores; dark pigmentation
most intense on lateral most fin rays, more
so distally. Pelvic fin with pattern of dark
chromatophores on distal portion of un-
branched rays and more central sections of
branched rays. Individual patches of dark
pigmentation forming broad, interrupted,
dark band across pelvic fin. Adipose fin
dark distally in smaller, more lightly-pig-
mented individuals, dark throughout in
larger specimens.

Coloration in life-—Photos of specimens
taken soon after capture show that the spe-
cies has the same pattern of dark pigmen-

VOLUME 117, NUMBER 4

tation as described above, but with the hy-
aline regions of head, body, and fins of pre-
served specimens having a rosy tint in life.

Remarks.—Vari (1979:332) noted that
the monophyly of Nannocharax had yet to
be demonstrated. That author further com-
mented that although Nannocharax and
Hemigrammocharax share a series of hy-
pothesized synapomorphies (Vari 1979:
331), the single feature that has been uti-
lized to distinguish Nannocharax from
Hemigrammocharax (the possession of a
completely- versus incompletely-pored lat-
eral line, respectively) may not serve to de-
limit monophyletic assemblages in light of
the independent reduction in the degree of
development of the lateral line in various
groups of characiforms. The possession of
various derived characters in a subset of the
species of Nannocharax and Hemigram-
mocharax to the exclusion of other mem-
bers of each genus (Vari & Géry 1981:
1082), furthermore, apparently delimits a
monophyletic lineage. That hypothesis sug-
gests that both Nannocharax and Hemi-
grammocharax as now defined are non-
monophyletic.

More recently, Coenen & Teugels (1989:
317) documented that population samples
of some nominal species within the Nan-
nocharax-Hemigrammocharax clade dem-
onstrated a continuum between distinctly-
shortened and fully-developed lateral lines.
Such continuity in the degree of develop-
ment of the poring of the lateral-line scale
series bridges the gap that purportedly dis-
tinguished Nannocharax from Hemigram-
mocharax, thereby casting further doubt on
the utility of a complete versus incomplete
lateral line as a generic delimiter for these
taxa. Above and beyond the uncertainty
about the naturalness of Nannocharax and
Hemigrammocharax, we are also encum-
bered by the limitation that the phylogenet-
ic relationships within the clade formed by
these two genera are yet to be critically ex-
amined within the context of a comprehen-
sive analysis. In the absence of such a phy-
logenetic study, we follow current taxo-

S57)

nomic practice and assign the new species
to Nannocharax on the basis of its com-
pletely-pored lateral line, in conjunction
with the possession of the synapomorphies
for the clade formed by Nannocharax and
Hemigrammocharax (see Vari 1979:331,
synapomorphies 96 to 107).

Distribution.—All examined population
samples of Nannocharax reidi were col-
lected in the upper portions of the Cross
River basin in Cameroon.

Etymology.—The specific name, reidi, is
in honor of the collector of the specimens
that served as the basis of the description
of the species, Dr. Gordon McGregor Reid,
of the North of England Zoological Society,
who first reported that these samples might
represent an undescribed form and who has
contributed broadly to our knowledge and
conservation of African freshwater fishes.

Ecology.—Nannocharax reidi was typi-
cally captured in the swiftly-flowing main-
stream portions of rivers, usually in asso-
ciation with submerged logs and branches.
In all such localities the water is clear and
brown-tinged. The new species was col-
lected together with N. fasciatus at the type
locality and at three other localities in the
upper Cross River. Those two species of
Nannocharax were sympatric with N. lati-
fasciatus at two of those sites.

Contact Organs and Breeding Tubercles in
Species of Nannocharax

Examination of other species of Nanno-
charax revealed the presence of hook-
shaped contact organs on the pectoral-fin
rays and on the scales in the region of the
body proximate to the insertion of the pec-
toral fin, along with the occurrence of
breeding tubercles on the head, body, and
fins of at least some species. The hook-
shaped contact organs on the fin rays and
breeding tubercles of these species of Nan-
nocharax had been previously reported
within the Characiformes only in some
Neotropical members of the order. The
hook-shaped contact organs on some scales

558

of the body in the genus are unknown in
any other member of the Characiformes.
Presence of hooks on pectoral-fin rays.—
In their recent analysis of the phylogenetic
relationships of various groups of Neotrop-
ical characids, Malabarba & Weitzman
(2003:73) enumerated a series of generic
and suprageneric taxa within the Characi-
formes that bear hook-shaped processes on
various combinations of the paired and un-
paired fins, including the pectoral fin. These
hook-shaped processes were termed contact
organs by some authors (e.g., Wiley & Col-
lette 1970, Collette 1977), who discussed
the distribution and possible functions of
these structures. In their commentary on
contact organs on the fins of characiforms,
Malabarba & Weitzman (2003) noted that
such bony processes on individual segments
of lepidotrichia were known to be present
in diverse Neotropical components of the
order, but were unknown in Old World
characiforms, including the family Disti-
chodontidae. Although Malabarba & Weitz-
man (2003) were correct that contact organs
on the fins had not been previously reported
for Old World characiforms, we found that
larger, apparently male, individuals of Nan-
nocharax reidi have a series of hook-
shaped, distally slightly anteriorly-bent
bony processes arranged in a single series
along the dorsal surface of the basal one-
half to two-thirds of the medial rays of the
pectoral fin. As is the case with many Neo-
tropical characids, each lepidotrichium of
the pectoral-fin rays with a hook-shaped
contact organ bears a single such process.
The extent of the field of hook-shaped
contact organs on the pectoral-fin rays dif-
fers both among specimens of N. reidi and
between species of Nannocharax. In N. rei-
di, the hook-shaped processes on the dorsal
surface of the pectoral-fin rays are more
highly-developed in larger individuals, but
even at their maximum observed degree of
development these structures are limited to
the basal one-half to two-thirds of the sec-
ond unbranched and first through fifth
branched pectoral-fin rays. Mature males of

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

N. rubrolabiatus (MRAC 95-022-P-001-
007), in contrast, have hook-shaped contact
organs on a greater number of fin rays (sec-
ond unbranched through eighth branched)
and have these processes nearly to the distal
tips of the fin rays. The presence of these
hook-shaped contact organs on the pectoral
fin in specimens of N. rubrolabiatus is cor-
related with other apparently breeding-as-
sociated modifications of the scales and fins
(the presence of hook-shaped contact or-
gans on the lateral surface of the scales in
the region medial to the pectoral fin and the
possession of epidermal breeding tubercles
distributed over the head, body, and fins;
see discussion of next two characters). This
correlation of apparently sexually-dimor-
phic features in conjunction with the rela-
tively few examined specimens of the spe-
cies of Nannocharax that demonstrate these
modifications indicate that the hook-shaped
contact organs on the pectoral-fin rays of N.
reidi and N. rubrolabiatus may be restricted
to mature males only during the height of
the breeding season.

Hook-shaped processes on scales.—TYhe
holotype and larger male paratypes of Nan-
nocharax reidi possess a form of contact
organ (sensu Wiley & Collette 1970, Col-
lette 1977) involving an elaboration of the
scales in the region of the body medial to
the adpressed pectoral fin that is apparently
unique not only within the Characiformes,
but perhaps throughout bony fishes. The
typical form of the scales among members
of the Distichodontidae (sensu Vari 1979)
is a laterally-unelaborated, relatively flat os-
sification with the posterior margin of the
main body of each scale bearing a series of
smaller, independent ossifications (see Vari
1979, figs. 38b and c), that form a distinct-
ly-serrate posterior margin to the scale.
These independent ossifications, which con-
stitute true cteni (sensu Johnson 1984, Rob-
erts 1993:70), vary both in number and
form across the members of the Disticho-
dontidae, but such elaborations, nonethe-
less, are nearly invariably limited to the
posterior margin of the scale. The single ex-

VOLUME 117, NUMBER 4

559

Fig. 2. Nannocharax rubrolabiatus, MRAC 95-022-P-001-007, 56.1 mm SL; showing breeding tubercles on
the head, anterior two-thirds of the body, and dorsal, pectoral, and pelvic fins. Arrow indicates the midlateral
region of the body lacking breeding tubercles, but with hook-shaped contact organs on the lateral surface of the

scales.

ception to that generality that we have dis-
covered, involves the form of the scales on
the portion of the body medial to the pec-
toral fin in some species of Nannocharax.
In larger specimens of N. reidi the scales of
the portion of the body medial to the basal
portion of the pectoral fin (see description
above), particularly those scales in the re-
gion of the body immediately dorsal to the
posteriorly-directed process of the cleith-
rum, have the cteni along their posterior
margins complemented by hook-shaped
processes arising from the lateral surface of
the scales. These scale processes have the
form of moderately elongate spines with
slightly anteriorly-directed distal hooks. Al-
though such processes are obvious in the
larger examined specimens of WN. reidi, they
nonetheless are somewhat scattered across,
and fail to completely cover, the lateral sur-
face of the involved scales.

A dramatically greater degree of devel-
opment of such contact organs in the region

of the body posterior to the insertion of the
pectoral fin characterizes mature males of
Nannocharax rubrolabiatus (Fig. 2). Con-
trary to the situation in all other examined
distichodontids, males of N. rubrolabiatus
lack distinct cteni along the posterior mar-
gins of the scales on the anterior portion of
the midlateral surface of the body. More
strikingly, the specimens in this population
sample have the lateral surface of the scale
variously covered by fields of laterally-di-
rected, elongate, hook-shaped contact or-
gans with anteriorly-curved distal tips.
Continuity between the fields of hook-
shaped processes of adjoining scales varies
across the portion of the body with such
lateral elaborations of the scales. Those
scales positioned closer to the posterior
margin of the pectoral girdle have patches
of contact organs that together with those
of adjoining scales form a nearly uninter-
rupted, brush-like expanse continuing ap-
proximately five scales posteriorly from the

560

posterior margin of the pectoral girdle and
extending dorsally to the horizontal running
through the dorsal margin of the opercular
opening. Farther posteriorly, the hook-
shaped processes on the lateral surface of
the scales are restricted to the posterior one-
half of the exposed portion of the scale and,
thus, form discrete patches of such contact
organs, with these patches distinctly sepa-
rated from each other. These posteriorly-po-
sitioned scales with separate patches of con-
tact organs on their lateral surfaces also dif-
fer from the more anterior scales character-
ized by the possession of such processes in
retaining independent, variably posteriorly-
directed cteni along at least a portion of the
posterior margin of the scale. Such cteni
are, however, often somewhat more later-
ally-directed than are the homologous os-
sifications in other members of the Disti-
chodontidae.

We are unaware of any laterally-posi-
tioned, hook-shaped contact organs of a
comparable morphology on the body scales,
elsewhere either within the order Characi-
formes or among other groups of fishes.
The only other report of an African fresh-
water fish with laterally-directed hook-
shaped processes on the scales involves the
gonorhynchiform Phractolaemus ansorgéi,
an ostariophysan that is phylogenetically
distant from the Characiformes. Phracto-
laemus differs significantly from Nanno-
charax in the distribution, morphology, and
number of such hook-shaped processes per
scale (see Thys van den Audenaerde 1961a,
fig. 2; 1961b, fig. 2) and the elaborations of
the scales in the two genera, thus, are ap-
propriately considered to be non-homolo-
gous. As a consequence of their apparent
unique morphology, the presence of the
dense patches of hook-shaped processes on
the anterior portion of the midlateral scales
of Nannocharax is a likely synapomorphy
for at least a subunit of that genus, albeit
one perhaps restricted to fully mature, sex-
ually-active males during the height of the
breeding season.

Breeding tubercles.—The presence of

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF W ASHINGTON

epidermal breeding tubercles has been re-
ported in a number of New World members
of the Characiformes including the families
Characidae, Parodontidae, and Lebiasinidae
(Wiley & Collette 1970:164—167, Collette
1977:236—241), but not within any of the
African families within that order, an ap-
parent absence that included the Disticho-
dontidae. One series of Nannocharax rub-
rolabiatus (MRAC 95-022-P-001-007) ex-
amined during this study has, however, very
well-developed epidermal breeding tuber-
cles distributed over the head, body and fins
(Fig. 2). The degree of development of the
tubercles correlates somewhat, albeit not
absolutely, with the size of the specimens.
The smallest specimen in the lot (45.5 mm
SL) has both fewer tubercles than most of
the larger conspecific individuals captured
with it and, furthermore, those tubercles are
proportionally less-developed than those in
larger specimens. In larger, apparently
male, individuals of N. rubrolabiatus, the
breeding tubercles are broadly distributed in
large numbers across the snout and the dor-
sal and lateral surfaces of the head (Fig. 2).
On the ventral surface of the head, the tu-
bercles are arranged in discrete rows along
the ventral surfaces of the branchiostegal
rays. Scales on the surface of the body have
one to four tubercles, other than those
scales medial to the pectoral fin whose sur-
faces are covered with the hook-shaped
contact organs (described in the previous
section and indicated by white arrow of Fig.
2). When present, the tubercles on the
scales are positioned toward the posterior
margin of the scale, and when three or four
tubercles occur on an individual scale, these
structures are arranged in an arch parallel-
ing the posterior margin of the scale. The
size and number of tubercles tend to be re-
duced on the scales of the ventrolateral por-
tion of the body. An extensive series of tu-
bercles occurs, however, on scales of the
prepelvic region of the body, with a less
concentrated field of tubercles present in
the region from the insertion of the pelvic
fin to the origin of the anal fin.

VOLUME 117, NUMBER 4

Breeding tubercles are present on all fins
of Nannocharax rubrolabiatus with the ex-
ception of the adipose fin. The tubercles on
the caudal fin are less developed than those
on the remaining fins, being apparent solely
as small, raised areas along the basal and
middle portions of the caudal-fin rays. Tu-
bercles are present on all of the dorsal-fin
rays with the exception of the first un-
branched and last branched rays. At their
maximum degree of development, such
breeding tubercles extend along nearly the
entire length of each dorsal-fin ray. Some
larger examined specimens of N. rubrola-
biatus have indications of poorly-developed
breeding tubercles on the basal portions of
the second unbranched anal-fin rays, with
better-developed tubercles present on all but
the terminal branched anal-fin ray. The pec-
toral fin has tubercles on the dorsal surface
of the unbranched rays, but tubercles are
absent on the portions of the second un-
branched through eighth branched rays with
anteriorly-directed, hook-shaped contact or-
gans. The ventral surface of the pectoral fin
has at most a few tubercles distributed
along the unbranched rays, but such struc-
tures are completely absent in some indi-
viduals. Variably-developed series of tuber-
cles extend along the length of the ventral
surfaces of each of the branched pectoral-
fin rays. The pelvic fin has a series of tu-
bercles arranged along the dorsal surface of
the branched rays, and along the ventral
surfaces of the last unbranched fin ray and
all of the branched fin rays with the excep-
tion of the medialmost branched ray.

Our comparative studies failed to reveal
any comparably well-developed breeding
tubercles in the other examined species of
Nannocharax. Larger examined specimens
of N. reidi do, however, have a pattern of
small, papillae-like processes on the upper
lip, snout, and dorsal surfaces of the head
that have an arrangement comparable to the
pattern of the breeding tubercles that occur
in those regions in most examined speci-
mens of N. rubrolabiatus. It will be nec-
essary to examine additional population

561

samples of N. reidi captured during the
height of the breeding season in order to
determine whether the papillae-like pro-
cesses present in that species would devel-
op into the distinctly larger breeding tuber-
cles that typify the examined sample of UN.
rubrolabiatus. Broader comparative studies
would possibly also yield insight in the
range of the distribution of breeding tuber-
cles across the species of Nannocharax.

