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PROCEEDINGS

of the

Biological Society of

Washington

VOLUME 112
1999

Vol. 112(1) published 23 March 1999 Vol. 112(3) published 17 September 1999
Vol. 112(2) published 15 June 1999 Vol. 112(4) published 29 December 1999

WASHINGTON
PRINTED FOR THE SOCIETY

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EDITOR

C. BRIAN ROBBINS

ASSOCIATE EDITORS

Classical Languages Invertebrates
FREDERICK M. BAYER STEPHEN L. GARDINER
FRANK D. FERRARI
RAFAEL LEMAITRE
Plants Vertebrates

DAVID B. LELLINGER GARY R. GRAVES

Insects
WAYNE N. MATHIS

All correspondence should be addressed to the
Biological Society of Washington, Smithsonian Institution
Washington, D.C. 20560

ALLEN PRESS INC.
LAWRENCE, KANSAS 66044

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OFFICERS AND COUNCIL
of the
BIOLOGICAL SOCIETY OF WASHINGTON
FOR 1998-1999

OFFICERS

President
RICHARD P. VARI

President-Elect
BRIAN F. KENSLEY

Secretary
CAROLE C. BALDWIN

Treasurer
T. CHAD WALTER

COUNCIL

Elected Members
MICHAEL D. CARLETON RAFAEL LEMAITEE
W. DUANE HOPE ROY W. MCDIARMID
SUSAN L. JEWETT JAMES N. NORRIS

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TABLE OF CONTENTS

Volume 112

Asakura, Akira. Jcelopagurus tuberculosis, a distinct new hermit crab species (Crustacea: Decap-
a a METAL PN EASIEN RII ANL ee siete See ene cane ce one co ee ae odes sama enn Qn eee eeum case acccesevencnecece

Blazewicz, Magadalena, and Richard W. Heard. First record of the family Gynodiastyldae Stebbing,
1912 (Crustacea: Malacostraca: Cumacea) from Antarctic waters with the description of Gyno-
MIN EEAICE NUK IP MARTA W SPENT ote oe CL etek dct. coche ue ce eae e tes iuerecdscauccvcccasceuss

Bolstad, Kathrin S., and Brian Kensley. Two new species of Hansenium (Crustacea: Isopoda:
Oe EE Spat ects Pain (eos G70 rch Mtn ee

Boyko, Christopher B. The Albuneidae (Decapoda: Anomura: Hippoidea) of the Hawaiian Islands,
EE eam TTIE WV USIICCIES NUS E ILL. ee oe seks Sees Coa cules edn Wav dle Shale bday dinbiswew es Seeese

Brown, W. C., A. C. Alcala, P. S. Ong, and A. C. Diesmos. A new species of Platymantis (Am-
phibia: Ranidae) from the Sierra Madre Mountains, Luzon Island, Philippines ...................

Cai, Yixiong, Nguyen Xuan Quynh, and Peter K. L. Ng. Caridina clinata, a new species of
freshwater shrimp (Crustacea: Decapoda: Atyidae) from northern Vietnam .......................

Campos, Ernesto. Inclusion of the austral species Pinnotheres politus (Smith, 1869) and P. garthi
Fenucci, 1975 within the genus Calyptraeotheres Campos, 1990 (Crustacea: Brachyura:
ta Aa ee er Re Ps ee PN eae ore ik ape avo eaidin Shs ats opidigs Oe Ke ened wee

Campos, Martha R. A new species of freshwater crab of the genus Strengeriana Pretzmann, 1971,
Mame mommia (Crustacea. Decapoda: Pseudothelphusidac) ..... 2... i... .cescee.-e ccc eccceeeecccans

Campos, Martha R., and Rafael Lemaitre. Two new freshwater crabs of the genus Ptychophallus
Smalley, 1964 (Crustacea: Decapoda: Brachyura: Pseudothelphusidae) from Panama, with notes
2B PEC ERILD YS DU Pee OT Sil set eemepeel penal op Alaa na ars ea aaa ile eae nm Eee

Carleton, Michael D., Robert D. Fisher, and Alfred L. Gardner. Identification and distribution of
cotton rats, genus Sigmodon (Muridae: Sigmodontinae), of Nayarit, México ......................

Chen, H.-L., and Peter K. L. Ng. Description of a new spider crab, Maja gracilipes, from the
South China Sea, with notes on the taxonomic validity of M. brevispinosis Dai, 1981 (Crustacea:
2 FILLE LS: EDS Peels SF eS es ee Sg a ee a

Erdmann, Mark V., and Raymond B. Manning. A new species of Siamosquilla from Indonesia
Deeeate Ae OM aOpouday ETOLOSGUUINGAG) ~ 0725. el occ ea oe eet cnc cnedecccceccusecncteseues

Erséus, Christer. Parvidrilus strayeri, a new genus and species, an enigmatic interstitial clitellate
eT te TTI WU ACCES 1H NAAT 1. es oe os canes oe ness esses sesncecesscenesvcctucucecncacecs

Espinasa, Luis. A new genus of the subfamily Cubacubaninae (Insecta: Zygentoma: Nicoletiidae)
from a Mexican cave

i

Espinasa, Luis. Two new species of the genus Anelpistina (Insecta: Zygentoma: Nicoletiidae) from
es ene Stat FCUCSEMPMOM OF Le SEMUS 1... sco. 2s se. sees qag eases cues ic ek eceanae ee naseececcee

Fautin, Daphne G., and Brian R. Barber. Maractis remicarivora, a new genus and species of sea
anemone (Cnidaria: Anthozoa: Actiniaria: Actinostolidae) from an Atlantic hydrothermal
vent

ee

Fraser, Thomas H. A new species of cardinalfish (Perciformes: Apogonidae) from the Bay of
Bengal, Indian Ocean

ee

Gillet, Patrick. A new species of Orbiniella (Orbiniidae: Polychaeta) from Marion Island, Indian
Ocean

Graves, Gary R. Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 7. Probable parentage of
RE eRe ad MORE SITELA DSS oe es as hk we was eis Se wlsewn cn name twewenececaveteesunceas
Graves, Gary R. Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 8. A provisional hypoth-
Eismonne uybid onein of Zodalia elyceria (Gould, 1858) -...... 2.2.2.2... e ese c ee eee eee ce ee cee
Graves, Gary R. Taxonomic notes on hummingbirds (Aves: Trochilidae). 2. Popelairia letitiae
eae HeCCeIVISES AGT) TGI2 YAS A VALI SPECIES... 0.522. ane re Foote cence eee cer ecneetcsscaaescnnanancas
Gutiérrez-Aguirre, M. A., and E. Sudrez-Morales. The freshwater centropagid Osphranticum la-
bronectum Forbes, 1882 (Crustacea: Copepoda: Calanoida) in Mexico with description of a new
ee el ne ene MA Ee Se LOE eee ee, tre ence nc cen cc ceboeeccmsscactswaecssauwestiestunns
Harasewych, M. G., and Yuri I. Kantor. A revision of the Antarctic genus Chlanidota (Gastropoda:
eae Sma cP Me CEPI EE IS ee ic aod we ee casas ee} Scie wen Ceuta a veweucucebehsessnccrbacsune
Hauswaldt, J. Susanne, and Katherine E. Pearson. Urticina mcpeaki, a new species of sea anemone
(Anthozoa: Actiniaria: Actiniidae) from the North American Pacific coast ..................0-005-
Heyer, W. Ronald. A new genus and species of frog from Bahia, Brazil (Amphibia: Anura: Lep-
todactylidae) with comments on the zoogeography of the Brazilian campos rupestres ...........

Vii

381-395

362-367

164-174

145-163

510-514

531-535

536-540

405409

553-561

813-856

754-758

94-96

327-337

52-58

59-69

624-63 1

40-44

593-597

443-450

491-502

804-812

687-694

253-303

652—660

19-39

Vill

Heyer, W. Ronald, and Anna M. Munoz. Validation of Eleutherodactylus crepitans Bokermann,
1965, notes on the type locality of Telatrema heterodactylum Miranda-Ribeiro, 1937, and de-
scription of a new species of Eleutherodactylus from Mato Grosso, Brazil (Amphibia: Anura:
Beptedactylidae) «a. ses.c.55 Uszdcsnaetneg coeties east airs eee et ie seieee see ae arnbite ene oe ien oc ce eee

Humes, Arthur G. Collocherides brychius, a new species (Copepoda: Siphonostomatoida: Aster-
ocheridae) from a deep-water hydrothermal site in the northeastern Pacific .......................

Jara, Carlos G., and Victor L. Palacios. Two new species of Aegla Leach (Crustacea: Decapoda:
Anomura: Aeglidac) trom SOUGHerR, CRIES ek nica ie es ees oaths seaciidinndiais a Ae epee

Jensen, Gregory C., and Rachel C. Johnson. Reinstatement and further description of Eualus subtilis
Carvacho & Olson, and comparison with E. lineatus Wicksten & Butler (Crustacea: Decapoda:
Even by ti ae) fin coi sece se occa st gh pened Sogn Se See es RR eT EA oe eee AN SM ie ae

Karanovic, Tomislav. A new stygobitic Calanoida (Crustacea: Copepoda) of the genus Stygodiap-
tomus Petkovski, 1981 from the. Balkan-Pemmsula) 2222.02. teeee ese 0+ 200 gape ackivepeeees Saye eeee

Kensley, Brian, and Marilyn Schotte. New records of isopods from the Indian River Lagoon,
Blorida (Crustacea: Peracarida)! vcs ct Os on aes oo ch eh Oo oo

Koenemann, Stefan, and John R. Holsinger. Megagidiella azul, a new genus and species of cav-
ernicolous amphipod crustacean (Bogidiellidae) from Brazil, with remarks on its biogeographic
and. pbylocenchc TelatiOnSbIPS, s.cc5400 oan thes cain sce Pee eae eee ee

Lawson, Dwight P. A new species of arboreal viper (Serpentes: Viperidae: Atheris) from Cameroon,
PSIAC A, fis ioc Ce ules tw mod ates wate ARTE ee UE Oc OR earache eae aera aor va ees Ae oc Oe

Lazo-Wasem, Eric A. A new species of Chevalia (Crustacea: Amphipoda: Corophiidae) from the
Indian Ocean with remarks on Chevalia carpenteri and the C. aviculae_ superspecies
COMIDION, oon 2h ioe se les oe te Rees IS ge oie we Sep a rer ae

Le6én-Gonzalz, Jestis Angel de, Vivianne Solis-Weiss, and Victor Ochoa Rivera. Nereidids (Poly-
chaeta) from the Caribbean Sea and adjacent Coral Islands of the southern Gulf of Mexico

Lewis, Julian J. Caecidotea simulator, a new subterranean isopod from the Ozark Springfield Plain
(Crustacea: Isopoda: Ascilidac)) so. ase: eo boon te Ste ee ase cae ae

Locke, J. M., and K. A. Coates. Redescriptions of Grania americana, G. bermudensis and descrip-
tions of two new species of Grania (Annelida: Clitellata: Enchytraeidae) from Bermuda .......

Long, Douglas J., and John E. McCosker. A new species of deep-water skate, Rajella eisenhardti,
(Chondrichthyes: Rajidae), from. the Galapacos, Islands oo). see) oe Be eee Se

Malabarba, Luiz R., and Stanley H. Weitzman. A new genus and species of South American fishes
(Teleostei: Characidae: Cheirodontinae) with a derived caudal fin, including comments about
INSEMINAtNeg -CHEMOGOMEMNES. <...,cic5.a-0sen nse oh oes Eeesben vo ¥s 008 eano sone pees ee eee

Mayer, Gregory C., John A. W. Kirsch, James M. Hutcheon, Francgois-Joseph Lapointe, and
Jacinthe Gingras. On the valid name of the lesser New Zealand short-tailed bat (Mammalia:
MGTIO PICEA) a oft oko os teoccs s Sioe S Sae ark et ee e Ue ee e

McCranie, James R., and Larry David Wilson. Two new species of the Eleutherodactylus rugulosus
group (Amphibia: Anura: Leptodactylidae) from Honduras

Migotto, Alvaro E., and Antonio C. Marques. Hydroid and medusa stages of the new species
Ectopleura obypa (Cnidaria: Hydrozoa: Tubulariidae) from Brazil .................... 6.02. c eee eee

Miquelarena, Amalia M., and Adriana E. Aquino. Taxonomic status and geographic distribution
of Bryconamericus eigenmanni Evermann & Kendall, 1906 (Characiformes: Characidae) ......

Mori, Atsushi. Caprella kuroshio, a new species (Crustacea: Amphipoda: Caprellidae), with a rede-
scription of Caprella cicur Mayer, 1903, and an evaluation of the genus Metacaprella Mayer ...

Murano, Masaaki. A new genus, Neodoxomysis (Crustacea: Mysidacea: Mysidae: Leptomysini),
with description of two new species

Ng, Peter K. L., and H.-L. Chen. On the identities of two Pacific species of deep-water porter
crabs, Hypsophrys longirostris Chen, 1986, and Homologenus donghaiensis Chen, 1986 (Crus-
bacea: Biecapoda: Brachyrray Homolidac). oy. ts nrc io eee ee rate ae ae ose oe

Ng, Peter K. L., and S. H. Tan. The Hawaiian parthenopid crabs of the genera Garthambrus Ng,
1966, and Dairoides Stebbing, 1920 (Crustacea: Decapoda: Brachyura) .....................+++05-

Nishe, Eijiroh, Tomoyuki Miura, and Michel Bhaud. A new species of Spiochaetopterus (Chae-
topteridae: Polychaeta) from a cold-seep site off Hatsushima in Sagami Bay, central Japan ....

Okuno, Junji. Paleomonella hachijo, a new species of shrimp (Crustacea: Decapoda: Palaemonidae)
from a submarine cave in southern Japan

rd

Olson, Sterrs L. A new species of pelican (Aves: Pelecanidae) from the Lower Pliocene of North
Caroling. and “PlOnda: .2. 32): Sd: oc tegaecase tie mee e. - bee eee ee ee eee

Peters, Daniel J., and Jean E. Pugh. On the entocytherid ostracods of the Brazos River basin and
adjacent coastal region of Texas

ee ee er er |

1-18

181-188

106-119

133-140

682-686

695-713

572-580

793-803

562-571

667-68 1

175—180

598-623

45-51

410-432

470-490

515-522

303-312

523-530

722-738

352-361

759-767

120-132

210—215

739-745

503-509

338-351

Rambla, Juan Pablo Blanco, and Rafael Lemaitre. Neocallicirus raymanningi, a new species of ghost
shrimp from the northeastern coast of Venezuela (Crustacea: Decapoda: Callianassidae)

Reid, Janet W., and Luis Moreno D. The western and southern distribution of Mesocyclops edax
eee) (omisinced: WoOpesada © YCIGHOIGA) 220) oa < jew nti es anne oe eens dete cde ee ccceesneee

Richart, Eric A., Jennifer A. Mercier, and Lawrence R. Heaney. Cytogeography of Philippine bats
oo PRR E eS CRED Pee ee toch ee ee Rr es ee Se ee
Rios, Rubén, and J. Emmett Dufy. Description of Synalpheus williamsi, a new species of sponge-
dwelling shrimp (Crustacea: Decapoda: Alpheidae), with remarks on its first larval stage ......
Robinson, Harold. Two new _ subtribes, Stokesiinae and Pacourininae, of the Vernonieae
2 JES DETE neopanshhs ase So hes See eee eae Sa Ne ae cae Se eee
Robinson, Harold. Revisions in paleotropical Vernonieae (Asteraceae) ................ 2. eee e ee eee eee
Rojas, Yolanda, Fernando Alvarez, and José Luis Villalobos. A new species of crayfish of the
genus Procambarus (Crustacea: Decapoda: Cambaridae) from Veracruz, Mexico ................
San Martin, Guillermo, and David Bone. Two new species of Dentatisyllis and Branchiosyllis
eeeeta Stace Sy liiac) MhOl VENEZMCIA aoa )od oc cece cae snt einen ea sate dine can atheadeaecennans
Savage, Jay M., James R. McCranie, and Larry David Wilson. A new species of rainfrog of the
Eleutherodactylus cruentus group from eastern Honduras (Amphibia: Anura: Leptodactylidae)
Smith, David G., and Carole C. Baldwin. Psilotris amblyrhynchus, a new seven-spined goby (Te-
leostei: Gobiidae) from Brazil, with notes on settlement-stage larvae .......................220005-
Solis-Marin, Francisco A., and Alfredo Laguarda-Figueras. Cucumaria flamma, a new species of
sea cucumber from the central eastern Pacific (Echinodermata: Holothuroidea) ..................
Stark, Bill P. Anacroneria from northeastern South America (Insecta: Plecoptera: Perlidae) ......
Suarez-Morales, E., and R. Palomares-Garcia. Cymbasoma californiense, a new monstrilloid (Crus-
Eacea: Ce apepeda: Monstrilloida) from Baja California, Mexico .......... 2.2.0... 0ccceeeeceenceeesee
Tavares, Marcos. Deilocerus captabilis, a new species of cyclodorippid crab from southeastern
Bia emistacea: Mecapoda: Brachyura: Cyclodorippidac) ..........-..-..2..0eseecceecesseeccenees
Thoma, Roger F., and Raymond F. Jezerinac. The taxonomic status and zoogeography of Cambarus
bartonii carinirostris Hay, 1914 (Crustacea: Decapoda: Cambaridae) ....................2..02-00e-
Villalobos, José Luis, and Fernando Alvarez. A new species of Macrobrachium (Crustacea: De-
capoda: Palaemonidae), with abbreviated development, from Veracruz, Mexico .................
Wetzer, Regina, and Niel L. Bruce. A new genus and species of sphaeromatid isopod (Crustacea)
CLLR a LEN ELE SDS LOTS el ee ee eR SR el ery eae

White, Tracy R., April K. Wakefield Pagels, and Daphne G. Fautin. Abyssal sea anemones (Cni-
daria: Actiniaria) of the northeast Pacific symbiotic with molluscs: Anthosactis nomados, a new
ee EM ONMCIIN VEST (GraVicr, L9TB)) 222. oc. 2s ec aecaacecaedencscudemesnsccesszeneascaveess

Williams, Jason D., and Christopher B. Boyko. A new species of Pseudostegias Shino, 1933
(Crustacea: Isopoda: Bopyridae: Atheliginae) parasitic on hermit crabs from Bali ...............
Winston, Judith E., and Stace E. Beaulieu. Striatodoma dorothea (Cheilostomatida: Tessaradomi-
dae), a new genus and species of bryozoan from deep water off California .......................
Young, Richard E., and Michael Vecchione. Morphological observations on a hatchling and a
paralarva of the vampire squid, Vampyroteuthis infernalis Chun (Mollusca: Cephalopoda) .....

Zottoli, Robert. Early development of the deep-sea ampharetid (Polychaeta: Ampharetidae) De-
cemunciger apalea Zottoli

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768-777

581-592

453469

541-552

216-219
220-247

396-404

319-326

787-792

433-442

778-786
70-93

189-198

141-144

97—105

746-753

368-380

637-651

714-721

313-318

66 1—666

199-209

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INDEX TO NEW TAXA

VOLUME 112

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

SAREE EEIRAN ITO CTE AAG Poteet te dee es | Re et le Apo elegy aby tarts
EN URC S pes ee ee ee ee ee ee es ON ee her ieee es

aarp ce rea uP oe S| My Bees Pte eee oth ag Pe ee et I es
TA TIY CLELAND DL, ep a a Ee ee

OL RE, STEREO ee ee Re SS a ee SE

RS ROSS RL Se eee a i ee SU ee Ue ee eS Ss Ey OS LED SSS
SESE Bey © oe a Ee ee ra, aca Pee ne ee eee ee ee ns LAr ts 1

MOLLUSCA

A Some CE Pe Seren eV 2075177 Se Se er ne Se a ee ee eee rene eee eres

ANNELIDA
Polychaeta

SLES ALAS GREE oie cele EE ae ae ea ee ee

ELIE RECTED Pen eee 0 7a RY) ea oN ana vepas Ge neeey aathd:
ce LL Et, PESTS SST ED ee SS SEERA OR IESE

NIN AE IMIR SINE SNITCH SUS Fs Sens Ae

Oligochaeta

Grania hylae
laxartus

ARTHROPODA

Crustacea

Aegla cholchol
hueicollensis

Albunea danai

Beatricesphaera
ruthae

Calyptraeotheres garthi n.c.
politus n.c.

Caprella kuroshio

Caridina clinata

640
304

625
625

653

313
S15

289

320
593

xi

Xil

Chevalia Airsuia A oe ee ae ee ee eee
Collocherides: brychins 2.08 eh a

Cymbasoma: cnliforiense: 2.02 ee ee

Detlocerus capiahiis | (2.020 te SEE ee ES bk pi 2 TE wi A eck eee

Gynodiastylis feel we skgh ae a ee

Hansentum:thoniast? Be eee Pee eee AS SENSE ee) IS I Pe

EFUOPION ne Nw ee

Nixiss (igabiaie es) o> oe ee ei A ae ectsx Ja, No. Je Se eee ee

Wee@lopagunruns seeders ao he eo

Neacrobmechitvm Jsdeacnse: 6222 ee ee

Whija sractles.. 28. 2 es Se Ee ee

WCB ONT CHa e828 ee Se ee eee rising oz aul L
azul

Neocallichirus raymanningi ____ 2

Neodoxomysis - x eae Vian.” SONI ee WM Demet Se 8

CROCS a 2 Vise TS IE Bow A Bae 2S zm fl

SGBAICRSIS ~ 22 Soe Se ee RE, Pee

Osphranticuss labronectim: mexicanum 22.2 2 ee ee ee

Palaemonella hachijo __— pas ewes =
Procambarus (Austrocambarus)-citlaiiepell 222 ee Eee

Pscudostesias maocdermothl 2s Sense 2

Piychophaltws: Kymid: te. a 2s ee ee ee eee
UNGINGHLES ) 2a ee a ee ee

Siamosquilla sexava

Sphaeromyopsis sanctaluciae

SERS CEAalA CONGIASE .cs2 0 3 ee oe a

EVE OUMDIOMIUS JOFUS oe Yost

Synalpheus williamsi =e

Insecta
Ancroneuria achagua ________ ath Sonin te or ea ras fem a
ER ns SG SEI Pe td Nl = Scene sees i =

chaima __ ae -

CTAUEIOG. © lt, We oe ae seme

GATT AIEE ne FS oo A 2 2 Poin lath A eo ae pee
PRGRAISTET to mee oe eee Fe i ee ee : : ee

POTS v=: ened EIT PETRA te teh. 2 Patt
periyja — aes Me A te I ee faz

timote _ RES ak OS RO RSET IRE OR Pe AS Gea NTI RIO AEN TE OPE
WOE ES TIE! Oe eee is ee ee
PRN SISEGEA CHIN ILORID, © ORS RN ee Ee Re renee oe eee ee =e 4 x
IRAP DCNICONE| 22s a PES Ee ee paces Ee Bauer es
NT TEES Te © 7 lt aE Ng AE Ee te RTT a a A oa gt
TREC AR TE ON i ER IO Te FoF Dc el EE eT _ sect espe tial

ECHINODERMATA

MS MNRAS TA OBIE D2 ES it A

562
182
190
141

362

169
165

697
382
746
754

572
S73

769

352
333
356

688
739
398
715

555
554

94
707
405
682
542

70
72
73
74
SD
ei
80
80
83
88

64
60

53
56

779

CHORDATA

Pisces

Eo TRG ARE TIIOTT oh EE DS ND De ES SESE ot NS IS RG Oe Pe Wee COT ee ae
A ECAB ERTL CURIS TIIICD Tessier a ie ea ep pir

SEU D GU OOO te a a eae RANE rh EE
aN SSA SMUD P ELI UNI ALECVO EOS pci map ee te es

IMIR TCAD NECUEAPCAE CA Uinta NO he MO

he SPU RVER EC DCMS COAST ets ee a A DN oe ee
ELIE DI le a a a se 2 eS I
DS PCOD IOS pct te la Ba tae ee a OE NAF, SS: SR A Oe TR
EDEL, a aE a a a ep PI RIN Be Si AEE

CTE TY co cig hg a sd er ee a eee Mee eee
ERIE DIIG I cel i 8 9 Be, 2A ON gt Sr eT RAE AAI EAs PN IO NEE EO LS. REE OO

ue F RS PRIN SS OU CT a0, CETTACLCLN CLS Seyi re oe eR tn ah de i a owes ca LL AN AN

ANTES (Ee G AEN SE SS EE a Net 1s re oe ee SS

EIU EATS SCURREME STAD os Re ner RE See eh Ra

AL TEDE ELS ESOS TR Coo EIR ee eee eRe ee el ea ne
clivorum n.c.
CS ZS IAIEN TG C eae S Ee P e eA
Soest EMULE AIS USI Cpa ore es Bis ae al Mn eer ee ten ett ete th IO eT
Tae EAA TR UK SUG TN TL Cy a a a a el etree, tote SES
saligna n.c.
Bea LISS OUD CoN Smee SAB ak Ea el sete ot ON, AMIE
silhetensis n.c.
spirei n.c.

Bechium nudicaule n.c.
rhodolepis n.c.

Centrapalus acrocephalus n.c.
SEF SBS AS a(S isa OS ae ae a I oa ee
chthonocephalus n.c.
denudatus n.c.
kirkii n.c.

Xill

X1V

origi | |. nee ea eA IS we Reon s fuer Meee ene wetea ve = Ten Os eS

@yanthilinum: albicams: 0:C... :.. ee ee
COMYZOIGES: TC. SE
hookerianwmMD.¢.) se ee ee
vernonioides n.c.

Wis feepotncanns: Hee een ya Mga a

Erlangeinae, new subtribe
Gymnantheminae, new subtulbe <2 =. eee im

Goynimanithiem ind. Ate eps 12 i A aa ee
ANGETSOMI NG: os
andrangovalense N.C. ¢ 25 oe 5 ee ee ee
antanalus n.c.

appemedicral atin Coe 5 ae duass
XE UOUCUMENN Co a hh ee oe
Wea Cg a a SE Se
belkin gina msc. =e ert re er a ee ST ee See ev OY bi
bochkiantim Niel ee SS eee
ehapelieri n.C. 222 te
OMAN OOS UUENN Le Se cr oo
COUFSU MG.) 2 ee a I ee eee eee
CPatae uhm wy. lp ee eo
CUMINGIANUIE Ihe, 2295. eet he ee eee ee eee
@y LUMENS SS Vk a eR
Cassscolumty asa 00s 5 oo ero 8 ee Sh ae és

esculentum n.c.

exsertiflorum n.c.
GSRSERUMIANCs. co ee ce se ole a ee ee
Slabertimiinit. 056), oo ee i eee ee es
hildebrandtii n.c.
humblotii n.c.
louvelii n.c.

miespilsfoliwan nic. St oe ee
Mayrianthim iC. oe Se ee 23
pectiniforme n.c.
PECIORAIE Mee! 0 ne ee ee
pleistanthum n.c.
rieppellti mG. ec! ee ee ee ee
secundifolium n.c.

theophrastifolium n.c.
thomsonianum n.c.

weghtianum n.c.
zanzibarense N.c.
zeylanicum n.c.

calyculata n.c.
hirsuta n.c.

leopoldii n.c.
FL COS re Ice Ali i Ot teh ei te AO el a Dt AIR EAT BAS wy ES Be eA
oligocephala n.c.
pinifolia n.c.

SPTLSCV ELST) TP et ar

PTT ISTE et ee
PANGRST Te ea

1 TTTREYD PSY BUL ES TS NS RE ee eae

Pa RENE AD) AP EDIE S = TCSNA UBIO EVTE@ ELA UEC ge se
LSTA SANTEE CLG Sg ek Soe a NS A a ne
(ETP SPT SLT TS ie, ee es ee ae ee eee

© OD, SE AVSTO UTS: 1 CR Ss ee ee ee ee eee
MBREERGTGHEESERI CEE GORIEVSS RTE G: 0 esate recs re lee ees ee EE er
CUE PES ETT STS Se ee i I Se I ea ee
CTS LESS TT 2S os Bek Min SUE ey ls a ee Ee eae
SE ETSELES SS TC i A en se a Ea a ee Ri ie a SE

Si IRN EPO RI al occ ey NS aie NSE ee Se
ECU LESS Cs Cee a ee ee eee eee

ERT a nN a ER ES a ee
SEL ICRA TS ON TES SoD Pe eee 2 oe ie ee

ae A Bec AE ce MEsE ART EE LEN ECTS SE ye nd Pe ae ee
stenocephala n.c.

aa ae aE oT wv 9 SL OO a ace ths ee he
cinerascens n.c.
homilanthus n.c.
karaguensis n.c.
undulatus n.c.

Pacourininae, new subtribe

Phyllocephalum microcephalum n.c.

Polydora angustifolia n.c.
Pee a St recat Ph ie Pe
chloropappa n.c.
SRN TI hin (one eee een Pee Son A er nas i ee Ne SS) a a
EAS ESTES, hip a Se a IS eR Va
serratuloides n.c.
steetziana N.c.
sylvicola n.c.

Stokesiinae, new subtribe

(TUS PERCE ce Nk I A eee eae a elk gn ee ee Re ee
aemulans n.c.
ambiguum n.c.
latifolium n.c.
musofense n.c.
SSSR Thy Gea) Ee a nee Rt NE ee ce UY OR ee a
ugandense n.c.
uncinatum n.c.
viatorum n.c.

XV

| fSeaper mee !

=,
i. a el al ae ;
£ pases n) a ‘ : ’ al al ca 0 whe odanaten,s
er la ; a See ing aN
a org — 4 a a ee eh meee late 6 e Sa ii» ation 5 |. oe
a a > = ia > @ _ ae
View aaa ten “ ; LN amy a? ee,
. ay : a ae hg ry ge) nn a oer tol ale
7 a omy, a
ae + Tle Sod ~ A —o2- ——~s - i :
,. g ( yy — > ms i 7 = —
+ ep >
a] a le -
» . = - sae bl i # i > &.
int : ;
= - __
mh ; 7
o. .
. 6 i =
¥ . j ;
=
oa i
) e “7@" =
4 - 1
}
% i
.
~The wars
; ie a's.
th.) sleeswniTr ey ee
i : =. ple : ww
Y. ibe
3
* i
f 5
—r 2A Zhe
: ie
Ss). Siem
1 - Tes oe

|

THE BIOLOGICAL SOCIETY OF WASHINGTON
1998-1999
Officers

President: Richard P. Vari Secretary: Carole C. Baldwin
President-elect: Brian F. Kensley Treasurer: T. Chad Walter

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This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(1):1-18. 1999.

Validation of Eleutherodactylus crepitans Bokermann, 1965, notes on
the types and type locality of Telatrema heterodactylum Miranda-
Ribeiro, 1937, and description of a new species of Eleutherodactylus
from Mato Grosso, Brazil (Amphibia: Anura: Leptodactylidae)

W. Ronald Heyer and Anna M. Mufoz

(WRH) Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian
Institution, Washington, D.C. 20560-0109, U.S.A.;
(AMM) Department of Biology, New Mexico State University, Las Cruces, New Mexico 88003,
U.S.A.; Current address: 1306 Alvarado, Carlsbad, New Mexico 88220, U.S.A.

Abstract.—Examination of the types of Eleutherodactylus crepitans Boker-
mann, 1965, indicates that they represent a valid species, which should be
removed from the synonymy of Eleutherodactylus fenestratus (Steindachner
1864). Eleutherodactylus crepitans is only known from the type material. Cer-
tain features of the types of Telatrema heterodactylum Miranda-Ribeiro, 1937,
are described and the type locality clarified. Analysis of the advertisement call
and morphology of a series of specimens from Chapada dos Guimaraes indi-
cates that the specimens represent a species distinct from E. fenestratus (Stein-
dachner 1864), for which no name is available. We describe this new species
as Eleutherodactylus dundeei. The frog fauna of Chapada dos Guimaraes con-
tains four distinct historical units: cerrado, chaco, Amazonian hylaea, and en-

demic.

The advertisement call of what has been
identified as Eleutherodactylus fenestratus
from Chapada dos Guimaraes, Mato Gros-
so, Brazil, was recently recorded by WRH
as part of PRODEAGRO. PRODEAGRO is
an ecological-agricultural survey of the
State of Mato Grosso, undertaken by the
State Secretariat of Planning, funded by the
World Bank, and contracted to the Sado Pau-
lo firm of consultants “‘Consorcio Nacional
de Engenheiros Consultores’”” (CENEC). As
part of the National Museum of Natural
History’s Research Training Program,
AMM analyzed the advertisement call of
the Chapada dos Guimaraes form and com-
pared it to the advertisement call of Eleu-
therodactylus fenestratus from the Amazon
basin. The results indicate that the Chapada
dos Guimaraes form is a different species
from the Amazonian E. fenestratus. To de-
termine whether there is an available name
for the Chapada dos Guimaraes form and

whether it has a distribution beyond the re-
gion of the town of Chapada dos Guima-
raes, we examined other specimens of
Eleutherodactylus from Mato Grosso, in-
cluding the type material of Telatrema het-
erodactylum Miranda-Ribeiro, 1937 and
Eleutherodactylus crepitans Bokermann,
1965. The purpose of this paper is to report
our findings.

Methods and Materials

The first recording of the Chapada dos
Guimaraes form is from USNM Tape 320,
Cut 2, Brazil, Mato Grosso, Chapada dos
Guimaraes, Estancia Monarca, recorded 30
September 1996, between 1835-1930 h, air
temperature 21.6°C, unvouchered, by
WRH, one call recorded and analyzed. The
second recording of the Chapada dos Gui-
maraes form is from USNM Tape 320, Cut
3, same data as Cut 2, except recorded at

2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1930 h, voucher USNM 507899, one call
recorded and analyzed. The recordings for
Eleutherodactylus fenestratus are from
USNM Tape 206, Cut 2, Peru, Madre de
Dios, Tambopata, recorded 6 January 1989,
1850 h, air temperature 24.0°C, voucher
USNM 342993, by Reginald B. Cocroft III,
four calls analyzed, and USNM Tape 266,
Cut 19, Peru, Madre de Dios, Tambopata,
recorded 15 November 1990, 1815 h, air
temperature 27.0°C, unvouchered, by Re-
ginald B. Cocroft III, six calls analyzed.

Call recordings were digitized using
Sound Image software at a sampling fre-
quency of 22.05 KHz and 16-bit resolution.
The digitized calls were converted into au-
dio data files using a wave converter pack-
age and analyzed as audiospectrograms and
wave forms using Computerized Speech
Research Environment software (AVAAZ
Innovations Inc., Version 4.2).

Eleven parameters were analyzed for
each call following the definitions of Heyer
et al. (1990): call duration, call rate, notes
per call, note duration, note repetition, note
pulsation, fundamental frequency, dominant
frequency, change in dominant frequency,
peak frequency, and characterization of har-
monics.

The audiospectrograms used in Fig. 1
were produced using Canary software (Cor-
nell Laboratory of Ornithology, Version
iD):

Color pattern and external morphological
data were recorded using a system of
sketches to represent all distinctive patterns
or states. These data were then summarized
and categorized into written character
states. Measurements were taken with dial
calipers and recorded to the nearest 0.1 mm
following Heyer (1984). Adult males were
determined by presence of vocal slits. Adult
females were determined by presence of
mature ova visible through the body wall.
Unclear sex determinations based on exter-
nal examination were verified by dissection.
Multivariate discriminant function analyses
were performed using SYSTAT 7.0 for
Windows.

Specimens examined are listed in Appen-
dix 1. Museum abbreviations follow Levi-
ton et al. (1985).

Results

Advertisement calls.—The ideal calls to
compare the Chapada dos Guimaraes form
with are those from topotypic Eleuthero-
dactylus fenestratus males. Eleutherodac-
tylus fenestratus (Steindachner 1864) was
described from specimens from Rio Ma-
moré, Rond6Onia and Borba, Amazonas,
Brazil. The geographically most proximate
adequate recordings we know for Amazo-
nian E. fenestratus are from Tambopata,
Peru. We use these recordings to represent
E. fenestratus for our comparisons.

As there is some variation among the
four individual frogs recorded from Peru
and Brazil, the data are summarized by in-
dividual caller (Table 1).

The frog from Tambopata, USNM Tape
206, Cut 2, has a mean call duration of 0.31
sec; 2.75 mean notes per call; and a mean
note duration of 0.07 sec. The dominant fre-
quency is the second harmonic; additional
harmonics are present, but weak. (Fig. 1a).

The second frog from Tambopata,
USNM Tape 266, Cut 19, has a mean call
duration of 0.18 sec; and a mean note du-
ration of 0.07 sec. The dominant frequency
is the second harmonic; additional harmon-
ics are present, but weak.

The single call of the frog from Chapada
dos Guimaraes, USNM Tape 320, Cut 2,
has a relatively short (0.02 sec) and weak
first note, the other 4 notes are stronger and
have a duration range of 0.03—0.04 sec. The
notes are pulsatile. The dominant frequency
is the second harmonic; additional harmon-
ics are present, but weak.

The single call of the second frog from
Chapada dos Guimaraes, USNM Tape 320,
Cut 3, has a relatively short (0.02 sec) and
weak first note, all other notes are stronger
with a duration of 0.04—0.05 sec. The notes
are pulsatile. The dominant frequency is the

VOLUME 112, NUMBER 1

10

Kilohertz

Frequency i

0.4

1.0

0.8

0.6

Time in Seconds

Fizs I.

A. Advertisement call of Eleutherodactylus fenestratus, USNM Tape 206, Cut 2. B. Advertisement

call of Chapada dos Guimaraes form, USNM Tape 320, Cut 3.

second harmonic; additional harmonics are
present, but weak. (Fig. 1b).

There is some variation between individ-
ual male vocalizations from the same lo-
cality. The variation observed between the
recordings from Tambopata might be ac-
counted for by differences of temperature
of calling males, at least in part. As a gen-
eral rule for frogs, increases in temperature
result in decreased call and note duration

and increased note repetition rate (Schnei-
der et al. 1984; Duellman & Trueb 1986).
The three degree difference of ambient tem-
perature between the two recordings of the
Peruvian specimens is in the predicted di-
rection for call and note duration and note
repetition rate. Note, however, that the re-
cordings from Chapada dos Guimerdaes
were recorded at lower temperatures than
those of Peru, yet these Brazilian calls had

4 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Advertisement call data for four Eleutherodactylus specimens from Tambopata, Peru and Chapada

dos Guimaraes, Brazil.

Geographic area Peru
USNM Tape and Cut 206, 2
Call duration (sec) 0.21—0.36
Number of notes/call 2-3
Note duration (sec) 0.06—0.09
(Mean) note repetition rate per sec tis:
Fundamental frequency range 1636-2411
Peak fundamental frequency 1720
Dominant frequency range 2842-4392
Peak dominant frequency Sis Sil

shorter note durations and higher note rep-
etition rates than the Peruvian calls, indi-
cating that temperature differences do not
explain the differences between the Brazil-
ian and Peruvian calls.

The calls from Brazil and Peru do not
differ in terms of carrier frequencies; they
are broadcasting in the same frequency
band. The calls differ markedly in terms of
temporal packaging (Table 1), such as num-
ber of notes per call, individual note dura-
tion, and note repetition rate. Penna (1997)
among others, has experimentally demon-
strated that the kinds of temporal differenc-
es found between the Tambopata and Cha-
pada dos Guimaraes frogs are recognized as
species differences by frogs.

Unfortunately, to date, there are no re-
cordings of what we consider to be Ama-
zonian Eleutherodactylus fenestratus from
the hylaea of northern Mato Grosso. In fact,
the only other published recordings for E.
fenestratus are from Amazonian Bolivia
(Marquez et al. 1995) and are very similar
to the recordings from Tambopata. Ronald
I. Crombie made three recordings in Ron-
d6énia, Brazil in which a single call of an E.
fenestratus is in the background. In two of
them (USNM Tape 55, Cut 5 and USNM
Tape 56, Cut 5, both from Alto Paraiso),
the call sounds as though it consists of two
notes, but can not be confirmed on an au-
diospectrogram due to the relative weak-
ness of the call and strength of the back-
ground noise. The call on USNM Tape 56,
Cut 8 from Santa Cruz da Serra is a bit

Peru

Brazil

Brazil

266, 19 320, 2 320, 3
0.16—0.20 0.29 0.53
2 5) 8
0.05—0.08 0.04 0.05
OT 14.7 1337
1464—2497 1894—2239 1808-2497
2067 2067 2153
2842—4220 3014—4220 3273-4737
3617 3692 3875

stronger and the audiospectrogram indicates
it is comprised of two notes at a dominant
broadcast frequency of 3300 Hz. The calls
from Rond6nia, as expected, match the
calls from Tambopata rather than the calls
from Chapada dos Guimaraes.

The advertisement call evidence is con-
sistent with recognizing the Chapada dos
Guimaraes form as a species distinct from
Eleutherodacytlus fenestratus.

Morphology.—There are two available
names for Eleutherodactylus from Mato
Grosso that potentially could apply to the
Chapada dos Guimaraes form. There are
also a number of Eleutherodactylus fenes-
tratus-like specimens from Mato Grosso.
The purpose of this section is to compare
the Chapada dos Guimaraes form with
these specimens to determine whether the
Chapada dos Guimaraes form occurs more
broadly within the State of Mato Grosso
and to determine whether there is an avail-
able name for it.

The materials for comparison comprise
the following (Fig. 2 for Mato Grosso lo-
calities; Appendix 1 for specimen lists): the
2 types of Telatrema heterodactylum Mi-
randa-Ribeiro, 1937; the 3 type specimens
of Eleutherodactylus crepitans Bokermann,
1965; 11 specimens from Chapada dos Gui-
maraes; 2 specimens from Fazenda Santa
Edwiges; 1 specimen from Jacubim; 2 spec-
imens from Barra do Tapirapés; and speci-
mens considered to be E. fenestratus; 3
specimens from Sao Jose do Rio Claro; 8
specimens from Apiacas, 48 specimens

VOLUME 112, NUMBER 1

60
8 e
10 lo, 9
La
|
16 °
60

Fig. 2.
city of Cuiaba.

from Juruena (measurement data taken for
10 females and 1 male), 1 specimen from
Alto Juruena, 46 specimens from Aripuana
(measurement data taken for 10 males and
females), and 28 specimens from Tambo-
pata and Pakitza, Madre de Dios, Peru (the
specimens from the nearby locality of Pak-
itza were added to those from Tambopata
to bring the sample sizes up to at least 10
males and 10 females).

Patterns: The dorsal snout patterns dem-
onstrate as much intra- as inter-population
variation and are not discussed further. The
other patterns analyzed demonstrate some
levels of interpopulation variation (Tables
2-8). In some cases, the sample sizes are

3
52
fe)
6
6
5
@
e 2
®
“16
4 52
@

Outline map of the State of Mato Grosso. See text for key to locality numbers. Open square is the

adequate to conclude that the variation is
meaningful (e.g., the differences in loreal
region pattern between the Mato Grosso
populations of Eleutherodactylus fenestra-
tus (Table 3, samples 7—10) and the Peru-
vian sample of E. fenestratus. In other cas-
es, the differences are suggestive, but any
conclusions are compromised by sample
sizes. For example, there are two aspects of
light mid-dorsal stripes that are impacted by
small sample size. The first is that there is
variation in the presence condition of the
stripe. It may either be narrow or broad, and
it may extend from the tip of the snout to
the vent or from the interocular area to the
vent. Because there were relatively few in-

ON

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Interorbital markings. Locality numbers refer to those in Fig. 2 (9a not mappable). A = Almost
absent; B = Outlined only, not with expansion; C = Solid, not with expansion; D = Solid with single prominent
posterior expansion; E = Solid with bifid prominent posterior expansion. Data are number of individuals dem-

onstrating the various states (question mark indicates specimen faded, state uncertain).

C. dos E. fenestratus

Taxa E. heterodactylus E. crepitans Guimaraes ? ue iy

Locality 2 + 5 6 7 8 9 9a 10 Peru
A 5
B Z 12 9 2
€ iF 3 11 1 1 a 8) 6 9 1 11 9
D 6
E 6

4 From illustration 2a (Miranda-Ribeiro 1937).

dividuals with light mid-dorsal stripes, the
information had to be collapsed for com-
parison. Second, the sample size for the
Chapada dos Guimaraes form is too small
to conclude that it lacks light mid-dorsal
stripes, even though all 11 specimens ex-
amined lack them. There are three adequate
samples of E. fenestratus to compare with
the Chapada dos Guimaraes form: Aripua-
na, Juruena, and Peru. For Aripuana, the
probability of an individual having a mid-
dorsal stripe is 0.35, thus the probability of
not having a stripe is 0.65. If we use that
probability to ask whether the Chapada dos
Guimaraes population could have the same
occurrence of stripes, the answer is no:
0.65'' = P < 0.001 (exact probability test).
However, when the population data from
Juruena and Peru are used, the answer is
yes: for Juruena, the probability of not hav-
ing a stripe is 0.91, for Peru 0.92, with P
= 0.35 and 0.40 respectively. Thus, based
on the low frequency of occurrence of mid-
dorsal stripes in the Juruena and Peru sam-

ples, if the Chapada dos Guimaraes form
has mid-dorsal stripes at those same fre-
quencies of occurrence, then the sample
size is not adequate to have found individ-
uals with that condition.

Morphological features: All but three in-
dividuals scored for tarsal fold variation
had discernible tarsal folds (Table 9). It is
likely that the three individuals thus scored
actually have them but are preserved in
such a manner that they are not discernible.
The two individuals of Eleutherodactylus
crepitans with discernible tarsal folds have
a very different morphology from all other
specimens examined (Table 9). The varia-
tion in toe fringe condition is continuous
(Table 10). The fringed condition is weakly
fringed, with noticeable basal toe webbing
(the fringe is not as well-developed as in
some species of Leptodactylus, for exam-
ple). The variation in tubercles on the sole
of the foot (Table 11) suggests that E. cre-
pitans is distinctive (in contrast to com-
pletely distinct) from the other samples, and

Table 3.—Loreal region patterns. Locality numbers refer to those in Fig. 2 (Locality 9a not mappable). A =
No pattern; B = Incomplete stripe in front of eye; C = Dark stripe from eye to nostril, entire loreal region may
be dark. Data are number of individuals exhibiting various states. Pattern not discernible for specimen from

locality 6.
C. dos E. fenestratus
Taxa E. heterodactylus E. crepitans Guimaraes ? z
Locality 2 6 7 8 9 9a 10 Peru
A 8
B 1 2 4 9 1 1 13
C 1 a) 2 3 8 12 20 7

VOLUME 112, NUMBER 1

~—

Table 4.—Supratympanic fold patterns. Locality numbers refer to those in Fig. 2 (9a not mappable). A =
Fold not accentuated with dark brown band; B = Fold accentuated with interrupted dark brown band; C = Fold
accentuated with solid dark brown band. Data are number of individuals demonstrating the various states (ques-
tion mark indicates state uncertain due to fading or perspective of illustration).

C. dos E. fenestratus

Taxa E. heterodactylus’ E. crepitans Guimaraes is ? oe
Locality 1 2 3 6 f 8 9 9a 10 Peru

A ue 2

B 3 2 12

C 9 Z ? 3 8 pi 1 21 14

@ From illustration 2a (Miranda-Ribeiro 1937).
the variation in the Chapada dos Guimaraes Taxonomy

specimen tubercles is distinctive from the
variation observed in the E. fenestratus
samples.

Measurements: Male and female sizes
vary among the samples (Table 12). Two
points are worth noting, although additional
samples are necessary to verify them statis-
tically. There is no sexual size dimorphism
in the Eleutherodactylus crepitans sample,
whereas sexual size dimorphism is pro-
nounced in all other samples that contain
both sexes. The Chapada dos Guimaraes
form is smaller than the Eleutherodactylus
fenestratus samples analyzed.

The results of multivariate discriminant
function analyses for males and females
(analyzed separately) agree in that the
Eleutherodactylus crepitans specimens are
clearly distinct from all other individuals
analyzed (Fig. 3). The rest of the popula-
tions analyzed indicate that there is differ-
entiation among them, but the degree of
variation is consistent with both intra- and
inter-specific variation based on similar
studies in other groups of frogs.

Telatrema heterodactylum Miranda-Ri-
beiro, 1937.—Alipio Miranda-Ribeiro de-
scribed this species from “‘Matto-Grosso—
Caceres, 2 exemplares colhidos na gruta
dita ‘Fazendinha’ Comm. Rondon (1937:
67).’’ His son Paulo Miranda-Ribeiro des-
ignated specimen 106A in the collection of
the Museu Nacional, Rio de Janeiro, as the
lectotype (1955:411). The second speci-
men, the paralectotype, which also had the
number 106 originally, was subsequently
recatalogued as 5089.

Dr. P. E. Vanzolini (pers. comm.) was
able to locate A. Miranda-Ribeiro’s locality.
Miranda-Ribeiro was a member of the Ron-
don Commission expedition and he collect-
ed the specimens in question himself. Cac-
eres, as used by Miranda-Ribeiro, refers to
the municipality, which at the time of the
Rondon Commission expedition was very
extensive. Vanzolini found in the Rondon
Commission itinerary that Miranda-Ribeiro
travelled to “‘Fazendinha”’ and returned to
Caceres in the same day and that ‘‘Fazen-
dinha’’ was near a place called Pirizal. Van-

Table 5.—Mid-dorsal stripe patterns. Locality numbers refer to those in Fig. 2 (9a not mappable). A = Absent;
B = Present. Data are number of individuals demonstrating the states.

C. dos E. fenestratus
Taxa E. heterodactylus E. crepitans Guimaraes ? ? wv
Locality 1 2 = 6 7 8 = 9a 10 Peru
A 1 3 11 2 2 5 8 44 l 34 26
B 4 12 2

(o/6)

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 6.—Dorsal patterns. Locality numbers refer to those in Fig. 2 (9a not mappable). A = Indistinct or
uniform; B = Dark symmetrical spots; C = Dark chevron in medial scapular region followed by dark blotches
posteriorly; D = Dark W-shaped mark in medial scapular region followed by dark blotches posteriorly; E =
Extended dark W-shaped pattern (additional strophes to next in series from inverted V to W) in medial scapular
region followed by dark blotches posteriorly. Data are number of individuals demonstrating the various states

(question mark indicates uncertain condition due to fading).

C. dos E. fenestratus

Taxa E. heterodactylus E. crepitans Guimaraes ? uy

Locality I 2 3 + 6 7 8 9 9a 10 Peru
A 2 rm 1 3 Wi 1 9 iS
B pe 1 3 4
C L 4 2 2
D 1 9 2 9 6 ES
EB D

4 From illustration 2a, b (Miranda-Ribeiro 1937).

zolini located a “‘Fazendinha”’ that not only
meets these conditions, but is in an area
known to have sandstone caves (grutas =
caves) at 16°00’S, 57°36’W (Fig. 2, Local-
iyo)

Lynch & Duellman (1997:225) recog-
nized Eleutherodactylus heterodactylus
(Miranda-Ribeiro, 1937) as a distinct spe-
cies and included it in the Eleutherodacty-
lus binotatus species-group. As Lynch and
Duellman did not examine the types, we of-
fer the following observations.

Both types are faded such that most fea-
tures of any color patterns are no longer
discernible. The lectotype is in poor con-
dition, the paralectotype is in worse condi-
tion. The paralectotype is very brittle and
fragile and disintegrates more each time it
is handled. The lectotype is the (noticeably)
larger of the two, and the only specimen
measured for purposes of this paper. WRH
recorded the measurements (in mm) of the
lectotype as: SVL 24.6 (contrasted to Mi-

randa-Ribeiro’s measurement of 27); head
length 8.6; head width 8.5; eye—nostril dis-
tance 3.5; eye—eye distance 5.1; greatest
tympanum diameter 2.4; thigh 13.9; shank
14.1; foot 12.0; width of 3rd finger disk 1.4;
width of 4th toe disk 0.7. The lectotype has
slits in the floor of the mouth, but they ap-
pear to be cuts, not vocal slits. The sex of
the lectotype can not be determined with
certainty without dissection, which given
the poor condition of the specimen was
considered inappropriate.

The following support recognition of Te-
latrema heterodactylum as a distinct and
valid species of Eleutherodactylus, confirm-
ing Lynch & Duellmans’ (1997) assess-
ment.

The disks on fingers 3 and 4 are very
large, much larger than the disk on the
fourth toe, and the shape is ovate (Fig. 4a).
The disks on fingers 3 and 4 of the other
Eleutherodactylus examined are expanded,
but either the same size or just larger than

Table 7.—Posterior face of thigh patterns. Locality numbers refer to those in Fig. 2 (9a not mappable). A =
Uniform; B = Finely mottled around vent, rest uniform; C = Finely mottled; D = Mottled. Data are number

of individuals demonstrating the various states.

C. dos Guima- E. fenestratus
Taxa E. heterodactylus’ E. crepitans raes Y ?
Locality I 2 3 4 6 7 8 gy) 9a 10 Peru
A 2 7 Z 3 df 18 20 5
3 3 1 3 1 1 8

B
C 1
D

VOLUME 112, NUMBER 1

\O

Table 8.—Belly patterns. Locality numbers refer to those in Fig. 2 (9a not mappable). A = Almost immaculate;
B = Noticeably flecked or mottled. Data are number of individuals demonstrating the states. Specimen from

locality 6 too faded to score.

C. dos

E. fenestratus

Taxa E. heterodactylus E. crepitans Guimaraes us ? ?

Locality 1 2 5 6 7 8 2 9a 10 Peru
A 1 3 10 2 l 6 10 11 28
B 1 S 2 11 1 10

the disks on toe 4 and they are broadly tri-
angular in shape (Fig. 4b, c).

The type illustrations (most likely that of
the lectotype, Miranda-Ribeiro 1937, Fig.
2a—b) show a pair of ill-defined light lon-
gitudinal stripes from behind the eye to the
sacrum. This feature is not seen in any of
the other Eleutherodactylus examined for
this study.

The type locality is characterized by a
very open vegetation formation lacking gal-
lery forests. Eleutherodactylus fenestratus
is a forest denizen and the Chapada dos
Guimaraes form inhabits well-developed
gallery forests. Eleutherodactylus crepitans
also occurs in open habitats, but is morpho-
logically very distinct from FE. heterodac-
tylus.

One other morphological feature will re-
quire fresh material to evaluate. There is no
indication of any tarsal fold or other tarsal
decoration on either the lectotype or para-
lectotype, suggesting that, if in fact the tar-
sus lacks a fold, that would be another fea-
ture distinguishing the species from all oth-
er Mato Grosso Eleutherodactylus.

Eleutherodactylus heterodactylus (Miran-
da-Ribeiro 1937) is thus far known only
from the original two specimens collected
by Miranda-Ribeiro from ‘“‘Fazendinha.”’
All other members of the Eleutherodactylus
binotatus group, to which Lynch & Duell-
man (1997) assigned E. heterodactylus, oc-
cur in eastern Brazil, primarily in the At-
lantic Forest Morphoclimatic Domain. In-
clusion of E. heterodactylus in this group
does not make zoogeographic sense.

Status of Eleutherodactylus crepitans
Bokermann, 1965.—Werner C. A. Boker-
mann described the species based on three
specimens from Sao Vicente, Mato Grosso.
The holotype and allotype are now in the
collections of the Museu de Zoologia,
Universidade de Sao Paulo and the other
paratype is in the Museu Nacional, Rio de
Janeiro.

John Lynch (1980:8) synonymized
Eleutherodactylus crepitans Bokermann,
1965 with E. fenestratus (Steindachner
1864). Lynch did not examine the speci-
mens of E. crepitans (1980:6). Lynch ar-
gued that because Bokermann was unaware

Table 9.—Tarsal fold states. Locality numbers refer to those in Fig. 2 (9a not mappable). A = Absent; B =
Short, oblique, lying more than length of inner metatarsal tubercle from inner metatarsal tubercle; C = Mod-
erately short, less than 2 length of tarsus, lying less than length of inner metatarsal tubercle from inner metatarsal
tubercle; D = Long, greater than % length of tarsus, lying less than length of inner metatarsal tubercle from
inner metatarsal tubercle. Data are number of individuals demonstrating the various states. Data not taken for
one desiccated individual from locality 10.

C. dos
Taxa E. heterodactylus E. crepitans Guimaraes ? ? ?
Locality 1 2 3 = 5 6 7 8 9 9a 10

E. fenestratus

Peru

A 1

NO —
—

B
Cc 10 Z ] ] 5 8 21 ] ee 28
D

10 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 10.—Toe fringe states. Locality numbers refer to those in Fig. 2 (9a not mappable). A = Toes free;
B = Toes with lateral ridges; C = Toes with modest lateral fringes. Data are number of individuals demonstrating

the states.
C. dos E. fenestratus
Taxa E. heterodactylus’ E. crepitans Guimaraes Y ey
Locality 2 3 4+ 6 7 8 9 9a 10 Peru
A 1 3 1 5 5
B 3 5 8 6 I7
es 8 be Ps 3 5) 12 1 10 6

that E. fenestratus was a valid species, Bok-
ermann compared his new species with in-
appropriate species from Ecuador, from
which they were very distinct. However,
had Lynch examined the type material of E.
crepitans, he would have found that they
differ from E. fenestratus as well.

The following argue for recognizing
Eleutherodactylus crepitans as a valid spe-
cies.

Eleutherodactylus crepitans has _ short,
oblique tarsal folds, separated from the in-
ner metatarsal tubercle by a distance greater
than the length of the fold. Eleutherodac-
tylus heterodactylus apparently lacks tarsal
folds. Eleutherodactylus fenestratus and the
Chapada dos Guimaraes form have longer
(but still relative?y short) tarsal folds, par-
allel to the long axis of the tarsus, separated
by a short distance (less than length of the
fold) from the inner metatarsal tubercle.

Eleutherodactylus crepitans does not
have marked sexual dimorphism in size.
Both Eleutherodactylus fenestratus and the
Chapada dos Guimaraes form show marked
sexual size dimorphism.

The overall body shape of Eleutheroda-
cytlus crepitans differs markedly from that

of E. fenestratus and the Chapada dos Gui-
maraes form (Fig. 3).

Bokermann (1965:264) made a point of
Stating that the specimens of Eleutherodac-
tylus crepitans were obtained in grass-shrub
vegetation in the middle of large blocks of
granite from such a dry habitat that he
would never have expected to find frogs
there. The immediate vicinity of Sao Vi-
cente does have a granitic intrusion of rath-
er small extent and is the only such granitic
habitat in Mato Grosso (Brasil 1982). It
would not be surprising if E. crepitans is
limited to this distinctive habitat.

Bokermann (1965:264) indicated that
males called at night from sandy hiding
places among rocks. He described the call
as consisting of two notes in sequence, like
a short and coarse click, repeated many
times but irregularly (translation by P. E.
Vanzolini). The two note call of Eleuther-
odactylus crepitans is more similar to
Eleutherodactylus fenestratus than the Cha-
pada dos Guimaraes form, but the frequent
calling of E. crepitans is different from the
very infrequent calls of both E. fenestratus
and the Chapada dos Guimaraes form.

Questionable status specimens.—The

Table 11.—Maximum number of tubercles on sole of foot. Locality numbers refer to those in Fig. 2 (9a not
mappable). Data are number of individuals demonstrating the states.

C. dos
Taxa E. heterodactylus’ E. crepitans Guimaraes ?
Locality 2 4
6) 2
1 8 pe
Z 1
3 1 1
4 2

E. fenestratus

6 7 8 9 9a 10 Peru
1
2 3) 9 9 23
1 pe 9 9 >
1 3 1 2:

VOLUME 112, NUMBER 1

11

Table 12.—Adult size. Locality numbers refer to those in Fig. 2 (9a not mappable). Data are SVL ranges in mm.

E. hetero- C. dos
Taxa dactylus_ E. crepitans Guimaraes ?
Locality 1 2 3 4 5
Females 30 34-36
Males 28-29 22-27 30
Largest juveniles 25 30

name Hylodes gollmeri bisignata Werner,
1899, which based on geography should be
considered as an available name for the
frogs considered in this paper, and speci-
mens of Eleutherodactylus from three lo-
calities in Mato Grosso can not be unequiv-
ocally assigned to either Eleutherodactylus
crepitans, fenestratus, heterodactylus, or
the Chapada dos Guimaraes form. Adver-
tisement calls are not available at present
for any of the specimens involved. At this
time, they are best left as indeterminate.

Werner (1899) gave a brief description of
Hylodes gollmeri bisignata without locality
information in the type description. Haupl
& Tiedemann (1978) indicated that the ho-
lotype, NMW 16502, in the Naturhisto-
rischen Museums Wien collection was from
Chaco, Bolivia. Lynch & Duellman (1997)
considered bisignatus to be a synonym of
Eleutherodactylus fenestratus. Based on the
fact that the Chaco and Amazonian hylaea
frog faunas are virtually distinct from each
other at the species level, the validity of E.
bisignatus should be reconsidered. The
original description is too brief to be of help
in evaluating this suggestion. Dr. Lynch ex-
amined the type of H. g. bisignata, but his
original notes are not in the appropriate
folder in his archives (pers. comm.). The
Naturhistorisches Museum Wien now has a
policy of not loaning types, precluding our
examination of the type for this paper. We
suspect that bisignatus is a species of
Eleutherodactylus distinct from E. fenestra-
tus. If it is the same as any species that
occurs in Mato Grosso, it might be conspe-
cific with E. heterodactylus, based on most
similar (but still quite different) habitat oc-
currences.

—

E. fenestratus

~~

6 7 8 9 9a 10 Peru
34 38-41 37-40 42 37-44 40-52
30 26-32 27-34
29

The two specimens from the Panatanal
locality of Fazenda do Santa Edwiges (Fig.
2, Locality 4) could either be the same as
the Chapada dos Guimaraes form or a dis-
tinct species. The smaller specimen
(MZUSP 71103) has a tarsal fold like that
of the Chapada dos Guimaraes form, but
the larger specimen (MZUSP 71104, which
looks like it died before it was preserved)
appears to have a much longer tarsal fold.

The single specimen from Jacubim
(MZUSP 4277) has completely mottled
thighs with a large pattern, such that the
white blotches almost cover the same total
area as the darker ground hue. The thigh
pattern differs markedly from other speci-
mens discussed in this paper, but is closer
to that of specimens identified as Eleuth-
erodactylus cf peruvianus (e.g., MZUSP
80854, 80856-80857, 80859, 80863) from
the Amazon forest locality of Apiacas. Ja-
cubim, however, is in an area of Cerrado-
seasonal forest contact (Fig. 2, Locality 5).

The two specimens from Barra do Tapi-
rapés are very faded and their proper tax-
onomic allocation may never be ascertained
until fresh specimens from the locality are
available. The fading extends to the poste-
rior surfaces of the thighs, but it does ap-
pear that the thighs are mottled in a similar
fashion to the pattern observed in MZUSP
4277 from Jacubim, with which they may
be conspecific.

Status of the Chapada dos Guirardes
form.—As documented above, the Chapada
dos Guimaraes Eleutherodactylus repre-
sents a distinct species from all other
known Eleutherodactylus, which we de-
scribe as:

12 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

MALES FEMALES
ee 6
e

4
ew
te = Q
O B
©

O
Te

; )
-10 -5 O 5 Te)

FACTOR |

Fig. 3. Discriminant function analysis results for males and females. Minimum polygons contain all individuals
for sample sizes >3. Dots = Eleutherodactylus crepitans; A = sample from Aripuana, Mato Grosso; B = individual
from Barra do Tapirapés, Mato Grosso; C = sample from Chapada dos Guimaraes, Mato Grosso; J = individual
(male) and sample (females) from Juruena, Mato Grosso; P = sample from Peru; 5 = individual from Jacubim,

Mato Grosso.

Eleutherodactylus dundeéi,
new species
Figs. 1b, 5

Holotype.—MZUSP 79834, adult male
from Brazil: Mato Grosso; Chapada dos
Guimaraes, Veu da Noiva, Rio Coxipozin-
ho, approximately 15°25’S, 55°47'W. Col-
lected by Miriam H. Heyer, W. Ronald Hey-
er, and Liliam Patricia Pinto on 25 Sep
1996.

Paratopotypes.—MZUSP 79835-79837
(1 female, 2 males), same data as holotype.

Paratypes.—All from Brazil: Mato Gros-

A B C

3

Fig. 4. Diagrammatic outlines of shape of third
finger disk based on free-hand sketches. A. Holotype
of Telatrema heterodactylum, actual width of disk 1.4
mm. B. Holotype of Eleutherodactylus crepitans, ac-
tual width 0.9 mm. C. Chapada dos Guimaraes form,
actual width 1.3 mm (based on USNM 507897).

so; Chapada dos Guimaraes; MNRJ 19785
(female), collected by Erminia (UFF) on 24
Oct 1987; MZUSP 76237 (Guvenile), an
originally unnumbered specimen from the
Werner C. A. Bokermann collection col-
lected by A. Sebben and A. Schwartz on 15
Jun 1988; MZUSP 85614 (male), originally
WCAB 15546 collected by M. Alvarenga,
E M. Oliveira, and Werner C. A. Boker-
mann from 15-25 Nov 1963; USNM
507897—-507898 (female, male), Casa de
Pedra, collected by Miriam H. Heyer, W.
Ronald Heyer, Liliam Patricia Pinto on 28
Sep 1996; USNM 507899 (male, call
voucher for Fig. 1b), Estancia Monarca,
collected by Miriam H. Heyer, W. Ronald
Heyer, Liliam Patricia Pinto on 30 Sep
1996; USNM 507900 (female), Escola
Evangelica de Buriti, collected by Miriam
H. Heyer, W. Ronald Heyer, Liliam Patricia
Pinto on 1 Oct 1996.

Diagnostic definition (scheme established
by Lynch, e.g., 1979, of numbered charac-
ter states followed here for ease of com-
parison).—Skin of upper surfaces finely

VOLUME 112, NUMBER 1

Aare

*
oo

‘
he 23
Sayan el .
4, cent ae
~
r

ry " “Si 44 ‘ ak

*

O32»
Th 8
Oh

ver)
s) pyeces

Fig. 5.

Dorsal view of paratype of Eleutherodactylus dundeei, new species, USNM 507899. This specimen

has the most contrasting pattern among the type series. Other specimens have similar, but fainter markings.

pebbled with few to many scattered white
fleshy tubercles, tubercles denser posteri-
orly, skin of throat, chest, and anteriormost
belly smooth, rest of belly weakly areolate;
tympanum distinct, its horizontal diameter
about %—¥; eye opening diameter; snout su-
belliptical in dorsal view, rounded in pro-
file; upper eyelid width equal to or (usually)
slightly smaller than interorbital distance,

with same fine pebbling of back with scat-
tered, low, flattened, light tubercles; no cra-
nial crests; vomerine teeth in two small
patches well posterior to choanae, separated
from each other by about the width of a
single tooth patch; males with vocal slits
and single subgular vocal sac, either notice-
ably expanded and wrinkled in preservation
or not; male nuptial pads of two weak light

13

14 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

glandular patches; first finger about same
size or just longer than second; fingers bear-
ing disks (largest on III and IV); largest
disks broader than long; fingers free or with
weak lateral keels; 2—7 light spots indicat-
ing weakly developed ulnar tubercles (tu-
bercles themselves visible in only some in-
dividuals); short tarsal fold, about % length
of tarsus, otherwise tarsus smooth; heel
smooth to granular, lacking tubercle(s); 2
metatarsal tubercles, inner oval, 2—3 times
size of outer; 0—2 fleshy plantar tubercles;
toes free or (usually) bearing weakly to well
developed lateral keels; individuals with
well-developed keels with basal toe web-
bing; predominantly brown frog, no flash
colors; posterior surfaces of thighs flesh
color-purplish in life, tan/brown in preser-
vative; adults small, males 22—27 mm, fe-
males 34—36 mm SVL.

Eleutherodactylus dundeei is most simi-
lar to and most likely to be confused with
Eleutherodactylus fenestratus. Eleuthero-
dactylus dundeei is smaller (males 22—27
mm, females 34—36 mm SVL) than E. fe-
nestratus (males 26—34 mm, females 37—52
mm SVL). The belly of E. fenestratus is
either entirely smooth; or weakly areolate
only on the lateralmost portions, but not
areolate posteriorly; the belly of E. dundeei
is weakly areolate at least posteriorly. Male
E. dundeei have two weakly developed
light glandular nuptial asperities on each
thumb; male E. fenestratus have a single,
better developed light glandular nuptial as-
perity on each thumb.

Description of holotype.—Snout shape
subelliptical in dorsal outline, rounded in
profile; canthus rostralis sharply rounded;
lip weakly flared; upper eyelid width about
equal interocular distance; no external in-
dications of cranial crests; tympanum dis-
tinct, horizontal diameter including annulus
about % eye opening diameter, horizontal
diameter < vertical diameter; vomerine
teeth in two small patches, well posterior to
rounded choanae, separated from each other
by about width of a single tooth patch; vo-
cal slits present, elongate; vocal sac single,

subgular, slightly expanded externally in
preservation; finger lengths I just
>II<IV<III deft finger IV mostly miss-
ing); finger tips on fingers I and II slightly
expanded, small, round, lacking circumfer-
ential grooves, fingers III and IV with ex-
panded disks, triangular, broader than long,
with circumferential groove and upper sur-
face weakly to noticeably notched; fingers
lacking noticeable lateral keels; no finger
webbing; palmar tubercle large, ovate-
heart-shaped, larger than proximate ovoid
thenar tubercle, one well-developed super-
numerary palmar tubercle associated with
each digit; subarticular tubercles distinct,
rounded, single, each thumb with 2 weakly
developed but distinct patches of light glan-
dular nuptial excrences; outer ulnar region
with a series of 6—7 light spots, apparently
with low tubercles in life, but only one
weakly visible on each arm; supratympanic
fold distinct; no other distinct glands or
folds on body; upper eyelid weakly pebbled
with several light tubercles, rest of back
texture weakly pebbled with scattered light
fleshy tubercles; throat smooth, discoidal
belly fold distinct, chest and adjoining belly
to about mid-belly smooth, sides of and
posterior belly weakly areolate, ventro-pos-
terior surfaces of thighs areolate, rest of
ventral limb surfaces smooth; toe lengths
I<WI~WI<V<IV (condition B as used in
Lynch & Duellman 1997, appendix III);
disks on toes I and V small, round, disks
on toes II, II, IV increasing in size, largest
disks triangularly ovate, broader than long,
upper surfaces emarginate or weakly
notched; sides of toes keeled, strongest on
inner sides of toes II, III, IV, forming basal
web, not encompassing the basal subarti-
cular tubercles, best developed between
toes II, II, [V; outer round metatarsal tu-
bercle about 4 size of ovate inner metatar-
sal tubercle; tarsal fold distinct, short, about
% length of tarsus, separated from inner
metatarsal tubercle by a distance of about
% length of tarsal fold, rest of outer tarsus
smooth; heel smooth, lacking tubercle(s) or
calcar; plantar surface with one prominent

VOLUME 112, NUMBER 1

fleshy tubercle; subarticular tubercles mod-
erately developed, weakly pungent.

Coloration in alcohol: A tan frog with
brown markings. Tip of snout lightest, ex-
panding into somewhat darker but light tri-
angle bounded by canthus rostralis and dark
interorbital mark, with a few brown dots
and lines. Dark canthal stripe in front of eye
only extending about % distance to nostril.
Loreal region almost uniform brown. Upper
lip indistinctly barred. Supratympanic fold
dark brown. Faint interorbital bar, defined
by straight solid brown line anteriorly, very
weakly defined posteriorly as a shallow U
with a few dark dots in the interorbital
mark. A faint dark W-shaped mark in the
pre-scapular area followed by a median,
faint, inverted Y-shaped mark between
scapular and sacral region, followed by two
median irregular sacral chevrons. Notice-
ably dark pair of lateral spots behind eyes
in scapular region. Upper limbs tan with ir-
regular darker cross bands. Series of 6—7
light spots on outer ulnar region. Throat
speckled with melanophores. Chest and bel-
ly light, almost immaculate, but with a very
few scattered melanophores. Posterior sur-
faces of thighs with very fine mottle pattern
on area just around vent, otherwise uniform
tan. Outer tarsus and sole of foot dark
brown.

Measurements of holotype.-—SVL 23.6
mm, head length 9.6 mm, head width 7.9
mm, eye opening diameter 3.0 mm, eye—
nostril distance 3.0 mm, eye—eye distance
4.9 mm, horizontal tympanum diameter 1.8
mm, vertical tympanum diameter 1.9 mm,
thigh length 11.6 mm, shank length 13.3
mm, foot length 11.7 mm, width of 3rd fin-
ger disk 1.0 mm, width of 4th toe disk 0.7
mm.

Variation.—Most details of variation
have been described in the preceding sec-
tions. Additional measurement variations
are (broken down by sex only when range
of one not subsumed in other): head length
39-42% SVL; head width 33-38% SVL;
eye—nostril distance 12-14% SVL; eye—eye
distance 20—22% SVL for males, 18-19%

15

for females; tympanum diameter 7—9%
SVL; thigh length 48-53% SVL; shank
length 52-57% SVL; foot length 47-54%
SVL for males, 46—53% for females; width
of 3rd finger disk 3—4% SVL; width of 4th
toe disk 3—4% SVL.

Color in life.—Iris bright bronze above,
dull bronze below; posterior face of thigh
flesh color-purplish; belly white with yel-
low wash; no flash colors (based on USNM
507897-—507898).

Advertisement call.—Described in pre-
vious section (Fig. 1b).

Etymology.—Named for Dr. Harold A.
Dundee for his personal, financial, and sci-
entific contributions to the field of herpe-
tology.

Ecological notes.—Data are available for
eight individuals. Five of them were col-
lected in the late morning on dead leaves
or under 5 cm diameter rocks on the
ground, four of these 3—6 m from a river
bank in rainforest-like gallery forest
(MZUSP 79834-79837), one of these on
dead leaves on the ground on the bank of
a tree-shaded pond (USNM 507900). Two
individuals were taken shortly after dark in
cerrado gallery forest 2 m from a 2 m wide
stream either on the surface of the soil
(USNM 507897) or on a 0.3 m diameter log
(USNM 507898). The call voucher (USNM
507899) was taken at night from secondary
cerrado vegetation on a leaf of a herbaceous
plant, less than 1 m above ground.

Distribution.—All known specimens
have been collected from within 25 km of
the town of Chapada dos Guimaraes in the
limited region identified as having meso-
philic seasonal forest vegetation cover in
the Cartograma de Vegetacao do Estado de
Mato Grosso, produced by the Cons6rcio
Nacional de Engenheiros Consultores S.A.,
published by the Governo do Estado de
Mato Grosso (Fig. 6). There are other is-
lands of similar vegetation cover in the
Same region (Fig. 6) which might be ex-
pected to contain Eleutherodactylus dun-
deei. We predict that either E. dundeei is
restricted to the known Chapada dos Gui-

16 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

54

Fig. 6. Major vegetation types in portion of the State of Mato Grosso surrounding Chapada dos Guimaraes.
Black areas are mesophilic seasonal forests. Stippled areas are dense cerrado (cerradao) and forest/cerrado contact
regions. Rest of region is covered by various forms of cerrado vegetation (except cerradao). Open square is the

city of Cuiaba.

Eleutherodactylus dundeei is thus far only known from the mesophilic seasonal forest block identified as 3
on the map, but other nearby blocks should be explored for the species. Map redrawn from “‘Cartograma de
Vegeta¢gao do Estado do Mato Grosso,”’ produced by Consércio Nacional de Engenheiros Consultores S.A.

maraes vegetation type block or to it and
one or more of the additional habitat islands
identified in Fig. 6.

Discussion

Eleutherodactylus fenestratus, with the
removal of Eleutherodactylus crepitans and
dundeei, has an Amazonian rainforest dis-
tribution (assuming Eleutherodactylus bi-
signatus is not a synonym of E. fenestra-
tus). There has not been an adequate study
of variation for E. fenestratus. The limited
data we analyzed indicate that there is at
least interpopulational variation in E. fenes-
tratus. Analyzing variation throughout the
entire range of the species would be an ob-

vious next step to understand the signifi-
cance of the variation we found.

Chapada dos Guimaraes has a very in-
teresting frog fauna from a zoogeographical
perspective. The landscape surrounding the
Chapada is cerrado. There are expected cer-
rado frog species at Chapada dos Guimar-
aes, such as Bufo paracnemis, Scinax fus-
covarius, Leptodactylus labyrinthicus, and
Physalaemus nattereri. There are also Cha-
co and Amazonian elements represented.
For example, the Chaco species Leptodac-
tylus chaquensis was collected syntopically
with the Amazonian species Leptodactylus
mystaceus in 1996. Finally, there are at least
two species thus far known only from Cha-
pada dos Guimaraes, Phyllomedusa cen-

VOLUME 112, NUMBER 1

tralis Bokermann, 1965 and Eleutherodac-
tylus dundeei. Obviously, the frog fauna of
Chapada dos Guimaraes represents a rich
and complicated history, worthy of much
more detailed study. There is actually a fair
amount of information available for frogs
from Chapada dos Guimaraes to base such
a study upon. Herbert Huntingdon Smith
collected materials reported on by E. D.
Cope (1887) and housed at the Academy of
Natural Sciences in Philadelphia. At least
one of the species reported, Leptodactylus
petersii, has not been collected since from
Chapada dos Guimaraes (Heyer 1994:79,
Cope’s Leptodactylus brevipes = L. peter-
sii). Boulenger (1903) listed nine species
collected by the Percy Sladen Expedition to
central Brazil. The species Boulenger listed
as Hylodes gollmeri is probably the species
we describe as Eleutherodactylus dundeei.
All other species of Boulenger’s list are
plausible (with taxonomic adjustments) ex-
cept for Ceratophrys cristiceps, which
should be re-examined. Werner C. A. Bok-
ermann and colleagues made collections
which now (for the most part) are at
MZUSP. MZUSP also has materials col-
lected by Drs. Rita Kloss and Francisca C.
do Val. Adao Cardoso also collected at
Chapada dos Guimaraes, and those speci-
mens are housed at the Universidade Estad-
ual de Sao Paulo in Campinas. As Chapada
dos Guimaraes is now a tourist attraction,
it is likely that there are other incidental
collections waiting study in various collec-
tions as well.

Acknowledgments

PRODEAGRO and CENEC made the
fieldwork possible during which the record-
ings of Eleutherodactylus dundeei were ob-
tained. Without these recordings, this paper
would not have been initiated. Drs. John D.
Lynch and P. E. Vanzolini critically re-
viewed the manuscript. AMM thanks the
National Museum of Natural History for
the opportunity to participate in the 1997
Research Training Program. WRH thanks

iw

Dr. P. E. Vanzolini and the Museu de Zool-
ogia da Universidade de Sao Paulo for char-
acteristic help and access to the collections.
Dr. Ulisses Caramaschi facilitated exami-
nation of specimens in the collections of the
Museu Nacional, Rio de Janeiro. WRH’s re-
search for this paper was supported by the
Neotropical Lowlands Research Program,
National Museum of Natural History, Dr.
Richard P. Vari, Principal Investigator.

Literature Cited

Bokermann, W. C. A. 1965. Trés novos batraquios da
regiao central de Mato Grosso, Brasil (Amphib-
ia, Salientia).—Revista Brasileira de Biologia
25:257-264.

Boulenger, G. A. 1903. List of the batrachians and rep-
tiles collected by M. A. Robert at Chapada,
Matto Grosso, and presented by Mrs. Percy
Sladen to the British Museum. (Percy Sladen
Expedition to Central Brazil).—Proceedings of
the Zoological Society of London 1903 II(1):
69-70.

Brasil. Ministério das Minas e Energia. 1982. Projeto
RADAMBRASIL. Folha SD.21 Cuiaba; geo-
logia, geomorfologia, pedologia, vegetacao e
uso potencial da terra. Levantamento de Recur-
sos Naturais, Vol. 26. Rio de Janeiro, 544 pp.,
5 maps.

Cope, E. D. 1887. Synopsis of the Batrachia and Rep-
tilia obtained by H. H. Smith, in the Province
of Mato Grosso, Brazil.—Proceedings of the
American Philosophical Society, Philadelphia
24:44-60.

Duellman, W. E., & L. Trueb. 1986. Biology of am-
phibians. McGraw-Hill Book Company, New
York, 670 pp.

Haupl, M., & E Tiedemann. 1978. Typenkatalog der
Herpetologischen Sammlung.—Kataloge der
wissenschaftlichen Sammlungen des Naturhis-
torischen Museums in Wien. Vertebrata. 2:1—35.

Heyer, W. R. 1984. Variation, systematics, and zoo-
geography of Eleutherodactylus guentheri and
closely related species (Amphibia: Anura: Lep-
todactylidae).—Smithsonian Contributions to
Zoology 402:1—42.

. 1994. Variation within the Leptodactylus pod-

icipinus-wagneri complex of frogs (Amphibia:

Leptodactylidae).—Smithsonian Contributions

to Zoology 546:1—124.