Comparative material examined.—Nan-
nocharax altus: MRAC 78-22-P-801-804, 4
specimens, Republic of the Congo, Mayala,
Niola Creek.

Nannocharax fasciatus: USNM 303754,
5 specimens; USNM 303756, 2 specimens;
USNM 303811, 1 specimen; USNM
303847, 3 specimens; USNM 303867, 2
specimens; USNM 303908, 4 specimens;
USNM 303995, 2 specimens; USNM
304081, 3 specimens; USNM 375192, 5
specimens; Cameroon, upper Cross River
system.

Nannocharax intermedius: CU 80570, 2
specimens, Gabon, Motoboi Village, Kine-
né Creek; CU 90276, 3 specimens, Gabon,
Okolville; MRAC 91-79-P-202-206, 4
specimens, Gabon, Riviere Loukénini;
MRAC A2-006-P-0826-0828, 3 specimens,
Gabon, Ivindo basin, Balé Creek.

Nannocharax maculicauda: USNM
224524, 3 paratypes; Gabon, upper Ivindo
River (1°20’N, 13°12’E); CU 80621, 1
specimen, Gabon, Woleu-Ntem, Ngomo
River (1°42'N, 11°38’E).

Nannocharax parvus: CU 80148, 19
specimens, Gabon (0°34’S, 10°12’E). CU
80163, 1 specimen; CU 80185, 2 speci-
mens; CU 80184, 3 specimens; Gabon, Bi-
roundou Creek (2°13’S, 11°28’E). CU
80191, 1 specimen, Gabon, Mimboumbou
Creek, near Franceville (1°38’S, 13°31’E).
CU 80279, 5 specimens, Gabon, Okolo-
ville. CU 80607, 6 specimens, Gabon,
stream at Okolville (1°29’S, 13°31’E).

Nannocharax rubrolabiatus: MRAC 95-
22-P-001-007, 7 specimens, Cameroon,
Sanaga River basin, Mi River (6°12'N,
14°23'E).

562

Acknowledgments

Research associated with this project was
supported by the Herbert R. and Evelyn
Axelrod Chair in Systematic Ichthyology in
the Division of Fishes of the National Mu-
seum of Natural History, Smithsonian In-
stitution. We thank Melanie L. J. Stiassny,
Scott A. Schaefer, Barbara Brown, and
Radford Arrindell (AMNH), John Friel
(CU), and Emmanuel Vreven and the late
Guy Teugels (MRAC) for the loan of spec-
imens and other assistance. Assistance at
USNM was provided by David Smith and
in particular Sandra Raredon who also pre-
pared Figs. 1 and 2. Gordon McG. Reid,
North of England Zoological Society, pro-
vided information on the collecting locali-
ties of the type-series and coloration of re-
cently captured specimens. The paper ben-
efitted from the comments and suggestions
of Thomas A. Munroe.

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Associate Editor: Edward O. Murdy

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(4):564—5S74. 2004.

Rhamdia guasarensis (Siluriformes: Heptapteridae), a new species of
cave catfish from the Sierra de Perija, northwestern Venezuela

Carlos DoNascimiento, Francisco Provenzano, and John G. Lundberg

(CDN) Seccién de Ictiologia, Museo de Historia Natural La Salle, Fundacion La Salle de

Ciencias Naturales, Apdo. 1930 Caracas 1010-A, Venezuela,
carlos.donascimiento @ fundacionlasalle.org.ve;

(FP) Laboratorio de Biosistematica de Peces, Instituto de Zoologia Tropical, Universidad Central

de Venezuela, Apdo. 47058 Caracas 1041-A, Venezuela, fprovenz @strix.ciens.ucv.ve;

(JGL) Department of Ichthyology, The Academy of Natural Sciences, 1900 Benjamin Franklin

Parkway, Philadelphia, Pennsylvania 19103, USA, lundberg @acnatsci.org

Abstract.—Rhamdia guasarensis n. sp. is described from subterranean waters
in the Rio Guasare drainage of northwestern Venezuela. The new species is
distinguished from congeners by its concave head profile; medially sutured
frontal bones; small, circular vestige of the anterior cranial fontanelle; and
troglomorphic characters such as absence of eyes and pigmentation, wide ce-
phalic laterosensory pores, and wide fossae of preoperculomandibular sensory
canal in preopercle and dentary. Cave catfish diversity in the Sierra de Perija
region of Venezuela is reviewed and compared to cave catfish diversity else-
where in South America.

Resumen.—Se describe Rhamdia guasarensis sp. n. proveniente de aguas
subterraneas de la cuenca del Rio Guasare en el noroccidente de Venezuela.
La nueva especie se diferencia de las restantes especies que conforman el
género por su perfil dorsal de la cabeza concavo; huesos frontales suturados
medialmente; fontanela craneal anterior reducida a un pequeno foramen cir-
cular; y caracteres troglomorficos tales como ausencia de ojos y pigmentacion,
poros cefalicos latero sensoriales anchos, fosas ensanchadas del canal sensorial
preoperculomandibular en el preopérculo y dentario. La diversidad de bagres
cavernicolas de la Sierra de Perija es revisada y comparada con la diversidad

de bagres cavernicolas de otras regiones de Suramérica.

The family Heptapteridae has invaded
and adapted to hypogean waters multiple
times. Among Neotropical catfish families,
heptapterids have the greatest diversity of
truly troglobitic taxa: Phreatobius cister-
narum, Pimelodella kronei, Rhamdia lalu-
chensis, Rhamdia laticauda typhla, Rham-
dia macuspanensis, Rhamdia quelen urichi,
Rhamdia redelli, and Rhamdia zongolicen-
sis. Trajano & Bockmann (2000) described
the ecology and behavior of Taunayia sp.,
a troglobitic catfish, inhabiting caves of
northeastern Brazil, but the species has not
been formally named. Pimelodella spelea

Trajano, Reis & Bichuette, 2004 is a re-
cently described troglophile without
marked specializations for hypogean life.
Taxonomic practice has shifted away from
assigning supra-specific rank to cave-dwell-
ing fishes solely on account of their trog-
lobitic adaptations. Among Heptapteridae,
the nominal monotypic genera Caecorham-
dia, Caecorhamdella, and Typhlobagrus
have long been treated as synonyms of
Rhamdia and Pimelodella respectively. Fur-
thermore, Silfvergrip (1996) synonymized
all cave populations of Rhamdia described
as separate species with R. quelen or R.

VOLUME 117, NUMBER 4

laticauda, both wide-ranging epigean spe-
cies.

In this paper we describe a new troglob-
itic heptapterid species in the genus Rham-
dia. Our placement of the new species is
more a matter of convenience than firm
phylogenetic resolution. Rhamdia is taxo-
nomically complex. In the latest revision of
the genus, Silfvergrip (1996) consolidated
its approximately 100 nominal species into
eight and he described three new species.
In 1998, Weber & Wilkens described the
blind species R. macuspanensis, and in
2003, Weber et al. described Rhamdia lal-
uchensis, another troglobitic species from
Mexico. In the most thorough phylogenetic
study of Heptapteridae to date, Bockmann
(1998) concluded that Rhamdia is non-
monophyletic but he did not attempt to re-
solve the genus into phylogenetically di-
agnosable units. As it stands, Rhamdia is a
non-monophyletic assemblage of common
fishes with an immense geographic distri-
bution in South and Middle America from
the lower Parana Basin in Argentina to cen-
tral México.

The new species, from a cave in the Si-
erra de Perija region of northwestern Ve-
nezuela, is distinct both in its typical trog-
lobitic specializations and other apomorph-
ic features, but overall it is most similar to
other Rhamdia. Discovering the relation-
ships of the new species and, more gener-
ally, resolving the relationships of Rhamdia
species are major problems quite beyond
our present scope. Our immediate concern
is to name and describe this previously un-
seen species that has a highly restricted dis-
tribution in a marginal and potentially frag-
ile habitat. We comment also on the sub-
terranean catfish fauna of the Perija region.

Material and Methods

Morphometric measurements follow the
criteria set out by Lundberg & McDade
(1986) and Bockmann (1994). Terminology
of cephalic laterosensory canals and
branches follows Arratia & Huaquin’s de-

565

scription of Diplomystes and Nematogenys
(1995). However, our numbering of sensory
pores in Rhamdia reflects anteroposterior or
mesiolateral pore order, and does not imply
individual homologies of pores among cat-
fishes. All measurements were made on the
left side of the specimens using a Mitutoyo
digital, needlepoint caliper at a precision of
0.1 mm. For osteological observation one
paratype (101.1 mm SL) was cleared and
stained using the method of Taylor & Van
Dyke (1985). A second paratype (93.4 mm
SL) was radiographed. Only these two
specimens were used for counts of verte-
brae, branchiostegal rays, ribs, and ptery-
giophores. The vertebral count includes the
first five vertebrae incorporated into the
Weberian apparatus whereas the compound
caudal centrum is counted as one. Institu-
tional abbreviations follow Leviton et al.
(1985). Other abbreviations are: SL—stan-
dard length, HL—head length, CS—cleared
and stained skeletal preparation, alc—
whole specimen preserved in alcohol.

Rhamdia guasarensis, new species
Figs. 1-4

Holotype.—MBUCV-V-29604: 106.8
mm SL; Surgencia del Tigre at 2.5 km W
of Cerro Yolanda, Rio Guasare basin, Sierra
de Perija, Estado Zulia, Venezuela
(10°52’53"N, 72°30'03”W). Elevation 200
m asl; collected by J. Lagarde, 3 April
1999.

Paratypes.—All collected with the ho-
lotype: MBUCV-V-29622, two specimens,
87.2—101.1 mm SL, the second cleared and
stained; ANSP 179878, one specimen, 93.4
mm SL.

Diagnosis.—Rhamdia guasarensis 1s dis-
tinguished from congeneric species by two
characters: dorsal profile of head concave
(Fig. 1, vs. convex or straight); and frontal
bones broadly sutured to each other anterior
to small, circular remnant foramen of an-
terior cranial fontanelle that is anteriorly ad-
jacent to epiphyseal bar (Fig. 2, vs. frontals
separated by anterior fontanelle -widely

566

Fig. 1.
of head; C, ventral view of head.

open from mesethmoid to epiphyseal bar).
Rhamdia guasarensis differs from all epi-
gean Rhamdia by the following troglo-
morphic characters: absence of eyes, com-
plete depigmentation, widened cutaneous
pores of the cephalic laterosensory system,
preoperculomandibular sensory canal form-
ing wide fossae in the dentary and preo-
percle (Fig. 3, vs. narrow pores and canals).

In addition to these characteristics, R.
guasarensis can be distinguished from other
species of the genus by the following com-
bination of characters: pectoral fins with a
spine and ten branched rays (vs. modally

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

i
i
j
i
i
if
i
hy
ie
I
|

Rhamdia guasarensis. Holotype MBUCV-V-29604, 106.8 mm SL. A, lateral view; B, dorsal view

eight or nine soft rays in other species, data
from Silfvergrip 1996); both lobes of the
caudal fin pointed (vs. at least one lobe
rounded); caudal skeleton with three hy-
pural plates, PH; 1 + 2; 3 + 4 + 5 Ws.
modally four PH; 1 + 2; 3 + 4; 5 in the
other species with the exception of R. lau-
kidi and R. jequitinhonha, see Silfvergrip
1996).

Description.—Morphometric data are
presented in Table 1. Body elongate, strong-
ly depressed anteriorly and gradually more
compressed from origin of pectoral fins to
caudal peduncle. Shape approximately tri-

VOLUME 117, NUMBER 4

Fig. 2. Rhamdia guasarensis. Skull roof illustrat-
ing reduction of the anterior cranial fontanelle and
midline contact of frontal bones. MBUCV-V-29622,
101.1 mm SL. Abbreviations: ACE anterior cranial
fontanelle; FR, frontal bone contact on midline; LET,
lateral ethmoid; MES, mesethmoid; PCE posterior cra-
nial fontanelle; PT, pterotic; SPH, sphenotic.

angular in transverse section at dorsal-fin
origin. Dorsal profile sinusoidal anterior to
dorsal fin, then approximately straight to
middle of adipose fin, then slightly concave
along caudal peduncle. Ventral profile near-
ly straight to anal-fin origin, then slightly
concave posteriorly.

Head depressed, its dorsal profile con-
cave, its lateral and ventral profiles nearly
straight. Mouth terminal, upper jaw slightly
in advance of lower jaw. Rictal folds little
developed. Upper and lower lips with weak
sulci, slightly evident in holotype, forming
single labial fold. Premaxillaries with single
band of diminutive teeth, arranged in ten
irregular tooth rows, the posterolateral cor-
ners rounded, not produced. Dentition of

567

Fig. 3.
culomandibular laterosensory canal and associated fo-
ramina. MBUCV-V-29622, 101.1 mm SL. Abbrevia-
tions: ANG, anguloarticular; D, dentary; HYO, hy-
omandibula; MPT, metapterygoid; OP, opercle; POP,
preopercle; Q, quadrate.

Rhamdia guasarensis. Enlarged preoper-

lower jaw similar to premaxillary teeth, in
six irregular tooth rows. Palatine and vomer
edentulous. Maxillary barbels long, extend-
ing beyond base of pelvic fins. Mental bar-
bels relatively short, inner mentals scarcely
reaching posterior border of branchial
membrane; outer mentals surpass pectoral
fin bases. Inner mental barbel bases inserted
slightly in advance of outer mental barbel
bases. Anterior nares tubular, near border of
snout. Posterior nares with elongated orific-
es, bounded anterolaterally by membrane of
fine skin. Internarial length less than width
between posterior nares. Eyes completely
absent. Branchial membranes overlapping
medially; united to isthmus only anteriorly.

Cephalic lateralis sensory system with
paired supraorbital (SO), infraorbital (IO),
preopercular (POP), mandibular (MA), otic
(OT), and post-otic (POT) canals, without
tubular commissure connecting supraorbital
canals. Sensory pores simple, not branched
and multiple. SO canal with six pores:
SO1—SO3 associated with nasal bone, SO1
medially adjacent to anterior naris, wide
and delimited by membrane of fine skin,
SO2 between anterior and posterior nares,
slightly closer to first, SO3 posteromedially
near posterior naris. SO4 near dorsal mid-
line at end of short medial tube and separate
from its counterpart of opposite side. SOS
lateral to its canal midway between SO4
and union of SO and IO canals. SO6 medial
to its canal a little posterior to union of SO
and IO canals.

568

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1—Measurement data for the type series of Rhamdia guasarensis. Measurement | expressed in mm.
Proportional measurements expressed as thousandths of standard length (2-19; 26—28) or head length (20—25).

Holotype
MBUCYV-V-29604

1. Standard length 106.8
2. Total length 1057
3. Body depth 162
4. Body width 171
5. Predorsal length 351
6. Preanal length 645
7. Prepelvic length 462
8. Preadipose length IS
9. Caudal peduncle length 211
10. Caudal peduncle depth 97
11. Dorsal fin spine length 85
12. Length of first branched dorsal fin ray 172
13. Dorsal fin base 115
14. Adipose fin length 387
15. Dorsal fin to adipose fin 99
16. Anal fin base 141
17. Pectoral fin spine length 121
18. Length of first branched pectoral fin ray 204
19. Pelvic fin length 169
20. Head length 259
21. Head width 661
22. Head depth 550
23. Internarial length 137
24. Anterior internarial width 173
25. Posterior internarial width 156
26. Maxillary barbel length 541
27. Outer mental barbel length 205
28. Inner mental barbel length 98

IO canal with four pores; [O1—3 wide
like SO1. IO1 posterior to anterior nostril;
[O02 emerges dorsal to groove for maxillary
barbel, posterior to base of barbel; IO3 near
point where IO canal curves dorsally; 104
at tip of short posterior tube near union with
SO canal. Holotype and one paratype (87.2
mm SL) have different single supernumer-
ary IO pores; extra pore of holotype from
left canal between the IO2 and IO3; extra
pore of paratype from right IO canal be-
tween IO3 and [O04.

POP canal with four pores; MA canal
with seven pores; all except MAI and
POP4 originate from much enlarged cavi-
ties in dentary and preopercular bones.
MAT in mental position near to midventral
line at tip of its branch from lower jaw sym-
physis.

Paratype
Paratype MBUCV-V-29622 Paratype
MBUCV-V-29622 (CS) ANSP 179878
87.2 101.1 93.4
1093 1083
178 167
180 168 173
367 352 359
653 659 656
458 490
545 530
210 214 224
104 105
86 108
176 183 187
121 104 113
411 432 394
2 68
127 127 143
111 123
197 5 216
153 159 169
262 256 276
645 631 633
S22 538 528
144 147
169 187
152 165
610 557 578
262 258 262
121 108 121

POT canal with two pores, POT1 over
pterotic dorsal to gill opening; POT2 dorsal
to supracleithrum and above main lateralis
canal at level of first pore. First pore of la-
teralis canal at end of ventral branch dorsal
to postcleithral process. Several following
pores also at tips of short postero-ventral
branches. Lateral line canal complete to
base of middle upper-lobe caudal rays.

Dorsal fin with a spinelet, spine and six
branched rays; its margin rounded. Dorsal
spine weakly developed, only its basal part
rigid and unsegmented; dentations diminu-
tive and scarcely visible, limited to basal
part of anterior margin. The distal two-
thirds of dorsal spine flexible and obliquely
segmented. Adipose fin long and low, its
origin near tip of depressed dorsal fin, and
extending posteriorly to approximately 80%

VOLUME 117, NUMBER 4

=S
x
au et
7

5 mm

Fig. 4. Rhamdia guasarensis. Pectoral spine in
dorsal view. Holotype MBUCV-V-29604, 106.8 mm
SL.

of caudal peduncle length; posterior end of
adipose fin adnate to caudal peduncle with-
out a free fleshly tab. Caudal fin deeply
forked; both caudal lobes pointed; upper
lobe slightly longer than lower; membrane
uniting innermost caudal rays complete.
Principal caudal rays 1,7-8,1, except 1,7-7,1
in one paratype. Anal fin with 12 rays, an-
teriormost two or three rays simple, others
branched; its margin rounded.

Pectoral fins with a spine and ten
branched rays. Pectoral spine (Fig. 4) with
weak dentations proximally on anterior
margin; distal half of spine flexible and
obliquely segmented. First branched pec-
toral-fin ray longest, posterior branched
rays diminishing in length. Postcleithral
process short, sharp. Pelvic fins with one
simple ray and five branched rays, its origin
posterior to end of dorsal fin.

Skull roof (Fig. 2) with anterior cranial
fontanelle extremely reduced to a small cir-
cular foramen located in front of epiphyseal
bar; mesethmoid posteriorly lacking con-
cave notch of fontanelle, and frontals meet-
ing medially along most of their length.
Posterior cranial fontanelle reduced to oval
foramen near center of supraoccipital. Su-
praoccipital process short, its length about
equal to length of supraoccipital body.

Anus and urogenital papillae separated,
anus located equidistant between medial
edge of pelvic-fin base and urogenital pa-
pilla, approximately at midlength along pel-
vic fins; urogenital papilla conspicuous and
elongated, located closer to base of anal fin
than to base of pelvic fin.

Total vertebrae 40—42; neural spines of
vertebrae 6—10 bifid; hemal arch closed in
vertebra 12 or 13, first hemal spine on ver-
tebra 14, 15 or 16; eight pairs of ribs borne
on vertebrae 6—13. Seven dorsal-fin ptery-

569

72W

Rio Guasare

Lago de
Maracaibo

Fig. 5.

The Lago de Maracaibo—Sierra de Perija
region, Venezuela, showing type locality (star) of
Rhamdia guasarensis. Map based on shaded relief im-
age PIA03388, Shuttle Radar Topography Mission,

National
(NASA).

Aeronautics and Space Administration

giophores preceded by small supraneural;
first dorsal-fin pterygiophore inserted be-
tween rami of neural spine of fourth ver-
tebra. Eleven anal-fin pterygiophores, first
inserting posterior to hemal spine of verte-
bra 21. Caudal skeleton with three hypural
plates: rectangular parhypural; triangular
hypurals 1 + 2; triangular hypurals 3 + 4
sD

Color in alcohol.—Body and fins com-
pletely depigmented; most of skin, rayed-
and adipose-fin membranes hyaline and
translucent; musculature appearing yellow-
ish, particularly jaw adductors and dorsal
trunk myomeres; parts of head and fin bases
whitish.

Distribution and habitat.—R. guasaren-
sis 1s known only from the Surgencia del
Tigre (Zu. 23), in the middle basin of the
Rio Guasare, north of the Sierra de Perija
in northwestern Venezuela (Fig. 5). The
cave is near the margin of Rio Guasare and
is the source of a spring during seasonal
rains (Sociedad Venezolana de Espeleologia

570

1991). The cave’s lower conduit has a 280
m course, 2—3 m wide and 1—2 m high, nar-
rowly communicating with the access gal-
lery. The underground river is permanently
fed by a spring about 60 m into the lower
gallery. At the time the cave was surveyed,
the average depth of this water course was
1.5 m with deeper pools along its course
where the catfishes were observed (Socie-
dad Venezolana de Espeleologia 1991).

Etymology.—The name is based on Rio
Guasare, parent stream of the subterranean
waters in which this endemic catfish species
lives.

Discussion

Rhamdia guasarensis is placed in Hep-
tapteridae by its possession of four syna-
pomorphies identified for the family (Lund-
berg & McDade 1986, Ferraris 1988, Bock-
mann & Guazelli 2003): posterior limb of
fourth transverse process expanded and
notched; posterodorsal corner of hyoman-
dibula greatly expanded for attachment of
levator operculi muscle; dorsal margin of
quadrate free, not sutured to hyomandibula
and metapterygoid; ventrolateral corner of
mesethmoid anteriorly recurved. However,
the new species lacks a fifth synapomorphy
of heptapterids: a straight-edged vertical
bony lamina on the Weberian complex cen-
trum. Instead, the vertical lamina has a con-
cave margin in R. guasarensis.

Except for the obvious lack of a free or-
bital rim, R. guasarensis possesses the
character combination presented as diag-
nostic of Rhamdia by Silfvergrip (1996:74).
This includes: three pairs of barbels, double
lip fold, vomer without teeth, transverse
processes of fourth vertebra expanded
branched distally, supraoccipital process
not contacting anterior nuchal plate, adi-
pose fin with free posterior margin, poste-
rior fontanelle closed and postcleithral pro-
cess well developed. However, none of
these are unambiguous synapomorphies of
the group of species comprising Rhamdia.
Instead, some characters are heptapterid or

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

higher level synapomorphies, and others,
some of uncertain polarity, have wider and
variable distributions among heptapterids.
Thus, placement of this new species in
Rhamdia must be considered provisional
because the genus has not been supported
as monophyletic. Bockmann’s (1998) phy-
logenetic analysis of Heptapteridae placed
one representative species, R. laticauda,
sister to Pimelodella but a second species,
R. quelen, is deeper in his cladograms. In
this context R. guasarensis has one derived,
although non-unique, feature listed by
Bockmann as diagnostic for R. quelen. This
is the highly reduced posterior cranial fon-
tanelle, long used as one of the diagnostic
characters of Rhamdia. Indeed, we find the
posterior fontanelle closed or reduced to a
small foramen in the supraoccipital in most
other Rhamdia examined: R. laukidi, R. ni-
caraguensis, R. quelen (including speci-
mens originally identified as R. guatema-
lensis, R. hilarii, R.vilsoni, R. wagneri) and
some R. laticauda. Silfvergrip (1996) re-
ported the fontanelle to be variably open or
reduced in R. laticauda, and our sample
also shows such variability among speci-
mens. We find that R. muelleri has an open
posterior fontanelle. The fontanelle is also
closed in the heptapterids Brachyglanis,
Brachyrhamdia, Leptorhamdia, and Myog-
lanis (Bockmann 1998, pers. obs.). Fur-
thermore, R. guasarensis has an uncinate
process on hypobranchial 1, unlike R. que-
len that lacks the process (listed as a second
non-unique derived feature of R. quelen by
Bockmann 1998). Accordingly, we do not
take the foregoing as evidence for a partic-
ularly close phylogenetic relationship be-
tween R. guasarensis and R. quelen. The
midline union of frontal bones (Fig. 2) and
concomitant extreme reduction of the an-
terior fontanelle are a distinctive apomorph-
ic character of R. guasarensis. Although not
all species have been examined for this fea-
ture, we have not observed it in any Rham-
dia nor has it been previously reported, and
in his description of the genus, Silvergrip
(1996:74) reported the anterior fontanelle to

VOLUME 117, NUMBER 4

be invariably open. This is at least a diag-
nostic autapomorphy of the species, al-
though these features are potentially infor-
mative about relationships. Bockmann
(1998) illustrated a variety of conditions of
anterior fontanelle narrowing and closure in
other heptapterids including Myoglanis,
Taunayia, Imparfinis pristos, and an unde-
scribed species, but all of these are struc-
turally different from that in R. guasarensis.

Another peculiar character of the new
species is the concave dorsal profile of the
head. In general, Rhamdia species, includ-
ing most cave populations, have convexly
rounded heads. The cave species R. macus-
Panensis recently described from Mexico
(Weber & Wilkens 1998) has a straight dor-
sal head profile, somewhat more similar to
that of R. guasarensis than to other con-
geners. Rhamdia macuspanensis is readily
distinguished from-R. guasarensis by its
strong development of pectoral spine den-
tations and rounded tips of the caudal lobes.

Rhamdia guasarensis possesses typical
reductive characteristics in common with
other cave-dwelling species and popula-
tions of the genus. Furthermore, the greater
relative length of the head and elevated
number of pectoral-fin rays are also shared
by other troglobitic species of the genus
(Weber 1996). It has been suggested that
larger head size is related to an increase in
the development of the cephalic laterosen-
sory system (Langecker & Longley 1993,
Weber 1996), and the greater number of
pectoral-fin rays is possibly correlated with
the increased mass of the anterior part of
the body, compensating this increase with a
greater fin area for hydrodynamic lift (We-
ber 1996). If there is a functional relation-
ship between head size and pectoral-fin area
in cave dwelling Rhamdia, it does not ex-
tend to the heptapterid genus Pimelodella,
wherein the large-headed P. kronei has
eight or nine pectoral-fin rays (Trajano
1997, Trajano & Britski 1992) and the
small-headed P. spelaea has ten pectoral-
fin rays (Trajano et al. 2004).

No Rhamdia species are known from the

571

surface waters of Rio Guasare, thus R.
guasarensis 1s not a cave-dwelling ecotype
of a proximate epigean species. Two Rham-
dia species have been reported from north-
western Venezuela. From the Lago de Ma-
racaibo Basin, Schultz (1944) published on
specimens now identified as R. quelen (Sil-
fvergrip 1996). Fernandez-Yépez & Martin
(1953) reported R. wagneri based on spec-
imens collected in the Rio Negro in the
southern part of the Perija range. One of us
(CDN) has reidentified these specimens at
the Museo de Historia Natural La Salle as
R. quelen. As noted above, there is no ev-
idence for a uniquely close relationship be-
tween R. guasarensis and R. quelen.

The fauna of troglobitic catfishes of the
Sierra de Perija region includes: Ancistrus
galani Pérez & Viloria, 1994, Trichomyc-
terus spelaeus DoNascimiento, Villarreal &
Provenzano, 2001, and Rhamdia guasar-
ensis. There is another hypogean population
of Trichomycterus, possibly an undescribed
species, living in a cave drained by the Rio
Yasa (Rio Negro system) in the southern
part of the Sierra de Perija (DoNascimiento,
in prep.).

The diversity of three cave catfishes of
the Rio Guasare system is among the high-
est of any Neotropical karst region, al-
though the species are not found syntopi-
cally in the same cave. By contrast Bichu-
ette & Trajano (2003) list five troglobitic
species in caves of the Sao Domingos karst
area, Goias, Brazil. Ancistrus cryptophthal-
mus and the trichomycterid /tuglanis pas-
sensis coexist in the Sao Vicente cave, To-
cantins Basin, Goiads, Brazil (Trajano &
Souza 1994, Fernandez & Bichuette 2002).
Also, the inundated caves of the Formosin-
ho karst region of Bodoquena, Mato Grosso
do Sul, southeastern Brazil, are co-inhabit-
ed by Ancistrus formoso and an unde-
scribed troglomorphic population of T7i-
chomycterus (Sabino & Trajano 1997).

Cave-dwelling and specialized troglobitic
neotropical catfishes belong to the families
Astroblepidae, Heptapteridae, Loricariidae,
and Trichomycteridae. Within the last three

572

of these families the genera Rhamdia, An-
cistrus, and Trichomycterus are most com-
monly represented in cave faunas. Their
prevalence suggests possession of morpho-
logical, physiological, behavioral, and eco-
logical features (preadaptations) that facili-
tate existence in cave waters (Eigenmann
1919, Norman 1926, Hubbs 1936).