. A. S: Rand, C: A. G..Cruz,.O, L. Peixoto, &
C. E. Nelson. 1990. Frogs of Boracéia.—Ar-
quivos de Zoologia 31:231—410.

Ceviion, A.B. kK” EH. Gibbs, Jr, E. ‘Heal, & C. E.
Dawson. 1985. Standards in herpetology and

18 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ichthyology: Part I. Standard symbolic codes
for institutional resource collections in herpe-
tology and ichthyology.—Copeia 1985:802-—
832.

Lynch, J. D. 1979. Leptodactylid frogs of the genus
Eleutherodactylus from the Andes of southern
Ecuador.—University of Kansas Museum of
Natural History Miscellaneous Publication 65:
1-62.

. 1980. A taxonomic and distributional synop-

sis of the Amazonian frogs of the genus Eleuth-

erodactylus.—American Museum Novitates

2696: 1-24.

, & W.E. Duellman. 1997. Frogs of the genus
Eleutherodactylus in western Ecuador: System-
atics, ecology, and biogeography.—The Univer-
sity of Kansas Natural History Museum Special
Publication 23:1—236.

Marquez, R., I. De la Riva, & J. Bosch. 1995. Adver-
tisement calls of Bolivian Leptodactylidae (Am-
phibia, Anura).—Journal of Zoology, London
237:313-336.

Miranda-Ribeiro, A. 1937. Alguns batrachios novos
das collecg¢des do Museu Nacional.—O Campo
May 1937:66—69.

Miranda-Ribeiro, P. 1955. Tipos das espécies e subes-
pécies do Prof. Alipio de Miranda Ribeiro de-
positados no Museu Nacional (Com uma rela-
cao dos géneros, espécies e subespécies descri-
tos).—Arquivos do Museu Nacional, Rio de Ja-
neiro 42:389-—417.

Penna, M. 1997. Selectivity of evoked vocal responses
in the time domain by frogs of the genus Ba-
trachyla.—Journal of Herpetology 31:202—217.

Schneider, H., T. S. Sofianidou, & P. Kyriakopoulou-
Sklavounou. 1984. Bioacoustic and morpho-
metric studies in water frogs (genus Rana) of

Lake loannina in Greece, and description of a
new species (Anura, Amphibia).—Zeitschrift
fiir zoologischen Systematik und Evolutionsfor-
schung 22:349—366.

Werner, FE 1899. Beschreibung neuer Reptilien und Ba-
trachier—Zoologischer Anzeiger 22:479—484.

Appendix 1
Specimens examined

Eleutherodactylus crepitans.—Brazil: Mato Grosso;
Sao Vicente, 15°38’S, 55°23’W, MNRJ 3985 (para-
type), MZUSP 73671 (allotype), 85628 (holotype).

Eleutherodactylus dundeei.—See species account.

Eleutherodactylus fenestratus.—Brazil: Mato Gros-
so; [Alto Juruena—see preceding Juruena], Apiacas,
09°34'S, 57°23'W, MZUSP 80851-80853, 80855,
80858, 80860-80862; Aripuana, 10°24’S, 59°19’W,
MZUSP 80643-80648, 80652-80654, 86410; Alto Ju-
ruena (locality is the upper Rio Juruena and cannot be
assigned specific coordinates), MZUSP 85540; Jurue-
na, 10°18’S, 58°32’W, MZUSP 86153-86200; Sao José
do Rio Claro, 13°26’S, 56°44’W, MZUSP 86358—
86360.

Peru: Madre de Dios; Pakitza, 11°58’S, 71°14’W,
USNM 333014, 342618—342622, 342851-—342853,
345172; Tambopata, 12°50’S, 69°17’W, USNM
222268, 247298-247304, 247630—247631, 268941—
268944, 314901, 332438—332439, 342993.

Eleutherodactylus heterodactylus.—Brazil: Mato
Grosso; Fazendinha, near Pirizal, 16°00’S, 57°36’W,
MNRJ 106 (lectotype), 5089 (paralectotype).

Eleutherodactylus sp.—Brazil: Mato Grosso, Barra
do Tapirapés, 10°39’S, 50°36’W, MZUSP 85645-
85646; Jacubim, 13°20’S, 52°09’W, MZUSP 4277; Fa-
zenda Santa Edwiges, 16°38’S, 55°11'W, MZUSP
71103-71104.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(1):19-—39. 1999.

A new genus and species of frog from Bahia, Brazil (Amphibia:
Anura: Leptodactylidae) with comments on the zoogeography of the
Brazilian campos rupestres

W. Ronald Heyer

Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560-0109, U.S.A.

Abstract.—A new genus and species of frog, Rupirana cardosoi, is described
from the northern Espinhacgo Range in the State of Bahia, Brazil. The new
genus shares most character states with the genus Thoropa, but cladistic anal-
ysis of morphological data indicates that most of these shared features are
primitive states. The cladistic analysis indicates that Rupirana and Thoropa do
not have a close sister-group relationship with each other. Rupirana cardosoi
is another addition to the many species endemic to the campos rupestres of the
Espinhacgo Range. The amphibians of the campos rupestres show a much stron-
ger biogeographical affinity with the Atlantic Forest biota than other groups
studied; these other groups show a much stronger affinity with the biota of the
diagonal of open formations running from northeast Brazil (caatingas-cerrados)
to Argentina and Paraguay (Gran Chaco).

Several years ago, Dr. Miguel T. Ro-
drigues brought my attention to a series of
unusual frogs he had collected at two lo-
calities in the State of Bahia, Brazil, asking
me if I knew what they were. I did not. Drs.
Rodrigues and P. E. Vanzolini kindly put
the specimens at my disposal for further
study, but for various reasons, detailed
study was delayed. Shortly after I had ex-
amined aspects of the myology and skele-
ton of the taxon, I had occasion to evaluate
some of the problematical specimens of the
Werner C. A. Bokermann collection as they
were being catalogued into the Museu de
Zoologia da Universidade de Sao Paulo col-
lection. I found a series of the same taxon
from a third locality in Bahia. All known
localities are in the distinctive campos ru-
pestres formation of Brazil. The purpose of
this paper is to demonstrate that the new
frog differs from all other known leptodac-
tylid genera, provide a description for it,
and briefly comment on its relationships
and zoogeography.

Methods and Materials

One male (MZUSP 65204) was superfi-
cially dissected to obtain myological infor-
mation on jaw, hyoid, and thigh muscula-
ture for the characters used in a previous
study of leptodactylid relationships (Heyer
1975). A second female specimen (MZUSP
68959) was cleared-and-stained using the
double-staining technique for cartilage and
bone (Dingerkus & Uhler 1977) to evaluate
osteological features.

For the species description, measurement
data were taken with a dial calipers to the
nearest 0.1 mm following the definitions in
Heyer et al. (1990).

Relationships were analyzed with PAUP
3.1 (Swofford 1993).

Comparison with Leptodactylid Genera

Lynch (1971) still provides the most
complete data set for leptodactylid genera,
providing a baseline for comparison with
subsequent studies. He recognized four
New World subfamilies of the family Lep-

20 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

gp
eee

oN

Fig. 1. Vertebral column of Rupirana cardosoi,
MZUSP 68959. Cartilage stippled. Scale line = 2 mm.

todactylidae: Ceratophryinae; Elosiinae;
Leptodactylinae, and Telmatobiinae. The
Australian and African leptodactyloids
Lynch treated as subfamilies of the family
Leptodactylidae are now recognized as the
separate families Myobatrachidae and He-
leophrynidae, respectively (Frost 1985;
Duellman & Trueb 1986).

The Bahia frog is not a member of the
subfamilies Ceratophryinae, Elosiinae, or
Leptodactylinae as defined by Lynch
(1971). Rather than document this state-
ment with a comparison of all states Lynch
used, only a single character is used as an
example. Ceratophryinae members have
widely expanded transverse processes of
the anterior presacral vertebrae; the Bahia
frog does not (Fig. 1). Elosiinae species
have a pair of dermal, scute-like glandular
pads on the dorsal surface of each digital
disk; the Bahia form lacks digital disks.
Leptodactylinae genera have a bony sternal
style; the sternum is a cartilaginous plate in
the Bahia frog (Fig. 2).

Lynch (1971:112—113) listed nine char-

acters defining the subfamily Telmatobiinae
(character numbers are those used by
Lynch): (1) sternum cartilaginous; (2) ver-
tebral shield lacking; (3) transverse pro-
cesses of the anterior presacral vertebrae
not widely expanded; (9) when present,
maxillary teeth blunt and pedicellate; (35)
mandible lacking odontoids; (39, the 38 is
a typographical error on p. 113) m. petro-
hyoideus anterior and m. sternohyoideus in-
sert on the lateral edge of hyoid plate; (48)
eggs laid in water, in terrestrial situations,
or in bromeliads. The egg deposition site is
unknown for the Bahia frog; it agrees in the
rest of the character states listed for the sub-
family.

Lynch (1971) defined five tribes of Tel-
matobiinae and one genus he was unable to
assign to any of these tribes: Alsodini;
Eleutherodactylini; Grypiscini; Odonto-
phrynini; Telmatobiini; and the genus Scy-
throphrys.

The Bahia frog cannot be assigned to the
Eleutherodactylini, Grypiscini, Odonto-
phrynini, Telmatobiini, or Scythrophrys.
Again, for sake of brevity, only examples
are used to document this statement. Mem-
bers of the Eleutherodactylini have rounded
sacral diapophyses and males lack cornified
nuptial asperities; the Bahia frog has flat-
tened sacral diapophyses and the males
have cornified nuptial asperities. In species
of the Grypiscini, the frontoparietal fonta-
nelle is not exposed; in the Bahia frog it is
(Fig. 3). Members of the Odontophrynini
have short transverse processes of the pos-
terior presacral vertebrae; the Bahia frog
has long processes (Fig. 1). Members of the
Telmatobiini have cervical cotyles that are
narrowly separated with two distinct artic-
ular surfaces (Lynch’s type I, 1971:54); the
Bahia frog has widely spaced cervical co-
tyles with two distinct articular surfaces
(Fig. 4, Lynch’s type I, 1971:53). Scythro-
phrys has rounded sacral diapophyses and
a concealed tympanum; the Bahia frog has
flattened sacral diapophyses and an ex-
posed, well-developed tympanum.

Lynch (1971:123—124) defined the Tribe

VOLUME 112, NUMBER 1

21

Bisa 2.
line = 1 mm.

Alsodini based on 16 characters (numbers
are those of Lynch): (3) transverse process-
es of posterior presacral vertebrae long; (4)
cervical cotylar arrangements type I or II;
(5) cervical and second vertebrae not fused;
(6) cranial bones not involved in dermos-
tosis; (7) omosternum present, moderately
large; (8) sacral diapophyses somewhat en-
larged; (9) maxillary teeth blunt, pedicel-
late; (12) facial lobe of maxilla deep, not
exostosed; (17) nasals not in contact with

Ventral portion of pectoral girdle of Rupirana cardosoi, MZUSP 68959. Cartilage stippled. Scale

frontoparietals; (18) frontoparietal fontanelle
exposed, moderate-sized; (19) frontopari-
etal not fused with prodtic; (21) temporal
arcade lacking; (37) alary processes of hy-
oid plate on narrow stalks; (42) male with
cornified nuptial asperities on thumb; (45)
outer metatarsal tubercle present, inner
metatarsal tubercle normal; (46) larvae with
median vent. Larvae are unknown for the
Bahia frog. In all other characters, except
(8), the Bahia frog matches exactly the def-

22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Ries yp
tanelle hatched. Scale line = 1 mm.

inition of the subfamily. In the Bahia frog,
the sacral diapophyses are flattened (not
rounded), but they are not really enlarged
(Fig. 1). However, the differences involved
in the shape of the sacral diapophyses are
minor and alone do not argue for a distinct
tribe status for the Bahia frog.

Lynch (1971) included four genera in his
concept of Alsodini: Batrachyla, Eupso-
phus, Hylorina, and Thoropa (in 1978 he
restricted the Tribe Batrachylini to the gen-
era Batrachyla and Thoropa). Of these, the
Bahia frog is most similar to Thoropa. Eup-
sophus and Hylorina have the type II cer-
vical cotylar arrangement; the Bahia frog

Dorsal view of skull of Rupirana cardosoi, MZUSP 68959. Cartilage stippled. Frontoparietal fon-

has type I. The maxillary arch of Batrachy-
la is incomplete and the quadratojugal is
absent; the maxillary arch of the Bahia frog
is complete (Figs. 3—4).

Lynch (1971:129—130) defined the genus
Thoropa by 31 characters (following his
numbers): (4) cervical cotylar arrangement
type I; (10) alary processes of premaxillae
directed dorsally and slightly anteriorly, rel-
atively narrow at base; (11) palatal shelf of
premaxilla very narrow with elongate pal-
atal process present [I could not find where
Lynch defined what a palatal process of the
maxilla was and I do not find such a struc-
ture for either Thoropa or the Bahia frog];

VOLUME 112, NUMBER 1

23

Fig. 4. Ventral view of skull of Rupirana cardosoi, MZUSP 68959. Cartilage stippled. Scale line = 1 mm.

(14) maxillary arch complete, quadratojugal
present; (15) nasals relatively large with
moderately long maxillary processes, sep-
arated medially; (16) nasals not in contact
with maxillae or pterygoids; (22) epiotic
eminences relatively long and narrow or
short and stocky, carotid artery passes dor-
sal to skull bones; (24) zygomatic ramus of
squamosal relatively short; (25) otic ramus
of squamosal moderately long, no otic
plate; (26) squamosal-maxillary angle 50—
70°; (27) columella present; (28) prevomers
relatively small, entire, separated medially,
toothed; (29) palatine long and narrow, ex-
panded laterally, separated medially; (30)
sphenethmoid entire, extending anteriorly

to posterior edge of nasals or not reaching
nasals; (31) anterior ramus of parasphenoid
broad, keeled medially, extending anteriorly
to prevomers; (32) parasphenoid alae ori-
ented at right angles to anterior ramus of
parasphenoid, relatively short, not over-
lapped laterally by median ramus of pter-
ygoids; (33) pterygoids large, anterior rami
in long contact with maxillae, not reaching
palatines; (34) occipital condyles large or
small, not stalked, moderately to widely
separated medially; (36) terminal phalanges
T-shaped; (40) m. depressor mandibulae in
two slips; (41) pupil horizontal; (42) males
with median subgular vocal sac; (43) body
lacking glands; (44) tongue large, oval, pos-

24 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

terior edge free; (45) toes lacking webbing,
bearing lateral fringes, digital tips bulbous,
somewhat dilated, first finger shorter than
second; (46) larvae with 2/3 tooth rows, la-
bial papillae broadly interrupted anteriorly;
(48) eggs large, few in number, laid in lotic
situations; (49) males 19-78, females 24—
70 mm SVL; (50) tympanum visible exter-
nally; (51) tadpoles with greatly flattened
and attenuate bodies and tails.

There is no egg placement or larval in-
formation for the Bahia frog.

Several features require further comment.
The Bahia frog does not have a medial keel
of the parasphenoid, nor does the paras-
phenoid extend anteriorly to the vomers
(Fig. 4), as stated for character 31 in Tho-
ropa. However, Lynch’s Fig. 86 (1971:130)
does not show the conditions he described.
In re-examining the material used by
Lynch, I find that in Thoropa miliaris (KU
92855), the parasphenoid has a weak me-
dial keel and does extend to the vomers, but
in both T. lutzi (KU 92908), and T. petro-
politana (KU 92862), the parasphenoid
lacks a median Keel and extends far short
of the vomers (as in the Bahia frog). For
character 32, in the Bahia frog the paras-
phenoid is overlapped by the median rami
of the pterygoids, contrasting with Lynch’s
statement for Thoropa. However, Lynch’s
Fig. 86 (1971:130) shows overlap, which I
also find in T. lutzi (KU 92908), 7. miliaris
(KU 92855), and 7. petropolitana (KU
92862). The terminal phalanges of the Ba-
hia frog, character 36, are not distinctly T-
shaped as in Thoropa, but either expanded
or weakly T-shaped (Fig. 5). Although the
shape of the terminal phalanges in the Ba-
hia frog differs from the distinct T-shaped
condition in Thoropa, this condition could
be interpreted as part of a morphological
continuum in degree of expansion of the
terminal phalanges. The depressor mandib-
ulae muscle, character 40, of the Bahia frog
is the DFSQat condition (three slips, the
third a small slip originating from the tym-
panic annulus), contrasting with the DFSQ
state listed for Thoropa by Lynch. I find

ras

Fig. 5. Terminal finger phalanges. Left to right,
Thoropa lutzi, from cleared-and-stained specimen KU
92908, third finger; Thoropa petropolitana, from
cleared-and-stained specimen KU 92862, third finger;
Rupirana cardosoi, from cleared-and-stained specimen
MZUSP 68959, third finger; Rupirana cardosoi, from
partially dissected wet specimen MZUSP 76035,
fourth finger. Scale line = 1 mm.

both the two and three slip conditions in
Thoropa, however (T. miliaris, DFSQ
[USNM 97765]; 7. petropolitana, DFSQat
[USNM 164135)]).

There have been two additional species
of Thoropa described since Lynch’s (1971)
publication (T. megatympanum, T. saxatilis)
which require modification of character 45
in that genus. Thoropa lutzi and T. petro-
politana have weak lateral toe ridges that
are less well-developed than in the Bahia
frog. Thoropa miliaris has weak ridges,
similar to the condition of the Bahia spe-
cies. The toe ridges of 7. megatympanum
are even weaker than those of 7. Jutzi and
T. petropolitana. Thoropa saxatilis lacks
any toe ridge or fringe. The digital tips of
the Bahia frog are narrow, not dilated. In
both 7. lutzi and T. saxatilis, the digit tips
are dilated and the finger tips are more di-
lated than the toe tips. In T. miliaris, the
digital tips are bulbous and somewhat di-
lated (Lynch’s description), and there is no
size difference between the finger and toe
tips. In both 7. megatympanum and T. pe-
tropolitana, the tips of the digits are slightly
dilated but not particularly bulbous and the
finger and toe tips are equal sized. The first
finger is the same length as the second in
the Bahia species, the same condition that
I find in T. miliaris (contra Lynch). The first
finger is shorter than the second in T. lutzi,
T. petropolitana, and T. saxatilis. In T. me-
gatympanum, the first finger is either just

VOLUME 112, NUMBER 1

shorter than the second or the first and sec-
ond are of equal length.

Summarizing thus far, the Bahia frog is
most similar to species in the genus Tho-
ropa, but differs consistently by two char-
acters used by Lynch (1971): shape of the
terminal phalanges (but see discussion of
character 36 above) and dilation of digit
tips.

In a preliminary analysis of relationships
of leptodactylid genera, I (Heyer 1975)
came to quite a different conclusion from
Lynch (1971) regarding the relationships of
Thoropa. 1 did not recognize formal taxo-
nomic groupings such as subfamilies and
tribes, but used informal grouping names,
which are equivalent in scope to Lynch’s
subfamilies. Specifically, my concept of a
grypiscine clade combined the content of
Lynch’s Grypiscinae plus Elosiinae with the
genera Paratelmatobius and Thoropa. The
reasons for this very different viewpoint are
due to two factors: although Lynch and I
analyzed most of the same characters, there
were some differences; and I attempted a
phylogenetic analysis, whereas Lynch’s
analysis was phenetic.

I analyzed 37 characters. For the states
that I listed for Thoropa (1975, table B, p.
51), the following differ from the condi-
tions found in the Bahia species. Character
5, state B, disks on toes (see discussion of
Lynch’s character 45 above). Character 6,
state A, no tarsal fold, flap, or tubercle. The
Bahia frog has a tarsal fold, however, re-
examination of 7. miliaris indicates that
specimens have weak tarsal folds as do
specimens of T. saxatilis. Character 9, state
D, toes with lateral fringe (see discussion
of Lynch’s character 45 above). Character
12, state B, depressor mandibulae origin
from dorsal fascia, squamosal, and otic re-
gion only (see discussion of Lynch’s char-
acter 40 above). Character 16, state B, omo-
hyoideus insertion on hyoid body and fascia
between posteromedial and posterolateral
processes. In the Bahia species, the omo-
hyoideus inserts only on the hyoid plate.
Character 17, state B, iliacus externus ex-

25

Pis..6.
MZUSP 68959. Cartilage stippled. Scale line = 1 mm.

Hyoid apparatus of Rupirana cardosoi,

tends %—% anterior on ilium. In the Bahia
species, the extension is >%4, about 90%.
Character 20, state B, adductor longus
poorly developed, inserting on adductor
magnus, covered by sartorius. In the Bahia
frog, the adductor longus is well-developed,
inserting on the knee. Character 30, state C,
anterior process of hyale absent. In re-ex-
amining material, I found that the process
is clearly absent in JT. miliaris (USNM
97765). In T. petropolitana (USNM
164135), there is a medial, but not an an-
terior swelling of the hyale at its most an-
terior extent, contrasting markedly with the
well-developed anterior process found in
the Bahia species (Fig. 6). Character 34,
state A, sacral diapophyses expanded. In
the Bahia frog, the sacral diapophyses are
flattened, but not expanded (Fig. 1). In re-
examining the two Thoropa species that are
similar in size to the Bahia species (T. lutzi,
KU 92908, T. petropolitana, KU (92862),
the sacral diapophyses are flat, as in the Ba-
hia species, and differ slightly from the Ba-
hia species in being expanded. Character
35, state B, terminal phalanges T-shaped is
the same as Lynch’s character 36 (see dis-
cussion above).

The Bahia species clearly shares most
features of all known leptodactylid genera

26 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

with the genus Thoropa. The question is
whether the definition of Thoropa should be
expanded to include the Bahia species, or
whether the Bahia frog should be treated as
a genus distinct from Thoropa.

The Bahia species differs from all known
Thoropa in seven characters (of the char-
acters examined for this study). For two of
these, the definition of the genus Thoropa
would only have to be slightly modified to
incorporate the Bahia frog: degree of ex-
pansion of the flattened sacral diapophyses
and shape of the terminal phalanges. The
other five characters would require a more
drastic redefinition of Thoropa: toe disks;
insertion of the omohyoideus muscle;
length of the iliacus externus muscle; de-
velopment of the adductor longus muscle;
anterior process of the hyale. These latter
five characters demonstrate different con-
ditions involving toe, hyoid, and thigh mor-
phologies. These kinds of differences are
consistent with the Bahia species being on
a separate evolutionary track from Thoro-
pa. Thus, if the Bahia form were described
as a species of Thoropa, evolutionary re-
lationships would be obscured (also see
section on relationships, below). Therefore,
in order to emphasize the evolutionary dis-
tinctiveness of the Bahia species, it is de-
scribed as:

Rupirana, new genus
Figs. 1-6

Type-species.—Rupirana cardosoi, new
species.

Diagnosis.—The only leptodactylid gen-
era that share the combination of sternum
cartilaginous, widely spaced cervical coty-
les with two distinct articular surfaces, long
transverse processes of posterior presacral
vertebrae, flattened sacral diapophyses, and
an exposed frontoparietal fontanelle are Ba-
trachyla, Thoropa, and Rupirana. Batra-
chyla has an incomplete maxillary arch; in
Rupirana the maxillary arch is complete.
Thoropa has well-developed T-shaped ter-
minal phalanges, dilated digit tips, and

lacks an anterior process of the hyale; Ru-
pirana does not have well-developed T-
shaped terminal phalanges, has narrow dig-
ital tips, and has an anterior process of the
hyale.

Definition.—Pupil horizontal; tympanum
distinct; vocal sac single, subgular, slightly
expanded externally; male thumb with one
extensive and one small patch of kerati-
nized sandpaper-like asperities; body with-
out well-defined glands; digital tips narrow,
not dilated; tarsus with a tarsal fold; outer
metatarsal tubercle small; inner metatarsal
tubercle large, ovoid, not cornified; toes
with lateral fringes, joined at base of toes.

Adductor mandibularis muscle condition
adductor mandibulae posterior subexternus
only; depressor mandibulae condition
DFSQat; geniohyoideus muscle contiguous
medially; anterior petrohyoideus insertion
on lateral edge of alary process and hyoid
plate; sternohyoideus muscle insertion en-
tirely near edge of hyoid body; omohyoi-
deus muscle insertion entirely on hyoid
plate; iliacus externus muscle extending al-
most to anterior tip of ilium; tensor fasciae
latae muscle insertion posterior to iliacus
externus muscle on iliac bone; interior and
exterior portions of semitendinosus muscle
uniting in a common distal tendon distally,
exterior portion larger than interior; adduc-
tor longus muscle about same size and
shape as sartorius muscle, inserting on knee
and adductor magnus.

Quadratojugal present, contacting maxil-
la; frontoparietals not meeting medially, ex-
posing moderate-sized fontanelle; vomerine

_teeth present; occipital condyles widely

separated; anterior process of hyale present;
alary processes of hyale on narrow stalks;
posterior sternum a cartilaginous plate, ter-
minally expanded and bifid; last presacral
vertebra just narrower than sacrum; sacral
diapophyses flattened, not noticeably ex-
panded; dorsal crest of ilium present; ter-
minal phalanges barely expanded, not or
weakly T-shaped.

Etymology.—From the Latin rupes, rock,
and rana, frog. The gender is feminine. The

VOLUME 112, NUMBER 1

ZF

Fig. 7.

name is to highlight the association of this
genus with the campos rupestres of Brazil.
Content.—Monotypic.

Rupirana cardosoi, new species
Figs. 7-8

Holotype.—MZUSP 65203, adult male,
27 Sep 1987, Mucujé, Bahia, Brazil,
13°00’S, 41°23'W, Miguel T. Rodrigrues.

Paratopotypes.—MZUSP 65202 (adult
male), 65204 (adult male), 65205 (juve-
nile), same data as holotype; MZUSP
68959 (cleared and stained adult female),
68960 Guvenile), 2 Oct 1990, Miguel T.
Rodrigues.

Paratypes.—MZUSP 68961-68962
(adult females), Morro do Chapeu, Bahia,

Dorsal view of male paratype of Rupirana cardosoi, MZUSP 76032.

Brazil, 3 Oct 1990, 11°33’S, 41°09’W, Mi-
guel T. Rodrigues; MZUSP 76017-76018
(adult males), 76023—76030 (adult males),
76031 (adult female), 76032-76034 (adult
males), 76035 (adult female, skull re-
moved), 76036 (adult male), 76037 (Guve-
nile male), 76038-76040 (adult males),
USNM _ 519755-519757 (adult males),
519758 (adult female), Andarai, Bahia,
Brazil, 12°48’S, 41°20’W, 19-22 Nov 1968,
Werner C. A. Bokermann, Francisco M.
Oliveira, and B. D. Silva.

Diagnosis.—As for genus.

Description of holotype.—Snout round in
profile and from above; nostrils anterolat-
eral, near tip of snout; canthus rostralis in-
distinct; loreal obtuse; tympanum distinct,

28 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fige 3:

rounded, about % diameter of eye; supra-
tympanic fold distinct from behind eye to
shoulder, bordering tympanum dorsally;
tongue elongate, triangular, with slight
emargination on anterior edge; vomerine
teeth in two small transverse patches in line
with posterior borders of small, round cho-
anae, vomerine tooth patches separated
from each other by about width of single
tooth patch; vocal slits elongate; vocal sac
single, subgular, indicated externally by lat-
eral skin folds/wrinkles; finger lengths
I~II~IV<II; fingers free of web, very
slight lateral ridges; tips of fingers rounded,
not expanded; palmar tubercle ovoid, just
smaller than ovoid thenar tubercle; subar-
ticular tubercles moderately developed,
slightly pungent; no supernumerary tuber-
cles; accessory palmar tubercles present,
two in line with each digit; thumb with ex-
tensive keratinized sandpaper-appearing as-
perity from penultimate phalanx to base of
thumb, a second ovoid patch on inner half
of thenar tubercle; forearm slightly hyper-
trophied; dorsum, including upper eyelid,
smooth with many scattered large white tu-
bercles; body lacking any obvious glands;
venter smooth except for areolate poster-
oventral thigh surfaces; belly disk weakly

Profile of head of holotype of Rupirana cardosoi, MZUSP 65203.

indicated; toe lengths I<II<V<III<IV; tips
of toes rounded, not expanded; toes with
weak lateral ridges expanded into weak
fringes at base of inner sides of toes Il and
III resulting in trace of strap-shaped basal
web between toes I, II, and III; inner meta-
tarsal tubercle small, round, about % size of
small, ovoid outer metatarsal tubercle; dis-
tinct but relatively weak tarsal fold extend-
ing about % length of tarsus; sole of foot
smooth; subarticular tubercles moderate,
pungent.

SVL 27.3 mm; head length 9.8 mm, head
width 10.3 mm; horizontal diameter of tym-
panum 2.1 mm, including annulus; distance
from eye to posterior edge of naris 2.8 mm;
internarial distance 2.5 mm; thigh length
12.5 mm; shank length 13.4 mm; foot
length 14.3 mm.

In preservative, dorsal body brown with
darker brown markings, upper limbs tan
with brown markings; upper snout with a
few irregular darker blotches and dots; ir-
regular interorbital bar connecting with li-
chenous network behind eyes, rest of dor-
sum with mostly elongate blotches and
broad stripes; upper limbs weakly cross-
banded; distinct broad dark brown canthal
stripe extending more faintly from nostril to

VOLUME 112, NUMBER 1 29

P no Ke) 7 tip of snout (Fig. 8); upper lip mostly uni-
J e <4 < = + form light tan; supratympanic fold dark
e : Lorn A ye brown, expanded into dark triangular blotch
= on wn +t posteriorly followed by series of small dark
spots on anterior half of otherwise light tan
x a cn ° flanks; throat, chest, and belly light tan with
x 2 ae = = = lighter spots lacking melanophores; sole of
s 0 foot brown; posterior surfaces of thighs uni-
c SS fog) +
= oo st ve on form tan.
Variation.—Available data do not indi-
. nn on ~ cate any marked variation in measurements
_- SS 2 2. among sites, with a suggestion of slight
= <u ie * on sexual dimorphism in size (Table 1). Mea-
= A os t ql surement variation of the entire sample of
adults (male n = 23, female n = 5) is
x eae = in (means in parentheses): SVL 27.3-—31.2
bile sores ? ee (29.6) mm for males, 30.2—34.4 (32.0) mm
ae Gi . . = for females; % head length/SVL 34-40
2 SS eta (37.3) for males, 32-39 (35.0) for females;
ra $ 5 at A % head width/SVL 36—40 (37.9) for males,
5 . ; cn os A As 34—39 (37.0) for females; % tympanum di-
on ae ones - ie cn ameter/SVL 7-9 (7.7) for males, 6—8 (7.2)
3 = Ao Q Q for females; % eye-nostril distance/SVL 9—
s 11 (10.0) for males, 9-12 (9.8) for females;
& e pete sie Aaet par Se % internarial distance/SVL 8-10 (9.1) for
ae ~ males, 8-10 (8.8) for females; % thigh
8 = hed are length/SVL 44—48 (45.7) for males, 42—44
® (43.2) for females; % shank length/SVL
= “ =7 © © 44-51 (48.2) for males, 44—47 (45.8) for
2 ee ea eee females; % foot length/SVL 47-54 (51.0)
= = : kas ey — for males, 45—49 (47.0) for females. The
iS = So Q o sample size for females is small, but it does
S appear as though the female legs are pro-
S x Q) 00 ° 7 portionately shorter than in males.
S 2 2 ==> SS % The most striking variation in addition to
ad ae J ite i ne the male secondary sexual characteristics
= 5 SO S a involving vocal slits, vocal sacs, thumb as-
5 perities, and forearm hypertrophy is the
ae x AQ + © sexual variation in dorsal texture. In fe-
: 4 ayes = C . males, the dorsum is either smooth or with
eae ) a weak shagreen, whereas the males have
= e co N m oa) :
2 = iS ol é many, large, white tubercles scattered pro-
: fusely over the back (Fig. 9). In the speci-
S aA 2a ge mens from Morro do Chapeu and Mucujé,
=. i x s i i i the male dorsal tubercles are largely limited
| Si Son o & to the upper eyelids and the back. The
2 ‘3 g 4 S a % % < males from Andarai have a much more ex-
= 3 $ 5 = 5 5 = 5 tensive distribution of tubercles, including
< = = the snout, outer arms, dorsal thigh surfaces,

30 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Hy WMH! ’

=
=

a

Bie:

a> 46 '
fst #, =% t¢ 2," a!
7 Ft F Eo,
wire Se

Skin texture of male and female Rupirana cardosoi. Portions illustrated are at the transition from

the dorsum to the flank above the arm insertion on the left side of the animal. Left illustration drawn from male
specimen, MZUSP 76032, right illustration drawn from female specimen, MZUSP 76031.

dorsal shank surfaces, outer tarsus, and var-
iably, the sole of the foot.

Other aspects of variation noted among
the specimens (males and females, except
as noted) include: tongue shape also ovoid
or round; vomerine tooth placement be-
tween to just posterior to choanae, separat-
ed from each other by <% distance of
length of a single tooth patch; finger lengths
also II just <IV just <I<III; some males
with no external indication of a vocal sac;
finger subarticular tubercles sometimes
pungent; outer tarsus sometimes shagree-
ned; tarsal fold occasionally extending %
distance of tarsus; the inner metatarsal tu-
bercle sometimes elevated: the sole of the
foot sometimes with series of small, fleshy
tubercles in line with digits; the dark supra-
tympanic stripe sometimes extending con-
tinuously to mid-flank; front of tympanum
sometimes bordered by dark brown spur ex-
tending from dark supratympanic stripe;
dorsum relatively uniform to longitudinally

striped; chin through the chest sometimes
scattered uniformly with melanophores
with anterior belly with mottled pattern and
posterior belly lacking melanophores to the
entire region from chin through belly light-
ly brown mottled.

MZUSP 76037, a 29.4 mm SVL speci-
men, is a juvenile male, although not the
smallest male in the sample. The specimen
does not have vocal slits, but it has a small
thumb patch asperity (on the thumb only,
not on the thenar tubercle) and no dorsal
tubercles. MZUSP 76039, a 30.2 mm SVL
male, has vocal slits that are in early de-
velopment, but has fully developed nuptial
pads and dorsal tubercles.

The cleared-and-stained female (MZUSP
68959) contained mature-appearing ova, of
varying sizes up to about 1.5 mm diameter
with melanophores on the animal pole.

Etymology.—Named in honor of Dr.
Ad4o José Cardoso, a colleague whose trag-
ic death is deeply felt both at a personal

VOLUME 112, NUMBER 1

level and as a premature halt to his signif-
icant contributions to our understanding of
the Neotropical amphibian fauna.

Distribution and habitat.—At present,
Rupirana cardosoi is known from three lo-
calities in the northern portion of the Es-
pinhaco Range in the State of Bahia, Brazil
(Fig. 10).

Dr. Rodrigues kindly provided the fol-
lowing habitat information. _MZUSP
65202-65205 were obtained during the day
on the bank of a small stream with white
sand and rocks. The stream crosses the city
of Mucujé. The water of the stream was
very clear but with a red-brown tint (as usu-
al in the Espinhacgo Range). The vegetation
was typical of quartzitic campos rupestres:
dominated by Velloziaceae, Euriocaulla-
ceae, Xiridaceae and other endemics. There
were many large and highly eroded quartz-
itic outcrops. MZUSP 68959-68960 were
collected during the day at a similar stream
habitat (2-3 m wide) less than 2 km from
Mucujé. The general habitat was the same
as for the Mucujé stream. Both of these
sites were also visited at night, but no Ru-
pirana cardosoi were found. MZUSP
68961-68962 were collected in a relictual
patch of white sands on a large red rocky
mountain not far from Morro de Chapeu.
The frogs were on the margins of two
small, drying ponds. The vegetation, dom-
inated by Mellastomataceae on the sand,
was typical for campos rupestres. Several
specimens of the sand-adapted Tropidurus
cocorobensis were also collected at the
same time, around 1500 h.

Relationships

Rupirana shares the most character states
with the genus Thoropa, as documented in
the section, “Comparison with leptodactyl-
id genera.”’ The evaluation of character
states in that section did not differentiate
between shared primitive and shared de-
rived states, however. If most of the states
shared by Rupirana and Thoropa are shared

31

primitive states, then there would be little
support for a close sister-group relationship.

In order to undertake a first approxima-
tion of the relationships of Rupirana within
leptodactylid frogs, the data assembled for
an earlier study (Heyer 1975) are used as a
basis for a cladistic analysis (Appendix 1).
As indicated above, the relationships of
Thoropa within the Leptodactylidae have
been disputed by Lynch (1971, 1978) and
me (Heyer 1975). In order to evaluate the
relationships of Rupirana relative to Tho-
ropa, the genus Batrachyla is included to
evaluate Lynch’s position that Batrachyla
and Thoropa are sister-taxa (Lynch 1978),
and the genera Cycloramphus and Megae-
losia are included to evaluate my position
that Thoropa is a member of a grypiscine
clade (Heyer 1975). Three additional rep-
resentatives of telmatobiine genera are in-
cluded to provide structure to test the alter-
native hypotheses: Eleutherodactylus, Eup-
sophus, and Hylorina. Three genera are
used as outgroups: Ceratophrys as a rep-
resentative of the South American cerato-
phryine clade and a representative each of
two subfamilies of the Australian Family
Myobatrachidae, Adelotus and Crinia. In all
cases except for Eleutherodactylus, the
character states analyzed are for genera. As
Eleutherodactylus is currently in the pro-
cess of being broken into smaller mono-
phyletic units, with the final configuration
of this effort far from clear, data for a single
species, E. coqui, are used for purposes of
this analysis.

The data were analyzed with as much or-
dering of character states as unambiguous
morphoseries would allow (Appendix 1).

Phyletic signal in the data set.—The
strength of the phyletic signal in the data
set is evaluated with two tests: the indirect
g, Statistic (Hillis 1991) and the more direct
Permutation Tail-Probabilities (PTP) test
(Faith and Cranston 1991).

Hillis (1991) explained the rationale for
the g, statistic and provided probability lev-
els for cases using 6, 7, or 8 taxa. As there
is no graphable correlation between the

32 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 10. The Espinhago Range in the States of Bahia and Minas Gerais. Stippling indicates areas above 1000
m. Dot is the type locality of Rupirana cardosoi; triangles indicate the other two known localities for R. cardosoi.
Map redrawn from Map 56, page 398 in Giuletti et al. 1997.

VOLUME 112, NUMBER 1

number of taxa and g, critical values, ex-
haustive PAUP analyses were run using 8
taxa with Adelotus or Crinia or Cerato-
phrys used as sole outgroup taxa and delet-
ing Hylorina. The g, statistics for the 8 taxa
are 0.31, 0.29, and 0.01 respectively when
Rupirana is coded as not having T-shaped
phalanges and 0.17, 0.39, and 0.08 respec-
tively when Rupirana is coded as having T-
shaped phalanges. The critical g, values for
8 taxa for P of 0.05 is —0.34 and P of 0.01
is —0.47. None of the g, statistic values ob-
tained in the six analyses approaches those
critical values. The distribution of trees is
not skewed.

The PTP test was run with Rupirana cod-
ed as not having T-shaped phalanges. The
P value is 0.06, which is not significant at
the traditional 0.05 level, but indicative that
there is likely some phylogenetic content in
the data.

Clearly, the phyletic signal in this data
set is not strong and the results must be
interpreted extremely conservatively.

Cladistic relationships.—The results of
PAUP analyses run on the entire data set
with a branch-and-bound search or with the
outgroup taxa run individually with ex-
haustive searches (using the data matrix in
Appendix 1) all have the same outcome:
multiple numbers of shortest trees, for
which the strict consensus tree is an entirely
unresolved polytomy. The branch-and-
bound search of the entire data set yields
19 shortest trees of tree length 112, with a
consistency index (CI) excluding uninfor-
mative characters of 0.52. The exhaustive
search of the data set using Adelotus as the
sole outgroup taxon gives 240 shortest trees
of tree length 86, with a CI value excluding
uninformative characters of 0.56. Similar
results for Crinia as the outgroup are 45
trees of length of 91, CI = 0.54 and for
Ceratophrys as the outgroup are 45 trees of
length 89, CI = 0.56.

In case the coding of the shape of the
terminal phalanges in Rupirana might be
pivotal, exhaustive searches were also run
using Adelotus, Crinia, and Ceratophrys in-

33

dividually as outgroup taxa with Rupirana
coded as having T-shaped terminal phalan-
ges. Again the analyses resulted in multiple
shortest trees with the strict consensus tree
of each being an entirely unresolved poly-
tomy. The results for using Adelotus as the
outgroup are 240 trees of shortest length 85,
CI (excluding uninformative characters) =
0.57; for Crinia, 150 trees of shortest length
91, CI = 0.55; for Ceratophrys, 45 trees of
shortest length 88, CI = 0.56.

These results certainly do not resolve any
questions about the relationships of Tho-
ropa within the Family Leptodactylidae.
However, there is one result that is so con-
sistent that a statement about relationships
can be made: Rupirana and Thoropa do not
share a close relationship. In fact, Rupirana
and Thoropa never form a sister-group re-
lationship in any trees, and Thoropa either
demonstrates a basal relationship with all or
most other taxa or (usually) forms a sister-
group relationship with Batrachyla (e.g., 17
out of 19 trees in the trees based on the
entire data set). This Batrachyla-Thoropa
sister-group relationship, repeated in most,
but not all trees, lends support for Lynch’s
proposal of relationships. However, the po-
tential problem of long-branch attraction
(Swofford et al. 1996:427) would have to
be ruled out to support the Batrachyla-Tho-
ropa relationship.

The results of the above analyses indicate
that the similarities analyzed between Ru-
pirana and Thoropa are due to shared prim-
itive state conditions, not shared derived
states, and that there is no close sister-group
relationship between the two genera.

Zoogeography

The campos rupestres contain a notably
endemic biota at the species level (for a bo-
tanical introduction, see Giuletti & Pirani
1988), including amphibians and at least
some reptiles (e.g., Vanzolini 1982, Ro-
drigues 1988). As I have extremely limited
experience with the campos rupestres, the
following comments are very superficial.

34 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Hopefully these comments will encourage
an honest zoogeographical study of the am-
phibians of the campos rupestres, because I
think the amphibians show a pattern that
differs from those based on study of other
groups.

The campos rupestres occur above 800—
900 m and, as the common name indicates,
are rocky places with poor soils (see Giu-
letti & Pirani 1988 and Giuletti et al. 1997
for better characterizations and maps).
Plants have been the most throroughly stud-
ied campos rupestres group (Giuletti & Pir-
ani 1988), otherwise, there has been little
research on the distributions of other groups
of organisms found in the campos rupestres
with the exception of certain lizards (for
Tropidurus see Rodrigues 1987, 1988, for
other lizards see Vanzolini 1982). Most of
the plant and lizard species found in the
campos rupestres are either endemic to
them or show their major central distribu-
tions on them. Species such as Polychrus
acutirostris and Tropidurus hispidus, which
occur in both the campos rupestres and
more broadly beyond them, seem to be ex-
ceptional cases. Even though the endemic-
ity at the species level is rather astounding,
there are few endemic genera of plants
(Giuletti & Pirani 1988). The affinities for
the campos rupestres endemic species are
primarily with cerrado and caatinga species
(for plants and Tropidurus), and cerrados
and caatingas likely served as the primary
source of species which differentiated in the
campos rupestres environment. Whereas the
campos rupestres could be considered a part
of the diagonal of open formations (caatin-
gas-cerrados-Gran Chaco), first noticed by
K. P. Schmidt (e.g., Schmidt & Inger 1951),
the campos rupestres do not fit comfortably
within the diagonal from a zoogeographic
perspective. Another common affinity for
both plants and Jropidurus is with the res-
tingas (sand beaches) scattered along the
east coast of Brazil. In this case, the restin-
ga associated elements are thought to be de-
rived from campos rupestres ancestors
(Giuletti & Pirani 1988, Rodrigues 1988).

The plants also show affinities that the cam-
pos rupestres lizards do not, i.e., with the
mountain biotas of northern South America
(stronger) and the Andes (weaker) (Giuletti
& Pirani 1988, Harley 1988). There is a mi-
nor affinity with Atlantic Forest plants and
lizards; the plant genus Pleurostima (Velo-
Ziaceae) occurs on the tops of some granitic
outcrops in the Atlantic Forests, and the At-
lantic Forest lizard genera Enyalius, Pla-
cosoma, and perhaps Heterodactylus have
representatives in the campos rupestres.
Most of our knowledge of campos ru-
pestres amphibians comes from the work of
Werner C. A. Bokermann and Ivan Sazima,
who published several papers describing
new species of amphibians from the Serra
do Cip6, southern Espinhacgo Range, in the
State of Minas Gerais. Many of the new
species described from Serra do Cip6 be-
long to well delineated species groups or
genera for which the general distributions
are known. For most of these, the Serra do
Cip6 endemics are related to Atlantic Forest
groups from which they must have been de-
rived. For example all species of the genera
Hylodes, Phasmahyla, and Thoropa and the
Scinax catherinae species group occur only
in the Atlantic Forests except for disjunct
endemic species at Serra do Cip6 (H. ota-
vioi, P. jandaia, T. megatympanum, and S.
machadoi). Determining the faunistic origin
of several other species from the Serra do
Cip6 is not as clear, but it would seem that
the majority of them were derived from At-
lantic Forest lineages and a smaller number
from cerrado groups. Some of the non-en-
demic species that occur at Serra do Cip6
are likely cerrado species, such as Lepto-
dactylus furnarius, L. jolyi, and Physalae-
mus cuvieri. Some of the Serra do Cip6 en-
demics may also have been derived from
cerrado ancestors, for example Leptodacty-
lus camaquara, L. cunicularius, Physalae-
mus evangelistai and Proceratophrys cu-
ruru. The frog fauna of the campos rupes-
tres thus has a strong faunal element de-
rived from Atlantic Forest ancestors and

VOLUME 112, NUMBER 1

this pattern is different from that found in
plants or lizards.

Remnants of the Atlantic Forest vegeta-
tion occur today in the Espinhago Range
(e.g., Pico das Almas, Harley 1995:20-—24).
The Atlantic Forests certainly were more
extensive in the Espinhaco Range and con-
tinuous with the coastal forests during more
mesic times. The campos rupestres, a very
open formation habitat, would be expected
to show biotic affinities with adjacent open
formation habitats, as is true for plants and
lizards. The campos rupestres would not be
expected to show a primary affinity with
the Atlantic Forest biota, as the basic ad-
aptations for those two environments are
profoundly different.

One aspect of anuran ecogeography
could explain why many campos rupestres
frogs were derived from the Atlantic Forest
frog fauna rather than entirely from the cer-
rado/caatinga frog fauna. There are small
streams (mostly seasonal at this stage of the
interglacial) in the campos rupestres land-
scape. There are, as far as I know, no strict-
ly stream-adapted frogs in the cerrados or
caatingas, whereas many of the Atlantic
Forest frogs are stream-adapted. Thus, the
most probable source of stream frogs in the
campos rupestres would come from the At-
lantic Forests rather than the cerrados or
caatingas. If, during periods of drier cli-
mates, the Atlantic Forests underwent grad-
ual change, first contracting to gallery for-
ests and then degrading to even more open
vegetation, the stream frogs may have been
able to adapt to the more open habitat con-
ditions and occupy the campos rupestres.

In summary, from the available data, it
would seem that the unique campos rupes-
tres frog fauna has been derived from only
two sources: Atlantic Forest (strongest);
and cerrado/caatinga (the conclusion of
Heyer 1988, that present data are inade-
quate to determine whether the caatingas
harbor a frog fauna distinct from that of the
cerrados is unfortunately still true).

The discovery of Rupirana cardosoi has
one expected and one unusual zoogeo-

35

graphical consequence. The Espinhaco
Range is comprised of a series of individual
isolated mountains (Fig. 10), with many
species narrowly endemic to single moun-
tain systems within the entire Range (Giu-
letti & Pirani 1988). Finding a new species
in the Chapada Diamantina, which is still
poorly sampled for its frog fauna, is to be
expected. What is unusual is that the new
species also represents a new genus.

The fact that all but one of the endemic
campos rupestres frogs are endemic only at
the species level suggests that the differ-
entiation of the campos rupestres biota oc-
curred in a single episode and relatively re-
cently in terms of geological time. Discov-
ery of an endemic campos rupestres genus
of frogs requires that the single episode of
faunal differentiation requires emendment
or explanation. There seem to be three most
likely alternatives.

Rupirana is an endemic campos rupestres
genus of frog. If this is true, that suggests
that it had an earlier episode of differenti-
ation than other campos rupestres endemic
frogs. Although this alternative requires an
additional episode of differentiation for
frogs, this is not unusual for the rest of the
campos rupestres biota, as there are endem-
ic genera of campos rupestres plants (e.g.,
Barbacenia (Velloziaceae), Morithamnus
(Compositae), Pseudotrimezia (Iridaceae),
and Raylea (Sterculiaceae), Giuletti and
Pirani 1988) and fishes (Copionodon and
Glaphryopoma (Trichomycteridae), Pinna
1992).

Rupirana represents a sampling problem.
It could be that Rupirana had or has a dis-
tribution with other species occurring in the
Atlantic Forests (I think this more likely
than in the cerrados/caatingas). If this is
true, then either the Atlantic Forest species
of the genus have become extinct and/or
they are still extant but have not been
found. The frog fauna of the northern At-
lantic Forests is not well known and I
would not be surprised if additional species
of Rupirana were found there.

Recognition of Rupirana as a distinct ge-

36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

nus is a taxonomic error. If this is the case,
the new species does not require an histor-
ical zoogeographical explanation different
from that for other campos rupestres en-
demic frogs.

Acknowledgments

Drs. Miguel T. Rodrigues (MZUSP), P. E.
Vanzolini (MZUSP), and George R. Zug
(USNM) critically reviewed the manuscript.
Drs. Rodrigues and Vanzolini steered me to
the pertinent literature and made substan-
tive contributions to my zoogeographic un-
derstanding of the biota of the Espinhacgo
Range. Steve Gotte (Biological Resources
Division, U.S. Geological Service stationed
at the Smithsonian Institution) made the
cleared-and-stained preparation of Rupi-
rana cardosoi.

Research for this paper was facilitated by
the Museu de Zoologia da Universidade de
Sao Paulo, especially from Dr. P. E. Van-
zolini. Additional support was provided by
the Neotropical Lowland Research Pro-
gram, Smithsonian Institution, Dr. Richard
P. Vari, Principal Investigator.

Figures 7, 8, and 9 were rendered by Re-
bekah St. John.

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graphic distribution of some plant species from
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, & R. M. Harley. 1997. Espinhaco
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bridge, UK, 562 + xiv pp.

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iope (Labiatae), and its relatives, in Brazil. Pp.
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dens, Kew, 853 pp.

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tergeneric relationships of the frog family Lep-
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. 1988. On frog distribution patterns east of the

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, A. S. Rand, C. A. G. Cruz, O. L. Peixoto, &
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38 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Appendix 1.— Character state descriptions and data matrix for phylogenetic analysis of selected genera of
leptodactylid frogs.

Data from Heyer (1975) supplemented by taxa examined for this study as indicated in body of text.
Multistate characters ordered by criterion of morphoseries when reasonable. Character states not polarized.
Character 1. Pupil shape. State O—round; State 1—horizontal.

Character 2. Tympanum visibility. State 0O—external, visible; State 1—partially concealed; State 2—concealed.
Character state order 0-1-2.

Character 3. Male thumb. State 0—no asperities; State 1—-pad; State 2—-spines. Character states not ordered.

Character 4. Body glands. State 0—none; State 1—paratoids; State 2—inguinals. Character states not ordered.

Character 5. Toe disks. State 0O—none; State 1—disks but no grooves; State 2—disks with circumferential
grooves; State 3—disks with dorsal scutes. Character states not ordered.

Character 6. Tarsal decoration. State 0—none; State 1—fold; State 2—flap. Character state order 0-1-2.

Character 7. Toe webbing. State 0—web; State 1—fringe; State 2—ridge; State 3—-none. Character state order
Q-1-2-3.

Character 8. Life history. State 0—tadpole with 2/3 denticle formula; State 1—tadpole with >2/3 denticle
formula; State 2—tadpole with <2/3 denticle formula; State 3—direct development. Character states ordered in
a triangular relationship with the following connections: 2-0-1; 2-3; 0-3; 1-3.

Character 9. Depressor mandibulae muscle. State O—three slips (DFSQAT to dfsqat); State 1—two slips
(DFSQ to dfsq); State 2—one slip (SQ). Character state order 0-1-2.

Character 10. Sternohyoideus muscle insertion. State 0—on lateral edge of hyoid plate; State 1—on lateral
edge and midline of hyoid body; State 2—on midline of hyoid body. Character state order 0-1-2.

Character 11. Omohyoideus muscle insertion. State O—muscle absent; State 1—insertion on hyoid body and
fascia between posteromedial and posterolateral processes; State 2—insertion on hyoid body only; State 3—
insertion on hyoid body adjacent to posteromedial process. Character states unordered.

Character 12. Iliacus externus muscle. State 0—extends <% ilium length; State 1—extends %—% ilium length;
State 2—extends %4—full length of ilium. Character state order 0-1-2.

Character 13. Tensor fasciae latae muscle insertion. State 0—posterior to anterior extent of iliacus externus
muscle; State 1—at same level as anterior extent of iliacus externus muscle.

Character 14. Semitendinosus muscle. State 0—External head smaller than internal head, external head attached
by tendon to internal head; State 1—as state 0, except internal and external heads displaced from each other;
State 2—external head absent. Character states unordered.

Character 15. Adductor longus muscle. State 0—well developed, insertion on or near knee; State 1—poorly
developed, insertion on adductor magnus muscle; State 2—absent. Character state order 0-1-2.

Character 16. Quadratojugal. State 0O—present, contacting maxilla; State 1—absent.

Character 17. Nasal contact with maxilla. State 0—present; State 1—absent. ,

Character 18. Nasal contact with frontoparietal. State 0—absent; State 1—present, not fused; State 2—present,
fused. Character states unordered.

Character 19. Frontoparietal fontanelle exposed. State O—not exposed; State 1—exposed.

Character 20. Squamosal. State 0O—zygomatic ramus about same length as otic ramus; State 1—otic ramus
developed into plate; State 2—as State 1 and zygomatic ramus articulates with maxilla; State 3—otic ramus
much smaller than zygomatic ramus, latter articulates with maxilla; State 4—-otic ramus much larger than
zygomatic ramus. Character states unordered.

Character 21. Vomerine teeth. State O—present; State 1—absent.

Character 22. Median contact of vomers. State 0—absent; State 1—present.

Character 23. Prootic fusion with frontoparietal. State O—absent; State 1—present.

Character 24. Occipital condyles. State 0O—confluent; State 1—approximate; State 2—-widely separated. Char-
acter state order 0-1-2.

Character 25. Anterior process of hyale. State 0—present; State 1—absent.

Character 26. Posterior sternum. State 0—cartilaginous plate, broadens posteriorly; State 1—cartilaginous plate,
parallel or narrow; State 2—as State 1 with mesosternal mineralization; State 3—-sternal style. Character state
order O-1-2-3.

Character 27. Length of last presacral transverse processes relative to sacrum. State 0—about equal; State 1—
last presacral processes < sacrum.

Character 28. Sacral diapophyses. State 0—expanded; State 1—rounded.

Character 29. Terminal phalanges. State O—simple, knobbed, or claw-shaped; State 1—T-shaped.

Character 30. Ilial dorsal crest. State 0—absent; State 1—present.

Character 31. Diploid number of karyotype. State 0—-greater than or equal to 26; State 1—24.

VOLUME 112, NUMBER 1

Character

25

22 23 24

15 16 17 18 19 20 21

13 14

12

ioe]

OO 10°-0" O30, 2

Adelotus
23-020

Crinia

1 O O&1

O

1

3 0

0)

0 0&2 2

I

O O&1&2
O O0&1&2

O02]. 0 3

Ceratophrys
Batrachyla

0

1 0&1 O

]

l

0

l

O&2&3 O

0&2 20 O

2

Cycloramphus

270 0 0 0 0-1

0

Eleutherodactylus

Eupsophus
Hylorina

0 0 0&1

0

E OScl 0201 {Oa oF 2

L700 OF a

1

O 0&3 O&1 O&]1

l

0

O&2

0

l

QO O&1&2

000 0 4

Megaelosia
Rupirana

1&3 O

I

0

|

O&2 O 0&1 O

1 O&1

1&2 O

0)

Thoropa

39

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(1):40—44. 1999.

A new species of cardinalfish (Perciformes: Apogonidae) from the
Bay of Bengal, Indian Ocean

Thomas H. Fraser

W. Dexter Bender and Associates, Inc., 2052 Virginia Avenue, Fort Myers, Florida 33901, U.S.A.

Abstract.—A new species, Apogon oxina, is described from the Bay of Ben-
gal, Indian Ocean. The preserved color pattern suggests a close relationship
with Apogon ventrifasciatus from the West Pacific. The new species lacks the
white spot at the posterior base of the second dorsal fin present in several other
species and has darker and sharper stripes on the head and upper body and
well-developed bars on the sides reaching to or above the lateral line.

New species of the large cardinalfish ge-
nus Apogon continue to be recognized and
described at a surprisingly high rate. Most
of these species are from the Indo-Pacific
region, with the majority in the large sub-
genus Ostorhinchus. There are about 111
valid species in this subgenus, of which
about 21 undescribed species are known to
me. Relationships within this large group
are not well understood. Gon (1996) pro-
posed phyletic relationships within his con-
cept of the Apogon subgenus Jaydia. His
detailed analysis illustrates the complexities
of developing a robust hypothesis for rela-
tionships within species groupings of Apo-
gon prior to testing with an inclusive anal-
ysis all of proposed subgenera, other spe-
cies groupings of Apogon and other genera.
Fraser (1972) suggested that a more com-
plete understanding of supra-specific rela-
tionships, within and without Apogon, was
dependant on well developed systematics of
the species. Such a critical mass of infor-
mation and material is emerging.

Color patterns and variations of those
patterns appear to be the best distinguishing
features of closely related species of Apo-
gon. New species usually are first recog-
nized by unique color patterns. Consistent
color pattern differences between closely
related species may be accompanied by a
few morphometric differences, such as

body depth, caudal peduncle length, spine
lengths, soft fin-ray lengths, jaw length and
eye diameter, or modally different counts
for gill rakers and pectoral fin-rays. Other
differences have been noted by many au-
thors, such as meristic changes in the spine
count of the first dorsal fin and soft rays of
the second dorsal and anal fins; loss of the
supramaxilla, supraneurals at the anterior
end of the dorsal fin, one pair of epipleural
ribs and uroneurals in the caudal skeleton;
fusion of some hypurals in the caudal skel-
eton; character of spination of the postem-
poral, preopercle, infraorbitals; coloration
of the stomach, intestine and peritoneum;
presence or absence of known or potential
bioluminescent ability; shape of the caudal
fin; elongation of individual vertical fin
rays; position of the anal opening between
the pelvic and anal fins; general scale size,
elaboration of pore architecture on scales as
a simple pore, complex pores, or pit in
scale; and the degree of ossification of the
preopercle. Apogon and the family, in gen-
eral, are replete with internal and external
character variation not usually seen within
a single percoid family. Nevertheless, some
allopatric species pairs, for example, Apo-
gon abrogramma-Apogon exostigma and
Apogon taeniopterus-Apogon menesemus
proposed by Fraser & Lachner (1985) differ
only in color pattern. No morphometric, in-

VOLUME 112, NUMBER 1

ternal, or meristic differences within those
species pairs are known.

The new species belongs in the subgenus
Ostorhinchus Lacepéde, 1802 as defined by
Fraser (1972) using the name Nectamia Jor-
dan, 1917. Gon (1987) designated a neo-
type for Apogon fleurieu (Lacepéde 1802)
and effectively relegated Nectamia as a
synonym.

Methods

Methods of taking and recording meristic
data and measurements are given in Fraser
& Lachner (1985). All measurements are in
millimeters to the nearest 0.1. The follow-
ing acronyms are used to designate insti-
tutions and collections cited and follow
general usage given in Eschmeyer (1998);
BPBM Bernice P. Bishop Museum, Hono-
lulu; CAS California Academy of Sciences,
San Francisco; MNHN Muséum National
d’Histoire Naturelle, Paris; USNM collec-
tions of the former United States National
Museum, deposited in the National Muse-
um of Natural History, Smithsonian Insti-
tution, Washington, D.C. Field station num-
bers are listed for additional collection in-
formation, for example PCH-69-279.

Apogon oxina, new species
Fig. 1

Material examined.—Holotype: Apogon
oxina CAS 33959, 55.9 mm SL; India, Ma-
dras; in 15—22 m; April-June 1975; K. V.
Rama Rao. Paratypes: India: CAS 98101;
7(43—71); data same as holotype. Sri Lanka:
USNM 213364 55(44—58); Trincomalee; 10
m; PCH 69-277; 29 Sep 1969; Phillip C.
Heemstra. USNM 213345 (43-44); PCH
69-279; 30 Sep 1969; Phillip C. Heemstra.
Other material: Sri Lanka: USNM 213346
4(19-—28); Trincomalee; CCK 69-135; 10—
20 m; Christopher C. Koenig; 4 Apr 1970.

Comparative material.—Paratypes Apo-
gon ventrifasciatus BPBM 34136 7(35—49);
Indonesia, Flores I., Maumere Bay; 8 m; J.
E. Randall; 19 Sep 1988. BPBM 34085
5(27—41); Indonesia, Flores I., Maumere

4]

Bay; 3—4 m.; J. E. Randall; 17 Sep 1988.
Other material. USNM 328265 (45); Papua
New Guinea, D’Entrecasteau Is., Normand-
by I.; 16 Dec 1993. BPBM 30132 (40-53);
Indonesia, Lombok I., Sorongjunkung; J. E.
Randall; 21 Feb 1974. Holotype Apogon
moluccensis MNHN 8707; 60.3 mm SL, 76
mm TL; Amboina; Quoy and Gaimard.

Diagnosis.—An Apogon of the subgenus
Ostorhinchus with 14 pectoral fin-rays, usu-
ally 23—24 total gill rakers on the first arch,
no white spot behind the second dorsal fin,
dark vertical bars present on body reaching
to or past the lateral line, dark stripes pre-
sent on head and upper body, and caudal
peduncle length 20—25% SL.

Description.—For general body shape
see Fig. 1. Range of proportions (as per-
centages of standard length) with the ho-
lotype in parentheses: greatest body depth
36-39 (39); head length 38-41 (41); eye
diameter 12—13 (12.2); snout length 8—9
(9.5); bony interorbital width 7—8 (7.5); up-
per jaw length 18—20 (19); caudal peduncle
depth 15-17 (16); caudal peduncle length
20-25 (23); first dorsal-fin spine length
2.8—3.8 (2.8); second dorsal-fin spine length
7.9-9.7 (8.0); third dorsal-fin spine length
17—20 (20); fourth dorsal-fin spine length
16—21 (21); second dorsal spine 12-16
(15); first anal-fin spine length 1.9-—3.3
(2.3); second anal-fin spine length 11-12
(11); pectoral fin length 23—27 (23); pelvic
fin length 23—26 (26).

Dorsal fin VII—1,9; anal fin II,8, last anal
ray much longer than preceding fin-ray;
pectoral fin-rays 14—14; pelvic fin 1,5; prin-
cipal caudal fin-rays 9 + 8; pored lateral
line scales 24; transverse scale rows above
lateral line 2; transverse scale rows below
lateral line 6; median predorsal scales 3—4
(4); circumpeduncular scale rows 12
(5+2+5); total gill rakers 23-24 (24), usu-
ally 20—22 (21) well developed 1—2+4-5
upper, 15—17+0-—2 lower (2+5-—16+1).

Villiform teeth in a band on premaxilla;
two to three rows on dentary; one row on
palatine and vomer; none on ectopterygoid,
endopterygoid or basihyal.

42 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs of,

Vertebrae 10 + 14. Five free hypurals,
one pair of slender uroneurals, three epur-
als, a free parhypural. Three supraneurals,
two supranumerary spines on first dorsal
pterygiophore. Basisphenoid present. Su-
pramaxilla absent. Posttemporal serrate on
posterior margin. Preopercle serrate on ver-
tical and horizontal margins. Infraorbital
shelf present on third bone. Scales ctenoid.
Simple pored lateral-line scales from post-
temporal to caudal fin, pored scales usually
24.

Life colors.—Unknown.

Preserved color pattern.—In 70% ethyl
alcohol: Adult pattern: five dark stripes on
each side of head with one (unpaired) mid-
nape from snout to origin of first dorsal fin,
two stripes over eye extending onto body
above lateral line, the dorsal stripe along
base of first dorsal fin reaching to or just
beyond origin of second dorsal fin, the ven-
tral one reaching well past origin of second
dorsal fin; two stripes behind eye, neither
extending onto body, the upper one reach-
ing below the posttemporal and the lower
one from snout, continuing behind eye
along midline of head to edge of opercle
above pectoral fin; one stripe extending
from lower lip below eye onto opercle end-
ing near pectoral fin base; four or five nar-
row dark bars from abdomen to first stripe

The holotype of Apogon oxina, 55.9 mm SL, from Madras, India, CAS 33959.

above lateral line on body, first bar just be-
hind and below pectoral fin reaching to lat-
eral line, second bar beginning on abdomen
near mid-pelvic fin length reaching past lat-
eral line merging with first stripe above lat-
eral line, third bar beginning on abdomen
near posterior tip of pelvic fin reaching
above lateral line merging with first stripe
above lateral line, fourth bar beginning
above anterior end of anal fin reaching
above lateral line, and a faint indication of
a fifth bar beginning about mid-anal fin
base reaching above lateral line; first dorsal
fin dusky; melanophore patterns indicate
stripe may be present in second dorsal fin
and in anal fin; faint dark mark at posterior
base of second dorsal; edges of caudal fin
pale, melanophore patterns indicate remains
of mid-stripe on caudal fin; outer edge of
pelvic fin dusky; stomach and intestine
black, peritoneum pale. Juvenile pattern:
mid-body lateral stripe extending from
snout, continuing behind eye along midline
of head and body onto caudal fin; 3—5 ver-
tical bars reaching nearly to stripe just
above lateral line; no melanophore patterns
in the second dorsal or anal fin; other pre-
served patterns similar to adults.

Distribution. —Known from Madras, In-
dia and Sri Lanka.

Etymology.—The Greek word oxina

VOLUME 112, NUMBER 1

meaning rake or harrow, a feminine noun
in apposition, referring to the vertical bars
on the side reaching the stripe.

Remarks.—This new species of Apogon
has a combination of characters suggesting
a close relationships with Apogon ventrifas-
ciatus Allen, Kuiter & Randall, 1994. Apo-
gon oxina differs in lacking the white spot
behind the second dorsal fin, having vertical
bars reaching to or past the lateral line, a
shorter caudal peduncle length (20—25% SL
versus 25-31% SL), stronger and darker
Stripes and bars at all sizes, and an appar-
ently larger size (71 mm SL versus 53 mm
SL). Total gill-raker counts for the first arch
overlap at 23—24 for Apogon oxina and 22—
25 for Apogon ventrifasciatus. Both species
have 20—22 well developed gill-rakers and
have 15-17 well developed lower arch gill
rakers.

Allen et al. (1994) compared their new
species, Apogon ventrifasciatus, with Apo-
gon moluccensis Valenciennes, 1832 men-
tioning some similarities in color pattern
and a difference in the well developed low-
er arch gill raker count of 15-17 for Apo-
gon ventrifasciatus and 20—21 for Apogon
moluccensis. However, the type of Apogon
moluccensis has 16 well developed lower
arch gill rakers with no remaining color pat-
tern other than a white spot at the posterior
base of the second dorsal fin based on my
examination. Valenciennes (1832) describes
the type of Apogon moluccensis as having
stripes on the nape, blackish lips, first dor-
sal fin with blackish tip and a reddish body
lacking any marks or spots on or near the
caudal fin. Two species with basal second
dorsal-fin white spots and differing gill-rak-
er counts exist and more species may be
present based on color patterns. There are
two Bleeker names which need to be con-
sidered in this group besides Apogon ven-
trifasciatus: Apogon chrysosoma Bleeker,
1852 and Apogon monochrous Bleeker,
1856. Gon (1987) identified Apogon mol-
uccensis from the Maldives at Rasdu Atoll.
He described the 38 mmSL specimen as
having 6—7 vertical rows of spots on the

43

body from the lateral line to the abdomen,
Stripes in the soft dorsal and anal fin, a
white spot at the posterior base of the sec-
ond dorsal fin, 21 well developed gillrakers
on the first arch and 24(6+ 18) total gill rak-
ers. The redescription of types and valid
species of this group are under study by J.
E. Randall, G. R. Allen and me. None of
these white-spot nominal or possibly un-
described species have the combination of
characters of Apogon oxina.

Acknowledgments

For the loan of material and the use of
museum facilities I extend thanks to Arnold
Y. Suzumoto (BPBM), David Catania, Wil-
liam N. Eschmeyer and Tomio Iwamoto
(CAS), M. L. Bauchot (MNHN), Susan L.
Jewett, David G. Smith, Jeffery T. Williams
(USNM) aided in curatorial processes. Jef-
fery T. Williams helped with photography
and David G. Smith took radiographs.
Leonard P. Schultz funds were provided by
Victor G. Springer (USNM) for several
study trips to the National Museum. John
E. Randall (BPBM), Gerald R. Allen
(WAM) and anonymous reviewers provided
useful comments on drafts of this manu-
Script.

Literature Cited

Allen, G. R., R. H. Kuiter, & J. E. Randall. 1994. De-
scriptions of five new species of cardinalfishes
(Apogonidae: Apogon) from Maumere Bay,
Flores, Indonesia and surrounding regions.—
Revue francaise d’Aquariologie 21(1994) 1-2:
27-38.

Bleeker, P. 1852. Bijdrage tot de kennis der ichthijol-
ogische fauna van de Moluksche Eilandan.
Visschen van Amboia en Ceram.—Natuurkun-
dig Tijdschrift voor Nederlandsch-Indié 3:229-—
309.

. 1856. Beschrijvingen van nieuwe of weinig
bekende vischsoorten van Manado en Makassar
grotendeels verzameld op een rijs naar den Mo-
luksen Archipel in het gevolg van den Goov-
erneur-Generaal Duymaer van Twist.—Acta So-
cietatis Scientiarum Indo-Neerlandicae 1:1—80.

Eschmeyer, W. N., 1998. Introduction. Pp. 16—22 in
Catalog of Fishes. Volume I. W. N. Eschmeyer,
ed., California Academy of Sciences.

44 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fraser, T. H. 1972. Comparative osteology of the shal-
low water cardinal fishes (Perciformes: Apo-
gonidae) with references to the systematics and
evolution of the family.—Ichthyological Bulle-
tin of the J. L. B. Smith Institute of Ichthyology
34:1-105.

, & E. A. Lachner. 1985. A revision of the car-
dinalfish subgenera Pristiapogon and Zoramia
of the Indo-Pacific region (Teleostei: Apogoni-
dae).—Smithsonian Contributions to Zoology
412:1-47.

Gon, O. 1987. Redescription of Apogon (Ostorhin-
chus) fleurieu (Lacepéde, 1802) with notes on
its synonymy.—Japanese Journal of Ichthyolo-
gy 34(2):138-145.

. 1987. The cardinal fishes (Perciformes: Apo-

gonidae) collected in the Maldive Islands during

the Xarifa expedition (1957/58).— Special Pub-

lication, J. L. B. Smith Institute of Ichthyology

42:1-18.

. 1996. Revision of the cardinalfish subgenus
Jaydia (Perciformes, Apogonidae, Apogon).—
Transactions of the Royal Society of South Af-
rica 51:147-194.

Jordan, D. S. 1917. Notes on Glossamia and related
genera of cardinal fishes.—Copeia 44:46—47.

Lacepéde, B. G. E. 1802.— histoire Naturelle des pois-
sons. Paris 4:1—728.

Valenciennes, A. 1832. Descriptions de plusieurs es-
péces nouvelles de poissons du genre Apo-
gon.—Nouvelles Annales Muséum Histoire Na-
turelle, Paris 1:51—60.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(1):45-51. 1999.

A new species of deep-water skate, Rajella eisenhardti,
(Chondrichthyes: Rajidae) from the Galapagos Islands

Douglas J. Long and John E. McCosker

(DJL) Department of Ichthyology, (JEM) Department of Aquatic Biology, California Academy of
Sciences, Golden Gate Park, San Francisco, California 94118, U.S.A.

Abstract.—A new species of skate, Rajella eisenhardti, is described from
two specimens collected from deep-slope (757—907 m) areas off the Galapagos
Islands. It is similar to another eastern Pacific species, Rajella nigerrima de
Buen 1960, but differs in part by having more precaudal vertebrae and more
pectoral radials, a wider and longer disc, a longer snout, larger gill openings
with wider interspaces, longer anterior and shorter posterior pelvic lobes, and

in coloration and spination.

The skate fauna of the eastern central Pa-
cific, extending from southern Mexico to
northern Peru, contains at least 12 species
(McEachran & Miyake 1984, 1990; Mc-
Eachran 1995). Galapagos records are lim-
ited to those of Bathyraja spinossima, Gur-
gesiella furvescens and Raja velezi (Mc-
Eachran & Compagno 1979; McEachran
1995; Grove & Lavenberg 1997). Mc-
Eachran and Miyake (1984) also reported a
specimen of Bathyraja cf. richardsoni from
near the Galdpagos Islands (00°11.3’S,
97°27.7'W, 1710 m), but not within the ar-
chipelago. The recent capture of two spec-
imens of an undescribed Rajella using the
manned submersible Johnson Sea-Link dur-
ing deep-water dives around the Galapagos
Islands (McCosker et al. 1997) has prompt-
ed this paper.

Methods

Abbreviations: CAS = California Acad-
emy of Sciences, ichthyological collections;
TCWC = Texas Cooperative Wildlife Col-
lection; TL = total length.

Measurements and meristic counts fol-
low current methodology in McEachran &
Miyake (1984, 1988) and Stehmann (1995).
Measurements were made with dial calipers
and recorded to the nearest 0.1 mm. Ver-

tebral and radial counts were made from ra-
diographs.

Rajella eisenhardti, new species
Figs. 1—2; Table 1

Diagnosis.—The new species can be dis-
tinguished from other Rajella species by a
combination of its moderately long tail (tail
length greater than distance from snout to
cloaca) that is largely free of denticles on
the ventral side; three rows of caudal thorns
that begin at level of anterior pelvic lobe;
67—68 predorsal caudal vertebrae; 68—69
pectoral radials; disc width 55.0—58.5% of
TL and disc length 50.0-51.2% of TL;
snout-vent length 44.1—46.4% of TL; pre-
orbital snout length 13.4—14.1% of TL; and
ventrum darker than dorsum, with distinct
light areas around mouth and nasal areas,
gill openings, abdomen, cloaca, and tips of
rostrum, tail, and pectoral fins.

Holotype: CAS 86817, an immature male
(384.5 mm TL) caught with a siphon hose
on the bottom at 757 m off Cabo Rosa
(01°04.74'S, 91°11.08'W), Isla Isabella, Ga-
lapagos Islands, by J. E. McCosker abroad
the submersible Johnson Sea—Link on 11
Nov 1995.

Paratype: CAS 86561, an immature male
(254.1 mm TL) caught with a siphon hose

46 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1.
mm TL); C—D dorsal and ventral views of paratype, CAS 86561 (254.1 mm TL).

on the bottom at 907 m off James Bay
(0°26.46’S, 90°19.04’W), Isla San Salvador
(Santiago Island), Galapagos Islands, by J.
E. McCosker aboard the submersible John-
son Sea—Link on 26 Nov 1995.
Comparative material: Rajella nigerrima,

Preserved types of Rajella eisenhardti. A—B, dorsal and ventral views of holotype, CAS 86817 (384.5

TCWC 3881-01, 273 mm TL, juvenile fe-
male, trawled off Chile (35°53.5’S,
72°44'’W) at 780-925 m, Anton Brunn
cruise 18A, sta. 40, 9-10 Aug 1966; TCWC
3885-01, 239 mm TL, juvenile female,
trawled off Peru (03°15’S, 80°55’W) at

VOLUME 112, NUMBER 1

Fig. 2. Rajella eisenhardti. Underwater photo-
graph of living paratype (CAS 86561) taken from the
submersible Johnson Sea-Link on the bottom at 907 m
off James Bay, Isla San Salvador (Santiago Island) Ga-
lapagos Islands.

945—960 m, Anton Brunn cruise 18A, sta.
120, 10 Sep 1966.

Description.—See Table 1 for meristics
and measurements. Meristic values de-
scribed below are listed for the holotype,
and values for the paratype are shown in
parentheses.

A medium-sized species of Rajella (sen-
su Stehmann 1970, 1978; McEachran &
Dunn 1998), its disc heart-shaped and mod-
erately broad, with broadly-rounded outer
corners. Disc length 0.88 (0.91) in disc
breadth. Anterior margin of disc weakly
concave near tip of rostrum, straight to
slightly convex to widest part of pectorals.
Axis of greatest width 0.83 (0.84) times dis-
tance from tip of snout to axil of pectoral
fins. Snout to mid-cloaca 2.15 (2.26) times
in total length.

Preorbital length 4.0 (3.5) times orbit
length; preoral length 1.87 (1.98) times in-
ternarial distance. Interorbital distance 1.16
(0.96) times orbit length; orbit length 1.48
(1.85) times spiracle length. Anterior nasal
lobe laterally expanded to form a nasal cur-
tain, weakly convex anteriorly and finely-
fringed laterally and posteriorly. Posterior
nasal flap well-developed, extending to near
corner of mouth; a single projection on lat-
eral side of flap with fine fringes on the
lateral posterior edges, with a smooth mar-
gin medially; length of flap 0.32 (0.29), and

47

width 0.59 (0.57), into preoral length; space
between fimbriae 0.30 (0.29) into preoral
length.

Distance between first gill slits 1.95
(2.15) times distance between nares; dis-
tance between fifth gill slits 1.22 (1.33)
times distance between nares; length of first
gill slit 1.20 (1.11) times length of fifth gill
slit. Third gill slit largest, 0.26 (0.27) times
distance between nares, fifth gill slit 0.16
(0.19) times distance between nares, only
slightly smaller than first, 0.19 (0.21) times
distance between nares.

Pelvic fins deeply incised; anterior lobe
long and moderately narrow, bluntly ta-
pered and rounded distally; length of ante-
rior lobe 1.59 (1.46) times length of pos-
terior lobe. Tail long, gradually tapering to
tip; distance from mid-cloaca to caudal tip
1.15 (1.26) times distance from snout to
cloaca. Base of tail moderately narrow,
9.3% (9.2%) of cloaca to caudal tip length,
and moderately convex dorsally, weakly
concave ventrally becomming flattened dis-
tally. Lateral folds originating well behind
the base of tail and continue along length
to just anterior of caudal tip where it ends
as small lobed flap. Dorsal fins low and
rounded; first fin only slightly larger than
second and similarly shaped, no space be-
tween dorsal fins or between second dorsal
and epichordal caudal lobe.

Dorsum entirely covered with fine den-
ticles, including tail and dorsal fins, except
on the lateral and posterior margins of the
pectoral fins; pelvic fins free of denticles
and anterior pelvic lobes smooth, but base
of pelvic with fine denticles in the paratype.
Larger denticles and thorns with oval or ob-
long bases, often with weak longitudinal
ridges and anteriorly curving tips. Larger
denticles on rostral shaft; one to two thorns
anterior to each orbit, three posterior to
each orbit, and one thorn on each side of
the inner orbit. Five thorns on nuchal mid-
line, three on each scapular lateral to the
midline, 6—7 thorns on post-scapular mid-
line anterior to the tail. Three rows of
thorns on the dorsal surface of the tail be-

48 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Measurements (in mm) and percentages of total length (in parentheses) of Rajella eisenhardti
compared to percentages of total length of R. nigerrima (5 males, 6 females; data from McEachran & Miyake,
1984).