Wilkens (1986) proposed a correlation
between degree of morphological reduction
and time of subterranean evolution based
on a neutral mutation model for the regres-
sive evolution of eyes and pigmentation in
cave fishes and crustaceans. The subterra-
nean catfishes of the Sierra de Perija, es-
pecially Trichomycterus spelaeus and R.
guasarensis, are highly advanced in their
troglobitic features, suggesting that they are
not new arrivals in their subterranean en-
vironment. Ocular and pigmentation reduc-
tion of R. guasarensis and T. spelaeus are
complete. These species exhibit additional
autapomorphies such as extremely elongate
barbels in Trichomycterus and much en-
larged head laterosensory organs in Rham-
dia. These characters, too, may indicate a
long period of hypogean evolution. Indirect
evidence suggests the availability of an am-
ple period of time for the evolution of the
Perija cave fishes. Paleogeographic recon-
structions of northwestern Venezuela sug-
gest that uplift of the Sierra began in the
early Cenozoic (Gonzalez de Juana et al.
1980). It is reasonable to assume that these
fishes originated in situ after subterranean
waters carved out their habitat within Cre-
taceous limestones of the Sierra de Periya.
On the fish side of the equation, the only
fossil record of Rhamdia are fin spines of
relatively young Pleistocene age (Cione
1982). However, based on much older Mio-
cene fossils of phylogenetically related pi-
melodid and pseudopimelodid catfishes, the
heptapterids are expected to have originated
and diversified long before the late Pleis-
tocene (Lundberg 1998). Thus it is possible
that subterranean aquatic habitats and cave
fishes have been present in this region for
tens of millions of years.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Comparative material.—Rhamdia lati-
cauda: ANSP 104034, one specimen, X-ray
and alc, 86 mm SL, Panama, Cocle:
UMMZ 197078, two dry skeletons, 131—
146 mm SL, Honduras. R. laukidi: ANSP
139184, one of three specimens, X-ray and
alc, 127 mm SL, Colombia, Meta. R. muel-
leri: ANSP 162521, two of four specimens,
X-ray and alc, 109-110 mm SL, Venezuela,
Amazonas. R. nicaraguensis: ANSP 8444,
one specimen, alc, 135 mm SL, Nicaragua,
Lago Nicaragua. Rhamdia quelen: ANSP
141578, two of five specimens, X-ray and
alc, 100-105 mm SL, Venezuela, Bolivar;
ANSP 45365 (original identification R.
guatemalensis), one specimen, X-ray and
alc, 120 mm SL, Panama, Canal Zone;
ANSP 172138 (original identification R. hi-
larii), two of 37 specimens, X-ray and alc,
107-110 mm SL, Brazil, Minas Gerais;
ANSP 16020 (original identification R. vil-
soni), one specimen, alc, 200 mm SL, Trin-
idad; ANSP 71621 (original identification
R. wagneri), one specimen, X-ray and alc,
125 mm SL, Colombia, Magdalena; DU-
F1021, one dry skeleton, 202 mm SL;
MBUCV-CT-561, eight specimens, CS, 23—
57 mm SL, Venezuela, Zulia; MHNLS-
1645, two specimens, alc, 59-223 mm SL,
MHNLS-1734, three specimens, alc, 107—
228 mm SL, Venezuela, Zulia.

Acknowledgments

We are indebted to the members of So-
ciedad Venezolana de Espeleologia, espe-
cially O. Villarreal who both brought us the
specimens described here and assisted with
illustration of the skull. K. Luckenbill ably
prepared the final figures. H. H. Ng gener-
ously provided us with character data on
comparative skeletal specimens at UMMZ.
We are indebted to two anonymous and
careful reviewers for many useful com-
ments on the manuscript. N. Milani de Ar-
nal photographed the holotype, and M. Litt-
mann radiographed the ANSP paratype.
Partial support of publication costs was pro-
vided by the All Catfish Species Inventory

VOLUME 117, NUMBER 4

(NSF DEB-0315963) and an NSF research
award to JGL (DEB-0089612).

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Associate Editor: Edward O. Murdy

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
117(4):575—-581. 2004.

Taxonomic review of the fossil Procellariidae (Aves:
Procellariiformes) described from Bermuda by R. W. Shufeldt

Storrs L. Olson

Division of Birds, National Museum of Natural History, Smithsonian Institution, Washington,

D.C. 20560, U.S.A., e-mail: olsons@si.edu

Abstract.—The literature and specimens relevant to the three new species of
petrels (Procellariidae) proposed by R. W. Shufeldt from Quaternary fossils
from Bermuda were re-examined. A case is made for citing all three binomials
as dating from Shufeldt’s earlier preliminary publication (1916) rather than his
later monograph (1922). Aestrelata vociferans Shufeldt, 2 October 1916, was
correctly synonymized with Aestrelata cahow Nichols & Mowbray, 31 March
1916, and a lectotype is designated here. Puffinis mcgalli Shufeldt, 1916, was
correctly synonymized with Puffinus puffinus Briinnich, 1764, with the holo-
type evidently representing a casual occurrence. A lectotype is designated for
Puffinus parvus Shufeldt, 1916. This taxon is not synonymous with Puffinus
lherminieri.:Lesson, 1839, being much smaller, and is provisionally retained
until its status relative to other taxa in the Puffinus assimilis/lherminieri com-

plex can be assessed.

Because seabirds of the family Procellar-
lidae are usually the most prevalent mem-
bers of the fossil avifaunas recovered in
Bermuda, it is desirable to resolve several
taxonomic and nomenclatural problems that
were introduced in two papers by R.W.
Shufeldt (1916, 1922) in which he named
three new species of petrels and shearwa-
ters from fossil remains of uncertain age
obtained in several caves in Bermuda. Al-
though his names were all subsequently
synonymized, these actions were taken
without reference to Shufeldt’s original ma-
terial, most of which is now to be found in
the Carnegie Museum of Natural History,
Pittsburgh (not the British Museum, as sur-
mised by Brodkorb, 1963). The objectives
of this review are: (1) to establish the orig-
inal citation for each of Shufeldt’s names;
(2) to attempt to identify at least parts of
the type series upon which each species was
based and designate lectotypes where ap-
propriate; and (3) to determine autoptically
the identity and validity of each of Shu-
feldt’s taxa.

Considering the deficiencies of the com-
parative osteological material available to
Shufeldt, his studies of Bermudan fossils
are quite exemplary. Regardless of the ul-
timate fate of Shufeldt’s names, his analysis
of the specimens and his conclusions were
for the most part meritorious—something
that cannot be said for many of his other
studies of fossil birds. Shufeldt’s first con-
tribution to Bermudan paleontology (Shu-
feldt 1916) was intended only as a prelim-
inary introduction to a larger work. He had
progressed at least as far as mounting the
plates for this proposed monograph, as at
this point he refers specifically to the plate
and figure numbers of the unpublished larg-
er manuscript. The figure numbers men-
tioned at this time correspond exactly with
those published later (Shufeldt 1922), al-
though the plates were renumbered accord-
ing to the sequence necessitated by the jour-
nal in which they appeared. Publication of
the definitive paper was originally to have
been through the American Museum of
Natural History, but this never took place;

576

the paper was delayed (7 years) and even-
tually was issued in the Carnegie Museum
series. That a delay was forthcoming must
have been apparent to Shufeldt in 1916, as
he included an addendum to his preliminary
paper in which he named his new taxa, al-
though the descriptions accompanying the
names were very spare. Some of the names
have been construed as nomina nuda at this
point (e.g., Brodkorb 1963:246), but for
reasons given below I consider all of Shu-
feldt’s names to date from the 1916 publi-
cation.

There were several collections of Ber-
mudan fossils upon which Shufeldt based
his descriptions of Aestrelata vociferans,
Puffinus mcgalli, and P. parvus. The orig-
inal one, upon which he had been invited
to work by EF A. Lucas “Director of the
American Museum of Natural History”
(Shufeldt 1916:623), had been obtained ‘by
L. L. Mowbray. Material from this collec-
tion was identified by Shufeldt (1922) as
being from the American Museum
(AMNH). Another collection was obtained
by Edward McGall and was referred to in
Shufeldt (1922) as the McGall Collection.
Apparently the AMNH material was never
returned and most of Shufeldt’s material
that has been traced so far is in the collec-
tions of the Carnegie Museum. Further-
more, at least one specimen identified in
Shufeldt (1922) as coming from the AMNH
collection was exchanged from the Carne-
gie Museum to the Smithsonian Institution
in 1932 (USNM 320059, accession no.
117209). (All USNM and CM catalog num-
bers refer to series in the ornithological
rather than paleontological collections.)

Identifying Shufeldt’s type material is
made more difficult by the fact that none of
the specimens involved had been cataloged
or numbered. It should be noted that
McGall and Anthony Tall evidently sent ad-
ditional specimens to Harvard University,
the British Museum, and perhaps elsewhere
(Shufeldt 1922:384), but Shufeldt never ex-
amined these specimens and they certainly
have no claim as types.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Pterodroma cahow (Nichols & Mowbray,
1916)

Aestrelata cahow Nichols & Mowbray,
1916 (31 March):194.
Aestrelata vociferans Shufeldt, 1916 (2 Oc-
tober):633, Shufeldt, 1922:365.
Oestrelata vociferans: Lambrecht,
DY Ne

Pterodroma cahow: Bent, 1922 (19 Octo-
ber):112 (new combination with A. vo-
ciferans in synonymy); Brodkorb, 1963:
246.

IQ32

Lectotype (here designated).— Aestrelata
vociferans Shufeldt 1916, skull (neurocra-
nium with attached maxillary rostrum and
right quadratojugal) included with USNM
320059. Measurements: total length 74.7
mm; cranium length 40.2, cranium width at
postorbital processes 29.5, cranium depth
21.1; least width interorbital bridge 10.4,
width at naso-frontal hinge 10.3; length of
rostrum from naso-frontal hinge 36.2;
length of nostril 11.4; length of premaxilla
anterior to nostril 20.0.

This specimen can be identified unequiy-
ocally as the fossil of Aestrelata vociferans
illustrated in Shufeldt (1922) as Figure 5 on
Plate 16, by the shape of the small flange
of bone projecting ventrally nearly across
the ventral interorbital fenestra. This flange
is extremely variable in Pterodroma cahow
and may range from a small pointed pro-
jection to a continuous bridge across the fe-
nestra. The distinctive shape in USNM
320059 is exactly as shown in Shufeldt’s
figure (Fig. la, b), and all other variations,
such as positions of small foramina, corre-
spond exactly as well. In Shufeldt (1916:
635) it is stated that ‘““The differences in the
osseous mandibles of a Petrel (4strelata
vociferans) and a Shearwater (Puffinus
lherminieri) are easily appreciated upon
comparing those parts in figs. 5 & 6 of pl.
i.” This reference is to figures in the then
unpublished manuscript. The plates were
renumbered in Shufeldt 1922, so that plate
1 became plate 16g in which Fig. 5 is the
specimen designated here as lectotype. In

VOLUME 117, NUMBER 4 577

Fig. 1. A, lectotype of Aestrelata cahow Shufeldt (1916), USNM 320059; the quadratojugal and quadrate
were separated from the rest of the skull subsequent to Shufeldt’s photograph and may not have been rejoined
in exactly the same position; the quadrate is not necessarily from the same individual as the skull and is not to
be considered as part of the lectotype. B, Shufeldt’s illustration (1922: fig. 5, plate 16) of the same specimen;
arrow indicates the diagnostic flange of bone in the interorbital foramen that identifies the photograph with
USNM 320059. C, left humerus of Puffinus Iherminieri USNM 428934 from Bermuda. D, left humerus, lectotype
of Puffinus parvus Shufeldt (1916), CM 16539. E, Shufeldt’s illustration (1922: fig. 56, plate 25) of the same
specimen; the markings on the shaft and bit of matrix in the olecranal fossa identify the photograph with CM

16539.

the legend, this was identified as being part
of the series that was supposed to be in
AMNH (see above).

USNM 320059 was received from the
Carnegie Museum in exchange in 1932.
The label with this specimen reads “‘Skel-
eton of adult ‘Cahow’ | Aéstrelata vociferans

sp. nov. Shuf. | Made as perfect as the
bones in the | collection would allow R. W.
S[hufeldt]. | 11 Dec. ‘15.”
Paralectotypes.—Because of adhering
matrix, discolorations, or individual osteo-
logical variation, the following specimens
can be identified with photographs in Shu-
feldt (1922) and are therefore unequivocally
part of his type series. Shufeldt’s figure
number follows the current museum num-
ber: skulls CM 16533 (fig. 1), 16534 (fig.

2), 16535 (fig. 3); sterna 16537 (fig. 26),
16538 (fig. 27). Skull CM 16536 may be
the one illustrated in fig. 4, but if so, both
quadratojugals are now lacking and I did
not detect any peculiarity of the specimen
that would allow it to be certainly identified
with the figure.

Remarks.—Of the new names for Ber-
mudan petrels introduced by Shufeldt, the
citation for Aestrelata vociferans presents
the most difficulties, as no characters of the
species itself are actually mentioned and no
specimens were illustrated in Shufeldt
(1916). Nevertheless, he did discuss osteo-
logical characters of the fossils that defi-
nitely refer them to Aestrelata (= Ptero-
droma) as opposed to Puffinus. Only one
species of Pterodroma has ever been found

578

in fossil deposits on Bermuda, and Shufeldt
identified his new species with the “‘ca-
how” of legend, which was later definitely
established as being a species of Ptero-
droma (Murphy & Mowbray 1951). Fur-
thermore, Shufeldt specifically refers to
bones of the new species illustrated in
plates prepared for his monograph pub-
lished later (Shufeldt 1922) and unequivo-
cally identifies them by figure number and
plate number. Therefore, it is now possible
to identify particular specimens of Shu-
feldt’s new species based on information
given in the 1916 publication. Thus, it may
be argued, as I believe, that Aestrelata vo-
ciferans is valid as of Shufeldt 1916 rather
than Shufeldt 1922. It is a moot point, how-
ever, as A. vociferans Shufeldt 1916 is still
a junior synonym by 6 months of A. cahow
Nichols & Mowbray, 1916. If A. vociferans
is dated from Shufeldt 1922, Bent (1922:
114), who had access to Shufeldt’s manu-
script, effectively synonymized Shufeldt’s
name 17 days later by saying that it was
“apparently the same bird” as A. cahow of
Nichols & Mowbray.

The unravelling of the identity of the bird
known to Bermuda’s early settlers as the
“cahow” is well summarized by Murphy &
Mowbray (1951). This bird was once in-
credibly abundant and provided the early
colonists with a ready supply of food. But
it was so overexploited by man and intro-
duced mammals that it had seemingly dis-
appeared before its identity could be made
known to naturalists. A living example of a
Pterodroma was taken in Bermuda in 1906
by L. L. Mowbray, but was referred to a
species that breeds in New Zealand (Brad-
lee 1906). Not until a decade later was this
specimen described as the type of a new
species, Aestrelata cahow (Nichols &
Mowbray 1916), almost simultaneously
with Shufeldt’s (1916) preliminary note.
Shufeldt deserves a fair amount of credit
for developing our knowledge of the Ca-
how, as his paleontological studies were as
seminal as any in providing documentation
that the Cahow was one of the gadfly pet-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

rels now recognized in the genus Ptfero-
droma.

Puffinus puffinus puffinus (Briinnich, 1764)

Puffinus puffinus bermudae Nichols &
Mowbray, 1916 (31 March):195.

Puffinus mcgalli Shufeldt 1916 (2 October):
630; Shufeldt, 1922:354.

Puffinus puffinus puffinus: Dwight 1927:
243 (with P. p. bermudae in synonymy).

Puffinus puffinus: Wetmore, 1931:407 (foot-
note; suggested synonymy of P. mcgalli);
Lambrecht, 1933:269; Wetmore, 1962:
16; Brodkorb, 1963:246.

Holotype.—Puffinus mcgalli Shufeldt
1916, sternum CM 16531, with a split in
the carina from which a piece of bone is
missing, also lacking the tip of the carina
and tips of some of the posterior processes.

Referred specimen.—In an addendum,
Shufeldt (1922:381, footnote) identified
what he believed to be a pedal phalanx 2.8
cm in length that he thought “belonged to
an adult specimen of Puffinus mcgalli, and
possibly to the same individual” as the hol-
otypical sternum. This specimen (CM
16532) is still in the same box with the ho-
lotype and measures 28.7 mm. It is actually
the left tibiotarsus of a juvenile passerine
bird with the proximal end quite porous and
incompletely ossified. It has no status what-
soever as a type.

Remarks.—Shufeldt (1916) based Puffi-
nus mcgalli on a sternum that was stated to
be larger than that of P. lherminieri and
smaller than that of P. major (= P. gravis),
in addition to which a measurement of the
holotype was provided. This is quite suffi-
cient to establish the name P. mcgalli at this
point. Wetmore (1931:407), presumably on
the basis of size and geographical proba-
bility, suggested that P. mcgalli was prob-
ably synonymous with P. puffinus and was
followed by Lambrecht (1933). Later, Wet-
more (1962:16) considered that Shufeldt’s
figures of the sternum of P. mcgalli “‘agree
exactly with a sternum of a female Puffinus
puffinus puffinus.” Brodkorb (1963) fol-

VOLUME 117, NUMBER 4

lowed Wetmore’s lead, but no one since
Shufeldt had ever critically examined the
specimen.

The shape of the manubrial area, the an-
gle of the sterno-coracoidal processes, and
other features establish that the holotype is
correctly referred to the genus Puffinus, as
opposed to Pterodroma. In size, it is within
the range of Puffinus puffinus puffinus:
length along midline 58.0 mm, width across
posteriormost costal facets 25.4 mm. In a
series of 10 skeletons of Puffinus puffinus
puffinus the length was 52.2—58.0 (avg.
55.1) and width 23.9—27.2 (avg. 25.7). This
is larger than Puffinus Iherminieri but
smaller than any of the other Atlantic spe-
cies of Puffinus. Thus Puffinus mcgalli Shu-
feldt, 1916, was correctly synonymized
with Puffinus puffinus Briinnich, 1764.

This occurrence of Puffinus puffinus as a
fossil in Bermuda is unique, as no other fos-
sils of the species have ever been encoun-
tered among the thousands of bones of sea-
birds collected so far. Although this species
is a common offshore visitor to Bermuda,
there are only three records of attempted
breeding (Bradlee et al. 1931, Bourne
1957). The first was a specimen “‘captured
while sitting on its solitary egg in a rocky
hole on a small island in Castle Harbor, in
April, 1864” (Reid 1884:274). The second
record, more doubtful, was another bird sit-
ting on an egg in an island in Castle Harbor
in May 1877 tentatively recorded as Puffi-
nus opisthomelas (Reid 1884:276). The fi-
nal record was a specimen taken “March
10, 1905, sitting on a single white egg in a
crevice in Gurnet Head Rock’’ (Nichols &
Mowbray 1916). This was described as a
new subspecies, Puffinus puffinus bermudae
Nichols & Mowbray, 1916, that was later
definitively synonymized with Puffinus puf-
finus puffinus by Dwight (1927).

In an instance perhaps similar to those on
Bermuda, a single incubating Manx Shear-
water was found in June 1973 on Penikese
Island, Massachusetts, west of Martha’s
Vineyard (Bierregaard et al. 1975), but
breeding evidently did not continue there

579

(Lee & Haney 1996). The first North Amer-
ican breeding colony of the species was es-
tablished in 1977 on Middle Lawn Island,
southern Newfoundland, and by 1981 the
population had grown to an estimated 350
individuals (Storey & Lien 1985). There is
no evidence that Puffinus puffinus was ever
able to establish such a colony on Bermuda
at any time in the last 400,000 years and all
the records, including the fossil sternum de-
scribed as Puffinus mcgalli, appear to have
resulted from single individuals or pairs.

Puffinus parvus Shufeldt, 1916

Puffinus parvus Shufeldt, 1916:632; Shu-
feldt, 1922:356.

Puffinus lherminieri: Wetmore, 1931:407
(footnote; suggested synonymy of P. par-
vus); Lambrecht, 1933:270; Wetmore,
1962: Brodkorb, 1963:246.

Lectotype (here designated).—Puffinus
parvus Shufeldt, 1916, left humerus, CM
16539 (fig. 56 of Shufeldt 1922). Measure-
ments: Total length 58.8 mm; proximal
width 10.7, depth of head 3.3, width and
depth of shaft at midpoint 3.8 X 2.6, distal
width 7.9.

Paralectotypes (figure numbers from
Shufeldt 1922 in parentheses).—CM 16540
right humerus (fig. 55), 16541 right humer-
us, 16542 left humerus, 16543 left humer-
us, 16544 right ulna (fig. 43), 16545 right
ulna, 16546 left ulna (fig. 44), 16547 left
radius (fig. 45), 16548 right carpometacar-
pus (fig. 67), 16549 right phalanx 1 of ma-
jor alar digit (fig. 74), 16550 left coracoid
(fig. 92), 16551 incomplete furcula (fig.
79), 16552 right tibiotarsus (fig. 119),
16553 left tibiotarsus (fig. 120), 16554 right
tarsometatarsus (fig. 107), 16555 right fe-
mur, 16556—58 left innominates.

Remarks.—TYhe name Puffinus parvus
dates from Shufeldt (1916), as there this
taxon was specifically characterized as be-
ing smaller than P. [herminieri and as be-
longing to a group of small shearwaters
having a short, rather than an elongate ster-
num. The type material he listed (p. 632)

580

as 12 bones from what he called the AMNH
series (of which only one certainly, and
three probably, can now be accounted for)
and the following from the McGall collec-
tion: “five perfect humeri, three ulnae, a ra-
dius, a carpo-metacarpus, a proximal joint
of an index digit, a coracoid, an inferior
mandible, an imperfect os furculum, a tar-
so-metatarsus, an Os innominatum of the
left side; subsequently there also came to
light an imperfect cranium.” These lists
were repeated nearly verbatim in Shufeldt
(1922:356) save that the last imperfect cra-
nium is omitted and that specimen is no
longer present, so perhaps he subsequently
re-identified it. In an addendum, Shufeldt
(1922:385) listed and identified a further se-
ries of 77 specimens of Puffinus parvus col-
lected by McGall and Tall that also was de-
posited in the Carnegie Museum, where all
but the 5 sterna and 2 of the fragmentary
furculae may still be found. It is very clear
from Shufeldt’s statements (e.g., 1922:385),
however, that the first two collections con-
stituted the type series and that the addi-
tional specimens were referred only subse-
quent to his 1916 paper and thus have no
status as types.

In the CM collections was a container of
bones labelled in Shufeldt’s hand ““McGall
Collection | Puffinus parvus Shuf. sp. nov |
Noy. 27 1915 | Fragile.” This series cor-
responds exactly to Shufeldt’s list of this
collection, less the cranium mentioned
above, except that it has been augmented
by a right and left tibiotarsus, a right femur,
and an additional two innominate bones.
Although no tibiotarsus was listed for the
McGall collection in either of Shufeldt’s
publications, the legend for Shufeldt’s
(1922) fig. 119 of a right tibiotarsus iden-
tifies it as being from the McGall collection,
whereas the left tibiotarsus in fig. 120 is
identified as being from the AMNH series,
in which there was only a single tibiotarsus.
The femur and the additional two innomi-
nates are doubtless the femur and two of
the four innominates listed for the AMNH
series, which has otherwise disappeared.

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I think that there can be no question that
all 21 of these bones may be safely regard-
ed as syntypes of Puffinus parvus Shufeldt.
Several can be identified with photographs
in Shufeldt (1922) and from these I have
selected as lectotype a humerus with dis-
tinctive markings making it individually
identifiable (Fig. Id, e). All of the remain-
ing bones in this series may be considered
paralectotypes and have been listed above
with their current catalog numbers and ref-
erence to the figure numbers in Shufeldt
(1922) where appropriate.

Without having seen the material, Wet-
more (1931) suggested in a footnote that
Puffinus parvus was probably the same as
the living Audubon’s Shearwater Puffinus
lherminieri Lesson, 1839, in which he was
followed by Lambrecht (1933). Later, in ex-
amining a few remains of small Puffinus
found in 1958 on Cockroach Island, Har-
rington Sound, Bermuda, Wetmore (1962)
noted what seemed to be two size classes
but considered that the smaller one consist-
ed of juveniles. Although he stated (p. 16)
that ““Shufeldt (1916 p. 632) noted two ap-
parent size groups and named the smaller
one Puffinus parvus,” 1 cannot interpret
anything in Shufeldt’s publication as indi-
cating that he thought there were two size
classes. Wetmore also noted that Shufeldt’s
(1922) photographs of the bones of P. par-
vus were not to the scale indicated, as Shu-
feldt himself had pointed out, however (p.
362 footnote). Wetmore concluded that P.
parvus was not a valid taxon and synony-
mized it with P. lherminieri, and he was
followed by Brodkorb (1963).

After having examined Shufeldt’s type-
series and much more extensive fossil ma-
terial from Bermuda dating from the middle
Pleistocene onward, I have concluded that
Puffinus parvus is indeed a much smaller
species than P. /herminieri (Fig. 1c, d). The
systematics of the Puffinus lherminieri/P.
assimilis assemblage is very complex and
imperfectly understood. Puffinus parvus
needs comparison with the Atlantic taxa
known as Puffinus affinis baroli, which oc-

VOLUME 117, NUMBER 4

curs in the Azores, Madeira group, and Ca-
nary Islands, and Puffinus lherminieri boydi
of the Cape Verde Islands (Jouanin &
Mougin 1979). Unfortunately, there is al-
most no skeletal material of these taxa
available for comparison. Apparently, P.
parvus was exterminated after human arriv-
al in Bermuda, after which P. lherminieri
was able to colonize the island for a brief
period before it became extinct itself as a
breeding: bird in the late 20th century. [ron-
ically, both species are present in the Cock-
roach Island material. Further investigation
of the small shearwaters of Bermuda is un-
der way, but for now Puffinus parvus Shu-
feldt, 1916, is retained as a taxon that is
clearly distinct from P. lherminieri.

Acknowledgments

I thank Kenneth _C. Parkes and Robin
Panza, Carnegie Museum of Natural His-
tory, Pittsburgh (CM), for making Shu-
feldt’s material available and for supplying
catalog numbers. The figure is by Brian
Schmidt, Division of Birds, National Mu-
seum of Natural History, Smithsonian In-
stitution (USNM).

Literature Cited

Bent, A. C. 1922. Life histories of North American
petrels and pelicans and their allies —United
States National Museum Bulletin 121:1—343.

Bierregaard, R. O. Jr., A. B. David, Il, T. D. Baird, &
R. E. Woodruff. 1975. First northwestern Atlan-
tic breeding record of the Manx Shearwater.—
Auk 92:145-147.

Bourne, W. R. P. 1957. The breeding birds of Bermu-
da.—Ibis 99:94—105.

Bradlee, T. S. 1906. Audubon’s Shearwater and Peale’s
Petrel breeding in Bermuda.—Auk 33:217.

, L. L. Mowbray, & W. EF Eaton. 1931. A list

581

of birds recorded from the Bermudas.—Pro-
ceedings of the Boston Society of Natural His-
tory 30:279-382.

Brodkorb, P. 1963. Catalogue of fossil birds. Part 1
(Archaeopterygiformes through Ardeifor-
mes).—Bulletin of the Florida State Museum,
Biological Sciences 7:179—293.

Dwight, J. 1927. The “new” Bermuda shearwater
proves to be Puffinus puffinus puffinus.—Auk
44:243.

Jouanin, C., & J.-L Mougin. 1979. Order Procellari-
iformes. Pp. 48-121 in E. Mayr & G. W. Cot-
trell, eds., Check-list of Birds of the World. Vol-
ume 1, 2nd ed. Cambridge, Massachusetts, Mu-
seum of Comparative Zoology, 547 pp.

Lambrecht, K. 1933. Handbuch der Palaeornithologie.
Gebrueder Borntraeger, Berlin, 1022 pp.

Lee, D. S., & J. C. Haney. 1996. Manx Shearwater
Puffinus puffinus—Birds of North America
257:1—28.

Murphy, R. C., & L. S. Mowbray. 1951. New light on
the Cahow, Pterodroma cahow.—Auk 68:266—
280.

Nichols, J. T., & L. L. Mowbray. 1916. Two new forms
of petrels from the Bermudas.—Auk 33:194—
195.

Reid, S. G. 1884. The birds of Bermuda.—U.S. Na-
tional Museum Bulletin 25:163—279.

Shufeldt, R. W. 1916. The bird-caves of the Bermudas
and their former inhabitants.—Ibis series 10, 4:
623-635.

. 1922. A comparative study of some subfossil
remains of birds from Bermuda, including the
“Cahow”’.—Annals of the Carnegie Museum
13:333-418.

Storey, A. E., & J. Lien. 1985. Development of the
first North American colony of Manx Shear-
waters.—Auk 102:395—401.

Wetmore, A. 1931. The fossil birds of North America.
Pp. 401—472 in Check-list of North American
Birds, 4th ed. American Ornithologists’ Union,
Lancaster, Pennsylvania.

. 1962. Bones of birds from Cockroach Island,

Bermuda. Pp. 15-17 in A. Wetmore, Notes on

fossil and subfossil birds. Smithsonian Miscel-

laneous Collections 142(2):1—-17.

Associate Editor: Gary R. Graves

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(4):582-593. 2004.

Revision of the genus Squamigera (Insecta: Zygentoma: Nicoletiidae)
with descriptions of two new species

Luis Espinasa and Bethany Burnham

Natural Sciences, Shenandoah University, 1460 University Drive, Winchester, Virginia 22601,
lespinas@su.edu or espinas!@yahoo.com

Abstract.—The genus Squamigera was described in 1999 from a single male.
Understanding of the genus was therefore limited. After several unsuccessful
expeditions, new material has finally been collected from the same cave. New
material of related Squamigera species was also found while reviewing mu-
seum collections. From these specimens two new species, S. cumcalcaris and
S. jaureguii, are described, and a better description of the diagnostic characters

of the genus is provided.

In 1988, a single male thysanuran was
collected by R. Espinasa-Closas in a Mex-
ican cave (Cueva de las Pozas Azules). The
specimen is unique in many ways. Measur-
ing 22 mm, it is one of the largest speci-
mens in the family Nicoletiidae, but more
diagnostically, it has spines on the cerci and
scales cover its body and head. All other
members of the subfamily Cubacubaninae
lack this combination. Despite many sub-
sequent visits to the same locality, no other
specimens were found. Eleven years after
the original discovery, the specimen was
described (Espinasa 1999a) and the new ni-
coletiid genus Squamigera, was established.
By necessity, the description of Squamigera
lacked a description of the female mor-
phology or of postembryonic development.
Comparison of the genus with other mem-
bers of the subfamily was difficult because
it was unclear which characters were unique
to the specimen (species variation) and
which characters had phylogenetic/taxo-
nomic value.

Fortunately, the situation has changed. A
revision of the nicoletiid collection of the
American Museum of Natural History pro-
vided a single female from a surface local-
ity collected in 1976 by Reddell and Grubs.
Also, the Sbordoni collection of cavernicole
organisms from Chiapas provided two

males and one female from two caves. And
finally, an additional male has been col-
lected from the type locality. This male is
considerably larger than any other Ameri-
can nicoletiid described.