Raja eisenhardti R. nigerrima (n = 11)

Holotype Paratype

CAS 86817 CAS 86561 Range Mean
Total length 384.5 254.1 139-374 —
Snout to mid-cloaca 178.5 (46.4) 112.3 (44.1) 40-44 43
Disc width 2235:0,(58.5) 140 (55.0) 51-53 a2
Disc length 197:0°61.2) 127.0 (50.0) 46-49 48
Head length 104.8 (27.3) 63.0 (24.8) NA NA
Snout length (preorbital) 54.3 (14.1) 34.3 (13.4) 10.2-12.8 11.0
Snout length (preoral) 57.5 (14.9) 35.9 (14.1) 12.6—14.5 13.9
Snout to maximum width 132.0,.(34.3) 78.0 (30.6) 31-35 32
Snout to pectoral axis 164.5 (42.8) 104.8 (41.2) NA NA
Prenasal length 47.7 (12.4) 31.4.@12:3) 9.6-11.5 11.0
Orbit diameter 13.) Bp) 9.8 (3.8) 3.74.8 4.3
Interorbital distance 15.7 (4.0) 93 G7) 3.3-3.6 3.5
Spiracle length 9:18 33) 53.8: (2.2) 1.52.3 1.8
Orbit and spiracle length 19.4 (5.0) 12.3 (4.8) 3.45.2 4.6
Distance between spiracles 28:06 2) 17.7 (6.9) 6.16.8 6.3
Mouth width 28.7 (7.4) 18.7 (7.3) 6.2—7.2 6.9
Distance between nostrils 30.8 (8.0) Loot C7) 6.5-—7.1 7.0
Nasal curtail length 18.6 (4.8) 10.3 (4.1) NA NA
Nasal curtail width 34.0 (8.8) 20.5 (8.1) NA NA
Width of Ist gill opening 5.951 5) 3.8 (1.4) 1.0—1.5 eZ
Width of 3rd gill opening ro [a a1 Gy 4.9 (1.9) 1.0-1.5 12
Width of 5th gill opening 4.9 (1.2) BA les) 0.6—1.1 0.8
Distance betw. Ist gill openings 60.3 (15.6) 38.9 Cl des) 13.0—14.5 13.8
Distance betw. 5th gill openings 37.6 (9.7) 24.1 (9.4) 8.1-8.8 8.5
Length of anterior pelvic lobe 63.9 (16.6) 41.3 (16.2) 10.4-14.7 12.9
Length of posterior pelvic lobe 40.2 (10.4) 2825: 11.8—16.0 14.9
Cloaca to Ist dorsal fin 142.8 (37.1) 100.2 (39.4) 41-44 42.0
Cloaca to caudal origin 188.5 (49.0) 134.3 (52.8) 47-57 54.0
Cloaca to caudal tip ZOGALO2S) 141.8 (55.8) 50-60 57.0
Caudal lobe length 11.6 (3.0) 8.9 (3.5) NA NA
lst dorsal height DT 6) 15.0 (5.9) NA NA
lst dorsal base 22 (SS) 16.1 (6.3) NA NA
2nd dorsal height 21.1 (3:5) 12.6 (5.0) NA NA
2nd dorsal base 22.8 (5.9) 16.5 (6.5) NA NA
Tail width at base 19.2% .0) 1322) NA NA
Tail width at Ist dorsal origin 6.9 (1.8) 5:0°(2.0) NA NA
Tail thickness at base 1316165) 782@G)) NA NA
Tail width at 1st dorsal origin Ae) (12) 32 (3) NA NA
No. of upper/lower tooth rows 48/32 46/42 36—46/36—46 42.4
No. of trunk vertebrae 29 28 27-30 28.6
No. of predorsal caudal vert. 67 68 59-65 62.6
No. of pectoral radials 69 68 61-65 63.0
No. of pelvic radials 19 ¥7) NA NA

ginning at the mid-level of the pelvic fins type entirely smooth and bare of denticles;
and continuing to first dorsal fin; median ventral surface of paratype smooth except
caudal thorns 24 (23); lateral caudal thorns for some small denticles on ventrolateral
numbering 38—39 (19-20), thorns diminish- margins of tail.

ing along the tail. Ventral surface of holo- Upper and lower jaws slightly arched;

VOLUME 112, NUMBER 1

teeth in quincunx arrangement with small
short median crowns becoming more blunt
and flattened distally; upper tooth rows 48
(46), lower rows 32 (42). Trunk vertebrae
29 (28), predorsal caudal vertebrae 67 (68).
Pectoral radials 69 (68), pelvic radials 19
Cry).

Color after preservation: Holotype pur-
plish gray and paratype light brownish gray
dorsally; both with whitish blotch on the
dorsal edge of the pectoral fin tip; ventral
surface darker except for distinct whitish
areas on the anterior tip of the rostrum, an-
terior and posterior to the mouth, around
the nostrils, around each gill opening,
around the cloaca, on the abdomen, on the
tips of the anterior pelvic fin lobes, and at
the tip of the tail; light area between ab-
domen and gill openings connected by a V-
shaped mark. Thorns and large denticles
whitish, smaller denticles same color as sur-
rounding skin. Color in life (based on an
underwater photo of the paratype): dorsal
interior of disc, pelvic base, and tail pale
brown to grayish; rostrum, dorsal fins, and
margins of pectoral and pelvic fins blue-
gray.

Etymology.—Named in honor of E. Roy
Eisenhardt, director emeritus of the Cali-
fornia Academy of Sciences, who has gen-
erously assisted us and our colleagues.

Discussion

The subgenus Rajella was recently ele-
vated to full generic status (McEachran &
Dunn 1998), and includes 14 species found
in the Atlantic, Pacific, and Indian oceans.
These are: R. annandalei Weber, 1913; R.
barnardi (Norman 1935); R. bathyphilia
(Holt & Byrne 1908); R. bigelowi (Steh-
mann 1978); R. caudaspinosa (Von Bonde
& Swart 1923); R. dissimilis (Hulley 1970);
R. fuliginea (Bigelow & Schroeder 1954);
R. fyllae (Liitken 1887); R. kukujevi (Dol-
ganov 1985); R. leopardus (Von Bonde &
Swart 1923); R. nigerrima (de Buen 1960);
R. purpuriventralis (Bigelow & Schroeder
1962); R. ravidula (Hulley 1970); and R.

49

sadowskii (Krefft & Stehmann 1974). Our
specimens differ from all of these species
by a combination of morphometrics, mer-
istics, spination, and coloration. It most
closely resembles the only other eastern Pa-
cific Rajella, R. nigerrima, a species that
ranges from Peru to Chile (McEachran &
Miyake 1984).

Rajella nigerrima has a checkered taxo-
nomic history. It was first placed in the ge-
nus Breviraja (de Buen 1960; Stehmann &
Seret 1983), later placed in the subgenus
Malacoraja (McEachran & Compagno
1982) which was subsequently elevated to
generic rank (McEachran 1984; McEachran
& Miyake 1984; Ishihara & Ishiyama
1986), later reclassified in Raja, subgenus
Rajella (McEachran & Miyake 1990; Pe-
queno & Lamilla 1993), and has recently
been elevated to generic status (McEachran
& Dunn 1998). The types of R. nigerrima
are believed lost and a neotype was desig-
nated by McEachran & Miyake (1984).

Our comparisons with the original de-
scription of R. nigerrima, specimens de-
scribed by McEachran & Miyake (1984),
and preserved specimens we examined,
show significant meristic and morphologi-
cal differences (Table 1). Raja nigerrima
has fewer predorsal caudal vertebrae and
fewer pectoral radials. There are several im-
portant proportional differences between R.
eisenhardti and R. nigerrima including the
latter’s smaller preorbital length in relation
to orbit length, and its first gill slit is longer
than its fifth gill slit. And (in relation to
total length) R. nigerrima has a shorter
snout, a shorter snout to mid cloaca length,
a reduced disc length and width, a narrower
mouth, narrower interorbital and interspi-
racular distances, shorter anterior pelvic
lobes, larger posterior pelvic lobes, and nar-
rower third and fifth gill openings.

Rajella nigerrima has three rows of dor-
sal thorns posterior to the suprascapular, but
R. eisenhardti has three rows beginning at
the middle edge of the pelvic fin; R. niger-
rima has three dorsal and lateral rows of
thorns from the base of the tail to the first

50 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

dorsal fin, but in R. eisenhardti the caudal
thorns begin at the level of the anterior pel-
vic lobes. The underside of the tail in R.
nigerrima has many small, sharp denticles
except for a very narrow strip along the
midline, which is bare; in R. eisenhardti,
the underside is completely bare (holotype)
or has spinelets only on the lateral margins
of the tail’s underside (paratype).

The color of R. nigerrima is brown, with
some indistinct lighter areas on the under-
side around the mouth, vent, abdomen, and
around the gills; in R. eisenhardti, the body
color is gray to brownish gray with very
distinct whitish areas around the nares,
mouth, gills, vent, and abdomen, and on the
tips of the rostrum, tail, and pectoral fins
and pelvic lobes.

Acknowledgements

For assistance and permission to study in
Ecuador, we sincerely thank: Ing. O. Aguir-
re, Subdirector de Pesca de Galapagos; C.
V. Zambrano, Bidl. H. Miller, Dr. E Or-
maza-Gonzalez, Instutito Nacional de Pes-
ca; Lcdo. A. Izurieta Valery and Biol. E.
Amador, Parque Nacional Galapagos; Ten-
iente A. Villacis, Capitan de Puerto de Isi-
dro Ayora; and Dr. C. Blanton, Director de
Estacion Cientifica Charles Darwin. Many
other individuals have assisted us in this
project, and in particular we wish to thank:
R. G. Gilmore and the staff of the Harbor
Branch Oceanographic Institution for the
operation of the Vessel Seward Johnson and
its submersible, the Johnson Sea—Link; J. D.
McEachran of Texas A&M University for
loan of comparative material and critical re-
view of an earlier draft of this manuscript;
E. R. Eisenhardt, D. Lin, and B. E. Carlson
for photographic assistance; W. Eschmeyer,
D. Catania, and J. D. Fong of the California
Academy of Sciences; and the David and
Lucile Packard Foundation and the Discov-
ery Channel for grants and other assistance.

Literature Cited

Bigelow, H. B., & S. C. Schroeder. 1954. A new fam-
ily, a new genus, and two new species of batoid

fishes from the Gulf of Mexico.—Breviora 24:

1-16.

Ge . 1962. New and little known batoid
fishes from the western Atlantic.—Bulletin of
the Museum of Comparative Zoology 128:162—
244.

de Buen, FE 1960. Tiburones, rayas y quimeras en la
Estacién de Biologia Marina de Montemar,
Chile.—Revista de Biologia Marina 10(1—3):3-
SW)

Dolganov, V. N. 1985. Raja (Rajella) kukujevi sp. n.
(Elasmobranchii, Rajidae) from the North-At-
lantic Ridge.—Zoologicheskii Zhurnal 64(2):
304-307.

Grove, J. S., & R. J. Lavenberg. 1997. The Fishes of
the Galapagos Islands. Stanford University
Press, Stanford. 863 pp.

Holt, E. W. L., & L. W. Byrne. 1908. Second report
on the fishes of the Irish Atlantic slope.—Fish-
eries Ireland Scientific Investigations 5:141—
201.

Hulley, P A. 1970. An investigation of the Rajidae of
the west and south coasts of southern Africa.—
Annals of the South African Museum 55(4):
151-220, 13 pls.

Ishihara, H., & R. Ishiyama. 1986. Systematics and
distribution of the skates of the north Pacific
(Chondrichthyes, Rajoidei). Pp. 269-280 in T.
Uyeno, R. Arai, T. Taniuchi, and K. Matsuura,
eds., Indo-Pacific Fish Biology: Preceedings of
the Second International Conference on Indo-
Pacific Fishes. Ichthyological Society of Japan,
Tokyo.

Krefft, G., & M. Stehmann. 1974. Ergebnisse der
Forschungsreisen des FFS ‘“Walther Herwig”
nach Sudamerika: XXXIII. Raja (Rajella) sa-
dowskii spec. nov. (Chondrichthyes, Batoidei,
Rajidae), ein weiterer neuer Roche vom siid-
westatlantischen Kontinentalabhang.—Ar-
chiv Fiir FischereiWeissenschaftliche 25:33-—
SW

Liitken, C. E 1887. Korte bidrag til nordisk ichthy-
ographi. VI. En for Grgnlandshavet ny Rokke-
art.—Videnskabelige Meddelelser Naturhistorik
Forening K6benhaven 1887:1—4.

McCosker, J. E., G. Merlen, D. J. Long, R. G. Gilmore,
& C. Villon. 1997. Deepslope fishes collected
during the 1995 eruption of Isla Fernandina,
Galapagos.—Noticias de Galapagos 58:22—26.

McEachran, J. D. 1984. Anatomical investigations of
the New Zealand skates Bathyraja asperula and
B. spinifera, with an evaluation of their classi-
fication within the rajoidei (Chondrichthyes).—
Copeia 1984(1):45—58.

. 1995, Rajidae. Pp. 773-777 in W. Fischer, F

Krupp, W. Schneider, C. Sommer, K. E. Car-

penter and V. H. Niem, eds., Guia FAO Para la

Identificaci6n de Especies para los Fines de la

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Pesca. Pacifico centro-oriental. Vol. II. Verte-
brados—Parte 1:647—1200.
, & L. J. V. Compagno. 1979. A further de-
scription of Gurgesiella furvescens with com-
ments on the interrelationships of Gurgesielli-
dae and Pseudorajidae (Pisces: Rajoidei).—Bul-
letin of Marine Science 29(4):530-—553.
, & . 1982. Interrelationships of and
within Breviraja based on anatomical structures
(Pisces: Rajoidei).—Bulletin of Marine Science
32(2):399—425.
, & K. A. Dunn. 1998. Phylogenetic analysis
of skates, a morphologically conservative clade
of elasmobranchs (Chondrichthyes: Rajidae).—
Copeia 1998 (2):271—290.
, & T. Miyake. 1984. Comments on the skates
of the tropical eastern Pacific: one new species
and three new records (Elasmobranchii: Raji-
formes).—Proceedings of the Biological Soci-
ety of Washington 97(4):773-787.
,& . 1988. A new species of skate from
the Gulf of California (Chondrichthyes, Rajoid-
ei).—Copeia 1988(4):877-886.
ye . 1990. Zoogeography and bathym-
etry of skates (Chondrichthyes, Rajoidei). Pp.
305-326 in H. L. Pratt, S. H. Gruber and T.
Taniuchi, eds., Elasmobranchs As Living Re-
sources. National Oceanographic and Atmo-
spheric Administration Technical Report 90.
Norman, J. R. 1935. Coast fishes. Part I. The south
Atlantic.— “Discovery” Reports 12:1—58.
Pequeno, G. & J. Lamilla. 1993. Batoideos comunes a
las costas de Chile y Argentina-Uruguay (Pi-

51

sces: Chondrichthyes).—Revista Biologia Ma-
rina, Valparaiso 28(2):203-—217.

Stehmann, M. 1970. Vergleichend morphologische und
anatomische Untersuchungen zur Neuordnung
der Systematik der nordostatlantischen Rajidae
(Chondrichthyes, Batoidei).—Archiv fiir Fis-
cherei- und Meeresforschung Wissenschaftliche
21:73-164.

. 1978. Raja “‘bathyphilia”’, eine doppelart des

subgenus Rajella: wiederbeschreibung von R.

bathyphilia Holt & Byrne, 1908 und Raja bi-

gelowi spec. nov. (Pisces, Rajiformes, Raji-
dae).—Archiv fiir Fischerei- und Meeresfor-
schung Wissenschaftliche 29(1/2):23-58.

. 1995. First and new records of skates (Chon-

drichthyes, Rajiformes, Rajidae) from the west

African continental slope (Morocco to South

Africa), with descriptions of two new species.—

Archive of Fishery and Marine Research 43(1):

1-119.

, & B. Seret. 1983. A new species of deep-
water skate, Breviraja africana sp. n. (Pisces,
Batoidea, Rajidae), from the eastern Central At-
lantic slope, and remarks on the taxonomic sta-
tus of Breviraja Bigelow & Schroeder, 1948.—
Bulletin de Muséum National d’Histoire Natu-
relle, Paris, 4th series, section A, 3:903—925.

Von Bonde, C., & D. B. Swart. 1923. The Plagiostoma
(skates and rays) collected by the S. S. “Pick-
le’’.—Union of South Africa Fisheries and Ma-
rine Biological Survey 3(5):1—22.

Weber, M. 1913. Die fische der Siboga-Expedition.—
Siboga Expedition 57:1-—710.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(1):52-58. 1999.

A new genus of the subfamily Cubacubaninae
(Insecta: Zygentoma: Nicoletiidae) from a Mexican cave

Luis Espinasa

CEAMISH, Universidad Aut6énoma del Estado de Morelos Av. Universidad #1001, Col.
Chamilpa, Cuernavaca, Mor. C.P. 62210, México

Abstract.—A new genus and species (Squamigera latebricola) of the sub-
family Cubacubaninae from a Mexican cave is described. The new genus is
distinguished from almost all the genera of this subfamily by having scales. It
differs from Texoreddellia, the only other genus of this subfamily with scales,
by the morphology of scales and other characters, which are discussed.

The family Nicoletiidae is a frequently
encountered group in caves of México and
Central America (Reddell 1981), but with
few exceptions, most species await descrip-
tion. During exploration of several caves in
the states of Guerrero and Morelos, Méxi-
co, several specimens of this family were
collected. A specimen collected from ‘“‘Po-
zas Azules”’’ cave is a previously undescri-
bed genus with scales of the subfamily Cu-
bacubaninae. Available material is of par-
ticular interest because most members of
the subfamily lack scales. Only two genera
have them: Texoreddellia (Wygodzinsky
1973) and the new genus described here.
However, the morphology of the scales
among other characteristics is drastically
different for these two genera; therefore,
they are probably not closely related.

Study Area

The material was collected in ‘“‘Pozas
Azules”’ cave (Taxco de Alarc6én Munici-
pality, Guerrero State, México, 18°36'40’N,
99°33'25”"W). The cave is 1399 m long and
+53 m deep. This cave is the resurgence of
a cave system formed by Isote cave and
Cueva de las Pozas Azules. Total length of
system is almost three kilometers and depth
is 230 m (for a detailed description of both
caves’ topography, see Espinasa-Perefia
1989).

This cave system has a very peculiar fau-
na. Apart from the new genus described
here, a freshwater polychaete has been de-
scribed as a troglobite (Solis-Weiss & Es-
pinasa 1991). Some undescribed copepods
living in association with the polychaete
will probably be characterized as troglobi-
tes. Other organisms collected in the system
are troglophile ricinulids (Cryptocellus bo-
neti), collembola of the families Entomo-
bridae and Sminthuridae, beetles, two spe-
cies of spiders, millipedes, mites, protozoa
and bacteria (Salmonella, Klebsiella, and
Bacillus). To date, physical connection be-
tween Isote cave and Cueva de las Pozas
Azules has not been explored because both
ends of the caves end in sumps. Connection
is assumed in part on geohydrological evi-
dence (Espinasa-Perefia 1989), by the prox-
imity of the two final sumps found in each
cave (less than 20 m apart), and most im-
portantly, by the shared fauna mentioned
above.

Cave formation.—The limestone rock
was formed during the lower Cretaceous
period (Albian, Cenomanian, and Turoni-
an). Elevation occurred at the end of Cre-
taceous (Maastrichtian), when erosion be-
gan. Presence of an igneous rock (Riolite
tilzapotla) overlaying the limestone from
which the water that formed the cave
comes, seems to indicate that the cave was
not formed until late Oligocene (L6pez-Ra-

VOLUME 112, NUMBER 1

mos 1974). It is also known that the cave
was already formed in the Quaternary be-
cause there is evidence in cave sediments
of climatic changes related to this period
(Espinasa-Perefia, pers. comm.). Thus, if
the cave was formed at some point between
Oligocene and Quaternary, there is reason
to assume that the troglobites in this cave
system separated from the outside commu-
nities at more or less the same time.

Materials and Methods

The specimen was found crawling on the
cave wall. It was placed into a vial with
70% alcohol. Dissections were made with
the aid of a stereo microscope and the dif-
ferent parts of the body were mounted in
fixed preparations with Hoyer’s solution.

All illustrations were made with the aid
of a camera lucida attached to a compound
microscope.

The type is deposited in the following
collection: LESM-DB-MEX (Laboratory of
Ecology and Systematic of Microarthro-
pods. Department of Biology, Faculty of
Sciences, UNAM. México D. EF) Catalog
number: ZYG-2.

Squamigera, new genus

Diagnosis.—A member of the subfamily
Cubacubaninae with scales. Cerci of male
with modified spines. Parameres with a
cleft on apex.

Description.—Body robust, long, ap-
proximately parallel-sided, thorax slightly
but distinctly wider than abdomen. Scales
present, numerous, and multiradiate. Form
mucronate to emarginate, with borders
slightly serrated. Head, thorax, and abdo-
men with scales and setae. Legs with scales
on proximal articles. Mouthparts and ab-
dominal stylets only with setae.

Pedicellus of male with unicellular
glands and a small spur on its base. Mouth-
parts not specialized. Mandibles strongly
sclerotized apically and with usual teeth.
Galea apically with several sensory pegs.
Lacinia heavily sclerotized distally; first

53

process of lacinia pectinate. Labium with-
out prominent lateral lobes.

Tarsi with 4 articles. Praetarsi with 3 sim-
ple claws, median claws glabrous, slender,
and smaller than lateral claws.

Abdominal sterna II (apparently)-III- VII
subdivided into coxites and sternite. Sterna
VIII and IX of male entire. Sterna II—VII
with 2 + 2 macrochaetae. Coxites on seg-
ments II-IX with stylets. Eversible vesicles
on segments II-VI, pseudovesicles on VII.
Urosternum VIII with a wide and not too
deep posterior emargination. Posterior pro-
jections acute to slightly rounded, pointing
slightly outward. Tergum X protruding, al-
most straight on posterior border, posterior
angles with 2 or 3 subequal macrochaetae.
Urosterna III and IV simples. Cerci of male
with modified spines.

Parameres with a cleft and specialized se-
tae on apex. Opening of penis longitudinal.

Type species.—Squamigera latebricola.

Etymology.—Latin Squamigera = scale
bearing, scaly. Here treated as a noun in the
nominative singular. Gender, feminine.
Makes reference to its body covered with
scales.

Distribution.—The only individual found
is from ‘“‘Cueva de las Pozas Azules’’ cave,
Taxco de Alarcén Municipality, Guerrero
State, México, 18°36’40"N, 99°33’25’"W.

Remarks.—Squamigera belongs to a
group of nicoletiid genera, the Cubacuban-
inae (Mendes 1988), characterized by sub-
divided abdominal sterna IJ—VII and fused
coxites of VIII and IX abdominal segments
of the male. Squamigera is distinguished
from almost all genera of this subfamily by
having scales. It differs from Texoreddellia,
the only other genus with scales, by the
morphology of scales, having scales on
head, morphology of parameres, and mod-
ified spines in male cerci.

The genus shares with Anelpistina (Sil-
vestri 1905 as subgenus), Prosthecina (Sil-
vestri 1933), and Cubacubana (Wygodzin-
sky & Hollinger 1977), the presence of
spines in cerci and a posterior emargination
on urosternum VIII, which are absent in

54 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1 ma
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Fig. 1. Squamigera latebricola new genus. (Male): A, Head; B, Basal article and pedicellus of antenna; C,
Small spur of pedicellus; D, Labial palp and labium; E, Mandible; EK Thoracic tergum (scales and setae partially
shown); G, Posterior border of metanotum (scales partially represent).

VOLUME 112, NUMBER 1 55

ih. A, Bee “al ic \
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Fig. 2. Squamigera latebricola new genus. (Male): A, Hind leg, microchaetae partially shown, scales not
shown; B, Urosternum VII and VIII (scales and setae partially shown); C, Genital area; D, Scales of head.

56 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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Squamigera latebricola new genus. (Male): A, Scales and spines of the posterior border of the

metanotum; B, Scales of urotergum I; C, Urotergum X (scales and setae partially shown); D, Cercus basal

portion.

Texoreddellia. It is possible that Sqguami-
gera has closer affinities with this group of
species than with Texoreddellia.

Squamigera latebricola, new species
Figs. 1A—G, 2A—D, 3A—D

Material examined.—México. Guerrero:
Taxco de Alarc6n, Cueva de las Pozas Azu-

les (+53 meters deep, 1399 meters long). 3
Dec 1988, R. Espinasa-Closas col. Male ho-
lotype.

Description.—Maximum body length
22.0 mm. Maximum length of antennae un-
known (broken), of caudal appendages 28.0
mm. Body and legs robust. Head, thorax,
abdomen, and proximal articles of legs cov-

VOLUME 112, NUMBER 1

ered by scales (Figs. 2D, 3A, B). General
color light yellow to white.

Head with macrochaetae and microchae-
tae as shown in Fig. 1A. Basal article of
antennae in males has a small projection
with 2 macrochaetae. Pedicellus of anten-
nae of male as shown in Fig. 1C, with 5
clusters of unicellular glands and a small
sclerotized spur. Mouthparts relatively long.
Apex of maxillary palp similar to Texored-
dellia, but with a third very small extra con-
ule. Labial palp long, apical article barely
longer than wide, labium and first article of
labial palp with macrochaetae (Fig. 1D).
Mandible without very small pegs over the
bigger tooth. Setae as shown in Fig. 1E.
Legs as shown in Fig. 2A, relatively long;
hind tibia approximately 5 times longer
than wide; claws of normal size.

Cerci of male with a longer than wider
basal article and a very long second article,
followed by short articles of subequal size.
In the very long article the spines start al-
most at its base until they become a more
prominent group; the relatively long spines
are inserted on distinct tubercles (Fig. 3D).
Compared with other genus of the subfam-
ily, spines are relatively small.

Thoracic terga with long macrochaetae
(Fig. 1F), very abundant on all borders.
Also, on the posterior border they show a
series of small sclerotized spines (Figs. 1G,
3A). Tergum X protruding, almost straight
on the posterior border (Fig. 3C).

Urosterna III and IV of male without
modifications. Urosternum VIII with a wide
and not too deep posterior emargination.
Posterior projections acute to slightly
rounded, pointing slightly outward (Fig.
2B).

Urosternum IX of male slightly curved
behind and centrally on the posterior border
with macrochaetae slightly more sclerotized
(Fig. 2C). Point of insertion of parameres
in urosternum IX is deep, with internal face
of coxal processes with macrochaetae more
sclerotized (Fig. 2C) similar to some mem-
bers of the genus Anelpistina.

Stylets IX bigger than the others, with 6

a7

macrochaetae and an extra subapical pair.
The other stylets only have 3 macrochaetae
and an extra subapical pair. In males, styles
IX without other modifications.

Penis and parameres as in Fig. 2C. Par-
ameres curved outward, attaining less than
Y% the length of stylets IX. Surface of par-
ameres with a cieft and short setae.

Etymology.—Latin latebricola = One
who dwells in lurking places. Gender mas-
culine. Makes reference to the cavernicole
habitat where this species dwells.

Remarks.—Despite several trips to Pozas
Azules cave, only a single male was col-
lected. Because of its size, this specimen is
probably an adult with all the secondary
sexual characters developed. Individuals at
different stages of the postembryonic de-
velopment might differ in characters, such
as the number and size of spines in cerci.

Many caves in this area of Guerrero,
México, have been explored and have
yielded many individuals of the family Ni-
coletiidae (Espinasa 1991), but all from dif-
ferent genera. The one locality where S. /a-
tebricola has been found is in this single
cave. Therefore it is probably a troglobite
with a very restricted geography.

Acknowledgments

I thank Dr. José G. Palacios Vargas, di-
rector of “Laboratorio de Ecologia y Sis-
tematica de Microarstro6podos,’’ where most
of the descriptive work was done and to the
Director of CEAMISH-UAEM, Dr. Oscar
Dorado, for the support to publish this man-
uscript. Thanks are also due to Ramon Es-
pinasa-Perefia, geologist at the ‘‘Universi-
dad Nacional Aut6noma de México’’ for
providing the information regarding the
date of formation of the cave, and to Dr.
Luis E Mendes and Monika Baker for re-
viewing the manuscript.

Literature Cited

Espinasa, L. 1991. Descripci6n de una nueva especie
del género Cubacubana (Zygentoma: Nicoleti-
idae) y registro del género para América Con-

58 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

tinental.—Folia Entomoldégica Mexicana 82:5—
16.

Espinasa-Perena, R. 1989. El resumidero del Isote y la
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Lépez-Ramos, E. 1974. Geologia de México. Third
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Mendes, L. FE 1988. Sur deux nouvelles Nicoletiidae
(Zygentoma) cavernicoles de Gréce et de Tur-
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LAPS

Reddell, J. R. 1981. A review of the cavernicole fauna
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Silvestri, EF 1905. Materiali per lo studio dei Tisanuri.
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. 1933. Nuovo contributto alla conoscenza dei
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Solis-Weiss, V., & L. Espinasa. 1991. Lycastilla cav-
ernicola, a new freshwater nereidid from an in-
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Wygodzinsky, P. 1973. Description of a new genus of
cave thysanuran from Texas (Nicoletiidae, Thy-
sanura, Insecta).—American Museum Novitates
2518:1-8.

, & A. M. Hollinger. 1977. A study of Nico-

letiidae from Cuba (Thysanura).—Resultats des

Expéditions Biospéleologiques Cubano-Rou-

maines a Cuba 2:313-—324.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(1):59-—69. 1999.

Two new species of the genus Anelpistina (Insecta: Zygentoma:
Nicoletiidae) from Mexican caves, with redescription of the genus

Luis Espinasa

CEAMISH, Universidad Auténoma del Estado de Morelos, Av. Universidad #1001 Col.
Chamilpa, Cuernavaca, Mor. C.P. 62210. México

Abstract.—Two new species of the genus Anelpistina are described, A. in-
appendicata and A. cuaxilotla. Of these two, A. inappendicata lacks the artic-
ulated submedian appendages found in urosternite ITV of males, which are di-
agnostic characters of genus Anelpistina. Analysis of other characters shows
that this species has close affinities with members of genus Anelpistina, and
therefore a new re-description of the genus is provided to include this species.

Insects of the family Nicoletiidae are
common inhabitants of caves in México.
Numerous species of troglobites and trog-
lophiles have been collected from this area,
but with few exceptions, they await descrip-
tion (Reddell 1981). During the exploration
of several caves in the states of Guerrero
and Morelos, México, several specimens of
this family were collected and are described
here, thus contributing to the knowledge of
the cave fauna of the region.

Definition of genus Anelpistina by Sil-
vestri (1905) is based on the articulated
submedian appendages found in urosternite
IV of males. Organisms who are closely re-
lated to species of genus Anelpistina, but
lack such appendages, would be incorrectly
placed in genus Cubacubana (Wygodzin-
sky & Hollinger 1977). For this reason, ge-
nus Anelpistina is redescribed using more
characters to include species, such as the
one described in this paper, that lack the
appendages.

Materials and Methods

Detailed descriptions of the caves can be
found in Hoffman et al. (1986), Diamant &
Espinasa-Perefia (1991), and in Espinasa-
Perefia (1989). Organisms collected were
crawling on floor or under rocks. They were
placed into a vial with 70% alcohol or were

taken alive to the laboratory. Dissections
were made with aid of a stereo microscope
and the different parts of the body were
mounted in fixed preparations with Hoyer’s
solution.

Comparison with A. boneti was done
with specimens collected by author from
their type localities of Juxtlahuaca cave,
Colotlipa, Guerrero, México, and from a
new locality, the cave of “‘Iglesia-Mina su-
perior’’, San Juan, Tepoztlan, Morelos,
México. Comparison with other species
was based on published material. All illus-
trations were made with aid of a camera
lucida attached to a compound microscope.

Types were deposited in the following
collection: LESM-DB-MEX (Laboratory of
Ecology and Systematic of Microarthro-
pods. Department of Biology, Faculty of
Sciences, UNAM. México D. E). Catalog
numbers: ZYG-3 for Anelpistina inappen-
dicata and ZYG-4 for Anelpistina cuaxilot-
la.

Anelpistina Silvestri, 1905

Diagnosis.—Member of subfamily Cu-
bacubaninae without scales. Submentum
without conspicuous lateral lobes bearing
glandular pores. Urosternite IV of adult
male with 1 + 1 articulated submedian ap-
pendages and/or with point of insertion of

60 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

parameres in urosternum [IX deep, normally
with internal face of coxal processes with
macrochaetae more or less sclerotized.

Description emending.—Body slender,
approximately parallel-sided, thorax slight-
ly but distinctly wider than abdomen. An-
tennae and body are of similar length. Head
with approximately 8 + 8 macrochaetae in
border of insertion of antennae. Pedicellus
of male normally with 3 clusters of unicel-
lular glands, one of them very long.

Mouthparts not specialized. Mandibles
strongly sclerotized apically and with usual
teeth, typically with more than 4 macro-
chaetae. Galea apically with several sensory
pegs. Lacinia heavily sclerotized distally;
first process of lacinia pectinate. Labium
without prominent lateral lobes. Next to last
article of labial palp more or less with a
rounded bulkiness.

Tarsus longer or equal in length to tibia.
Tarsi with 4 articles. Praetarsi with 3 simple
claws, median claws glabrous, slender, and
smaller than lateral claws.

Abdominal sterna II—-VII subdivided into
coxites and sternite. Sterna VIII and IX of
male entire. Coxites on segments II-IX
with stylets. Eversible vesicles on segments
II-VI, pseudovesicles on VII. Urosterna III
simple. Urosterna IV in males with or with-
out | + 1 articulated submedian append-
ages. Point of insertion of parameres in
urosternum IX generally deep except in A.
weyrauchi (Wygodzinsky 1959) and to cer-
tain degree in A. ruckeri (Silvestri 1905),
normally with internal face of coxal pro-
cesses with macrochaetae more or less
sclerotized. Cerci of male with modified
spines.

Normal parameres, not subdivided or
somewhat constricted apically. With spe-
cialized setae on apex. Opening of penis
longitudinal.

Type species.—Anelpistina wheeleri Sil-
vestri 1905.

Remarks.—According to Silvestri’s
(1905) definition of genus Anelpistina, the
main difference between this genus and the
other members of the subfamily Cubacu-

baninae is the articulated submedian ap-
pendages in the urosternite IV of males.
Specimens that are closely related to spe-
cies of the genus Anelpistina, but lack such
appendages could incorrectly be placed in
the genus Cubacubana. Therefore a broader
re-description is given to include these
specimens.

With the new re-description, genus Anel-
pistina differs from Texoreddellia (Wygod-
zinsky 1973) by the absence of scales and
of a conspicuous process of the pedicellus
in males, as well as by the presence of mod-
ified spines on the cerci of males. From AI-
lonicoletia (Mendes 1992) by the presence
of stylets in urosternite Il. From Neonico-
letia (Paclt 1979) by the aspect of the en-
dopodium. Prosthecina (Silvestri 1933) and
Cubacubana agree with Anelpistina in the
possession of modified spines on the cerci
of males, but Prosthecina has a submentum
with conspicuous lateral lobes bearing nu-
merous glandular pores. Cubacubana lacks
1 + 1 articulated submedian appendages in
urosternite IV and the point of insertion of
parameres in urosternum IX is shallow
without more or less sclerotized macro-
chaetae in the internal face of coxal pro-
cesses. Anelpistina has either or both the
appendages and the deep insertion.

Anelpistina inappendicata, new species
Figs. 1A—K 2A-—G

Type material.—México, Guerrero State,
Tetipac municipality, Agua Brava cave (17
meters deep, 819 meters long). ex soil, 7
Aug 1988, 25 Mar 1989 and 15 Mar 1991,
L. Espinasa col. Male holotype, 4 male par-
atypes and ten female paratypes.

Description.—Maximum body length
13.5 mm. Maximum length of antennae 9.0
mm, of caudal appendages 9.0 mm. When
complete, antennae and caudal appendages
measure more or less the length of body.
General color light yellow to white.

Head with macrochaeta and microchaeta
as shown in Fig. 1E with approximately 8
+ 8 macrochaetae on border of insertion of

VOLUME 112, NUMBER 1

5 mm
[-oaaa aman nnnnnanann]

-1l mm.

[oo eae I

1 mn.
[------------]

1 mn.

(eS)

is N hil
(ay \
As ' Se
a : Sy
BS
a ee.
ue eS
iN wlan
'

61

-5 mn.

[sge- See -s-- =)

Fig. 1. Anelpistina inappendicata n. sp.: A, Mandible; B, Apical portion of maxilla; C, Labium with palp;
D, Hind leg, microchaetae partially shown; E, Male. Basal portion of antenna; K Head.

antennae. Basal article of antennae in males
without projections. Pedicellus of antennae
of male as shown in Fig. 1E, with numerous
clusters of unicellular glands. Female basal
articles of antennae simple.

Mouthparts relatively long, apex of max-
illa as shown in Fig. 1B. Labial palp long,
apical article longer than wide. Labium and
first article of labial palp with macrochae-
tae. Mandibles with approximately 8 ma-

62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

-5 mm.
[-nae wn an nna nennnnnn)

[ono 222-229 ------- ==]

[-oonnnnnnnnn nn nnnnn]

F
" . 4;

ye

SZ
Bn.

[-ooenn nanan]
-5 mm.
[ --------------------]

SO\ |
Ss

O\

SS

[-oraaanennn nanan ==o]

1 mn.
[--eonena-- ==]

Vile FSi G

Fig. 2. Anelpistina inappendicata n. sp.: A-B, Male. Genital area; C, Ovipositor and subgenital plate; D,
Subgenital plate; E, Basal portion of cercus; K Modified spines of cercus, higher magnification; G, Thoracic
nota. dorsal.

VOLUME 112, NUMBER 1

crochaetae, some very small pegs on larger
tooth (Fig. 1A). Legs as shown in Fig. 1D,
moderately elongate; hind tibia approxi-
mately 7X longer than wide. Claws of nor-
mal size.

Cerci of male with a longer than wide
basal article, a very long second one bear-
ing numerous spines, followed by numer-
ous short articles of simple chaetotaxy.
Spines along most of second article and of
similar size. Spines with medial enlarge-
ment in form of ring (Fig. 2E, F). Female
cerci simple.

Thorax with short macrochaetae, very
abundant on lateral borders, 1 + 1 subme-
dian distinct macrochaetae, apart from sev-
eral setae of varied sizes, in posterior border
of pro-, meso- and metanotum. 2 + 2 also
on anterior border of pronotum (Fig. 2G).

Urotergite X almost straight in both sex-
es, posterior angles with few macrochaetae
of varied sizes, length of inner macrochae-
tae almost equal to distance between them.

Abdominal terga and sterna as in other
members of genus. Abdominal sterna H-—
VII subdivided into coxites and sternite.
Sterna VIII and IX of male entire. Uroster-
num III and IV of male without modified
coxites. Urosternum VIII of male shallowly
emarginate on posterior margin. Uroster-
num IX of male straight behind without a
row of sensory cones, but point of insertion
of parameres in urosternum IX is deep, with
internal face of coxal processes with ma-
crochaetae more sclerotized (Fig. 2A, B).
Stylets H—VIII as usual for subfamily. Sty-
lets IX larger than others, with 2 or 3 ma-
crochaetae and an extra subapical pair. Ter-
minal spine with small teeth. In males and
females styles without modifications.

Penis and parameres as shown in Fig.
2A, B. Parameres attaining % of stylets IX,
divergent and slightly concave. Surface of
parameres with short setae. Subgenital plate
of female triangular, twice as wide as it is
long (Fig. 2C, D). Ovipositor barely sur-
passing apex of stylets IX. Gonapophyses
with approximately 12 articles.

Postembryonic development not very

63

complex, with younger instars almost iden-
tical to older ones except for size. In small-
er sized males (9 mm.), spines on cerci lack
medial enlargement in form of a ring (it is
unknown if younger instars than those col-
lected lack spines). In females, length of
Ovipositor increases proportionally with
body size until at body length of 9.5 mm,
Ovipositor barely surpasses apex of stylets
IX. Longer body sizes of up to 13.5 mm.
did not bring an increase of size of ovipos-
itor when compared to stylets IX.

Etymology.—In/appendicata = without/
appendages. Makes reference to the lack of
articulated submedian appendages in this
species, which are normally found in the
urosternite IV of males in the genus Anel-
pistina.

Remarks.—The characters that allow us
to identify A. inappendicata as more closely
related to species of the genus Anelpistina
than to Cubacubana, even though it lacks
such appendages are: Antennae and body
are of similar length, as in Anelpistina,
while in Cubacubana the antennae are
twice as long; Pedicellus of antennae with
three clusters of unicellular glands, one of
them very long. This is in general the case
in Anelpistina. The norm in Cubacubana is
four clusters bordered by a “U”’ shaped
row of microchaetae; Head with approxi-
mately 8 + 8 macrochaetae in the border of
insertion of the antennae, as in some Anel-
pistina, while Cubacubana normally has 5
+ 5; Lateral borders of thoracic nota with
approximately eight macrochaetae. This is
closer to most Anelpistina, with approxi-
mately five macrochaetae, than to Cubacu-
bana, with approximately three; Mandibles
with more than four macrochaetae, as in
some Anelpistina, while in Cubacubana
there are typically four; The next to last ar-
ticle of the labial palp without a rounded
bulkiness. In this character A. inappendi-
cata is, on the contrary, more similar to
species of the genus Cubacubana whose ar-
ticle is straighter, while in Anelpistina they
are bulky. This character could be ex-
plained by a loss of bulkiness in A. inap-

64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

pendicata; Tarsus longer than the tibia. Al-
though this is generally the case for Cuba-
cubana, in Anelpistina it is longer to equal;
Point of insertion of parameres in
urosternum IX deep, with internal face of
coxal processes with macrochaetae sclero-
tized. This deep insertion and the sclero-
tized macrochaetae are absent in Cubacu-
bana. On the contrary, the deep insertion is
present in all the species of Anelpistina, ex-
cept A. weyrauchi, and the sclerotized ma-
crochaetae are present in most of the spe-
cies.

Anelpistina inappendicata and the other
North American species of Anelpistina have
in common a deep insertion of the paramers
and other intermediate characteristics,
which are absent in the South American
species (A. weyrauchi). It is likely therefore
that the divergence of A. inappendicata
from the species of this genus happened af-
ter the separation of A. Weyrauchi from the
group, but before the radiation of the other
described North American species of Anel-
pistina.

Anelpistina decui (Wygodzinsky & Hol-
linger 1977), new combination Cubacu-
bana decui Wygodzinsky & Hollinger,
1977:320—322, figs. 4-6; Mendes, 1986:
341; Espinasa, 1991:6, 14.

Remarks.—Al\though A. decui lacks the
articulated submedian appendages on uros-
ternite IV of the male, this species has clos-
er affinities with members of Anelpistina
than it does to Cubacubana because its
point of insertion of parameres in uroster-
num IX is deep and the internal face of cox-
al processes has sclerotized macrochaetae.

An analysis of other characters present in
A. decui that allow us to identify it as more
closely related to species of genus Anelpis-
tina than to Cubacubana, even though it
lacks such appendages, is given based on
the illustrations and description of Wygod-
zinsky (1977): Antennae twice as long the
length of the body, as in Cubacubana,
while in Anelpistina antennae are of similar
length. Even though this character would
appear to make it closer to Cubacubana, its

longer antennae may be due to a secondary
adaptation to its cavernicole environment;
Pedicellus of antennae with two long clus-
ters of unicellular glands and no “U”
shaped row of microchaetae. In Anelpistina,
normally there are three clusters, one of
them very long. The norm in Cubacubana
is four clusters bordered by a “U”’ shaped
row of microchaetae; Lateral borders of
thoracic nota with approximately five ma-
crochaetae, as in most Anelpistina, while
Cubacubana approximately has three; The
next to last article of the labial palp with a
somewhat rounded bulkiness, as in Anelpis-
tina, while in Cubacubana it is straighter;
Cerci of male with four spines; a small one,
a long and strong one, another small and a
very long curved one. Although in both
Anelpistina and Cubacubana there is diver-
sity in morphology of spines, this particular
arrangement of four spines is exactly the
same for most species in genus Anelpistina.

Other characteristics that could further
establish to whom the species is more
closely related, such as the number of ma-
crochaetae on mandibles and on the border
of the insertion of antennae, regrettably
could not be obtained from the illustrations
of Wygodzinsky (1977).

Anelpistina cuaxilotla, new species
Figs. 3A—G, 4A—D

Type material.—Type locality: México.
Guerrero: Apetlanca, Cuaxilotla town, in
the penumbra zone (Entrance to 120 m) of
‘‘Cuaxilotla’’ cave, under rocks. 6 Jun 1987
and 23 Aug 1987. L. Espinasa col. Male
holotype, ten male paratypes and 12 female
paratypes. Other localities: México, More-
los state, Tepoztlan municipality, San Juan
town, ‘“‘San Juan’’ or ‘“‘Sistema Ferrocarril-
Mina inferior’? volcanic cave system, under
rocks. 21 Jan 1979. R. Garcia col. One male
paratype.

Description.—Maximum body length
12.0 mm. Maximum length of antennae 6.0
mm, of caudal appendages 5.0 mm. When
complete, antennae and caudal appendages

VOLUME 112, NUMBER 1 65

see
- ‘8 = fe)
Un - N ‘ey.
NS fetoch g
: V es ! i
at t 7) i a
ai Nyy a
ol ‘é »* wl
. ' Bs . '
1 == ! {
: =)
uy 1 raid
E Fee
D
a alts
Hihag
i Te a
j lly j il 7
4 Riteoy\. Vet 25.5.) bs
di Ga ae wri |
MTT ATA, .
ie Wek ~ehL Av
' > ~ . .
| | 38 a
: Us aN a4
ie Ve A iN
\ Vo :
!

=

mn.
[-seanenseeseantaas=]

i
1 >
1
w H ~
. - a mm
+ Ala
a 7
2 ai
j ! —
l rill
1 ! PS
i !
iro aren
| tes Ae
4 5
J
H

Fig. 3. A-—C, Anelpistina boneti (Wygodzinsky), D—-G, Anelpistina cuaxilotla n. sp.: A, Male, basal portion
of antenna; B, Female, basal portion of antenna; C, Mandible. D, Male, basal portion of antenna; E, Female,
basal portion of antenna; F Mandible; G, Male, abdominal sternum VIII; H, Male from “‘San Juan”’ cave, genital
area; I, Male from “‘Cuaxilotla” cave, genital area; J, Ovipositor and subgenital plate.

66 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

5 eae
[-onnn anne nnn

-5 mm.
[-o-nn naan nanan

Fig. 4. Anelpistina cuaxilotla n. sp.: A, Male from “‘Cuaxilotla’’ cave, appendages of urosternite IV; B, Male

from “San Juan” cave, appendages of urosternite IV; C, Male from ‘“Cuaxilotla’” cave, modified spines of
cercus; D, Male from “‘San Juan’’ cave, modified spines of cercus.

measure less than length of the body. Setae

on body strong, long and very abundant.
General color light yellow to white.

Head with macrochaetae and microchae-

tae similar to Fig. 1E Basal article of an-
tennae in males slightly longer than pedi-
cellus, but shorter than in A. boneti. In the
female it is also shorter than in A. boneti

VOLUME 112, NUMBER 1

(Fig. 3A, B, D—E). Pedicellus of male as
shown in Fig. 3D, with numerous clusters
of unicellular glands. Female basal articles
of antennae simple.

Mouthpart appendages relatively long,
very similar to A. boneti. Apex of maxillary
palp with two conules, one longer than wid-
er and the other wider than longer. Labial
palp long, apical article barely longer than
wide and only slightly longer than next to
last article. This penultimate article with
bulkiness with two macrochaetae. Labium
and first article of labial palp with macro-
chaetae. Mandibles without very small pegs
on bigger tooth and chaetotaxy as in Fig.
3E Legs as in A. boneti, with sclerotized
macrochaetae on tibia. Claws of normal
SiZe.

Cerci of male with a longer than wide
basal article, second article wider than lon-
ger and a very long third one bearing 4
spines in distal portion, followed by nu-
merous short articles of simple chaetotaxy.
Spines consist of a very small one, a strong,
subacute one, another very small, and a
long, acute and slightly curved one (Fig.
4C, D). Cerci of female simple.

Abdominal sterna and terga as in other
members of the genus. Setae long and nu-
merous. Thorax with shorter setae than on
urosterna. Lateral borders as in A. boneti,
with 4—5 macrochaetae. Urotergite X long
and almost straight in both sexes, posterior
angles with 2 + 2 macrochaetae and a few
relatively strong setae.

Appendages of urosternum IV of male
very long and robust, acute on apex. Their
longest diameter equals the length of the
stylets of this segment (Fig. 4A, B).

Urosternum VIII of male long and shal-
low shallowly emarginate on the posterior
margin (Fig. 3G), slightly deeper than in A.
boneti.

Urosternum IX of male with setae abun-
dant. Behind insertion of parameres, in the
center, with a small group of short, sclero-
tized and spiniform setae. Internal face of
coxal processes with irregular row of 5 spi-
niform macrochaetae highly sclerotized

67

(Fig. 31). In the only male individual col-
lected from cave of San Juan (Sistema Fer-
rocarril-Mina inferior), this row is of ten
macrochaetae (Fig. 3H). It is not known if
this difference is due to population differ-
ences or because the individual of San Juan,
being the biggest (12.0 mm) corresponds to
an older instar.

Stylets similar to A. boneti. Stylets IX
bigger than the others. Penis and parameres
as shown in Fig. 3H, I. Parameres attain the
length of stylets IX. Surface of parameres
with short setae.

Subgenital plate of female rounded, with
the apex slightly flat (Fig. 3J). Ovipositor
surpassing apex of stylets IX by twice the
length of stylets. Gonapophyses with ap-
proximately 17 articles.

Postembryonic development not very
complex with younger instars almost iden-
tical to older ones except for size. In male,
appendages of urosternite IV appear very
small at a length of approximately 6.0 mm
and acquire adult morphology at a length of
7.0 mm. In the longest individual (12.0
mm) only a slight increase in the propor-
tions of sexual secondary characters is ob-
served.

Female development not well known be-
cause all females collected measure more
than 7.5 mm, when length of ovipositor al-
ready surpasses apex of stylets IX by one
and a half to two times its size.

Etymology.—Cuaxilotla. Makes refer-
ence to the type locality, ‘““Cuaxilotla”’ cave.

Remarks.—The closest species to Anel-
pistina cuaxilotla are A. bolivari, A. ano-
phtalma, and A. boneti. In males of A. cuax-
ilotla, urosternum VIII is shallowly emar-
ginate on posterior margin, while in A. bo-
livari, emargination is narrow and deep.
Other characteristics that differentiate these
two species are the spines in the cerci and
the abundance of setae in terga and sterna.
From A. anophtalma it differs because in
this species, urotergite X is deeply emar-
ginate on the posterior margin, appendages
of urosternite IV are thin and ovipositor is
short, while in A. cuaxilotla, urotergite X is

68 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON .

almost straight, appendages are broad and
ovipositor is long. For A. boneti, the closest
species to A. cuaxilotla, the main difference
is that in A. boneti the appendages of uros-
ternite IV are thin, while in A. cuaxilotla
they are broad. Other characteristics that
differentiate them are that in the former, the
cerci have three spines (although some in-
dividuals from ‘‘Iglesia-Mina superior’’
cave in Morelos have 4) while in A. cuax-
ilotla they have four; the basal article of the
antennae in males and females is propor-
tionally longer than the pedicellus in A. bo-
neti than it is in A. cuaxilotla (Fig. 3A, B,
D, E); and the chaetotaxy of the mandible
(Figs. 3C and F). To establish these differ-
ences with precision, a large number of in-
dividuals of A. boneti collected from the
type locality by the author were compared
to the individuals of A. cuaxilotla.

The somewhat overlapping distribution
of A. cuaxilotla and A. boneti is intriguing.
The latter was reported to a wide distribu-
tion comprising the states of Guerrero, Mo-
relos, D.E and even Habana island in the
Gulf of California (Wygodzinsky 1946),
while A. cuaxilotla has been found in Guer-
rero and Morelos. Wygodzinsky (1946) ini-
tially distrusted such a big geographic dis-
tribution of A. boneti, although repeated
studies failed to show differences between
samples of different origins (“‘Desconfia-
mos inicialmente da grande distribu¢cao da
espécie, repetidos exames detalhados fal-
haram de demostrar diferengas entre os ex-
emplares de procedéncia diferente’’).

Regardless of whether organisms from
the Gulf of California are indeed A. boneti,
the author of the present paper has found a
locality where both species can be found in
close proximity. Individuals that clearly be-
long to A. boneti have been collected in the
cave system of the “‘Iglesia-Mina superior”’
(Morelos, Tepoztlan, San Juan). This cave
system is only tens of meters away from the
cave system of Ferrocarril-Mina inferior,
where an individual of A. cuaxilotla was
found. Individuals of both species from
these two caves can clearly be differentiated

morphologically, especially because sam-
ples of both contain mature males.

It is currently not known if the presence
of four spines on the cerci in males of A.
boneti from Iglesia-Mina superior cave, in-
stead of three spines as it is the norm for
this species, reflects variation within the
species, or if it instead reflects local hybrid-
ization between both species in these neigh-
boring caves.

Acknowledgments

I wish to thank Dr. José G. Palacios Var-
gas, director of “‘Laboratorio de Ecologia y
Sistematica de Microarstr6podos”’, where
most of the descriptive work was done and
to the Director of CEAMISH-UAEM, Dr.
Oscar Dorado, for the support to publish
this manuscript. Thanks are also due to Dr.
Luis E Mendes and Monika Baker for re-
viewing the manuscript.

Literature Cited

Diamant, R., & R. Espinasa-Perena. 1991. La cueva
de Agua Brava.—Tepeyollotli: Gaceta de la So-
ciedad Mexicana de Exploraciones Subterradneas
5:46—48.

Espinasa, L. 1991. Descripcién de una nueva especie
del género Cubacubana (Zygentoma: Nicoleti-
idae) y registro del género para América Con-
tinental.—Folia Entomolédgica Mexicana 82:5—
16.

Espinasa-Perefia, R. 1989. Proyecto Chilacachapa, re-
porte del progreso.—Tepeyollotli: Gaceta de la
Sociedad Mexicana de Exploraciones Subterra-
neas 4:32—40.

Hoffman, A., J. G. Palacios-Vargas, & J. B. Morales-
Malacara. 1986. Manual de bioespeleologia.
Universidad Nacional Aut6énoma de México,
México, 274 pp.

Mendes, L. E 1986. Nouvelles données sur le Zygen-
toma (Insecta) de 1’Amérique Centrale et du
Mexique.—Bulletin du Muséum national
d’ Histoire naturelle., Paris (4)8(A)(2):333-342.

. 1992. Novos dados sobre os tisanuros (Micro-
coryphia e Zygentoma) da América do Norte.—
Garcia de Orta 16(1—2):171-193.

Paclt, J. 1979. Neue Beitrage zur Kenntnis der Apter-
ygoten-Sammlung des Zoologischen Instituts
und Zoologischen Museums der Universitat
Hamburg. VI. Weitere Doppel- und Borstensch-
wanze (Diplura: Campodeidae; Thysanura: Lep-
ismatidae und Nicoletiidae).—Entomologische

VOLUME 112, NUMBER 1 69

Mitteilungen aus dem zoologischen Museum Wygodzinsky, P. 1946. Sobre Nicoletia (Anelpistina)

Hamburg 6(105):221—228. Silvestri 1905 e Prosthecina Silvestri, 1933.—
Reddell, J. R. 1981. A review of the cavernicole fauna Ciencia T:AS—25.
of México, Guatemala and Belize.—Texas Me- . 1959. Contribution to the knowledge of the

morial Museum Bulletin 27:204-205. me eae, eiaeuilidae” “(insecta).—
Revista Brasileira de Biologia 19(4):441—457.
. 1973. Description of a new genus of cave
Thysanura from Texas (Nuciletiidae, Thysanu-
ra, Insecta).—American Museum Novitates

Silvestri, E 1905. Materiali per lo studio dei Tisanuri.
VI. Tre nuove specie di Nicoletia appartenenti
ad un nuovo sottogenero.—Redia (Firenze) 2:

2a 2518:1-8.

. 1933. Nuovo contributo alla conoscenza dei Wygodzinsky, P, & A. M. Hollinger. 1977. A study of
tisanuri de Mesico.—Bolletino del Laboratorio Nicoletiidae from Cuba (Thysanura).—Resul-
di Zoologia general e agraria di Portici 27:127— tats des Expéditions Biospéleologiques Cubano-

144. Roumaines a Cuba 2:313-324.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
112(1):70-93. 1999.

Anacroneuria from northeastern South America
(Insecta: Plecoptera: Perlidae)

Bill P. Stark
Department of Biology, Mississippi College, Clinton, Mississippi 39058, U.S.A.

Abstract.—Descriptions are given for ten new species of Anacroneuria from
Venezuela, Guyana, and Suriname, and records are given for ten species pre-
viously described from this region. The following 10 new species are described
(type locality in parenthesis): Anacroneuria achagua (Venezuela. Portuguesa:
Guanare), A. arawak (Suriname. Brokopondo: Brownsberg Naturpark, Maza-
roni Plateau), A. chaima (Venezuela. Sucre: Rio Cocollar, 1.5 km SE Las Pie-
dras de Cocollar), A. claudiae (Venezuela. Zulia: Rio Yasa (3 km E Kasmera),
Estacion Biologica), A. karina (Venezuela. Sucre: Rio Cocollar, 1.5 km SE Las
Piedras de Cocollar), A. makushi (Guyana. Kanuku Mountains, Moco Moco
River), A. paria (Venezuela. Sucre: Rio Cocollar, 1.5 km SE Las Piedras de
Cocollar), A. perija (Venezuela. Zulia: Parque Nacional Perija, Rio Negro, To-
romo), A. timote (Venezuela. Tachira: Quebrada Los Mirtos, 8 km S El Cobre),
and A. wapishana (Guyana. Potaro River, Kaieteur Falls). Anacroneuria phan-
toma (Banks) and A. pictipes Klapalek are redescribed and a modified Key to

regional males is presented.

Stark (1995) provided descriptions of Ve-
nezuelan Anacroneuria based on collec-
tions made primarily through 1985. Re-
cently a series of Venezuelan and Guyanan
specimens collected in 1994—97 were made
available by O. S. Flint and R. Holzenthal,
and a few specimens were provided by the
California Academy of Science. This ma-
terial includes nine new species, new re-
cords for eight previously described spe-
cies, and a few unassociated females de-
scribed under informal designations. Also
included are redescriptions of A. pictipes
Klapalek and A. phantoma (Banks) from
holotypes and an additional new species
from Suriname. Holotypes are deposited in
the National Museum of Natural History,
Washington, D.C. (USNM) or the Califor-
nia Academy of Science, San Francisco,
CA (CAS). Paratypes and other specimens
are deposited in the Universidad Central de
Venezuela, Maracuy (UCV), the University
of Minnesota, Saint Paul (UMSP) or the
collection of the author (BPS). Additional

material was examined from the Museum
of Comparative Zoology, Harvard Univer-
sity (MCZ) and the National Museum of
Natural History, Prague (NMBH).

Anacroneuria achagua, new species

Adult habitus.—Head with an obscure
brown pattern over anterior half of frons;
lappets brown. Median pronotal stripe pale,
broad irregular lateral bands brown (Fig. 1).
Wing membrane transparent, veins pale.

Male.—Forewing length 8 mm Hammer
thimble shaped, height greater than basal
diameter (Fig. 2). Aedeagal apex short, with
wide, emarginate tip offset from shoulders;
ventral aspect with an obscure pair of mem-
branous lobes; dorsal keel triangular; shoul-
ders somewhat quadrate, hooks stout (Figs.
3-5).

Female.—Unknown.

Nymph.—Unknown.

Etymology.—The the

name honors

71

VOLUME 112, NUMBER 1

joe os wae
i‘

‘ ee
Cr

2 os
TN he Oe erent,
RA a)

ventral. 4. Ae-

°

gus

dorsal. Scales: 0.6 mm (1), 0.3 mm (2), 0.15 mm (3-5).

>

A. achagua structures. 1. Head and pronotum. 2. Male sternum 9. 3. Aedea

Figs. 1-5.
deagus, lateral. 5. Aedeagus

72 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 6-11.
Aedeagus, ventral. 10. Aedeagus, lateral. 11. Aedeagus, dorsal. Scales: 0.6 mm (6), 0.3 mm (7, 8), 0.15 mm
(9-11).

Achagua people of Venezuela and is used
as a noun in apposition.

Types.—Holotype ¢ and ¢ paratype
from Venezuela, Portuguesa, Guanare, 10—
13 Sep 1957, B. Malkin (CAS).

Diagnosis.—This species has a V-shaped
aedeagal keel and apex somewhat like that
of A. pictipes but these species are easily
distinguished by the distinctive wing and
pronotal pigmentation of that species (Figs.
48, 53).

TY Pls
Ae (LLY Pea ;
IS / iF M¢ itta\
d pe Lo
eee wi. —

\\AS aa
NUE
5 y,

A. arawak structures. 6. Head and pronotum. 7. Female sterna 8 and 9. 8. Male sternum 9. 9.

Anacroneuria arawak, new species

Adult habitus.—Head with diffuse yel-
low brown area extending forward from
ocelli; pronotum with diffuse brown lateral
bands and a narrow pale median band (Fig.
6). Wing membrane transparent, veins pale.

Male.—Forewing length 9 mm. Hammer
thimble shaped, height subequal to basal di-
ameter (Fig. 8). Aedeagal apex slender,
scoop shaped with a pair of subapical ven-

VOLUME 112, NUMBER 1

tral membranous lobes. Apex curved slight-
ly dorsad; hooks slender, dorsal keel long
and narrow (Figs. 9-11).

Female.—Forewing length 10.5 mm. Su-
bgenital plate four lobed, median lobes
shorter than lateral lobes; inner margins of
lateral lobes bearing a short setal fringe.
Sternum 9 with a broad transverse sclerite
and a trilobed median setal patch; median
lobe clothed with minute setae, lateral lobes
with prominent setae (Fig. 7).

Nymph.—Unknown.

Etymology.—The name honors the Ara-
wak people of Guyana and Suriname and is
used as a noun in apposition.

Types.—Holotype ¢ and @ paratype
from Suriname, Brokopondo District,
Brownsberg Naturpark, Mazaroni Plateau,
400-500 m, 16 Aug 1982, W. E. Steiner
(USNM).

Diagnosis.—This species does not ap-
pear to be closely related to other regional
Anacroneuria. The aedeagal shape and pale
habitus are somewhat similar to A. isleta
(Stark 1994) but that species has a trian-
gular keel and a shorter and more rounded
aedeagal tip. In dorsal aspect the aedeagus
is similar to that of A. bari (Stark 1995),
but that species does not have membranous
ventral lobes on the aedeagus. The female
is distinctive by virtue of the setal fringe on
the lateral subgenital plate lobes (Fig. 7).

Anacroneuria aroucana Kimmins

Anacroneuria aroucana Kimmins, 1948:
105. Holotype 6, Arouca River, Trinidad.
Anacroneuria aroucana: Stark, 1994:173.

Material.—Venezuela: Sucre, Quebrada
Zapateral, 1.5 km SE Las Piedras de Co-
collar, 810 m, 9 Apr 1995, R. Holzenthal,
O. S. Flint, 4 ¢ (USNM, UCV). Sucre, Rio
Cocollar, 1.5 km SE Las Piedras de Cocol-
lar; 810 m, 7-8 Apr 1995, R. Holzenthal,
O. S. Flint, 10 6 (USNM).

Comments.—These are the first mainland
records for this species.

73

Anacroneuria bari Stark

Anacroneuria bari Stark, 1995:226. Holo-
type 6, El Tucuco, 45 km SW Mac-
hiques, Zulia, Venezuela.

Material.—Venezuela: Guarico, Parque
Nacional Guatopo, Quebrada Guatopo, 0.5
km NE La Colina, 600 m, 22 Jan 1994, R.
Holzenthal, C. Cressa, Rincén, 3 6 (UMSP.
UCV). Trujillo, Quebrada Potrerito, 7.5 km
NE Bocono, 1530 m, 29-30 Apr 1995, R.
Holzenthal, C. Cressa, Gutic, 2 ¢ (BPS).

Comments.—This species was previously
reported from the Venezuelan states of Bar-
inas, Miranda, and Zulia (Stark 1995).

Anacroneuria bifasciata (Pictet)

Perla bifasciata Pictet, 1841:231. Holotype
2, Moritz, Colombia.

Anacroneuria bifasciata: Zwick, 1972:
1154.

Anacroneuria bifasciata: Stark, 1995:239.

Material.—Venezuela: Distrito Federal,
Rio Camuri Grande, 1 km S Camuri (nu-
cleo USB), 30 m, 24 Jan 1994, R. Holzen-
thal, C. Cressa, Rincén, 1 6, 1 2 (BPS).

Comments.—This species was previously
reported from the Venezuelan states of Ar-
agua, Distrito Federal, and Lara (Stark
1995).

Anacroneuria caraca Stark

Anacroneuria caraca Stark, 1995:228. Ho-
lotype ¢, Rio Limon, Parque Nacional
Henri Pittier, Aragua, Venezuela.

Material.—Venezuela: Guarico, Parque
Nacional Guatopo, Quebrada Guatopo, 0.5
km NE La Colina, 600 m, 22 Jan 1994, R.
Holzenthal, C. Cressa, Rincon, 1 ¢
(UMSP).

Comments.—This species was previously
reported from the Venezuelan states of Ar-
agua and Barinas (Stark 1995).

Anacroneuria chaima, new species

Adult habitus.—Head with a wide brown
area forward of ocelli extending to M-line;

74 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

lappets and anterior margin of head brown.
Median pronotal stripe pale; broad lateral
stripes dark brown (Fig. 12). Wing mem-
brane and veins brown.

Male.—Forewing length 8 mm. Hammer
cylindrical, height greater than basal diam-
eter (Fig. 13). Aedeagal apex simple, scoop
shaped, slender and rounded at the tip; ven-
tral aspect with a small circular pair of
membranous lobes; dorsal aspect with a
low wide keel; hooks slender (Figs. 14—16).

Female.—Unknown.

Nymph.—Unknown.

Etymology.—The name honors the Chai-
ma people of Venezuela and is used as a
noun in apposition.

Types.—Holotype ¢ (pinned) from Ven-
ezuela, Sucre, Rio Cocollar, 1.5 km SE Las
Piedras de Cocollar, 810 m, 7—8 Apr 1995,
R. Holzenthal, O. S. Flint (USNM).

Diagnosis.—This species is similar to A.
perija in general aedeagal structure but dif-
fers in having small, circular membranous
aedeagal lobes (Fig. 14) and in having the
arms of the dorsal aedeagal keel divergent
(Fig. 16). The dark color pattern of A. per-
ija is also much more prominent, particu-
larly over the occiput (Fig. 38).

Anacroneuria chorrera Stark

Anacroneuria chorrera Stark, 1995:230.
Holotype 3, La Chorrera Canyon, Meri-
da, Venezuela.

Material.—Venezuela: Merida, Rio La
Gonzalez, between Merida and Jaji, 1870
m, 25 Apr 1995, R. Holzenthal, C. Cressa,
Gutic, 3 ¢ (UCV). Merida, Rio Albarregas,
1 km NW Universidad de los Andes, 1980
m, 17 Jan 1994, R. Holzenthal, C. Cressa,
Rincon, 3 6 (UMSP). Merida, Parque Na-
cional Sierra Nevada, Quebrada La Mucuy,
7 km E Tabay, 2200 m, 18 Jan 1994, R.
Holzenthal, C. Cressa, Rincén, 4 d (BPS).
Trujillo, Quebrada Potrerito, 7.5 km NE
Bocon6, 1530 m, 20—30 Apr 1995, R. Hol-
zenthal, C. Cressa, Gutic, 4 6 (UMSP).
Tachira, tributary to El Valle, 3.8 km SE El
Zumbador, 2730 m, 21 Apr 1995, R. Hol-

zenthal, C. Cressa, Gutic, 7 6 (UMSP).
Tachira, Quebrada Los Mirtos, 8 km S El
Cobre, 2400 m, 22 Apr 1995, R. Holzen-
thal, C. Cressa, Gutic, 1 6 (UCV).

Comments.—This species was previously
reported from the Venezuelan states of Ar-
agua, Distrito Federal, and Merida (Stark
1995).

Anacroneuria claudiae, new species

Adult habitus.—Head yellow with dif-
fuse brown pattern extending from ocelli to
anterior margin but interrupted by three
pale areas at M-line; lappets brown. Median
pronotal stripe pale, lateral stripes dark, an-
terolateral margins with pale spot (Fig. 17).
Wing membrane transparent, veins brown.

Male.—Forewing length 8 mm. Hammer
thimble shaped, height greater than basal
diameter (Fig. 18). Aedeagal apex simple,
scoop shaped, slender and truncate at the
tip; ventral aspect with a small pair of mem-
branous lobes; dorsal aspect with a narrow
keel, hooks slender (Figs. 19-21).

Female.—Unknown.

Nymph.—Unknown.

Etymology.—The matronym honors
Claudia Cressa of the Universidad Central
de Venezuela, Caracas.

Types.—Holotype ¢ from Venezuela,
Zulia, Rio Yasa, 3 km E Kasmera (Estacion
Biologica), 150 m, 14 Jan 1994, R. Holzen-
thal, C. Cressa, Rincén (USNM). Paratype
6 from Venezuela, Zulia, Parque Nacional
Perija, Rio Negro, Toromo, 360 m, 15 Jan
1994, R. Holzenthal, C. Cressa, Rincén
(UCV).

Diagnosis.—This species is similar to A.
chiquita in general aedeagal structure but
differs from that species in having small
paired membranous ventral lobes (Fig. 17)
and in having unmodified aedeagal hook
apices (Fig. 17).

Anacroneuria cruza Stark

Anacroneuria cruza Stark, 1995:231. Ho-
lotype 6, Exp. Culebra, N Duida, Terri-
torio Federal Amazonas, Venezuela.

a

VOLUME 112, NUMBER 1

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a

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.

.

.

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.
.

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ov?

.

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.

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. Aedeagus, ventral. 15.

Male sternum 9. 14

Head and pronotum. 13.

12

ima structures

A. cha

Aedeagus, lateral. 16. Aedeagus

Figs. 12-16.

16).

0.15 mm (14—

0.6 mm (12), 0.3 mm (13),

. Scales

dorsal

>

76 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Ta en ees iw tne a patie
. *°,
'

ty

CS CE ee secicvcs Pees
ot een ee me en So emssee lt ae
. : ; Cite: eee

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4

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Figs. 17-21. A. claudia structures. 17. Head and pronotum. 18. Male sternum 9. 19. Aedeagus, ventral. 20.
Aedeagus, lateral. 21. Aedeagus, dorsal. Scales: 0.6 mm (17), 0.3 mm (18), 0.15 mm (19-21).

VOLUME 112, NUMBER 1

Material.—Guyana: Kanuku Mountains,
Kumu River, 28-30 Apr 1995, O. S. Flint,
2 6 (USNM). Kanuku Mountains, Moco
Moco River, 29 Apr 1995, O. S. Flint, 1 d
(USNM). Venezuela: Sucre, Rio Cocollar,
1.5 km SE Las Piedras de Cocollar, 810 m,
7-8 Apr 1995, R. Holzenthal, O. S. Flint,
4 6 (USNM, UCV).

Comments.—This species was previously
known from Territorio Federal Amazonas,
Venezuela (Stark 1995).

Anacroneuria karina, new species

Adult habitus.—Head yellow with a dif-
fuse brown spot forward of ocelli; lappets
brown. Irregular midlateral pronotal bands
dark brown, median field pale, grading to
pale brown (Fig. 22). Wing membrane pale
brown, veins brown, R and Sc dark brown.

Male.—Forewing length 11 mm. Ham-
mer thimble shaped, height greater than
basal diameter (Fig. 23). Aedeagal apex
simple, scoop shaped and broadly rounded;
ventral membranous lobes narrow; dorsal
keel well developed; hooks slender (Figs.
24-26).

Female.—Unknown.

Nymph.—Unknown.

Etymology.—The name honors the Kar-
ima people of Venezuela and is used as a
noun in apposition.

Types.—Holotype 6 (USNM) and 7 6
paratypes (USNM, UCV) from Venezuela,
Sucre, Rio Cocollar, 1.5 km SE Las Piedras
de Cocollar, 810 m, 7—8 Apr 1995, R. Hol-
zenthal, O. S. Flint.

Diagnosis.—Many species of Anacro-
neuria share the color pattern of A. karina
and the aedeagus also bears a general sim-
ilarity to that of other species such as A.
bari and A. arcuata (Stark 1995). Anacro-
neuria karina, however, lacks the transverse
dorsal aedeagal keel found in A. arcuata
and the aedeagal apex is much longer and
narrower in A. bari.

Anacroneuria llana Stark

Anacroneuria llana Stark, 1995:234. Ho-
lotype ¢, La Escalera, 108 km S Rio Cu-
yuni, Bolivar, Venezuela.

77

Material.—Guyana: Potaro River, Kaie-
teur Falls, 1350’, 21-23 Aug 1997, O. S.
Flint, 9 ¢6, 58 2 (USNM).

Comments.—This species was previously
known from the holotype. The Guyana
males agree with the holotype in all re-
spects except in having falcate aedeagal
hooks. The females collected with these
males share the same color pattern (Fig. 63)
and are associated on that basis. These fe-
males have eggs with a terminal spine sim-
ilar to those of A. blanca (Stark 1995) and
they also are similar in color pattern, size
and in having an intersegmental band of mi-
crotrichia.

Female.—Forewing length 13—14 mm.
Subgenital plate four lobed, outer lobes
slightly longer. Median sclerite of sternum
9 densely hirsute, hairs slightly longer lat-
erally. Intersegmental membrane covered
with a dense microtrichia band (Fig. 64).

Anacroneuria makushi, new species

Adult habitus.—Head yellow; pronotum
with diffuse midlateral brown bands (Fig.
27). Wing membrane transparent, veins
pale.

Male.—Forewing length 9.5 mm. Ham-
mer laterally compressed, height greater
than basal diameter (Fig. 28). Aedeagal
apex scoop shaped, tip blunt; ventral aspect
with a pair of subapical membranous lobes;
dorsal aspect with a long, low keel; dorsal
sclerite basal to keel bearing an arcuate pro-
cess; hooks irregularly scalloped on inner
margin (Figs. 29-31).

Female.—Unknown.

Nymph.—Unknown.

Etymology.—The name honors the Mak-
ushi people of Guyana and is used as a
noun in apposition.

Types.—Holotype 6 (pinned) and 2 ¢
paratypes from Guyana, Kanuku Moun-
tains, Moco Moco River, 29 Apr 1995, O.
S. Flint (USNM).

Diagnosis.—A. makushi is similar to A.
baniva in aedeagal structure (Stark 1995).
The most conspicuous difference between

78

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

er

—~
Sever on
a

et “peece
dd

Py

' < U
CON TOC, pO oe hee eres ee mre owe

77

2
© -
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% 28,

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3

Figs. 22—26. A. karina structures. 22. Head and pronotum. 23. Male sternum 9. 24. Aedeagus, ventral. 25.
Aedeagus, lateral. 26. Aedeagus, dorsal. Scales: 0.6 mm (22), 0.3 mm (23), 0.15 mm (24-26).

VOLUME 112, NUMBER 1 79

. enews mw en
=_—% OF ——
ar re EA a i

e ge

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tee

Figs. 27-31. A. makushi structures. 27. Head and pronotum. 28. Male sternum 9. 29. Aedeagus, ventral. 30.
Aedeagus, lateral. 31. Aedeagus, dorsal. Scales: 0.6 mm (27), 0.3 mm (28), 0.15 mm (29-31).

80 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

these species is the arcuate process on the
dorsal aedeagal sclerite in A. makushi (Fig.
SE):

Anacroneuria paleta Stark

Anacroneuria paleta Stark, 1995:236. Ho-
lotype 6, 4 km S Santo Domingo, Mer-
ida, Venezuela.

Material.—Venezuela: Trujillo, Quebra-
da Potrerito, 7.5 km NE Bocono, 1530 m,
29-30 Apr 1995, R. Holzenthal, C. Cressa,
Gutic, 31 d6 (UMSP, UCV, BPS).

Comments.—This species was previously
recorded from the Venezuelan states of Bar-
inas and Merida (Stark 1995).

Anacroneuria paria, new species

Adult habitus.—Very diffuse brown pat-
tern over ocelli and midlaterally on prono-
tum (Fig. 32). Wing membrane pale with
distinctive longitudinal brown band extend-
ing from anal cell to tip; band narrow in
proximal half, expanded at the cord, but in-
terrupted by a large circular ““window”’ be-
yond the cord (Fig. 37).

Male.—Forewing length 10 mm. Ham-
mer thimble shaped, height less than basal
diameter (Fig. 33). Aedeagal apex with five
dorsal lobes and a short narrow keel; ven-
tral aspect of mesal lobe scoop shaped,
without membranous lobes; hooks slender
(Figs. 34—36).

Female.—Forewing length 12 mm. Su-
bgenital plate four lobed; lateral lobes larg-
er than inner lobes, mesal notch deep and
V-shaped. Transverse sclerite of sternum 9
wide, sinuate and prominent, mesal sclerite
T-shaped; lateral lobes of mesal sclerite
with prominent setae, stalk covered with
minute setae (Fig. 54).

Nymph.—Unknown.

Etymology.—The name is based on the
Paria Peninsula and is used as a noun in
apposition.

Types.—Holotype d (pinned) and 9 2?
paratypes from Venezuela, Sucre, Rio Co-
collar, 1.5 km SE Las Piedras de Cocollar,

810 m, 7—8 Apr 1995, R. Holzenthal, O. S.
Flint (USNM). Additional paratypes; Ven-
ezuela: Sucre, Parque Nacional Peninsula
de Paria, Uquire, Rio La Vidua, 15 m, 30
Mar—1 Apr 1995, R. Holzenthal, O. S. Flint,
C. Cressa, 3 2 (UCV). Sucre, Peninsula de
Paria, Puerto Viejo, “Rio el Pozo’’, 20 m,
3 Apr 1995, R. Holzenthal, O. S. Flint, C.
Cressa, 6 2 (USNM).

Diagnosis.—This species is similar to A.
bifasciata (Pictet) and A. vistosa Stark in
displaying a distinctive wing pigmentation
pattern, however A. paria is readily distin-
guished from both species on the basis of
color pattern, aedeagal shape and details of
female sternum 9 (Stark 1995). Anacroneu-
ria bifasciata adults have tiny ocelli, a
prominent dark mesal pronotal band and the
wing pigmentation is separated into three
distinct transverse bands. The aedeagal
apex of A. bifasciata also bears five dorsal
lobes but the mesal lobe lacks a keel and
the mesal sclerite of the female 9th sternum
is triangular and evenly setose. The aedea-
gal apex of A. vistosa is trilobed and the
mesal sclerite of the female 9th sternum is
evenly setose.

Anacroneuria perija, new species

Adult habitus.—Center of frons bright
yellow, anterior head margin, posterolateral
head margins, and ocellar area dark brown.
Median pronotal stripe pale, lateral third
brown (Fig. 38). Wing membrane brown,
veins dark brown, Sc pale.

Male.—Forewing length 9 mm. Hammer
cylindrical, height greater than basal diam-
eter (Fig. 39). Ventral aedeagal apex a short
scoop covered with a pair of membranous
lobes; dorsal aspect with a wide keel and a
low U-shaped region; hooks slender (Figs.
40-42).

Female.—Unknown.

Nymph.—Unknown.

Etymology.—The name is based on the
type locality and is used as a noun in ap-
position.

Types.—Holotype ¢ from Venezuela,

VOLUME 112, NUMBER 1 81

Figs. 32-37. A. paria structures. 32. Head and pronotum. 33. Male sternum 9. 34. Aedeagus, ventral. 35.
Aedeagus, lateral. 36. Aedeagus, dorsal. 37. Forewing. Scales: 1.2 mm (37), 0.6 mm (32), 0.3 mm (33), 0.15
mm (34-36).

82

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

eet te
eee
vs.)

~ . .
meme oe,
ie a ee

ar es
a

’
me we. ld <
. =
. poe

42

Figs. 38-42. A. perija structures. 38. Head and pronotum. 39. Male sternum 9. 40. Aedeagus, ventral. 41.
Aedeagus, lateral. 42. Aedeagus, dorsal. Scales: 0.6 mm (38), 0.3 mm (39), 0.15 mm (40-42).

VOLUME 112, NUMBER 1

Zulia, Parque Nacional Perija, Rio Negro,
Toromo, 360 m, 15 Jan 1994, R. Holzen-
thal, C. Cressa, Rinc6én (USNM).

Diagnosis.—This species keys to A. ban-
iva in Stark (1995) but the aedeagal apex is
much shorter and narrower than in that spe-
cies, and the dorsal keel and subapical U-
shaped process of A. perija also will distin-
guish the two. Although the condition of A.
baniva type material left much of the color
pattern obscured, it is clear that species
lacks the distinctive head pattern of A. per-
ija.

Anacroneuria phantoma (Banks)

Neoperla phantoma Banks, 1914:609. Ho-
lotype 6, Mallali, Guyana.

Adult habitus.—Head yellow with dif-
fuse yellow brown lappets. Pronotum with
pale median stripe and broad, diffuse brown
midlateral bands; lateral margins pale (Fig.
43). Wing membrane and veins pale.