From these specimens, two new species
are described and a revision of the taxo-
nomic characters for the genus is provided.

Materials and Methods

The live specimen was found crawling
on the cave wall and was preserved in 96%
ethanol. Dissections were made with a ste-
reo microscope and the body parts were
mounted in fixed preparations with Hoyer’s
solution. The female and juvenile male
from Chiapas, and the new Pozas Azules
specimen were not dissected. All illustra-
tions were made with aid of a camera lucida
attached to a compound microscope. The
types were deposited in the Zygentoma col-
lection of the American Museum of Natural
History.

Squamigera Espinasa, 1999

Diagnosis (amended).—A member of the
subfamily Cubacubaninae with mucronate
to emarginate scales with smooth to serrate
borders. Cerci of males with modified

VOLUME 117, NUMBER 4

spines. Parameres without a cleft on the
apex.

Description (amended).—Body propor-
tions normal to robust. Head, thorax, ab-
domen, and proximal articles of legs with
scales and setae. Distal articles of legs,
mouthparts and abdominal stylets only with
setae. Scales numerous and multiradiate,
their form mucronate to emarginated, with
smooth to highly serrated borders.

Pedicellus of adult males with unicellular
glands and apparently with a spur on its
base. Mouthparts not specialized. Mandi-
bles strongly sclerotized apically with usual
teeth. Galea apically with several sensory
pegs. Lacinia heavily sclerotized distally.
First process of lacinia pectinate. Labium
without prominent lateral lobes.

Tarsi with four articles. Praetarsi with
three simple claws. Middle claw glabrous,
slender and smaller than lateral claws.
Urosterna II—VII subdivided into coxites
and sternite. Urosterna VIII and IX of male
entire. Middle portion of sternites with 1 +
1 sublateral macrochaetae at hind borders,
as well as | + 1 near suture at about middle
of segment. Coxites on segments II-LX with
stylets. Eversible vesicles on segments [—
VI, pseudovesicles on VII. Urosterna III of
adult males sometimes with modified cox-
ites. Urosterna IV apparently without artic-
ulated submedian appendages. Urosterna
VIII with a wide and not too deep posterior
emargination. Posterior projections acute to
slightly round, pointing slightly outward.
Tergum X very protruding, almost straight
on posterior border. Posterior angles with
several subequal macrochaetae.

Point of insertion of parameres relatively
deep and with modified setae on internal
face of coxal processes. Parameres with
specialized setae on apex, but without a clef
or other modifications. Stylets [IX apparent-
ly without spines. Opening of penis longi-
tudinal. Cerci of male with modified spines.
Median filament with or without spines. Fe-
males with a subgenital plate and gonapo-
physes of adult females apparently with nu-
merous articles.

583

Rica le
(larger individual, dorsal view) and Squamigera sp.,
juvenile male (smaller individual, ventral view). Com-
parison of body proportions to illustrate the large size
of S. latebricola.

Squamigera latebricola, male topotype

Type species.—Squamigera latebricola
(Fig. 1).

Distribution.—All specimens to date
come from south-central Mexico. It is cur-
rently unknown but likely that members of
the genus occur in South America and the
Antillean islands. Their distribution is prob-
ably restricted to the neotropics.

Remarks.—Several amendments were
made to the original description of the ge-
nus: 1. Body proportions are not always ro-
bust. 2. Scales are not only slightly serrated,
but can be highly serrated. 3. Size of spur
on male pedicellus can be variable. 4. Uros-
terna II subdivided into coxites and sternite.
In the fixed preparation of the holotype it
was unclear if the urosterna was divided. 5.
Urosterna III of adult males can have mod-
ified protuberances similar to those found
in some Cubacubana (Espinasa 1991) and
Prosthecina (Espinasa 2000). 6. Number of
macrochaetae in posterior angles of tergum
X can be variable. 7. Point of insertion of
parameres relatively deep and with modi-
fied setae on internal face of coxal process-
es. 8. Parameres without a cleft. In the fixed
preparation of the holotype, the parameres
were broken as an artifact of the prepara-
tion, giving the impression of a cleft (The

584

cleft/break was not represented in the orig-
inal figures, it was only mentioned in the
text). 9. Central filament sometimes with
spines. 10. Females with a subgenital plate
and gonapophyses of adult females appar-
ently with numerous articles. There were no
female samples available when the original
description was made.

Squamigera belongs to a group of nico-
letiid genera, the Cubacubaninae (Mendes
1988), characterized by subdivided uroster-
na II-VII and fused coxites VIII and IX of
males. Squamigera is distinguished from al-
most all genera of this subfamily by having
scales. It differs from Texoreddellia (Wy-
godzinsky 1973), the only other genus with
scales, by the morphology of scales (in Tex-
oreddellia scales have three pointed borders
instead of smooth to serrated borders), and
by having scales in the head and modified
spines in cerci, which are both absent in
Texoreddellia.

Squamigera cumcalcaris, new species
Figs. 2A—G, 3A—E

Type material.—“*Grotta I Finca S. Ani-
ta’? cave, Finca S. Anita, Simojovel de Al-
lende, Chiapas, México. 830 m above sea
level. 10/LX/1973 V. Sbordoni col. Male ho-
lotype, female paratype.

Description.—Body length 15 mm. Max-
imum conserved length of antennae 12 mm
and of caudal appendages 11 mm. General
color: light yellow to white. Morphology of
the body as in generic description. Scales
similar to Fig. 5B.

Male antennae as in Fig. 2A—B. Pedicel-
lus slightly more than one half as long as
basal article. On ventral side with approxi-
mately six clusters of unicellular glands ar-
ranged in two long rows, surrounded by mi-
crochaetas forming a “U” shape. Outside
this microchaetae, another two clusters of
unicellular glands and a downward pointing
robust spine, opposite to an extension of the
basal article (Fig. 2B). Base of female an-
tennae simple and pedicellus half as long as
basal segment.

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Head with approximately 5 + 5 macro-
chaetae on border of insertion of antennae
(Fig. 2A). Mouthpart appendages relatively
short. Labial palp as in Fig. 2D. Apical ar-
ticle slightly wider than long and barely
longer than penultimate article. Penultimate
article with bulge containing macrochaetae.
Labium and first article of labial palp with
macrochaetae. Maxilla as shown in Fig. 3D.
Last article slightly longer than penulti-
mate. Apex of maxillary palp with two con-
ules of similar width and a 3rd minute extra
conule similar to Fig. 4G. Two teeth on la-
cinia. Mandibles chaetotaxy as in Fig. 2C.

Thoracic nota with scales and macro-
chaetae on lateral borders apart from sev-
eral setae of varied sizes (Fig. 3B), but no
small sclerotized spines on posterior bor-
ders. Legs relatively short and stout. Tibia
on 2nd leg with five macrochaetae, some of
them stout, and approximately 3.5 longer
than wide and % shorter than tarsus (Fig.
2E). Tibia on 3rd leg with five macrochae-
tae, and approximately 4.5 longer than
wide and ¥; shorter than tarsus (Fig. 2F).
Claws relatively short.

Urosterna III and IV of male without
modifications in the samples examined. It is
currently unknown if more fully adult spec-
imens will develop them. Urosterna VIII
posterior projections acute to slightly
rounded, subtriangular (Fig. 3A). Urotergite
X posterior angles with several long macro-
chaetae and setae of different sizes. On the
borders some prominent scales (Fig. 2G).

Urosterna IX of male similar to some
Anelpistina; point of insertion of parameres
deep and setae slightly more sclerotized on
internal face of coxal processes and above
insertion of parameres (Fig. 3A). Stylets [IX
bigger than the others, with 4—5 macro-
chaetae and an extra subapical pair. In both
males and females without spines or other
modifications. Other stylets have only three
macrochaetae plus the subapical pair.

Penis and parameres as shown in Fig.
3A. Parameres attaining % of stylets IX.
Parameres globular and with a distinct
group of microchaetae on the tip. Overall

VOLUME 117, NUMBER 4 585

Ni {

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wea

TARAS

Fig. 2. Squv umigera cumcalcaris. Male holotype. Scales and microchaetae partially shown; A, Head; B,
Pedicellus; C, IWandible; D, Labial palp and labium; E, Tibia of 2nd leg; K 3rd leg; G, Urotergum X.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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586

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Fig. 3. Squamigera cumcalcaris. Male holotype except C, female paratype. Scales and microchaetae partially
shown; A, Genital area; B, Thoracic tergum; C, Ovipositor; D, Maxilla; E, Median filament (left) and cercus

(right).

VOLUME 117, NUMBER 4

appearance similar to some Prosthecina
(Wygodzinsky 1946). Subgenital plate of
female parabolic (Fig. 3C). Ovipositor sur-
passing apex of stylets IX by thrice the
length of stylets (Fig. 3C). Gonapophyses
with approximately 38 articles.

Male caudal appendages as in Fig. 3E.
Inner side of cerci in males with spines of
varied sizes. Some spines arranged even in
a double row. The central filament also with
spines of subequal size arranged on multi-
ple rows facing both cerci. Female caudal
appendages without modifications.

Postembryonic development unknown
because of the scarcity of samples. It is as-
sumed that specimens examined are adult
based on the development of sexual sec-
ondary characters. Comparison to other
species within the subfamily indicates that
in younger instars we could expect that
spines, modifications of antenna, and size
of parameres to be reduced in younger
males. In females a smaller ovipositor could
be expected.

Known range.—Known only from the
type locality.

Etymology.—The name is derived from
the Latin “‘cum-+calcaris” for with+spur,
alluding to the prominent curved spur in the
pedicellus of the antennae in males.

Remarks.—Squamigera cumcalcaris can
be differentiated from all species of sub-
family Cubacubaninae by the spines on the
central filament. Such spines until now
were described only in Nicoleiids in the
subfamilies Coletiniinae and Subnicoleti-
inae (Mendes 1988). Adult males can be
further differentiated by the large curved
spur oriented toward the base of antennae
in the pedicellus, which in S. latebricola is
reduced to a small spine. Adult females can
be differentiated from S. jaureguii by a par-
abolic instead of trapezoidal subgenital
plate and by a considerably less subdivided
gonapophyses.

Squamigera jaureguii, new species
Figs. 4A—-H, 5A-K 6A—D

Type material.—Puente Actopan, 5 km
SE Actopan, Veracruz, Mexico. 25 Dec

587

1976. J. Reddell and A. Grubbs cols. Fe-
male holotype.

Description.—Body length 9.5 mm. An-
tennae and caudal appendages broken.
Maximum conserved length of antennae 4
mm and of caudal appendages 5 mm. Body
proportions as in Fig. 4A. General color:
light yellow to white. Morphology of the
body as in the generic and S. cumcalcaris
descriptions, unless otherwise stated. Scales
as in Figs. 4D and 5B.

Antennae as shown in Fig. 4B. Basal ar-
ticle without projections. Pedicellus slightly
less than one half as long as the basal ar-
ticle. Head with approximately 8 + 8 ma-
crochaetae on border of insertion of anten-
nae (Fig. 4C). Labial palp as in Fig. 4E.
Maxilla as shown in Fig. 4F—G. Last article
¥. longer than penultimate. Apex of maxil-
lary palp with two conules of similar width
and a 3rd minute extra conule (Fig. 4G).
Mandibles chaetotaxy as in Fig. 4H. Tho-
racic nota as in fig. 5A. Legs relatively
short and stout (Fig. SC—D). Tibia on 2nd
leg with five macrochaetae, some of them
stout, and approximately 3.2 longer than
wide and % shorter than tarsus, and five ma-
crochaetae. Hind leg broken in the speci-
men. Claws relatively short (Fig. 5E).

Urosterna I and II as in Fig. 5K Uroter-
gite X posterior angles with 2—3 long ma-
crochaetae and setae of different sizes and
on borders some prominent scales (Fig.
6A). Stylets [X bigger than the others, with
5—6 macrochaetae and an extra subapical
pair (Fig. 6B). Subgenital plate of female
trapezoid, outer border almost straight (Fig.
6B). Ovipositor surpassing apex of stylets
IX by thrice the length of stylets (Fig. 6B).
Apex as in Fig. 6C. Gonapophyses with ap-
proximately 53 articles. Cerci without mod-
ifications (Fig. 6D).

Males unknown. Postembryonic devel-
opment unknown because only a single fe-
male individual could be examined. It is as-
sumed that this individual is an adult based
on its large ovipositor. Comparison to other
species within the subfamily indicates that

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

588
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REEFS

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Fig. 4. Squamigera jaureguii. Female holotype. Scales and microchaetae partially shown. A, Body; B, Basal
portion of antennae; C, Head; D, Scales on head; E, Labial palp and labium; EK Maxilla; G, Apical portion of

maxilla; H, Mandible.

VOLUME 117, NUMBER 4

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bs a GRE ==
4 ? :

& ,? x Pn RB a OMe ONG
ee Se ‘ 4 ae eT oN
LAW 2x5 fi Ti V, x
y fp 7. ef ae xX

nee | ie

Fig. 5. Squamigera jaureguii. Female holotype. Scales and microchaetae partially shown. A, Thoracic ter-
gum; B, scales of urotergum I; C, 2nd leg; D, Apex of 2nd tibia; E, Claws of 2nd leg; K Urosternum I and II.

590 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

oy)

} ee

Fig. 6. Squamigera jaureguii. Female holotype. Scales and microchaetae partially shown. A, Urotergum X:
B, Subgenital plate and ovipositor; C, Apex of ovipositor; D, Caudal appendages.

VOLUME 117, NUMBER 4

in younger instars we could expect a small-
er Ovipositor.

Known range.—Known only from the
type locality.

Etymology.—this species is dedicated to
Sergio Jauregui to recognize his enthusias-
tic, long time participation in cave nicole-
tiid collecting and field work.

Remarks.—Squamigera jaureguii can be
differentiated from other described Squa-
migera by having more macrochaetae on
the head at the border of the insertion of
antennae, and a shorter body and append-
ages. Adult females can be differentiated
from S$. cumcalcaris by the trapezoidal in-
stead of parabolic subgenital plate and by a
considerably more subdivided gonapophys-
es. No S. latebricola females are available
for comparison. .

Squamigera latebricola Espinasa
Fig. 7A—G

Topotype.—““Cueva de las Pozas Azu-
les”’ cave (Espinasa-Perefia 1989), Taxco de
Alarcon Municipality, Guerrero State, Méx-
ico, 18°36'40"N, 99°33'25”’W. April 2001.
L. Espinasa col. Male.

Description.—Body length 29 mm. Max-
imum conserved length of antennae 29 mm
and of caudal appendages 35 mm. Body
proportions as in Fig. 1. General color: light
yellow to white. Morphology of body sim-
ilar to S. cumcalcaris and S. jaureguii, un-
less otherwise stated. Scales with slightly
less serrated borders.

Pedicellus with clusters of unicellular
glands and a small spur (Espinasa 1999a;
Fig. 1C) instead of long hooked spine of S.
cumcalcaris. Mouthpart appendages rela-
tively thin and long. Apical article of labial
palp barely longer than wide and barely
shorter than penultimate (Espinasa 1999a;
Fig. 1D). Penultimate article’s bulge not too
prominent. Maxilla as in Fig. 7D. Last ar-
ticle shorter than penultimate. Apex of
maxillary palp with two conules of similar
width and a 3rd small extra conule (Fig.
TB).

591

Thoracic nota with small sclerotized
spines on lateral and posterior borders (Es-
pinasa 1999a; Fig. 3A). Legs relatively long
(Espinasa 1999a; Fig. 2A). Tibia on 2nd leg
with seven thin macrochaetae, and approx-
imately 4.5 longer than wide and ¥% short-
er than tarsus. Tibia on 3rd leg with eight
thin macrochaetae, and approximately
slightly over 5X longer than wide and 4
shorter than tarsus. Trochanter on 3rd leg
with a protuberant spine projection (Fig.
7B) which is not present in the smaller
sized (22 mm) holotype. Claws of normal
Size.

Coxites in urosterna III (Fig. 7C) with
protuberances similar to those found in
some Cubacubana (Espinasa 1991) and
Prosthecina (Espinasa 2000). Urosterna III
in smaller holotype also with a slight pro-
tuberance (not reported in original descrip-
tion), similar to nascent protuberance found
in some immature individuals of the afore-
mentioned Cubacubana and Prosthecina.
Urosterna IV without modifications (Fig.
7A). Urosternum IX as in Fig. 7E In this
specimen the point of insertion of paramer-
es is slightly deeper than in the holotype
and closer in appearance to some Anelpis-
tina (Espinasa 1999b). Stylets [X with five
macrochaetae and an extra subapical pair
but otherwise without any other modifica-
tions. Penis and parameres as shown in Fig.
7F Parameres attaining less than % of sty-
lets [IX and curved outward. Cerci as in Fig.
7G. Females unknown.

Postembryonic development only partial-
ly understood since only two fairly large
male individuals are available, the holotype
(22 mm) and this new topotype (29 mm).
In the smaller specimen, projections of
urosterna III are only starting to develop,
spines in cerci are less prominent and tro-
chanter of hind leg has no projection.

Known range.—Known only from the
type locality.

Remarks.—Being 3 cm in length (10 cm
if antennae and caudal appendages are in-
cluded), S. latebricola can easily be differ-
entiated from all species of the subfamily

592 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 7. Squamigera latebricola. Male topotype. Scales and microchaetae partially shown. A, Urosternum IV;
B, 3rd leg. Notice projection in trochanter (scales not shown); C, Urosternum III; D, Maxilla; E, Apical portion
of Maxilla; EK Genital area; G, Spines in cercus.

VOLUME 117, NUMBER 4

Cubacubaninae by its large size. This is the
longest species among the nicoletiids,
which typically measure 1 cm or less. En-
largement of body and appendages is com-
mon among cave adapted organisms and it
is certainly the case for this species. This
species can further be differentiated from S.
cumcalcaris and S. jaureguii by the small
sclerotized spines on lateral and posterior
borders on thoracic nota (Espinasa 1999a;
Figs. 1G and 3A), and by the morphology
of its sexual secondary characters.

Squamigera sp.
Fig. 1

Material examined.—**E] Chorreadero”’
cave, Chiapa de Corzo Municipality, Chia-
pas, México. 650 m above sea level. 10/1 1-
VIII- 73. V. Sbordoni col. Male.

Description.—Body length 10 mm. An-
tenna and caudal appendages broken. Mid-
dle filament missing. Scales as in other
members of the genus. No apparent spines
in pedicellus, sterna III, or cerci. Parameres
curved outward, similar to the holotype of
S. latebricola (Espinasa 1999a, Fig. 2C),
but attaining less than Y; of stylets IX. This
single individual is probably not a mature
adult. Chorreadero cave is visited relatively
often by speleologists and hopefully more
samples will be available one day for a for-
mal description of this population.

Acknowledgments

We thank Dr. Randall T. Schuh, curator
and chair of the Division of Invertebrate
Zoology of the American Museum of Nat-
ural History, for kindly giving access to the

593

museum collection and facilitating exami-
nation of the specimens. We also thank Val-
erio Sbordoni for facilitating acquisition of
specimens from Chiapas. Work was done
with support from CEAMISH-Universidad
Aut6noma del Estado de Morelos, in facil-
ities of the American Museum of Natural
History and Shenandoah University.

Literature Cited

Espinasa, L. 1991. Descripci6n de una nueva especie
del género Cubacubana (Zygentoma: Nicoleti-
idae) y registro del género para América Con-
tinental.—Folia Entomolo6gica Mexicana 82:5—
16.

. 1999a. A new genus of the subfamily Cuba-

cubaninae (Insecta: Zygentoma: Nicoletiidae)

from a Mexican cave.—Proceedings of the Bi-
ological Society of Washington 112(1):52—58.

. 1999b. Two new species of the genus Anel-

pistina (Insecta: Zygentoma: Nicoletiidae) from

Mexican caves, with redescription of the ge-

nus.—Proceedings of the Biological Society of

Washington 112(1):59—-69.

. 2000. A new species of the genus Prosthecina
(Insecta, Zygentoma, Nicoletiidae).—Pedobiol-
ogia 44:333-341.

Espinasa-Perefia, R. 1989. El resumidero del Isote y la
Cueva de las Pozas azules.—Tepeyollotli: Gac-
eta de la Sociedad Mexicana de Exploraciones
Subterraneas 4:24—27.

Mendes, L. EF 1988. Sur deux nouvelles Nicoletiidae
(Zygentoma) cavernicoles de Gréce et de Tur-
quie et remarques sur la systématique de la fam-
ille—Revue Suisse de Zoologie 95(3):751—
V2:

Wygodzinsky, P. 1946. Sobre Nicoletia (Anelpistina)
Silvestri 1905 e Prosthecina Silvestri, 1933.—
Ciencia 7:15—25.

. 1973. Description of a new genus of cave thy-

sanuran from Texas (Nicoletiidae, Thysanura,

Insecta)—American Museum Novitates 2518:

1-8.

Associate Editor: Wayne Mathis

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(4):594—596. 2004.

BIOLOGICAL SOCIETY OF WASHINGTON
131st Annual Meeting, 15 June 2004

President Roy McDiarmid called the
meeting to order at 10:30 a.m. in the Waldo
Schmitt Room, National Museum of Natu-
ral History (NMNH). Council members and
editorial staff present: Marilyn Schotte
(Elected Council), Ron Heyer (Acting Pres-
ident Elect), Chad Walter (Treasurer), Car-
ole Baldwin (Secretary), Richard Banks,
Stephen Cairns, Bruce Collette, and Storrs
Olson (Past Presidents), and Steve Gardi-
ner, Carol Hotton, and Ed Murdy (Associate
Editors).

Minutes of the 130th Annual Meeting
were summarized by Secretary Baldwin.
Those minutes are scheduled to appear in
Volume 117(1) of the Proceedings, which
had not been published at the time of the
annual meeting. Following approval of the
minutes, McDiarmid summarized recent
Society activities. McDiarmid announced
that Proceedings Editor Richard Sternberg
submitted his resignation as Editor on 9 Oc-
tober 2003 but agreed to remain in the po-
sition until a replacement could be found.
Past President Richard Banks has agreed to
serve as interim Editor beginning | July
2004. In view of declining manuscript sub-
missions to the Proceedings, McDiarmid is
appointing a committee to investigate elec-
tronic publishing. To investigate declining
Society membership, he is re-establishing a
Membership Committee. McDiarmid also
noted that he met recently with NMNH Di-
rector, Cristian Samper, and new NMNH
Associate Director for Research and Col-
lections, Hans Sues, to inform them of the
existence of the Society and its historical
relationship with and support from the mu-
seum. Those administrators are in favor of
the museum’s continued support of the So-
ciety and are interested, in principal, in
hosting the Society’s website on the muse-

um server, but they indicated that a final
decision about the website should not be
made until the museum’s new information-
technology director is hired. Associate Ed-
itor Steve Gardiner, who has produced the
Society’s web pages on the server at Bryn
Mawr College, announced that his institu-
tion is agreeable to leaving the Society’s
website on its server if necessary. Mc-
Diarmid concluded his summary of recent
Society activities by noting that the Society
will publish a special Bulletin this year en-
titled Study of the Dorsal Gill-Arch Mus-
culature of Teleostome Fishes, with Special
Reference to the Actinopterygii, by Victor
G. Springer and G. David Johnson. This
800+ page Bulletin will comprise two vol-
umes and be published in an 8%” X 11”
format.

President McDiarmid then called on
Chad Walter for the Treasurer’s Report (Ta-
ble 1). Income for the period 1 January
2003 to 31 December 2003 was
$93,105.92, and expenses for the same pe-
riod were $74,024.02. Total Society assets
as of 15 April 2004 were $99,705.40. The
value of the endowment fund increased by
$12,307 in 2003. The Audit Committee,
Don Wilson and Neal Woodman, indicated
that they had reviewed the books and led-
gers of the Treasurer and found all financial
records to be accurate and in good order.
The Treasurer’s report was approved.

Proceedings Editor Richard Sternberg re-
ported at the Society’s Council meeting on
17 May 2004 that four issues of Volume
116 were published comprising 75 papers
and 1007 pages. As of 1 June 2004, there
were 34 submissions, but neither 117(1) nor
117(2) had yet been published. Issue 117(1)
was submitted in January 2004, but because
of the low number of submissions (8), that

VOLUME 117, NUMBER 4

Table 1.—Summary Financial Statement for 2003.

General Fund

Assets: January 1, 2003 18,365.23
Total Receipts for 2003 78,473.46
Total Disbursements for 2003 71,698.63
Assets: December 31, 2003 25,140.06
Net Changes in Funds 6,774.83

a: Annual gain in value of Endowment.
b: Annual loss in value of Endowment.

issue would have been unusually small.
Publication was delayed until more manu-
scripts were ready for publication. A deci-
sion was then made to split 117(1) into
117(1) and 117(2), both to be published ap-
proximately the same time and very soon.
Sternberg acknowledged that the decrease
in submissions: reported last year finally
caught up with us. Furthermore, he noted
that the delayed publication of the first is-
sues of volume 117 also was attributable to
slower-than-normal production of page
proofs and page-proof mailing errors. Issue
117(3) is on track for timely production.
The Editor’s report was approved by the
Council.

Custodian of Publications Storrs Olson
reported little activity with back issues but
noted that he had filled a few orders. The
print run of the Proceedings was reduced
previously from 1000 to 850 copies, but
since membership is 730, the print run
could be reduced again, perhaps to 800.

Frank Ferrari noted that the Finance
Committee (Stephen Cairns, Oliver Flint,
Chad Walter, and Ferrari) had consulted an
attorney about tax laws regarding member
contributions to the Society. The Finance
Committee recommends three categories of
gifts: Contributor ($100—$499), Sponsor
($500—$999), and Benefactor ($1000 and
higher). Donors will receive a letter from
the Society that indicates the donor re-
ceived nothing for his/her contribution, and
names of donors will be listed in four con-
secutive issues of the Proceedings. The
Committee is currently working on an an-
nouncement of the gift-fund categories.

Endowment Fund Total Assets
66,277.07 84,642.30
14,631.94a 93,105.40

2,325.39b 74,024.02
78,583.62 103,723.68
12,306.55 19,081.38

Secretary Baldwin indicated that a vote
was needed on a change to Article 8 of the
Bylaws proposed last year by the Finance
Committee. Regarding the Society’s En-
dowment Fund, the first sentence of Article
8 currently states: “There shall be an En-
dowment Fund which shall consist of con-
tributions from members, miscellaneous
gifts, and surplus funds from operations.”
The proposed change would remove “and
surplus funds from operations,” and the
amended first sentence would read: ““There
shall be an Endowment Fund which shall
consist of gifts from members and miscel-
laneous gifts.’ The proposed change was
unanimously approved.

Baldwin also noted that results of the
2004 election of officers could not be an-
nounced as usual at the annual meeting be-
cause the ballots, which are part of Pro-
ceedings issue 117(1), have not been
mailed. Results of the election will be add-
ed as an addendum to these minutes.

President McDiarmid then announced
that in hopes of increasing attendance at the
annual meetings, he had decided this year
to add a program at the conclusion of the
annual meeting. Uncharacteristically for
Society meetings, the Waldo Schmitt room
was nearly full as McDiarmid introduced
the scheduled program. McDiarmid re-
marked that the purpose of the program was
to honor Past President Bruce Collette,
whose activities in our Society are a reflec-
tion of his attitude, activities, and devotion
to promoting science. The program began
with Division of Fishes ichthyologist David
Smith presenting The Natural History of

596

Bruce B. Collette, a talk Dave had written
as an introductory talk for a symposium
honoring Collette held at the May 2004
meetings of the American Society of Ich-
thyology and Herpetologists in Norman,
Oklahoma. Following Dave’s thoughtful
and entertaining presentation, Bruce was in-
vited by McDiarmid to speak. Bruce’s com-
ments reflected his love of his job and the
thrill of being able to work on so many in-
teresting fish groups, such as tunas (“‘warm-
blooded fish!”’). At the core of his message
was the fact that seeking answers to simple
natural history questions, rather than hy-
pothesis testing, had led him to so many
years of study and a lot of publications.
Bruce concluded with a plea for Society
members to work to conserve diversity and
habitats. The meeting was adjourned at 12:
15 p.m.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Respectfully submitted,
Carole C. Baldwin
Secretary

Addendum to Minutes.—Results of the
2004 election of officers are as follows:
President-Elect—W. Ronald Heyer; Secre-
tary—Carole C. Baldwin; Treasurer—T.
Chad Walter; Elected Council—Michael D.
Carleton, W. Duane Hope, Marilyn Schotte,
E Christian Thompson, Jeffrey T. Williams,
and Neal Woodman. Additionally, a pro-
posed amendment to Article 5 was passed.
This amendment allows the Council to elect
a replacement President-Elect to finish the
term if the President-Elect is unable to car-
ry out the duties of office, and then both
the President and President-Elect positions
are voted on at the next scheduled election.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(4):597—600. 2004.

THE BIOLOGICAL SOCIETY OF WASHINGTON
CONSTITUTION AND BYLAWS

Adopted 3 December 1884
(As amended August 2004)

Article 1. Name

The name of this Society shall be the Bi-
ological Society of Washington

Article 2. Purpose

The purpose of this Society shall be for
the furtherance of taxonomic study of or-
ganisms and for the increase and diffusion
of biological knowledge among interested
persons.

Article 3. Membership

Membership in this Society shall be open
to persons and organizations interested in
the promotion of systematic biology. The
following classes of members shall be rec-
ognized: Associate Members, Active Mem-
bers, Life Members and Emeritus members.
Changes of status of membership may be
effected at any time by the payment of ap-
propriate dues. Membership shall become
effective upon payment of dues.

Associate Members shall pay annual
dues, shall receive notices of meetings and
be eligible to vote at meetings of the So-
ciety and in ballots by mail.

Active Members shall pay annual dues,
shall receive the publications of the Society,
shall receive notices of meetings, and shall
be eligible to vote at meetings of the So-
ciety and in ballots by mail.

Life Members shall be recognized as
such by the payment of a fee established by
the Council. This fee shall be paid either in
one lump sum or in four equal, consecutive
annual installments. During their lifetime,
Life Members shall receive the publications
of the Society, shall receive notices of

meetings and shall be eligible to vote at
meetings of the Society and in ballots by
mail.