Male.—Forewing length 9-10 mm.
Hammer laterally compressed, height great-
er than basal diameter (Fig. 44). Aedeagal
apex bluntly pointed, shoulders undevel-
oped. Dorsal keel long, ventral membra-
nous lobes large. Hooks scalloped along in-
ner margins (Figs. 45—47).

Female.—Unknown.

Nymph.—Unknown.

Material.—Guyana: Mallali, Mar, H. S.
Parish, 1 3 (holotype) (MCZ). Potaro Riv-
er, Kaieteur Falls, 1350’, 21-23 Aug 1997,
O. S. Flint, 4 ¢ (USNM).

Diagnosis.—This species is quite similar
to and perhaps synonymous with A. baniva
(Stark 1995). The subtle differences in ae-
deagal structure include a small apical
notch, decurved apex and low dorsal keel
in A. baniva. The holotype of A. phantoma
has a blunt aedeagal apex, straight in lateral
aspect and a more pronounced dorsal Keel
(Figs. 45-47). Anacroneuria makushi rep-
resents another member of this complex
with a more robust aedeagal apex and a
transverse arcuate keel (Figs. 29-31).

Comments.—The type locality above

83

Linden on the Demerara River is one of the
H. S. Parish sites of 1912—13 (Adams, pers.
comm.).

Anacroneuria pictipes Klapalek

Anacroneuria pictipes Klapalek, 1923:21.
Holotype 2, Haut-Carsevenne, Guyana.

Adult habitus.—(Modified from Klapa-
lek, 1923) Body ochre yellow, pronotum
darker brown along the posterior margins
and near lateral margins (Fig. 48). Wing
membrane and veins brown; R dark brown,
costal border pale (Fig. 53).

Male.—Forewing length 10 mm. Ham-
mer laterally compressed at tip, height
greater than basal diameter (Fig. 49). Ae-
deagal apex truncate to emarginate, arising
from low broadly rounded shoulders; dorsal
keel triangular, hooks slender (Figs. 50—52).

Female.—Forewing length 11 mm. Su-
bgenital plate weakly four-lobed. Median
field of sternum nine weakly sclerotized
and sparsely hirsute. Posterior margin of
nine without transverse sclerite (Fig. 55).

Nymph.—Unknown.

Material.—Guyana [French Guiana ?]:
Haut-Carsevenne, 1878, FE Geay, 1 2 (ho-
lotype) (NMH). Guyana: Kanuku Moun-
tains, Moca Moca River, 29 Apr 1995, O.
S. Flint, 2 ¢6 (USNM). Paramaketoi Village,
2350', 24-25 Aug 1997, O. S. Flint, 3 ¢
(USNM).

Comments.—This association of males
with the holotype female is based on the
distinctive wing pigmentation pattern. Fe-
males collected with the males at Moca
Moca River represent another species, de-
scribed below as “‘GU-1” which lacks this
wing pattern.

Anacroneuria timote, new species

Adult habitus.—Head yellow with dif-
fuse brown area forward of ocelli and M-
line; lappets brown. Broad median pronotal
stripe pale, irregular midlateral stripes
brown, lateral margins pale (Fig. 56). Wing
membrane transparent, C and Sc veins pale,
R vein dark brown.

84 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

'

ee
ie 2
Fin
14
ua
Paik
By

’
“yee?
se
Pd;
tyr ee
‘
.

Figs. 43-47. A. phantoma structures. 43. Head and pronotum. 44. Male sternum 9. 45. Aedeagus, ventral.
46. Aedeagus, lateral. 47. Aedeagus, dorsal. Scales: 0.6 mm (43), 0.3 mm (44), 0.15 mm (45-47).

VOLUME 112, NUMBER 1 85

ef h=

<<

WAKER

Figs. 48-53. A. pictipes structures. 48. Head and pronotum. 49. Male sternum 9.50. Aedeagus, ventral. 51.
Aedeagus, lateral. 52. Aedeagus, dorsal. 53. Forewing. Scales: 1.2 mm (53), 0.6 mm (48), 0.3 mm (49), 0.15
mm (50-52).

86

iY
wt) =e
ets

Figs. 54-55. Anacroneuri

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

40
tec as a a e\*
aa eee
LER Nee OF CE
et gee eee is eo ew pce 8
oe eS Be ee eo ee ee “it
Oe ee ee ee ge ae
em Oia ir iN Nie 6 ee hy po ae
= ee ee a , . Vit She! Wiese sad oe ee fe fe
ie = om = Se 5 E reece! - — ae a;
a oe Ele) ee ee
ees | ree dee Be x NA oes Ee (ee
* hi. Ag if - 2 [Lo & =e C3 [ ‘e ya is “a BS
oye s 3 de : me [# sale BY t ee ay A
iN ae ea Lem va a ae
eye l- | i lg ea [ [ .- Sn iee is ce
z.. -4 eas : a a ere ea
eat ; i 4 ue! Aqyeis

55

a female sterna 8 and 9. 54. A. paria. 55. A. pictipes. Scale: 0.3 mm.

VOLUME 112, NUMBER 1

t
ae

SY Da thlad ak

‘

Tikereys

Figs. 56-60. A. timote structures. 56. Head and pronotum. 57. Male sternum 9. 58. Aedeagus, ventral
Aedeagus, lateral. 60. Aedeagus, dorsal. Scales: 0.6 mm (56), 0.3 mm (57), 0.15 mm (58-60).

87

=, ee

88 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Male.—Forewing length 17 mm. Ham-
mer thimble shaped, height less than basal
diameter (Fig. 57). Aedeagal apex simple,
scoop shaped with broadly rounded tip;
ventral aspect with an inconspicuous pair of
membranous lobes; dorsal keel well devel-
oped, hooks slender (Figs. 58—60).

Female.—Unknown.

Nymph.—Unknown.

Etymology.—The name honors the Tim-
ote people of Venezuela and is used as a
noun in apposition.

Types.—Holotype 6 (USNM) and 4 ¢
paratypes (UCV, UMSP) from Venezuela,
Tachira, Quebrada Los Mirtos, 8 km S El
Cobre, 2400 m, 22 Apr 1995, R. Holzen-
thal, C. Cressa, Gutic.

Diagnosis.—The head and pronotal pat-
tern, size, and general aedeagal structure of
this species is similar to A. shamatari (Stark
1995). The most conspicuous difference in
aedeagal structure involves the membra-
nous area between the hooks. In A. sha-
matari this area terminates well short of the
aedeagal base whereas in A. ftimote it ex-
tends almost to the aedeagal base (Fig. 58).

Anacroneuria wapishana, new species

Adult habitus.—Head dark brown except
for anterior margin and callosities. Prono-
tum dark brown except for V-shaped me-
dian band and small anterolateral areas
(Fig. 61). Fore and mid femora and tibiae
dark brown; hind femora yellow in basal
two thirds, dark brown apically. Wing
membrane brown except for transparent
window at cord; veins dark brown except
for pale costal area; wing tips dark brown.

Male.—Unknown.

Female.—Forewing length 11 mm. Su-
bgenital plate four lobed; outer lobes small,
notches shallow; median notch deep. Mesal
sclerite of sternum 9 with short broad me-
dian field of fine setae, lateral fields with
long thick bristles; transverse posterior
sclerite indistinct (Fig. 62).

Nymph.—Unknown.

Etymology.—The name honors the Wap-

ishana people of Guyana and is used as a
noun in apposition.

Types.—Holotype ¢° from Guyana, Po-
taro River, Kaieteur Falls, 1350’, 21—23
Aug 1997, O. S. Flint (USNM).

Diagnosis.—The exceptional color pat-
tern of the holotype is the basis for naming
this species from a female. The species
bears some similarity to A. dourada (Jewett
1960) but is much smaller and has a more
extensive area of dark pigment on the frons.

Unassociated Females
Anacroneuria GU-1

Adult habitus.—Head yellow, without
dark pattern. Pronotum with irregular
brown midlateral bands and a broad yellow
median band. Wing membrane transparent,
veins pale.

Female.—Forewing length 11 mm. Su-
bgenital plate with four subequal lobes. Me-
sal sclerite of sternum 9 with broad median
field of short, fine setae, lateral field with
longer thick bristles; posterior margin with
a narrow U-shaped median sclerite (Fig.
65).

Material.—Guyana: Kanuku Mountains,
Moco Moco River, 29 Apr 1995, 4 2, O.
S. Flint (USNM). Kanuku Mountains,
Kumu River and falls, 28—30 Apr 1995, 1
2, O. S. Flint (USNM).

Anacroneuria GU-2

Adult habitus.—Head yellow brown, lap-
pets pale. Pronotum with diffuse brown
midlateral bands. Wing membrane transpar-
ent, veins pale.

Female.—Forewing length 11 mm. Su-
bgenital plate four lobed; lateral lobes wide
and offset from narrow median lobes by
shallow acute notches. Mesal sclerite of
sternum 9 with median field covered with
fine short setae which form a sagittate patch
separated from lateral patches covered with
longer bristles; posterior sclerite absent
(Fig. 66).

Material.—Guyana: Dubulay Ranch, Ar-

VOLUME 112, NUMBER 1

89

2 site Ws

SY me Lie NES eT
\ XY ai; Cee ley (:

eo
on

Ate 6

eae
tet

Figs. 61-64. Anacroneuria female structures. 61. A. wapishana head and pronotum. 62. A. wapishana sterna
8 and 9. 63. A. llana head and pronotum. 64. A. /lana sterna 8 and 9. Scales: 0.6 mm (61, 63), 0.3 mm (62,

64).

amatani Creek, 15—18 Apr 1995, 2 2, 0.S.
Flint (USNM).

Anacroneuria GU-3

Adult habitus.—Head yellow with dif-
fuse brown pigment forward of ocelli; lap-
pets brown. Pronotum with irregular diffuse

brown spots scattered over midlateral area
(Fig. 67). Tibiae banded, basal and apical

bands brown, median band pale. Wing
membrane transparent except for pale
brown tips which accent an obscure win-
dow at the cord; veins brown, R darker, cos-
tal area pale.

Female.—Forewing length 10 mm. Su-
bgenital plate four lobed; outer lobes small,
notches shallow, median notch deep and U-
shaped. Median sclerite of sternum 9 with

90

Aca CE:
Ba & a “

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

He

Ric f
fe. af eee =| seas od . . al piltes lh A | x ‘ S IEA 2
spent laches Ge eh CEL G
Da MRR PAL ey nupeat a
. « ‘ a ; “| ats AVE Ds :
de a den UAE A a

‘ rao ear fone fl ek
Oe eae re we t i ie
: i . a aN ihe ° oa . = \ : a i “i | it. IM it lak fe 4 “|

-

CEE ETA. th ue CalEe, lease | eC

66

Figs. 65-66. Anacroneuria female sterna 8 and 9. 65. Anacroneuria GU-1. 66. Anacroneuria GU-2. Scale

0.3 mm.

VOLUME 112, NUMBER 1

.
se
Salt
‘
“«
ene ane
'

oe fs

iach hie
Ao

1
at 2
‘ : a
Bs

91

ri? OL
WHS
OZ

Figs. 67-68. Anacroneuria GU-3 female structures. 67. Head and pronotum. 68. Sterna 8 and 9. Scales: 0.6

(67), 0.3 (68).

long narrow stem; stem and mesal field
with short, fine setae, lateral areas with
thicker setae. Transverse sclerite sinuate
(Fig. 68).

Material.—Guyana: Mazaruni-Potaro
District, Takutu Mountains, 20 Dec 1983, P
J. Spangler, 1 2 (USNM). Same location,
12 Dec 1983, P. J. Spangler, R. A. Faitoute,
P. D. Perkins, 1 2 (USNM).

Provisional Key for Male Anacroneuria
from Venezuela, Guyana, and Suriname
(Modified from Stark 1995)

1. Wings banded in amber and dark brown

Wings variable, but without bands... 3
2. Dark pigment separated into basal, me-
dian and apical bands ..... A. bifasciata
Dark pigment forming a narrow lon-
gitudinal band from base to cord (Fig.

LO A rae ee A. paria
3. Forewing length greater than 13.5 mm
MR DS eid ehh! etait, wan wc od 8 +
Forewing length less than13 mm .._ 11
4. Hammer low, scarcely elevated above
LEAD TT de eee A. chorrera
Hammer length equal to apical diam-
eee eects Clee urease xe ia ce ss 5

5. Hammer apex quadrate; aedeagal hook
apices foothke ty. 0.010% A. cuadrada

10.

11.

12.

Hammer apex circular; aedeagal hooks

without Tootlike apices’ < ....-..2.<,.....: 6
Dorsum of aedeagus with a transverse,
subapical arcuate lobe ..... A. arcuata
Dorsum of aedeagus without arcuate
LODE: 4 oc ERE ee. bare phipooene 7
Hammer tiny; aedeagal apex notched
TUES Sowede eee an aan: & 2B A. muesca
Hammer thimble shaped; aedeagal
SDEX TOMNBEOA Gs 28s oc ebias 5 3 tes 8

Aedeagal apex strongly trilobed; pale
mesal pronotal band not extending lat-
erally beyond ocelli A. fenestrata
Aedeagal apex simple; pale mesal
pronotal stripe wide, extending later-

OR he: a

allysbeyend ‘acelliget\. caus wants alsin 9
Aedeagal apex abruptly narrowed at
SOUTER G Sec ee) a ee ee eee 10
Aedeagal apex gradually tapered ..

ee tents $21 ..0F 2b shiw ocak A. paleta

Membranous area between hooks ex-
tends almost to aedeagal base (Fig. 58)
A. timote

i 21S eeu tere)» Se fa le ef we. wae iste le ye fe \ele

as @ Seuss en Sete ae = tee setae & me

Medeacal apex. simple). :..20. 0.4... ...% 12

Aedeagal apex multilobed ........ 25
Aedeagal hooks straight, daggerlike . .

PMOOIS:: U2 SARDINES Aah. es A. cruza
Aedeagal hooks curved, scythelike... 13

92

13.

14.

ix

16.

U7:

18.

19.

20.

i

yap Lp

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Apex of aedeagal hooks fingerlike, ae-
deagal apex surmounting a distinct
neck .hs 2535 <3 A. digitata
Apex of aedeagal hooks not fingerlike,
subapical area of aedeagus without
HECK . 4 0c-s Lee ee 14
Ventral aspect of aedeagus with a con-
spicuous pair of membranous lobes .. 15
Ventral aspect of aedeagus with mem-
branous lobes absent or inconspicuous
Bet AR eR ek Tee Meg 1 ei Bet ae 22
Dorsal keel formed by a pair of widely
spaced ridges (Fig. 16); head with a
conspicuous dark area at ocelli (Fig.
Ne a, Se eee ks FCA E eo oa. 16
Ridges of dorsal keel closely spaced
(Fig. 31); head with diffuse or pale

ocellar pisment (Bie627) «2 bh: 17
Keel arms divergent anteriorly (Fig.
16); occipital area pale (Fig. 12) ....
Ee eso sek Se Btn SN ty RAE A. chaima
Keel arms convergent anteriorly (Fig.
42); occipital area dark (Fig. 38) ....
op ateha® Dnt oehee ite, SORES EE, OB A. perija
Inner margins of aedeagal hooks scal-
loped: (Pia), 29) Wet ots, eke 18
Inner margins of aedeagal hooks
smoothly curved (Fig. 24) ........ 20
Aedeagus with a dorsomesal arcuate
keels (Pia, SO ease 5 ahi: A. makushi
Aedeagus without dorsomesal arcuate
Keehn 5 ee ee 19
Aedeagal apex notched, decurved in
lateral aspees Hi hse 2 Tory A. baniva

Aedeagal apex rounded, straight in lat-
eral aspect (Figs. 45—46) A. phantoma
Membranous ventral aedeagal lobes
wider than long (Fig. 24); submarginal
pronotal bands narrow (Fig. 22) .....
Membranous ventral aedeagal lobes
longer than wide (Fig. 19); submargin-
al pronotal bands wide (Fig. 17) ... 21
Aedeagal apex truncate (Fig. 19); ven-
tral membranous aedeagal lobes small
(Fig. 19) A. claudiae
Aedeagal apex pointed (Fig. 9); ventral
membranous aedeagal lobes large (Fig.
2) nO >: te Met 9) e. A. arawak
Aedeagal apex gradually narrowed to
a pOmite ng) . ities Jose Thee: A. bari
Aedeagal apex truncate or broadly
rounded

Ke

24.

29:

26.

ake

28.

Jeg)

30.

at;

a2.

Aedeagus with a small ventral mem-
branous lobe A. chiquita
Aedeagus without ventral membranous
lobe og oon 3 Spee os Sa 24
Projecting apex of aedeagus subequal
to shoulder A. llana
Projecting apex of aedeagus about
twice as long as shoulder .... A. blanca
Aedeagal apex with a dorsolateral pair
of small hornlike processes ....... 26
Aedeagal apex without hornlike pro-
cesses
Hornlike aedeagal processes acute and
conspicuous in ventral aspect

2,50) oe * @ -e (6 = 6 ‘« 6 —=

Rieawe. ©. (ecm esa tain aie os 6

Hornlike aedeagal processes rounded
and inconspicuous in ventral aspect ..

A. aroucana
Dorsal aedeagal keel V-shaped (Fig.
92) 2 3. dh se ee 28
Dorsal aedeagal keel not V-shaped... 29
R and basal half of Sc vein covered
with dark brown pigment (Fig. 53)

A. pictipes
R and basal half of Sc without con-
strasting pigment band A. achagua
Ventral aspect of aedeagus with a large
membranous lobe A. pequena
Ventral aspect of aedeagus without
membranous lobe
Lateral aedeagal lobes scarcely pro-
jecting A. menuda
Lateral aedeagal lobes distinctly pro-
jecting
Lateral aedeagal lobes about as wide

as median lobe... . i. -" 7 ae eee 32
Lateral aedeagal lobes about half as
wide as median lobe ...... A. sp VZ-10
Wings with a large transparent spot be-
yond cord, and an irregular transparent
costal band A. vistosa
Wings rather uniformly pigmented ...

oso 6 o a 6: ane! 6 @@ « @) 6 2. a) w one, 6) aye

eo ae 6 te 2 0, a eek es) mie, (6) Le) fa ie eo eee
Pe ee

ee @ © © © tes © © & @ 0s) » © © 06 © la ia ae

Acknowledgments

I thank R. Holzenthal, University of Min-

nesota, and O. S. Flint and N. Adams, Unit-
ed States National Museum, for the loan of
specimens. I also thank P. Zwick and cu-
rators at the California Academy of Science
and the Museum of Comparative Zoology

VOLUME 112, NUMBER 1

for their help in arranging loans of type ma-
terial.

Literature Cited

Banks, N. 1914. New neuropteroid insects, native and
exotic.—Proceedings of the Academy of Natu-
ral Science of Philadelphia 66:608—632.

Jewett, S. G. 1960. Two new species of Anacroneuria
(Plecoptera) from Goids, Brazil—bLos Angeles
County Museum Contributions in Science 36:
1-4.

Kimmins, D. E. 1948. A new species of Anacroneuria
(Plecoptera, Perlidae) from Trinidad.—The Pro-
ceedings of the Royal Entomological Society of
London, Series B 17:105—106.

93

Klapalek, F 1923. Plécoptéres nouveaux.—Annales de
la Sociéte Entomologique de Belgique 63:21-
29.

Pictet, E J. 1841. Histoire naturelle générale et parti-
culiére des insectes Névroptéres. Famille des
Perlides. 1. Partie: 1-423.

Stark, B. P. 1994. Anacroneuria from Trinidad and To-
bago (Plecoptera: Perlidae).—Aquatic Insects
16:171-175.

Stark, B. P. 1995. New species and records of Anacro-
neuria (Klapalek) from Venezuela (Insecta, Ple-
coptera, Perlidae).—Spixiana 18:211—249.

Zwick, P. 1972. Die Plecopteren Pictets und Burmeis-
ters, mit Angaben tiber weitere Arten (Insec-
ta).—Revue Suisse de Zoologie 78:1123-—1194.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(1):94—96. 1999.

A new species of Siamosquilla from Indonesia
(Crustacea: Stomatopoda: Protosquillidae)

Mark V. Erdmann and Raymond B. Manning

(MVE) Department of Integrative Biology, University of California at Berkeley, Berkeley,
California 94720, U.S.A.; (RBM) Department of Invertebrate Zoology, National Museum of
Natural History, Smithsonian Institution, Washington, D.C. 20560-0163, U.S.A.

Abstract.—Siamosquilla sexava, the second species of the genus, is de-
scribed from Indonesia. It can be distinguished from the type species, S. hyl-
lebergi Naiyanetr, from Thailand, by the shorter median rostral spine, the larger
ocular scales, the much broader telson, and differences in ornamentation of the
fused sixth abdominal somite and telson.

Among the stomatopods collected by one
of us (M. V. E.) during a six-year field study
in Indonesia was a minute protosquillid
which proved to be the second known spe-
cies of Siamosquilla Naiyanetr, 1989. It is
described below.

Abbreviations used in the account in-
clude: LON, Lembaga Oseanologi Nasional
(National Institute of Oceanoraphy), Jakar-
ta, Indonesia; TL, total length, measured on
the midline; USNM, National Museum of
Natural History, Smithsonian Institution,
Washington, D.C., U.S.A.

The holotype and some paratypes have
been deposited in the USNM; two paraty-
pes are in the LON.

Family Protosquillidae Manning, 1980
Siamosquilla Naiyanetr, 1989
Siamosquilla sexava, new species
Fig. 1

Material examined.—Indonesia: Moro-
maho, Tukang Besi: 12, TL 18 mm (ho-
lotype, USNM 260927), 222, TL 16—17
mm (paratypes, LON).—Gili Meno, Lom-
bok: 16, TL 17 mm, 12, TL 18 mm (par-
atypes, USNM 260928).—Melanguane,
Sangihe-Talaud: 12, TL 17 mm (paratype,
USNM 260929).—Taupun, Togian Islands:
1d, TL 13 mm (paratype, USNM 260930).

Diagnosis.—Size very small, TL less

than 20 mm in adults. Cornea broadened,
set obliquely on stalk. Ocular scales well
developed, produced into triangular lobes
laterally, extending nearly to lateral rostral
Spine in adults.

Rostral plate sharply trispinous, median
spine distinctly upturned distally, extending
to base of corneas; ventral projection of ros-
tral plate large, obtusely rounded ventrally.
Lateral rostral spines extremely long, slen-
der and recurved, length nearly two-thirds
that of median spine. Basal part of rostral
plate very thin. Anterior margins of lateral
plates of carapace concave, anterolateral an-
gles strongly produced to a sharp point, ex-
tending anteriorly to base of rostral plate.

Mandibular palp absent. Five epipods
present.

Raptorial claw with inflated part of outer
margin of dactylus notched. Propodus with
single movable spine proximally on inner
margin.

Anterior 4 abdominal somites smooth,
unarmed, not carinate dorsally. Fifth ab-
dominal somite smooth medially, with sin-
gle low longitudinal carina laterally above
lateral margin, separated from margin by a
groove, armed with posterolateral spine.
Sixth abdominal somite entirely fused with
telson in adults, dorsal surface rough, with
shallow, irregularly curved grooves, lacking
carinae entirely.

VOLUME 112, NUMBER 1

95

Fig. 1.

Siamosquilla sexava, new species, female holotype, TL 18 mm. a, Anterior part of body, dorsal view

(apex of left ocular scale damaged); b, Rostral plate, lateral view; c, Fused sixth abdominal somite and telson

and left uropod, dorsal view. Scales = 1 mm.

Telson much wider than long, dorsal sur-
face rough, slightly inflated, with 3 very in-
distinct bosses. Median boss outlined by a
shallow groove in dorsal surface of telson;
boss rounded anteriorly, converging poste-
riorly; outline of submedian bosses even
less distinct, extending posteriorly almost to
posterior margin of telson. Median fissure
completely fused, no longer visible. Three
pairs of marginal teeth, submedians with
moveable apices arising submarginally un-
der submedian marginal projections. Inter-
mediate teeth very low and rounded, almost
indistinguishable in larger specimens. Lat-
eral teeth more distinctly produced, with
rounded apices. Separation between three
pairs of marginal telson teeth shallow and
rounded. Submedian teeth with 12—14 den-
ticles on either side of midline, increasing
noticeably in length laterally, intermediate
teeth each with 2 fixed mesial denticles, lat-
erals each with 1 denticle. Lateral margins
of telson straight, rounded distally.

Uropods stout, proximal segment of ex-

opod with 9 short movable spines laterally,
distalmost extending beyond midlength of
distal segment, with fixed distal spine ven-
trally. Inner margin of uropodal exopod se-
tose; uropodal endopod with normal com-
plement of setae. Inner spine of basal pro-
longation of uropod much shorter than out-
er.

Size.—Males (n = 2), TL 13-17 mm. Fe-
males (n = 5), TL 13-18 mm.

Remarks.—This small species has a very
distinctive rostral plate, with long, recurved
lateral spines, and a telson that is most re-
markable for its lack of distinguishing fea-
tures; the sixth abdominal somite is fused
to the telson, which has almost indiscern-
ible bosses and a fused median fissure. Sia-
mosquilla sexava differs from S. hyllebergi
in numerous features: the median rostral
spine is shorter, extending to, rather than
beyond, the cornea and the lateral rostral
spines are longer; the ocular scales are
broader; in adults the anterolateral angles of
the carapace are sharper and more pro-

96 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

nounced; the dorsal sculpture of the fused
sixth abdominal somite and telson is much
more elaborate; and the telson is much
broader.

The fixed projections on the posterior
margin of the telson are much more pro-
nounced in S. hyllebergi than in S. sexava,
and the submedian projections in the former
species probably obscure the submarginal,
movable submedian teeth of the telson,
which are clearly visible in S. sexava.

In S. sexava, the size of the ocular scales
increases allometrically, and in very small
specimens of S. sexava the scales resemble
those of larger S. hyllebergii; they increase
in width with increasing TL.

Distribution.—Known only from eastern
Indonesia, where it is relatively widespread.
Recorded from five regions: Lombok, San-
gihe-Talaud, Togian Islands, and Tukang
Besi Archipelago.

Etymology.—The species name is from
the word sexava, which means “‘mantis”’ in
Bahasa Talaud, the local language in one of
the collection localities (Melanguane, San-
gihe-Talaud). The local fishermen, school
teachers and children there were most help-
ful in assisting with collecting, and were
fully cognizant of the differences between
stomatopods and other decapod shrimp,
comparing stomatopods to the “‘sexava”’
without any solicitation. Their assistance is
most appreciated.

Acknowledgements

We thank the Indonesian Institute of Sci-
ences for sponsoring Erdmann’s project,
and Kalam Sebayang and Ibu Dewi Soen-
arijadi for their assistance with research vi-
sas. Dr. Mohammad Kasim Moosa was the
Indonesian sponsor of this research, and
Erdmann gratefully acknowledges his men-
torship on stomatopod morphology and his
continual support of this project. We also
thank Dr. Sukarno, Dr. Suharsono, and Dr.
Anugerah Nontji of the Indonesian Institute
of Sciences.

Funding for Erdmann’s research was pro-
vided by grants from the UC Pacific Rim
Research Program, the NSF International
DDIG (#9503060 and #9704616) Program
and a visiting scientist appointment from
the Smithsonian Institution. Lilly King
Manning prepared the figure. This is con-
tribution no. 467 from the Smithsonian Ma-
rine Station at Fort Pierce, Florida; the sup-
port of that program for Manning’s studies
of stomatopod systematics is gratefully ac-
knowledged. We gratefully acknowledge
the reviews of Shane Ahyong, David K.
Camp, and Roy K. Kropp as well as the
cooperation of Rafael Lemaitre.

Literature Cited

Naiyanetr, P. 1989. Siamosquilla hyllebergi, a new ge-
nus and new species of stomatopod from Thai-
land. Jn E. A. Ferrero, ed., Biology of stomato-
pods. Selected symposia and monographs,
U.Z.1., 3:281—284. Muchi, Modena.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(1):97-105. 1999.

The taxonomic status and zoogeography of Cambarus bartonii
carinirostris Hay, 1914 (Crustacea: Decapoda: Cambaridae)

Roger E Thoma and Raymond EF Jezerinact

(RFT) Ohio Environmental Protection Agency, 2110 Aurora Rd., Twinsburg, Ohio 44087, U.S.A.

Abstract.—Historically, Cambarus bartonii carinirostris was considered a
subspecies of Cambarus bartonii. Recent studies indicate that C. b. cariniros-
tris is indistinguishable from the nominate species and should be relegated to
synonymy with C. bartonii. In other studies this assignment is not accepted
and subspecific status has been maintained for C. b. carinirostris when re-
porting on crayfish closely related to C. bartonii. Work in the area of northern
Appalachian crayfishes has made it apparent to us that C. b. carinirostris should
be elevated to full species status based on its unique meristic and morphometric

characteristics.

For nearly two decades we have studied
the crayfishes of the northern Appalachians.
Of the many questions associated with this
area, the proper identity and distribution of
C. b. bartonii (Fabricius 1798) has been and
continues to be a principal concern. Ort-
mann’s (1905, 1906, 1931) studies remain
the foundation on which the distribution
and systematics of northern Appalachian
crayfish are presently understood. After
Ortmann’s death in 1933, Horton H. Hobbs,
Jr., shifted the center of crayfish systematic
studies to the southern Appalachians (and
other southern areas) and the identity of C.
b. bartonii and its subspecies remained un-
resolved. In the early 1970s we became in-
terested in the ecology of crayfish in our
home state, Ohio. We were immediately
confronted with the problem of C. b. bar-
tonii and the subspecies associated with it,
since both C. b. carinirostris (Hay 1914)
and C. b. cavatus (Hay 1902) are reported
from Ohio (Hobbs 1974, 1989; Thoma &
Jezerinac 1982). With the unresolved taxo-
nomic problems and a domination by mem-
bers of the subgenus Cambarus in Ohio, it
became apparent that a taxonomic study

+ Deceased, 21 April 1996.

was needed before the study of crayfish
ecology could commence.

Materials and Methods

Specimens examined came from the Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington, D.C.
(USNM), and the Ohio State University
Museum of Biological Diversity, Colum-
bus, Ohio (OSU-MBD). The paratypes,
housed at the Museum of Comparative Zo-
ology (MCZ) were not examined. Field col-
lections (now house at OSU-MBD) were
made using a 1.3 X 2 m seine or by hand.
A total of 95 specimens of C. carinirostris
(33 Form I males, 27 Form II males, and
35 females) and 191 specimens of C. b.
bartonii (79 Form I males, 43 Form II
males, and 69 females) were measured for
this study. Measurements were made to the
nearest 0.1 mm using a vernier caliper and
followed Hobbs (1981) and Jezerinac
(1985). Measurements of regenerated body
parts were avoided. Analysis was per-
formed using SYSTAT 5.2.1. Principal
component analysis used only Form I male
specimens.

98 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Cambarus (Cambarus) carinirostris Hay,
new status

Cambarus bartonii carinirostris Hay, 1914:
384.

Cambarus bartonii montanus.—Faxon,
1914:386 (in part)—Newcombe, 1929:
286 (in part).

Cambarus montanus montanus.—Ortmann,
1931:106 (in part).

Cambarus (Cambarus) bartoni cariniros-
tris.—Ortmann, 1931:107.

Cambarus (Cambarus) bartonii cariniros-
tris.—Hobbs 1969: 109, fig. 19m; 1974:
11, fig. 24; 1989: 13 fig. 30.—Thoma,
1982:875.—Thoma & Jezerinac, 1982:
136.—Jezerinac, 1983: 4.—Jezerinac &
Thoma, 1984: 120 figs. 8—9.—Jezerinac
& Stocker, 1989: 2.—Jezerinac & Stock-
er, 1990: 1.—Jezerinac, Stocker & Tarter,
1995: 76-83, fig. 35-38.

Diagnosis.—Body pigmented. Carapace
subcylindrical, slightly flattened dorsoven-
trally. Eyes slightly reduced. Rostrum with
parallel or slightly concave margins, mar-
gins thickened, lacking marginal spines or
tubercles, rostrum curved abruptly cephali-
cally and terminating in upturned corneous
tubercle; frequently with a median carina.
Areola 3.4—12.0 times longer than wide
(median = 5.7), comprising 25.2—40.4% of
total length of carapace (median = 37.9%),
bearing 3 to 4 punctations across narrowest
part. Cervical spine absent or reduced to
blunt tubercle. Cervical grove uninterrupt-
ed. Suborbital angle acute. Postorbital ridge
lacking cephalic spine or tubercle. Bran-
chiostegal spine reduced to small knob. An-
tennal scale approximately 1.5 times as
long as broad, with mesial and lateral mar-
gins subparallel near and at midlength; dis-
tomesial margin strongly sloping. Basio-
podite of antenna lacking spine. Ischiopod-
ite of antenna with blunt spine. Chela
smooth, robust, length 79.6—111.9% of total
carapace length (median = 94.0%) in Form
I males (71.8-88.4%, median = 78.8% in
females), bering 1 row of 5 to 8 (median =
7.0) adpressed squamous tubercles along

mesial margin of palm and 5 to 7 puncta-
tions dorsal to such tubercles (occasionally
distal 1 to 4 punctations replaced by tuber-
cles). Width of gape of fingers of Form I
male 17.4—46.3% of palm length (median
= 27.2%), less so in Form II males (median
= 16.3%) and females (median = 17.2%).
Lateral margin of fixed finger weakly cos-
tate; moderately developed dorsomedial
ridges on both fingers flanked by parallel
rows of punctations; fixed finger impressed
at dorsal and ventral lateral bases; dactyl
0.9—2.5 (median = 2.1) times longer than
mesial margin of palm; palm width 41.5—
51.1% (median = 46.1%) of chela length;
third or fourth tubercle of mesial margin of
fixed finger enlarged; 1—2 tubercles usually
present on mid-subpalmar surface; never
with elongated setae at base of fixed finger.
Dorsomesial margin of carpus of chela with
1 distal spine and 1 proximal blunt tubercle;
ventral surface with 1 or 2 conical tubercles
at distal margin. Ventrolateral ridge of mer-
us usually with 2—3 spines. Hook only on
ischium of third pereiopod of male. Basal
boss on coxa of fourth pereiopod well de-
veloped. First pleopods of Form I male con-
tiguous at base, with 2 short terminal ele-
ments bent at approximately 90° to main
shaft; corneous central projection truncated
distally, bearing subapical notch; mesial
process inflated, tapering distally; central
projection of Form II male pleopods non-
corneous, club-shaped. Females with an-
nulus ventralis slightly embedded in ster-
num, asymmetrical, subrhomboid, slightly
movable, lacking cephalolateral promi-
nence; first through fifth pleopods similar
in shape.

Color notes.—For the most part, this spe-
cies is dorsally a uniform brown ranging
from chestnut to tan with ventral surfaces
fading to cream. Some populations have a
greenish hue. The thickened rostral margins
and postorbital ridges are a brownish red
and the larger tubercles and spines orange.
No banding or striping evident.

Types.—‘““Type” and paratypes USNM
23962 (1 Form I male, 7 Form II male, 15

VOLUME 112, NUMBER 1

female); paratypes, MCZ 7399 (1 Form I
male, 1 Form II male, 1 female). The Form
I male housed at the USNM is herein des-
ignated the lectotype of the species.

Type locality—lHay (1914:385) stated
““Gandy Creek, Oceola, Randolph Co., W.
Va.”’ in his original description. A visit to
this area and discussions with local resi-
dents indicated Oceola (38°42'50’N,
79°38'00"W) was the location of a now
non-extant school house at the site of a his-
toric lumber camp. The old school house
was located near the Sinks of Gandy. We
made a topotype collection from Gandy
Creek upstream of County Road 40, just
west of County Road 29/1 (38°43'22’N,
79°37'38"W). This site is the first road
crossing downstream of the mouth of the
Sinks of Gandy.

Range.—Found throughout the drainages
of the Allegheny and Monongahela rivers
in Pennsylvania, New York and West Vir-
ginia; tributaries of the Ohio River up-
stream of Sunfish Creek in Ohio and Fish
Creek in West Virginia; southern tributaries
of Lake Erie and Lake Ontario from the
Grand River, Ohio, to the Gennesse River,
New York; throughout the Greenbrier River
and tributaries of the New River upstream
of the Greenbrier; upper Elk River, West
Virginia.

Variation.—In the upper reaches of the
Monongahela basin (Cheat and Tygart/
Buckhannon basins) C. carinirostris prob-
ably retains its most plesiomorphic state.
This is the only portion of the range in
which no other stream dwelling forms of
Cambarus are found. Here, C. carinirostris
attains its greatest degree of sculpturing (in-
cluding the carina) and approaches the body
size of members of the subgenera Camba-
rus and Puncticambarus that inhabit larger
rivers. The plesiomorphic appearance is
likely in part due to the larger sizes at-
tained. In the remainder of the range, Cam-
barus (Puncticambarus) robustus Girard,
1852, or Cambarus (C.) sciotensis Rhoades,
1944, occupy the larger mainstem streams
and C. carinirostris is confined to the

99

smaller tributaries. In the Casselman River
of the Youghiogheny River, Pennsylvania,
we have seen specimens with body forms
reminiscent of the subgenus Erebicambarus
Hobbs, 1969, in that they display a more
tubular, sausage-shaped carapace that is less
dorsoventrally compressed. The chelae re-
main decidedly within the range of C. car-
inirostris, though they have the least
amount of gape between the fingers. Spec-
imens from the southernmost extent of the
range display reduced inflation of the ros-
tral margins and lack the 90° angle at the
rostral tip. The rostra in this population
most closely resemble the probable ple-
siomorphic state.

Occasionally, a second row of slightly
produced tubercles can be found on the
palm of the chela. This character does not
exhibit a defined geographical pattern but
appears in some individuals in most collec-
tions. All other populations exhibit the nor-
mal characteristics given in the Diagnosis.

Size.—Mature specimens range from 26
to 48.8 mm (median = 34.9 mm) total car-
apace length (median: Form I males 36.9
mm, Form II males = 31.9 mm, females =
35.4 mm).

Life history notes.—Jezerinac et al.
(1995) reported Form I males from late
April through early September and oviger-
ous females from July through mid August.
This study found Form I males as late as
October and ovigerous females in the re-
ported range of dates. No information exists
on longevity, growth rates, thermal prefer-
ences or dietary habits.

Habitat and ecology.—The normal hab-
itat occupied by C. carinirostris is pools
and riffles of high gradient first and second
order streams. Populations can be found in
intermittent streams but the abundance is
greatly reduced. Some burrowing occurs,
mostly sub-boulder, in mid-stream or on the
edges of streams. This species is a second-
ary burrower. Cambarus carinirostris is ca-
pable of expanding its niche in the absence
of other species of Cambarus that are pri-
mary burrowers or mainstem inhabitants.

100 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Often caves, if present, yield specimens,
usually in the vicinity of the mouth. This
species is sensitive to excess silt, bed load
sediments, nutrient enrichment, acid mine
impacts, and habitat alterations that reduce
cobble and boulder abundance.

Taxonomic status.—Numerous authors
have commented on the distribution and
taxonomic status of C. carinirostris. In his
original description Hay (1914: 385) re-
ported the species from Tygart Valley and
Cheat River in Randolph County, West Vir-
ginia, stating that C. carinirostris was “‘...
a well marked subspecies ...” differing
from the nominate species in that “‘... the
carapace is a little more cylindrical, the ros-
trum broader and flatter, and always fur-
nished near the tip with a median longitu-
dinal carina.’’ Faxon (1914: 385), com-
mented that the rostral carina was a rather
elusive character and reported additional re-
cords for this taxon from the Greenbrier
River basin (West Virginia). The next report
on C. carinirostris was Ortmann (1931:
139) in which he retained the subspecific
status for C. carinirostris, and stated C. car-
inirostris differs from C. b. bartonii only in
the presence of the carina. After Ortmann’s
death, no further comments were published
on the taxonomy of C. carinirostris until
Hobbs (1972: 111) commented that it prob-
ably should not be recognized. Bouchard
(1976: 588) recognized no subspecies of C.
bartonii, reporting that no characters sepa-
rate the currently recognized subspecies. He
also noted the instability of the rostral char-
acters. Thoma (1982), Thoma & Jezerinac
(1982), Jezerinac (1983), and Jezerinac et
al. (1995) continued to use subspecific sta-
tus for C. carinirostris.

We agree with Faxon (1914), Ortmann
(1931), and Bouchard (1976) that the carina
of C. carinirostris is an elusive trait, fre-
quently absent or greatly reduced. In our
study of the species we found the popula-
tion in the vicinity of the type locality ex-
hibited the carina most strongly and fre-
quently, and in this respect one could con-
clude that C. carinirostris is a local varia-

®

tion. When we examined the bartonii
complex for other traits, we noted clear dif-
ferences in chela structure between popu-
lations of the Atlantic and Mississippi
drainages in Pennsylvania (especially in
form I males). Thoma & Jezerinac (1982:
137) reported that C. carinirostris could be
distinguished from C. bartonii bartonii (Fa-
bricius, 1798) by the thickened and fre-
quently concave rostral margins; an abrupt-
ly ending rostrum that forms a 90° angle
with the base of the acumen; more strongly
developed postorbital ridges; stronger de-
velopment of the lateral impression and
dorsal ridges of the chela; a slight devel-
opment of a second row of 2 or 3 tubercles
on the mesial margin of the palm; reduced
gape between the dactyl and propodus; and
an enlarged third tubercle on the mesial
margin of the opposable propodus. Contin-
ued work throughout the ranges of the two
forms (and the rest of the range of the sub-
genus) revealed that the combination of
thickened rostral margins, an enlarged third
(or fourth) tubercle on the mesial margin of
the opposable finger of the propodus, mod-
erate development of the lateral impression
and dorsal ridges of the chela, and less than
2 full rows of palmar tubercles is sufficient
to distinguish this species from the nomi-
nate species and all other taxa presently
known in the bartonii complex. The partial
second row of palmar tubercles (not con-
sistently present), 90° angled rostral mar-
gins, strong postorbital ridges, and dactyl
gape are useful in distinguishing this spe-
cies individually from other species of the
bartonii complex. Within the subgenus two
forms (C. b. cavatus Hay, 1902, and C. sci-
otensis Rhoades, 1944) consistently display
the development of a second row of tuber-
cles on the mesial margin of the palm.
Cambarus sciotensis displays a full com-
plement of 5 or 6 tubercles in the second
row. Thoma & Jezerinac (1982) reported
that C. b. cavatus consistently had 3 to 5
tubercles in the second row of palmar tu-
bercles, noninflated rostral margins, and a
narrower areola (the narrower areola may

VOLUME 112, NUMBER 1

relate to the burrowing habits of C. b. ca-
vatus).

Principal component analysis of morpho-
metric data for C. carinirostris and C. b.
bartonii (Fig. 1) illustrates slight differenc-
es in body structure. The gape of the chela
fingers, dactyl width, and areola width are
the strongest loading variables (Table 1).
Areola and dactyl width tend to be wider
and chela gape narrower (Fig. 2) in C. car-
inirostris. No single body proportions are
sufficient to separate the two taxa on a con-
sistent basis. The most reliable character
traits (using Form I males only) for consis-
tently separating the two taxa are develop-
ment of an enlarged third tubercle on the
inner margin of the opposable propodus,
lateral impression of the chela, and dorsal
ridges on the chela fingers of C. cariniros-
tris (Fig. 2). The northern form of C. b.
bartonii (Atlantic drainage from the Poto-
mac basin northward) lacks these chela
characters altogether while the southern
complex of C. b. bartonii lacks the lateral
impression and thickened rostral margins.
Some southern representatives of C. b. bar-
tonii have an enlarged third tubercle on the
opposable dactyl. This is accompanied by
an enlarged first and fourth (occasionally
third) tubercle on the opposable finger of
the propodus, a characteristic lacking in C.
carinirostris.

No intergrade populations have been
found (using the characters we employed)
between C. carinirostris and any other
closely related taxon. For these reasons we
elevate C. carinirostris to full species sta-
tus. Important to note is that the above char-
acteristics are most reliable in Form I
males, preferably of a large size.

Evolution, zoogeography and associated
variation.—Cambarus carinirostris 1s pri-
marily an inhabitant of the still extant por-
tions of the preglacial Pittsburgh River
(Leverett 1902, Tight 1903) and (in part)
the New River basin. This distribution is a
reflection of the evolutionary history of the
species. Cambarus carinirostris was most
likely derived from an ancestral stock of the

101

bartonii complex that inhabited the New
River (Kanawha basin). This stock was in-
tern derived from populations inhabiting
the upper Tennessee River basin, in partic-
ular the Clinch and Powell systems. Hobbs
(1969) concluded that the subgenus Cam-
barus originated in the Tennessee basin and
entered the Kanawha River basin from a
northward migrating member of the “‘extra-
neus group.’ We originally believed the
subgenus Cambarus originated in the Ka-
nawha basin (Jezerinac et al. 1995), but the
recent discovery of a previously unknown
archetypal Cambarus species (a description
of this species and a discussion of its evo-
lutionary significance is in development) in
an isolated portion of what was once the
Tennessee basin and now part of the Cataw-
ba River basin, confirms Hobbs’ conclu-
sion.

Upon entering the New River basin
through interdigitating headwater streams,
probably in the Burk’s Garden area of Vir-
ginia, the ancestral C. carinirostris spread
steadily through the system. Cambarus car-
inirostris probably was the first Cambarus
to enter the upper New River system, thus
having uninhibited dispersal opportunities.
Access was gained to the preglacial Pitts-
burgh River basin in the headwaters of the
Greenbrier River, again before other mem-
bers of the subgenus Cambarus. No further
dispersal occurred until postglacial times
when C. carinirostris followed the retreat-
ing glaciers into the new Laurencian basin,
where it established populations from the
Gennesse River of New York in the east to
the Grand River of Ohio in the west. Dis-
persal on the western edge of the range was
thwarted by the presence of crayfish popu-
lations (presently assigned to C. b. cavatus)
that occupied the preglacial Teays River
system (Jezerinac 1983). Cambarus b. ca-
vatus most likely entered the Teays basin
via the Big Sandy River. The Teays River
population was a stream dwelling species of
the subgenus Cambarus that apparently had
ecological preferences similar to C. carini-
rostris. In the Allegheny Plateau portion of

102 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Factor 3

oe)

A

2 C. carinirostrs @ et
& a @ e%
we,

Factor 2
7 Gape (mm)
6 B -
a)
Ss
: 8 8
oO e
4 e@ e a =
C.b.bartonii Ors C. carinirostris
O
3 a, °
O
a” .
1 O °
6)

10 20 30 40 50 60
Chela Length (mm)

Fig. 1. A, Principal Component Analysis of Cambarus carinirostris (black circles, N = 33) and C. bartonii
bartonii (open circles, N = 79) using Factors 2 and 3 (Table 1), Form I males only; B, Gape of chela fingers
by length of chela for C. carinirostris (black circles, N = 33) and C. b. bartonii (open circles, N = 79), Form
I males only.

VOLUME 112, NUMBER 1

Table 1.—Factor loadings for Principal Component
Analysis of morphometric data of Form I male speci-
mens of C. b. bartonii and C. carinirostris.

Measurement Factor 2 Factor 3
Chela length 0.066 —0.044
Chela width 0.017 0.032
Chela depth 0.030 0.037
Gape of fingers 0.227 (0.326
Palm length 0.039 = O.051
Dactyl length 0.088 —0.042
Dactyl width W095 0.394
Propodus finger length 0.045 —0.078
Propodus finger width O76 0.120
Carapace length* 0.040 0.027
Carapace width 0.016 0.035
Areola length 0.110 0.023
Areola width —0.667 =O: 189

* Measurement includes rostrum.

the New-Kanawha River basin and those
streams draining to the Ohio River down-
stream of the Sardis Col, C. b. cavatus pre-
vailed. In the newly formed Ohio River, the
two species are presently found on their re-

103

spective preglacial sides of the Sardis Col,
with neither having been able to advance
into the other’s range. To the north and to
some degree to the east, the dispersal of C.
carinirostris was inhibited by the presence
of C. b. bartonii. It appears that the post-
glacial dispersal of C. b. bartonii (in the
Atlantic slope drainage) was much more
rapid than that of C. carinirostris, with C.
b. bartonii effectively excluding C. carini-
rostris in those areas. Few opportunities ex-
isted for eastward dispersal of C. cariniros-
tris in the Greenbrier, Monongahela, and
Allegheny river basins because of physical
barriers. Where potential stream piracies
have been identified, no C. carinirostris
could be found in Atlantic drainage
streams. The ecological preferences of C.
carinirostris and C. b. bartonii appear to be
very similar.

Taxonomic distinction.—No other spe-
cies in the subgenus Cambarus possesses
the character combination of thickened ros-

Fig. 2._ A, Dorsal view of pereiopod I chela of Cambarus carinirostris; 1\—enlarged third tubercle on mesial
margin of immovable finger, 2—lateral impression, 3—dorsal ridge; B, Dorsal view of carapace of Cambarus
carinirostris,; 4—rostral carina, 5—thickened rostral margins; C, Dorsal view of pereiopod I chela of Cambarus

b. bartonii.

104 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

tral margins, an enlarged third tubercle on
the mesial margin of the opposable finger
of the propodus, no enlarged tubercles on
the mesial margin of the dactyl, moderate
development of the lateral impression and
dorsal ridges of the chela, and 1 row of pal-
mar tubercles.

Acknowledgments

We would like to thank the many indi-
viduals who helped us collect the crayfish
used for this paper (including those who
went before us). A special posthumous
thanks to Dr. H. H. Hobbs, Jr. and to the
National Museum of Natural History,
Smithsonian Institution, for access to the
crayfish collections. We thank Dr. Joseph EF
Fitzpatrick, Jr., of the University of South
Alabama, Michael Bolton of the Ohio En-
vironmental Protection Agency and two
anonymous reviewers for evaluating the
manuscript.

Literature Cited

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crayfishes of the Cumberland Plateau and Cum-
berland Mountains, Kentucky, Virginia, Tennes-
see, Georgia and Alabama. Part II. The Genera
Fallicambarus and Cambarus. Pp. 585—605 in
J. W. Avault, Jr., ed., Freshwater Crayfish, Lou-
isiana State University Division of Continuing
Education.

Fabricius, J. C. 1798. Supplementum Entomologiae
Systemicae. Hafniae: Proft et Storch, 527 pp.

Faxon, W. 1914. Notes on the crayfish in the United
States National Museum and the Museum of
Comparative Zoology with descriptions of new
species and subspecies to which is appended a
catalogue of the known species and subspe-
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Girard, C. 1852. A review of the North American As-
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Hay, W. P. 1902. Observations on the crustacean fauna
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1914. “‘Cambarus bartonii carinirostris

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Hobbs, H. H., Jr. 1969. On the distribution and phy-
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. 1972. Crayfishes (Astacidae) of North and

Middle America, Identification Manual 9, Biota

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. 1974. A checklist of the North and Middle

American crayfishes (Decapoda: Astacidae and

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. 1981. The crayfishes of Georgia.—Smithson-

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. 1989. An illustrated checklist of the American
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Jezerinac, R. EF 1983. Possible correlations of present
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. 1985. Morphological variations of Cambarus

(Cambarus) bartonii cavatus (Decapoda: Cam-

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Ohio form.—Ohio Journal of Science 85(3):

131-134.

, & G. W. Stocker. 1989. Distribution of the

stream crayfishes of the genus Cambarus (De-

capoda: Cambaridae) in West Virginia.—Ohio

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, & . 1990. Distribution of crayfishes

(Decapoda: Cambaridae) of West Virginia. Part

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Sciences 62(1):7—8.

, & R. E Thoma, 1984. An illustrated key to

the Ohio Cambarus and Fallicambarus (Deca-

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, G. W. Stocker, & D. C. Tarter. 1995. The cray-

VOLUME 112, NUMBER 1

fishes (Decapoda: Cambaridae) of West Virgin-
ia.—Ohio Biological Survey Bulletin New Se-
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(1):106—119. 1999.

Two new species of Aegla Leach (Crustacea:
Decapoda: Anomura: Aeglidae) from southern Chile

Carlos G. Jara and Victor L. Palacios

Instituto de Zoologia “‘Dr. Ernst Kilian’’, Universidad Austral de Chile, Casilla 567,
Valdivia, Chile

Abstract.—Two new species of the genus Aegla Leach from southern Chile
are described. Aegla cholchol, new species, a medium to large sized spinulated
aeglid, has the River Chol-Chol (Cautin Province) as its type locality. It is
probably more related to A. rostrata, from subandean lakes at the Toltén and
Valdivia River basins than to A. bahamondei, from the Tucapel River Basin on
the western slope of the Nahuelbuta Coastal Cordillera. The other, A. hueicol-
lensis, new species, a small to medium sized non-spinulated aeglid, has the
River Pichihueicolla (Valdivia Province), on the western slope of the Pelada
Coastal Cordillera, as its type-locality. A. hueicollensis is similar to A. abtao
with which it shares more morphological attributes than with other non-spi-
nulated aeglids of the same geographic region (i.e., A. alacalufi, A. manni, and

A. concepcionensis).

Continental Chile is a long and narrow
strip of land extending from Arica to Cape
Horn, characterized by a series of climatic
and biogeographical regions (see Di Castri
et al. 1968). Three main topographical fea-
tures, namely, the Andes Cordillera on the
east, the Coastal Cordillera on the west,
and the Central Valley between both rang-
es dominate the landscape from north to
south (Borgel 1983). The central southern
part of Chile, between the cities of Con-
cepci6n (36°55’S) and Puerto Montt
(41°28’S), harbors the highest diversity of
aeglids and parastacids in the country (Ba-
hamonde & Lopez 1963). This diversity
appears associated to the many rivers and
lakes that constitute the drainage system of
this area.

Aeglids are distributed along a stretch of
about 2000 kilometers, from the Choapa
River (31°38’S) in the north down to Ma-
dre de Dios Island (50°02’S) in the south
(Bahamonde & Lépez 1963, Jara & Lopez
1981). At least ten species and two sub-
species of Aegla (A. concepcionensis

Schmitt, 1942a, A. expansa Jara, 1992, A.
pewenchae Jara, 1994, A. bahamondei
Jara, 1982, A. spectabilis Jara, 1986, A.
rostrata Jara, 1977, A. abtao Schmitt,
1942a, A. d. denticulata Nicolet, 1845, A.
d. lacustris Jara, 1989, A. manni Jara,
1980, A. araucaniensis Jara, 1980, and A.
alacalufi Jara & Lopez, 1981) have been
described for the area between Concepcion
and Puerto Montt, several associated to the
Coastal Cordillera. In fact, three, A. con-
cepcionensis, A. bahamondei, and A. man-
ni, are restricted to small basins on its
western slope.

In this paper two new species of Aegla,
both from localities in the Chilean Coastal
Cordillera between Concepcién and Puerto
Montt, are described.

Specimens are deposited in the Crusta-
cean Collection of the Instituto de Zoologia,
Universidad Austral de Chile (ZUA OC),
Valdivia, Chile. The size of the specimens
was recorded as carapace length (CL), Le.,
distance between rostral apex and posterior
margin of cephalothorax.

VOLUME 112, NUMBER 1

<g™
Mal) OP :

Bag. /i.

Aegla cholchol, new species
Figs. 1, 2a—i

Aegla bahamondei.—Jara, 1982:232 (in
part), Fig. 1 (see Remarks).

Type locality.—Chol-Chol River, 100 m
downriver from bridge in the town of Chol-
Chol, 38°36'40"S, 72°51'05”W, 29 km NW
of Temuco, Province of Cautin, Chile.

Type material.—Holotype: female (24.5
mm CL), C. G. Jara coll., 22 Dec 1982,
IZUA C-328-B. Allotype: male (22.4 mm
CL). Paratypes: 9 females (17.5—30.3 mm

107

Aegla cholchol, new species, male allotype, Chol-Chol River, IZUA C-328-B, dorsal view.

LC), 6 males (14.9-27.2 mm LC), same
data as holotype.

Other material.—3 females (21.5—23.5
mim). “Queper (River .:38°51/40°S,
72 37 02 WoC) A. Viviani. coll., 24 Feb
1969, IZUA C-008-B. 3 males (24.3—26.1
mm LC), 2 females (23.8—25.6 mm LC),
Quepe River at Boroa, 39°49'06’S,
I253' 02 W.»G 4G. Jara colli; 21. Dec 1982,
IZUA C-297-B. 7 females (18.1—23.4 mm
LG),.° Traiguén. River, §38°14'45'S,
72°39'40"W, C. G. Jara & R. Navarro coll.,
22 Dec 1982, IZUA C-316. 3 males (26.9-—

108 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Aegla cholchol, new species, male allotype, Chol-Chol River, IZUA C-328-B; a, anterior part of
cephalothorax, dorsal view; b, anterior part of cephalothorax, lateral view; c, third and fourth sterna; d, second
abdominal epimeron; e, telson plate; f, ischium of left cheliped, ventral view; g, carpus of left cheliped, ventral
view; h, left cheliped, dorsal view; 1, merus of cheliped, lateral view. Scales: d = 1 mm; a, b, c, e, f, g, i = 5
mm; h = 10 mm.

VOLUME 112, NUMBER 1

18.9 mm LC), 5 females (26.4—15.5 mm
LC), Colpi River near Galvarino,
38°19'52”"S, 72°47'10"W, C. G. Jara & R.
Navarro coll., 22 Dec 1982, IZUA C-317-
B. 2 males (19.3—29.3 mm LC), 6 females
(23.6—26.0 mm LC), Quepe River,
38°51'40"S, 72°37'02’"W, C. G. Jara & R.
Navarro coll., 21 Dec 1982, IZUA C-320-
A. 8 males (13.6—30.8 mm LC), 9 females
(14.3—21.8 mm LC), Lumaco River at Lu-
maco, 38°09'46"S, 72°54’21”"W, C. G. Jara
& R. Navarro coll., 22 Dec 1982, IZUA C-
321.5 males (15.8—25.5 mm LC), 9 females
(17.3—24.1 mm LC), Cautin River, Temuco,
38°44'30"S, 72°35'00"W, C. G. Jara & R.
Navarro coll., 23 Dec 1982, IZUA C-322-
B. 8 males (11.7—21.9 mm LC), 8 females
(17.8—24.7 mm LC), Chol-Chol River near
Galvarino, 38°36'40"S, 72°51'05”"W, C. G.
Jara & R. Navarro coll., 22 Dec 1982,
IZUA C-324-C. 12 males (12.4—27.3 mm
LC), 9 females (12.5—21.2 mm LC), Lu-
maco River at Pellahuén, 38°25’40’S,
72°56'00’"W, C. G. Jara & R. Navarro coll.,
22 Dec 1982, IZUA C-326-A. 1 juvenile
(5.6 mm LC), Pichicautin River near Lu-
maco, 38°09’25"S, 72°54'15”W, T. Gonser
coll., 16 Jan 1983, IZUA C-424.
Diagnosis.—Carapace slightly longer
than wide, sparsely setose, moderately ex-
panded at branchial level. Rostrum long,
subtriangular, acute, styliform at distal half.
Orbital spine prominent; extraorbital sinus
wide, U-shaped. Anterolateral angle of first
hepatic lobe spiniform. Anterior branchial
margin subdenticulated; posterior finely
serrated, dorsally upturned. Anterolateral
angle of second abdominal epimeron spi-
niform. Fourth thoracic sternum with blunt
medial tubercle, occasionally with 1 or 2
scales on frontal border. Telson divided.
Adult males markedly heterochelous; fe-
males weakly so. Chelae robust; propodus
inflated though dorso-ventrally flattened;
dactylar lobe blunt; palmar crest subrect-
angular or arcuate, expanded, dentated; pal-
mar lobe low, blunt, with scales only on
frontal edge; medial scale largest; carpal

109

lobe with prominent conical acute spine.
Middorsal carpal ridge tubercles blunt.
Description.—Carapace slightly longer
than wide, somewhat setose especially on
branchiostegal and pterygostomial areas.
Abdominal terga and middorsal line of mer-
us of second to fourth pereopods also se-
tose. Precervical portion narrower than
postcervical; apex of acute pyramidal bran-
chial lobe separated from margin of third
hepatic lobe by wide notch. Rostrum long,
acute, narrow, styliform, slightly expanded
at its proximal 7%th but conical and some-
what flattened on distal th, gently re-
curved. Rostral tip with acute scale. Rostral
borders scarcely marginated, bearing mi-
nute scales along 7th of rostrum length, na-
ked distally. Rostral carina prominent and
scaly on proximal half of rostrum length,
absent on distal half; scales small, button-
like, in 2 subparallel rows. Epigastric prom-
inences forming low, smooth, arcuate ridge,
broadest just behind deepest point of orbital
sinus. Protogastric prominences forming
low tuberculiform area bearing short
oblique row of tiny stud-like scales. An-
terolateral lobe of carapace and hepatic
lobes well defined; former elongated as
straight conical spine reaching up to middle
of adjacent cornea. Gastric area broad, pos-
terior more protuberant. Anterolateral angle
of first hepatic lobe spiniform; spine point-
ing straight forward. Branchial lobe pyra-
midal, elongated, acutely tipped, well de-
tached from margin of third hepatic lobe
and from anterolateral angle of anterior
branchial area; with single irregular row of
acute scales on outer border. Branchial mar-
gins moderately expanded; anterior subden-
ticulated, posterior finely serrated; denticles
on anterior margin short, ill-defined, except
anteriormost which is as large as branchial
lobe; each denticle carrying 3 to 4 tiny
acute scales, mingled with short stiff setae.
Dorsum of anterior branchial area flat to
slightly concave; posterior branchial area
and adjacent marginal cardial area dome
shaped; free border of posterior branchial
area recurved and upturned. Anterolateral

110

angle of second abdominal epimeron spi-
niform; spine apex scarcely detached from
and at level with posterior branchial mar-
gin. Lateral angle of third to fifth abdominal
epimera spiniform. Fourth thoracic sternum
with blunt median tubercle, occasionally
carrying 1 or 2 scales on frontal border. Tel-
son longitudinally articulated. Chelae ro-
bust; propodus inflated though dorso-ven-
trally flattened. Dactylar lobe, at proximo-
dorsal end of dactylus, blunt, with 1 to 4
tiny scales in a row along apex. Palmar
crest subrectangular or arcuate, moderately
expanded, border coarsely dentated; teeth
subtriangular, recurved, apices point front-
dorsally, especially the most distal. Palmar
lobe, just behind dactylus-propodus joint,
blunt, low, with acute scales only on frontal
edge, its medial scale generally largest;
frontal border of palmar crest overhanging
palmar lobe, separated by deep notch. Car-
pal lobe prominent, massive, supporting ro-
bust, conical, acute, somewhat recurved
spine. Dorsointernal border of carpus, be-
hind carpal lobe, with neat row of 2 to 3
large conical acute spines, decreasing in
size towards rear. Carpal ridge as arcuate
row of blunt knob-like tubercles topped by
minute scale and/or short setae. Carpus ven-
tral face with robust conical spine. Merus
of cheliped with row of blunt to spiniform
tubercles, distalmost largest; dorsodistal an-
gle of merus tuberculiform, bearing group
of short stiff setae, and 2 to 4 scales of
which median is largest.

Color.—Live, with dorsum of carapace
and abdomen olive green; sternal surface
and ventral face of pereopods creamy
white. Dorsum of pereopods with alternat-
ing yellow green and dark green bars. Tip
of largest spines pale yellow to creamy,
darkening to green at their base. Joints of
pereopods bright red, minutely spotted
white. In alcohol, carapace and abdomen
smoky tan to creamy white.

Geographic range (Fig. 3).—Found on
the Chol-Chol River basin, from Lumaco to
Chol-Chol; also in the Cautin River, in front
of Temuco, and in the Quepe River near

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Quepe; all these are tributaries of the Im-
perial River basin which discharge into the
Pacific Ocean, at Puerto Saavedra.

Habitat.—Aegla cholchol is found in me-
dium sized rivers (order 2 to 3, after Strah-
ler, 1957) with coarse gravel and boulder
substrates where water current velocity
varies between 0.1 to 0.7 m/sec. As a con-
sequence of intensive agricultural activity,
the fluvial environment of some stretches of
the Chol-Chol River basin have undergone
severe alterations; the riparian vegetation,
mostly non-native species, is in a hetero-
geneous state of conservation. At these
sites, specimens of A. cholchol, new spe-
cies, were collected on coarse gravel to fine
quartzitic micaceous sand, sharing the bio-
tope with A. spectabilis.

Etymology.—The specific name is de-
rived from the name of the type locality, as
a latinized noun used in apposition. It is
treated as indeclinable, for the purposes of
Article 31b of the International Code of
Zoological Nomenclature (International
Trust for Zoological Nomenclature 1985).

Affinities.—Aegla cholchol, new species,
A. rostrata, and A. bahamondei are similar
in appearance but the new species is more
similar to A. rostrata than to A. bahamon-
dei. Common to the three species is a wide
carapace with expanded branchial areas and
denticulated or subdenticulated branchial
borders; steep inward downward inclination
of the branchiostegal surface; dorsum of an-
terior branchial area subconcave; well de-
fined branchial lobe; protuberant gastric and
cardiac areas with elevated middorsal line
but non-keeled as in A. denticulata; acute
first hepatic angle and anterolateral angle of
carapace; ample front and orbital sinuses
limited by acute orbital spines, one on the
plane of the orbital border and one just be-
low the former, usually not visible from
above; narrow, slightly troughed, elongated
and conically tipped rostrum. In each of the
three species, the appendages, especially
the chelipeds, present a basic morphologi-
cal pattern with proper modifications. The
new species differs from A. rostrata in hav-

VOLUME 112, NUMBER 1 111

— INTERNATIONAL BORDER
REGIONAL BORDER

Fig. 3. Map showing the geographical distribution of Aegla cholchol, new species. Asterisks indicate local-
ities where specimens were collected. In some cases, asterisk indicates more than one sampling site.

12 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ing: blunt tubercles on the carpal ridge; dor-
sodistal angle of merus of chelipeds, sec-
ond, and third pereopods blunt, or at most,
with small round tubercle tipped by minute
scales and/or setae; dactylar or palmar lobes
not spiniform; edge of the anterior branchi-
al area subdenticulated, teeth apressed non
spine-like; fourth thoracic sternum with me-
dian tubercle. Aegla cholchol, new species,
differs from A. bahamondei by having: epi-
branchial lobe monocuspidate; dactylar
lobe low and blunt; protogastric promi-
nences little protuberant; tubercle on fourth
thoracic sternum acute but non-spiniform;
posteroventral edge of merus of second to
fourth pereopods smooth, non-denticulated;
surface of carapace and chelae with few mi-
nute scales.

Remarks.—A. cholchol, new species, A.
rostrata, and A. bahamondei share a series
of complex characters. In particular, a ten-
dency to have every protruding angle of the
carapace, abdomen, and appendages, as a
pointed conical tubercle or spine. However,
the spines and spiniform tubercles are cer-
tainly more developed in A. rostrata than
in A. cholchol, new species, and A. baha-
mondei. In the latter, the angles of the car-
apace are acute but never spiniform. Jara
(1986:40) argued that “the trend toward
profuse spinulation seems correlated with
living in lentic environments.” In this con-
text, both A. cholchol, new species, and A.
bahamondéi are restricted to rivers while A.
rostrata is found mostly in lakes. Jara
(1982:237) indicated that “‘some specimens
... found in Quepe River near Temuco...
could prove to be conspecific with ... [A.
bahamondei|’’. On reexamination, those
specimens proved to be A. cholchol, new
species (IZUA C-008-B, formerly C-328).

Aegla hueicollensis, new species
Figs. 4, 5a-i

Type locality.— Pichihueicolla River, 0.5
km SE from its outlet into the Pacific Ocean
(40°10’S, 73°40’W), on the western slope

of Cordillera Pelada (Coastal Range), Prov-
ince of Valdivia, Chile.

Type series.—Holotype: male (16.5 mm
CL), 26 Jan 1983, C. G. Jara coll., IZUA
C-245. Allotype: female (16.5 mm CL).
Paratypes: 7 males (10.9-19.7 mm LOC), 3
females (14.1-14.6 mm LC), 2 juveniles
(7.3—7.7 mm LC), same data as holotype.

Other material.—3 males (10.7-14.1
mm LC), 3 females (19.3—20.7 mm LO),
Chiveria, Cordillera Pelada, 40°04’50’S,
73°10'50”"W, R. Formas coll., 14 Jan 1971,
IZUA C-025. 5 females (12.1—-18.1 mm
LC), MHueicolla River, 40°08’30’S,
73°37'42"W, R. Formas coll., 1974, IZUA
C-026. 14 males (8.9-18.3 mm LC), 7 fe-
males (11.7—16.4 mm LC), Amargos Creek,
Corral, Valdivia, 39°52'24"S, 72°26'25’W,
C. G. Jara coll., 19 Mar 1974, IZUA C-042.
1 male (13.4 mm LC), 3 females (6.8—13.2
mm LC), Trainel River, Lago Huillinco,
Chiloé, H. J. Wetzlar coll, 07 Oct 1975,
IZUA C-131. 20 males (10.1—24.8 mm
LC), 11 females (7.8—21.3 mm LC), San
Carlos Creek, San Carlos, Corral, Valdivia,
39°51'40"S, 73°26'25"W, C. G. Jara coll, 20
Mar 1976, IZUA C-175. 9 males (11.7—
24.2 mm LC), 4 females (18.3-19.9 mm
LC), Chiveria, Cordillera Pelada,
40°04'20”"S, 73°10’30"W, R. P. Schlatter
coll., 21 Jan 1977, IZUA C-184. 3 males,
(12.3-18.3 mm LC), 9 females (10.9-17.6
mm LC), Hueicolla River, 40°08’30’S,
73°39'05’"W, R. P. Schlatter coll., 22 Jan
1977, IZUA C-185. 18 males (10.5—-16.8
mm LC), 8 females (10.2-14.6 mm LC),
Manzano River, Todos los Santos Lake,
Llanquihue, 41°07'20"S, 72°24’50’W, G.
Milhe coll., 26 Apr 1974, IZUA C-196. 3
males (11.3-14.6 mm LC), 4 females
(11.3-15.9 mm LC), Refugio River, Chiloé,
R. P. Schlatter coll., 11 Oct 1977, IZUA C-
197. 2 males (8.3—23.6 mm LC), 1 female
(26.4 mm LC), Roblental Creek, eastern
slope of Cordillera Pelada, 40°16’S,
73°12'W, C. G. Jara coll., 12 Feb 1978,
IZUA C-206. 1 male (14.8 mm LOC), 3 fe-
males (15.7—21.4 mm LC), Pichihueicolla
River, 40°09'40"S, 73°39'50”W, C. G. Jara

VOLUME 112, NUMBER 1

113

Fig. 4.

coll., 26 Jan 1982, IZUA C-246. 27 males
(4.5-15.9 mm LC), 22 females (7.9—17.6
mm LC), Hueicolla River, 40°09'07’S,
73°38 25%.W, C. G. Jara coll., 27 Jan. 1982,
IZUA C-247. 5 males (10.3—19.7 mm LC),
9 females (10.1—22.1 mm LC), Chaihuin
River, 40°05'07”"S, 73°21'51”W, C. G. Jara
coll., 15 Feb 1984, IZUA C-363. 10 males
(4.2—27.2 mm LC), 22 females (6.9—26.3
man ©) fieColin «River :40°02'31'S,
73°31'26"W, C. G. Jara coll., 15 Feb 1984,

Aegla hueicollensis, new species, male holotype, Pichihueicolla River, IZUA C-245, dorsal view.

IZUA C-364. 6 males (7.0—17.2 mm LC),
6 females (8.5—15.1 mm LC), Hueicolla
River, 40°08’30"S, 73°31'38”"W, H. FE Jara
coll, 07 Feb 1992, IZUA C-507. 6 males
(9.6-16.7 mm LC), 1 female (13.9 mm
LC), Hueicolla River, 40°08'27'S,
73°30'39’"W, H. FE Jara coll, 08 Feb 1992,
IZUA C-508.

Diagnosis.—Carapace ovoidal, not ex-
panded at branchial areas. Epibranchial
lobe pyramidal, acute, borders scaly. Ros-

114 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Aegla hueicollensis, new species, male holotype Pichihueicolla River, IZUA C-245; a, anterior part
of cephalothorax, dorsal view; b, anterior part of cephalothorax, lateral view; c, third and fourth sterna; d, second
abdominal epimeron; e, telson plate; f, ischium of left cheliped, ventral view; g, carpus of left cheliped, ventral
view; h, left cheliped, dorsal view; i, merus of cheliped, lateral view. Scales: c, d, f, i = 1 mm; a, b, e, g,h =
5 mm.

VOLUME 112, NUMBER 1

trum short, wide at base, neatly triangular,
both sides of low-profiled carina barely
troughed, conical at apex, acute. First he-
patic lobe blunt. Orbital and extraorbital si-
nus present. Branchial margin smooth,
slightly nodulated. Second abdominal epi-
meron acute but not spiniform. Telson di-
vided. Fourth thoracic sternum smooth.
Male chelae elongated, pyriform, inflated at
propodus; palmar crest flabelliform, sub-
rectangular, dentate; dactylar lobe subspi-
niform; carpal lobe spiniform, apex mark-
edly displaced towards distal end of article,
frontal edge with row of tufts of short setae
mingled with scales; tubercles of carpal
ridge blunt, low, topped by transversal row
of scales and setae. Tubercles on dorsal
edge of merus of chelipeds blunt, tipped by
2 to 4 scales roughly ordered in transversal
oblique row. Ventroexternal border of is-
chium, ventral face of carpus, and ventral
borders of chelae with tufts of long setae in
row.

Description.—Carapace surface finely
punctated, each pit with a minute scale and
1 or 2 short stiff setae; highest density of
setae on abdominal epimera. Precervical
carapace almost as wide as postcervical.
Epibranchial lobe pyramidal, acute, its edg-
es lined with short acute stout scales; sim-
ilar scales along outer borders of carapace
except for posterior border and external half
of orbital sinus. Rostrum short, wide at
base, neatly triangular, barely troughed;
low-profiled carina merges into rostrum
body at proximal half; rostral apex conical,
with stout acute apical scale encircled at
base by 3 to 4 minor scales; dorsum of ca-
rina with double row of tiny scales proxi-
mally, and single unordered row distally;
ventral carina not particularly prominent, its
deepest point at base. Epigastric promi-
nences nodular, blunt, topped with 3 to 4
scales in transverse row; protogastric prom-
inences barely discernible, marked by small
field of button-like scales mingled with stiff
setae. Anterolateral lobe of carapace later-
ally expanded, flattened proximally, its ex-
ternal border slightly sinuose; anterolateral

115

angle of carapace short, acute, its apex
reaching up to proximal third of adjacent
cornea. Orbital sinus wide, delimited by an
inconstantly present orbital spine; extraor-
bital sinus variable in depth and width, gen-
erally narrowly V-shaped. First hepatic lobe
blunt, well separated from base of antero-
lateral lobe; second and third lobes scarcely
defined, separated by shallow notches in-
terrupting marginal row of scales. Branchial
margin narrow, smooth, slightly nodulated;
posterior slightly upturned. Areola broad,
convex, elevated over remaining cardial
area. Thoracic sterna flat; fourth sternum
with frontal fringe of long stiff setae; sel-
dom with low, blunt, median tubercle, car-
rying 1 or 2 scales. Anterolateral angle of
second abdominal epimeron acute, dorsum
subcarinate behind anterolateral angle; la-
teroventral face of epimeron deeply con-
cave, profusely covered by short stiff setae;
free ventral border with dense row of long
plumose setae, also present over posterior
branchiostegal surface, edge of remaining
abdominal epimera, and borders of uropods
and telson. Telson longitudinally articulat-
ed.

Males heterochelous; left hand largest.
Chelae robust but not particularly massive,
subovoidal in outline, surface microtuber-
culate; each tubercle carrying a minute
acute scale and 2 short stiff microsetae,
most evident on distal half of propodus and
dactylus. Molar process well developed on
cutting edge of fixed finger and dactylus of
left chela but negligible or absent on night
one. Dorsum of dactylus with robust spi-
niform lobe on proximal end. Ventral face
of chelae with prominent ridge along ven-
troexternal border of propodus, marked by
row of pits and tufts of long setae; pits more
numerous along dactylus and fixed finger.
Base of palmar crest on ventral side of che-
lae parallel to short tuberculate ridge with
large pits and tufts of setae. Central part of
ventral face of chelae broadly convex,
forming longitudinal ridge between center
of propodus-carpus joint and distalmost
dactylus-propodus joint, marked by large

116

pits and setae. Palmar crest prominent,
thick, its border denticulated with scattered
small acute scales between contiguous larg-
er denticles. Distal end of palmar crest sep-
arated from predactylar lobe of carpus by
almost right-angled deep notch; predactylar
lobe elevated, its free border with irregular
row of short acute scales; largest scale at
anterodorsal angle. Carpus robust, with two
dorsal ridges; outermost lower, distinguish-
able as slightly arcuate row of broad low
tubercles; apex of tubercles with row of 3
to 6 short stiff setae mingled with minute
scales; innermost prominent, formed by
row of blunt tubercles topped by tranverse
row of 3 to 5 acute scales and setae. Carpal
lobe spiniform, asymetrically subtriangular,
apex leveled with frontal border of same
lobe. Inner border of carpus with row of 3
large spines proximally decreasing in size.
Dorsum of anteriormost spine with 2 to 3
scales or setae protruding from large pits in
a row along spine axis. Same arrangement
On minor spines or at least, 1 pit and setae
present. Ventral surface of carpus with ro-
bust conical spine; between it and inner
border of carpus several thick, long, simple
setae, uneven in size. Dorsal edge of merus
with row of subacute tubercles tipped by 1
or 2 scales, 3 or 4 on distalmost tubercle;
frontal side of tubercles with tufts of long
simple setae; frontodorsal angle blunt,
thick, little prominent, with patch of short
stiff setae and scales; ventral borders with
row of tubercles tipped with scale and/or
seta; distal angles as short conical spine.
Dorsal edge of merus of second, third and
fourth pereopods with fringe of long plu-
mose setae covering minute spiniform tu-
bercles; carpus, propodus and dactylus with
fringe of short stiff simple setae. Postero-
ventral edge of merus scabrous, subserrat-
ed, with narrow band of setae and scales.
Color.—Not observed live. In alcohol,
carapace, abdomen, and appendages uni-
formly smoky-tan to creamy white.
Geographic range (Fig. 6). Small
streams and creeks flowing from east to
west on the western slope of Cordillera Pe-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

lada (Coastal Range between Corral to the
north and the outlet of the Bueno River to
the south). Specimens collected are from:
Amargos Creek (Corral, Valdivia), Coltin,
Chaihuin, Hueicolla, and Pichihueicolla riv-
ers. Also found in Manzano River, at Todos
Los Santos Lake, in the Andean district of
the Llanquihue province.

Habitat.—Small rivers and fast flowing
streams on moderate to steep slopes, cov-
ered or surrounded by cold-temperate Val-
divian rain forest. Specimens are found
among pebbles and boulders on the stream
bottom, at sites where a heterogeneous
mass of vegetal detritus accumulates serv-
ing as shelter for the crabs.

Etymology.—The specific name derives
from Hueicolla, a small town on the Pacific
coast about 40 km to the south from Corral,
where the rivers Hueicolla and Pichihuei-
colla discharge into the Pacific Ocean.

Affinities.—Aegla hueicollensis, new spe-
cies, shares a series of features with other
species of Aegla. In general, the new spe-
cies closely resembles A. abtao. Several
morphological similarities suggest a close
ancestral relationship between the two. A.
hueicollensis, new species, also displays
morphological similarity to A. alacaluf.
Both share the almond-shaped outline of
the carapace, the short triangular acute ros-
trum, and the pronounced distalward dis-
placement of the apex of the carpal lobe.
However, A. alacalufi has an undivided tel-
son plate, and lacks minute scales on the
carapace surface. The new species also
shares features with A. manni and A. neu-
quensis Schmitt, 1942b, such as: the ireg-
ular row of scales and/or setae along the
dorsal axis of the larger spines on the inner
border of carpus of chelipeds and the trans-
verse row of scales on the tubercles of the
carpal ridge of chelae. The new species dif-
fers from A. manni in having the rostral ca-
rina and rostral margins scaly; from A. neu-
quensis by having the anterolateral angle of
second abdominal epimeron subspinose.
The low, somewhat concave, profile of the
rostral carina and the protuberant gastric

VOLUME 112, NUMBER 1 i

-Colico
CSS
d t
was ewais\

ya
J
Lolafquen
at (MS.

| H
—<--——"_ INTERNATIONAL BORDER
—-:-— REGIONAL BORDER

Fig. 6. Map showing the geographical distribution of Aegla hueicollensis, new species. Asterisks indicate
localities where specimens were collected. In some cases, asterisk indicates more than one sampling site.