Emeritus Members. Any member who
has been an Active or Associate Member
may, at the discretion of the Council, be
accorded the privileges of Emeritus Mem-
bership. These persons shall then be granted
the same status as a Life Member.

An organization which is a member may
designate a representative who may cast a
single vote in its behalf.

Article 4. Dues

Annual dues for Associate and Active
Members shall be fixed by the Council and
may be changed by the Council.

Article 5. Officers and Elections

The Officers shall be a President, a Pres-
ident-Elect, a Secretary and a Treasurer.
The President-Elect shall succeed the Pres-
ident upon the expiration of the latter’s term
of office. The President-Elect, Secretary,
and Treasurer shall be elected for a term of
two years by a majority of the members
voting by means of a mail ballot. The of-
ficers shall take office at the end of the an-
nual business meeting. A slate of candidates
shall be prepared by a Nominating Com-
mittee appointed by the President. Ballots
shall be mailed to all members at the time
of billing for annual dues in an election
year.

If, for any reason, the President shall be
unable to carry out the duties of the office,
he/she shall be succeeded by the President-
Elect until the Council judges him/her to be
competent to resume the duties of the of-

598

fice. If, for any reason, the President-Elect
shall be unable to carry out the duties of
office, the Council will elect a replacement
to fill the term; both the President and Pres-
ident-Elect positions will be voted on at the
next scheduled election. Vacancies in the
other offices shall be filled temporarily or
until the next election by a majority vote of
the Council.

Article 6. Council

The Council shall consist of the Presi-
dent, the President-Elect, the Secretary, the
Treasurer, the Chairmen of Standing Com-
mittees, the ex-Presidents, and six addition-
al members who shall be nominated by the
Nominating Committee and elected at the
same time the Officers are elected and have
two year terms of office.

The Council shall be the governing body
of the Society. It shall be responsible for
matters of policy and procedure. It shall
meet at the discretion of the President and
shall always meet prior to the annual busi-
ness meeting. It shall receive and act on re-
ports from the Committees of the Society.
It shall receive and act on the annual budget
prepared by the Finance Committee. It shall
fix the time and place of the annual busi-
ness meeting.

Actions of the Council may be emended
at any annual meeting of the Society by a
three-fourths vote of the members present.
Actions of the Council may be approved or
rejected at any annual meeting of the So-
ciety by a majority vote of the members
present.

The President, with the approval of the
Council, shall appoint ad hoc committees
which shall report to the Council.

Article 7. Meeting

The Society shall hold at least one sched-
uled meeting each year except in an emer-
gency as decided by a three-fourths vote of
the Council members present.

Reports of Standing Committees, the
Treasurer, the Auditor, and the Council shall

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

be presented to the members at the annual
meeting.

Should the Council declare an emergen-
cy, these reports may be made to the mem-
bers in printed form. The Council shall in-
stall officers in the event there shall be no
annual meeting.

Article 8. Publications

The publications of the Society shall be
The Proceedings of the Biological Society
of Washington and any other publication
that the Council authorizes. The Proceed-
ings shall be managed by an Editorial Com-
mittee, consisting of an Editor, who shall
serve as Chairman, and not less than three
Associate Editors.

Article 9. Bylaws

The Society may enact bylaws which in-
terpret and implement this Constitution.
Such bylaws, when approved by the Coun-
cil, may be adopted, amended, or repealed
by a two-thirds majority of those voting at
an annual meeting of the Society or in a
mail ballot, provided that in either case, no-
tice of the proposed action shall have been
sent to each voting member of the Society
at least thirty (30) days before the date of
the vote.

Article 10. Amendments

This Constitution may be amended by a
two-thirds majority of members voting, ei-
ther at an annual meeting of the Society, or
in a mail ballot, provided that in either case
notice of the proposed action, when ap-
proved by the Council, shall have been sent
to each voting member of the Society at
least thirty (30) days before the date of the
vote.

Article 11. Limitation

The purposes of the Society are listed in
Article 2 of the Constitution. Lobbying or
activities specifically designed to influence
legislation are not among the objectives of

VOLUME 117, NUMBER 4

the Society and no official group within the
Society shall engage in such activity.

Article 12. General Prohibitions

Notwithstanding any provision of the
Constitution or Bylaws which might be sus-
ceptible to a contrary construction:

a. The Biological Society of Washington
shall be organized exclusively for sci-
entific and educational purposes;

b. The Biological Society of Washington
shall be operated exclusively for scien-
tific and educational purposes;

c. No part of the net earnings of the Bio-
logical Society of Washington shall or
may under any circumstances inure to
the benefit of any private shareholder or
individual;

d. No substantial part of the activities of the
Biological Society of Washington shall
consist of carrying on propaganda, or
otherwise attempting to influence legis-
lation;

e. The Biological Society of Washington
shall not participate in, or intervene in
(including the publishing or distribution
of statements) political campaigns on be-
half of any candidate for public office;

f. The Biological Society of Washington
shall not be organized or operated for
profit;

g. The Biological Society of Washington
shall not: 1) lend any part of its income
or corpus; without the receipt of ade-
quate security and a reasonable rate of
interest to; 2) pay any compensation, in
excess of a reasonable allowance for sal-
aries or other compensation for personal
services actually rendered, to; 3) make
any part of its services available on pref-
erential basis, to; 4) make any purchase
of securities or any other property, for
more than adequate consideration in
money or money’s worth from; or 6) en-
gage in other transactions which result
in substantial diversions of its income or
corpus to; any officer, member of the
Council, or substantial contributor to the

599

Biological Society of Washington. The
prohibitions contained in this subsection
(g) do not imply that the Biological So-
ciety of Washington may make such
loans, payments, sales or purchases to
anyone else, unless such authority be
given or implied by any other provisions
of the Constitution or Bylaws.

Article 13. Distribution or Dissolution

Upon dissolution of the Biological So-
ciety of Washington, the Council shall dis-
tribute the assets and accrued income to one
or more organizations as determined by the
Council, but which organization or organi-
zations shall meet the limitations prescribed
in subsections (a)—(g) inclusive, of Article
12, immediately preceding.

BYLAWS

1. Quorum. Five Council members shall
constitute a quorum at a meeting of the
Council.

2. The Secretary. The Secretary shall keep
minutes of the meetings of the Council
and of the Society and shall present a
yearly summary to the Society and
Council. He/she shall issue notices for
the meetings of the Society and the
Council, shall notify members of their
election, and shall conduct the corre-
spondence of the Society and Council.

3. The Treasurer. The Treasurer shall be
in charge of the funds and keep the fi-
nancial records of the Society. He/she
shall be authorized by the Council to
make necessary disbursements, within
the limits set by the budget. The Trea-
surer shall preserve a receipted bill, or
bill and cancelled check for each pay-
ment. He/she shall present a statement of
financial accounts, audited by the Fi-
nance Committee at the time of the an-
nual business meeting.

4. The President. The President shall pre-
side at meetings of the Council and of
the Society, and perform such other
functions that may adhere to the office.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

5. The President-Elect. In cases of illness,

incapacities, death or absence of the
President, the President-Elect shall as-
sume all duties incumbent on the Presi-
dent until the Council judges the Presi-
dent to be competent to resume the du-
ties of the office. In the event of the
death of the President, the President-
Elect shall automatically become Presi-
dent.

. Budget. Prior to the annual meeting, a
budget for the next year shall be pre-
pared by the Finance Committee and
submitted to the Council for action. No
financial obligation against the Society
may be contracted by any officer or
member except as specified in the annual
budget or as provided for by special ac-
tion of the Council upon recommenda-
tion of the Finance Committee.

. Committees. The Society shall maintain
the following committees. They shall be
provided with such needed financial sup-
port, to be designated in the budget, as
the funds of the Society may warrant.
The chairmen of the standing commit-
tees shall be appointed by the President
following the annual meeting.

A. The Finance Committee shall con-
sist of the Treasurer and two members
to be appointed by the President. The
Treasurer shall not serve as the Chair-
man. It shall prepare the annual budget
for submission to the Council and shall
advise the Council in all matters affect-
ing the finances of the Society, including

the deposit and investment of funds, en-
dowments, and long-term financial pol-
icies.

B. The Editorial Committee shall be
composed of the Editor, who shall serve
as Chairman, and not less than three As-
sociate Editors. This Committee shall
advise the Council on all matters affect-
ing publication. The Editor shall be ap-
pointed by the Council. The Associate
Editors shall be appointed by the Editor.
Terms for the members of the Editorial
Committee shall be at the discretion of
the Editor.

C. The Membership Committee shall
consist of a Chairman and not less than
three members and shall be responsible
for the Society’s effort to increase or
maintain the membership. The Chairman
shall be appointed by the President, the
other members shall be appointed by the
President upon recommendation of the
Chairman.

. Endowment Fund. There shall be an

Endowment Fund which shall consist of
contributions from members and miscel-
laneous gifts. At the discretion of the
Council, the principal of this fund may
be used in publishing the Society’s jour-
nal or for the general operations of the
Society. At the discretion of the Council,
the principal of this fund may also be
used in the publication of symposia,
monographic studies, or other special
publications; however, such a decision
must be reached only during a regularly
scheduled meeting of the Council.

PROCEEDINGS

of the

Biological Society of

Washington

VOLUME 117
2004

Vol. 117(1) published 1 June 2004 Vol. 117(3) published 7 December 2004
Vol. 117(2) published 4 August 2004 Vol. 117(4) published 20 December 2004

WASHINGTON
PRINTED FOR THE SOCIETY

EDITOR

RICHARD V. STERNBERG
RICHARD C. BANKS

ASSOCIATE EDITORS

Classical Languages Invertebrates

FREDERICK M. BAYER STEPHEN L. GARDINER
CHRISTOPHER B. BOYKO
JANET W. REID

Plants Vertebrates

CarROoL HOTToNn Gary R. GRAVES
CAROLE C. BALDWIN
EDWARD O. Murpy

Insects Invertebrate Paleontology
WAYNE N. MaTHIS GALE A. BISHOP

All correspondence should be addressed to the
Biological Society of Washington,
National Museum of Natural History
Washington, D.C. 20013

Printed by
ALLEN PREss INC.
LAWRENCE, KANSAS 66044

OFFICERS AND COUNCIL
of the
BIOLOGICAL SOCIETY OF WASHINGTON
FOR 2004—2005

OFFICERS

President
ROY W. McDIARMID

President-Elect
W. RONALD HEYER

Secretary
CAROLE C. BALDWIN

Treasurer
T. CHAD WALTER

COUNCIL

Elected Members
MICHAEL D. CARLETON F. CHRISTIAN THOMPSON
W. DUANE HOPE JEFFREY T. WILLIAMS
MARILYN SCHOTTE NEAL WOODMAN

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INFORMATION FOR CONTRIBUTORS
See the Society’s web page— www.biolsocwash.org

Submission of manuscripts—Manuscripts may be submitted in one of three ways. You may mail
three paper copies of the manuscript complete with tables, figure captions, and figures (do not submit
original figures unless/until the manuscript is accepted for publication) to the Editor, Dr. Richard C.
Banks, MRC-116, National Museum of Natural History, P. O. Box 37012, Washington, DC 20013-7012
USA. Manuscripts (in Word or WordPerfect) and figures may be sent on separate computer diskettes or
CDs to the same address. For courier delivery, the museum address is 10¢4 and Constitution Ave., NW,
Washington, DC 20560 (telephone no. 202-633-0783). Finally, manuscripts and figures may be submit-
ted electronically, as attachments to email, to Banksr@si.edu. Mailing and email addresses of the corre-
sponding author must be clearly indicated. Most communication about the manuscript will be by email.

Review.—The Society strives to publish peer-reviewed research results of its members promptly. The
Editor evaluates manuscripts for general content and appropriateness and sends them to an Associate
Editor, who in turn sends them te at least two referees who are knowledgeable in the particular taxonom-
ic area. Associate Editors may return manuscripts to authors with reviews and suggestions for improve-
ment.

Publication charges.—The Society is a non-profit organization with limited funds. Depending on
available financial resources, the Society may subsidize up to 12 pages per author per year (one author
per paper). This 12-page waiver is not automatic with membership, but must be justified in a request to
the editor and treasurer when a manuscript is accepted. Even with a waiver, the author will be required
to pay full costs of figures, tables, author’s changes in proofs, and reprints. Payment of full costs is
strongly encouraged and will facilitate speedy publication.

Costs.—Printed pages @ $65.00 each, figures @ $10.00 each, tabular material @ $3.00 per printed
column inch, author’s changes in proofs $3.00 each.

Bulletins —The Society publishes appropriate monographic length manuscripts as Bulletins on an
irregular basis. Please send inquiries about possible Bulletins to the Editor.

Front cover—from this issue, p. 546.

CONTENTS

Studies on western Atlantic Octocorallia (Coelenterata: Anthozoa). Part 5. The genera Plumarella Gray,
1870; Acanthoprimnoa, n. gen.; and Candidella Bayer, 1954
Stephen D. Cairns and Frederick M. Bayer
A new species of the sea anemone Megalactis (Cnidaria: Anthozoa: Actiniaria: Actinodendridae) from
Taiwan and designation of a neotype for the type species of the genus
Adorian Ardelean and Daphne Gail Fautin
A new genus and new species of crab of the family Xanthidae MacLeay, 1838 (Crustacea: Decapoda:
Brachyura) from the southwestern Gulf of Mexico Ana Rosa Vazquez-Bader and Adolfo Gracia
A new anchialine shrimp of the genus Procaris (Crustacea: Decapoda: Procarididae) from the Yucatan

Peninsula Richard v. Sternberg and Marilyn Schotte
Macrobrachium patheinense, a new species of freshwater prawn (Crustacea: Decapoda: Palaemonidae)
from Myanmar Hla Phone and Hiroshi Suzuki
A new species of Enhydrosoma Boeck, 1872 (Copepoda: Harpacticoida: Cletodidae) from the Eastern
Tropical Pacific Samuel Gomez
New record of Ophiosyzygus disacanthus Clark, 1911 (Echinodermata: Ophiuroidea: Ophiomyxidae)
in the Caribbean Sea Giomar Helena Borrero-Pérez and Milena Benavides-Serrato
Sunagocia sainsburyi, a new flathead fish (Scorpaeniformes: Platycephalidae) from northwestern
Australia Leslie W. Knapp and Hisashi Imamura

A new species of Nannocharax (Characiformes: Distichodontidae) from Cameroon, with the descrip-
tion of contact organs and breeding tubercles in the genus
Richard P. Vari and Carl J. Ferraris, Jr.
Rhamdia guasarensis (Siluriformes: Heptapteridae), a new species of cave catfish from the Sierra de
Perija, northwestern Venezuela
Carlos DoNascimiento, Francisco Provenzano, and John G. Lundberg
Taxonomic review of the fossil Procellariidae (Aves: Procellariiformes) described from Bermuda by

R. W. Shufeldt Storrs L. Olson
Revision of the genus Squamigera (Insecta: Zygentoma: Nicoletiidae) with descriptions of two new
species Luis Espinasa and Bethany Burnham

Minutes of the 2004 Annual Meeting
Constitution and Bylaws

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