118 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

area of A. hueicollensis, new species, re-
semble those of A. concepcionensis. These
similarities led Schmitt (1942b:504) to re-
port specimens of Aegla from Corral
(northern limit of Cordillera Pelada) as A.
concepcionensis. Similarly, Haig (1955:
504) classified as A. concepcionensis spec-
imens from Ancud (230 km to the south of
Corral), Chiloé Island. To determine if these
specimens are conspecific with A. hueicol-
lensis, new species, they would have to be
re-examined.

Remarks.—The geographic range of A.
hueicollensis, new species, includes a series
of localities on the western slope of the
Coastal Cordillera and one locality (Man-
zano River) on the western slope of the An-
des Cordillera, about 100 km southeast of
the type locality. There is no hydrological
connection between these areas at present.
This fact leads to the assumption that, in
the past, the geographic range of the new
species must have been broader, enclosing
localities on the intervening territory be-
tween the Andean and the Coastal ranges.
Connected to this is the fact that southern
Chile, from paralell 38°S southward, was
severely affected by Quaternary glaciations.
Illies (1960) found evidence that during
glaciations the ice-sheet front advanced
west from the Andean highlands completely
covering the Central Valley; the ice front
pushed upon the eastern slope of the Coast-
al Range but never surmounted it. The
western slope of the Coastal Range which
was never completely covered by ice (Illies
1960) could have served as a refugial area
for A. hueicollensis, new species. Its pres-
ence in the small Manzano River basin,
amid the Andean massif, presumes that
somehow the site was unaffected by de-
structive glacial effects, thus preserving the
preexisting population.

Acknowledgments

The authors thank Drs. R. Formas, T.
Gonser, H. E Jara, R. Navarro, and R. P.
Schlatter for the presentation of valuable

material; Dr. Rafael Lemaitre and two
anonymous reviewers for their efforts to
improve the readability of the manuscript.
The present paper was financed by grants
91-0900 of the Fondo de Investigacién
Cientifica y Tecnold6gica (Fondecyt, Chile)
and S-91-4 of the Direccié6n de Investiga-
cidn y Desarrollo, Universidad Austral de
Chile.

Literature Cited

Bahamonde, N., & M. T. Lépez. 1963. Decapodos de
aguas continentales en Chile.—Investigaciones
Zoolégicas Chilenas 10:123—149.

Borgel, R. 1983. Geomorfologia. in Geografia de
Chile, 2. Instituto Geografico Militar, Santiago,
Chile, 182 pp.

Di Castri, EF 1968. Esquisse écologique du Chili. Biol-
ogie de l’Amérique Australe. Pp. 7—82 in C.
Delamare Deboutteville & E. Rapoport, eds.,
Biologie de 1’ Amérique Australe, 4. Editions du
Centre National de la Recherche Scientifique,
Paris, 685 pp.

Haig, J. 1955. The Crustacea Anomura of Chile.—Re-
ports of the Lund University Chile Expedition
1948-49, 20. Lund Universitets Arsskrift. N.E
Avd. 2, 51(12):1—68.

Illies, H. 1960. Geologie der Gegend von Valdivia
(Chile).—Neues Jahrbuch fiir Geologie und Pa-
laentologie 111:30—110. (Translated into Span-
ish by J. Walper. Universidad Austral de Chile,
Valdivia, 1970, 50 pp.)

International Trust for Zoological Nomenclature. 1985.
International Code of Zoological Nomenclature,
Third edition, London, 338 pp.

Jara, C. 1977. Aegla rostrata, n. sp. (Decapoda, Ae-
glidae), nuevo crustaceo dulceacuicola del Sur
de Chile.—Studies on Neotropical Fauna and
Environment 12:165—176.

. 1980. Dos nuevas especies de Aegla Leach

(Crustacea, Decapoda, Anomura) del sistema

hidrografico del Rio Valdivia.—Anales del Mu-

seo de Historia Natural de Valparaiso 13:255-—

266.

. 1982. Aegla bahamondei, new species (Crus-

tacea: Decapoda: Anomura) from the Coastal

Mountain Range of Nahuelbuta, Chile.—Jour-
nal of Crustacean Biology 2(2):232—238.

. 1986. Aegla spectabilis, a new species of

freshwater crab from the eastern slope of the

Nahuelbuta Coastal Cordillera, Chile.—Pro-

ceedings of the Biological Society of Washing-

ton 99:34-41.

. 1989. Aegla denticulata lacustris, new sub-

species, from Lake Rupanco, Chile (Crustacea:

Decapoda: Anomura: Aeglidae).—Proceedings

VOLUME 112, NUMBER 1

of the Biological Society of Washington 102:
385-393.

. 1992. Aegla expansa, new species (Crustacea:
Decapoda: Anomura: Aeglidae), from the Low-
er Bio-Bio river basin, Concepcién, Chile.—
Gayana (Zoologia) 56(1—2):49-—57.

. 1994. Aegla pewenchae, new species of Cen-
tral Chilean freshwater decapod (Crustacea: An-
omura: Aeglidae).—Proceedings of the Biolog-
ical Society of Washington 107:325-—339.

, & M. T. Lopez. 1981. A new species of fresh-
water crab (Crustacea: Anomura: Aeglidae)
from insular South Chile.—Proceedings of the
Biological Society of Washington 94:88—93.

119

Nicolet, H. 1849. Crustaceos. Pp. 113-318 in C. Gay,
Historia Fisica y Politica de Chile, Zoologia 3,
547 pp.

Schmitt, W. L. 1942a. Two new species of Aegla from
Chile.—Revista Chilena de Historia Natural
44(1940):25-31, pl. 1.

. 1942b. The species of Aegla, endemic South
American freshwater crustaceans.—Proceedings
of the United States National Museum 91:431—
a74U)

Strahler, A. N. 1957. Quantitative analysis of water-
shed geomorphology.—Transactions of the
American Geophysicists Union 38:13—920.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(1):120—132. 1999.

The Hawaiian parthenopid crabs of the genera
Garthambrus Ng, 1996, and Dairoides Stebbing, 1920
(Crustacea: Decapoda: Brachyura)

Peter K. L. Ng and S. H. Tan

Department of Biological Sciences, National University of Singapore, Lower Kent Ridge Road,
Singapore 119260, Republic of Singapore

Abstract.—The taxonomy of four species of deep-water parthenopid crabs,
Garthambrus stellata (Rathbun, 1906), G. lacunosa (Rathbun, 1906), G. com-
planata (Rathbun, 1906) and Dairoides kusei (Sakai, 1938) from Hawaii is
treated. Dairoides kusei is recorded from Hawaii for the first time. The iden-
tities of G. stellata, G. lacunosa and G. complanata are clarified, with detailed
comparisons and figures provided. Despite earlier reports, there are no inter-
mediates between these three species. A key to the genus Garthambrus is

provided.

Rathbun (1906), in her treatment of the
Hawaiian Brachyura, had originally recog-
nized eight parthenopid species from these
islands. One species, Harrovia truncata
Rathbun, 1906, was transferred to a sepa-
rate genus (Cyrtocarcinus Ng & Chia,
1994) in the Xanthidae by Ng & Chia
(1994). Another species, Parthenope (Pla-
tylambrus) stellata Rathbun, 1906, was
originally established with three subspecies,
P. (P.) stellata stellata, P. (P.) stellata la-
cunosa Rathbun, 1906, and P. (P.) stellata
complanata Rathbun, 1906. Garth (1993)
reappraised the validity of these three taxa
and regarded them as distinct species. He
briefly commented that the features used by
Rathbun (1906) to separate the three vari-
eties were good species characters. Al-
though Garth examined the type series of
all three taxa, he did not comment much
about them or figure any of the types. This
despite the fact that one taxon, P. (P.) com-
planata, had never been figured before. Nor
did he comment on specimens which Rath-
bun (1906) had regarded as being interme-
diate between the three species. In describ-
ing a new species (Parthenope cidaris)
from Australia, Garth & Davie (1995) pro-

vided photographs of the types of P. (P.)
stellata, P. (P.) lacunosa and P. (P.) com-
planata but did not make any additional
comments other than again stating briefly
that all three taxa were good species.

Sakai (1938, 1976) had commented that
P. (P.) lacunosa (as a subspecies) was a
junior synonym of Lambrus (Parthenopo-
ides) pteromerus Ortmann, 1893, a species
which he transferred to the genus Tutankh-
amen Rathbun, 1925. Garth (1993), how-
ever, disagreed, commenting that P. (P.) la-
cunosa lacked the lamellar ridges lining the
afferent channels on the carapace found on
Tutankhamen cristatipes (A. Milne Ed-
wards, 1880), the type and only species of
the genus. On the basis of Garth’s (1993)
redescription of P. (P.) stellata and his
comments on P. (P.) lacunosa and P. (P.)
complanata, Ng (1996) subsequently trans-
ferred all three species to the genus Gar-
thambrus Ng, 1996.

Nine species of Parthenopidae are cur-
rently known from the Hawaiian islands,
Garthambrus stellata (Rathbun, 1906), G.
lacunosa (Rathbun, 1906), G. complanata
(Rathbun, 1906), Platylambrus nummifera
(Rathbun, 1906), Rhinolambrus lamelligera

VOLUME 112, NUMBER 1

(White, 1847), Aulacolambrus hoplonotus
(Adams & White, 1848), Aulacolambrus
whitei (A. Milne Edwards, 1878), Pseudo-
lambrus calappoides (Adams & White,
1848), and Daldorfia horrida (Linnaeus,
1758). The generic classification used here
follows that proposed by Ng & Rodriguez
(1986) and Ng (1996).

The present paper reports on the deep-
water Parthenopidae from Hawaii primarily
based on material deposited in the Bernice
P. Bishop Museum, Honolulu; and National
Museum of Natural History, Smithsonian
Institution, Washington, D.C. Garthambrus
stellata, G. lacunosa and G. complanata are
rediagnosed, and detailed figures of their di-
agnostic characters provided for the first
time. Using new characters identified in this
study, the ‘‘intermediates’’ reported by
Rathbun (1906) can easily be assigned to
the three species. A key to the genus Gar-
thambrus is provided. The unusual deep-
water species, Dairoides kusei (Sakai,
1938), previously known only from Japan,
is also reported from Hawaii for the first
time.

The terms used here essentially follow
those used by Garth (1958, 1993). The ab-
breviations cw and cl are for the carapace
width and length, and Gl and G2 are for
the male first and second pleopods respec-
tively. Specimens are deposited in the Bern-
ice P. Bishop Museum, Honolulu (BPBM);
National Museum of Natural History,
Smithsonian Institution, Washington, D.C.
(USNM); Nationaal Naturhistorish Muse-
um, Leiden (ex Rijksmuseum van Natuur-
lijke Historie, RMNH); and the Zoological
Reference Collection, National University
of Singapore (ZRC).

Systematic Account
Genus Garthambrus Ng, 1996

Garthambrus Ng, 1996: 156 (type species:
Garthambrus posidon Ng, 1996, by orig-
inal designation).

Diagnosis.—Carapace subtriangular in
shape, broader than long; angle between an-

121

tero- and posterolateral margins strongly
produced; dorsal surfaces granulose, spi-
nose or rugose; progastric, mesogastric,
metagastric, mesobranchial, metabranchial,
cardiac and intestinal regions strongly in-
flated; gastric and branchial regions sepa-
rated by deep grooves. Median lobe of ros-
trum prominent, sub-spatulate, deflexed
downwards. Hepatic region and margin
separated from anterolateral margin by
cleft, notch or large tubercle. Posterolateral
margin and metabranchial regions without
long or prominent teeth or spines. Cheli-
peds at least 2.5 times carapace length. G1
relatively stout, not armed with long spines
or stiff hairs. Distal segment of G2 elon-
gate, subequal to or distinctly longer than
basal segment (modified from Ng 1996:
156):

Remarks.—Ng (1996) removed a number
of species previously assigned to Parthen-
ope, Platylambrus or Parthenope (Platy-
lambrus) to a separate genus which he char-
acterised by their triangular carapaces and
long distal segment of the second male ple-
opod. All eight species (see key) currently
placed in Garthambrus are essentially deep-
water species.

One unusual species which requires men-
tion is the Japanese species Parthenope
pteromerus (Ortmann, 1893). It is very sim-
ilar in appearance to G. lacunosa, which
Sakai (1976) synonymised under P. pter-
omerus. Sakai (1976) also transferred P.
pteromerus to the genus Tutankhamen
Rathbun, 1925, arguing that the afferent re-
Spiratory channels on its carapace are lined
with lamellar ridges. Garth (1993) refuted
this synonymy because G. lacunosa and all
Garthambrus species do not have such la-
mellar ridges. Subsequently, Ng (1996)
suggested that P. pteromerus was not as-
signable to Tutankhamen, and probably be-
longed in a separate genus. While P. pter-
omerus resembles many Garthambrus spe-
cies superficially, the possession of lamellar
ridges lining the afferent respiratory chan-
nels on its carapace argues against its in-
clusion in Garthambrus as presently de-

122 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

fined. Parthenope pteromerus is also a rel-
atively shallow-water species compared to
known species of Garthambrus (unpub-
lished data).

Key to Garthambrus species

la. Carapace and legs covered with numer-
ous long, sharp spines; margin of bran-
chial region with deep, distinct cleft ..
Si Mai i la Mi gene Me G. mironovoi
1b. Carapace and legs not covered with nu-
merous long, sharp spines; margin of
branchial region entire or almost so .. 2
2a. Granules on branchial, cardiac and gas-
tric regions of carapace fused to various
degrees forming distinct granular
patches G. allisoni
2b. Carapace dorsal surface almost smooth
or if granules present, those on bran-
chial, cardiac and gastric regions never
distincily, fuseds:--5.:.5, awtaanh ees: 3
3a. Margins of ambulatory meri distinctly
ih i: ae ooo es ane G. lacunosa
3b. Margins of ambulatory meri not dis-
tinctly cristate, but lined with spines,
teeth or granules
4a. Carapace and cheliped surfaces without
pitsen McCune e.g oe G. complanata
4b. Carapace and cheliped surfaces with

distinct pits and/or lacunae ......... =i
5a. Margins of ambulatory meri with
prominent rounded granules ........ 6

5b. Margins of ambulatory meri with un-
even granules, tubercles or small spines 7
6a. Median protuberance of rostrum large,
margins of ambulatory propodi and car-
pi distinctly granulated ...... G. poupini
6b. Median protuberance of rostrum small,
margins of ambulatory propodi and car-
pi smooth, unarmed '5.". 5/20. G. posidon
7a. Dorsal surface of carapace densely
granulated all over; metabranchial re-
gions strongly swollen, appears peak-
like from frontal view G. stellata
7b. Dorsal surface of carapace with rugae
and granules but never densely packed;
metabranchial regions high but not very
swollen, not peak-like from frontal
view G. cidaris

Garthambrus stellata (Rathbun, 1906)
(Figs. 1A—D, 6A—C)

Parthenope (Platylambrus) stellata Rath-
bun, 1906:884 (part), pl. 15 Figs. 1, 2.—
Garth, 1993:786, Figs. 3, 4.

Parthenope (Platylambrus) stellatus.—Se-
rene, 1968:60.

Parthenope stellata.—Garth & Davie,
1995-225, Fis. 2B.

Garthambrus stellata.—Ng, 1996:158.

Material examined.—Holotype, 1 male,
cw 48.6 mm, cl 32.8 mm (USNM 29839),
south coast of Oahu Island, Hawaii, 435—
461 m, station 3811, coll. Albatross Expe-
dition, 27 Mar 1902. Paratype, 1 male, cw
32.9 mm, cl 23.1 mm (USNM 29840), Ka-
waihae, Hawaii Island, Hawaii, 20°01’45”N
155°54'15”"W, 269-362 m, station 4045, 8
hemp tangles gear, coll. Albatross Expedi-
tion, 11 Jul 1902. Others—1 female
(USNM 239148), Pearl Harbor, Mamala
Bay, Oahu Island, Hawaii, disposal site,
21°16'48"N 157°56'30"W, 366 m, coll. R/V
Hurl, 30 Apr 1983. 1 male, cw 70.2 mm,
cl 45.6 mm (ZRC 1997.441), Off Pearl Har-
bor, Oahu Island, Hawaii, station 82-105, in
dredge spoil site, on sediment bottom in vi-
cinty of outcrop, 366 m, coll. R/V Hurl,
Makali’i Dive, D. M. Devaney & B. Bart-
ko, 1 Sep 1982. 1 female (carapace only)
(BPBM 1978.417), Off Makupuu Point,
Molokai Channel, Hawaii, 12 km straight
cast, 21°02’N 157°32’W, coll. Valkryrien,
Capt. S. Rayner, 20 May 1967. 1 male, ca.
cw 65.5 mm, cl 43.4 mm (left edge of car-
apace broken) (BPBM 1976.259), Entrance
to Pearl Harbor, Oahu Island, Hawaii, 2.5
km off buoy 1,338 m, coll. in shrimp traps,
Easy Rider, E. H. Chave, 27 Sep 1976. 2
females, cw 21.6 mm, cl 16.5 mm, ca. cw
16.0 mm, cl 12.5 mm (right side of cara-
pace crushed) (BPBM 5508), Oahu Island,
1.9 km off Kahala, Hawaii, 46 m, coll.
Brock, 14 Apr 1949.

Diagnosis.—Surface of carapace densely
and granulated uniformly throughout, in-
cluding grooves and depressions; metabran-
chial regions strongly swollen, appearing

123

VOLUME 112, NUMBER 1

A, Car-

Pas ce

USNM

1906). Holotype male (48.6 by 32.8 mm) (
Left chela.

>

Garthambrus stellata (Rathbun

Fig. 1.
apace; B, Abdomen; C

Front; D,

5

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig, 2
Carapace; B, Abdomen.

hee

Fig. 3. Male telson and sixth abdominal segment.
A, Garthambrus stellata, holotype male (USNM
29839): B, Garthambrus lacunosa, holotype male
(USNM 29842).
(USNM 29842b).

C, Garthambrus lacunosa, male

Garthambrus lacunosa (Rathbun, 1906). Holotype male (30.9 by 21.8 mm) (USNM 29842). A,

peak-like in frontal view (Fig. 1C), highest
point with sharp granule; sub-branchial,
suborbital, subhepatic and branchiostegal
regions distinct, densely covered with dis-
tinct small granules. Anterolateral margin
with sharp, acutely triangular teeth, lateral
margins of teeth with numerous accessory
spinules and/or small, sharp granules. Che-
liped surfaces (especially merus, carpus,
propodus and dactylus) with well devel-
oped sharp spines, spinules and granules
(Fig. 1D); merus with oblique ridge of
strong spines about 4% from proximal end;

VOLUME 112, NUMBER 1

Fig. 4. Garthambrus complanata (Rathbun, 1906). Lectotype male (17.3 by 12.7 mm) (USNM 29845a). A,

Carapace; B, Abdomen.

surfaces never with pits or lacunae. Margins
of ambulatory legs unevenly cristate to den-
tate, with numerous accessory spinules; sur-
faces densely covered with small granules,
never with pits or lacunae. Anterior thoracic

sternites densely covered with small gran-
ules, without pits or lacunae (Fig. 1B). Ab-
dominal surfaces densely covered with
small granules, never with pits or lacunae;
lateral margins of male telson gently con-

126

vex. Gl relatively slender, tip distinctly
turned inwards towards median part of tho-
racic sternum.

Remarks.—Rathbun (1906) described
Parthenope stellata on the basis of ten
specimens from various parts of Hawaii. In
her discusion on variation within the spe-
cies, however, she noted that only two spec-
imens conformed with P. stellata sensu
stricto, and the other eight specimens ap-
peared to belong to two other varieties or
were intermediate in form (see discussion
for next two species). One specimen
(USNM 29839) was designated as the
““‘type’’ (= holotype) of P. stellata. How-
ever, Rathbun (1906:884), noted that the
smaller specimen (USNM 29840) “...
shows the tubercles and spines all sharp in-
stead of blunt pointed, and lacks the hair
near the ends of the legs.’’ She regarded
this smaller specimen as belonging to the
‘“‘sharp-spined variety”’ of P. stellata sensu
stricto, but did not apply any name. Ex-
amination of a relatively good series of
specimens has shown that differences noted
by Rathbun (1906) between her two type
specimens can easily be accounted for by
infraspecific variation. Edmondson (1951:
213) had reported two small specimens
from shallow waters in Hawaii and Ng
(1996:159) commented that “... his spec-
imens are probably misidentified’ as the
carapace proportions were different, the
rostrum was not trifid and the ambulatory
dactylus was more styliform. We have re-
examined these two specimens (BPBM
5508), and they are clearly juvenile G. stel-
lata. The differences in carapace propor-
tions are almost certainly due to their small
size and the ambulatory dactylus is actually
not as styliform as depicted in Edmondson’s
(1951:Fig. 18d) illustration. The bifid ros-
trum shown for the specimen figured by
Edmondson (1951:Fig. 18a) is such because
the median lobe is broken. In the other
specimen, the rostrum is clearly trifid.

The relatively good series of specimens
of various sizes and sexes shows that the
characters identified here are consistent and

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

are useful for defining this species. The G1
shows slight variation, in that the distal part
may appear slightly more flared or slightly
more bent inwards (towards the median part
of the thoracic sternum). The density of the
granules does not change much with
growth. However, with an increase in size,
the individual granules do become larger,
more prominent and distinctly more stellate
in structure.

Garthambrus stellata is known from Ha-
wali, Tuamotu and Taiwan (Garth 1992,
Poupin 1996:32, Tan et al. 1999). In Ha-
wail, the species is recorded up to depths
of 461 m, although juveniles have been
found in shallower waters of only about 46
m. The record from Tuamotu is of an adult
from about 300 m depth. The specimen fig-
ured by Poupin (1996: pl. 15f) from Tua-
motu is a dirty white overall and speckled
with numerous fine brown spots, and the
fingers are pigmented brown.

Garthambrus lacunosa (Rathbun, 1906)
(Figs. 2A—B, 3A—C, 6D—P)

Parthenope (Platylambrus) stellata lacu-
nosa Rathbun, 1906:884, pl. 15 Fig. 7.
Parthenope (Platylambrus) lacunosa.—
Garth, 1993:788.

Parthenope lacunosa.—Garth & Davie,
1995:220; Pigs SA.

Garthambrus lacunosa.—Ng, 1996:158

Material examined.—Holotype, 1 male,
cw 30.9 mm, cl 21.8 mm (USNM 29842),
Kawaihae, west coast of Hawaii Island, Ha-
waii, 269-362 m, 20°01’45"N 155°54’
15”W, station 4045, 8 hemp tangles gear,
coll. Albatross Expedition, 11 Jul 1902.
Paratypes, 1 male, cw 30.4 mm, cl 31.6 mm
(USNM 29843a), Maui Island, Lipoa Point,
Pailolo Channel, Hawaii, 238—276 m,
21°05'30"N__ 156°40'30’W, station 4100, 8
hemp tangles gear, coll. Albatross Expedi-
tion, 23 Jul 1902, (1 other paratype male
specimen transferred to Stanford University
according to labels, not examined). 1 fe-
male (USNM 29844), northwest coast of
Oahu Island, Hawaii, 282—357 m, coll. Al-

VOLUME 112, NUMBER 1

batross Expedition. 1 female, cw 30.2 mm,
cl 21.3 mm (USNM 29841), south coast of
Molokai Island, Hawaii, off Lae-O-Ka Laau
Lighthouse, 309-333 m, 64°00’N
00°13'42’W, station 3835, coll. Albatross
Expedition, 3 Apr 1902. Others—1 male,
cw 31.4 mm, cl 21.5 mm (USNM 29843b),
Maui Island, Lipoa Point, Pailolo Channel,
awa, 238-276 m, 21°05'30"N
156°40’30’W, station 4100, 8 hemp tangles
gear, coll. Albatross Expedition, 23 Jul
1902.

Diagnosis.—Surface of carapace slightly
rugose, with scattered to relatively numer-
ous granules; metabranchial regions mod-
erately raised but not strongly swollen, not
peak-like from frontal view, highest point
unarmed; sub-branchial, suborbital, subhe-
patic and branchiostegal regions distinct,
covered with numerous but scattered small
granules. Anterolateral margin with lobi-
form to truncate teeth, lateral margins of
teeth lined with numerous accessory spi-
nules or small, sharp granules especially
along anterior margin. Outer surfaces of
chelipeds (especially merus, carpus, pro-
podus and dactylus) with well developed
spines, spinules and granules; merus with
oblique ridge of strong spines about % from
proximal end; outer surfaces with numerous
pits, often with well developed lacunae.
Margins of ambulatory legs distinctly cris-
tate, entire to uneven; surfaces almost
smooth or with very small, scattered gran-
ules, usually with distinct pits and/or lacu-
nae. Anterior thoracic sternites with numer-
ous scattered small granules and pits, often
with lacunae. Abdominal surfaces covered
with numerous scattered small granules and
pits, often with lacunae; lateral margins of
male telson concave. G1 relatively slender,
tip turning slightly inwards towards median
part of thoracic sternum.

Remarks.—Rathbun (1906) regarded five
specimens of the original nine specimens of
Parthenope (Platylambrus) stellata belong-
ing to a separate variety P. (P.) stellata la-
cunosa. She has also mentioned that the dif-
ferences of these five specimens are so dif-

127

ferent from the type specimen of G. stel-
lata, that they could be a different species
altogether and she mentioned several char-
acters: ‘““The branchio-cardiac depression is
deep, and another depression runs along the
outer side of the branchial region, adjacent
to the marginal teeth. The elevated part of
this region has a row of large pits through
its middle, and similar line of pits dividing
the gastric region in three and roughening
the chelipeds. The granules are in large part
confluent and thus obliterated, especially on
the higher parts of the carapace and the che-
lipeds. The legs have smooth surfaces, thin
cristate margins which are somewhat cre-
nate or dentate in the merus and are desti-
tute of long hair. Along with two of this
variety from station 4100 is one which is
intermediate between the typical (P. stellata
stellata) and varietal form (P. stellata la-
cunosa), the stellate granules being every-
where fairly well shown, and also the lines
of pits.”” (Rathbun, 1906:884).

One of the original nine type specimens
of Garthambrus stellata was considered as
intermediate between P. stellata and P. la-
cunosa. This specimen (USNM 29843b) is
similar to P. stellata sensu stricto in having
smaller granules overall, even on the cara-
pace depressions and grooves, compared to
typical P. lacunosa. The lacunae on the car-
apace and chelipeds are also smaller and
less developed. A close examination of this
supposedly intermediate specimen, howev-
er, Shows that it only represents the extreme
end of the variation in P. lacunosa. In the
presence of lacunae on the carapace and
chelipeds, truncate anterolateral teeth, dis-
tinctly cristate ambulatory meri, low, non-
peaked metabranchial regions which lack a
sharp median granule and gently concave
male telson (Fig. 3C), the specimen clearly
represents P. /acunosa. Even though this
specimen is more granulated than typical P.
lacunosa, it is still much less granulated
overall compared to any specimen of P.
stellata sensu stricto we have examined,
even specimens of similar sizes.

Rathbun (1906) identified one specimen

128 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(USNM 29842) as the “‘type’’ (= holotype)
of Garthambrus lacunosa. The other four
specimens listed by Rathbun (1906) are
thus paratypes. The supposedly intermedi-
ate specimen (USNM 29843b) was not re-
garded as belonging to P. (P.) stellata la-
cunosa by Rathbun (1906) when she orig-
inally named the taxon, and thus, is not part
of the type series.

Garthambrus complanata (Rathbun, 1906)
(Figs. 4A—B, 6G—I)

Parthenope (Platylambrus) stellata com-
planata Rathbun, 1906:884.

Parthenope (Platylambrus) stellata com-
planataus Seréne, 1968:60.

Parthenope (Platylambrus) complanata.—
Garth, 1993:789.

Parthenope complanata.—Garth & Davie,
1995:226, Fis. 3B.

Garthambrus complanata.—Ng, 1996:158.

Material examined.—Lectotype, herein
designated, 1 male, cw 17.3, cl 12.7 mm
(USNM 29845a), Hanamaulu Bay, Kauai
Island, Hawaii, 470-570 m, 22°01'30"N
150°21'10’”W, station 4132, 8 foot Blake
Beam trawl gear, coll. Albatross Expedi-
tion, 1 Aug 1902. Paralectotype, 1 male
(USNM 29845b), same data as lectotype.

Diagnosis.—Surface of carapace smooth
to gently rugose, not granulated; metabran-
chial regions raised but not strongly swol-
len, not peak-like from frontal view, highest
point unarmed; sub-branchial, suborbital,
subhepatic and branchiostegal regions dis-
tinct but almost smooth except for a few
scattered granules. Anterolateral margin
with liboform to truncate teeth, lateral mar-
gins of teeth lined with scattered accessory
spinules or small, sharp granules especially
along anterior margin. Surface of chelipeds
(especially merus, carpus, propodus and
dactylus) with few, scattered granules, but
with distinct simple spines; merus without
distinct oblique ridge of spines; surfaces
with few, scattered simple granules. Mar-
gins of ambulatory legs smooth, not cristate
but with well spaced spines and/or teeth;

surfaces smooth, without lacunae. Anterior
thoracic sternites almost smooth, without
pits or lacunae. Abdominal surfaces almost
smooth, without pits or lacunae; lateral
margins of male telson concave. G1 rela-
tively stout, straight.

Remarks.—For her third variety of P.
(P.) stellata, Rathbun (1906:884) noted that
“Still a third type seems worthy of a dis-
tinguishing name, P. (P.) stellata complan-
ata. It differs from the type of P. (P.) stel-
lata stellata in the surface of carapace and
chelipeds being smooth to the naked eye,
though under the lens finely punctate and
roughened; the tubercle or spine at the inner
third of the postero-lateral margin is repre-
sented by a triangular nodule; tubercle at
each end of posterior margin large and
round; antero-lateral teeth broader and more
dentiform than in other forms; no teeth nor
spines at outer end of postero-lateral mar-
gin, but a nodule on the dorsal surface at
that point may represent them; marginal
spines of chelipeds inclining to sharp; legs
approaching the type in roughness; margins
prominently spinate, without long hair.”
Most of these differences are valid, and G.
complanata can easily be separated from
congeners by the carapace sculpturation, ar-
mature of the antero-lateral margin, absence
of an oblique ridge on the relatively smooth
merus of the cheliped, structure of the am-
bulatory merus, and form of the thoracic
sternites and abdominal surface.

Garth & Davie (1995:Fig. 3B) provided
a photograph of the “‘holotype”’ of this spe-
cies, but no measurements were provided.
However, Rathbun (1906) listed two male
specimens (USNM 29845a) only as
‘types’? without designating a holotype;
thus, both are syntypes. Garth (1993) had
earlier examined both specimens and listed
them as syntypes. The larger specimen
(17.3 by 12.7 mm) (USNM 29845a) in bet-
ter condition, is here designated as the lec-
totype. It is the same specimen figured by
Garth & Davie (1995).

VOLUME 112, NUMBER 1

129

Fig. 5.

Genus Dairoides Stebbing, 1920

Dairoides Stebbing, 1920:233 (type spe-
cies: Dairoides margaritus Stebbing,
1920, by monotypy).

Astherolambrus Sakai, 1938:341 (type spe-
cies: Astherolambrus kusei Sakai, 1938,
by monotypy).

Remarks.—Sakai (1938) established
Astherolambrus for a new species from Ja-
pan, A. kusei. Sakai (1965) later synony-
mised Astherolambrus under Dairoides
Stebbing, 1920, which had been established
for D. margaritus Stebbing, 1920, a species
known only from South Africa. Takeda &
Ananpongsuk (1991) subsequently de-
scribed a third species, D. seafdeci, from
the Andaman Sea in the Indian Ocean.

Dairoides kusei (Sakai, 1938)
(Figs. 5, 6 J—L)

Astherolambrus kusei Sakai, 1938:341, pl.
41 Figs. 5, 6.

Dairoides kusei (Sakai, 1938). 1 male (52.2 by 47.3 mm) (ZRC 1997.447).

Dairoides kusei.—Sakai, 1965:99, pl. 45
Fig. 2.—Sakai, 1976:288, pl. 97 Figs. 1,
2.

Material examined.—1 male, 2 females
(BPBM 1980.194), Oahu Island, off Bar-
ber’s Point, Hawaii, 117—128 m, trapped by
gill nets laid overnight, coll. Teritu, T.
Clarke, 19-20 Apr 1971. 1 male, cw 52.2
mm, cl 47.3 mm (ZRC 1997.447), Hawaii,
coll. E. Bilderback, 17 Jan 1979. 1 male
(dried), cw 54.8 mm, cl 44.6 mm (RMNB),
Wagu, Kii Peninsula, Mie Prefecture, Ja-
pan, coll. N. Yamashita, 1978—79. 2 males,
1 female (RMNH 32004), Wagu, Kii Pen-
insula, Mie Prefecture, Japan, coll. N. Ya-
mashita, 1978-79.

Remarks.—Dairoides kusei had _ previ-
ously been reported only from Japanese wa-
ters. The specimens reported here agree
well with the descriptions and figures of
this species by Sakai (1938, 1965, 1976).
We have also examined four specimens of
D. kusei from Japan in the RMNH donated

130 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Gls. A-C, Garthambrus stellata (Rathbun, 1906), holotype male (48.6 by 32.8 mm) (USNM 29839);
D-F G. lacunosa (Rathbun, 1906), holotype male (30.9 by 21.8 mm) (USNM 29842); G—I, G. complanata
(Rathbun, 1906), lectotype male (17.3 by 12.7 mm) (USNM 29845a); J—L, Dairoides kusei (Sakai, 1938), 1
male (52.2 by 47.3 mm) (ZRC 1997.447).

VOLUME 112, NUMBER 1

by the late Tune Sakai. They agree with the
specimens from Hawaii in all major as-
pects, including the structures of their gon-
opods.

Acknowledgments

The first author is most grateful to Lu
Eldredge for his excellent hospitality during
his visit to the BPBM. Thanks are also due
to Rafael Lemaitre and Lipke Holthuis for
their kind assistance in examining the spec-
imens in the USNM and RMNH respec-
tively. This study has been partially sup-
ported by grant RP 3972371 of the National
University of Singapore to the first author.

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ies, Taipei (in press).

White, A. 1847. Descriptions of new Crustacea from
the eastern seas.—Proceedings of the Zoologi-
cal Society of London 15:56—58.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(1):133-140. 1999.

Reinstatement and further description of Eualus subtilis Carvacho &
Olson, and comparison with E. lineatus Wicksten & Butler
(Crustacea: Decapoda: Hippolytidae)

Gregory C. Jensen and Rachel C. Johnson

(GCJ) School of Fisheries, Box 357980, University of Washington, Seattle, Washington 98195,
U.S.A.; (RCJ) National Marine Fisheries Service, Southwest Fisheries Center,
Tiburon, California 94920, U.S.A.

Abstract.—Eualus subtilis Carvacho & Olson 1984 is presently considered
a synonym of E. lineatus Wicksten & Butler 1983. Comparisons of the two
forms revealed differences in the armature of the antennular peduncle, stylo-
cerite, and pereopods, as well as differences in color and maximum size, in-
dicating that E. subtilis is a valid species. Furthermore, EF. subtilis, unlike E.
lineatus, exhibits marked sexual dimorphism, and characteristics of the previ-
ously undescribed, diminutive males of this species are provided.

The genus Eualus Thallwitz, 1891 is
comprised of relatively small caridean
shrimps that occur primarily in the higher
latitudes. Butler (1980) illustrated a small,
striped shrimp as E. herdmani (Walker,
1898), and mentioned that it could be
trawled in small numbers in Departure Bay,
Canada. However, subsequent examination
of the holotype and only known specimen
of FE. herdmani by Wicksten & Butler
(1983) revealed that E. herdmani belonged
in the genus Heptacarpus (Holmes, 1900)
and the specimens illustrated by Butler
(1980) were described as a new species, E.
lineatus Wicksten & Butler, 1983. One of
the characters distinguishing E. lineatus
was the presence of three moderate dorso-
distal spines on the first article of the an-
tennular peduncle (hereafter referred to as
“‘antennular spines’’).

A second description of a small eualid, E.
subtilis Carvacho & Olson, 1984 was pub-
lished based on a single specimen trawled
off Baja California. As Carvacho & Olson
did not mention E. lineatus, and still referred
to “E. herdmani’’, they were evidently un-
aware of Wicksten & Butler’s (1983) paper.
Eualus subtilis strongly resembled E. linea-
tus in the size and armature of the rostrum,

but had only a single, stout, dorsolateral an-
tennular spine. Wicksten (1988) considered
E. subtilis to fall within the range of varia-
tion of E. lineatus and thus a junior syno-
nym, and used the record as a southern range
extension for E. lineatus.

Subtidal sampling in the Puget Sound re-
gion revealed an extremely abundant small
eualid that matched the description of E.
subtilis. However, out of hundreds of spec-
imens examined, all had a single antennular
spine. Based on the length of the rostrum
some of these specimens keyed out to E.
pusiolus (Kr@yer, 1841), yet they bore sev-
eral strong, distal spines on the merus of
the walking legs while E. pusiolus has only
a single spine (Squires 1990). Furthermore,
this species exhibited marked sexual di-
morphism, whereas there is no distinct dif-
ferences between sexes in E. pusiolus
(Greve 1963). In view of these observa-
tions, a study was undertaken to determine
if E. subtilis is distinct from E. lineatus, and
provide information about the unusual,
small males.

Materials and Methods

Due in part to the ambiguity of existing
keys, museum specimens of these shrimps

134

have been variously cataloged under the
names E. herdmani, E. pusiolus, or E. li-
neatus. These specimens needed to be re-
examined. Sixty-one specimens from the
British Columbia Provincial Museum col-
lection were examined, as was the holotype
of E. lineatus (AHF 4129) deposited in the
Natural History Museum of Los Angeles
County and the paratype from the National
Museum of Natural History, Smithsonian
Institution, Washington, D.C. (USNM). The
two paratypes reportedly deposited in the
National Museum of Canada were not sent
at the time E. lineatus was described and
now cannot be located. Additional material
was acquired from Friday Harbor Labora-
tories of the University of Washington (43
specimens), the California Academy of Sci-
ences (8 specimens), and our own collec-
tions ranging from the Pribilof Islands to
Puget Sound (45 specimens). Twenty-two
North Atlantic specimens of E. pusiolus
from the Royal Norwegian Society of Sci-
ences were examined for comparison.

Live specimens having a single anten-
nular spine were abundant in shell rubble
and easily collected while diving by care-
fully placing dead bivalve shells in a fine-
meshed bag. Hand-operated suction devices
were used to sample in rock crevices.

Measurements were taken using an im-
age analysis system (Optimus®) on a Wild
MC3 dissecting microscope, with additional
measurements taken using an ocular micro-
meter. Carapace length (cl) was measured
from the posterior margin of the orbit to the
middorsal posterior margin of the carapace;
rostrum length, from the same position on
the orbit to the tip. Sex, rostral formula,
number of distal spines on the basal article
of the antennular peduncle, and relative
lengths of the stylocerite and rostrum were
also noted, as was the number of meral
spines on pereopods 3—5. Drawings were
made with the aid of a camera lucida on a
Wild® M5 microscope.

Results

Of the 159 shrimp measured, only 17
were found bearing multiple antennular

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

spines, and these were compared with 93
specimens that had only a single antennular
Spine and multiple spines on the merus of
the third pereopod. The remaining 49 spec-
imens were identified as E. pusiolus based
on published descriptions and comparison
with the specimens from Norway.

The 17 specimens with multiple anten-
nular spines reached substantially larger
sizes than those bearing a single spine (Fig.
la, b). Multiple antennular spines were also
correlated with a reduced number of meral
spines on the third pereopod. Those with
multiple antennular spines typically had
only one meral spine, although some spec-
imens had a second, much reduced spine
present on at least one side. Shrimp with a
single antennular spine had significantly
more meral spines (x? = 81.21; p<0.001),
the great majority bearing three strong mer-
al spines and the remainder varying from
two to five (Fig. 2).

Those with multiple antennular spines
also had a curved, dorsal tooth near the
base of the stylocerite not previously noted
in the literature (Fig. 3); that tooth was
lacking in those with a single antennular
spine. A blunt suborbital carapace spine
was also present in those with multiple an-
tennular spines, and missing in those with
a single antennular spine and more than one
spine on the merus.

Males with a single antennular spine typ-
ically had a very thin, bifid rostrum (Fig.
4c); the largest male (2.1 mm cl) was small-
er than any of the ovigerous females (2.6—
3.8 mm). Unlike that of females, the pro-
podus of pereopods 3—5 of males was dis-
tinctly broadened distally and armed with
two rows of spines forming a dense comb
on the flexor margin, with spines increasing
in length distally (Fig. 4a); the male dac-
tylus was also armed with an unusual series
of compound spines on the flexor margin
(Fig. 4b). The appendix masculina was sub-
equal in length to the appendix interna, and
tipped with two long spinules and five short
ones. Males comprised only 10% of the
samples.

112, NUMBER 1

0.90

0.80

0.70

0.60

0.50

0.9

0.8

<
fe)
:
rostrum length/carapace length rostrum length/carapace length 45

Big. J.

135

5 6 7 8

postorbital carapace length (mm)

b

5 6 7 8

postorbital carapace length (mm)

Scatterplot of postorbital carapace length vs. proportional rostrum length for specimens having a

single distodorsal spine on the basal article of the antennular peduncle (a), and specimens with multiple disto-
dorsal spines on the basal article of the antennular peduncle (b).

Only two males with multiple antennular
spines were available for examination, but
other than the presence of an appendix mas-
culina they did not appear to differ from
females with multiple spines. The appendix
masculina in these specimens was slightly
more than half the length of the appendix
interna and tipped with eight long spinules,
as previously described (Wicksten & Butler
1983; Fig. 2d).

The form with only a single antennular
spine consistently displayed the color pat-
tern described for E. lineatus (Wicksten &
Butler, 1983) as shown in Butler (1980:
plate 1C), having thin red diagonal lines on
the carapace and first two abdominal seg-
ments and red spotting on the remainder of
the abdomen. In contrast, a live specimen
of the form that has multiple antennal
spines was boldly marked on both the car-

136
80
Y
Y number of
60 Y antennular
Y spines
a. ZY multiple
5 Uy, 0D single
2 40 Y
j
a baal
Y
Y
0 1 2 3 4 5
number of leg 3 merus spines
Fig. 2. Bar graph showing number of lateral spines

on the merus of the third pereopod for specimens with
single (n = 85) or multiple (n = 12) distodorsal spines
on the first article of the antennular peduncle.

apace and abdomen with broad orange
bands against a translucent background
(Figw 5):

The holotype of E. lineatus has multiple
antennular spines, a suborbital carapace
spine, dorsal tooth on the stylocerite, and
two spines on the merus of the third pereo-
pod, whereas the USNM paratype of E. li-
neatus has only a single antennular spine,
no suborbital spine or stylocerite tooth, and
three strong spines on the merus of the third
pereopod.

Discussion

The combination of size, color and mor-
phological differences clearly indicate that
E. subtilis should be considered a valid spe-
cies, distinctly separate from E. lineatus.
Eualus subtilis lacks the multiple dorsodis-
tal spines on the basal article of the anten-
nular peduncle, the suborbital carapace
spine, and the dorsal tooth on the stylocerite
that are all present in E. lineatus. Further-
more, the largest E. subtilis barely exceed
half the length of E. lineatus, and E. subtilis
exhibits marked sexual dimorphism while
E. lineatus does not. The number of spines

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

on the merus of the third walking leg is
usually reliable for differentiating the two
species, but there is some slight overlap,
with E. subtilis varying from 2 to 5 (usually
3) spines whereas E. lineatus varies from 0
to 2 (usually 1). The type series for E. li-
neatus consists of a mix of both of these
species, but since the specimen designated
as the holotype has multiple antennular
spines and a spine on the stylocerite, this is
the form that should retain the name E. /i-
neatus.

Much of the confusion regarding these
species is due to variability in the length of
the rostrum of E. subtilis (Fig. 4c—f). Most
keys continue to follow the pattern estab-
lished by Rathbun (1904) of separating E.
pusiolus and “‘E. herdmani’” (= E. lineatus)
solely on the basis of whether the rostrum
overreaches the second article of the anten-
nular peduncle. We found many cases
where, depending on the length of the ros-
trum, specimens of E. subtilis from the
same haul had been cataloged as E. pust-
olus and E. herdmani or E. lineatus.

The presence of multiple antennular
spines makes E. lineatus very easy to dif-
ferentiate from E. subtilis and E. pusiolus.
The latter two species can be reliably sep-
arated by the number of meral spines on the
pereopods: E. subtilis has 2 to 5 strong, dis-
tal spines on the merus of the third pereo-
pod (and normally multiple spines on the
fourth), whereas E. pusiolus has only a sin-
gle spine on each of these pereopods. Eu-
alus pusiolus has a small, rounded subor-
bital carapace spine, while E. subtilis has
none. In this respect E. subtilis resembles
the South American species E. dozei (A.
Milne Edwards, 1891), but this species also
has only single meral spines (Holthuis
1952)

We found that some of the characters
given by Carvacho & Olson (1984) to dif-
ferentiate E. subtilis were not useful. The
ventral spines on the abdomen (considered
by these authors to be unique to this spe-
cies) were present on all males and most
non-ovigerous females of E. pusiolus and

VOLUME 112, NUMBER 1

137

Fig. 3.

Eualus lineatus Wicksten & Butler, 1983. Lateral view of anterior region of carapace, showing curved

tooth on the base of the stylocerite and multiple spines on the basal article of the antennular peduncle. Scale

bar is | mm.

E. lineatus. Furthermore, the basicerite of
E. pusiolus has only one (not two) lateral
spines, so this character is not useful for
separating this species from E. subtilis.
Eualus subtilis is quite possibly the most
abundant shrimp in Puget Sound, occurring
subtidally on virtually any bottom type
from mud to solid rock. It sometimes oc-
curs in the low intertidal and has been col-
lected in trawls to at least 74 m, and to date
has been found from Barkley Sound, Brit-
ish Columbia (this study) to Bahia de Todos
Santos, Punta Banda, Baja California (Car-
vacho & Olson 1984). Males of E. subtilis
are fairly uncommon in collections, proba-

bly due to their very diminutive size. Al-
though males were always much smaller
than ovigerous females, the presence of
small females suggests that the species is
not strictly protandric. The unusual modi-
fications to the pereopods bear some resem-
blance to those described by Bauer (1986)
for another small hippolytid, Thor manningi
Chace, 1972, a species that exhibits a novel
reproductive strategy involving nearly
equal proportions of protandric individuals
and primary males.

Less is known about the habits and hab-
itat of the much rarer E. lineatus. Speci-
mens have been collected at depths of 12-

138 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

z

Fig. 4. Eualus subtilis Carvacho & Olson, 1984. a, Third pereopod of male, lateral view; b, dactyl of third
pereopod of male, lateral view; c, lateral view of male carapace; d, lateral view of female carapace; e, rostrum
variation, female, lateral view; f, rostrum variation, female, lateral view.

VOLUME 112, NUMBER 1

139

Fig. 5. Eualus lineatus Wicksten & Butler, 1984. Dorsal view of live specimen (male, 3.5 mm postorbital

carapace length) showing pattern of broad orange bands.

120 m from Juneau, Alaska to at least Santa
Cruz Island, California. This species may
associate with sponges since the holotype
was collected from “‘mud and sponge bot-
tom,’’ and the trawl collection records for
the specimens examined often indicated
that the shrimp were removed from sponge
cavities, or noted the presence of sponges
in the haul. The live specimen collected by
one of us (GCJ) was found in a suction
sampler that had been used in both crevices
and small sponges on a vertical rock face,
but it is not known at which point in the
dive it was captured.

Given the confusion that has surrounded
these species, any records should be con-
sidered suspect until the specimens have
been reexamined. It is likely that at least
some of the depth and range records will
be revised, and perhaps new information on
the habitat of E. lineatus will come to light.

Acknowledgments

We are grateful for the help and support
of the staff of the Shannon Point Marine
Center, where part of this study was sup-
ported by the National Science Foundation
Research Experiences for Undergraduates
program, and J. Orensanz for the loan of
his camera lucida. The Royal Norwegian
Society of Sciences, Trondheim, provided
specimens of FE. pusiolus; K. Sendall, K.
Reid, R. Van Syoc, and J. W. Martin gra-
ciously provided specimens from their re-
spective museum collections, J. Price of the
Canadian Museum of Nature went to great
lengths to try to find the missing paratypes,
and P. Jensen provided invaluable assis-
tance in field sampling.

Literature Cited

Bauer, R. T. 1986. Sex change and life history pattern
in the shrimp Thor manningi (Decapoda: Cari-

140

dea): a novel case of partial protandric her-
maphroditism.—Biological Bulletin 170:11-—31.

Butler, T. H. 1980. Shrimps of the Pacific coast of Can-
ada.—Canadian Bulletin of Fisheries and
Aquatic Sciences 202:1—280.

Carvacho, A., & Y. R. Olson. 1984. Nuevos registros
para la fauna carcinoldgica del noreste de Méx-
ico y descripcién de una nueva especie: Eualus
subtilis, n.sp. (Crustacea: Decapoda: Natan-
tia)—The Southwestern Naturalist 29(1):59—
g cil

Chace, FE A., Jr. 1972. The shrimps of the Smithsonian-
Bredin Caribbean expeditions with a summary
of the West Indian shallow-water species (Crus-
tacea: Decapoda: Natantia) Smithsonian Con-
tributions to Zoology 98:1—179.

Greve, L. 1963. The genera Spirontocaris, Lebbeus,
Eualus and Thoralus in Norwegian waters
(Crustacea, Decapoda).—Sarsia 11:29—42.

Holmes, S. J. 1900. Synopsis of California stalk-eyed
Crustacea—Occasional Papers of the California
Academy of Sciences 7:1—262.

Holthuis, L. B. 1952. The Crustacea Decapoda Macru-
ra of Chile-—Reports of the University of Chile
Expedition 1948—49, 5:1—110.

Krgyer, H. 1841. Udsigt over de nordiske Arter af
Slaegten Hippolyte.—Naturhistorisk Tidsskrift
3:570—-579.

Milne Edwards, A. 1891. Crustaces. Mission scienti-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

fique du Cap Horn 1882-1883, volume 6, Zo-
ology, part 2F:1—54.

Rathbun, M. J. 1904. Decapod crustaceans of the
northwest coast of North America.—Harriman
Alaska Expedition Series 10:1—210.

Squires, H. J. 1990. Decapod Crustacea of the Atlantic
coast of Canada.—Canadian Bulletin of Fish-
eries and Aquatic Sciences 221:1—532.

Thallwitz, J. 1891. Decapoden-Studien, ibessondere
basirt auf A. B. Meyer’s Sammlungen im Ostin-
dischen Archipel, nebst einer Aufzahlung der
Decapoden und Stomatopoden des Dresdener
Museums. Abhandlungen und Berichte des
K6niglichen Zoologischen und Anthropolo-
gisch-Ethnographischen Museums zu Dresden
1890—91(3):1-55.

Walker, A. O. 1898. Crustacea collected by W. A.
Herdman in Puget Sound, Pacific Coast of
North America, Sept. 1897.—Transactions of
the Liverpool Biological Society 12:268—287.

Wicksten, M. K. 1988. New records and range exten-
sions of shrimps and crabs from California,
U.S.A. and Baja California, Mexico.—Califor-
nia Fish and Game 74(4):236—248.

, & T. H. Butler. 1983. Description of Eualus

lineatus new species, with a redescription of

Heptacarpus herdmani (Walker) (Caridea: Hip-

polytidae). Proceedings of the Biological Soci-

ety of Washington 96:1—6.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(1):141-144. 1999.

Deilocerus captabilis, a new species of cyclodorippid crab from
southeastern Brazil (Crustacea: Decapoda: Brachyura:
Cyclodorippidae)

Marcos Tavares

Universidade Santa Ursula, ICBA, Rio de Janeiro 22231-040, Brasil

Abstract.—Deilocerus captabilis, new species, the third western Atlantic
species and seventh representative of the genus is herein described based on
an adult female collected on calcareous algae nodules from 50 m depth, south-
eastern Brazil. Deilocerus captabilis, new species, is the only species in the
genus with three teeth on the lateral margin of the carapace.

Resumo.—Deilocerus captabilis nova espécie, terceira a ocorrer no Atlantico
sul ocidental das sete conhecidas para o género, é descrita e ilustratada. A nova
espécie €é baseada em uma fémea adulta coligida a 50 m de profundidade em
fundos de nddulos de algas calcareas. A nova espécie se distingue facilmente
das demais espécies do género por ser a Unica a apresentar trés dentes na

margem lateral da carapaga.

As part of an ongoing project (REVI-
ZEE) of the Brazilian government’s depart-
ment “‘Ministério do Meio Ambiente’’ to
evaluate the country’s marine living re-
sources, a biological survey of the conti-
nental shelf and slope from Salvador (12°S)
to Cabo de S40 Tomé (23°S) was conducted
in 1997 onboard the “‘Astro Garoupa’’.

The samples yielded an adult female of
a new species of cyclodorippid crab, Deil-
ocerus captabilis, described herein.

The holotype is deposited in the Museu
Nacional, Rio de Janeiro (MNRJ). Descrip-
tive terminology follows Tavares (1991,
1996). Abbreviations: Mxp1-3, first to third
maxillipeds; P2—P5, second to fifth pereo-
pods, P1, cheliped; cl, carapace length; cw,
carapace width; mm, millimeters.

Deilocerus captabilis, new species
Fig. 1

Material examined.—Brazil: Espirito
Santo. “‘Astro Garoupa’’, REVIZEE Cen-
tral II, st. 34C, 3 Nov 1997, 20°24’S,
39°49'W, 50 m: female holotype cl 1.7 mm,
cw 2.0 mm (MNRJ 7303).

Type locality.—Brazil: Espirito Santo
(20°24’S, 39°49’W, 50 m).

Description.—Carapace slightly broader
than long. Dorsal surface ornamented with
very fine scattered granules, denser near
margins, except on smooth, shallow
grooves defining gastric regions. Ventrolat-
eral surfaces of carapace almost smooth;
subhepatic region densely covered with
rounded coarse granules, coarser on dorsal
surface. Frontolateral tooth rounded, dense-
ly covered with small granules; exorbital
tooth blunt. Hepatic and anterolateral teeth
short, blunt, covered with rounded granules;
anterolateral tooth smallest. Laterobranchial
tooth present as low lobe. Anterolateral
margin (from exorbital tooth to branchial
tooth) rounded, about as long as posterolat-
eral margin (measured from branchial tooth
to posterior margin). Posterolateral margin
Straight, well defined by row of small gran-
ules.

Ocular peduncle covered with small
rounded tubercles, anterodistal tubercles
more acute; cornea pigmented. Antenna
very small, hidden in dorsal view; articles

142 PROCEEDINGS

OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Biss ls.

Deilocerus captabilis, new species, Espirito Santo, Brazil, 20°24’S, 39°49’W, 50 m: female holotype

cl 1.7, cw 2.0 mm (MNRJ 7303). A, dorsal view of whole crab. B, right cheliped outer view.

1—4 ornamented with small granules; sec-
ond article slightly flattened; fifth article al-
most smooth; flagellum obsolete.
Endostomial channel visible dorsally be-
tween frontolateral teeth. Third maxilliped
with ischium and merus, each more than 2

times as long as broad, with outer surfaces
covered with rounded granules; palp with-
out granules, articulated on inner surface of
merus.

Chelipeds densely covered with small tu-
bercles, much less dense and smaller on in-

VOLUME 112, NUMBER 1

ner surfaces; inner surfaces of merus, car-
pus and chela forming concave surface fit-
ting closely against walls of carapace; dor-
sal and ventral margins of merus, carpus,
palm and fingers well defined. Fingers ter-
minating in sharp tips, cutting edge with
few small acute teeth. Dactyl smaller than
palm, set obliquely relative to palm axis.
Fixed finger about 2 times broader proxi-
mally than distally. Palm about 2 times lon-
ger than broad, ornamented longitudinally
with 2 rounded protuberances, proximal
one largest.

P2 longer than P3, otherwise similar. P2
and P3 laterally flattened; both legs with
propodus, carpus, and merus densely orna-
mented with tubercles on dorsal and ventral
margins, flanks almost smooth; dactyl cy-
lindrical, with minute granules.

P4 and P5 generally similar, subdorsal,
subcheliform, much smaller than P2 and
P3; P4 with ischium about 3 times longer
than in P5; dactylus and propodus short,
strongly curved; propodus twisted, dactylus
flexing on its lateral surface. Ornamentation
on propodi, carpi and meri less pronounced
on P4—5 than on P2-3.

Female abdomen with 6 segments dense-
ly covered with small tubercles diminishing
in size and density from pleotelson to first
segment. Pleotelson as wide as fifth seg-
ment, lateral margins broadly rounded.

Distribution.—Known only from the
type-locality Espirito Santo, Brazil
(20°24'S, 39°49’'W, 50 m).

Etymology.—The specific name, capta-
bilis (Latin, that can take), refers to the sub-
cheliform P4 and PS.

Remarks.—The genus Deilocerus Tava-
res, 1993, is strictly American in distribu-
tion. The genus is represented in the west-
ern Atlantic by two species, and in the east-
ern Pacific by four (Tavares 1993, 1996).
Two groups of species are recognizable.
The first group includes species with only
one anterolateral tooth on the margin of the
carapace: D. perpusillus (Rathbun, 1900)
and D. analogus (Coelho, 1973), from the
western Atlantic, and D. laminatus (Rath-

143

bun, 1935) from the eastern Pacific. The
second group is found only in the eastern
Pacific and encompasses species with two
teeth (one hepatic and one anterolateral),
along the margin of the carapace: D. planus
(Rathbun, 1900), D. decorus (Rathbun,
1933) and D. hendrickxi Tavares, 1993.
Deilocerus captabilis, new species, falls
into the second group and is, therefore, the
first western Atlantic representative of that
group. Deilocerus captabilis, new species
is, so far, the only species in the genus with
three teeth on the lateral margin of the car-
apace (hepatic, anterolateral, and latero-
branchial): the first two teeth being well de-
veloped, and the laterobranchial represented
by a low lobe. Deilocerus captabilis, new
species, can also be readily recognized from
the remaining six species of the genus by
the shape and ornamentation of the fronto-
orbital margin of the carapace.

Although no carrying behavior (Guinot
et al. 1995) has been observed in Deiloce-
rus captabilis, new species, it is possible
that the crab can hold an object over its
carapace using the last two pairs of legs. P4
and P5 are subchelate, with the propodus
and especially the dactyl strongly curved;
the propodus is twisted so the dactyl closes
on the lateral surface of the propodus. This
structure allows a small object, such as a
piece of shell, to be held over the carapace.
Carrying behavior has been observed in
other species of the genus, such as D. lam-
inatus (see Garth 1946, Tavares 1994) and
D. planus (see Schmitt 1921, Wicksten
1982).

Acknowledgments

I am grateful to Daniéle Guinot (Muséum
national d’ Histoire naturelle, Paris), Rafael
Lemaitre (Smithsonian Institution, Wash-
ington, D.C.) and Peter K. L. Ng (National
University of Singapore) for reading a draft
of the manuscript and providing helpful
suggestions, and to Maria Helena Pinheiro
(Universidade Santa Ursula) for preparing
drawings. This research has been funded by

144 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Ministério do Meio Ambiente, SECIRM,
Programa REVIZEE, and Universidade
Santa Ursula, Rio de Janeiro. Thanks are
due also to CNPq for supporting studies on
the systematic of decapod crustaceans in the
form of ongoing grant 30.09.15/97-—7.

Literature Cited

Coelho, P. A. 1973. Descri¢ao preliminar de Clythro-
cerus analogus, n. sp., do litoral brasileiro
(Crustacea, Decapoda, Dorippidae).—Ciéncia e
Cultura 25(6)(suppl.):343-344.

Garth, J. S. 1946. Littoral brachyuran fauna of the Gal-
apagos Archipelago.—Allan Hancock Pacific
Expedition 5(10):i-iv + 341-600.

Guinot, D., D. Doumenc, & C. C. Chintiroglou. 1995.
A review of the carrying behaviour in Brachy-
uran crabs, with additional information on the
symbioses with sea anemones.—Raffles Bulle-
tin of Zoology 43(2):377—416.

Rathbun. M. J. 1900. Synopses of North American in-
vertebrates. X. The Oxyrhynchous and Oxysto-
matous crabs of North America.—American
Naturalist 34:503—520.

. 1933. Preliminary descriptions of nine new

species of oxystomatous and allied crabs.—Pro-

ceedings of the Biological Society of Washing-

ton 46:183—186.

. 1935. Preliminary descriptions of seven new

species of oxystomatous and allied crabs.—Pro-

ceedings of the Biological Society of Washing-
ton 48:1—4.

Schmitt, W. L. 1921. The Marine Decapod Crustacea
of California with special reference to the Deca-
pod Crustacea collected by the United States
bureau of Fisheries Steamer “‘Albatross”’ in
connection with the Biological Survey of San
Francisco Bay during the years 1912—1913.—
University of California Publications in Zoolo-
gy 23:1—470.

Tavares, M. 1991. Révision préliminaire du genre Ty-
molus Stimpson, avec la description de Tymolus
brucei sp. nov. d’ Australie occidentale (Crus-
tacea, Brachyura, Cyclodorippoidea).—Bulletin
du Muséum national d’Histoire naturelle, Paris
(4), 13, sect. A, (3—4):439—456.

. 1993. Crustacea Decapoda: Les Cyclodorip-

pidae et Cymonomidae de |’ Indo-Ouest-Paci-

fique a l’exclusion du genre Cymonomus. In A.

Crosnier, ed., Résultats des Campagnes MU-

SORSTOM, 10.—Mémoires du Muséum na-

tional d’Histoire naturelle, Paris 156:253-313,

Fig. 1-20.

. 1994. Description préliminaire de quatre nou-

veaux genres et trois nouvelles espéces de Cy-

clodorippoidea américains.—Vie Milieu 43(2—

3):137-144.

. 1996. Révision systématique des Cyclodor-
ippidae américains (Crustacea, Decapoda,
Brachyura).—Bulletin du Muséum national
d’Histoire naturelle, Paris (4), 18, sect. A, (1—
2):233-295.

Wicksten, M. K. 1982. Behavior in Clythrocerus plan-
us (Rathbun, 1900) (Brachyura, Dorippidae).—
Crustaceana 43(3):306—307.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(1):145-163. 1999.

The Albuneidae (Decapoda: Anomura: Hippoidea) of the Hawaiian
Islands, with description of a new species

Christopher B. Boyko

Department of Invertebrates, American Museum of Natural History, Central Park West at 79th
St., New York, New York 10024, U.S.A. and Department of Biological Sciences, University of
Rhode Island, Kingston, Rhode Island 02881, U.S.A.

Abstract.—Albunea danai, a new species that has been confused with A.
speciosa Dana, is described from Oahu, Hawaii. The new species actually
resembles A. carabus (Linnaeus) from the Mediterranean and western Africa.
The discovery of this new species brings the total number of Indo-West Pacific
members of the genus to eight. Albunea speciosa is broadly distributed in the
Indo-West Pacific and is not a Hawaiian endemic as previously believed. As
the identities of the two Hawaiian species have been confused, A. speciosa is
here redescribed from new material and a neotype is selected. Both A. danai
and A. speciosa are discussed using newly defined morphological characters.

Albunea speciosa Dana, 1852, is the only
species of albuneid heretofore known from
the Hawaiian Islands and has been consid-
ered a Hawaiian endemic (Seréne 1973). In
the course of examining specimens of sand
crabs for a worldwide revision of the family
Albuneidae, material from Hawaii was
found labeled ‘‘Albunea speciosa’’ and
*‘“Albunea symnista’’ [sic] in the collections
of the Western Australian Museum (WAM),
but that was clearly not conspecific with A.
speciosa. Later, additional Hawaiian speci-
mens were obtained from the Bernice P.
Bishop Museum that were labeled ‘“‘Albu-
nea thurstoni’’ that clearly are not A. thur-
stoni Henderson, 1893, but instead are the
same taxon as the WAM material. Since
these specimens cannot be placed in any
known species of albuneid, they are de-
scribed here as a new species. In addition,
my examination of specimens that are re-
ferable to A. speciosa revealed that this spe-
cies is not endemic to Hawaii, but has a
broad range in the Indo-West Pacific. Be-
cause the faunal composition and biogeog-
raphy of island groups has attracted much
attention of late and the fact that more than
one species has been repeatedly identified

with A. speciosa, that species is redescribed
from new material and a neotype is desig-
nated herein in order to fix its identity. This
description of the new species brings the
total number of Indo-West Pacific species
of Albunea to eight: A. symmysta (Linnae-
us, 1758), A. speciosa, A. microps Miers,
1878, A. thurstoni, A. elioti Benedict, 1904,
A. steinitzi Holthuis, 1958, A. madagascar-
iensis Thomassin, 1973, and A. danai new
species.

Materials, Methods, and Morphological
Terminology

Materials.—Specimens for this study
came from the collections of the American
Museum of Natural History, New York
(AMNH), Bernice P. Bishop Museum, Hon-
olulu, Hawaii (BPBM), California Acade-
my of Sciences, Invertebrate Zoology, San
Francisco (CASIZ), Musée Royal de
|’ Afrique Centrale, Tervuren, Belgium
(MRAC), Queensland Museum (QM), Uni-
versity Museum of Zoology, Cambridge
(UMZC), National Museum of Natural His-
tory, Smithsonian Institution, Washington,
D.C. (USNM), Western Australian Muse-

146

um, Perth (WAM), and Yale Peabody Mu-
seum, New Haven, Connecticut (YPM).

Methods.—Carapace length (CL), as
measured from the midpoint of the anterior
margin (including rostrum) to the midpoint
of the posterior concavity, is provided as an
indicator of specimen size. In the list of
synonyms, asterisks refer to publications
citing material examined during the present
study. Absence of an asterisk in a specific
entry does not imply that the identifications
therein are in doubt, but only that it was not
possible to examine the material cited in
that publication.

Illustrations were created using a modi-
fied approach of Harvey & De Santo
(1997). Specimen images were first cap-
tured on a Macintosh® computer with a
digital camera connected to a Wild M8 dis-
secting microscope. These images were
then prepared using the programs Adobe
Photoshop™ and Adobe Illustrator®. I at-
tempted to record the position and size of
setae in these drawings as accurately as
possible, although for clarity of presenta-
tion the plumules of plumose setae were not
drawn.

Morphological terminology.—During the
course of this study and that of Boyko &
Harvey (in press), several important diag-
nostic morphological features were encoun-
tered that have not been described previ-
ously for albuneids. Although these, along
with other features, are discussed in Boyko
& Harvey (in press), they are also consid-
ered here to facilitate a clearer understand-
ing of the descriptive terms used in this pa-
per.

The front of the carapace of albuneids
bears a broad mat of very short, dense, sim-
ple setae. This mat, hereafter called the se-
tal field, varies in shape and extent across
genera and species, but appears to be rela-
tively invariant within species. The cara-
pace also possesses numerous transverse,
setose grooves. Although carapace grooves
(CG) have been scarcely mentioned by pre-
vious authors, 11 major grooves (numbered
1-11, Fig. 1) have been identified which

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Setal field

Fig. 1. Diagrammatic albuneid carapace based on
Albunea microps Miers, 1878, showing setal field and
setose carapace grooves (CG 1-11) discussed in text.

can be recognized across albuneid genera.
Variability in the presence and the degree
of fragmentation of specific grooves, in the
anterior-posterior displacement of individ-
ual fragments, and in the texture of the
grooves (e.g., smooth, crenulate) tends to
be conservative within species, and thus
carapace grooves are useful in recognizing
species.

Because several of the specimens exam-
ined during this study were incorrectly
sexed, brief remarks on the determination
of male and female identity in albuneids are
presented here. As in most decapod crus-
taceans, albuneid females have gonopores
on the coxae of the third pereopods, where-
as males have gonopores on the coxae of
the fifth pereopods. However, in some al-
buneid genera (e.g., Lepidopa Stimpson,
1858 and some Albunea, including both
species from Hawaii), males also have a
small pore on the coxa of the third pereo-
pod in a position analogous to that of the
female gonopore. The precise nature and
function of this pore is unknown.

In albuneids, females have well devel-
oped uniramous pleopods on abdominal so-

VOLUME 112, NUMBER 1

Fig. 2.
view, showing terms used in species accounts for land-
marks on pereopod dactyli.

Pereopod II dactyl of Albunea sp., lateral

mites II-V. Male albuneids have tradition-
ally been considered to lack pleopods (Ef-
ford & Haig, 1968). However, rudimentary
or small pleopods were found on abdominal
somites II-V of male specimens with well
developed gonopores on the fifth pereopods
in species of several albuneid genera (e.g.,
Lepidopa). In some Albunea species (e.g.,
A. microps, A. speciosa), specimens with
large pores on the fifth pereopods and with
small pores on the coxae of the third pereo-
pods, show no signs of pleopod develop-
ment. In those species in which the male
pore occurs, it is always smaller than gon-
opores of same-sized females; likewise, the
pleopods of females are always much more
developed than those of males. Males are
most reliably recognized by the presence of
a gonopore on the fifth pereopod and the
rudimentary degree of development of the
pleopods or lack thereof. In small speci-
mens, however, the presence or absence of
the male gonopore is a more reliable indi-
cator of sex than is pleopod development
because both males and females may have
small pleopod buds as juveniles.

The shape of the dactylus of the pereo-
pods, particularly the third pereopod, has
been used to distinguish among species of
albuneids. To facilitate the description of
the complex shape of this segment, several
terms are used to refer to important land-
marks (Fig. 2). The “‘base’’ of the dactylus
is the ventroproximal angle; the “‘heel”’ cor-
responds to the dorsoproximal angle, which

147

is often strongly produced. The dorsal mar-
gin is almost always concave, sometimes
smoothly so; in most species, however, the
dorsal margin has a distinct angle, the apex
of which is referred to as an “‘indent.’’ The
dactylus terminates in a “‘tip,’’ which is
somewhat rounded and lacking in a corne-
ous nail.

In some species of albuneids, certain seg-
ments of the pereopods bear a large trans-
parent, decalcified area, hereafter called the
‘“‘window,”’ that has not been previously re-
ported in this family. This area, when pre-
sent, is most prominent on the lateral sur-
face of the merus, where it is comparable
to the “‘leg membranes”’ of porcelain crabs
(Porcellanidae) discussed in detail by Still-
man & Somero (1996). These windows can
also be found to a lesser degree on other
pereopod segments, both laterally and me-
sially.

Superfamily Hippoidea Latreille, 1825
Family Albuneidae Stimpson, 1858
Albunea Weber, 1795
Albunea speciosa Dana, 1852
Figs. 3, 4

Albunaea speciosa Dana, 1852: 405—406;
1855: pl. 25, figs. 6a—f.—Stimpson,
1858: 230 (list).

Albunea speciosa.—Miers, 1878: 315 (list),
331 (after Dana, 1852).—Ortmann, 1896:
223 (key), 225 (list), 239 (table).—Gor-
don, 1938: 187 (list).—Edmondson,
1946: 266.—Seréne, 1973: 262-263, pl. 2.

‘““?Albunea_ speciosa’’.—Borradaile, 1904:
fos eae

Material examined.—Neotype: Hawaii,
Oahu, 27 May 1938: 1 male, 9.0 mm CL
(USNM 260868); same data as neotype: 2
males, 9.5—10.4 mm CL, 1 female, 14 mm
CL (USNM 287087); Kailua, Oahu, Mar
P9382) } “male, 104. sam “CL. (BPBM
S$11781); Honolulu Harbor, Oahu, coll. E.
M. Ehrhon, Dec 1916: 2 males, 6.4—9.9 mm
CL, 1 female, 7.3 mm CL (CASIZ 109240);
Halonu Blow Hole dive site, south shore,
Oahu, coll. R. Holcom, 3 Aug 1997: 1 fe-

148 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Albunea speciosa Dana, 1852. A, male, 9.0 mm CL, USNM 260868, neotype; B—J, male, 10.4 mm
CL, USNM 287087. A, carapace and eyes, dorsal view; B, eyes, dorsal view; C, left antennule, lateral view; D,
left antenna, lateral view; E, left mandible; mesial view; F right maxilla, lateral view; G, left maxillule, lateral

view; H, left maxilliped I, lateral view; I, right maxilliped II, lateral view; J, left maxilliped III, lateral view.
Scale = 1.6 mm (B, E, G), 2.2 mm (1), and 3.3 mm (A; C, D, E H, 3).

VOLUME 112, NUMBER 1

149

Fig. 4. Albunea speciosa Dana, 1852. A-E, male, 10.4 mm CL, USNM 287087; E male, 9.0 mm CL, USNM
260868, neotype; G, female, 14 mm CL, USNM 287087. A, right pereopod I, lateral view; B, right pereopod
II, lateral view; C, right pereopod III, lateral view; D, right pereopod IV, lateral view; E, abdominal somites I—
VI, dorsal view; E telson of male, dorsal view; G, telson of female, dorsal view. Scale = 3.0 mm (KE G), 4.0

mm (A-—D), and 4.2 mm (BE).

male, 6.6 mm CL, 1 broken unsexable/un-
measurable specimen (QM W22284); Hal-
onu Blow Hole dive site, south shore,
Oahu, 12.2—13.7 m, coll. R. Holcom, 4 Apr
1997: 2 males, 6.4—7.1 mm CL, 3 females,
5.7-9.5 mm CL, 6 juveniles, 3.3—4.1 mm
CL (QM W22285); Oahu, coll. R. Holcom,
Apr 1997: 2 ovigerous females, 7.5 mm CL
(QM W22286); ‘‘Hawaii,’’ coll. C. M.
Cook Jr., 1897: 1 female, 10 mm CL (YPM
P1933):

Australia: Bay on north side of Point

Cloates, lee side of reef, Western Australia,
113°38’E, 22°41'S, depth 3.7 m, coll. WAM
Ningaloo Expedition, 23 Aug 1968: 1 male,
8.7 mm CL, 1 carapace, 9.1 mm CL (WAM
489-97); southwest of Point Cloates, West-
ern Australia, 113°39'30”E, 22°43'30’S,
coll. WAM Ningaloo Expedition, 7 Sep
1968: 1 ovigerous female, 9.7 mm CL
(WAM 490-97).

Seychelles: Mahé, coll. Mission Zoolo-
gique MRAC-ULB, July—Sep 1966: 1 ovig-
erous female, 9.9 mm CL (MRAC 53.894).

150

Maldives: Hulule, Male Atoll, coll. J. S.
Gardiner: 1 male, 8.4 mm CL (UMZC).

Type locality.—Sandwich Islands (= Ha-
waiian Islands) (Dana, 1852) herein re-
stricted by neotype selection to Oahu, Ha-
waii, USA, Pacific Ocean.

Type material.—lIt is unclear exactly how
many specimens of this species Dana
(1852) had before him when writing the de-
scription of this species, but the description
suggests that there was only one. No type
material of this species is extant in either
USNM, the Museum of Comparative Zo-
ology, Harvard University, or the British
Museum (Natural History) (Evans 1967;
Boyko, pers. obs.). Because these institu-
tions are the only known repositories for
Dana’s extant type material (Evans 1967),
the type material of A. speciosa must be
considered lost. In light of the new infor-
mation about the range of A. speciosa, and
the discovery of a new species of albuneid
in the Hawaiian Islands which has been re-
peatedly misidentified as A. speciosa, it is
appropriate to follow the suggestion of Se-
rene (1973) and select a neotype for the
species. A male, 9.0 mm CL (USNM
260868) is herein designated as the neotype
for Albunea speciosa.

Diagnosis.—Carapace slightly longer
than wide, covered with strongly setose
grooves. Anterior margin with 13-17 teeth
on either side of ocular sinus. Setal field
with narrow lateral elements and concave
anterior margin; posterior lateral elements
extending to posterior lateral elements of
CG1.CG1 with separate posterior lateral el-
ements but with anterior and posterior ele-
ments united by posterior elements of setal
field; CG4 with 2-3 short anteriorly dis-
placed medial elements; CG5 entire, nearly
reaching margins of CG6; CG6 and CG7
separate; CG8 with 1—2 median elements
separated from lateral elements; CG11 pres-
ent. Rostrum reaching just beyond proximal
margin of ocular plate. Ocular plate sub-
quadrate. Ocular peduncles dorsoventrally
flattened and elongate, rounded at tip, ap-
proximate along mesial margin; lateral mar-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

gin concave; mesial margin straight. Cornea
at tip of ocular peduncle. Antennule with
48-53 flagellar exopod and 2 endopodal
segments. Antenna with 5—6 flagellar seg-
ments; acute spine on dorsolateral surface
of peduncle segment I. Dactyli of pereo-
pods II, III with heels low and smoothly
rounded. Coxa of pereopod III of males
with small male pore. Telson of male spat-
ulate, laterally expanded, dorsoventrally
flattened; produced slightly at tip. Telson of
female flattened, rounded at tip.
Redescription.—Carapace (Fig. 3a)
slightly wider than long. Anterior margin
concave on either side of ocular sinus, be-
coming convex laterally with 10—12 large
and 3—4 small spines along length. Rostrum
a small acute tooth, reaching just beyond
proximal margin of ocular plate. Ocular si-
nus smoothly concave, unarmed. Frontal re-
gion smooth; setal field broad posteriorly,
narrowing anteriorly, with narrow anterior
lateral elements and concave anterior mar-
gin; posterior lateral elements reaching to
posterior lateral elements of CG1. Medial
portion of CG1 parallel to anterior margin
of carapace, sinuous, slightly crenulate, di-
vided into medial fragment and curved pos-
teriorly-displaced lateral elements, but with
medial and lateral elements connected by
posterior lateral elements of setal field. Me-
sogastric region smooth; CG2 short, with
1—2 elements; CG3 broken into 2 longer
lateral elements and 1-3 short medial ele-
ments; CG4 with 2—3 short medial elements
displaced anteriorly, with gap at midline be-
tween short elements. Hepatic region
smooth with long setose groove at median
of lateral margin. Epibranchial region ap-
proximately triangular, smooth; posterolat-
eral margin with 3 short rows of setae. Me-
tagastric region smooth; CG5 ranging from
entire to 4 elements, nearly reaching mar-
gins of CG6. CG6 strongly crenulate,
strongly anteriorly concave medially and
sloping out to anteriorly convex lateral
thirds. CG7 nearly straight relative to an-
terior margin of carapace and separate from
CG6. Cardiac region smooth; CG8 with 1-—

VOLUME 112, NUMBER 1

2 median elements separated from lateral
elements. CG9 present as 2 lateral grooves
with short gap at midline. CG10 present as
2 curved lateral fragments, with gap be-
tween fragments about half length of single
fragment. CG11 present. Branchial region
with numerous short, transverse rows of se-
tae. Posterior margin deeply and evenly
convex, with submarginal groove reaching
about half-way either side of posterior con-
cavity. Branchiostegite with short anterior
submarginal spine; anterior region with
scattered short transverse lines ventral to
linea anomurica; with many short rows of
setae and sparsely covered with long plu-
mose setae ventrally; posterior region mem-
branous, with numerous irregular frag-
ments, and sparsely covered with long plu-
mose setae.

Ocular plate (Fig. 3b) subquadrate, with
broad median indentation; proximal ocular
segments (Fig. 3b) reduced to small round-
ed calcified area on either side of ocular
plate. Ocular peduncles (Fig. 3b) elongate,
with medially concave lateral margins, ta-
pering to rounded distal corneae; mesial
margins approximated along entire length;
mesial and ventral margins of peduncle
with sparse row of long plumose setae; tuft
of plumose setae at proximal lateral ventral
angle.

Antennule (Fig. 3c) segment III narrow
proximally, expanding distally to twice
proximal width; with plumose setae on dor-
sal and ventral margins; dorsal exopodal
flagellum with 48-53 segments and long
plumose setae on dorsal and ventral mar-
gins; ventral endopodal flagellum short,
with 2 segments and plumose setae on dor-
sal and ventral margins. Segment II medi-
ally inflated in dorsal view, with plumose
setae on dorsal and ventral margins, and se-
tae scattered on ventrolateral third of sur-
face. Segment I wider than long, unarmed;
dorsal third of lateral surface faintly rugose
with long plumose setae; long plumose se-
tae on dorsal and ventral margins.

Antenna (Fig. 3d) segment V about 3
times longer than wide, with long plumose

151

setae on dorsal and ventral margins; flagel-
lum 5—6 segmented, with long plumose se-
tae on dorsal, ventral and distal margins.
Segment IV expanded distally, with long
plumose setae on dorsal, ventral and distal
margins, and row of setae on dorsolateral
margin. Segment III with long plumose se-
tae on ventral margin. Segment II short,
widening distally, with plumose setae on
margins; antennal acicle long, thin, reach-
ing to distal margin of segment IV, with
long plumose setae on dorsal margin. Seg-
ment I rounded proximally, flattened ven-
trolaterally with long plumose setae on
margins; lateral surface with acute spine
dorsally, with low semicircular dorsolateral
lobe ventrodistal to spine; segment with
ventromesial antennal gland pore.

Mandible (Fig. 3e) incisor process with
1 tooth; cutting edge with 1 tooth. Molar
process with 1—2 teeth. Palp 3-segmented,
with plumose setae on margins and long,
thick, simple setae arising from bend in sec-
ond segment.

Maxilla (Fig. 3f) exopod evenly rounded,
with plumose setae along distal margin.
Scaphognathite bluntly angled on posterior
lobe, with plumose setae. Endopod and en-
dites without distinctive characters.

Maxillule (Fig. 3g) distal endite proxi-
mally narrow, widening to inflated distal
end, with thick simple setae on distal mar-
gin. Proximal endite with thick simple setae
on distal margin. Endopodal external lobe
truncate distally and curled under; internal
lobe reduced, with 3 thick setae at distola-
teral margin.

Maxilliped I (Fig. 3h) epipod with plu-
mose setae on distal margin and distolateral
surface. Endite tapered distally, subequal to
first segment of exopod. Exopod with 2
segments; proximal segment narrow, mar-
gins parallel and with plumose setae; distal
segment spatulate, about as long as wide,
broadest medially, margins with long plu-
mose setae. Endopod flattened and elon-
gate, reaching to distal end of proximal ex-
opodal segment; with plumose setae on
margins.

152

Maxilliped II (Fig. 31) dactylus evenly
rounded, length equal to width, with thick
simple setae distally. Propodus 1.5 times
wider than long, with plumose setae on dor-
sal margin and long simple setae on distal
margin. Carpus not strongly produced dor-
sodistally, about 2 times longer than wide,
with long simple setae on dorsal margin.
Merus about 3 times longer than wide, mar-
gins parallel, with simple setae on ventro-
lateral margin and plumose setae on dor-
solateral margin. Basi-ischium incomplete-
ly fused, with plumose setae on margins.
Exopod % times longer than merus, with
flagellum 1-segmented.

Maxilliped III (Fig. 3j) dactylus evenly
rounded; with long plumose setae on dorsal
margin and lateral surface. Propodus with
longitudinal median row of plumose setae
on lateral surface; dorsal margin with plu-
mose setae. Carpus slightly produced onto
propodus; lateral surface with row of plu-
mose setae ventromedially; plumose setae
on dorsal margin. Merus unarmed, with
plumose setae on all margins. Basi-ischium
incompletely fused, without crista dentata.
Exopod 2-segmented, proximal segment
small; distal segment styliform, tapering,
approximately % length of merus, with plu-
mose setae scattered on surface; without
flagellum.

Pereopod I (Fig. 4a) subchelate. Dactylus
curved and tapering; lateral and mesial sur-
faces smooth; dorsal margin with long plu-
mose and short simple setae; ventral margin
with short simple setae. Propodus lateral
surface with numerous short, transverse
rows of setose rugae; dorsal margin un-
armed; ventral margin produced distally
into acute spine; cutting edge lacking teeth,
lined with long plumose setae; dorsal mar-
gin with long plumose setae, ventral margin
with short simple setae. Carpus dorsodistal
angle produced into strong corneous-tipped
spine, dorsal margin with few large and
small spines posteriorly along distal third;
dorsal and distal margins with long plu-
mose setae; lateral surface with small distal
rugose area, with few transverse setose

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ridges on distal half of surface; mesial sur-
face smooth with few scattered rows of
long plumose setae, dorsal and ventral mar-
gins with long plumose setae. Merus un-
armed; lateral surface with scattered trans-
verse rows of long plumose setae, dorsal,
ventral and distal margins with long plu-
mose setae; mesial side with few short rows
of setae. Basi-ischium incompletely fused,
unarmed. Coxa unarmed.

Pereopods II-IV dactyli laterally com-
pressed and dorsoventrally expanded.

Pereopod II (Fig. 4b) dactylus smooth;
with base to heel straight, heel smoothly
rounded, heel to tip with rounded broad in-
dent, tip acute, tip to base broadly convex;
lateral surface smooth, with several small
tufts of short setae in approximately straight
line across medioproximal surface, several
widely-spaced submarginal tufts of short
setae dorsodistally; mesial surface smooth,
ventral margin with long plumose setae,
dorsal margin with short simple setae, with
patch of long plumose setae at base. Pro-
podus dorsal surface smooth, ventral mar-
gin inflated and rounded; oblique row of
long plumose setae on distal margin of lat-
eral surface; distal and ventral margins with
long plumose setae; dorsolateral surface a
narrow, oblique, flattened shelf, with short
setae on dorsal margin and long plumose
setae on ventral margin; mesial surface with
elevated, curved, setose ridge from ventral
junction with dactylus almost to ventral
proximal junction with carpus. Carpus
slightly produced dorsodistally; lateral sur-
face nearly smooth, with irregular broken
row of rugae and submarginal elevated
ridge ventrally, rugae and ridge with long
plumose setae; dorsodistal projection with
mat of short setae on lateral surface; mar-
gins with long plumose setae; mesial sur-
face smooth with long plumose setae in
scattered patches on surface, and on mar-
gins. Merus lateral surface with large de-
calcified window and few scattered setae on
surface and margins; mesial surface nearly
smooth with few setae. Basi-ischium in-

VOLUME 112, NUMBER 1

completely fused, unarmed. Coxa with
small spine on anterior margin.

Pereopod III (Fig. 4c) dactylus with base
to heel straight, heel broadly rounded and
low, heel to tip with broad evenly rounded
indent, tip acute, tip to base smoothly con-
vex to straight; lateral surface smooth, with
several small tufts of short setae in roughly
straight line across medioproximal surface,
dorsodistal margin with tufts of short setae;
ventromesial margin with long plumose se-
tae, dorsal margin with short simple and
plumose setae; mesial surface smooth, with
plumose setae proximally at junction with
propodus. Propodus not inflated dorsoven-
trally; lateral surface smooth, with long plu-
mose setae distally, with simple setae on
dorsal margins, and long plumose setae on
ventral margin; dorsolateral surface narrow,
oblique, flattened; mesial surface with scat-
tered long setae on and near distal margin.
Carpus produced dorsodistally, exceeding
proximal margin of propodus by about 4%
length of propodus, pointed but not acute;
dorsolateral margin unarmed; lateral sur-
face slightly rugose dorsodistally, with mat
of short setae and 2 longer rows of setae
ventrally; mesial surface smooth, with long
plumose setae on margins and scattered on
surface. Merus smooth with large decalci-
fied window; dorsal and ventral margins
unarmed, with long plumose setae; later-
odistal margin with long plumose setae;
mesial surface smooth. Basi-ischium in-
completely fused and unarmed. Coxa un-
armed. Female with large gonopore on an-
terior mesial surface of coxa, surrounded
with short plumose setae; male with small
pore on coxa.

Pereopod IV (Fig. 4d) dactylus with base
to tip proximally convex becoming con-
cave, heel and indent absent, tip acute, tip
to base straight distally becoming convex
proximally; lateral surface smooth, ventral
margin with long plumose setae, dorsal
margin with short simple setae; mesial sur-
face with dorsal decalcified window, de-
marcated ventrally by longitudinal elevated
ridge bearing row of short setae; with setose

153

punctae ventral to decalcified window. Pro-
podus expanded dorsally and ventrally;
ventral expansion exceeds ventral margin of
dactylus, margin with long plumose setae;
dorsal expansion with row of long plumose
setae medially; lateral and mesial surfaces
smooth. Carpus not produced dorsodistally;
lateral and mesial surfaces smooth; dorsal
margin with short simple and long plumose
setae; ventral margin with short simple se-
tae; mesial surface with decalcified win-
dow. Merus lateral surface with scattered
short transverse rows of setae, dorsal and
ventrodistal margins with long plumose se-
tae; mesial surface with large decalcified
window proximoventrally. Basi-ischium in-
completely fused and unarmed. Coxa un-
armed.

Pereopod 5 reduced, slender, lacking dis-
tinctive features. Coxa of male with large
mesioproximal gonopore.

Abdomen (Fig. 4e) somite I approxi-
mately as long as wide, widest posteriorly;
dorsal surface with anterior margin straight;
posterior margin straight, with elevated
submarginal row of short setae; with small
transverse decalcified windows laterad to
segment midline. Somite II dorsal surface
with submarginal transverse ridge anterior-
ly; with small transverse decalcified win-
dows laterad to segment midline just ante-
rior to submarginal ridge; with tuft of setae
at posterolateral angle, extending onto pleu-
ra posteromesially; posterior margin with
indistinct punctate submarginal groove lat-
erally; pleura expanded and directed slight-
ly anteriorly; lateral margins rounded, an-
terior and lateral margins with long plu-
mose setae, posterior margin with short se-
tae. Somite III similar to somite II, but
narrower, shorter, and lacking anterior sub-
marginal ridge; small tuft of short thick se-
tae on posterolateral angle; pleura thinner
and shorter than on somite II, directed an-
terolaterally, with setae as in somite II; an-
terolateral angle acute; dorsal surface
obliquely flattened anterolaterally. Somite
IV similar to somite III, but thinner and
shorter; dorsal surface with thick setae pos-

154 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

terolaterally; pleura thinner and shorter than
on somite III, directed posterolaterally; dor-
sal surface obliquely flattened anterolater-
ally; margin with long plumose setae. So-
mite V narrower than somite IV; lateral
margins with short plumose setae; pleura
absent. Somite VI subequal to somite V in
width but longer; dorsal surface with short
transverse rows of setae laterad to midline
anteriorly; lateral margins with long plu-
mose setae; pleura absent.

Females with uniramous, paired pleopods
on somites II-V; males lacking pleopods.

Uropods lacking distinctive features.

Telson of male (Fig. 4f) spatulate, later-
ally expanded, with length subequal to
width, produced into short rounded tip dis-
tally; weakly calcified except for large tri-
angular anterior plate; median longitudinal
groove long, extending to distal end of cal-
cified plate, lined with long thin simple se-
tae; calcified plate slightly elevated medi-
ally but without ridge. Telson of female
(Fig. 4g) ovate, longer than wide, broadly
triangular, dorsal surface smooth, with me-
dian longitudinal groove anteriorly; with
row of setose punctae lateral to midline
from median of longitudinal groove to dis-
tal end of groove; margins with long plu-
mose setae.

Coloration.—Off-white with whitish se-
tae in life and in preservative.

Distribution.—Indo-West Pacific: Ha-
wail; Western Australia; Seychelles; Mal-
dives; 3.7—13.7 m.

Remarks.—This species can easily be
separated from all of the other Indo-West
Pacific species of Albunea, except A. mad-
agascariensis, by the concave shape of the
lateral margins of the eyes. Although a di-
rect comparison of A. speciosa with A.
madagascariensis material is desirable, the
types (and only known specimens) of A.
madagascariensis apparently were not de-
posited in the Muséum National d’ Histoire
Naturelle (MNHN), contrary to Thomassin
(1973) (Nguyen, pers. comm.). Neverthe-
less, based on Thomassin’s (1973) descrip-
tion and illustrations, the two species can

tentatively be separated by several charac-
ters. A. speciosa has a short rostrum that
does not reach the distal margin of the oc-
ular plate, a CG4 comprised of two medial
elements, a more truncate heel of pereopod
Ill, and a rounded distal tip on the telson
of the male, while A. madagascariensis has
a long rostrum that well exceeds the distal
margin of the ocular plate, a CG4 of 1 me-
dial element, a rounded heel of pereopod
Ill, and a pointed distal tip on the telson of
the male. Given a larger sample size, par-
ticularly from the westernmost Indo-Pacific,
all of these characters may prove to repre-
sent only intraspecific variation of A. spe-
ciosa, in which case A. madagascariensis
would become a synonym of A. speciosa.
Two of Thomassin’s specimens exhibited a
peculiar orange and brown banding pattern
(Thomassin 1973: 268, pl. 1). No other spe-
cies of Albunea have been reported with
anything but almost uniform coloration,
suggesting the possibility that this may
have been an artifact of the environment.
More study is needed to determine whether
this is similar to the type of environmen-
tally-induced color changes observed in
mole crabs of the genus Hippa Fabricius,
1787 (e.g., Bauchau & Passelecq-Gérin
1987).

Little is known about the biology of this
species other than the few records of ovig-
erous females herein reported. Several A.
speciosa specimens (QM W22285) were
collected together with the holotype of Al-
bunea danai new species, but it is unknown
if the two species regularly coexist.

Specimens of A. speciosa have been re-
ported only three times in the literature
(Dana 1852, Borradaile 1904, Seréne
1973). Examination of Borradaile’s (1904)
specimen from the Maldives confirms its
identity as this species, making Borradaile’s
(1904) record the first from outside Hawaii,
although his paper was overlooked by sub-
sequent researchers.

Because A. speciosa is now known to be
a wide-ranging Indo-Pacific species, rather
than a Hawaiian endemic, it would be use-

VOLUME 112, NUMBER 1

ful to understand the origin and subsequent
distribution of this species in order to better
understand albuneid biogeography. The Ha-
waiian specimens of A. speciosa are far re-
moved from other populations of this spe-
cies, a phenomenon that has been shown for
other wide-ranging Hawaiian shallow-water
fauna with western Indo-Pacific affinities
(Newman 1986), and they probably reached
the relatively young Hawaiian Islands by
long distance dispersal. However, the clos-
est locality where this species has been re-
ported is not from the Philippines or nearby
islands in the northern Indo-Pacific, but
rather from Western Australia to the south.
Unless the species is found on the east coast
of Australia or in the island groups to the
east, such as the New Hebrides or Society
Islands, it is difficult to hypothesize the
eastward dispersal route by which A. spe-
ciosa reached the Hawaiian Islands. A
northern or southern dispersal route across
the Pacific appears equally likely, given the
evidence currently available.

Albunea danai, new species
Figs 5, 6

Type material.—Holotype: Hawaii, Hal-
onu Blow Hole dive site, south shore,
Oahu, 12.2—13.7 m, coll. R. Holcom, 4 Apr
1997: 1 male, 16.7 mm CL (QM W23105).
Allotype: Kailua, Oahu, Mar 1938: 1 fe-
male, 16.8 mm CL (BPBM S11782). Para-
types: Waikiki, Oahu, 22.7 m, coll. Smith
and Allen, 23 May 1948: 1 male, 11.6 mm
CL (BPBM S5343); off Waikiki, Oahu, 6.1
m, coll. Allen and Smith, 30 May 1948: 1
female, 10.8 mm CL (AMNH 17716); Ka-
hana Bay, Oahu, 7.6—9.1 m, coll. “‘Pele”’
Expedition, 25 Jul 1959: 1 male, 8.6 mm
CL (BPBM S6775); off Sand Island, Oahu,
4.8-7.6 m, coll. ‘“‘Pele’’? Expedition, 17 July
1959: 1 male, 10.4 mm CL (AMNH
17717); Diamond Head, Oahu, 7.6—13.6 m,
coll. “‘Pele’’ Expedition, 9 Sep 1959: 1 fe-
male, 13.0 mm CL (BPBM S6777); [Ma-
mala Bay], off Honolulu, Oahu, 27.4—40.2
m, coll. T. Richert, Feb—Mar 1962: 1 male,

155

foe mm «Cl. li female, 15.5 mm CL
(WAM 481-97); [Mamala Bay], off Ewa
Beach, near Pearl Harbor, Oahu, 27.4 m,
coll. B..R- Wilson on R/V “Pele,” 5 Jul
1964: 1 ovigerous female, 10.8 mm CL
(WAM 143-70).

Additional material examined (non
type).—Hawaii: Kanoehe Bay, Oahu, 1924:
1 male (poor condition), 4.0 mm CL
(BPBM S7806).

Type locality.—Halonu Blow Hole dive
site, south shore, Oahu, Hawaii, USA, Pa-
cific Ocean.

Diagnosis.—Carapace slightly longer
than wide, covered with lightly setose
grooves. Anterior margin with 8—9 teeth.
Setal field with narrow lateral elements and
slightly concave anterior margin; posterior
lateral elements not extending to posterior
lateral elements of CG1. CG1 with separate
posterior lateral elements; CG4 with 2—4
short medial elements; CG5 divided into 2
lateral elements, not nearly reaching mar-
gins of CG6; CG6 and CG7 separate, but
almost approximate; CG8 with 1-2 poste-
riorly displaced median elements separated
from lateral elements; CG11 absent. Ros-
trum present, not reaching proximal margin
of ocular plate. Ocular plate subquadrate.
Ocular peduncles dorsoventrally flattened
and elongate, pointed at tip, approximate
along mesial margin; lateral margin convex;
mesial margin straight proximally, convex
distally. Cornea at lateral margin of tip. An-
tennule with 87—92 flagellar exopodal and
3—4 endopodal segments. Antenna with 7
flagellar segments; acute spine on dorsolat-
eral surface of peduncle segment I. Dactyli
of pereopods II, III with heels low and
smoothly rounded. Coxa of pereopod III of
males with small male pore. Telson of male
triangular, dorsoventrally flattened laterally
and distally, inflated medially. Telson of fe-
male flattened, rounded at tip.

Description.—Carapace (Fig. 5a) slightly
wider than long. Anterior margin slightly
concave on either side of ocular sinus, be-
coming convex laterally, with 8—9 large
spines along length. Rostrum a small acute

156 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Albunea danai, new species. A, male, 16.7 mm CL, QM W23105, holotype; B—EK H—J, ovig. female,
10.8 mm CL, WAM 143-70; G, male 11.6 mm CL, BPBM 5343. A, carapace and eyes, dorsal view; B, eyes,
dorsal view; C, left antennule, lateral view; D, right antenna, lateral view; E, left mandible, lateral view; EF left
maxilla, lateral view; G, left maxillule, lateral view; H, left maxilliped I, lateral view; I, left maxilliped I, lateral
view; J, left maxilliped III, lateral view. Scale = 1.2 mm (C, D, E H, J), 1.6 mm (B, E, G), 2.2 mm (J), and

6.7 mm (A).

tooth, extending only half the distance be-
tween distal margin of ocular sinus and oc-
ular plate. Ocular sinus smoothly concave
and unarmed. Frontal region smooth; setal
field broad posteriorly, narrowing anterior-
ly, with narrow anterior lateral elements and
slightly concave anterior margin; posterior

lateral elements thin and not reaching to
posterior lateral elements of CG1. CG1 me-
dial portion parallel to anterior margin of
carapace, faintly sinuous, strongly crenu-
late, divided into medial fragment and
curved, posteriorly displaced lateral ele-
ments. Mesogastric region smooth; CG2

VOLUME 112, NUMBER 1

157

Fig. 6. Albunea danai, new species. A, KE male, 16.7 mm CL, QM W23105, holotype; B—E, male 11.6 mm
CL, BPBM 5343; G, female, 16.8 mm CL, BPBM S11782, allotype. A, left pereopod I, lateral view; B, right
pereopod II, lateral view; C, right pereopod III, lateral view; D, right pereopod IV, lateral view; E, abdominal
somites I-VI, dorsal view; E telson of male, dorsal view; G, telson of female, dorsal view. Scale = 3.0 mm (EF
G) and 4.4 mm (A-E).

absent; CG3 broken into 6 short elements
approximately equally spaced between pos-
terior lateral elements of CG1; CG4 with
2—4 short medial elements spaced approx-
imately equally between longer lateral ele-
ments of CG4. Hepatic region smooth with
oblique setose groove at median of lateral
margin. Epibranchial region roughly trian-

gular, smooth; posterolateral margin with 2
short rows of setae. Metagastric region
smooth; CG5 divided into 2 short lateral el-
ements. CG6 strongly crenulate, strongly
anteriorly concave medially and sloping out
to anteriorly convex lateral thirds, median
and lateral thirds separated by short setae-
free gap lateral to small depressions. CG7

158

oblique, almost reaching lateral margins of
median segment of CG6. Cardiac region
smooth; CG8 present as 2 very short later-
omedial elements displaced posteriorly
from longer lateral elements. CG9 present
as 2 short lateral grooves with gap at mid-
line. CG10 present as 2 curved lateral frag-
ments, with gap between fragments approx-
imately equal to length of single fragment.
CG11 absent. Branchial region with nu-
merous short, transverse rows of setae. Pos-
terior margin deeply and evenly convex,
with submarginal groove reaching about
half-way either side of posterior concavity.
Branchiostegite with short anterior submar-
ginal spine; anterior region with scattered
short transverse lines ventral to linea ano-
murica; with many short rows of setae and
sparsely covered with long plumose setae
ventrally; posterior region membranous
with numerous, irregular fragments and
sparsely covered with long plumose setae.

Ocular plate (Fig. 5b) subquadrate with
narrow indentation; proximal ocular seg-
ments (Fig. 5b) reduced to small rounded
calcified area on either side of ocular plate.
Ocular peduncle (Fig. 5b) elongate, with
medially convex lateral margins, tapering to
rounded distal cornea located in lateral
notch; mesial margin approximate almost
all of length; mesial and proximolateral
margins of segment with sparse row of long
plumose setae; tuft of plumose setae at
proximolateral ventral angle.

Antennule (Fig. 5c) segment III narrow
proximally, expanding distally to twice
proximal width; with plumose setae on dor-
sal and ventral margins and sparsely scat-
tered on lateral surface; dorsal exopodal fla-
gellum with 87—92 segments and long plu-
mose setae on dorsal and ventral margins;
ventral endopodal flagellum short, with 3—
4 segments and plumose setae on dorsal and
ventral margins. Segment II medially in-
flated in dorsal view, with plumose setae on
dorsal and ventral margins and scattered se-
tae on ventrolateral third of surface. Seg-
ment I wider than long, unarmed; dorsal
third of lateral surface rugose with long plu-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

mose setae; long plumose setae on dorsal
and ventral margins.

Antenna (Fig. 5d) segment V about 2
times longer than wide, with long plumose
setae on dorsal margin and scattered setae
on distal half of lateral surface; flagellum
7-segmented, with long plumose setae on
dorsal, ventral and distal margins. Segment
IV expanded distally with long plumose se-
tae on dorsal, ventral and distal margins,
and 2 rows of setae on dorsolateral surface.
Segment III with long plumose setae on
dorsal and ventral margin. Segment II short,
widening distally, with plumose setae on
margins and scattered on lateral surface; an-
tennal acicle long, thin and exceeding distal
margin of segment IV by % the length of
segment IV, with long plumose setae on
dorsal margin. Segment I rounded proxi-
mally, flattened ventrolaterally, with long
plumose setae on margins; lateral surface
with acute spine dorsally, with low semi-
circular dorsolateral lobe ventrodistal to
spine; segment with ventromesial antennal
gland pore.

Mandible (Fig. 5e) incisor process with
1 tooth; cutting edge with 1 tooth. Molar
process with 2-3 teeth. Palp 3-segmented,
with plumose setae on margins and long,
thick, simple setae arising from bend in sec-
ond segment.

Maxilla (Fig. 5f) exopod evenly rounded,
with plumose setae along distal margin.
Scaphognathite bluntly angled on posterior
lobe, with plumose setae. Endopod and en-
dites without distinctive characters.

Maxillule (Fig. 5g) distal endite proxi-
mally narrow, widening to inflated distal
end, with thick simple setae on distal mar-
gin. Proximal endite with thick simple setae
on distal margin. Endopodal external lobe
truncate distally, and curled under; internal
lobe reduced, with 3 thick setae at distolat-
eral margin.

Maxilliped I (Fig. Sh) epipod with plu-
mose setae on margins, distolateral surface
and mesial surface (epipod shown curled in
Fig. 5h). Endite tapered distally and sube-
qual to first segment of exopod. Exopod

VOLUME 112, NUMBER 1

with 2 segments; proximal segment narrow,
margins parallel, margins with plumose se-
tae; distal segment spatulate, about as long
as wide, broadest medially, margins and
mesioventral surface with long plumose se-
tae. Endopod flattened and elongate, reach-
ing to distal end of proximal exopodal seg-
ment; plumose setae on margins and me-
dian of lateral surface.

Maxilliped II (Fig. 51) dactylus evenly
rounded, length equal to width, with thick
simple setae distally and on distolateral sur-
face. Propodus 2 times wider than long,
slightly produced at dorsodistal angle, with
plumose setae on dorsal margin and long
simple setae on dorsodistal margin. Carpus
not produced dorsodistally, about 2 times
longer than wide; long simple setae on dor-
sal and distal margins. Merus about 3 times
longer than wide, margins parallel; with
simple setae on ventrolateral margin and
plumose setae on dorsolateral margin. Basi-
ischium incompletely fused, plumose setae
on margins. Exopod % times longer than
merus, with flagellum 1-segmented.

Maxilliped III (Fig. 5j) dactylus with
rounded tip; with long plumose setae on
dorsal margin and lateral surface. Propodus
with longitudinal median row of plumose
setae on lateral surface; dorsal margin with
plumose setae. Carpus slightly produced
onto propodus; lateral surface with row of
plumose setae ventromedially; plumose se-
tae on dorsal margin. Merus unarmed, with
plumose setae on dorsal and ventral mar-
gins and scattered on lateral surface. Basi-
ischium incompletely fused, with weak
crista dentata of about 2 teeth. Exopod 2-
segmented: proximal segment small; distal
segment styliform, tapering, approximately
43 length of merus; with plumose setae on
surface; without flagellum.

Pereopod I (Fig. 6a) subchelate. Dactylus
curved and tapering; lateral and mesial sur-
faces smooth; dorsal margin with long plu-
mose and short simple setae; ventral margin
with short simple setae. Propodus lateral
surface with numerous short, transverse
rows of setose rugae; dorsal margin un-

159

armed; ventral margin produced distally
into acute spine; cutting edge lacking teeth,
lined with long plumose setae; dorsal mar-
gin with long plumose setae, ventral margin
with short simple setae. Carpus with dor-
sodistal angle produced into strong corne-
ous-tipped spine; dorsal margin otherwise
unarmed; dorsal and distal margins with
long plumose setae; lateral surface with
small distal rugose area, with few trans-
verse setose ridges on distal half of surface;
mesial surface smooth with few median
rows of setae, margins with long plumose
setae. Merus unarmed; lateral surface with
scattered transverse rows of long plumose
setae, margins with long plumose setae;
mesial side with few short rows of setae.
Basi-ischium incompletely fused, unarmed.
Coxa unarmed.

Pereopods [I-IV with dactyli laterally
compressed and dorsoventrally expanded.

Pereopod II (Fig. 6b) dactylus smooth;
base to heel straight, heel smoothly round-
ed, heel to tip with wide acute indent, tip
acute, tip to base broadly convex distally
and slightly concave proximally; lateral sur-
face smooth, with several small tufts of
short setae in roughly straight line across
medioproximal surface, several widely
spaced submarginal tufts of short setae dor-
sodistally; mesial surface smooth, ventral
margin with long plumose setae, dorsal
margin with short simple setae, with patch
of long plumose setae at base. Propodus
dorsal surface smooth, ventral margin in-
flated and rounded; oblique row of long
plumose setae on distal margin of lateral
surface; distal and ventral margin with long
plumose setae; dorsolateral surface a nar-
row, oblique, flattened shelf, with short se-
tae on dorsal margin and long plumose se-
tae on ventral margin; mesial surface with
elevated, curved setose ridge from ventral
junction with dactylus almost to ventral
proximal junction with carpus. Carpus pro-
duced and gently rounded dorsodistally,
dorsal margin unarmed; lateral surface
smooth, with irregular broken row of rugae
and submarginal elevated ridge ventrally,

160

rugae and ridge with long plumose setae;
margins with long plumose setae; mesial
surface smooth with long plumose setae in
scattered patches on dorsal half of surface
and on margins. Merus with large median
decalcified window covering nearly all of
lateral surface, with few scattered setae on
surface and margins; mesial surface nearly
smooth with few setae, with decalcified
area on proximal % near junction with basi-
ischium. Basi-ischium incompletely fused
and unarmed. Coxa with small spine on an-
terior margin.

Pereopod III (Fig. 6c) dactylus with base
to heel straight, heel broadly rounded and
slightly produced, heel to tip with broadly
concave indent, tip acute, tip to base
smoothly convex distally to straight proxi-
mally; lateral surface smooth, with several
small tufts of short setae in approximately
straight line across medioproximal surface,
dorsodistal margin with tufts of short setae;
ventral margin with long plumose setae,
dorsal margin with short simple and plu-
mose setae; mesial surface smooth with
plumose setae proximally at junction with
propodus. Propodus not inflated dorsoven-
trally; lateral surface smooth, with long plu-
mose setae distally, with simple setae on
dorsal margins; dorsolateral surface narrow,
oblique, flattened; mesial surface with scat-
tered long setae on and near distal margin.
Carpus produced dorsodistally, exceeding
proximal margin of propodus by about %
length of propodus, rounded; dorsolateral
margin unarmed; lateral surface slightly ru-
gose dorsodistally, with mat of short setae
and row of setae ventrally; mesial surface
smooth, with long plumose setae on mar-
gins and scattered on surface. Merus
smooth, with large decalcified window cov-
ering nearly half of lateral surface medially;
dorsal and ventral margins unarmed, with
long plumose setae; laterodistal margin
with long plumose setae; mesial surface
smooth. Basi-ischium incompletely fused
and unarmed. Coxa with tubercle on ante-
rior margin. Female with large gonopore on
anterior mesial margin of coxa, surrounded

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

with short plumose setae; male with small
pore on coxa.

Pereopod IV (Fig. 6d) dactylus with base
to tip proximally convex becoming distally
concave, heel and indent absent, tip acute,
tip to base straight distally, becoming con-
vex proximally; lateral surface smooth,
ventral margin with long plumose setae,
dorsal margin with short simple setae; me-
sial surface with dorsal decalcified window,
demarcated ventrally by longitudinal ele-
vated ridge with row of short setae; with
setose punctations ventral to decalcified
window. Propodus expanded dorsally and
ventrally; ventral expansion exceeds ventral
margin of dactylus, margin with long plu-
mose setae; dorsal expansion with row of
long plumose setae medially; lateral and
mesial surfaces smooth. Carpus not pro-
duced dorsodistally; ventral %4 of lateral sur-
face and mesial surface smooth, dorsal 4 of
lateral surface with mat of short setae; me-
sial surface with decalcified window; dorsal
margin with short simple and long plumose
setae; ventral margin with short simple se-
tae. Merus lateral surface with scattered
short transverse rows of setae, dorsal and
ventrodistal margins with long plumose se-
tae; mesial surface with large decalcified
window proximoventrally. Basi-ischium in-
completely fused and unarmed. Coxa un-
armed.

Pereopod 5 reduced, slender, lacking dis-
tinctive features. Coxa of male with large
mesioproximal gonopore.

Abdomen (Fig. 6e) somite I approxi-
mately as long as wide, widest posteriorly;
dorsal surface with anterior margin straight;
posterior margin straight with elevated sub-
marginal row of short setae; with small
transverse decalcified windows laterad to
segment midline. Somite II dorsal surface
with submarginal transverse ridge anterior-
ly; with small transverse decalcified win-
dows laterad to segment midline just ante-
rior to submarginal ridge; with tuft of setae
at posterolateral angle, extending onto pleu-
ra posteromesially; pleura expanded and di-
rected slightly anteriorly; lateral margins

VOLUME 112, NUMBER 1

rounded, anterior and lateral margins with
long plumose setae, posterior margin with
short setae. Somite III similar to somite I,
but narrower, shorter, and lacking anterior
submarginal ridge; small tuft of short thick
setae on posterolateral angle; pleura thinner
and shorter than on somite II, directed pos-
terolaterally, with setae as in somite II; an-
terolateral angle acute; dorsal surface
obliquely flattened anterolaterally. Somite
IV similar to somite III, but thinner and
shorter; dorsal surface with few thick setae
posterolaterally; pleura thinner and shorter
than on somite III, directed posterolaterally;
dorsal surface obliquely flattened anterolat-
erally; margins with long plumose setae.
Somite V subequal to somite IV; lateral
margins with plumose setae; pleura absent.
Somite VI subequal to somite V in length
but wider; dorsal surface with short trans-
verse rows of setae laterad to midline an-
teriorly and posteriorly; lateral margins
with long plumose setae; pleura absent.
Females with uniramous, paired pleopods
on somites II—-V; males without pleopods.
Uropods lacking distinctive features.
Telson of male (Fig. 6f) triangular, slight-
ly longer than wide, with smoothly rounded
tip; proximal half heavily calcified, distal
half weakly calcified except for large me-
dian region; median longitudinal groove ex-
tending to distal end of calcified area, line
with long thin simple setae; junction of
proximal and distal regions demarcated by
strong line of long setae laterally; calcified
plate slightly elevated medially but without
ridge. Telson of female (Fig. 6g) ovate, lon-
ger than wide, rounded distally; dorsal sur-
face smooth, with median longitudinal
groove anteriorly; with row of setose punc-
tae lateral to midline from posterior end of
longitudinal groove to %4 length of telson;
margins with long plumose setae.
Coloration.—In life, brownish with red-
dish-brown setae. Preserved, uniform off-
white to tan.
Distribution.—Known only known from
Oahu, Hawaii; 4.8—40.2 m.

Etymology.—Named after James D.

161

Dana (1813-1895), famed carcinologist and
describer of A. speciosa, and many other
species of Indo-Pacific Crustacea. Gender:
masculine.

Remarks.—This species is most similar
to A. carabus (Linnaeus, 1758), from the
Mediterranean and western Africa, in the
shape of the dactyli of pereopods II-IV and
telson morphology. Albunae carabus can be
easily separated from A. danai, new spe-
cies, by its CG8 of four medial elements,
more strongly crenulated CGs, more pro-
nounced heel on the dactyli of pereopods II
and III, and a less inflated maxilliped III
merus. A. danai new species can be distin-
guished from other Indo-West Pacific spe-
cies by the triangular shape of the telson of
the male, the rounded dactyl of pereopod
III, and setal patterns on the carapace of
both sexes. All other species of Albunea
from the Indo-West Pacific region with
smoothly rounded heels on pereopod III
(e.g., A. speciosa) also have much more
strongly setose carapace grooves than A.
danai, new species, and distinctive male
telson morphologies.

As previously indicated, the holotype
was collected with specimens of A. specio-
sa (QM W22285).

This species, unlike A. speciosa, appears
to be a true Hawaiian endemic based on all
available data. However, this conclusion
should be accepted cautiously, given the
‘“‘endemic’”’ label applied to A. speciosa pri-
or to this study and that endemicity appears
to be the exception, rather than the rule, in
the Hawaiian biota (Newman 1986). As a
whole, the Hawaiian albuneids seem to fit
in the category of attenuated Indo-West Pa-
cific fauna (Newman 1986), given the
markedly greater diversity of albuneids (17
Species in six genera) throughout the rest of
the Indo-West Pacific.

Acknowledgments

I thank R. Van Syoc (CASIZ), R. Jocqué
(MRAC), R. Symonds (UMZC), R. Lemai-
tre (USNM), J. Gnffith and M. Hewitt

162 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(WAM), and E. Lazo-Wasem (YPM) for
making specimens available for study.
Also, thanks to Nguyen Ngoc-Ho (MNHN)
for trying to locate the types of A. mada-
gascariensis. L. Eldredge loaned specimens
from the collections of BPBM and gener-
ously offered to deposit two paratypes of A.
danai in the AMNH. Special thanks to P.
Davie (QM) for allowing me to study the
holotype specimen of A. danai. Color pho-
tographs of both species were generously
provided by R. Holcom of Hawaii. A. Har-
vey (Georgia Southern University) was in-
strumental in providing training in comput-
er aided biological illustration, co-produced
Figs. 1 and 2, and reviewed the manuscript.
Two anonymous reviewers also contributed
greatly to the final product.

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logical color changes in anomuran decapods of
the genus Hippa.—Indo-Malayan Zoology 4:
135-144.

Benedict, J. E. 1904. A new genus and two new spe-
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Borradaile, L. A. 1904. Marine crustaceans. XIII. The
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Boyko, C. B., & A. W. Harvey. 1999. Crustacea De-
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. 1855. United States Exploring Expedition.
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Edmondson, C. H. 1946. Reef and shore fauna of Ha-
waii.—Bernice P. Bishop Special Publication
22:381 pp.

Efford, I. E., & J. Haig. 1968. Two new genera and
three new species of crabs (Decapoda: Ano-
mura: Albuneidae) from Australia.—Australian
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Evans, A. C. 1967. Syntypes of Decapoda described
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Fabricius, J. C. 1787. Mantissa insectorum sistens eor-
um species nuper detectus adjectis characteribus
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observationibus. Hafniae. 1:348 pp.

Gordon, I. 1938. A comparison of the two genera AI-
bunea and Lepidopa (Crustacea, Anomura),
with description of a new species from Singa-
pore.—Bulletin of the Raffles Museum 14:186—
1ST:

Harvey, A. W., & E. M. De Santo. 1997. A new spe-
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ton 110(1):65—68.

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cinology.—Transactions of the Linnaean Soci-
ety of London, 2nd ser. 5(10):325—458, pls. 36—
40.

Holthuis, L. B. 1958. Crustacea Decapoda from the
northern Red Sea (Gulf of Aqaba and Sinai Pen-
insula) IJ. Hippidea and Brachyura (Dromiacea,
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eries Research Station Bulletin 17(9):41—54.

Latreille, P A. 1825. Familles naturelles du régne an-
imal, exposées succinctement et dans un ordre
analytique, avec l’indication de leurs genres.
Paris: J.-B. Bailliére. 570 pp.

Linnaeus, C. 1758. Systema naturae per regna tria na-
turae, secundum classes, ordines, genera, spe-
cies, cum characteribus, differentis, synonymis,
Locis. 824 pp.

Miers, E. J. 1878. Revision of the Hippidea.—Zoolog-
ical Journal of the Linnean Society 14(76):312-
356; ple-5:

Newman, W. A. 1986. Origin of the Hawaiian marine
fauna: dispersal and vicariance as indicated by
barnacles and other organisms. Jn R. H. Gore
& K. L. Heck, eds. Crustacean Biogeography.
Crustacean Issues 4. Rotterdam: A. A. Balkema.
pp. 21-49.

Ortmann, A. E. 1896. Die geographische verbreitung
der decapodengruppe der Hippidea.—Zoologis-
che Jahrbucher, Abteilung fiir Systematik, Geo-
graphie und Biologie der Thiere 9(2):219—243.

Seréne, R. 1973. A new species of Decapoda Hippi-
dea: Albunea mariellae nov. sp. from the Banda
Sea.—Crustaceana 24(3):261—264, 2 pls.

Stillman, J. H., & G. N. Somero. 1996. Adaptation to
temperature stress and aerial exposure in con-
generic species of intertidal porcelain crabs (ge-

VOLUME 112, NUMBER 1

nus Petrolisthes): correlation of physiological
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tribution.—Journal of Experimental Biology
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Stimpson, W. 1858. Prodromus descriptionis animal-
ium evertebratum, quae in Expeditione ad
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lica Federata missa, Cadwaladaro Ringgold et
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of Philadelphia 10:225—252.

163

Thomassin, B. A. 1973. Albunea madagascariensis n.
sp., nouvelle espéce d’ Hippidea (Decapoda, An-
omura) des sables coralliens de la région de Tu-
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24(3):265—274.

Weber, E 1795. Nomenclator entomologicus secundum
entomologiam systematicam ill. Fabricii adjec-
tis speciebus recens detectis et varietatibus. Chi-
lonii et Hamburgi: Carolum Ernstum Bohn. viii
172 pp:

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(1):164—174. 1999.

Two new species of Hansenium (Crustacea: Isopoda: Asellota) from
Madang, Papua New Guinea

Kathrin S. Bolstad and Brian Kensley

Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian
Institution, Washington, D.C. 20560, U.S.A.

Abstract.—Two new species of Hansenium, H. tropex and H. thomasi, are
described from Paddock Reef, Madang, Papua New Guinea. H. tropex is char-
acterized by the possession of a broad oar-shaped lobe of the carpus of pereo-
pod 1 in the male. H. thomasi is characterized by a narrow tapering mesially-
directed lobe of the carpus of pereopod 1 of the male. The genus is redefined,
with its chief character being the presence of a large carpal lobe on the first
male pereopod. The eight stenetriid genera are compared on the basis of six

characters.

The shallow water marine crustaceans
(with the exception of the Amphipoda) of
the Papua New Guinea region are poorly
known. The Isopoda have received scant at-
tention (in all, only about 15 species have
been recorded from the region.) Stebbing
(1900), recorded three cirolanids, one cy-
mothoid and one sphaeromatid from New
Britain, New Guinea, Nobili (1905) de-
scribed a corallanid and a bopyrid, while
Nierstrasz (1931) lists three cymothoids,
two cirolanids, three corallanids, and one
sphaeromatid in his catalogue of isopod re-
cords. Bruce (1982, 1993, 1994) has de-
scribed cirolanids from the Madang area,
while Keable (1997) added further cirolanid
records; Jones et al. (1983) described three
species of corallanids; Williams & Bunk-
ley-Williams (1992) described two new
species of cymothoids. No asellote isopods
have been reported. The finding of two very
distinctive stenetriids was, therefore,
thought to be well worth recording.

This paper was written while the authors
participated in the Mentorship Program of
the Thomas Jefferson High School for Sci-
ence and Technology in Alexandria, Virgin-
ia.

The material described in this paper was
collected by J. D. Thomas and J. Clark-

Walker of the National Museum of Natural
History, Smithsonian Institution. The col-
lectors were carrying out a survey of the
shallow water marine invertebrate fauna of
Madang, Papua New Guinea, in coopera-
tion with the Christensen Research Insti-
tute. Jebb & Lowry (1995) provide a useful
description of the habitats of Madang La-
goon.

Suborder Asellota
Family Stenetriidae Hansen, 1905
Genus Hansenium Serov & Wilson, 1995

Hansenium Serov & Wilson, 1995: 72.

Diagnosis.—Lateral tooth of cephalon
moderately well developed, antennal tooth
subequal to lateral, or rounded. Rostrum
short, rectangular, anterior margin truncate.
Eyes reniform, of 13—19 ommatidia. Pereo-
pod 1 in male with carpus produced pos-
terodistally into lobe; propodus longer than
width of palm, latter often with few teeth
close to articulation of dactylus. Pleopod 2
in male with rounded protopodal lobe dis-
tally, appendix masculina of endopod dis-
tally somewhat broadened, truncate, often
ringed by cuticular hairs.

Type species.—Stenetrium hanseni No-
bili, 1906.

VOLUME 112, NUMBER 1

Remarks.—Serov & Wilson (1995), di-
vided the genus Stenetrium into five sepa-
rate genera, and reviewed the remaining
three genera in the family Stenetriidae.
They also listed the constituent species for
each of the genera. Examination of the two
present species, as well as undescribed ma-
terial from the Indian Ocean has led us to
revise the diagnosis of Hansenium, as given
above. The presence of a variously devel-
oped posterodistal lobe on the carpus of pe-
reopod 1 in the male would seem to be the
most characteristic synapomorphy of the
genus, along with the presence of one or
more teeth on the short propodal palm. The
extreme development of this lobe into the
expanded and flattened oar-like structure
seen in H. tropex and H. wilsoni (Miller,
1991a) is approached in two undescribed
species from the Indian Ocean. A review of
the 13 species of Hansenium listed by Ser-
ov & Wilson (1995) shows that only Ste-
netrium bowmani Kensley, 1984, and S. gil-
Lbertense Nordenstam, 1946 do not possess
a carpal lobe on pereopod 1 in the male.

Table 1 summarizes the six characters
thought most significant in defining the
eight genera of the Stenetriidae.

Species included in Hansenium.
Hansenium caicosense (Kensley & Heard,

1991). Turks & Caicos Islands.
Hansenium dodo (Miiller, 1991b). Réunion

Island.

Hansenium entale (Nordenstam, 1946). Gil-
bert Islands.

Hansenium hanseni (Nobili, 1906). Tua-
motu Islands.

Hansenium monodi (Nordenstam,
Gulf of Suez; Seychelles.

Hansenium spathulicarpus (Kensley, 1984).
Belize.

Hansenium stebbingi (Richardson, 1902)
(= Stenetrium antillense Hansen, 1904,
and = Stenetrium occidentale Hansen,
1904). Bermuda; Belize; St. Thomas,
West Indies.

Hansenium thomas n. sp. Madang, Papua
New Guinea.

1946).

165

Hansenium tropex n. sp. Madang, Papua
New Guinea.

Hansenium wilsoni (Miller, 1991a). Moo-
Fea.

Hansenium tropex, new species
Figs. 1-2

Material examined.—Holotype, USNM
253348, do tl 4.8 mm, Paratypes, USNM
253349, 2 ovigerous 2 tl 3.5 mm (dam-
aged), 5.0 mm, Paddock Reef, Madang,
Papua New Guinea, coral rubble, 3—4 m,
coll. J. D. Thomas, 14 Jan 1989.

Description.—Body slender, about 4
times as long as wide, cephalon about twice
as wide as long. Rostrum short, rectangular,
apically truncate, subequal in length to an-
tennal teeth. Antennal teeth elongate, acute.
Lateral teeth pronounced, subequal to an-
tennal teeth. Frontal margin of cephalon be-
low rostrum slightly convex. Eyes reniform,
consisting of about 14 ommatidia.

Antennule, flagellum of about 20 articles;
1 aesthetasc on first article, 2 on second ar-
ticle. Mandible with 4-cusped incisor; 3-
cusped lacinia mobilis; left mandible with
Spine row of 4 comb setae; right mandible
Spine row with 7 comb setae and 3 simple
setae; molar with 8 plumose setae below
marginal serrations. Maxilla 1 inner lobe
with 2 large fringed setae and 2 smaller
simple setae; outer lobe with 8 fringed se-
tae. Maxilla 2 outer lobe with 4 stout
fringed setae and 1 simple seta; middle lobe
with 5 stout fringed setae; inner lobe with
8 simple setae and 4 stout plumose setae on
mesial margin. Maxilliped palp with 3
proximal articles wider than 2 distal arti-
cles, latter longer than wide; endite broad
with 6 broad flattened fan setae distally;
mesial margin with 5 coupling hooks.

Pereonites 1—4 decreasing in length pos-
teriorly, with anterolateral margins acute;
pereonites 5—7 increasing in length poste-
riorly; pereonites 5—6 with rounded pos-
terolateral margins; pereonite 7 with angu-
lar posterolateral margins. Male pereopod 1
elongate, slender; dactylus equal in length

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

166

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VOLUME 112, NUMBER 1 167

Uy y

Fig. 1. Hansenium tropex, new species. A, 6, dorsal view; B, Pereopod 1 36; C, Pereopod 1 2; D, Pereopod
2; E, Pereopod 7; EK Operculum 2; G, Pleopod 1 ¢; H, Pleopod 3.

to propodal palm; short unguis extending projecting anteroventrally, about 0.8 times
beyond proximal spine of propodal palm; length of propodus (Fig. 1B). Female pe-
propodal palm serrate; carpus with large reopod 1 significantly smaller than in male,
stout distally rounded densely setose lobe with sparser setae; dactylus equal in length

168 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig.2. Hansenium tropex, new species. A, Right mandible; B, Left mandible; C, Mandibular palp; D, Maxilla
1; E, Maxilliped; EK Maxilla 2; G, Antennule.

VOLUME 112, NUMBER 1

to propodal palm, with 10 short simple se-
tae on posterodistal margin; propodal palm
with 6 short simple setae, terminating in 2
long setae; carpal lobe much less prominent
than in male.

Pleon having 2 vestigial pleonites plus
pleotelson; posterolateral spines pro-
nounced; postanal margin nearly semicir-
cular. Male pleopod 1 biramous with rami
elongate; mesial margins of rami parallel;
lateral margins rounded and sparsely setose.
Female pleopod 2 1.4 times longer than
wide, pentagonal, with outer margins
sparsely setose. Pleopod 3 operculiform,
with outer ramus broad, sparsely setose; in-
ner ramus narrow and nearly triangular,
with 4 terminal setae.

Remarks.—Hansenium tropex bears the
closest resemblance to H. wilsoni (Miller,
1991a), but several features distinguish the
two. In H. tropex, the antennal spines are
clearly longer than the lateral spines, while
in H. wilsoni, the antennal spines are re-
duced, rounded, and shorter than the lateral
spines. The anterior margin of the cephalon
in H. tropex is convex, straight in H. wil-
soni. In H. tropex the maxilliped has six
flattened fan setae on the endite where H.
wilsoni has three. In pereopod 1 of the male
in H. tropex, the carpal process extends
about %4 the length of the propodus, the pro-
podal palm terminates in a simple blunt
seta, the merus is not produced and the is-
chium is sparsely setose; in H. wilsoni, the
carpal lobe extends beyond the propodus
and the dactylus distally, the propodal palm
has no articulated seta, the merus is slightly
produced, and there is a dense cluster of
setae on the ischium. Pleopod 1 in H. tro-
pex females is nearly pentagonal, with setae
at regular intervals on the lateral margins;
in H. wilsoni, the female pleopod 1 oper-
culum has only four setae and is more tri-
angular in shape. Pleopod 1 in the male of
H. tropex is nearly semicircular, tapering
proximally and distally, while in H. wilsoni
the male pleopod 1 only narrows distally.

Etymology.—The specific name is de-
rived from the Greek ‘tropex’, an oar, refers

169

to the paddle-like carpal lobe of the male
pereopod 1, and is used as a noun in ap-
position.

Hansenium thomasi, new species
Figs. 3-5

Material examined.—Holotype, USNM
253350, 3 tl 4.3 mm, Paratypes, USNM
253351, 3 3, 8 ovigerous ¢ tl 3.4—4.5 mm,
4 juveniles, Paddock Reef, Madang, Papua
New Guinea, 1.5—4 m, coll. J. D. Thomas,
14 Jan 1989.

Diagnosis.—Body about 3.5 times longer
than wide, width of cephalon about 1.5
times length. Rostrum short, rectangular,
anterior margin truncate; frontal margin of
cephalon convex posterior to rostrum. An-
tennal spines equal in length to rostrum; lat-
eral spines subequal in length to antennal
spines. Eyes reniform with about 19 om-
matidia.

Antennule of about 8 articles; 2 aesthe-
tascs on terminal article; 1 aesthetasc on
subterminal; 1 aesthetasc on following ar-
ticle. Mandible with 4-cusped incisor and
3-cusped lacinia mobilis; left mandible with
spine row of 4 comb setae, left molar with
8 fringed setae; right mandible with spine
row of 7 comb setae, 2 plumose setae, and
2 simple setae; right molar with 5 plumose
setae. Maxilla 1 inner ramus with 3 stout
plumose setae, 1 shorter simple seta; outer
ramus with 5 stout comb setae and 5 stout
simple setae. Maxilla 2 outer lobe with 2
long fringed terminal setae, 2 shorter simple
setae, mesial margin with 6 simple setae;
middle lobe with 3 long fringed terminal
setae, 1 simple terminal seta, 6 simple setae
on mesial margin; inner lobe with 6 fringed
setae and 6 simple setae. Maxillipedal en-
dite broad; distal margin with 6 round flat-
tened fan setae and 5 narrow fringed setae,
with 1 short stout simple terminal seta; me-
sial margin with 4 coupling hooks, 4
fringed setae.

Pereonites 1—3 increasing in length pos-
teriorly; pereonite 4 longer than pereonite
5; pereonite 6 longer than pereonite 7; cox-

170 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Hansenium thomasi, new species. A, 6, dorsal view; B, Antennule; C, Left mandible; D, Right
mandible; E, maxilla 2; K Maxilla 1; G, Maxilliped.

VOLUME 112, NUMBER 1 171

412 4d

Oe AS
LG JAN
Go Uji
\ J y
\ ase f
<2 Fond
x oe UY

ZZ,
ge
G0
Hy Uy

} l é

eggs

\
A aA Api

AST NARA
ALA.

i
f

Fig. 4. Hansenium thomasi, new species. A, Pereopod 1 6; B, Pereopod 1 2; C, Pereopod 2; D, Pereopod
7; E, Uropod; E Pleopod 1 3; G, Pleopod 3; H, Operculum @; I, Pleopod 2 ¢.

172

a Ki i

ae visible on pereonites 1 and 3—5; pereon-
ites 1—2 with acute anterolateral angles; per-
eonites 3—4 with concave lateral margins;
pereonite 5 with flat lateral margins, round-
ed posterolateral angle; pereonites 6—7 lat-
erally broad and rounded.

Male pereopod 1 dactylus elongate, nar-
row with acute unguis, extending far be-
yond propodal palm; propodal palm broad,
with 3 sharp teeth, posterodistal angle
greatly produced into narrow lobe termi-
nating in simple stout seta; carpus with
strongly produced posterodistal setose lobe;
merus with small posterodistal projection
and tufts of setae on postero- and antero-
distal margins. Female pereopod 1 much
smaller than in male; dactylus equal in
length to broad propodal palm, with row of
short setae along posterodistal margin; pro-
podal palm broader than in male, with 7
teeth; carpal process much smaller than in
male; merus with setal tufts and low anter-
odistal lobe. Pereopods 2-7 similar, carpus
with 2 posterodistal setae; propodus with
4—6 stout setae on posterior margin.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Pleon with 2 vestigial pleonites plus
pleotelson, latter with single produced pos-
terolateral spine on each lateral margin;
postanal region gently convex. Male pleo-
pod 1 mesial margins finely setose, parallel;
lateral margin convex with 6—7 setae dis-
tally. Male pleopod 2 endopod simple, outer
margin with about 9 setae; protopod distally
subacute, exopod rounded. Female pleopod
2 operculum subtriangular, 1.3 times longer
than wide, sparsely setose. Pleopod 3 rami
together sub-circular, with lateral margin of
exopod setose; endopod nearly triangular
with 6 distal spines.

Remarks.—Hansenium thomasi bears
considerable resemblance to H. entale
(Nordenstam, 1946) from the Gilbert Is-
lands in the Pacific. Comparison of the
male pereopod 1, however, easily distin-
guishes the two species. The dactylus of H.
thomasi is less curved. There are three teeth
on the propodal palm of H. thomasi, more
apically acute than the four found on H.
entale. The propodal teeth of H. thomasi are
well separated from each other, whereas in

VOLUME 112, NUMBER 1

H. entale their fused bases join the propodal
palm. Hansenium thomasi has a large pos-
terodistal propodal lobe, separate from the
teeth, which terminates distally in a simple
stout seta. Hansenium entale has a similar
large toothlike process, but it is more di-
rectly a part of the propodal palm, close to
the other four teeth. Both H. thomasi and
H. entale have produced carpi in the male
pereopod 1, but the process is more setose
and distally rounded in H. thomasi. The
meral process in H. thomasi is much shorter
than in H. entale, where it extends almost
half the length of the carpus.

The anterior margin of the cephalon in
H. thomasi is gently convex, straight in H.
entale. Hansenium entale has a character-
istic pigment pattern, with a line across the
anterior cephalon and a broad band between
the eyes; H. thomasi apparently has no pig-
mentation. In H. entale, the coxae are vis-
ible on pereonites 3 and 4 only, while in S.
thomasi they are visible on pereonites 3—5.

Etymology.—The species is named for
James Darwin Thomas, amphipod special-
ist, who collected the specimens.

Acknowledgments

We thank Dr. Jim Thomas and Ms. Janice
Clark-Walker for collecting the material de-
scribed herein. The collectors’ work was
supported by funding from the National
Geographic Society, the Smithsonian Insti-
tution, and the Christensen Research Insti-
tute of Madang, Papua New Guinea. We
thank Dr. Niel Bruce, and two anonymous
reviewers for their valuable comments on
an earlier draft of this paper. We thank Ms.
Marilyn Schotte who assisted with SEM
preparation of specimens.

Literature Cited

Bruce, N. L. 1982. Records of isopod Crustacea (Cor-
allanidae, Cirolanidae) from Papua New Guin-
ea, with the description of a new species.—
Journal of Crustacean Biology 2:612-618.

. 1993. Two new genera of marine isopod crus-

taceans (Cirolanidae) from Madang, Papua New

173

Guinea.—Memoirs of the Queensland Museum

31(1):1-15.

. 1994. Cirolana and related marine isopod
crustacean genera (family Cirolanidae) from the
coral reefs of Madang, Papua New Guinea.—
Cahiers de Biologie Marine 35:375—413.

Hansen, H. J. 1904. On the morphology and classifi-
cation of the Asellota group of crustaceans, with
descriptions of the genus Stenetrium Hasw. and
its species.—Proceedings of the Zoological So-
ciety of London 19:302-331.

. 1905. On the morphology and classification
of the Asellota group of crustaceans with de-
scriptions of the genus Stenetrium Hasw. and its
species.—Proceedings of the Zoological Socie-
ty of London 1904, (2 Suppl. I):302-—331.

Haswell, W. A. 1881. On some new Australia marine
Isopoda. Part I.—Proceedings of the Linnean
Society of New South Wales 5:470—481.

Jebb, M. H. P, & J. K. Lowry. 1995. Natural history
of Madang Lagoon with an appendix to col-
lecting localities. Pp. 1-24 in J. K. Lowry, ed.,
The Amphipoda (Crustacea) of Madang La-
goon, Papua New Guinea, Part 1.—Records of
the Australian Museum, Supplement 22:1-174.

Jones, D. A., J. D. Icely, & S. M. Cragg. 1983. Some
corallanid isopods associated with wood from
Papua New Guinea, including three new species
(Isopoda: Corallanidae).—Journal of Natural
History 17:837—847.

Keable, S. J. 1997. The Cirolanidae (Crustacea: Iso-
poda) of Darwin Harbour, Northern Territory,
with additional records from northern Australia
and Papua New Guinea. Pp. 245-278 in J. R.
Hanley, G. Caswell, D. Megirian, and H. K.
Larson, eds., Proceedings of the Sixth Interna-
tional Marine Biological Workshop. The marine
flora and fauna of Darwin Harbour, Northern
Territory, Australia. Museums and Art Galleries
of the Northern Territory and the Australian
Marine Sciences Association, Darwin.

Kensley, B. 1984. The Atlantic Barrier Reef ecosystem
at Carrie Bow Cay, Belize, III. New marine Is-
opoda.—Smithsonian Contributions to Marine
Sciences 24:1-81.

, & R. Heard. 1991. Studies on the Crustacea
of the Turks and Caicos Islands, British West
Indies I. Four new marine isopod crustaceans
from the vicinity of Pine Cay.—Gulf Research
Reports 8(3):237—246.

Miiller, H. G. 1991la. The marine isopod family Ste-
netriidae from coral reefs at Bora Bora and
Moorea, Society Islands, with descriptions of
four new species (Crustacea).—Revue Suisse de
Zoologie 1991, 98(1):70—76.

. 1991b. Stenetriidae from coral reefs at Ré-

union Island, southern Indian Ocean. Descrip-

tion of three new species (Crustacea: Isopoda:

174 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Asellota).—Senckenbergiana Biologia 71(4/6):
303-318.

Nierstrasz, H. EK 1931. Die Isopoden der Siboga-Ex-
pedition III. Isopoda Genuina II. Flabellifera.—
Siboga-Expeditie Monografie 32C:121—232.

Nobili, G. 1905. Decapodi e isopodi della Nuova
Guinea tedesca, raccolti dal Sign. L. Biro —An-
nales Musei Nationalis Hungarici 3:480—507.

. 1906. Diagnoses préliminaires de crustacés,
décapodes et isopodes nouveaux recueillis par
M. le Dr. G. Seurat aux files Touamotu.—Bul-
letin du Muséum National d’ Histoire Naturelle,
Paris 1906, 5:256—270.

Nordenstam, A. 1946. Marine Isopoda from Professor
Dr. Sixten Bock’s Pacific Expedition 1917-
1918.— Arkiv f6r Zoologi 37A(7):26—29.

Richardson, H. 1902. The marine and terrestrial iso-
pods of the Bermudas, with descriptions of new
or little known species.—Proceedings of the
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Schultz, G. A. 1978. Protallocoxoidea new superfam-
ily (Isopoda Asellota) with a description of Pro-
tallocoxa weddellensis new genus, new species

from the Antarctic Ocean.—Crustaceana 34(3):

245-250.

. 1979. A new asellote (Stenetriidae) and two,

one new, Anthuridea (Anthuridae) from Ber-

muda (Crustacea, Isopoda).—Proceedings of

the Biological Society of Washington 91:904—

Salle

. 1982. Species of Protallocoxoidea and Ste-
netrioidea (Isopoda, Asellota) from the Antarc-
tic and southern seas. Biology of Antarctic Seas
10.—Antarctic Research Series, 32(2):17—62.

Serov, P. A., & G. D. E Wilson. 1995. A review of the
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82.

Stebbing, T. R. R. 1900. On Crustacea brought by Dr.
Willey from the South Seas.—Willey’s Zoolog-
ical Results 5:605—690.

Williams, E. H. Jr, & L. Bunkley-Williams. 1992.
Renocila loriae and R. richardsonae (Crustacea:
Isopoda: Cymothoidae), external parasites of
coral reef fishes from New Guinea and the Phil-
ippines.—Proceedings of the Biological Society
of Washington 105:299-309.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(1):175—180. 1999.

Caecidotea simulator, a new subterranean isopod from the Ozark
Springfield Plain (Crustacea: Isopoda: Asellidae)

Julian J. Lewis

217 W. Carter Avenue, Clarksville, Indiana 47129 U.S.A.

Abstract.—With the description of Caecidotea simulator, a phreatobitic iso-
pod reported herein from Arkansas and Kansas, a total of six species of sub-
terranean asellids is now known from the Springfield Plain section of the Ozark
Plateau. The presence of C. macropropoda in Arkansas, a subterranean species
previously known with certainty only from Oklahoma, is confirmed by two
collections examined from northeastern Arkansas.

The Springfield Plain is a region of gent-
ly rolling karst landscape formed on flat-
bedded limestones in the southwestern part
of the Ozark Plateau. This area of the cen-
tral United States includes parts of south-
western Missouri, northwestern Arkansas,
northeastern Oklahoma, and a tiny piece of
southeastern Kansas. Previously reported
from the Springfield Plain were the subter-
ranean asellids Caecidotea ancyla (Flem-
ing, 1972), C. antricola Creaser (1931), C.
macropropoda Chase & Blair (1937), C.
stiladactyla Mackin & Hubricht (1940), and
C. steevesi (Fleming 1972). The type spec-
imens of C. steevesi were examined and
three species were found to be present: (1)
C. steevesi from Carrico Cave, Dade Coun-
ty, Missouri (type-locality), and Gitten
Down Mountain Cave, Adair County,
Oklahoma; (2) C. antricola, also in the Car-
rico Cave collection; and (3) the new spe-
cies described below from Baxter Springs,
Cherokee County, Kansas.

Caecidotea simulator, new species
Figs. 1, 2a, c-f, 3a, b

Asellus steevesi Fleming 1972: 491-494
[Baxter Springs, Kansas record]; 1973:
295, 300 [in part].

Material examined.—Kansas: Cherokee
County, Baxter Springs, seeps off 7th Av-
enue, 12 Jun 1964, J. R. Holsinger, 10d d,

13 2 2 .—Arkansas: Washington County, O.
A. Lasterling’s well, 0.25 mile west High-
way 71, Fayetteville, 22 Jul 1965, E. H.
Schmitz, 15¢:6 2 ¢e

An 11.0mm 6 from Baxter Springs,
Kansas is designated as the holotype
(USNM 216971), with paratypes from Bax-
ter Springs (USNM 222477) and Laster-
ling’s well (USNM 216972). All of the ma-
terial has been deposited in the collection
of the National Museum of Natural History,
Smithsonian Institution.

Description.—Eyeless, unpigmented,
longest male to 19mm, female to 13mm.
Body slender, linear, about 6.5X as long as
wide. Margins of head, pereonites, and
pleotelson moderately setose. Head about
1.5X as wide as long, anterior margin con-
cave, postmandibular lobes moderately pro-
duced. Pleotelson about 2X as long as wide,
sides subparallel, caudomedial lobe slightly
produced.

Mandibles with 4-cuspate incisors and la-
cinia mobilis, palp with rows of plumose
setae on distal segments. Maxilla 1 with 5
robust plumose setae on inner lobe, 13
spines on outer lobe. Antenna 1 flagellum
to 18 segments, esthete formula 3-0-1-0-1.
Antenna 2, last segment of peduncle about
1.2X length of preceding segment, flagel-
lum of 19mm ¢ with 168 segments.

Male pereopod 1, propus 1.5 as long as

176 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Caecidotea simulator, male from Baxter Springs, Cherokee Co., Kansas: (a) habitus, (b) distal seg-
ments of antenna 1 flagellum, (c) right mandible, incisor, (d) left mandible, incisor and lacinia mobilis, (e)
mandibular palp, (f) maxilla 1, outer lobe, (g) same, inner lobe, (h) pleopod 3, (i) pleopod 5, G) pereopod 4.

VOLUME 112, NUMBER 1

177

Big.. 2.
steevesi, (b, g) male from Carrico Cave, Dade Co., Missouri: (a) pleopod 1, (b) same, (c) gnathopod propus,
palmar margin, (d) pleopod 2, (e) same, endopod tip, (f) same, caudal processes folded over anterior processes
under coverslip, (g) pleopod 2, endopod tip.

wide, palm with raised proximal spine, me-
dian triangular process and slightly bicuspid
distal process close together, sexual dimor-
phism absent. Pereopods 2—7 with moderate
setation as figured, sexual dimorphism for
clasping slight with male pereopod 4 carpus
2.6X as long as wide female 2.9.

Male pleopod 1, protopod about 0.6
length of exopod, with 4 retinacula. Exopod
about 1.6X as long as wide; lateral margin

Caecidotea simulator, (a, c—f) male from Baxter Springs, Cherokee Co., Kansas, and Caecidotea

concave, distal margin with 6—8 long plu-
mose setae. Pleopod 2, protopod with 3 me-
sial setae. Exopod, proximal segment with
about 5 lateral setae, distal segment with
about 15 long plumose setae along margin.
Endopod with distinct basal apophysis, en-
dopod tip twisted in appearance, processes
directed away from axis of endopod, can-
nula tapering to a stylet, mesial process ta-
pering and becoming digitiform, decurved

178 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Fourth pleopod exopods of related subterranean Caecidotea from the Springfield Plain: (a) C. sim-
ulator, Baxter Springs, Cherokee Co., Kansas, (b) C. simulator, Lasterling’s well, Washington Co., Arkansas,
(c) Caecidotea stiladactyla, small seeps 4.0 m. S. Boxley, Newton Co., Arkansas, (d) C. steevesi, Gittin Down
Mountain Cave, Adair Co., Oklahoma, (e) C. steevesi, Carrico Cave, Dade Co., Missouri, (f) C. steevesi, War
Eagle Cave, Madison Co., Arkansas.

VOLUME 112, NUMBER 1

mesiad, caudal process heavily sclerotized,
thickened laterally and mesially. Pleopod 3
exopod, proximal segment about 9.7X
length of distal segment, with about 6 plu-
mose setae on distal margin. Pleopod 4 with
up to 18 proximolateral setae, two false su-
tures present. Pleopod 5 with 2 sutures.
Uropods of male about 2 length of pleo-
telson, equal to length of pleotelson in fe-
male.

Etymology.—The noun ‘‘simulator’’,
from the Latin meaning imitator or pretend-
er, indicates the close resemblance of C.
simulator to C. steevesi. The vernacular
name suggested for this species is the
Springfield Plain groundwater isopod.

Habitat and range.—Caecidotea simu-
lator is known from the type-locality in
southeastern Kansas and one locality in the
adjacent corner of northwestern Arkansas.
From its vermiform, eyeless, unpigmented
appearance C. simulator is clearly an in-
habitant of subterranean waters. Caecidotea
simulator has not been found in caves and
presumably lives in the saturated soil inter-
Stices that supply groundwater to the two
sites from which it has been taken, a well
and a seep spring.

Relationships.—Caecidotea simulator
and C. steevesi are very similar morpholog-
ically. The gnathopods of both species are
nearly identical. The apex of the first ple-
opod exopod has elongate plumose setae in
both species, a characteristic also shared
with all species of the Hobbsi Group (Lew-
is, 1982). In C. simulator these setae occur
all the way across the distal margin of the
pleopod and are about 7—8 in number. In C.
steevesi the setae are located only on the
mesial half of the distal margin and number
4—5 (Fig. 2b).

Due to torsion of the second pleopod en-
dopod the taxonomically important tip pro-
cesses are difficult to interpret and compare
in C. simulator and C. steevesi, a charac-
teristic shared with C. macropropoda and
C. stiladactyla. The critical features are not
easily seen without applying a coverslip,
and the positions of the tip processes are

179

quite distorted under the pressure of the
glass (as the endopod twists). The endopod
tips illustrated in Fig. 2 are the result of
numerous attempts to gain the same per-
spective for both C. simulator and C. steev-
esi. The endopods appear to be fundamen-
tally similar, although side by side compar-
ison (Fig. 2e—g) illustrates minor differenc-
es of questionable importance. For
example, the mesial process in C. simulator
tapers to a narrow, cylindrical digitiform
process, while in C. steevesi it is wider and
obliquely truncate.

The best way to separate C. simulator
from C. steevesi is by the structure of the
fourth pleopod exopod (Fig. 3a—b, d—f),
which has two false sutures in C. simulator
(and C. macropropoda, Lewis, 1982, Fig.
3h), but only a single sigmoid suture in C.
steevesi (and C. stiladactyla, Fig. 3c).

Caecidotea macropropoda Chase & Blair,
1937

Material examined.—Arkansas: Carroll
County, White River at Beaver Down, G.
C. Kephart, Jul 1978, 4656, 52 2.—Wash-
ington County, spring 2.2 miles north
Dutch Mills, L. Hubricht, 21 May 1942,
633,522; seep, 1.5 miles north Winslow,
L. Hubricht, 22 May 1940, 31466, 722.

Remarks.—Lewis (1982) synonymized
C. ozarkana with C. macropropoda and re-
described the species. Dearolf (1953) re-
ported this species from two sites in Arkan-
sas, but the validity of the records was un-
known (Lewis, 1982). The records provided
here substantiate those of Dearolf for the
occurrence of C. macropropoda in Arkan-
sas. The species is endemic to the Spring-
field Plain, where it is found in cave
streams, springs, and seeps.

Acknowledgments

The late Dr. Thomas E. Bowman provid-
ed the loan of type-specimens of Caecido-
tea steevesi from the collections of the
Smithsonian Institution, read the manu-

180 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

script, and offered suggestions for its im-
provement.

Literature Cited

Chase, H. D., & A. P. Blair. 1937. Two new blind
isopods from northeastern Oklahoma.—Ameri-
can Midland Naturalist 18:220—224.

Creaser, E. P. 1931. A new blind isopod of the genus
Caecidotea, from a Missouri cave.—Occasional
Papers of the Museum of Zoology, University
of Michigan 222:1-7.

Dearolf, K. 1953. The invertebrates of 75 caves in the
United States ——Pennsylvania Academy of Sci-
ence 27:225-241.

Fleming, L. E. 1972. Four new species of troglobitic
asellids (Crustacea: Isopoda) from the United
States.—Proceedings of the Biological Society
of Washington 84(57):489—SO00.

Lewis, J. J. 1982. A diagnosis of the Hobbsi Group,
with descriptions of Caecidotea teresae, new
species, and C. macropropoda Chase and Blair
(Crustacea: Isopoda: Asellidae).—Proceedings
of the Biological Society Washington 95:338—
346.

Mackin, J. G., & L. Hubricht. 1940. Descriptions of
seven new species of Caecidotea (Isopoda,
Asellidae) from the central United States.—
Transactions of the American Microscopical So-
ciety 59:383-—397.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(1):181-188. 1999.

Collocherides brychius, a new species (Copepoda: Siphonostomatoida:
Asterocheridae) from a deep-water hydrothermal site in the
northeastern Pacific

Arthur G. Humes

Boston University Marine Program, Marine Biological Laboratory, Woods Hole, Massachusetts
02543, U.S.A.

Abstract.—Collocherides brychius is described from a depth of 2253 m at
a hydrothermal site on the Juan de Fuca Ridge in the northeastern Pacific. The
species is close to C. astroboae Stock 1971, but may be distinguished from
that species by its relatively small size (length of female 0.57 mm) and by the
inner terminal seta on the caudal ramus of the female being approximately as
long as the caudal ramus, instead of more than 3 times its length. Although its
three congeners are associated with ophiuroid echinoderms in shallow water,
the new species was recovered free in microfaunal samples in the deep sea.

The copepod genus Collocherides Stock
1971 (Siphonostomatoida: Asterocheridae),
contains three species. Collocherides astro-
boae Stock 1971, lives in the stomach of
the basket star Astroboa nuda (Lyman) at
Eilat in the Gulf of Aqaba and at the Dahlak
Archipelago (depth about 50 cm) in the Red
Sea (Stock 1971). It has also been recov-
ered from the stomach of Astroboa alba-
trossi Déderlein in Indonesia (precise lo-
cality unknown but probably Java Sea)
(Stock 1971). Large numbers of C. astro-
boae were found on A. nuda in 18 m at
Nosy Bé, northwestern Madagascar (Humes
1973). Collocherides singularis Humes
1986, lives with Astroboa nuda (depth 5 m)
at Poelau Gomumu, Moluccas, 01°50’00’S,
127°30'54"E (Humes 1986). Collocherides
bleptus Humes 1993, occurs intertidally on
the ophiuroid Macrophiothrix sp. at Nosy
Bé, northwestern Madagascar (Humes
1993):

Sixty-seven species of copepods have
been recorded in deep water from hydro-
thermal vents and cold seeps in the world’s
oceans (Humes & Segonzac 1998). These
species consist for the most part of siphon-
ostomatoids, with fewer calanoids, miso-

phrioids, cyclopoids, poecilostomatoids,
and harpacticoids. In this paper a new spe-
cies of the siphonostomatoid genus Col-
locherides is described from a deep-water
hydrothermal site in the northwestern Pa-
cific. With the addition of this new species,
a new erebonasterid poecilostomatoid re-
ported by Martinez Arbizu (1999), and a
new aegisthid harpacticoid described by
Conroy-Dalton & Huys (1999), the number
of copepods known from deep-sea hydro-
thermal vents and cold seeps rises to 70.

Materials and Methods

The copepods were collected at a low
temperature vent, where the highest reading
was 52° Celcius. The vent (Marker M) was
situated on new lava flows, and is believed
to be relatively young. (Vents at older ma-
ture lavas are believed to support more het-
erogeneous faunas, with more species, than
young vents (Milligan & Tunnicliffe
1994).) The diving submersible Alvin used
its arm or “‘claw’’ to gather tube worms
(Vestimentifera) and associated fauna and
deposit them in a biobox mounted on a bas-
ket. The sample was preserved for later

182

sorting, during which copepods were recov-
ered (Tsurumi, pers. comm.).

The copepods, which had been preserved
in ethanol, were cleared and dissections
made in lactic acid, using the wooden slide
method described by Humes & Gooding
(1964). All figures were drawn with the aid
of a camera lucida. The letter after the ex-
planation of each figure refers to the scale
at which it was drawn.

Siphonostomatoida Thorell, 1859
Asterocheridae Giesbrecht, 1899
Collocherides Stock, 1971
Collocherides brychius, new species
Figs. 1-3

Type material.—9 22, 3 36, in 2253
m, Juan de Fuca Ridge, Segment Cleft,
North Field, Vent Marker M, northeastern
Pacific, 44°58.97'N, 130°12.35’W, 28 Aug
1990. Holotype 2 (USNM 243645), allo-
type ¢6 (USNM 243646), and 6 paratypes
(5 22,1 35) (USNM 243647) deposited in
the National Museum of Natural History,
Smithsonian Institution, Washington, D.C.
Remaining specimens (dissected) in the col-
lection of the author.

Female.—Body slender (Fig. 1a), not
flexed between prosome and urosome (Fig.
1b). Average length (not including setae on
caudal rami) 0.57 mm (0.55—0.58 mm) and
average width 0.18 mm (0.17—0.19 mm),
based on 9 specimens. Dorsoventral thick-
ness (at level of slight protuberance be-
tween maxillipeds and first pair of legs) 127
4m. Epimera of metasomal somites round-
ed (Fig. 1b). Ratio of length to width of
prosome 2.03:1. Ratio of length of prosome
to that of urosome 1.31:1.

Urosome with 5 somites. Somite bearing
leg 5 (Fig. 1c) 47 X 83 wm. Genital double-
somite in dorsal view 81 wm long and 81
4m wide at widest part. In lateral view (Fig.
1d), this double-somite with dorsal and ven-
tral sides slightly rounded. Genital areas lo-
cated laterally in anterior half of double-
somite (Fig. lc, d). Each genital area with
small seta and minute spine (Fig. 1d). Three

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

postgenital somites from anterior to poste-
rior 36 X 53, 31 X 40, and 23 X 36 wm.
Genital double-somite and 3 postgenital so-
mites with pairs of small posterolateral
spinelike processes or spinules (Fig. 1d, e),
those on anal somite prominent.

Caudal ramus (Fig. le) moderately elon-
gate, unornamented, length including ter-
minal pointed process 44 ym, length with-
out process 35 wm, width 16 wm. Ratio of
length (without process) to width 2.19:1, ra-
tio including process 2.75:1. With 6 smooth
setae, 1 long and 5 short, longest seta 52
wm long, much longer than other setae and
slightly swollen proximally. Long seta
slightly longer than ramus, ratio 1.18:1.

Body surface with few sensilla on uro-
some, otherwise unornamented.

Egg sac (Fig. 1d), seen on only 1 female,
large, oval, 179 X 99 wm.

Rostral area (Fig. 1f) triangular, not pro-
truding. Antennule (Fig. 2a) 20-segmented,
187 wm long, with aesthetasc 60 wm long
on segment 18. Formula for armature: 1, 2,
Ly 2», 2m 25 25 23/0; 25 Dy 2,625 2p ee ee
1 aesthetasc, 2, and 9. All setae smooth.
Antenna (Fig. 2b) 99 wm long, including
terminal spine 22 «zm. First segment (coxa)
short and unarmed. Second segment (basis)
with 1 minute seta (exopod). Third segment
long and unarmed. Short fourth segment
bearing 2 short setae and 1 long terminal
Spine, its truncated tip with few extremely
minute setules.

Oral cone (Fig. 1f) short, oval in ventral
view, prominent in lateral view (Fig. 1b).

Mandible (Fig. 2c) with slender 2-jointed
palp 60 pm long. Gnathobase (Fig. 2d) 50
wm long., with several blunt terminal teeth.
Maxillule (Fig. 2e) with 1 seta on slender
outer lobe and 4 setae on stout inner lobe.
Maxilla (Fig. 2f) 2-segmented, first segment
unarmed, second segment bearing recurved
claw with truncated minutely spinulose tip.
Maxilliped (Fig. 2g) 5-segmented. First
segment with inner distal seta. Second seg-
ment elongate (52 1m long) with 1 seta on
inner margin. Third and fourth segments
short with 1 seta. Fifth segment elongate

VOLUME 112, NUMBER 1 183

Fig. 1. Collocherides brychius, new species. Female. a, body, dorsal (scale A); b, body, lateral (A); c,
urosome, dorsal (B); d, urosome, lateral (B); e, anal somite and caudal ramus, ventral (C); f, cephalosome,
ventral (B).

184 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

0.05 mm

Fig. 2. Collocherides brychius, new species. Female. a, antennule, ventral (scale C); b, antenna, dorsal (C);
c, mandible, ventral (C); d, gnathobase of mandible, flat view (D); e, maxillule, anterior (C); f, maxilla, posterior
(C); g, maxilliped, posterior (C); h, leg 1 and intercoxal plate (E); i, leg 2 and intercoxal plate, posterior (E); j,
leg 3 and intercoxal plate, anterior (E); k, leg 4 and intercoxal plate, anterior (E); 1, leg 5, ventral (C).

VOLUME 112, NUMBER 1

Fig. 3. Collocherides brychius, new species. Male. a, body, dorsal (scale A); b, urosome, dorsal (B); c,
antennule, ventral (C); d, leg 5, ventral (C); e, somites 1-3 of urosome, showing legs 5 and 6, ventral (E).

186 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

with 1 distal seta. Claw (26 wm long) with
truncated tip bearing minute spinules.

Ventral area between maxillipeds and
first pair of legs (Fig. 1f) slightly protuber-
ant (Fig. 1b).

Legs 1—4 (Fig. 2h—k) biramous with 3-
segmented rami. Formula for spines (Ro-
man numerals) and setae (Arabic numerals)
as follows:

P, coxaQ@=0> phases 1-0 expll; 741; -1,2,2
enpO:l:. 0-2; 25
P, coxa0-0 basis1-0 explI-l; I[-1; ULI4
enp@-l:- ~O-25 fe
P, coxa@-0-_-basisl1-0 expll; El; /MEES
enpO-1; O-2; 1,1,3
P, coxaG-0" basist-0 expl-i: (fel; ,.LES
enp 0-1 G-2¢ AT?

Inner coxal seta absent in all 4 legs. Out-
er margins of exopod segments of all 4 legs
with very small spinules.

Leg 5 (Fig. 21) 2-segmented. First seg-
ment with outer seta and produced medially
as broad triangular flap, pointed at tip, and
having inner and outer setules. Second seg-
ment 31 X 14 um, ratio 2.2:1, bearing 4
setae and ornamented along inner margin
with few setules.

Leg 6 represented by seta and small spi-
niform process on genital area (Fig. 1d)

Color of living specimens unknown.

Male.—Body (Fig. 3a) elongate, slender,
and, as in female, not flexed. Length (not
including setae on caudal rami) 0.50 mm
(0.50—0.51 mm) and greatest width 0.15
mm (0.15—0.16 mm), based on 3 speci-
mens. Greatest dorsoventral thickness at
level of small ventral protuberance 96 wm.
Ratio of length to width of prosome 1.98:
1. Ratio of length of prosome to that of uro-
some 1.36:1.

Urosome (Fig. 3b) with 6 somites. So-
mite bearing leg 5 32 X 70 wm. Genital
somite 60 X 83 wm, with rounded lateral
margins. Four postgenital somites from an-
terior to posterior 34 X 49, 32 X 39, 26 X
32, and 21° 3)+jm-.

Caudal ramus similar to that of female,

37 X 15 pm including terminal process. In-
ner terminal seta 143 pm long, relatively
much longer than in female, 3.86 times lon-
ger than ramus.

Rostral area like that of female. Anten-
nule (Fig. 3c) 18-segmented, geniculate.
Aesthetasc on segment 17 59 um long. Ar-
mature: 1, 2352/2, 2, 2; 2-276, 23232=——
2, 2, 1, 1 + aesthetasc, and 9. Antenna as
in female.

Oral cone, mandible, maxillule, maxilla,
maxilliped, and legs 1—4 like those of fe-
male.

Leg 5 (Fig. 3d) with second segment 20
x 12 wm, ratio 1.67:1, bearing 3 slender
outer setae and 2 broad, hyaline, inner se-
tae, 33 wm, with blunt tips.

Leg 6 (Fig. 3e) usual posteroventral flap
on genital somite bearing 2 setae.

Color of living specimens unknown.

Etymology.—The specific name bry-
chius, from Greek brychios, meaning from
the depths of the sea, refers to the depth at
which specimens of the new species was
found.

Remarks.—The female of Collocherides
brychius may be differentiated from its

three congeners as follows:

From C. astroboae Stock 1971, in which
the long terminal seta on the caudal ramus
is approximately 3.6 times longer than the
ramus, and the body length is 0.62 mm
(measurements based on an average of five
specimens from Astroboa nuda from the
Dahlak Archipelago (Zodlogisch Museum
Amsterdam, cat. no. 101.090).

From C. singularis Humes, 1986, in
which the caudal ramus lacks a terminal
process, the long terminal seta on the cau-
dal ramus is approximately 1.8 times longer
than the ramus, the second segment of the
endopod of legs 1—3 has one inner seta, and
the average body length is 0.64 mm.

From C. bleptus Humes, 1993, in which
the ratio of the innermost terminal seta on
the caudal ramus to the length of the ramus
itself is approximately 2.51:1, the genital
double-somite is 120 X 80 um, distinctly
longer than wide, the average body length

VOLUME 112, NUMBER 1

is 0.77 mm, and the ventral surfaces of the
genital and postgenital somites have nu-
merous small scalelike spines.

In C. brychius sexual dimorphism occurs
in the length of the inner terminal seta on
the caudal ramus, which in the female is 42
wm, but in the male is 143 wm and less
swollen proximally than in the female.

The four species of Collocherides may
be further distinguished by the following
key.

Key to females of the genus Collocherides

1. Genital double-somite distinctly longer
than wide, ratio 1.4:1; body length 0.77
mm (0.75—0.78 mm) C. bleptus
Genital double—somite quadrate or near-
ly so, ratio approximately 1:1; body
length not exceeding 0.67 mm

2. Long inner terminal seta on caudal ra-
mus 3.6 times longer than ramus
et ee ee ee ee C. astroboae
Longest seta on caudal ramus 1.18 times

ee © © © ee ee

ee e ee

cor less) longer than ramus .......... 3
3. Body length 0.57 mm (0.55—0.58 mm);

legs 1—3 with second segment of endo-

pod having 2 inner setae ..... C. brychius

Body length 0.64 mm (0.63—0.65 mm);
legs 1—3 with second segment of endo-
Poe with 1 inner scta....... C. singularis

The ecological and host relationships
within the genus Collocherides are poorly
known. Collocherides astroboae, C. singu-
laris, and C. bleptus are associated with
shallow-water ophiuroids, while C. bry-
chius lives in deep—water hydrothermal
sites and was recovered free in meiofaunal
samples. However, much more information
is needed to understand these relationships.
The possibility that there may have been
ophiuroids at the deep-sea vent site cannot
be excluded. Ophiuroids, known from abys-
sal depths (e.g., Lauermann and Kaufman
1998), have been reported from deep-sea
hydrothermal sites (Hessler and Smithey
1983; Grassle 1986; Tunnicliffe 1991,
1998; Segonzac 1992; Sibuet & Olu 1998).
A new genus and species of ophiuroid,
Ophioctenella acies, was described from

187

the Mid-Atlantic Ridge by Tyler et al.
(1995). However, no ophiuroids are yet
known from Juan de Fuca vents, where the
C. brychius was found (Tunnicliffe, pers.
comm.).

Acknowledgments

I thank Dr. Verena Tunnicliffe and Maia
Tsurumi, Department of Biology, Univer-
sity of Victoria, British Columbia, for send-
ing the specimens of C. brychius to me for
study. I also thank Dirk Platvoet, Zodlo-
gisch Museum, Amsterdam, for arranging
the loan of Stock’s specimens of Astroboa
nuda from Dahlak Archipelago.

Literature Cited

Conroy-Dalton, S., & R. Huys. 1999. A new genus of
Aegisthidae (Copepoda: Harpacticoida) from
hydrothermal vents on the Galapagos Rift.—
Journal of Crustacean Biology (in press).

Hessler, R. R., & W. M. Smithey, Jr. 1983. The distri-
bution and community structure of megafauma
at the Galapagos Rift hydrothermal vents.—ZJ/n
Hydrothermal processes and seafloor spreading
centers. P. A. Rona, K. Bostr6m, L. Laubier, &
K. L. Smith, Jr, eds., NATO conference series
4:735—770. Plenum Press, New York.

Humes, A. G. 1973. Cyclopoid copepods associated
with the ophiuroid Astroboa nuda in Madagas-
car.—Beaufortia 21:25-35.

. 1986. Two new species of Copepoda associ-

ated with the basket star Astroboa nuda (Ophiu-

roidea) in the Moluccas.—Zoologica Scripta 15:

323-332.

. 1993. Collocherides bleptus n. sp. (Copepoda:

Siphonostomatoida) associated with an intertid-

al ophiuroid in Madagascar.—Bijdragen tot de

Dierkunde 63:121—127.

, & R. U. Gooding. 1964. A method for study-

ing the external anatomy of copepods.—Crus-

taceana 6:238—240.

, & M. Segonzac. 1998. Copepoda from deep-
sea hydrothermal sites and cold seeps: descrip-
tion of a new species of Aphotopontius from the
East Pacific Rise and general distribution.—Ca-
hiers de Biologie Marine 39:51—62.

Lauermann, L. M. L., & R. S. Kaufman. 1998. Deep-
sea epibenthic echinoderms and a temporally
varying food supply: results from a one year
time series in the N. E. Pacific-—Deep-Sea Re-
search II 45:817-—843.

Martinez Arbizu, P. 1999. New Erebonasteridae (Co-
pepoda) from Vilkitzky Strait in the Arctic and

188 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

from a Pacific hydrothermal vent site (northern
Fiji Basin).—Journal of Crustacean Biology 19:
93-105.

Milligan, B. N., & V. Tunnicliffe. 1994. Vent and non-
vent faunas of the Cleft segment, Juan de Fuca
Ridge, and their relations to lava age.—Journal
of Geophysical Research 99:4777—4786.

Segonzac, M. 1992. Les peuplements associés a
Vhydrothermalisme océanique du Snake Pit
(dorsale médio-atlantique; 23°N, 3480 m): com-
position et microdistribution de la mégafau-
ne.—Comptes Rendus de 1’Académie des Sci-
ences Paris 314 (série 3):593—600.

Sibuet, M., & K. Olu. 1998. Biogeography, biodiver-
sity and fluid dependence of deep-sea cold-seep
communities at active and passive margins.—
Deep-Sea Research II 45:517—567.

Stock, J. H. 1971. Collocherides astroboae n. gen., n.
sp., a siphonostome cyclopoid copepod living
in the stomach of basket stars —Bijdragen tot
de Dierkunde 41:19-—22.

Tunnicliffe, V. 1991. The biology of hydrothermal
vents: ecology and evolution.—Oceanography
and Marine Biology 29:319—407.

, A. G. McArthur, & D. McHugh. 1998. A bio-
geographical perspective of the deep-sea hydro-
thermal vent fauna.—Advances in Marine Bi-
ology 34:353-—442.

Tyler, P. A., G. J. L. Paterson, M. Sibuet, A. Guille, B.
J. Murton, & M. Segonzac. 1995. A new genus
of ophiuroid (Echinodermata: Ophiuroidea)
from hydrothermal mounds along the Mid-At-
lantic Ridge.—Journal of the Marine Biological
Association of the United Kingdom 75:977—
986.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(1):189-198. 1999.

Cymbasoma californiense, a new monstrilloid (Crustacea: Copepoda:
Monstrilloida) from Baja California, Mexico

E. Sudrez-Morales and R. Palomares-Garcia

(ES-M) El Colegio de la Frontera Sur (ECOSUR)-Unidad Chetumal. A.P. 424. Chetumal,
Quintana Roo. 77000. Mexico; (RP-G) Centro Interdisciplinario de Ciencias Marinas
(CICIMAR). COFAA. A.P. 592. La Paz, B.C.S., Mexico

Abstract.—Cymbasoma californiense is described from three adult females
collected during three cruises carried out in an area adjacent to Bahia Mag-
dalena, a large coastal system on the southwest coast of the Baja California
Peninsula, and in the southern Gulf of California. The new species is very
closely related to at least three other Cymbasoma (C. longispinosum (Bourne)
and its allies), but can be distinguished by a combination of features including:
the presence of cuticular striations on the forehead, cephalothorax, and genital
somite; relatively long antennules; the structure of the fifth legs, and the relative
length of the ovigerous spines. Cymbasoma californiense is the first species of
this genus recorded in the Californian region and the Eastern Tropical Pacific.

Monstrilloids are protelean parasitic co-
pepods which are free-living only as first
nauplii and adults (Raibaut & Trilles 1993).
The first nauplius is the infective stage, but
later stages complete their development as
endoparasites of benthic organisms such as
polychaetes and prosobranch molluscs.
Adults are free-swimming, exclusively re-
productive, and have no feeding append-
ages (Isaac 1975; Davis 1984).

The number of described species has
been estimated as about 90 (Grygier
1994a), and they are grouped into three rec-
ognized genera (Monstrilla, Monstrillopsis,
and Cymbasoma). The generic nomencla-
ture of the group is still unclear, and efforts
have been made to solve this problem, par-
ticularly with regard to the genus Cymba-
soma (Grygier 1994a, 1997). This genus
contains now about 57 described species,
with about half of them being described un-
der the invalid genus name Thaumaleus
(Razouls 1996).

There are several (six) records of mon-
strilloid copepods in the Northeastern Pa-
cific region (Park 1967, Grygier 1995, Ra-
zouls 1996). However, only three species of

monstrilloid copepods have been recorded
in lower latitudes, such as the California re-
gion or the Eastern Tropical Pacific. All
three regional records are of the genus
Monstrilla. Monstrilla capitellicola Hart-
man, 1961 was recorded at La Jolla Canyon
as a parasite of a polychaete of the genus
Capitella (Hartman 1961), and was de-
scribed upon V stage copepodids. Monstril-
la spinosa Park, 1967, originally described
from the Strait of Georgia, was recorded by
Sudrez-Morales & WVdasquez-Yeomans
(1996) off Bahia de Todos Santos, Baja
California, and M. gibbosa Suarez-Morales
& Palomares-Garcia, 1995 was collected on
the southeastern coast of the Baja Califor-
nia Peninsula (Suadrez-Morales & Paloma-
res-Garcia 1995).

As part of a survey of the main coastal
systems along the southern portion of the
Baja California Peninsula, plankton sam-
ples were collected at Bahia de Magdalena,
a large bay located on the western coast.
Our taxonomic analysis of the copepods in
the samples revealed the presence of several
monstrilloids which turned out to belong to
an undescribed species. The description of

190

the new species is presented herein follow-
ing the standards set by Grygier & Ohtsuka
(1995) for descriptions of monstrilloid co-
pepods.

Materials and Methods

Plankton samples were collected during
the BAMA8611, BAMA9710, and CER-
RALVO9609 cruises. The first two were
carried out in Bahia Magdalena (BAMA),
a large coastal system on the southern por-
tion of the western coast of the Baja Cali-
fornia Peninsula, Mexico. The other cruise
took place in the southern portion of the
Gulf of California, along the southeastern
coast of the Baja California. One female of
the same previously undescribed species of
monstrilloid copepod as captured during
each of the three cruises: one at station K2
(BAMA9710), on 27 October 1997, another
at station R2 (BAMA9611), on 5 February
1996, and the third was collected at station
SIUG49 (CERRALVO9609), 20 September
1996 (Fig. 1). Samples were collected dur-
ing surface hauls with a standard plankton
net (0.333 mm mesh-size). Zooplankton
was fixed with a buffered formalin solution.
Monstrilloid copepods were sorted and
transferred to 70% ethanol. Observations
were made under a Zeiss microscope and
drawings were made with the aid of a cam-
era lucida. Standard terminology for cope-
pod morphology (Huys & Boxshall 1991)
and for monstrilloid antennular armature
(Grygier & Ohtsuka 1995) was followed.

Order Monstrilloida
Family Monstrillidae Dana, 1849
Cymbasoma Thompson, 1888
Cymbasoma californiense, new species,
Suarez-Morales

Material examined.—Holotype, 2.1 mm,
adult female, undissected, ethanol-pre-
served. Sta. K2 (coordinates in decimal no-
tation: 24.69°N, 112.07°W) BAMA9710
cruise, Bahia Magdalena, South Baja Cali-
fornia, Mexico, 27 Oct 1997. Vial deposited
in the Collection of Crustacea, National

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Museum of Natural History, Smithsonian
Institution, Washington D.C., under number
USNM-261422. Paratype adult female, par-
tially dissected, permanent slide, mounted
in CMC. Sta. SUG49 (24.27°N, 109.85°W)
CERRALVO9609 cruise, Isla Cerralvo,
Southern Gulf of California, Mexico, 20
Sep 1996. Deposited in the NMNH,
USNM-261422. Paratype adult female, to-
tal length: 2.03 mm, ethanol-preserved. Sta-
tion R2 (24.41°N, 111.65°W), BAMA8611
cruise, Bahia Magdalena, Baja California
Sur, Mexico, 5 Feb 1986. Vial deposited in
the Zooplankton Collection of El Colegio
de la Frontera Sur Unidad Chetumal, Méx-
ico, under number ECO-CH-Z-00369.

Type locality.—Bahia Magdalena, south-
ern portion of the eastern coast of the Baja
California Peninsula, Mexico. Water col-
umn.

Etymology.—The species is named after
the Californian region in which this species
was collected.

Description of female.—Average body
length of 3 individuals, 2.1 mm measured
from forehead to posterior margin of anal
somite. Cephalothorax (incorporating first
pedigerous somite) accounting for almost
65% of total body length (Fig. 2A, B).
Ventral margin of anteriormost portion of
cephalothorax slightly curved. Oral papilla
as in Fig. 2A, lying midventrally 0.21 of
way back along cephalothorax. Forehead
with strong cuticular striations mainly be-
tween antennule bases, and, more dorsally,
a circular, spiral pattern with two anterior
and two posterior whorls, present in all the
individuals. Nauplius eye present, weakly
developed, ocelli slightly pigmented with
rounded shape (Fig. 3E). Pair of low
bumps on ventral side of cephalothorax be-
tween antennule bases and oral papilla,
surrounded by circles with wrinkles ar-
ranged in spiral-shaped pattern. Cuticular
ornamentation also on lateral posterior
margin of cephalothorax: clear vertical
striations (Fig. 2B). Antennules four-seg-
mented, armed with 0, I; 1,V; 2, I; 10, VII
setae (Arabic numbers) and spines (Roman

VOLUME 112, NUMBER 1 191

25.0 32°
30°
24.9
28°
a) j
24.8 um
24°
24.7
PACIFIC OCEAN
22°
Magdalena Bay 1164 lam tas? 40" 108°
24.6 2
24.5
Almejas Bay
|
24.4 e
PACIFIC OCEAN SJ
24.3

Mook women tee 014119) |e aii die 1115

Cerralvo Island
24.3

24.2

24.1-

GULF OF CALIFORNIA

24.0
110.10 110.00 109.90 109.80

Fig. 1. Study area showing the sites in which specimens of the new species were collected.

2

Fig. 2.
spine.

numbers) (Fig. 3B, C). Terminal two
spines asymmetrical, forming chela-like
structure. Last segment with one small
aesthetasc on distal margin, large aesthe-
tasc (4aes) on proximal outer margin. Most
spines and setae of general pattern de-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

0.6 mm

Cymbasoma californiense n. sp., holotype female. A. habitus, lateral. B. habitus lateral. C. ovigerous

scribed by Grygier & Ohtsuka (1995) pre-
sent (Fig. 3C). Only setae b5 and b6 miss-
ing on last segment of right and left anten-
nules, setae bl1—4 unbranched. Ratio of
lengths of antennule segments: 11.3: 23.1:
14.3:51.3 = 100. Antennule slightly lon-

VOLUME 112, NUMBER 1 193

Fig. 3. Cymbasoma californiense n. sp., holotype female. A. genital complex and fifth legs, ventral. B right
antennule, dorsal. C. left antennule, dorsal, with armature following the nomenclature of Grygier & Ohtsuka
(1995). D. urosome and furcal rami, dorsal. E. forehead, dorsal.

194

ger than 17.5% of total body length, and
25.5% of cephalothorax length.
Incorporated first pedigerous somite and
three free succeeding pedigerous somites
(latter accounting for 22% of total length in
dorsal view) each bearing a pair of bira-
mous swimming legs (Fig. 4A—D). Endop-
odites and exopodites of legs 1—4 triarticu-
lated. Legs 1—4 decreasing in size posteri-
orly. Armature of swimming legs as:
basis endopodite
leg 1 1-0 0-1:0-1:1,2,2 1-1:0-1:L4,3

legs2—4 1-0 O-1;0-1;51,2,2 I-1;0-1;1,2,3

exopodite

Basis separated from coxa posteriorly by
diagonal articulation, with lateral hair-like
seta on legs 1—4; seta on leg 3 at least 1.5
times longer and noticeably thicker than the
others, and plumose, while others are sim-
ple. Inner seta on first exopodite segment of
legs 1-4 plumose, long, reaching to distal
margin of endopodite. Inner margin of first
endopodal segments and outer margin of
second and third exopodal segments of legs
1—4 with short hairs. Outer exopodal apical
seta of legs 1—4 with row of setules along
inner side, but bearing row of small, closely
spaced denticles along outer side. Most se-
tae biserially plumose.

Fifth legs joined medially; each with
rounded endopodal lobe. Exopodal lobe 2
times longer than wide, with two long,
equal plumose setae, and one short inner,
plumose seta, about 28% as long as the oth-
er two (Fig. 3A).

Urosome consisting of fifth pedigerous
somite, genital double somite, one free ab-
dominal (anal) somite, and furcal rami, al-
together representing 14.3% of total body
length. Genital double-somite slightly lon-
ger than one-third total length of urosome
(35%). Ratio of lengths of fifth pedigerous
somite, genital double somite, and free ab-
dominal somite: 37.5:35.5:26.6 = 100.
Genital somite with strong transverse cutic-
ular wrinkles on lateral margins and dorsal
surface (Fig. 3D). Medial portion of genital
double somite moderately swollen, bearing

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

long, basally conjoined, ovigerous spines.
Spines separate from level of posterior mar-
gin of furcal rami. Spines slender at their
base, swollen distally, in dorsal view, left
one of holotype shorter than right one (Fig.
4E). Spines about 1.4 times total body
length (Fig. 2C). Caudal rami short, 1.2
times longer than wide, moderately diver-
gent, bearing three strong, terminal setae, as
usual in genus. Middle seta slightly thicker
than others (Fig. 3D).

Male.—unknown.

Remarks.—The new species from Baja
California is assigned to the genus Cym-
basoma by virtue of the presence of two
urosomal somites in the female, with only
one free somite between the genital double
somite and the caudal rami (Isaac 1975).

According to the most comprehensive
key to the species of this genus (Isaac
1975), the Californian specimens would be
identified as females of C. longispinosum
(Bourne, 1890). However, other species
such as C. chelemense Suarez-Morales &
Escamilla, 1997, recently described from
the Gulf of Mexico, and C. morii Sekigu-
chi, 1982, redescribed by Grygier (1994b)
from Japan are closely related to and can
also be confused with C. longispinosum.
The female specimens of C. californiense
share several features with C. longispinos-
um, C. morri, and C. chelemense, such as
the relatively long cephalothorax, the gen-
eral structure of the fifth legs, with a round-
ed endopodal lobe, the position of the oral
papilla, the antennule/cephalothorax length
ratio, the dorsal striations on the genital
double segment, the swollen tips and the
asymmetry of the ovigerous spines, and rel-
atively long ovigerous spines conjoined at
the base. However, the new species differ
from these other species in several key
structures generally regarded as important
in monstrilloid taxonomy, such as the mor-
phological details of the fifth legs, the struc-
ture of the genital complex, and the body
proportions.

In the females of C. californiense the
cephalothorax represents 65% of the total

VOLUME 112, NUMBER 1 19S

Fig. 4. Cymbasoma californiense n. sp., holotype female. A. right leg 1, posterior. B. left leg 2, posterior.
C. right leg 3, posterior. D. right leg 4, posterior. E. ovigerous spines, detail of distal portion.

196

body length; the figure for C. chelemense is
68%, while in C. longispinosum it is less
than 65%, and in C. morii the reported
range (Grygier 1994b) is 66—73%. In the
new species the oral papilla is relatively
much smaller and shows a different aspect
with respect to those of C. chelemense and
C. morii, but is similar to that of C. longis-
pinosum. The strong cuticular protuberanc-
es on the forehead of the new species are
shared only by C. chelemense, but are dif-
ferent; in the latter, the striations show a
more complex pattern. The forehead is only
slightly rugose in C. morii (see Grygier
1994b). Cymbasoma californiense shows
anteroventral knobs (nipples), which were
described for C. morii, and C. chelemense
and are probably also present in C. longis-
pinosum. However, in the new species these
knobs are reduced, not protruding as far as
those described for the other species.

The fifth leg’s endopodal lobe in Cym-
basoma longispinosum has a clearly trian-
gular-elongated shape (Giesbrecht 1892;
Sars 1921; Isaac 1975), while the same
structure is completely rounded in the new
Species, as it is in both C. morii (Grygier
1994b) and C. chelemense (Suarez-Morales
& Escamilla 1997). However, this structure
is relatively smaller in C. morii than in the
other two species.

The fifth legs inner seta in C. califor-
niense is relatively short in comparison
with those of the other species. It is about
28% as long as the other two setae (3.5
times shorter), while this figure is 62.5% in
C. chelemense (1.6 times shorter), 40% (2.5
times shorter) in C. morii, and 66% (1.5
times shorter) in C. longispinosum.

The armature of the antennules is similar
in the four species. The new species shows
the same antennular armament pattern as
described in C. morii and C. chelemense,
with only slight differences in the size and
position of some of the smallest setae. The
largest aesthetasc on the distal segment is
relatively shorter in C. californiense than it
is in C. chelemense and in C. morii. Pro-
portions of antennular segments are similar

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

in these species, with the terminal segment
representing more than 50% of the total an-
tennular length. However, the antennules in
the new species are relatively longer than
in the other species compared herein, rep-
resenting 25.5% of the cephalothorax
length versus 21—22% in the other three
species considered herein.

In C. californiense and C. chelemense,
the genital double somite’s posterior margin
is about 40% wider than the anterior margin
of the succeeding free (anal) somite. From
Giesbrecht’s (1892) illustrations, in C. lon-
gispinosum the proportional value is less
than half (16%) that measured for the new
species; in C. morii the value is around
30%, and the posterior margin of the genital
double somite is slender and somewhat
convex. This margin is slender in the other
species and clearly irregular in the new spe-
cies.

The proportional length of the ovigerous
spines differs among the four species. In C.
longispinosum and C. chelemense the
spines are only 14% longer than the body
(Giesbrecht 1892; Sars 1921; Sudarez-Mo-
rales & Escamilla 1997). In C. morii they
are much longer, about twice as long as the
body, whereas in the new species they are
42% longer than the body. All four species
have a long common base for the two ovig-
erous spines.

Cymbasoma californiense shows cuticu-
lar wrinkles along the lateral margins of the
fifth pedigerous and anal somites, and on
the posterior lateral margins of the cepha-
lothorax. These features are absent in the
other species and could be considered as a
key character to recognize this species. The
total body length recorded for C. califor-
niense (2.2 mm) is slightly under the range
reported for C. longispinosum and C. che-
lemense (2.3 to 3.16 mm) (Isaac 1975, Sua-
rez-Morales & Escamilla 1997), but is
slightly over the average length of C. morii
(2.18 mm) (Grygier 1994b).

There are several species of Cymbasoma
with a long, slender cephalothorax: C. mo-
rii (Sekiguchi, 1982), C. longispinosum

VOLUME 112, NUMBER 1

197

Table 1.—Comparison of features present in females of two groups of species of Cymbasoma with a relatively

long cephalothorax.

Species C. gigas C. morii C. chelemense C. longispinosum C. californiense C. bowmani C. reticulatum
Lobes on fifth 2 2 2 2 1 1
leg
Inner lob ovate rounded rounded subtriangular rounded — —
Nipples ? present present present reduced absent absent
Ovig. spines ? joined at —joined at joined at base _ joined at separated separated at
base base base at base base
O.S. longer ry 2 times 14% longer 14% longer 43% longer _ shorter shorter
than body
Striation on & present present not described present absent absent
gen. som.
Striation on 7 absent absent not described present absent absent
anal som.
Striation on ? present present not described present present absent
forehead (light) (strong) (strong) (light)

(Bourne, 1890), C. chelemense (Suarez-
Morales & Escamilla), 1997, Cymbasoma
gigas (A. Scott, 1909), C. reticulatum
(Giesbrecht, 1892), and C. bowmani (Sua-
rez-Morales & Gasca), 1998. As suggested
by Suarez-Morales & Gasca (1998), these
Cymbasoma species can be divided into
two groups (see Table 1). In the first group,
with a bilobed fifth leg, nipple-like ventral
cephalic protuberances, striated genital so-
mite, and basally fused genital spines
which are longer (1.5—2 times) than the
body, C. morii, C. longispinosum, C. che-
lemensis, probably C. gigas, and now C.
californiense, could be included. The sec-
ond group shares a single-lobed fifth leg,
the absence of nipple-like ventral protu-
berances, and basally separated genital
spines that are shorter than the body (C.
reticulatum, C. bowmani).

The lack of comparative biogeographical
information on monstrilloids is related to
several factors: the scarcity of the group in
plankton collections, the unsolved taxono-
my of the group, the uncertainty in recog-
nizing the male and female of the same spe-
cies from plankton samples, and the irreg-
ular putative distributional patterns
throughout the group (Isaac 1975). How-
ever, it is relevant to mention the known
geographical ranges of the four species

compared herein. The Mediterranean, Great
Britain, Mindanao, Portugal, Black Sea,
Gulf of Suez, Vietnam, and the Arabian
Gulf for C. longispinosum (Isaac 1975); the
wide distribution reported for this species
could be the result of overlooking closely
related species. This has been suggested by
Grygier & Ohtsuka (1995) for Monstrilla
helgolandica Claus (1863). Cymbasoma
morii has been recorded from Japan, sup-
posedly Vietnam, and probably India (Gry-
gier 1994b); C. chelemense from the south-
ern Gulf of Mexico (Suarez-Morales & Es-
camilla 1997), and C. californiense from
Baja California. Presumably, the geograph-
ical isolation of the new species with re-
spect to the others would favour the con-
ception of a different taxon.

This new species of Cymbasoma repre-
sents the second record of the genus in the
Northeastern Pacific region; the only pre-
vious record was of C. rigidum Thompson,
1888, near the Kodiak Island, Alaska
(Threlkeld 1977). It is also the first record
of the genus in the California region.

Acknowledgments

One specimen was obtained during the
operation of the project ‘‘Potencial de la
Isla Cerralvo, Baja California Sur como

198

area de reserva pesquera’’. Comments of
three anonymous reviewers improved an
earlier version of this manuscript.

Literature Cited

Davis, C. C. 1984. Planktonic Copepoda (including
Monstrilloida). Pp. 67-91. in K. A. Steidinger
& L. M. Walter, eds., Marine plankton life cycle
strategies. C. R. C. Press. Florida.

Giesbrecht, W. 1892. Systematik und Faunistik der pe-
lagischen Copepoden des Golfes von Neapel
und der angrenzenden Meeres-Abschnitte.—
Fauna Flora Golf. Neapel 19:1-831 + Atlas
(pls. 1-54).

Grygier, M. J. 1994a [dated 1993]. Identity of Thau-
matoessa (=Thaumaleus). typica Kr@yer, the
first described monstrilloid copepod.—Sarsia
78:235-242.

. 1994b. Nomenclature, redescription, and new

record from Okinawa of Cymbasoma morii Sek-

iguchi, 1982 (Monstrilloida).—Hydrobiologia
292/293:23-29.

. 1995. Annotated chronological bibliography

of Monstrilloida (Crustacea: Copepoda).—Gal-

axea 12:1—82.

. 1997. Monstrilla Dana, 1849 and Thaumaleus

Krgyer, 1849 (Crustacea, Copepoda): con-

served.—Bulletin of Zoological Nomenclature

54:131-132.

, & S. Ohtsuka. 1995. SEM observations of the
nauplius of Monstrilla hamatapex, new species,
from Japan and an example of upgraded de-
scriptive standards for monstrilloid copepods.—
Journal of Crustacean Biology 15:703-719.

Hartman, O. 1961. A new monstrillid copepod para-
sitic in capitellid polychaetes in southern Cali-
fornia.—Zoologischer Anzeiger 167:325—334.

Huys, R., & G. A. Boxshall. 1991. Copepod evolu-
tion.—Ray Society, London, England 159:1-
468.

Isaac, M. J. 1975. Copepoda. Sub-order: Monstrilloi-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

da.—Fiches d’Identification du Zooplankton
144/145: 10 pp.

Park, T. S. 1967. Two unreported species and one new
species of Monstrilla (Copepoda: Monstrilloida)
from the Strait of Georgia.—Transactions of the
American Microscopical Society 86:144—152.

Raibaut, A., & J. P Trilles. 1993. The sexuality of
parasitic crustaceans.—Advances in Parasitolo-
gy 32:367—444.

Razouls, C. 1996. Diversité et répartition géographique
chez les copépodes pélagiques.2.—Platycopioi-
da, Misophrioida, Mormonilloida, Cyclopoida,
Poecilostomatoida, Siphonostomatoida, Harpac-
ticoida, Monstrilloida——Annales de 1 Institut
Océanographique 72:1—149.

Sars, G. O. 1921. An account of the Crustacea of Nor-
way with short descriptions and figures of all
the species. VIII. Copepoda Monstrilloida &
Notodelphyoida. Bergen Museum, Bergen, 91
pp, pls. IE-XXXVII.

Sekiguchi, H. 1982. Monstrilloid copepods from Ago
Bay, central Japan.—Proceedings of the Japa-
nese Society of Systematic Zoology 22:24—34.

Suarez-Morales, E., & J. B. Escamilla. 1997. An un-
described monstrilloid copepod (Copepoda:
Monstrilloida) from the northern Yucatan Pen-
insula, Mexico.—Bulletin of Marine Science
61:539-547.

, & R. Gasca. 1998. Cymbasoma bowmani sp.

nov., a new monstrilloid (Copepoda: Monstril-

loida) from a Caribbean reef, with notes on spe-

cies variation.—Journal of Marine Systems 15:

433-439.

, & R. Palomares-Garcia. 1995. A new species

of Monstrilla (Copepoda: Monstrilloida) from a

coastal system of the Baja California Peninsula,

Mexico.—Journal of Plankton Research 17:

745-752.

, & R. Vasquez-Yeomans. 1996. On Monstrilla
spinosa Park, 1967 (Copepoda, Monstrilloida)
in the eastern Pacific.—Crustaceana 69:288—
294.

Threlkeld, S. T. 1977. The distribution of five species
of Monstrillidae (Copepoda, Monstrilloida) near
Kodiak Island, Alaska.—Crustaceana 32:225—
228.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(1):199-—209. 1999.

Early development of the deep-sea ampharetid
(Polychaeta: Ampharetidae) Decemunciger apalea Zottoli

Robert Zottoli

Department of Biology, Fitchburg State College, Fitchburg, Massachusetts 01420, U.S.A.

Abstract.—Early development of the opportunistic, deep-sea ampharetid
polychaete Decemunciger apalea Zottoli is described from specimens removed
from wood panels placed on the deep-sea floor by Turner (1973). Larvae with
less than 8 setigers were not found. Larval spatulate notosetae and neuropodial
uncini on segments 3—6 formed during early development are eventually lost
by the 14-setiger stage making segment 7 the first uncinigerous thoracic seg-
ment in juveniles and adults. Early development of this species is generally
similar to that described for other shallow and deep-sea ampharetids.

The deep-sea ampharetid polychaete De-
cemunciger apalea Zottoli inhabits galleries
in wood originally excavated by bivalve
molluscs belonging to the genera Xylopha-
ga and Xyloredo (Family Pholadidae, sub-
family Xylophagainae) (Turner 1973, 1977;
Zottoli 1982). The wood was placed by R.
D. Turner, using the submersible DSRV AI-
vin, at four experimental bottom stations in
the North Atlantic to “test the hypothesis
that wood is an important source of nutri-
ents and contributes to diversity in the
deep-sea”’ (Turner 1973, 1977). This study
is based on a detailed examination of 393
complete juvenile and adult specimens from
wood panel N31 (DOS2) (Fig. 1). Early de-
velopment and further role of this species
in deep-sea ecosystems is discussed.

Materials and Methods

Three experimental islands, each with 12,
one foot spruce cubes, were placed by
Turner (1977), for a period of five years at
the following locations: Deep Ocean Sta-
tion 1 (DOS-1), 39°46’N, 70°41’W, 110
miles south of Woods Hole, Mass., in 1830
m; Deep Ocean Station 2 (DOS-2),
38°18.4’N, 69°35.6’W, 190 miles southeast
of Woods Hole, Mass., in 3506 m; Tongue

of the Ocean, Bahama Islands (TOTO Tow-
er 3), 24°53.2’N, 77°40.2'W, in 2066 m.

Each experimental island is surrounded
by wood panels (24” X 5” X 1”) which are
removed and replaced each time the islands
are visited. Each panel is enclosed in a
mesh bag when it is retrieved, to prevent
loss of specimens. The mesh bags and their
contents are then transferred to retrieval
boxes carried on the DSRV Alvin basket.
The contents of the bags may be preserved
at the time the boxes are closed for return
to the surface by puncturing plastic bags
containing glutaraldehyde, previously
placed in the retrieval boxes. In certain cas-
es, the panels were preserved immediately
upon reaching the surface. Wood panel N31
(DOS 2) was submerged by Alvin on dive
601, 5 Sep 1975, and removed on dive 777,
3 Aug 1977 harboring 393 whole amphar-
etids. Refer to Turner (1973, 1977) for a
more detailed description of the methods
used to place and retrieve the panels from
experimental islands.

Systematics

Ampharetid polychaetes are for the most
part widest anteriorly, tapering gradually to-
wards the posterior end. The prostomium, a
pre-segmental structure, is situated imme-

200

2.50-5.95 (Adults)

1.30-2.45 (Juveniles)

Length in mm

0-1.25 (Larvae)

0 20 40 60

|) bal
N31 (DOS2).

diately in front of the peristomium. Seg-
ments 1 and 2 that are fused together follow
behind the peristomium. Refer to Zottoli
(1974) for additional information on the
formation of these structures. The segmen-
tal numbering system used in this paper is
that of Malmgren (1865-1866) and Fauvel
(1927) who recognized two segments in
front of the paleal segment (segment 3).
The thorax begins on segment 4. Segment
3 in some species has one dorsolateral bun-
dle of paleal setae, or notopod with a bun-
dle of winged capillary setae, on each side.
Segment 3 is achaetous in adult Decemun-
ciger apalea; however, small notopods with
larval, hooded, flared (spatulate) notosetae
or winged capillary setae are present in ear-
ly stages. Adult worms have 13 thoracic se-
tigers, the last 10 of which are unciniger-
ous, and 14 abdominal uncinigerous seg-
ments (Zottoli 1982). In addition, 4 pairs of
smooth branchiae are present on the dorsal
surface of segments 3—5. Refer to Zottoli
(1982) for a more complete description of
adult characteristics.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

80 100 120 140 160 180 200
Number of individuals

Frequency histogram for length from a collection of 393 Decemunciger apalea from wood panel

Larval and juvenile stages of D. apalea
in this study are named according to the
total number of segments bearing spatulate
and/or capillary notosetae (setigers). Post-
setigerous segments are those that follow
these setigers and bear uncinigerous pin-
nules (neuropodia with uncini). For adults
the word setiger refers to any segment with
setae of either the capillary or uncinate
type. The number of thoracic segments with
both capillary notosetae and neuropodial
uncini is a diagnostic feature of juveniles
and adults but not larvae since larvae gain
and lose setae during development, and the
thorax is not fully formed.

Early Development

The eight female D. apalea with mature
eggs in their body cavities ranged from 2.5
to 5.6 mm in length and from 0.5 to 0.7
mm in maximum width, whereas the 13
males with mature sperm ranged from 2.6
to 5.95 mm in length and from 0.42 to 0.7
mm in maximum width. Therefore, speci-

VOLUME 112, NUMBER 1

mens with the full complement of adult se-
tigers and ranging from 1.3 to 2.45 mm in
length are considered juveniles. Worms less
than 1.3 mm in length and lacking the full
complement of adult setigers are considered
larvae. On this basis, 89 adults, 126 juve-
niles and 178 larvae were identified from
the 393 whole specimens removed from
Panel N31 (DOS 2) (Fig. 1). Sexually ma-
ture specimens of this species previously
studied (Zottoli 1982) ranged from 3.6 to
6.3 mm in length and from 0.54 to 0.9 mm
in maximum width.

&-Setiger Stage.—The 8-setiger stage
(Fig. 2) is named for its eight segments
bearing notosetae. Larvae with less than 8
setigers were not found. In the 8-setiger
stage (Fig. 2), larval ciliary bands were not
evident; a short, medial, ventrally ciliated
tentacle protrudes from the mouth. A mus-
cular ventral bulb is visible internally, just
below the pharynx and above the lower lip.
Pharynx, esophagus, stomach and intestine
are visible through the transparent body
wall. The digestive tract is filled with par-
ticulate matter as well as occasional Xylo-
phaga larvae, suggesting that they feed in
the manner described for Hobsonia florida
by Zottoli (1974): “larvae begin feeding on
microscopic plant and animal material at
about the two-setiger stage by forcing ma-
terial from the mud surface into the diges-
tive tract through the action of the ventrally
located buccal mass and by cilia on the up-
per lip.”’

Prostomial and pygidial pigment spots
are lacking. Fused segments | and 2 are
situated immediately behind the prostomi-
um. Segments 3—6 (setigers 1—4) bear 2
small dorsolateral notopods each with 1
spatulate (hooded, flared) seta (about 11 wm
long, 6 ~m maximum width) (Fig. 3A) and
1—2 smooth, winged, bilimbate capillary se-
tae (about 25 wm long, 2 wm wide basally)
(Fig. 3A). The number of smooth, winged,
bilimbate capillary setae per notopod in-
creases as the larva grows from the 8- to
the 14-setiger stage (Table 1).

One uncinus (about 6 wm long) is em-

201

- 4 .!
- ° a
Mee
- A Q; ee
F ie ed as
ee . ia YH
: oe ‘s » oD
: fis Meg 7, ab
7 r A ps
Je | at Va
e

%,

4
at
a NG: -
es VY a Sante

3s

28° yi ¥ Fe
s

Fig. 2. Decemunciger apalea 8-setiger larva. Lat-
eral view of entire worm, 0.6 mm long.

bedded in the epidermis, below the noto-
pod, on each side of segments 3—6 (setigers
1—4). Each of the first 4 pairs of uncini have
about 12 teeth above a basal prow (Fig.

202 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

C

Fig. 3. Decemunciger apalea 8-setiger larva. A. Single spatulate seta and 2 smooth, winged, bilimbate
capillary setae from the left notopod of the second setiger; B. Uncinus, frontal view, from the same setiger as
above; C. Uncinus, frontal view, from the left uncinigerous pinnule of the 5th setiger.

VOLUME 112, NUMBER 1 203

Table 1.—Number of smooth, winged, bilimbate capillary setae per notopodium on “‘setigerous’’ segments
of the 8- to 14-setiger stages of Decemunciger apalea. Note the loss of 1 setiger between the 14b- and 13a-
setiger stages.

Setiger stage

Segment Setiger 8 11 12 14a 14b 13a 13b 13c 13d
3 1 1 1 2 1 1 0) 0) 0) 0
4 2 2 3 3 2 5 3 3 6 +
5 3 2 3 5 3 3 3 4 6 =
6 = 2 4 5 3 5 3, 6 8 6
7 3) 3 3 3 4 6 6 | 8 8
8 6 2 2 3 3 6 6 6 8 8
9 7 2 2 3 3 5) 6 6 9 8

10 8 1 2 2 3 4 6 6 2 2
11 9 1 2 4 4 5 6 8 9
12 10 1 2 3 4 5 5 8 9
13 11 1 2 3 4 + 5 8 8
14 12 2 5 4 + 5 8 6
15 13 5 3 + 5 a 6
16 14 2 2 3 3 q 6

8 = 0.6 X 0.19 mm (1 pair br.), 11 = 1.0 X 0.19 mm (1 pair br.), 12 = 0.9 X 0.18 mm (2 pair br.), 14a =
1.4 X 0.28 mm (3 pair br.), 14b = 1.56 X 0.30 mm (3 pair br.), 13a = 1.75 X 0.30 mm (4 pair br.), 13b =
2.10 X 0.75 mm (4 pair br.), 13c = 3.50 X 0.50 mm ( pair br.), 13d = 5.00 X 0.80 mm (4 pair br.), 14b =
1.56 X 0.30 mm (3 pair br.), br = branchiae.

3B); they disappear in later stages making Larvae at this stage were found in mem-
segment 7 (setiger 5) the first uncinigerous branous tubes covered by fine particulate
thoracic segment in juveniles and adults. matter. The membranous lining is most
Segments 7—10 (setigers 5—8) bear 2 dorso- likely mucus secreted by the thick, glan-
lateral notopods each with one or two’ dular pads on the ventral surface of most
smooth, winged, bilimbate capillary noto- thoracic setigerous segments.
setae (about 25 wm long and 2 pm wide 11-Setiger Stage.—The 11-setiger stage
basally) (Fig. 2; Table 1). Uncini (about 8 (Fig. 4), named for its eleven segments
zm long) from segments 7—15 (setigers 5— bearing notosetae, is similar to the 8-setiger
8 and postsetigers 1-5), are borne on ven-_ stage except for an increase in number of
trolateral extensions of the body (uncini- segments and setae (Table 1), an increase in
gerous pinnules), one pair per segment (Fig. length of the median tentacle, addition of 2
2). Uncini have about 14 teeth above a bas- lateral tentacles, loss of larval spatulate se-
al prow (Fig. 3C). The numbers of uncini_ tae on setiger 4, and loss of larval uncini
per uncinigerous pinnule on all setigers,ex- on setigers 1 and 2 (Table 2).
cept those of segments 3—6 (setigers 1—4) 12-Setiger Stage.—By the 12-setiger
gradually increase from the 8- to the 14-_ stage, named for its twelve segments bear-
setiger stage (Table 2). ing notosetae, the larval uncini on setigers
There are 5 post-setigerous segments 3 and 4 (Table 2), and the larval spatulate
(segments 11—15) bearing uncinigerous pin-__ setae on setigers 1—3 disappear. There is an
nules but lacking notopodia and notosetae. increase in the number of smooth, winged,
The pygidium lies immediately behind seg- bilimbate capillary notosetae per notopo-
ment 15. One pair of branchiae is present dium (Table 1) and in the total number of
on the dorsoposterior edge of segment 3. ‘“‘setigerous’’ and ‘“‘post-setigerous”’ seg-
There is a narrow mid-dorsal gap between ments (Table 2). There is an increase in the
the branchium of each side. length of the median tentacle, and a second

204

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Number of uncini per uncinigerous pinnule on “‘setigerous’’ and ‘‘post-setigerous’’ segments of
the 8- to 14-setiger stages of Decemunciger apalea. Note the loss of 1 setiger between the 14b- and 13a-setiger

stages. Stage lengths are listed in Table 1.

Segment Setiger 8 11 12
3 ] ] 0) 6)
4 2. 1 6) 1
5 3 1 1 1
6 4 1 2; 1
Ji 3) 3 5 i
8 6 2 3 6
9 If 1 2 4

10 8 i 1 3
hill 9 1 1 3
| 10 1 1 Z
i) 11 1 1 1
14 12 1 1 1
i) 13 1 1 1
16 14 1 1
**Post setiger’’ (Abdomen)
17 US) 1
18 16
19 AG,
20 18
21 19
22 20
23 21
24 2D
25 23
26 24
Dal 25
28 26
29 ed |

pair of branchiae appears on the dorsolat-
eral, posterior surface between segments 4
and 5. Later, a third pair of branchiae arises
from the dorso-lateral surface of segment 5.

14-Setiger Stage.—By the 14-setiger
stage, all larval uncini on segments 3—6
have disappeared (Table 2, 14a, b). There is
a general increase in the number of noto-
setae (Table 1) and uncini (Table 2). A
fourth pair of branchiae arises from the an-
terior, dorsolateral surface of segment 6.
Notosetae are eventually lost on segment 3,
thus creating the 13-setiger stage described
below (Table 1).

13-Setiger Stage.—The early 13-setiger
stage (Fig. 5) (Tables 1—2, 13a) has the full
complement of adult thoracic setigerous

ms
>

=
NW fBPOADNDOWwWOUOrK LOCO OC

— eS RS RS KS —S LPO

Setiger stage

14b 13a

O 0) 0) O 0)
0) 0) 0) 0 0)
O 0) O 0) 0
0 0) 0) 0 0)
11 15 3) Js) 30
Tet 14 24 29 28
9 11 18 2M 26
8 13 18 25) 21
6 10 16 23 20
S) 8 KS 23 19
5 8 14 21 18
3 7 14 18 13
3 6 14 17 10

2 » 14 17
2 3 5 16 10
2D 3 5) 10 10
2 3 5 10 9
2 3 5 8 9
1 8 =) J 9
1 Z 5) 7 6
1 DZ 5 7 7
1 1 5) i) 5
1 5 5 4
1 4 5 4
+ 4
yo 3

3

segments of which 10 are uncinigerous.
Uncinigerous thoracic segments begin on
segment 7. Changes that occur between
this stage and adult worms (Tables 1-2,
13c, d) include an increase in number of
capillary notosetae per notopod, an in-
crease in number of uncini per unciniger-
ous pinnule, an increase in number of teeth
per uncinus to about 20, and an increase in
number of uncinigerous abdominal seg-
ments to 12—14.

Uncini are first formed in the dorsal por-
tion of each uncinigerous pinnule. They
range in length from 6—8 wm in an 8-se-
tiger stage to 6-13 pm in a 12-setiger stage
and from 12-18 wm in a 13-setiger stage.
As new, longer uncini are created, older,

VOLUME 112, NUMBER 1

205

Fig. 4. Decemunciger apalea 11-setiger larva. Lateral view of entire worm, 1.0 mm long.

and shorter ones are pushed ventrally and
eventually re-absorbed. This process is
similar to that in A. galapagensis (Zottoli
1983) and E. nebulosa (Bhaud & Grémaré

1988). Few of the originally formed uncini
remain in late juvenile and adult stages,
suggesting an overall rapid growth rate for
D. apalea.

206 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Decemunciger apalea 13-setiger juvenile. Lateral view of entire worm, 1.75 mm long.

VOLUME 112, NUMBER 1

Discussion

Larval ciliated bands, although not found
in D. apalea, disappear at about the 6-se-
tiger stage in H. florida (Hartman) (Zottoli
1974), the 7-setiger stage in Alkmaria rom-
ijni Horst (Cazaux 1982), and were not
found by Zottoli (1983) in later stages of
Amphisamytha galapagensis.

Spatulate setae similar to those described
in D. apalea are found in notopods of the
first 3 setigers of larval A. galapagensis
Zottoli (Zottoli 1983), H. florida (Hartman)
(Zottoli 1974), and Schistocomus sovjeticus
Annekova (Okuda 1947) and in the noto-
pods of the first 4 setigers of larval Am-
pharete acutifrons (Grube) (Clavier 1984)
and Melinna palmata (Grehan et al. 1991).
What appear to be hooded, flared notosetae
can be observed on Nyholm’s (1950) pho-
tograph of a 3-setiger Melinna cristata Sars.
Curiously, none were reported by Cazaux
(1982) for A. romijni. Russell (1987) de-
scribed hooded, flared setae in notopodia of
segments 3—6 from the paedomorphic am-
pharetid Paedampharete acutiseries Rus-
sell. He suggested that “‘the spatulate setae
of H. florida and A. galapagensis may rep-
resent a type of “‘spatulate’’ setae distinctly
different from that of S. sovjeticus, A. acu-
tifrons and P. acutiseries.”’ A detailed ex-
amination of spatulate setae from larval A.
galapagensis, D. apalea, and H. florida
shows a similar 3-dimensional form to that
described by Russell (1987). Notosetae,
which are generally similar in appearance
to ampharetid spatulate setae, have been de-
scribed, respectively, by the bracketed au-
thors on the first 6, 8 and 11 setigers of
Eupolymnia nebulosa (Bhaud 1988 and
Bhaud & Gremare 1988), Thelepus setosus
(Duchéne 1983), and Loima medusa (Wil-
son 1928). Hooded, flared, notosetae even-
tually disappear, with the possible exception
of P. acutiseries, in all of the species dis-
cussed above.

The initial formation of spatulate setae,
capillary notosetae and uncini and their
subsequent loss on anterior segments here,

207

are generally similar to that described for
the ampharetids A. galapagensis (Zottoli
1983), M. palmata (Grehan et al. 1991), H.
florida (Zottoli 1974), and the terebellids E.
nebulosa (Bhaud & Gremare 1988), Loima
conchilega (Kessler 1963), L. medusa (Wil-
son 1928), and Nicolea zostericola (Eckel-
barger 1974).

Larval and adult uncini are similar in D.
apalea, in contrast to A. galapagensis (Zot-
toli 1983) and H. florida (Zottoli 1974)
where multi-toothed larval uncini are re-
placed in most segments by uncini with a
single row of teeth.

Branchial formation in D. apalea is sim-
ilar to that of the ampharetids A. galapa-
gensis (Zottoli 1983), A. romjini (Cazaux
1982), H. florida (Zottoli 1974), and pos-
sibly P. acutiseries (Russell 1987).

Tubes, similar to those of D. apalea,
were formed by 3-setiger, 1.75—2 day old
H. florida larvae (Zottoli 1974), by 4-setig-
er M. palmata l\arvae (Grehan et al. 1991)
and by 8-setiger, 20 day old Alkmaria rom-
ijni larvae (Cazaux 1982), shortly after they
had abandoned the interior of the maternal
tube. Larvae of the terebellids, E. nebulosa
(Bhaud & Gremare 1988), L. conchilega
(Kessler 1963), N. zostericola (Eckelbarger
1974), and 7. setosus (Duchéne 1983),
formed similar tubes, respectively, at the 6-,
2-, 14- and 4-setiger stages. Tube formation
at such an early stage suggests that D. apa-
lea larvae are benthic, remaining in the
Same general area as their parents. This
does not preclude the possibility of larval
transport from place to place by bottom cur-
rents. Once larvae form their own tube,
they most likely remain permanently af-
fixed, reaching new feeding areas by tube
elongation as described by Fauchald & Ju-
mars (1979). Ecklebarger (1974) reported
that juvenile and adult N. zostericola,
forced from their tubes, moved by repeat-
edly folding the abdomen upon itself and
then straightening out. These actions lifted
worms off the bottom and into the water
column, transporting them short distances.
The author observed similar movements in

208

juvenile and adult H. florida forced from
their tubes. Because of the general similar-
ity in body shape and structure of amphar-
etids and terebellids, it is assumed that
most, including D. apalea, could move
short distances in the manner described
above if displaced from their tubes.

Panel N31 (DOS 2) was submerged for
about 2 years from 5 Sep 1975 to 3 Aug
1977. There are more larvae and juveniles
than adults (Fig. 1). Assuming that larvae
and juveniles of D. apalea remain in the
area where they were bred, Fig. 1 most like-
ly reflects the distribution of the species as
a whole. This assumption is supported by
the fact that worms live in attached tubes
in all but the earliest phases of their life and
that their body shape is not conducive to
sustained active locomotion in a pelagic en-
vironment. The presence of numerous lar-
vae and adults with gametes in their body
cavities suggests that breeding took place
Shortly before the time of retrieval. The
small number of large adults suggests a life
span of one year. This is also supported by
the presence of gravid adults, juveniles and
larvae in a panel (N68, DOS1) retrieved af-
ter approximately one year. If the life span
of D. apalea were more than one year, one
would expect a proportionately greater
number of adults.

Acknowledgments

Thanks are due to Dr. J. Fred Grassle,
Charlene D. Long, the late Dr. Meredith L.
Jones and Dr. Ruth D. Turner for making
specimens available. The study, conducted
by Turner (1977), was supported by the Of-
fice of Naval Research (ONR Contract No.
14-76-C-1281, NR 104-687 to Harvard
University).

Literature Cited

Bhaud, M. 1988. Change in setal pattern during early
development of Eupolymnia nebulosa (Poly-
chaeta: Terebellidae) grown in simulated natural
conditions.—Journal of the Marine Biological
Association of the United Kingdom 68:677—
687.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

, & A. Grémaré. 1988. Larval development of
the terebellid polychaete Eupolymnia nebulosa
(Montagu) in the Mediterranean Sea.—Zoolo-
gica Scripta 17(4):347-356.

Cazaux, C. 1982. Développement larvaire de
l’Ampharetidae Lagunaire Alkmaria romijni
Horst 1919.—Cahiers de Biologic Marine 23:
143-157.

Clavier, J. 1984. Description du cycle biologique d’
Ampharete acutifrons (Grube, 1860) (Annélide
Polychéte)—-Comptes Rendus Academie des
Sciences, Paris, Série III 299(3):59-62.

Duchéne, J-C. 1983. Développement larvaire et fixa-
tion chez Thelepus setosus (Annélide Polyché-
te) a Kerguelen, Province Subantarctique.—Vie
et Millieu 33(2):65-—77.

Eckelbarger, K. J. 1974. Population biology and larval
development of the terebellid polychaete Nico-
lea zostericola.—Marine Biology 27:101—113.

Fauchald, K., & P. A. Jumars. 1979. The diet of
worms: A study of polychaete feeding guilds.—
Oceanography and Marine Biology Annual Re-
view 17:193—284.

Fauvel, P. 1927. Polychétes Sédentaires——Faune de
France 16:1—494.

Grehan, A., C. Retiére, & B. Keegan. 1991. Larval
development in the ampharetid Melinna pal-
mata Grube (Polychaeta).—Ophelia Supple-
ment 5:321—332.

Kessler, Von M. 1963. Die entwicklung von Lanice
conchilega (Pallas) mit besonderer beriicksich-
tigung der lebensweise.—Helgolander Wissen-
schaftliche Meersuntersuchungen 8(4):425—
476.

Malmgren, A. J. 1865-1866. Nordiska Hafs-Annula-
ter.—Ofversigt Svenska Vetenskaps Akade-
miens Forhandlingar 22:181—192; 355—410.

Nyholm K-G. 1950. Contributions to the life history
of the ampharetid Melinna cristata.—Zoologis-
ka Bidrag fran Uppsala 29:79-91.

Okuda, S. 1947. On an ampharetid worm, Schistoco-
mus sovjeticus Annekova, with some notes on
it’s larval development.—Journal of the Faculty
of Science Hokkaido Imperial University, Series
6 9:321—329.

Russell, D. E. 1987. Paedampharete acutiseries, a new
genus and species of Ampharetidae (Polychae-
ta) from the North Atlantic Hebble area, exhib-
iting progenesis and broad intraspecific varia-
tion.—Bulletin of the Biological Society of
Washington 7:140—151.

Turner, R. D. 1973. Wood-boring bivalves, opportu-
nistic species in the deep sea—Science 180:
1377-1379.

. 1977. Wood, molluscs, and deep-sea food
chains.—Bulletin of the American Malacologi-
cal Union 1977:13-19.

Wilson, D. P. 1928. The post-larval development of

VOLUME 112, NUMBER 1

Loimia medusa Sav.—Journal of the Marine Bi-
ological Association of the United Kingdom 15:
129-147.

Zottoli, R. A. 1974. Reproduction and larval devel-
opment of the ampharetid polychaete Amphic-
teis floridus.—Transactions of the American
Microscopical Society 93:78-89.

. 1982. Two new genera of deep-sea polychaete

worms of the family Ampharetidae and the role

209

of one species in deep-sea ecosystems.—Pro-
ceedings of the Biological Society of Washing-
ton 95:48-—57.

1983. Amphisamytha galapagensis, a new
species of ampharetid polychaete from the vi-
cinity of abyssal hydrothermal vents in the Gal-
apagos Rift, and the role of this species in Rift
ecosystems.—Proceedings of the Biological So-
ciety of Washington 96:379-391.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(1):210-—215. 1999.

A new species of Spiochaetopterus (Chaetopteridae: Polychaeta) from
a cold-seep site off Hatsushima in Sagami Bay, central Japan

Eijiroh Nishi, Tomoyuki Miura, and Michel Bhaud

(EN) Natural History Museum and Institute, Chiba, Aoba 955-2, Chuo, Chiba 260-8682, Japan;
(TM) United Graduate School of Agricultural Science, Kagoshima University, 1-21-24, Korimoto,
Kagoshima, 890 Japan; (MB) Observatoire Océanologique de Banyuls, Université P. & M. Curie,

(Paris VI) Laboratoire Arago, B.P. 44, F-66651 Banyuls-sur-mer, France

Abstract.—Spiochaetopterus sagamiensis, a new species (Polychaeta: Chae-
topteridae), is described from two specimens collected from a cold-seep site
off Hatsushima in Sagami Bay, central Pacific side of Japan. In this species,
the large chaetae on chaetiger 4 (A4) are distinctive in having a triangular
profile of the head, a nearly straight ventral edge of the inflated distal part, and
two dorsolateral grooves on the shaft. The species has trilobed notopodia on
the first segment of the median region (B1), bilobed ones on the second (B2),
and entire ones on the third (B3) and on all succeeding segments. Uncinal
plates of neuropodium B2 have more than 30 teeth; the upper toothed edge is
smoothly curved. Anterior eyespots are absent. Spiochaetopterus sagamiensis
occurs in deep water (800—1100 m depth) and is the first member of this family
to be recorded in a deep-sea chemosynthetic community.

Chaetopterid polychaetes commonly oc-
cur on mud and sandy mud bottoms from
the intertidal to shallow shelf waters. In
Japanese waters, two species of Spiochae-
topterus are known: S. okudai Gitay, 1969
and S. costarum (Claparéde, 1870) (Okuda
1935, Gitay 1969, Nishi & Arai 1996).
Chaetopterids are poorly known from deep-
sea bottoms: only one species, the type spe-
cies S. typicus Sars, 1856, has been record-
ed from shallow water to a depth of 2700
meters off W. Greenland (Kirkegaard
1960), at depths of 20 to 1865 meters in
Arctic Seas and the Atlantic Ocean (Fauvel
1914), and at depths of 35 to 2030 meters
in the North Pacific and associated water-
bodies (Uschakov 1955). Levin et al.
(1991) listed unidentified chaetopterid poly-
chaetes present in the fauna of seamounts
from the eastern Pacific Ocean at depths of
1058 to 3353 meters. Other records from
deep-sea bottoms are worthy of re-exami-
nation (e.g., Levenstein 1961; Hartman

1971; Hartman & Fauchald 1971). This pa-
per describes a new species of Spiochae-
topterus found in a deep sea cold—seep off
Hatsushima in Sagami Bay (Miura 1988).
It is the first chaetopterid polychaete found
in a chemosynthetic community (cf. Des-
bruyéres & Segonzac 1997) and the third
species in the genus described from Japan.

Materials and Methods

During dive 115 of the Deep-sea Re-
search Vehicle (DSRV) Shinkai 2000 (ob-
server: K. Egawa) in Sagami Bay, two
chaetopterid specimens were collected with
three Calyptogena shells at depths of about
800-1100 m (Sugiura & Egawa 1985). The
specimens were fixed in 10% sea-water for-
malin on the mother ship and later trans-
ferred to 70% ethanol. The specimens were
sent to the second author for further study
of the faunal structure of the Hatsushima
cold-seep site (Miura 1988).

VOLUME 112, NUMBER 1

Parts of the specimens, such as segment
7 of the anterior region and large (cutting)
chaetae were removed dehydrated through
an alcohol series, air-dried, and observed
with scanning electron microscopy (Hitachi
S-800). Type series were deposited in the
Natural History Museum and Institute, Chi-
ba (CBM-ZW). For the description, the ter-
minology of Crossland (1904) and Bhaud
et al. (1994) is followed. In these papers,
three body regions are defined as region A
(anterior), region B (middle) and region C
(posterior); in each region, segments are
numbered from anterior to posterior: for in-
stance, the fourth segment of the anterior
region is termed segment A4 or simply A4.

Family Chaetopteridae Malmgren, 1867
Genus Spiochaetopterus Sars, 1856
Spiochaetopterus sagamiensis, new species
Figs. 1—2

Spiochaetopterus sp.—Miuura, 1988 (dive
numbers 177 and 115 inadvertently
switched in table 2).

Material examined.—Off Hatsushima,
Sagami Bay, DSRV Shinkai 2000 Dive
115, 5 Jun 1984, 35°01'N, 139°12’E, 800—
1100 m, collected with Calyptogena shells,
holotype (CBM-ZW-701), incomplete, with
a fragment of tube and paratype (CBM-ZW-
702).

Diagnosis.—Spiochaetopterus of small
size, eyes absent, with one pair of long
palps. Large (cutting) chaetae of segment
A4 with inflated head. Distal end of A4
chaetae triangular ventrally; shaft without
ventral groove, with two dorso-lateral
grooves. Segment A7 with brownish ventral
gland; A8 and A9 with whitish ventral
glands observed in alcohol preserved spec-
imens. Notopodia trilobed in segment B1,
bilobed in B2, entire in B3 and following
segments. Neuropodia entire in B1, bilobed
in all other middle segments. Uncini with
over 30, maximum 40 teeth on uncinal
plates. Tube unbranched, smooth, not an-
nulated, with serrated opening.

Description.—Both holotype and _ para-

yA Is |

type incomplete, lacking posterior region
(region C). Holotype 18 mm long excluding
palps, about 1 mm wide. Paratype 24 mm
long excluding palps, 0.8-1.0 mm wide.
Regions A and B (anterior and middle)
creamy white, dorsal side of region B light
brown in alcohol apart from glandular re-
gions.

Region A narrow, 3.5 mm long for nine
segments in holotype; 9 mm in paratype.
Prostomium ovoid. Peristomium horseshoe-
shaped, plump with prostomium on dorsal
side (Fig. 1B). Eyespots absent. Prostomial
antennae absent (Fig. 1B, C). Paired palps
long, grooved, 10 mm long in holotype (14
mm in paratype), arising from posterior
border of peristomium, near posterolateral
border of prostomium (Fig. 1A, B, C, E).
Dorsal groove ciliated, extending from base
of palpi in Al to A9 (Fig. 1B, E). Ventrum
of region A with a long slender plastron
(ventral glandular area) (Fig. 1A, C, EB),
with longitudinal white stripes, separated
into five portions (Fig. 1C); portion I, lon-
ger than others, overlaying from peristomi-
um to segment A2, white in colour, portion
II from A3 to A4, narrower at A4, cream-
colored; portion HI from A5 through ante-
rior half of A6, dark brown; portion IV
from posterior half of A6 through A7,
white; portion V from A8 to AQ, white.
Segments A1l—A3 short, parapodia with a
single row of 25—40 lanceolate chaetae; A4
elongate, with two large (cutting) chaetae,
and more than 20 lanceolate chaetae; A5—
A9 longer and wider than anterior three
segments, with single row of 35—40 lance-
olate chaetae (Fig. 2E—H).

Large (cutting) A4 chaetae obliquely tri-
angular, occurring singly on both side (Fig.
2A-F); overhang of ventral edge of head
distinct but weak. Distal part 300 pm long,
230 wm wide, with blunt distal tip and near-
ly horizontal ventral edge 200 wm wide;
shaft 1300 pm long, 10-15 wm wide in
middle portion, asymmetrical, nearly oval
in cross section, without ventral groove;
neck between head and shaft as wide as
shaft (Fig. 2A, B, E).

212 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Z. VW

a6 a5 a4 a3 a2 al

elie Museu

Fig. 1. Spiochaetopterus sagamiensis new species, drawn from holotype (A—E, J—M) and paratype (F—H).
For all drawings (A to M) anterior part of the body is on the right. A. Whole body, lateral view. B. Anterior
portion enlarged. C. Anterior portion including A, segment B1 and anterior B2, ventral view. D. Same, lateral
view. E. Regions A and B, dorso-lateral view. E A8 to B2, lateral view. G. B3 neuropodium, lateral view. H.
A6 to B8 showing rami of neuropodia, ventrolateral view. J. Segments A9 and B1, dorsal view. K. Segment
B2, dorsal view. L. Segment B3, dorsal view. M. Segment B4, dorsal view. Abbreviations.—I to V, indication

VOLUME 112, NUMBER 1

Fig. 2. Spiochaetopterus sagamiensis new species. Scanning electron micrographs of chaetae and tube of
paratype. A—D. Distal part of A4 large (cutting) chaeta, ventral view (A, B) and lateral view (C, D); scales equal
200 pm (A) and 50 pm (B-D). E & E Distal part of A4 large (cutting) chaeta, ventrolateral (E) and anterior
dorsolateral view (F); unlabeled arrows point to dorsolateral groove on shaft; scale equal 100 wm (E) and 50
wm (F). G & H. Chaetae from region A, close-up view of thoracic setae; scales equal 75 wm (G) and 50 um
(H). I. Uncini of B2 neuropodium; scale equals 10 wm. J. Part of tube, lower arrow shows undulation, upper
one serration; scale equals 750 wm.

<_

of ventral parts of region A, i.e., separated portion of ventral plastron and posterior peristomium; al to a9,
segments of region A; b1, first segment of region B; b2, b3, second or third segment of region B; n, notopodium;
ne, neuropodium; p, palp; pe, peristomium; po, prostomium. All scales equal 0.5 mm.

214

Head of A4 chaetae inflated, obliquely
truncate; upper tip of head tilted toward
longitudinal axis of body; consequently ex-
ternal lateral oblique edge longer than in-
ternal edge. Horizontal edge smoothly cir-
cular without ventral sinus. Dorsolateral
grooves on shaft (Fig. 2E, F). Ventral edge
of oblique plane perpendicular to axis of
shaft. Collar decreasing regularly in diam-
eter.

Remaining anterior segments (A5—A9)
large, inflated, shorter and wider ventrally;
parapodia with 35—40 lanceolate chaetae in
single rows; lanceolate chaetae asymmetri-
cal, longitudinally folded, slightly brown-
ish, with smooth surface and serrated edges
(Fig..2G; Fh):

Region B longer than anterior region,
with eight elongate glandular segments
(seven in paratype); paddle and cupule ab-
sent. Bl shorter than following segments
(Fig. 1E). Middle parapodia biramous,
along posterior margins of segments (Fig.
1E, E, H). Notopodia with one to three
lobes; neuropodia bilobed except in seg-
ment B1, with inflated uncinal plates. No-
topodia of segment B1 foliaceous, trilobed,
medial lobes resulting from a subdivision of
inner lobe; outer lobe cirriform (Fig. 1E, J).
Notopodia of B2 bilobed, with single, entire
inner lobes (Fig. 1E, K); those of segments
B3 and following entire (Fig. 1L, M). Neu-
ropodia of segment B1 entire, with only
lower lobe developed (Fig. 1F); other neu-
ropodia bilobed, with rounded upper lobes
and elongate lower lobes (Fig. 1G, H). On
neuropodia of B3, teeth on uncinal plate di-
rected backward on anterior smaller dorsal
lobe; that on larger ventral lobe directed
forward.

Uncini bluntly triangular, with single row
of 35 to 40 minute teeth (n = 6, average
38.2 = 1.94) (Fig. 21). Segment B4 of par-
atype with more than 40 uncini on outer
lobe and more than 15 on inner one. Outer
serrated edge of uncinal plate smoothly
curved.

Posterior body region (region C) lacking
in types.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Both examined specimens are mature,
with segments of region B distended; the
notopodia appear as distended, inflated
transverse bags.

Tube is fragile, slender, straight, smooth,
and not annulated. Collar folds occur at
subequal intervals along tube length. Tube
wall is thin, consisting of several layers of
secretion, and regularly undulated external-
ly (Fig. 2J).

Etymology.—The species epithet is de-
rived from the type locality, Sagami Bay.

Remarks.—Among the 11 described spe-
cies in Spiochaetopterus, 5 species lack
eyespots: S. sagamiensis, S. typicus Sars, S.
bergensis Gitay, S. okudai Gitay and S.
monroi Gitay. These five species may be
distinguished mainly on the shape of the
ventral edge of the oblique section of A4
setae: straight without protuberances in S.
sagamiensis, circular in S. typicus, with
three protuberances in S. bergensis (Bhaud,
1998). In these three cases the ventral edge
is horizontal, perfectly perpendicular to the
shaft Spiochaetopterus okudai and S. mon-
roi were reexamined. The ventral edge of
A4 chaetae is oblique relative to the lon-
gitudinal axis of the shaft in both species.
S. okudai lacks the sinus and S. monroi has
a marked sinus. Secondarily, Spiochaetop-
terus okudai and S. monroi have bilobed
notopodia in region B (Gitay 1969); how-
ever, most notopodia are entire in S. saga-
miensis. From observations by the third au-
thor of Atlantic-Mediterranean and Pacific
species, smoothly curved serrated edges of
uncinal plates of S. sagamiensis may be re-
garded as a discriminatory character.

In conclusion, four diagnostic character-
istics may be retained for this new species:
eyes are absent; the morphology of A4
chaetae with a triangular profile of the head,
a nearly straight ventral edge of the inflated
distal part, and two dorsolateral, and not
ventral, grooves on the shaft; the notopodia
on B3 have only one lobe; and, outer ser-
rated edges of uncinl plates smoothly
curved.

VOLUME 112, NUMBER 1

Acknowledgments

The first author wishes to thank the staff
of the marine laboratory of Natural History
Museum and Institute, Chiba; a facility yet
to be completed. This study was partly sup-
ported by the grant to young scientists from
the Ministry of Education, Sports and Cul-
ture of Japan (to E. N., No. 09740645). The
third author would like to thank the co-au-
thors for carrying out this work. For him,
it is an element of a larger program to ver-
ify the reality of species with a cosmopol-
itan distribution, to define the role of large-
scale larval dissemination and to identify
the spatial limits of successful recruitment.
His participation was funded by the French
National Programme on Determinism of
Recruitment. We are grateful to Dr. Kristian
Fauchald, Smithsonian Institution, for his
useful suggestions on the manuscript.

Literature Cited

Bhaud, M. 1998. The species of the genus Spiochae-
topterus (Polychaeta, Chaetopteridae) in the At-
lantic-Mediterranean biogeographic area.—Sar-
Sia (in press).

, M. C. Lastra, & M. E. Petersen. 1994. Re-
description of Spiochaetopterus solitarius (Rio-
ja, 1917), with notes on tube structure and com-
ments on the generic status (Polychaeta; Chae-
topteridae).—Ophelia 40:115-133.

Crossland, C. 1904. The polychaeta of the Maldive
Archipelago from the collection made by Stan-
ley Gardiner in 1899.—Proceedings of the Zoo-
logical Society of London 1:270-289.

Desbruyéres, D., & M. Segonzac. 1997. Handbook of
deep-sea hydrothermal vent fauna. Editions
IFREMER, Brest, France, 279 pp.

Fauvel, P. 1914. Annelides Polychetes non pelagiques
provenant des campagnes de |’Hirondelle et de
la Princess-Alice (1885—1910).—Resultats des

2AS

Campagnes Scientifiques du Prince de Monaco
46:1—432.

Gitay, A, 1969. A contribution to the revision of Spi-
ochaetopterus (Chaetopteriidae, Polychaeta).—
Sarsia 37:9—20.

Hartman, O. 1971. Abyssal polychaetous annelids
from the Mozambique basin off Southeast Af-
rica, with a compendium of abyssal polychae-
tous annelids from world-wide areas.—Journal
of Fisheries Research Board of Canada 28:
1407-1428.

, & K. Fauchald. 1971. Deep-water benthic
polychaetes annelids off New England to Ber-
muda and other North Atlantic areas, part 2.—
Allan Hancock Foundations, Monograph Series
6:1—327.

Kirkegaard, J. B. 1960. Polychaeta and Pogonophora
from the deepest part of the Skagerrak.—Vi-
denskabelige Meddelelser fra Dansk naturhis-
torisk Forening Kopenhaven 123:211-—226.

Levenstein, R. Y. 1961. Polychaeta from the Bering
Sea.—Trudy Instalagione Okeanologi Akade-
miia Nauk SSSR 46:147—178. (In Russian)

Levin, L. A., L. D. McCann, & C. L. Thomas. 1991.
The ecology of polychaetes on deep-seamounts
in the Eastern Pacific Ocean.—Ophelia, Supple-
ment 5:467—476.

Miura, T. 1988. Parasitic animals collected in a Calyp-
togena-dominant community developing off
Hatsushima, The Sagami Bay.—JAMSTEC
Deepsea Research 9 1988:239—244. (In Japa-
nese with English abstract)

Nishi, E., & Y. Arai. 1996. Chaetopterid polychaetes
from Okinawa.—Publications of the Seto Ma-
rine Biological Laboratory, Kyoto University
37:51-61.

Okuda, S. 1935. Chaetopteridae from Japanese waters.
Journal of the Faculty of Science, Hokkaido
University, ser. 6, Zoology 4:87—102.

Sugiura, A., & K. Egawa, 1985. Research of bottom
fishing grounds in Sagami Bay.—Technical Re-
ports of Japan Marine Science and Technology
Center 1985:67—72.

Ushakov, P. V. 1955. Polychaeta of the far eastern seas
of the U.S.S.R.—Izdatel’stvo Akademii Nauk
SSSR, Moskva-Leningrad, Translated from
Russian and published by Israel Program for
Scientific Translations, Jerusalem 1965, 430pp.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1I2@1):216-2192 1999"

Two new subtribes, Stokesiinae and Pacourininae, of the Vernonieae
(Asteraceae)

Harold Robinson

Department of Botany, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560, U.S.A.

Abstract.—New subtribes Stokesiinae and Pacourininae are established in
the Vernonieae (Asteraceae) for the Southeastern United States genus Stokesia
and the South American genus Pacourina

Revisions of the Vernonieae by the au-
thor have until recently involved mostly
Western Hemisphere members (Robinson
1996). In the absence of studies of Old
World members of the tribe, among which
relatives might have been discovered, the
monotypic Western Hemisphere genera Sto-
kesia L’Hér. and Pacourina Aubl. were left
unplaced as to subtribe. More recent studies
of Old World Vernonieae by the author
have made it clear that Stokesia and Pa-
courina are most closely related to other
Western Hemisphere genera, but that they
are deserving of separate subtribal status.
The new subtribes are described below.

The genus Stokesia has long been noted
for its mostly liguliform corollas, resem-
bling those of the tribe Lactuceae, Fitchia
Hook.f. of the Heliantheae, and some Mu-
tisieae such as Hyaloseris Griseb. (Espinar
1973). The tribal position has sometimes
been questioned, but placement has usually
been in its correct position in the tribe Ver-
nonieae (Bentham & Hooker 1873, Hoff-
mann 1890-1894). The genus contains a
single species, and has been promoted as a
possible crop plant useful as a source for
epoxy resins (Gunn & White 1974). The
plant is also widely cultivated as an orna-
mental. A drawing has been provided by
Gunn and White (1974), and a color pho-
tograph can be seen in Rickett (1967). The
pollen of the genus has a rather weak per-
forated tectum and a unique lophate pattern
with trisected colpi meeting at the poles

(Figs. 1-4). The genus has a chromosome
number that differs from most other Ver-
nonieae, especially other genera from the
Western Hemisphere. A first count of n =
9 (Jones, 1968) has been corrected by a se-
ries of six subsequent counts of n = 7
(Jones 1974). Almost all other New World
Vernonieae have n = 16 or 17. Most Old
World Vernonieae have n = 9 or 10.

Stokesiinae H.Rob., subtribus nov.

Type: Stokesia LHér., Sertum Angl. 27.
1789.

Plantae herbaceae perennes ad 0.5 m al-
tae, sparce pilosae, pilis longis simplicibus
non septatis. Folia plerumque rosulata al-
terna base anguste petioliformia in nodis
vaginata. Inflorescentiae pauce capitatae
laxe cymosae. Capitula pedunculata; brac-
teae involucri 40—5O in seriebus 3—4, brac-
teae exteriores in appendicibus longe foli-
iformes margine spinosae, bracteae inter-
iores angustiores in apicibus setiferae. Flo-
res 60—70 homogami; corollae azurae vel
albae plerumque late liguliformes in limbis
5-lobatae, corollae centrales ca. 3 actino-
morphae; thecae antherarum base rotunda-
tae, cellulis endothecialibus distincte linea-
tis, lineis in partibus longitudinalibus cet-
erum variabiliter arcuatis; appendices api-
cales antherarum breves glabrae in
parietibus cellularum tenues; basi stylorum
non noduliferi; rami stylorum glandulo-
punctati, papillis aciculiformibus argutis.

VOLUME 112, NUMBER 1

217

Figs. 1-4. Stokesia laevis (Hill) Greene, Knobloch 1426 (US), Mississippi. 1. Polar view, line = 20 wm. 2.
Colpar view, line = 20 um. 3, 4. Detailed views of muri with baculae and perforated tectum. 3. line = 3.8 um.
4. line = 1.76 wm.

Achenia 3-4-angulata plerumque prope bas-
em glandulo-punctata, cellulis subsuperfi-
cialibus porosis fibriformibus, raphidis sub-
nullis minutis breviter oblongis; squamae
pappi 4 aut 5 subulatae perfacile deciduae
8—9 mm longae. Grana pollinis triporata, la-
cunis colpi rhomboideis, muris minute
crenulatis (Figs. 1-4). Numerus chromoso-
matum n = 7.

The single species in the subtribe is Sto-
kesia laevis (Hill) Greene, which is native

to the Southeastern United States in south-
ern South Carolina, Georgia, Alabama and
Mississippi, western Florida and eastern
Louisiana.

Pacourina is a singularly distinctive
emergent aquatic plant of tropical America.
The inflorescence, with heads sessile in a
series of leaf axils, is reminiscent of the
Lepidaploa Group in the subtribe Vernoni-
nae. The sclerified apical anther appendage,
however, is totally foreign to that group and

218

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Figs. 5,6. Pacourina edulis Aubl., Killip and Smith 14576 (US), Colombia. 5. Whole grain, line = 15 pm.
2. Closer view showing lacuna with pore and lack of micropunctations on muri, line = 6.7 wm.

is extreme for even the Piptocarphinae and
Lychnophorinae. Triporate, psilolophate
pollen is known otherwise in the tribe Ver-
nonieae only in the Paleotropical subtribe
Erlangeinae, but the pollen of Pacourina is
larger than pollen in any members of that
group, and the inflorescence and anther ap-
pendages are totally different. Thus, the
combination of characteristics precludes
placement in any presently existing subtribe
of the Vernonieae. Closest relationships of
the new subtribe are not known, but they
are presumed to be Neotropical. The dis-
tinctive nature of the plant may derive to
considerable extent from is aquatic special-
ization.

Pacourininae H.Rob., subtribus nov.

Type: Pacourina Aubl., Hist. Pl. Guiane
228002 1775.

Plantae carnosae aquaticae, folia simpli-
cia alterna valde dentata. Inflorescentiae se-
riate cymosae, bracteis foliiformibus; capit-
ula sessilia axillaria solitaria late campan-
ulata homogama; bracteae involucri ca. 50
latae virides et margine albae; receptacula

epaleacea; flores ca. 50 in capitulo; corollae
purpureae, lobis distaliter valde scleroideis;
thecae anterharum base dentate appendicu-
latae; appendices apicales antherarum gla-
brae valde scleroideae; basi stylorum leniter
latiores, pilis stylorum acicularibus. Ach-
enia 10-costata suberose corticata in sulcis
idioblastifera; setae pappi breves multiser-
iatae deciduae, squamellis persistentibus.
Grana pollinis triporata psilolophata emi-
cropunctata (Figs. 5, 6).

The single species in the subtribe in Pa-
courina edulis Aubl. of Central America
and tropical South America. The species is
well illustrated in Nash and Williams (1976,
fig. 7, p. 461).

Acknowledgments

The SEM photographs were made by Su-
sann Braden of the National Museum of
Natural History SEM Laboratory using a
Hitachi S-570. Prints were prepared by
Sherry Pittam, previously of the Depart-
ment of Botany.

VOLUME 112, NUMBER 1

Literature Cited

Bentham, G., & J. D. Hooker. 1873. Compositae.—
Genera Plantarum. Vol. 2(1):163-—533. Reeve &
Co., London.

Espinar, L. A. 1973. Revision del genero Hyaloseris
(Compositae).—Kurtziana 7:195—211.

Gunn, C. R., & G. A. White. 1974. Stokesia laevis:
taxonomy and economic value.—Economic
Botany 28:130-—135.

Hoffmann, O. 1890-94. Compositae 4(5):87—387. In
A. Engler & K. Prantl, (eds.).—Die Natiirlichen
Pflanzenfamilien. Leipzig.

Jones, S. B. 1968. Chromosome numbers in South-
eastern United States Compositae, I—Bulletin
of the Torrey Botanical Club 95:393-—395.

a9

. 1974. Vernonieae (Compositae) chromosome
numbers.—Bulletin of the Torrey Botanical
Club 101:31—34.

Nash, D. L., & L. O. Williams. 1976. Flora of Gua-
temala.—Fieldiana: Botany 24(12):1—603.

Rickett, H. W. 1967. Wild flowers of the United States,
the southeastern states. 2(2):601, pl. 224. Mc-
Graw-Hill, New York.

Robinson, H. 1996. The status of generic and subtribal
revisions in the Vernonieae. Pp 511-526 in D.
J. N. Hind & H. J. Beentje, eds., Compositae:
Systematics.—Proceedings of the International
Compositae Conference, Kew, 1994. (D. J. N.
Hind, Editor-in-Chief). Vol. 1, Royal Botanic
Gardens, Kew.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
112(1):220-—247. 1999.

Revisions in paleotropical Vernonieae (Asteraceae)

Harold Robinson

Department of Botany, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560-0166, U.S.A.

Abstract.—The paleotropical subtribes Erlangeinae, Centrapalinae and Gym-
nantheminae are described as new. Fourteen paleotropical genera are described,
raised from lower rank, resurrected, enlarged or reduced with 110 new com-
binations. Cabobanthus (2 spp.), Hilliardiella (8 spp.), Orbivestus (4 spp.),
Vernoniastrum (8 spp.), Koyamasia (1 sp.), Brenandendron (3 spp.), Myan-
maria (1 sp.) and Manyonia (1 sp.) are described as new. Oocephala (S. B.
Jones) H. Rob. (2 spp.) and Lampropappus (O. Hoffm.) H. Rob. (3 spp.) are
elevated to generic rank. Four genera are resurrected, the Asian Acilepis D.
Don with 9 of 10 species newly combined, the African Bechium DC. (2 spp.),
Centrapalus Cass. (9 spp.), Linzia Sch. Bip. ex Walp. with 5 of 7 species newly
transferred, and Polydora Fenzl (8 spp.). One species is transferred to make a
total of 27 in Distephanus, 4 species are transferred to total 7 in Cyanthillium,
and 39 species are transferred to total 43 in Gymnanthemum. Lamprachaenium
Benth. is synonymized with Phyllocephalum Blume with 1 species transferred.

The present paper provides a limited re-
organization of the Eastern Hemisphere
Vernonieae for purposes of a projected ge-
neric review of the tribe. The need to sum-
marize the whole tribe presents a special
problem. The half of the tribe in the West-
ern Hemisphere has been rather well delim-
ited into workable and phyletically accept-
able genera (Robinson 1996). This includes
an accurate delimitation of the type genus
Vernonia Schreb. that is primarily found in
eastern North America. The Eastern Hemi-
sphere members of the tribe have thus been
left in a particularly untenable position,
with an excessively paraphyletic core genus
and mostly paraphyletic or polyphyletic
segregates, none of which are congeneric
Vernonia. In fact, none of the existing sub-
tribal names in the Vernonieae are appli-
cable to the large elements that are endemic
to the Eastern Hemisphere.

At the time of the summary of the New
World Vernonieae (Robinson 1966), only a
few Eastern Hemisphere elements have
been revised by the author. The name Bac-

charoides was resurrected for the members
of the Stengelia Group as early as Robinson
et al. (1980), and transfers were made into
Baccharoides and Cyanthillium (Robinson,
1990) mostly to accomodate species adven-
tive in the Western Hemisphere. Distephan-
us was resurrected and partially revised as
a last step in the removal of some of its
species from the tribe Senecioneae (Rob-
inson & Kahn 1986). In the latter study the
opportunity was taken to provide the proper
combination for the type species of Gym-
nanthemum Cass. that had originally been
named as a Eupatorium. Nevertheless, no
further work was planned in the Eastern
Hemisphere members of the Vernonieae as
recently as the time of the publication of
the preliminary Western Hemisphere revi-
sion in the Kew International Compositae
Conference volume (Robinson 1996).

The present study has built upon the ref-
erence works of many other authors. Hum-
bert (1960), Wild (1977, 1978), Wild &
Pope (1978a, 1978b), Jeffrey (1988), which
cover all the genera of the tribe in various

VOLUME 112, NUMBER 1

parts of Africa, have proven most useful in
the present study. The study of Jones
(1981), restricted to Vernonia sensu lato,
and Lisowski (1992) that excludes Vernon-
ia, have been less useful. The works of
Kirkman (1981), Pope (1983), Koyama
(1984), Isawumi (1993, 1995), and Isawumi
et al. (1996) have added important infor-
mation. The present study has used the
summary of secondary metabolite chemis-
try of the Vernonieae by Bohlmann and
Jakupovic (1990) and the various reports of
chromosome numbers by Jones (1974,
1979, 1982). DNA studies are as yet limited
to those mentioned by Keeley (1994, 1995)
and Kim et al. (1996). The final conclusions
of the present study are, nevertheless, ulti-
mately based on examination of specimens
in the United States National Herbarium
(US) and on some specimens kindly sent on
loan by Kew (K), British Museum (BM),
Bruxelles (BR) and Paris (P).

Jones (1977, 1981) noted basic differenc-
es between the Vernonieae in the Eastern
and Western Hemispheres, and this has
been generally accepted by others such as
Jeffrey (1988). These differences have been
reviewed to some extent by Isawumi
(1995). The Hemispheric trends noticed by
Jones (1977) are in the chromosome num-
bers and chemistry. The Western Hemi-
sphere species usually have a chromosome
number of N = 17, and the Eastern Hemi-
sphere species have mostly N = 9 or 10.
The chemicals cited were terpenoids and
flavonoids. Jones (1981) also stated that tri-
porate pollen grains were known only from
the Eastern Hemisphere. Since that time,
some examples of triporate grains have
been found in Latin America, Acilepidopsis
(Robinson 1989), Mesanthophora (Robin-
son 1992a), Pacourina (Robinson 1992b),
but at least the first two of these are con-
sidered to be related to Eastern Hemisphere
members of the tribe. Robinson and Kahn
(1986) and Isawumi et al. (1996) briefly
noted the apparent restriction of glandular
dots on the anthers and their appendages to
New World members of the Vernonieae. In

221

the present study, the raphids of the achene
wall in the Eastern Hemisphere species
have proven to be predominently elongate,
with only a few entities such as Gutenber-
gia having characteristically subquadrate or
short raphids. Most groups of Western
Hemisphere Vernonieae have characteristi-
cally subquadrate raphids. In their chemical
survey, Bohlmann and Jakupovic (1990)
found the distinctive 5-alkyl coumarins
only in Eastern Hemisphere genera and spe-
cies, groups that are all placed in this study
in the new subtribe Erlangeinae.

Inflorescences with scorpioid or well de-
veloped seriate cymes in the Vernonieae
seem almost entirely restricted to the New
World and New World genera like Stru-
chium that have become Pantropical. A no-
table exception, Manyonia, described below
as a new genus from East Africa, is consid-
ered to be a relative of New World genera,
and it is described in the subtribe Vernon-
linae.

Some doubts about the hemispheric dif-
ferences were expressed by Keeley and
Turner (1990). They mention some cases of
Western Hemisphere ‘‘Vernonia’’ with
chromosome numbers of N = 10 or 12, and
they call attention to similarities between
the groups referred to by the present author
as Piptocarphinae in the Americas and the
Gymnantheminae in the Eastern Hemi-
sphere. The latter groups both tend to be
woody, often have deciduous inner involu-
cral bracts and usually have blunt-tipped
sweeping hairs on their styles. Neverthe-
less, the possible relationship is considered
here to be above the generic and subtribal
levels. The doubts of Keeley & Turner
(1990) are justified to the extent that Ver-
nonia cannot be simply divided into two
genera, one for each Hemisphere. Many
segregates are needed for each hemisphere.

There is an important limitation to the
use of pollen structure as a phyletic char-
acteristic in the Vernonieae, a limitation al-
ready seen in both Western and Eastern
Hemisphere members of the tribe (Robin-
son 1996). First, the simple non-lophate or

222

sublophate pollen, Type A, often consid-
ered primitive in the tribe, was not consid-
ered primitive by the present author (Rob-
inson 1996). Second, some pollen varia-
tions are of more importance than others.
Differences between various lophate pat-
terns seem to correlate well with other char-
acters, but all groups seem able to revert
erratically to non-lophate or sublophate
forms, the Type A of Jones (1981). The tru-
ly triporate forms, with irregularly orga-
nized polar lacunae, are evidently not close-
ly related to other lophate types with dis-
tinct colpi. However, they may be very
closely related to colporate forms that are
non-lophate. This does not mean that the
difference between triporate and non-lophate
tricolporate cannot be used as a key char-
acter when the character happens to corre-
late with other features, as in Cyanthillium.

The present study has been restricted by
available time and specimens. The new sub-
tribes are kept to a minimum, although that
minimum seems to fit the Paleotropical ma-
terial rather well. The new genera and taxa
elevated to generic rank are those that seem
inescapable, and no attempt is made here to
dispose of some additional distinctive ele-
ments in the Eastern Hemisphere that do
not fit in the genera presently recognized.
Possibly, some smaller distinctive elements
have been completely missed in the present
survey. The number of species transferred
into the various segregate genera is neces-
sarily incomplete, but proper names are
now available for many species that should
not be retained in Vernonia. It is hoped that
other authors will continue the process of
making transfers. An attempt has been
made to avoid overly broadening the con-
cepts of the segregate genera. Even so,
Gymnanthemum has been interpreted to in-
clude many elements that differ from the
type by corolla lobe, style and achene char-
acters. This has been done with the convic-
tion that the broader concept of that genus
will prevail.

The synonymy given for most of the Af-
rican species follows Jeffrey (1988).

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

New Subtribes of the Eastern Hemisphere
Vernonieae

All presently named subtribes of the Ver-
nonieae are based on primarily Western
Hemisphere groups. None of these subtribal
names apply to the primarily Eastern Hemi-
sphere groups. To rectify this situation, the
following three new subtribes are named.

Erlangeinae H. Rob., subtribus nov. Type:
Erlangea Sch. Bip., Flora 36:34. 1853.
Type: E. plumosa Sch. Bip.

Vernonia sect. Tephrodes DC. Prodr. 5:
24. 1836. Type: Conyza cinerea L.
Bechium DC., 5:70. 1836. Type: B. scap-

iforme DC.

Vernonia sect. Lepidella Oliver & Hiern,
Fl. Trop. Afr. 3:267. 1877. Type: Ver-
nonia petersii Oliver & Hiern.

Vernonia subg. Orbisvestus S. B. Jones,
Rhodora 83:60. 1981. Type: Vernonia
karaguensis Oliver & Hiern.

Vernonia sect. Orbisvestus S. B. Jones,
Rhodora 83:61. 1981. Type: as above.

Vernonia subsect. Orbisvestus S. B.
Jones, Rhodora 83:61. 1981. Type: as
above.

Vernonia subsect. Hilliardianae S. B.
Jones, Rhodora 83:66. 1981. Type:
Webbia oligocephala DC.

Vernonia subsect. Tephrodes (DC.) S. B.
Jones, Rhodora 83:70. 1981.

Vernonia subsect. Lepidella (Oliver &
Hiern) S. B. Jones, Rhodora 83:72.
1981.

Vernonia subsect. Oocephalae S. B.
Jones, Rhodora 83:72. 1981. Type:
Vernonia oocephala Baker

Vernonia subsect. Bechium (DC.) S. B.
Jones, Rhodora 83:73. 1981.

Plantae herbaceae annuae aut perennes
vel frutescentes, pilis saepe symmetriciter
T-formibus. Folia alterna vel opposita vel
ternata pinnatinervata. Receptaculum epa-
leaceum vel raro paleaceum. Flores 3 vel
ca. 100 in capitulo; corollae lavandulae vel
purpureae; thecae antherarum base non vel
tenuiter caudatae; appendices apicales gla-

VOLUME 112, NUMBER 1

brae in parietibus cellularum tenues raro
tenuiter ornatae; base stylorum noduliferi;
rami stylorum aciculiformiter papillosi.
Achenis 4—6 vel 8-10 costata; raphidis
plerumque elongatis raro subquadratis; car-
popodia anguste cylindrica; pappus longe
setiformis vel abbreviatus vel coroniformis
saepe facile deciduus. Grana pollinis non
lophata et tricolporata vel lophata et tripor-
ata in formibus lophatis lacunis polaribus
irregularibter dispositis, tectis micropunc-
tatis vel emicropunctatis. Numerus chro-
mosomatum N = 9, 10, 20.

The name is chosen to conform with the
already established term ‘‘Erlangeoid’’
(Pope 1983). The subtribe is circumscribed
to include all the Vernonieae presently
known with triporate pollen (excluding Pa-
courina Aubl., Robinson 1992b) or 5-alkyl
coumarins (Bohlmann & Jakupovic 1990).
The unquestioned core of the subtribe in-
cludes genera with only 4—5 angled achenes
and herbaceous habits, but, at present, ad-
ditional forms with 10-ribbed achenes and
woody habits are also included. All the in-
cluded genera have acicular sweeping hairs
of the style. The genera included are the
New World Acilepidopsis H. Rob. (1989)
and Mesanthophora H. Rob. (1992a), and
Old World Acilepis D. Don, Ageratinastrum
Mattf., Ambassa Steetz, Bechium DC..,
Bothriocline Oliv. ex Benth., Brachythrix
Wild & G. V. Pope, Cyanthillium Blume,
Decastylocarpus Humbert, Dewildemania
O. Hoffm., Diaphractanthus Humbert, Er-
langea Sch. Bip., Ethulia L. f., Gossweilera
S. Moore, Gutenbergia Sch. Bip. ex Walp.,
Hystrichophora Mattf., Iodocephalus Tho-
rel ex Gagnep., Kinghamia C. Jeffrey, Lam-
prachaenium Benth., Msuata O. Hoffm.,
Phyllocephalum Blume, Omphalopappus
O. Hoffm., Rastrophyllum Wild & G. V.
Pope, and six genera newly named or ele-
vated below.

Centrapalinae H. Rob., subtribus nov.
Type: Centrapalus Cass., Dict. Sci. Nat.
ed. 2, 7:382. 1817. Type: Centrapalus
galamensis Cass.

223

Vernonia subsect. Stengelia Sch. Bip. ex
Walp., Repert. Bot. Syst. 2 [Suppl. 1]:
946. 1843. Type: Vernonia adoensis
Sch. Bip. ex Walp.

Vernonia sect. Stengelia (Sch. Bip. ex
Walp.) Benth. & Hook. f., Gen. Pl. 2:
227 60873.

Vernonia subsect. Centrapalus (Cass.) S.
B. Jones, Rhodora 83:69. 1981.

Vernonia sect. Azureae S. B. Jones, Rho-
dora 83:74. 1981. Type: Vernonia gla-
bra Vatke.

Plantae herbaceae perennes vel raro an-
nuae, pilis simplicibus multiseptatis vel
asymmetriciter T-formibus vel nullis. Folia
alterna pinnatinervata vel longitudinaliter
nervata. Bracteae involucri apice saepe ap-
pendiculatae vel plerumque herbaceae. Flo-
res numerosi in capitulo; corolla purpurea
vel azurea; thecae antherarum base rotun-
datae; appendices apicales glabrae vel raro
papilliferae in parietibus cellularum leniter
incrassatae; base stylorum noduliferi vel
non noduliferi; rami stylorum acute papil-
liferi. Achenis 10-costata, raphidis elongatis
vel interdum subquadratis; pappus pler-
umque setiformis. Grana pollinis tricolpor-
ata lophata vel non lophata micropunctata
vel non micropunctata in formibus lophatis
lacunis polaribus solitariis interdum prae-
sentibus. Numerus chromosomatum N = 9,
10.

The subtribe is typified by Centrapalus
Cass., and includes Adenoon Dalz., Aedesia
O. Hoffm, Baccharoides Moench, Cam-
chaya Gagnep., Lachnorhiza A. Rich., Lin-
zia Sch. Bip. ex Walp., Muschleria S.
Moore, Neurolakis Mattf. and Pleurocar-
paea Benth. The sesquiterpene constituents
include elemanolides (Bohlmann & Jaku-
povic 1990).

The genera of the subtribe are herba-
ceous or weakly shrubby, and the sweeping
hairs of the styles are acicular. The subtribe
includes elements with the distinctive Lin-
zia-type pollen cited by Jeffrey (1988) such
as Linzia and Aedesia and one with polar
lacunae on its pollen grains and a lack of

224

basal stylar nodes like Baccharoides (Isa-
wumi 1993; Isawumi et al. 1996).

Gymnantheminae H. Rob., subtribus nov.
Type: Gymnanthemum Cass., Bull. Soc.
Philom. Paris 1817:10. 1817. Type: Gym-
nanthemum cupulare Cass. [= G. color-
atum (Willd.) H. Rob. & B. Kahn].

Distephanus Cass., Bull. Soc. Philom.
Paris 1807-151. 18t7. Type: Diste-
phanus populifolius (Lam.) Cass.

Vernonia sect. Strobocalyx Blume ex
DC., Prodr. 5:21. 1836. Type: Vernon-
ia arborea Buch.-Ham.

Gongrothamnus Steetz in Peters., Reise
Mossamb. Bot. 336. 1864. Type: G.
divaricatus Steetz in Peters

Vernonia sect. Distephanus (Cass.)
Benth. & Hook. f., Gen. pl. 2:228.
1873.

Vernonia sect. Lampropappus O. Hoffm.,
Bol. Soc. Broter. 13:14. 1896. Type:
Vernonia lampropappa O. Hoffm.

Vernonia subsect. Strobocalyx (Bl. ex
DC.) S. B. Jones, Rhodora 83:64.

1981.

Vernonia subsect. Gongrothamnus
(Steetz) S. B. Jones, Rhodora 83:65.
1931.

Vernonia subsect. Pawekianae S. B.
Jones, Rhodora 83:66. 1981. Type:
Vernonia angulifolia DC.

Vernonia subsect. Urceolatae S. B.
Jones, Rhodora 83:67. 1981. Type:
Vernonia sphaerocalyx O. Hoffm.

Vernonia subsect. Turbinella S. B. Jones,
Rhodora 83:67. 1981. Type: Vernonia
lampropappa O. Hoffm.

Vernonia subsect. Distephanus (Cass.) S.
B. Jones, Rhodora 83:68. 1981.

Vernonia subsect. Glutinosae S. B. Jones,
Rhodora 83:73. 1981. Type: Vernonia
glutinosa DC.

Plantae fruticosae vel arborescentes vel
scandentes, pilus simplicibus et arachnoid-
eis vel L-formibus vel T-formibus. Folia al-
terna pinnatinervata vel trinervata. Bracteae
involucri 20—70 in seriebus 2—7 plerumque
gradatae interiores interdum deciduae; re-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ceptacula epaleacea vel paleacea. Flores 1—
40 (—75) in capitulo; corollae lavandulae
vel roseae vel flavae, lobis plerumque er-
ectis et longe triangularibus rariter elongatis
et arcte revolutis; thecae antherarum base
plerumque valde caudatae; appendices api-
cales induratae et glabrae in parietibus cel-
lularum aliquantum ornate incrassatae; basi
stylorum non noduliferi vel noduliferi in-
terdum abrupte noduliferi; rami stylorum
plerumque obtuse papilliferi. Achenia 5 aut
10-12-costata, raphidis plerumque elongatis
interdum subquadratis vel nullis, carpopo-
dia lata; pappus plerumque setiformis, Gra-
na pollinis tricolporata non lophata vel sub-
lophata vel raro lophata in formibus lopha-
tis lacunis polaribus solitariis nullis; tectis
micropunctatis. Numerus chromosomatum
N = 10, 2N = 30.

The subtribe is typified by the genus
Gymnanthemum Cass., but it also includes
Distephanus Cass., Centauropsis Boj. in
DC., Oliganthes Cass., and three genera
named or elevated below. The sesquiter-
pene lactone constituents include eleman-
olides (Bohlmann & Jakupovic 1990).

The subtribe includes all of the true large
shrub and tree Vernonieae in the Eastern
Hemisphere. The sweeping hairs of the
styles often have rather blunt tips. Inner in-
volucral bracts are persistent or deciduous,
and a few species of Gymnanthemum and
Brenandendron have long coiled corolla
lobes. These characteristics are generally
Shared by the Gymnantheminae and the
American Piptocarphinae, but deciduous
bracts and reflexed corolla lobes are much
more consistently present and blunt sweep-
ing hair much less consistently present, in
the Piptocarphinae. The Gymnantheminae
lack stellate hairs like those common in the
Piptocarphinae.

New Genera and New Combinations of
Eastern Hemisphere Vernonieae
Subtribe Vernoniinae

Manyonia H. Rob., gen. nov. (Vernoni-
inae).

VOLUME 112, NUMBER 1

Type: Vernonia peculiaris Verdc.

Plantae herbaceae perennes ad 1 m altae;
caules brunnescentes striati sparce hispidu-
li, pilis simplicibus multiseptatis uniseriatis,
internodis 4—6 cm longis. Folia alterna, pe-
tiolis 8-10 mm longis; laminae membran-
aceae ovatae 8-11 cm longae 3.2—5.0 cm
latae base anguste cuneatae margine biser-
ratae apice plusminusve acuminatae supra
sparce pilosae subtus dense glandulo-punc-
tatae, nervis secundariis utrinque 5 vel 6 su-
pra et subtus puberulis. Inflorescentiae se-
riate cymosae, ramis ad 14 cm longis, pe-
dunculis plerumque brevibus 2—3 (—15) mm
longis dense puberulis. Capitula campanu-
lata 6 mm alta et lata; bracteae involucri ca.
100; seriebus exterioribus 3—4 patentes
longe subulatae 2.5 mm longae base dila-
tatae 0.1—0.2 mm latae inferne margine
puberulae; bracteae intermediae oblongae 4
mm longae apice longe aristatae; bracteae
interiores anguste oblongae 5 mm longae et
2 mm latae apice acuminatae; receptaculum
glabrum in diametro ca. 1.8 mm. Flores ca.
35 in capitulo; corollae lilacinae ca. 5.5 mm
longae, tubis angustis ca. 2.5 mm longis su-
perne infundibularibus, faucibus ampliatis
ca. 1 mm longis, lobis oblongis ca. 1.5 mm
longis et 0.3 mm latis extus glanduliferis;
thecae antherarum ca. 2 mm longae base
non caudatae; appendices apicales ca. 0.4
mm longae in parietibus cellularum tenues
glabrae; basi stylorum disciformiter nodati;
rami stylorum in papillis aciculiformibus
obsiti. Achenia oblonga 1.5 mm longa 5-
costata inter costam breviter setulifera et
longitudinaliter multiseriate idioblastifera,
costis glabris prominentibus et induratis, ra-
phidis perdense dispositis subquadratis vel
polygonatis; setae pappi ca. 20 barbellatae
ca. 4 mm longae; squamulae exteriores ca.
25 minute ciliatae. Grana pollinis in dia-
metro ca. 35 w sublophata tricolporata.

The genus Manyonia has been consid-
ered by the author as a relative of New
World Vernonieae since he first saw the il-
lustration of the seriate cymes accompany-
ing the original description (Verdcourt

225

1956). This structure, otherwise lacking in
native Old World Vernonieae, is confirmed
in material borrowed from The Royal Bo-
tanic Gardens, Kew. This relationship
seems confirmed by a poorly researched
characteristic, the type of ornamentation of
the endothecial cells of the anther. The cells
in Manyonia have rather strong thickenings,
longitudinal in some cell rows, and in other
rows, arching across the lower end. This is
a pattern seen in many Western Hemisphere
Vernonieae. Eastern Hemisphere Vernon-
ieae have weaker ornate thickenings if any.
Further examination of specimens of Man-
yonia shows a strong resemblance to New
World genera such as Heterocypsela H.
Rob. and Dipterocypsela S. F Blake, es-
pecially in the wall of the achene with ex-
tremely crowded subquadrate or polygonal
raphids. The new genus differs from Het-
erocypsela and Dipterocypsela in lacking
the heteromorphic achenes and the glan-
duliferous anther appendages of its Ameri-
can relatives. The pollen of Manyonia is
sublophate, more like Dipterocypsela and
unlike the lophate form in Heterocypsela.

The generic name is derived from the
vernacular name Manyoni cited on Burtt
5119 (K). The needed combination is as
follows:

Manyonia peculiaris (Verdc.) H. Rob.,
comb. nov. basionym: Vernonia pecu-
liaris Verdc., Kew Bull. 1956:447. 1956.
Tanzania.

New Genera and New Combinations of
Eastern Hemisphere Vernonieae
Subtribe Erlangeinae H. Rob.

Acilepis D. Don, Prodr. Fl. Nepal. 169.
1825. Type: Acilepis squarrosa D. Don.

Lysistemma Steetz in Peters, Reise Mos-
samb. Bot. 340. 1864. Type: Lysistem-
ma dendigulense (DC.) Steetz.

Xipholepis Steetz in Peters, Reise Mos-
samb. Bot. 344. 1864. Type: Xiphole-
pis silhetensis Steetz.

Vernonia sect. Xipholepis (Steetz) Benth.
& Hook.f., Gen. pl. 2:229. 1873.

226

Erect perennial herbs; stems pentangular,
with hairs multiseptate at base and often
with long subfusiform apical cell. Leaves
alternate, obovate to oblong-ovate. Inflores-
cences of single heads, spiciform cymes, or
corymbose cymes with few to many heads.
Involucres funnelform to campanulate; bracts
50—200 in 6—12 series, persistent, apiculate to
subacute; receptacle epaleaceous. Heads with
25—80 florets; corollas lavender, tubes slender
below, funnelform above into throat, throat
half or less as long as anther thecae, lobes
long and narrow, with glandular dots; anther
bases blunt, not tailed; apical anther append-
ages glabrous, with thin-walled cells; style
base with node, style branches with acicular
sweeping hairs. Achenes 8—10 ribbed, setulae
with one cell long, other cell short, raphids
oblong with rhomboid tips; pappus whitish,
both series rather easily deciduous, with
many barbellate inner setae, outer setae short-
er, scarcely broader. Pollen triporate, lophate,
nearly psilate, emicropunctate, with ca. 20 la-
cunae.

The Asiatic Acilepis is distinct in the
rather simple stem hairs, the pedunculate or
separated heads, the triporate pollen and in
such details as the often totally deciduous
pappus and the highly unequal cells of the
setulae of the achene. Jeffrey (1988) rec-
ognized the group for an African species,
Vernonia polysphaera, treated below as the
new genus Cabobanthus. The latter has
basal tubers, sessile clustered heads, a more
persistent pappus and the cells of the setu-
lae of equal length.

The following ten species are recognized
in the genus:

Acilepis aspera (Buch.-Ham.) H.Rob.,
comb. nov. basionym: Vernonia aspera
Buch.-Ham., Trans. Linn. Soc. London
14:219. 1824.

Eupatorium pyramidale D. Don, Prodr.
FL Nepal. 1702/1825;

Vernonia roxburgii Less., Linnaea 6:674.
1831.

Xipholepis aspera (Buch.-Ham.) Steetz in

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Peters, Reise Mossamb. Bot. 345.
1864.

Vernonia pyramidalis (D. Don) Mitra,
Ind. For. 99:100. 1973. China, India,
Myanmar, Nepal, Thailand.

Acilepis clivorum (Hance) H. Rob., comb.
nov. basionym: Vernonia clivorum
Hance: J... Bot., 7.164. (869:

Aster coriaceiformis H. Lév. & Vaniot, Re-
pert. Spec. Nov. Regni Veg. 8:358. 1910.
China.

Acilepis dalzelliana (J. R. Drumm. &
Hutch.) H. Rob., comb. nov. basionym:
Vernonia dalzelliana J. R. Drumm. &
Hutch., Kew Bull. 1909:261. 1909.
India.

Acilepis dendigulensis (DC.) H. Rob.,
comb. nov. basionym: Decaneurum den-
digulense DC. in Wight, Contr. Bot. Ind.
7. 1834, not Vernonia dendigulensis DC.

Lysistemma dendigulense (DC.) Steetz in
Peters, Reise Mossamb. Bot. 341.
1864.

Vernonia indica C. B. Clarke, Comp. Ind.
16. 1876. Western peninsular India.

Acilepis nantcianensis (Pamp.) H. Rob.,
comb. nov. basionym: Vernonia bractea-
ta var. nantcianensis Pamp., Nouv.
Giorn. Bot. Ital., n.s. 18:98. 1911.

Vernonia silhetensis vat. nantcianensis
(Pamp.) Hand.-Mazz., Symb. Sin. 7:
1084. 1936.

Vernonia nantcianensis (Pamp.) Hand.-
Mazz., Noitsibl. Bot. Gart. Mus. Berl.-
Dahl. 13:608. 1937.

China.

Acilepis saligna (DC.) H. Rob., comb. nov.
basionym: Vernonia saligna DC., Prodr.
3: 33.01836.

Vernonia longicaulis DC., Prodr. 5:33.
1836.

Vernonia martinii Vaniot, Bull. Acad. In-
tern. Geogr. Bot. 12:124. 1903.

Vernonia sequinii Vaniot, Bull. Acad. In-
tern. Geogr. Bot. 12:241. 1903.
China, India, Myanmar.

VOLUME 112, NUMBER 1

Acilepis scariosa (DC.) H. Rob., comb.
nov. basionym: Decaneurum scariosum
DC. Prodr-7:264. 1838:

Vernonia scariosa Arn., Nova Acta Phys.
Med. Acad. Caes. Leop. Carol. Nat.
Cur. 18:346. 1836, hom. illeg., not V.
scariosa Poir., 1808.

Gymnanthemum scariosum (DC.) Sch.
Bip. ex Walp., Rep. 2:949. 1843.

Centratherum scariosum C. B. Clarke,
Comp. Ind. 4. 1876.

Vernonia lankana Grierson, Ceylon J.
ser, Biol. Sci. 10:43. 1972.

Sri Lanka.

Acilepis silhetensis (DC.) H. Rob., comb.
nov. basionym: Decaneurum silhetense
DC Prodr. 5:67. 1836.

Eupatorium glabrum Heyne ex Wallich,
mam List. Dr pl. 3283., 1831, nom.
nud.

Decaneurum glabrum DC., Prodr. 5:67.
1836.

Gymnanthemum glabrum (DC.) Sch. Bip.
ex Walp., Rep. 2:948. 1843.

Gymnanthemum silhetense (DC.) Sch.
Bip. ex Walp., Rep. 2:948. 1843.

Xipholepis silhetensis (DC.) Steetz in Pe-
ters, Reise Mossamb. Bot. 344. 1864.

Vernonia bracteata Wall. ex C. B.
Clarke, Comp. Ind. 17. 1876.

Vernonia silhetensis (DC.) Hand.-Mazz..,
Symb. Sin. 7:1084. 1936.

China, India, Thailand.

Acilepis spirei (Gandog.) H. Rob., comb.
nov. basionym: Vernonia spirei Gandog.,
Bull. Soc. Bot. France 54:194. 1907.
China, Laos, Vietnam.

Acilepis squarrosa D. Don, Prodr. Fl. Nepal
re. 1525.

Vernonia squarrosa (D. Don) Less., Lin-
naea 6:627. 1831.

Vernonia rigiophylla DC., Prodr. 5:15.
1836.

Vernonia teres Wall. ex DC., Prodr. 5:15.
1836.
China, India, Nepal, Sikkim, Thailand,
Vietnam.

227

Bechium DC., Prodr. 5:70. 1836. Type: Be-
chium scapiforme DC.

Erect or subhorizontally proliferating an-
nual or biennial herbs, 1—4 dm high; hairs
mostly of one long cell, appearing seri-
ceous, red stipitate glands with multicellular
tips on stems and bracts. Leaves alternate,
rosulate or subrosulate, subsessile, blades
oblong. Inflorescences scapiform with 1 to
many corymbosely disposed heads. Heads
shortly to longly pedunculate; involucral
bracts ca. 30 in ca. 3 series, with red stip-
itate glandular hairs; receptacle epalea-
ceous. Florets 25—50; corollas reddish-vio-
let, slender tubes funnelform above, throat
very short, lobes with sparse biseriate non-
glandular hairs, rarely with stipitate reddish
gland at tip; anther bases rounded, without
tails; apical appendage glabrous, with thin
cell walls; style base with distinct wide
node; style with sweeping hairs fat, pointed,
few to many septate. Achenes 10-costate,
with many unevenly pointed setulae, many
idioblasts, raphids elongate with rhomboid
tips; pappus with single series of easily de-
ciduous to subpersistent bristles and short
outer squamae, bristles narrowed below,
some wider distally. Pollen tricolporate,
non-lophate, echinate.

The genus is recognized primarily on the
basis of the herbaceous annual or biennial
habit, elongate raphids of the achene wall,
slender white pappus bristles, type A pol-
len, and the rather broad, pointed sweeping
hairs with few to many septations. The spe-
cies placed in the genus here are two of the
members of the Vernonieae in Madagascar
that have reddish glands with multicellular
caps on their peduncles and involucral
bracts. Both species need new combinations
since the oldest name for the type species
has never been transferred to the genus.
Bechium nudicaule (Less.) H. Rob., comb.

nov. basionym: Vernonia nudicaulis
Less., Linnaea 6:637. 1831.

Bechium scapiforme DC., Prodr. 5:71.
1836.
Vernonia scapiformis (DC.) Drake, Bull.

228

Soc. Bot. France 46:244. 1889.
Madagascar.

Bechium rhodolepis (Baker) H. Rob.,
comb. nov. basionym: Vernonia rhodo-
lepis Baker, J. Bot. 20:139. 1882.

Vernonia purpureo-glandulosa Klatt,
Bot. Jahrb: Syst!” 12°(Beible 27):20
1890.

Madagascar.

Cabobanthus H. Rob., gen. nov. (Erlangei-
nae).
Type: Vernonia polysphaera Baker.

Plantae herbaceae perennes base tuberosae.
Caules subglabri aut tomentosi, pilis simpli-
cibus uniseriatis multiseptatis. Folia alterna
sessilia vel subsessilia. Capitula in axillares
superiores ad 5 aggregata; involucra infun-
dibularia, bracteis ca. 35 in seriebus ca. 5
ovatis vel oblongis apiculatis; receptacula
epaleacea. Flores ca. 10 in capitulo; corollae
purpureae, tubis cylindricibus superne infun-
dibularibus; faucibus quam theceis dimidiis
brevioribus; thecae base breviter caudatae;
appendices antherarum apicales in parietibus
cellularibus tenues non glanduliferae; basi
stylorum noduliferi; rami stylorum aciculifor-
miter papillosi. Achenia 8—10-costatae, setulis
in cellulis aequales, raphidis minutis anguste
oblongis; setae pappi arcte barbellatae. Grana
pollinis triporata ca. 35 ym lacunosa emicro-
punctata.

The genus is notable for the erect stems
bearing axillary clusters of heads and the
triporate, emicropunctate pollen grains. The
following two species are recognized:

Cabobanthus bullulatus (S. Moore) H.
Rob., comb. nov., basionym: Vernonia
bullulata S. Moore, J. Bot. 65, suppl. 2:
44. 1927.

Zambia.

Cabobanthus polysphaerus (Baker) H.
Rob., comb. nov., basionym: Vernonia
polysphaera Baker, Kew Bull. 1898:148.
1898.

Vernonia humblei De Wild., Repert.
Spec. Nov. Regni Veg. 13:207. 1914.
Congo, Tanzania, Zambia.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Cyanthillium Blume, Bidjr. 889. 1826.
Type: Cyanthillium villosum Blume.

Isonema Cass., Bull. Soc. Philom. Paris
1817:152. 1817, hom. illeg., not Jso-
nema R. Br., 1810. Type: Isonema
ovata Cass.

Vernonia sect. Tephrodes DC., Prodr. 5:
24. 1836. Lectotype: Conyza cinerea
Blume (Jones 1981).

Cyanopsis Blume ex DC., Prodr. 5:69.
1836, nom. illeg. et superfl., not Cass.,
1817.

Claotrachelus Zoll. & Moritz ex Zoll.,
Natuur-Genrrsk. Arch. Ned. Indié 2:
263, 565. 1845. Type: Claotrachelus
rupestris Zoll. & Moritz ex Zoll.

Seneciodes L. ex Post & O. Kuntze, Lex.
Gen. Phan. 2:515. 1903. Type: Conyza
cinerea L.

Triplotaxis Hutch., Bull. Misc. Inform.
1914:355. 1914. Lectotype: Herderia
stellulifera Benth. in Hook.f. (Robin-
son 1990).

Vernonia subsect. Tephrodes (DC.) S. B.
Jones, Rhodora 83:70. (1981).

Annual or short-lived perennial herbs ca. 1
m tall; stem hairs asymmetrically and sym-
metrically T-shaped. Leaves alternate, nar-
rowly petiolate, blades thinly papery. Inflo-
rescences terminal, corymbose to pyramidal
cymes. Heads pedunculate; involucral bracts
papery, green with pale or purplish margins,
ca. 30 in 3(—5) gradate series, persistent; re-
ceptacle epaleaceous. Florets 15—94; corollas
bluish to lavender, funnelform with slender
lower tubes, throat a third as long to nearly
as long as lobes, lobes with simple hairs es-
pecially near tips; anthers without tails; apical
appendages glabrous, with thin cell walls;
style base with broad node; sweeping hairs
acicular. Achenes 5-angled or ribbed, or te-
rete, with idioblasts, sometimes with glands,
raphids elongate; pappus with many long,
fragile, slender-tipped bristles or squamellae,
persistent, with callose ring in one species.
Pollen triporate, echinolophate. N = 9, 11, 18
(Jones 1979, 1982).

The synonymy follows that in Robinson

VOLUME 112, NUMBER 1

(1990a) with the removal of Vernonia sub-
sections Orbisvestus and Hilliardianae. At-
tempts to interpret the genus more broadly to
include Gutenbergia Sch.Bip. ex Walp. (Rob-
inson, 1990b), are rejected here. At this time,
other more closely related species, with
shrubbier habits, non-lophate pollen, tailed
anthers and T-shaped hairs or no hairs on the
corolla lobes, are placed in separate genera
(see Hilliardiella and Orbivestus below). The
species of Cyanthillium are all annuals or
weak perennial subshrubs with triporate lo-
phate pollen, The following seven species are
presently placed in the genus.

Cyanthillium albicans (DC. in Wight) H.
Rob., comb. nov. basionym: Vernonia al-
bicans DC. in Wight, Contrib. Bot. Ind.
6. 1834.

Western peninsular India.

Cyanthillium cinereum (L.) H. Rob., Proc.
Biol. Soc. Wash. 103:252. 1990.

Conyza cinerea L., Sp. Pl. 862. 1753.

Vernonia cinerea (L.) Less., Linnaea 4:
291. 1829.

Seneciodes cinerea (L.) Post & O. Kuntze,
Lex. Gen. Phan. 2:515. 1903.
Throughout paleotropical region,
widely adventive in Neotropics.

Cyanthillium conyzoides (DC. in Wight) H.
Rob., comb. nov. basionym: Vernonia
conyzoides DC. in Wight, Contr. Bot. Ind.
6. 1834.

Western peninsular India.

Cyanthillium hookerianum (Arn.) H. Rob.,
comb. nov. basionym: Vernonia hooker-
tana Arn., Nov. Act. Nat. Cur. 18:346.
1836.

Sri Lanka.

Cyanthillium patulum (Ait.) H. Rob., Proc.
Biol. Soc. Wash. 103:252. 1990.

The synonymy is as in Robinson (1990a)
with the exclusion of Conyza chinensis
Lam. and its combinations.

Tropical Asia, Indonesia, Madagascar,

adventive in West Indies.

Cyanthillium — stelluliferum (Benth.) H.

229

Rob., Proc. Biol. Soc. Wash. 103:252.
1990.

Herderia stellulifera Benth. in Hook. f.
& Benth., Niger Fl., 425. 1849.

Triplotaxis stellulifera (Benth.) Hutch.,
Bull. Misc. Inf. Kew 1914:356. 1914.
West and central tropical Africa,
Uganda to Angola.

Cyanthillium vernonioides (Muschl.) H.
Rob., comb. nov. basionym: Erlangea
vernonioides Muschl., Bot. Jahrb. Syst.
46:62. 1911, not Vernonia vernonioides
(A. Gray) Bacigalupo, 1931.

Vernonia meiostephana C. Jeffrey, Kew
Bull. 43:225. 1988.

Hilliardiella H. Rob. gen. nov. (Erlangei-
nae). Type: Vernonia pinifolia Less.

Webbia DC., Prodr. 5:72. Oct. 1836, lec-
totype Vernonia pinifolia Less., hom.
illeg., not Webbia Spach, Jun 1836.

Vernonia subsect. Hilliardianae S. B.
Jones, Rhodora 83:66. 1981. Type: Ver-
nonia oligocephala (DC.) Sch. Bip.

Plantae herbaceae perennes ad 1 m altae;
caules pilosi, pilis aequaliter T-formibus.
Folia alterna; laminae subtus saepe dense
canescentiter pilosae. Inflorescentiae laxe
vel subdense corymbose cymosae; capitula
pedunculata; involucra campanulata; brac-
teae 25—40 ca. 3—4-seriatae persistentes; re-
ceptacula epaleacea. Flores 12—20 in capi-
tulo; corollae purpureae extus pauce vel
dense pilosae, pilis T-formibus leniter con-
tortis; tubis superne infundibularibus, fau-
cibus brevibus, lobis linearibus; thecae base
non vel breviter appendiculatae; appendices
apicales antherarum glabrae in parietibus
cellularum tenuis; basi stylorum noduliferi,
papillae ramorum aciculiformes. Achenia
4—5-costata dense setulifera et idioblasti-
fera, raphidis elongatis; carpopodia anguste
cylindrica; setae pappi albae barbatae ten-
ues subpersistentes, seriebus exteriores
breviter lanceolatis. Grana pollinis non lo-
phata tricolporata echinata. Numerus chro-
mosomatum N = 9, 10 (Jones 1942).

Chemistry reported for the genus in-

230

cludes germacranolides, hirsutanolides,
quaianolides and bisabolene derivatives
(Bohlmann & Jakupovic 1990). Most chro-
mosome counts have reported N = 10.

Hilliardiella is closely related to Cyan-
thillium, but it seems consistently different
by the more perennial habit, the non-lophate
pollen, and especially the T-shaped hairs of
the corolla. The group has been rather well
defined at various levels by Candolle
(1836) and Jones (1981).

The name is derived from the subsectional
name of Jones (1981), but the genus is de-
scribed as new to avoid any complications
from spelling. The name, as indicated by
Jones (1981), honors Dr. Olive M. Hilliard,
student of the Asteraceae of Natal. The fol-
lowing eight species are credited to the genus.

Hilliardiella aristata (DC.) H. Rob., comb.
nov. basionym: Webbia aristata DC.,
Prodr. 5:73. 1836, not Vernonia aristata
Less: 1829:

Vernonia natalensis Sch. Bip. ex Walbp.,
Rep. 2:947. 1843.
South Africa.

Hilliardiella calyculata (S. Moore) H.
Rob., comb. nov. basionym: Vernonia
calyculata S. Moore, J. Linn. Soc. Bot.
35:316. 1902.

Congo, Tanzania, Malawi, Mozambique,
Zambia.

Hilliardiella hirsuta (Sch. Bip. ex Walp.)
H. Rob., comb. nov. basionym: Vernonia
hirsuta Sch. Bip. ex Walp., Rep. 2:947.
1843.

South Africa.

Hilliardiella leopoldii (Vatke) H. Rob.,
comb. nov. basionym: Vernonia leopoldii
Vatke, Linnaea 39:478. 1878.

Ethiopia.

Hilliardiella nudicaulis (DC.) H. Rob.,
comb. nov. basionym: Webbia nudicaulis
DC Prodr. 527321836:

Vernonia dregeana Sch. Bip. ex Walp.,
Rep. 2:947. 1843.
South Africa.

Hilliardiella oligocephala (DC.) H. Rob..,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

comb. nov. basionym: Webbia oligoce-
phala DC., Prodr. 5:73. 1836.

Webbia elaegnoides DC., Prodr. 5:73.
1836, not Vernonia elaegnoides
H.B.K., 1818.

Vernonia oligocephala (DC.) Sch. Bip.
ex Walp., Rep. 2:947. 1843.

Vernonia krausii Sch. Bip. ex Walp.,
Rep. 2:947. 1843.

Tanzania south to the Cape Province.

Hilliardiella pinifolia (Less.) H. Rob.,
comb. nov. basionym: Vernonia pinifolia
Less., Linnaea 4:257. 1829.

Webbia pinifolia (Less.) DC., Prodr. 5:72.
1836.
South Africa.

Hilliardiella smithiana (Less.) H. Rob.,
comb. nov. basionym: Vernonia smithi-
ana Less., Linnaea 6:638. 1831.

West and Central Tropical Africa.

Orbivestus H. Rob., gen. nov. (Erlangei-
nae). Type: Vernonia karaguensis Oliv.
& Hiern.

Vernonia subg. Orbisvestus S. B. Jones,
Rhodora 83:60. 1981. Type: Vernonia
karaguensis Oliv. & Hiern.

Plantae suffrutescentes vel frutescentes;
caules pilosi, pilis T-formibus. Folia alterna
breviter petiolata; laminae sparse pilosae. In-
florescentiae laxe corymbose cymosae. Ca-
pitula pedunculata; involucra campanulata;
bracteae 25—30 ca. 4-seriatae persistentes; re-
ceptacula epaleacea. Flores 8—16 in capitulo;
corollae purpureae extus non vel perpauce pi-
losae, pilis raris sub-T-formibus, tubis super-
ne leniter infundibularibus, faucibus et lobis
subaequilongis, lobis longe triangularibus;
thecae antherarum base breviter vel longe
caudatae; appendices antherarum apicales
glabrae in parietibus cellularum tenues; basi
stylorum breviter et distincte noduliferi; pa-
pillae ramorum aciculiformes. Achenia 4—5-
costata setulifera multo idioblastifera, raphi-
dis elongatis; carpopodia anguste cylindrica;
setae pappi barbellatae facile deciduae,
Squamis exterioribus brevis persistentibus.
Grana pollinis non lophata tricolporata echin-

VOLUME 112, NUMBER 1

ata. Numerus chromosomatum N = 9, 20
(Mangenot & Mangenot 1962, Mehra et al.
1965, Jones 1982).

The genus Orbivestus is distinct from the
related Cyanthillium by the more shrubby
habit, the lack of simple hairs on the corolla
lobes, and the non-lophate pollen. It is dis-
tinct from Hilliardiella by the longer co-
rolla throats and more triangular lobes that
bear few or no hairs. The few hairs seen on
the corolla lobes in the type species are
asymmetrically T-shaped, quite different
from those in Hilliardiella. The genus has
more prominent tails on the anthers than in
either of the related genera. Chemical con-
stituents include 5-alkyl coumarins, bisa-
bolene derivatives and glaucolides (Bohl-
mann & Jakupovic 1990).

Although the subgeneric name by Jones
(1981) is the inspiration for the generic
name, the genus has been described as new
to avoid the consistent extra “‘s’’ in the
Jones spelling. Jones did not explain his
spelling, and the reason for his choice is
unknown. The spelling used here is that
found in Jeffrey (1988) and Bohlmann &
Jakupovic (1990). Still, it seems unneces-
sary to completely abandon the name
coined by Jones.

The following four species are placed in
the genus.

Orbivestus cinerascens (Sch. Bip. in
Schweinf.) H. Rob., comb. nov. basio—
nym: Vernonia cinerascens Sch. Bip. in
Schweinf., Beitr. Fl. Aeth. 162. 1897.

Vernonia tephrodioides Chiov., Fl. So-
italien: 25. 1932.
Senegal east to western India, south to
Angola and Botswana.

Orbivestus homilanthus (S. Moore). H.
Rob., comb. nov. basionym: Vernonia
homilantha S. Moore, J. Bot. 41:138.
1903.

Vernonia sennii Chiov., Fl. Somal. 2:256.
1932.
Kenya, Somalia.

Orbivestus karaguensis (Oliv. & Hiern) H.

231

Rob., comb. nov. basionym: Vernonia
karaguensis Oliv. & Hiern, Trans. Linn.
Soc. London 29:91. 1873.

Vernonia cistifolia O. Hoffm., Engl.
Pflanzenw. Ost-Afr. C 404. 1895.

Vernonia elliotii S. Moore, J. Linn. Soc.
Bot, 35:315. 1902.

Vernonia bothrioclinoides C. H. Wright,
Bull. Misc. Inf. 1906:108. 1906.

Vernonia porphyrolepis S. Moore, J. Bot.
46:39. 1908.

Vernonia campanea S. Moore, J. Bot. 54:
Zl el ONG:

Vernonia melanacrophylla Cufod., Nouv.
Giorn. Bot. Ital., n.s. 50:102. 1943.
Sudan south to Mozambique and Zim-
babwe, west to Nigeria.

Orbivestus undulatus (Oliv. & Hiern) H.
Rob., comb. nov. basionym: Vernonia
undulata Oliv. & Hiern, Fl. Trop. Afr. 3:
276. 1877.

West and central tropical Africa, north to
Sudan, south to Angola.

Oocephala (S. B. Jones) H. Rob., stat. nov.
(Erlangeinae).

Vernonia subsect. Oocephalae S. B.
Jones, Rhodora 83:72. 1981.

Type species: Vernonia oocephala Baker.

Low much-branched shrubs to 1 m high,
stems with weakly L-shaped simple hairs and
with multiseptate simple hairs. Leaves alter-
nate, subsessile, linear to elliptical, sometimes
serrate. Inflorescences corymbose cymes with
usually shortly pedunculate heads or with
heads sessile in apical clusters of leaves. In-
volucre ovoid or cylindrical; bracts 20—40 in
4-7 gradate series, ovate to oblong, ap-
pressed; receptacle without pales. Florets ca.
15 in a head; corollas white or lavender, tu-
bular to narrowly funnelform, throat as long
as lobes, tips without hairs or with few short
biseriate hairs; anther bases rounded, apical
appendages glabrous, with thin-walled cells;
style base with indistinct ring; style branches
with acicular sweeping hairs. Achenes weak-
ly 8-ribbed, sericeous with many setulae, id-
ioblasts numerous, raphids narrowly elon-

232:

gate; pappus biseriate, outer shorter and

broader, inner setiform, subplumose, glabrous

near base. Pollen triporate, lophate, minutely
papillate on murae, emicropunctate or weakly
micropunctate.

The genus is distinquished by subplumose
inner pappus, triporate pollen and stems with
weakly L-shaped hairs. Vernonia sect. Ooce-
phala is based on a single species with pe-
dunculate ovoid heads containing few tubular
florets. The additional species here referred
to Oocephala was described by Jeffrey
(1988) in his Group 3 subgroup A with Cen-
trapalus, but the achene is most like Ooce-
phala and Vernoniastrum, and a small corolla
remnant shows triporate pollen. The species
is not related to the Centraplinae, and 1s
placed here on the basis of its subplumose
pappus, in spite of considerable difference in
general appearance.

The genus contains the following two
species.

Oocephala agrianthoides (C. Jeffrey) H.
Rob., comb. nov. basionym: Vernonia
agrianthoides C. Jeffrey, Kew Bull. 43:
2271-1988,

Burundi, Congo, Tanzania.

Oocephala stenocephala (Oliv.) H. Rob.,
comb. nov. basionym: Vernonia steno-
cephala Oliv. in Hook., Ic. Pl. 14:35, t.
1349A. 1881.

Vernonia oocephala Baker, Bull. Misc.
Inf. 1895:68. 1895.

Vernonia luteoalbida De Wild., Repert.
Spec. Nov. Regni Veg. 13:207. 1913.
Nigeria east to Tanzania and south to
Mozambique.

Polydora Fenzl, Flora 27:312. 1844. Type
species: Polydora stoechadifolia Fenzl =
Webbia serratuloides DC.

Crystallopollen Steetz in Peters, Reise
Mossamb. Bot. 363. 1864. Type spe-
cies: Crystallopollen angustifolium
Steetz.

Mostly annuals; stems with one-armed T-
Shaped hairs. Leaves alternate. Inflores-
cence a lax thyrsoid panicle with corym-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

bosely cymose branches with pedunculate

heads or with single terminal head. Invo-

lucral bracts ca. 80 in ca. 7 series, often
with widely scarious margins and awned
often black tips, receptacles epaleaceous.

Florets ca. 30 in a head; corollas whitish to

purplish, tube longly funnelform, throat as

long as the narrow glabrous lobes; anther
bases plain, not tailed; apical appendage
glabrous, with cell walls thin, sometimes
weakly ornate; style base with distinct an-
nular node; branches with acicular sweep-
ing hairs. Achenes 5 or 8—10 ribbed, setu-
liferous, raphids narrowly elongate; pappus
with copious barbellate setae, greenish, yel-
lowish or tawny, rarely white; outer pappus
short, squamiform. Pollen triporate, scarce-
ly echinolophate to psilolophate, with or
without micropunctations. Chromosome
number N = 9 (Jones 1979, 1982). Report-
ed sesquiterpene lactones are germacranol-
ides, hirsutanolides and furanheliangolides

(Bohlmann & Jakupovic 1990).

The genus is distinct in the annual habit,
chromosome number of N = 9, the one-
armed T-shaped hairs, untailed anthers and
triporate pollen.

Some of the species of the Vernonia
chloropappa group are keyed by Pope
(1986).

The genus is here credited with the fol-
lowing eight species.

Polydora angustifolia (Steetz in Peters) H.
Rob., comb. nov. basionym: Crystallo-
pollen angustifolium Steetz in Peters, Re-
ise Mossamb. Bot. 366. 1864, not Ver-
nonia angustifolia Michx., 1803 or V. an-
gustifolia D. Don ex Hook. & Arn., 1835.

Vernonia erinacea H. Wild, Kirkia 11:2.
1978.
Tanzania and Mozambique, east to
Zambia and Zimbabwe.

Polydora bainesti (Oliv. & Hiern) H. Rob.,
comb. nov. basionym: Vernonia bainesii
Oliv. & Hiern in Oliv., Fl. Trop. Afr. 3:
2A2 STF.

Tanzania and Mozambique, east to Zam-
bia and Zimbabwe.

VOLUME 112, NUMBER 1

Polydora chloropappa (Baker) H. Rob.,
comb. nov. basionym: Vernonia chloro-
pappa Baker, Bull. Misc. Inf. 1898:146.
1898.

Vernonia kassneri De Wild. & Muschl.,
Bull. Soc. Bot. Belg. 49:242. 1912.
Vernonia smaragdopappa S. Moore, J.

Linn. Soc. Bot. 47:284. 1925.
Congo, Malawi, Tanzania, Zambia.

Polydora jelfiae (S. Moore) H. Rob., comb.
nov. basionym: Vernonia jelfiae S.
Moore, J. Linn. Soc. Bot. 47:262. 1925.
Angola, Burundi, Congo, Malawi, Tan-
zania, Zambia, Zimbabwe.

Polydora poskeana (Vatke & Hildebr.) H.
Rob., comb. nov. basionym: Vernonia
poskeana Vatke & Hildebr., Oesterr. Bot.
Zettsehr. 25:324. 1875.

Angola, Botswana, Nambia, Transvaal,
Zimbabwe.

Polydora serratuloides (DC.) H. Rob.,
comb. nov. basionym: Webbia serratu-
loides DC., Prodr. 5:72. 1836, not Ver-
nonia serratuloides H. B. K., 1818.

Vernonia perrottetii Sch. Bip. ex Walp.,
Rep. 2:947. 1843.

Polydora stoechadifolia Fenzl, Flora 27:
312. 1844.
West and central tropical Africa from
Sudan and Ethiopia south to Angola
and Zambia.

Polydora steetziana (Oliv. & Hiern) H.
Rob., comb. nov. basionym: Vernonia
steetziana Oliv. & Hiern in Oliv., FI.
Trop Afr. 3:278. 1877.

South Africa.

Polydora sylvicola (G. V. Pope) H. Rob.,
comb. nov. basionym: Vernonia sylvicola
G. V. Pope., Kew Bull. 41:395. 1986.
Angola, Congo, Malawi, Mozambique,
Tanzania, Zambia, Zimbabwe.

Vernoniastrum H. Rob., gen. nov. (Erlan-
geinae) Type: Crystallopollen latifolium
Steetz in Peters

Vernonia sect. Lepidella Oliv. & Hiern,
Fl. Trop. Afr. 3:267. 1877. Type: Ver-
nonia petersii Oliv. & Hiern, not Lep-

235

idella Tiegh., 1912 or Lepidella E. J.
Gilbert, 1925.

Vernonia subsect. Lepidella (Oliv. &
Hiern) S. B. Jones, Rhodora 83:72.
1981.

Plantae herbaceae perennes 0.3—1.0 m al-
tae; caules pilosi, pilis simplices in cellulis
apicalibus elongatis base leniter inaequalibus.
Folia alterna. Inflorescentiae uni- vel multo
capitatae. Involucra campanulata; bracteae ca.
50 in ca. 3 series gradatae persistentes; re-
ceptacula epaleacea. Flores ca. 50 in capitulo;
corollae rubro-purpurascentes, tubis anguste
infundibularibus, faucibus quoad lobis et the-
cis brevioribus, lobis distaliter pilosis; thecae
antherarum base acuminate vel acute cauda-
tae; appendices antherarum apicales glabrae
in parietibus cellarum tenues; base stylorum
noduliferi; rami stylorum aciculariter piliferi.
Achenia 4—5-angulata, idioblastis saepe in
seriebus transversalibus aggregatis, raphidis
elongatis; setae pappi interiores margine
dense barbellatae subpersistentes, squamae
exteriores lateriores persistentes. Grana pol-
linis triporata lophata micropunctata vel em-
icropunctata. Numerus chromosomatum N =
10 (Jones 1979, 1982).

Vernoniastrum seems closely related to
the foregoing Polydora Fenzl, but differs by
the perennial habit, the non-T-shaped hairs,
the tailed anther bases and the chromosome
number of N = 10. The core element of the
genus also has the idioblasts of the achene
in distinct transverse bands, a feature not
seen outside of the genus Vernoniastrum.

The genus is credited here with the fol-
lowing eight species:

Vernoniastrum aemulans (Vatke) H. Rob.,
comb. nov. basionym: Vernonia aemu-
lans Vatke, Oesterr. Bot. Zeitschr. 27:195.
1877.

Kenya, Tanzania.

Vernoniastrum ambiguum (Kotschy &
Peyr.) H. Rob., comb. nov. basionym:
Vernonia ambigua Kotschy & Peyr., Pl.
Hanes 35; t:l7 BY 186s:

West tropical Africa to Sudan and Tan-
zania and south to Angola.

234

Vernoniastrum latifolium (Steetz in Peters)
H. Rob., comb. nov. basionym: Crystal-
lopollen latifolium Steetz in Peters, Reise
Mossamb. Bot. 364, t. 48a. 1864, not V.
latifolia Lem., 1855.

Vernonia petersii Oliv. & Hiern, Trans.
Linn. Soc. London 29:90. 1873.
Vernonia eriocephala Klatt, Bull. Herb.
Boiss. 4:826. 1896.
Angola and Congo east to Mozam-
bique and Tanzania.

Vernoniastrum musofense (S. Moore) H.
Rob., comb. nov. basionym: Vernonia
musofensis S. Moore, J. Bot. 56:206.
1918:

Vernonia lappoides O. Hoffm., Bol. Soc.
Brot. 13:19. 1896, hom. illeg., not
Baker, 1873.

Vernonia miamensis S. Moore, J. Bot.
64:304. 1926.

Vernonia hoffmanniana Hutsch. & Dalz.,
Bly Wo iropsAte2> 167. [93s mom.
nov. for V. lappoides O. Hoffm.

Vernonia philipsoniana Lawalree, Expl.
Hydrobiol. Lac. Tanganyika (1946—
47) Rés. Sc. 4, 2:59. 1955; nom. nov.
superfl. for V. lappoides O. Hoffm.
Tropical Africa from Nigeria to Tan-
zania, south to Angola and Zimbabwe.

Vernoniastrum nestor (S. Moore) H. Rob.,
comb. nov. basionym: Vernonia nestor S.
Moore, J. Linn. Soc. Bot. 35:317. 1902.

Vernonia chariensis O. Hoffm., Bull.
Soc. Bot. France 55, mém. 8:40. 1908.
Vernonia cannabina Muschl., Bot. Jahrb.
Syst. 46:94. 1911.
West Africa east to Tanzania south to
Natal.

Vernoniastrum ugandense (S. Moore) H.
Rob., comb. nov. basionym: Vernonia
ugandensis S. Moore, J. Linn. Soc. Bot.
35:314. 1902.

Vernonia caput-medusae S. Moore, J.
Linn. Soc. Bot. 37:166. 1905.

Vernonia fontinalis S. Moore, J. Bot. 52:
90. 1914.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Vernonia punctulata S. Moore, J. Linn.
Soc. Bot. 47:262. 1925.

Vernonia proclivicola S. Moore, J. Linn.
Soc. Bot. 47:262. 1925.

Vernonia mgetae Gilli, Ann. Naturhist.
Mus. Wien 78:164. 1974.
Congo, Burundi and Tanzania south to
Angola, Zambia and Mozambique.

Vernoniastrum uncinatum (Oliv. & Hiern
ex Oliv.) H. Rob., comb. nov. basionym:
Vernonia uncinata Oliv. & Hiern ex
Ohiv., Fl. Trop Afr 3:277. 1877:

Vernonia amplexicaulis Baker, Kew Bull.
1895:216:21895:
Ethiopia, Kenya, Somalia.

Vernoniastrum viatorum (S. Moore) H.
Rob., comb. nov. basionym: Vernonia
viatorum S. Moore, J. Linn. Soc. Bot. 35:
315: 1912.

Malawi.

Koyamasia H. Rob., gen. nov. (Erlangei-
nae). Type: Camchaya calcarea Kitam.

Plantae herbaceae perennes ad 0.5 m al-
tae pauce ramosae; caules pilis simplicibus
multiseptatis obsiti. Folia alterna anguste pe-
tiolata. Inflorescentiae terminales et axillares.
Capitula longe pedunculata late campanulata;
bracteae involucri ca. 90 ca. 4—5-seriatae per-
sistentes in partibus majoribus herbaceae et
reflexae; receptaculm glabrum. Flores ca. 90
in capitulo; corollae tenuiter carnosae, tubis
angustis, faucibus abrupte campanulatis, lobis
quoad faucibus leniter longioribus non reflex-
is extus glanduliferis; thecae antherarum val-
de exertae saepe nigricans base rotundatae,
cellulis endothecialibus elongatis, in apicibus
noduliferis; appendices apicales antherarum
non longiores quam lateriores glabrae in par-
ietibus cellularum firmis; basi stylorum non
noduliferi. Achenia 10-costata glabra; raphi-
dis minutis anguste oblongis; setae pappi
paucae breves facile deciduae. Grana pollinis
triporata emicropunctata. Numerus chromo-
somatum 2N = 54 (Koyama 1984).

The genus Koyamasia is established for a
single Southeast Asian species occurring in
limestone areas. It was originally described

VOLUME 112, NUMBER 1

in Camchaya Gagnep., but was excluded
from it by Koyama (1984). The species is
similar to Camchaya in its geography and re-
duced pappus, but it differs in the triporate
rather than tricolporate lophate pollen and the
simple rather than T-shaped hairs. The genus
may be more closely related to Kinghamia C.
Jeffrey, which contrary to its original descrip-
tion, has triporate rather than tricolporate pol-
len, and is not closely related to Linzia. The
West African Kinghamia is similar, but is a
less robust plant with narrower leaf blades
and much smaller heads. Kinghamia may
represent convergent evolution in many of its
similarities, but it seems to differ from Koy-
amasia by only some technical characteristics
such as the reflexed mature corolla lobe tips,
the presence of a basal stylar node, the pale
and only partially exerted anthers, the longer
anther appendages and the subquadrate en-
dothecial cells without nodular thickenings.
The single species is as follows:

Koyamasia calcarea (Kitam.) H. Rob.,
comb. nov. basionym: Camchaya calca-
rea Kitam., Acta Phytotax. Geobot. 23:
71. 1968.

Thailand.

Phyllocephalum Blume, Bidjr. Fl. Ned. Ind.
888. 1826. Type: Phyllocephalum frutes-
cens Blume

Decaneurum DC. ex Wight, Contr. Bot.
Ind. 7-8. (1833), not Decaneurum
DC., 1836. Type: Decaneurum reti-
culatum DC. ex Wight [= Phylloce-
Phalum indicum (Less.) Kirkman]

Rolfinkia Zenker, Pl. Ind. 13. 1837. Type:
Rolfinkia centaureoides Zenker [=
Phyllocephalum indicum (Less.) Kirk-
man].

Lamprachaenium Benth, in Benth. &
Hook. f., Gen. pl. 2:225. (1873). Type:
Decaneurum microcephalum Dalzell

The herbaceous genera Phyllocephalum
Blume of India and Indonesia, with three spe-
cies (Kingham, 1981), and Lamprachaenium
Benth. of India, with one species, have been
examined and found to differ mostly by the
costate achenes of the former versus the shiny

235

dark ecostate achenes of the latter. In both
genera, the achenes are oblong and glabrous,
somewhat obcompressed, abruptly rounded
above to the narrow attachment of the corol-
la, the pappus is of short highly deciduous
setae and the raphids in the achene wall are
subquadrate. Both genera have lophate tri-
porate pollen with minutely spinulose mar-
gins of the muri as seen in Kirkman (1981)
and Robinson & Marticorena (1986). Both
have foliose lower involucral bracts or foliose
tips on the bracts. The genera are here con-
sidered the same and the needed combination
is as follows:

Phyllocephalum microcephalum (Dalzell)
H. Rob., comb. nov. basionym: Decaneu-
rum microcephalum Dalzell, Hooker’s J.
Bot. Kew Gard. Misc. 3:231. 1851. India.

New Genera and New Combinations of
Eastern Hemisphere Vernonieae
Subtribe Centrapalinae H. Rob.

Centrapalus Cass., Bull. Soc. Philom. Paris
1817:10. 1817. Type: Centrapalus gala-
mensis Cass.

Vernonella Sond., Linnaea 23:62. 1850.
Type: Vernonella africana Sond.

Annual or perennial, scapose or subsca-
pose herbs, with or without brown woolly
hair on crown of rootstock, anthesis often
prior to leaf emergence; stem hairs simple,
multiseptate. Leaves basal or cauline, alter-
nate, sessile. Inflorescences terminal on
stems and branches. Involucre hemispheri-
cal; bracts 125-150 in ca. 5 gradate series,
linear, green, often with small teeth on dis-
tal margin; receptacle epaleaceous. Florets
ca. 100 in a head; corollas light blue to blu-
ish purple, tube funnelform above, throat
nearly half as long as thecae, lobes some-
times fringed with long papillae; anther
base not tailed; apical anther appendage
glabrous, often colored, with slightly thick-
ened cell walls; style base with broad node.
Achenes weakly 10-costate, setuliferous,
raphids narrowly oblong; pappus setae
long, subpersistent, outer series setiform,

236

short. Pollen tricolporate, echinate, lophate
or non-lophate, without muri projecting into
colpi, micropunctate. Chromosome number
N = 9 (Jones 1974, 1979, 1982). Sesqui-
terpene lactones include elemanolides
(Bohlmann & Jakupovic 1990).

The name Centrapalus has been used
most recently within the broad concept of
Vernonia for a group of coarse annual or
perennial African herbs. The genus is of
some potential commercial importance for
the extraction of epoxy resins (Ayorinde et
al., 1990). The proper combination has not
previously been provided for the type spe-
cies, and an additional eight species are
placed in the genus here.

Centrapalus acrocephalus (Kliatt) H. Rob.,
comb. nov. basionym: Vernonia acroce-
phala Klatt, Ann. Hofmus. Wien 7:100.
1897.

Sierra Leone, Nigeria and Congo south
to Angola and Zimbabwe.

Centrapalus africanus (Sond.) H. Rob.,
comb. nov. basionym: Vernonella afri-
cana Sond., Linnaea 23:62. 1850.

South Africa.

Centrapalus chthonocephalus (O. Hoffm.)
H. Rob., comb. nov. basionym: Vernonia
chthonocephala O. Hoffm., Bol. Soc.
Bro 135175 1396:

Sierra Leone and Sudan south to Angola
and Malawi.

Centrapalus denudatus (Hutch. & Burtt)
H. Rob., comb. nov. basionym: Vernonia
denudata Hutch. & Burtt, Rev. Zool. Bot.
Abts 23:57 al 932-

Congo, Tanzania, Zambia.

Centrapalus kirkii (Oliv. & Hiern in Oliv.)
H. Rob., comb. nov. basionym: Vernonia
kirkii Oliv. & Hiern in Oliv., Fl. Trop.
Att. 3-274 AST

Vernonia swynnertonii S. Moore, J. Linn.
Soc. Bot. 40:107. 1911.

Vernonia zambesiaca S. Moore, J. Bot.
552102; 4917:
Tanzania south to Zambia and Moz-
ambique.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Centrapalus pauciflorus (Willd.) H. Rob.,
comb. nov. basionym: Conyza pauciflora
Willd., Sp. Pl. 3:1927. 1803.

Centrapalus galamensis Cass., Dict. Sci.
Nat. ed= 2, /2383, IiSi7-

Vernonia pauciflora (Willd.) Less., Lin-
naea 4:292. 1829, not (Pursh.) Poir.,
1817.

Vernonia afromontana R. E. Fr., Acta.
Hort. Berg. 9:116. 1929.

Vernonia zernyi Gilli, Ann. Naturhist.
Mus. Wein 78:165. 1974.

Vernonia petitiana A. Rich., Tent. FI.
Abyss. 1:373. 1848.

Sudan, Ethiopia, Somalia, Kenya,
Uganda, Tanzania, Malawi, Zambia,
Mozambique and West Africa.

Centrapalus praemorsus (Muschl.) H.
Rob., comb. nov. basionym: Vernonia
praemorsa Muschl., Bot. Jahrb. Syst. 46:
68. 1911
Angola, Congo, Malawi, Tanzania, Zam-
bia.

Centrapalus purpureus (Sch. Bip. ex
Walp.) H. Rob., comb. nov. basionym:
Vernonia purpurea Sch. Bip. ex Walp.,
Rep. 2:946. 1843.

Vernonia inulifolia Steud. ex Walp., Rep.
2:946. 1843.

Vernonia jaceoides A. Rich., Tent. FI.
Abyss. 1:376. 1848.

Vernonia rigorata S. Moore, J. Bot. 41:
155. 1903:

Vernonia scabrida C. H. Wright, Bull.
Misc. Inf. Kew 1906:21. 1906.

Vernnonia duemmeri S. Moore, J. Bot.
S29 VALLE

Vernonia pascuosa S. Moore, J. Linn.
Soc. Bot. 47:163. 1925.

Vernonia keniensis R. E. Fr., Acta Hort.
Berg. 9:114. 1929.
West, central and south tropical Africa
north to Sudan and Ethiopia.

Centrapalus subaphyllus (Baker) H. Rob.,
comb. nov. basionym: Vernonia suba-
phylla Baker, Bull. Misc. Inf. Kew 1895:
290. 1895.

VOLUME 112, NUMBER 1

Nigeria, Cameroon, Congo, Tanzania,
Angola, Malawi, Zambia.

Linzia Sch. Bip. ex Walp., Rep. 2:948.
1843. Type: Linzia vernonioides Sch.
Bip. ex Walp.

Vernonia sect. Azureae S. B. Jones, Rho-
dora 83:74. 1981. Type: Linzia glabra
Steetz in Peters

Perennial herbs; stems with simple mul-
tiseptate hairs. Leaves alternate, subsessile
to short-petiolate. Inflorescences corymbi-
form cymes or single heads with short to
long pecuncles. Involucre funnelform to
campanulate; bracts 50-150 in 5-6 series,
often pectinate-denticulate along distal mar-
gins, outer tips often elongate, green and
recurved; receptacle epaleaceous. Florets
ca. 20—50 in a head; corollas bluish, tube
very long, funnelform near throat, throat
very short, lobes apically stiffly hairy; an-
ther base rounded; apical anther appendage
glabrous, triangular, with thickened orna-
mentation in center; style base with small
annulus. Achenes strongly 10-costate, usu-
ally with rows of idioblasts along sides of
costae, setuliferous, raphids subquadrate to
short-oblong; pappus of many somewhat
persistent, long bristles, with short outer se-
ries. Pollen tricolporate, lophate, with muri
intruding into short colpi, single polar la-
cunae often present, with or without micro-
punctations. Chromosome number N = 10
(Jones 1979, 1982). Sesquiterpene lactones
include germacranolides and hirsutanolides
(Bohlmann & Jalupovic 1990).

Linzia has been recognized rather accu-
rately in recent literature because of the rather
characteristic bluish flowers (Jones 1981) or
distinctive pollen (Jeffrey 1988). The species
are related to Centrapalus, but they differ by
the more perennial habit, the very short
throats of the corollas, the stronger ribs on
the achenes, the idioblasts that are positioned
along those ribs in most species and the chro-
mosome number of N = 10.

The following seven species are credited
to the genus.

Linzia gerberiformis (Oliv. & Hiern in

237

Oliv.) H. Rob., comb. nov. basionym:
Vernonia gerberiformis Oliv. & Hiern in
Oly Flo Trop: Afe3:285.. 1877.

Vernonia macrocyanus O. Hoffm., Bol.
Soc. Brot. 13:20. 1896.

Vernonia nandensis S. Moore, J. Linn.
Soc.,.Bot. 55:323.,,1902.

Vernonia towaensis De Wild., Bull. Jard.
Bot. Brux. 5:96. 1915.
Angola, Burundi, Cameroon, Congo,
Malawi, Nigeria, Sudan, Tanzania,
Uganda, Zambia, Zimbabwe.

Linzia glabra Steetz in Peters, Reise Mos-
samb. Bot. 353. 1864.

Vernonia glabra (Steetz) Vatke, Oesterr.
Bot. Zeitschr. 27:194. 1877.

Vernonia obconica Oliv. & Hiern in
Oliv., Fl. Trop. Afr. 3:286. 1877.

Vernonia pogosperma Klatt, Ann. K.K.
Naturhist. Hofmus. 7:99. 1892.

Vernonia hindei S. Moore, J. Bot. 41:
13D 71 903:

Vernonia piovanii Chiov., Racc. Bot.
Miss. Cons. Kenya 61. 1935.

Vernonia roseopapposa Gilli, Ann. Na-
turhist. Mus. Wien 78:165. 1974.
Congo, Burundi, Kenya, Tanzania,
south to Angola, Namibia, Mozam-
bigue, Transvaal, Swaziland, Natal,
Madagascar.

Linzia infundibulariformis (Oliv. & Hiern
in Oliv.) H. Rob., comb. nov. basionym:
Vernonia infundibulariformis Oliv. &
Hiern in Oliv., Fl. Trop. Afr. 3:285. 1877.

Vernonia saussureoides Hutch., Bull.
Misc. Inf. Kew 1921:378. 1921.
Burundi, Congo, Cameroon, Nigeria,
Sudan, Tanzania, Uganda.

Linzia ituriensis (Muschl.) H. Rob., comb.
nov. basionym: Vernonia ituriensis
Muschl., Wiss. Ergebn. Deutsch. Zent.-
Afr. Exped. 1907-8 (2):364. 1911.

Vernonia hillii Hutch. & Dalz., Fl. W.
Trop. Afr. ed. 1, 2:165, 168. 1931.
Vernonia muhiensis Kalanda, Bull. Jard.

Bot. Nat. Belg. 52:125. 1982.

238

Burundi, Cameroon, Congo, Ethiopia,
Nigeria, Rwanda, Sudan, Tanzania.

Linzia melleri (Oliv. & Hiern in Oliv.) H.
Rob., comb. nov. basionym: Vernonia
melleri Oliv. & Hiern in Oliv., Fl. Trop.
Afr '3:282. AST?r

Vernonia superba O. Hoffm., Engl.
Pflanzenw. Ost-Afr. C. 406. 1895.
Vernonia scabrifolia O. Hoffm., Bot.
Jahrb. Syst. 30:424. 1901, nom. illeg.,

not Hieron., 1897.

Vernonia paludigena S. Moore, J. Bot.
52:91. 1914.

Vernonia vanmeelii Lawalrée, Expl. Hy-
drobiol. Lac Tanganyika. Rés. Sc. 6, 2:
59. 1955, nom. nov. for V. scabrifolia
O. Hoffm.

Angola, Congo, Malawi, Mozam-
bique, Rwanda, Tanzania, Zambia,
Zimbabwe.

Linzia vernonioides Sch. Bip. ex Walp.,
Rep. 2:948. 1843.

Vernonia quartiniana A. Rich., Tent. FI.
Abyss. 1:379. 1848.

Vernonia congolensis De Wild. &
Muschl., Bull. Soc. Bot. Belg. 49:237.
On.

Vernonia vernonioides (Sch. Bip. ex
Walp.) Cufod., Bull. Jard. Bot. Brux.
36, suppl.:1078. 1966, hom. illeg., not
(A. Gray) Bacigal, 1931.

Burundi, Congo, Ethiopia, Tanzania.
Linzia usafuensis (O. Hoffm.) H. Rob.,
comb. nov. basionym: Vernonia usafuen-

sis O. Hoffm., Bot. Jahrb. Syst. 30:425.

1901.

Vernonia candelabricephala Gilli, Ann.
Naturhist. Mus. Wien 78:161. 1974.
Tanzania.

New Genera and New Combinations of
Eastern Hemisphere Vernonieae
Subtribe Gymnantheminae H. Rob.

Distephanus Cass., Bull. Soc. Philom. Paris
1817. 151. 1817. Type: Conyza populi-
folia Lam.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

The genus characterization, synonymy,
and many of the species are given by Rob-
inson & Kahn (1986), who stressed the yel-
low flowers and trinervate leaves as impor-
tant generic characteristics. The type spe-
cies has been redescribed and illustrated
particularly well by Hind (1996), and it is
suggested in Scott (1993) and Hind (1996)
that the number of species in the genus
should be nearer to 40 instead of the 26
transferred by Robinson & Kahn (1986).
During the present study only the following
single additional species is transferred.

Distephanus henryi (Dunn.) H. Rob.,
comb. nov. basionym: Vernonia henryi
Dunn., J. Linn. Soc. Bot. 35:500. 1903.
China.

Gymnanthemum Cass., Bull. Soc. Philom.
Paris 1817:10. 1817. Type: G. cupulare
Cass. = Baccharis senegalensis = Gym-
nanthemum coloratum (Willd.) H. Rob.
& BKahn

Bracheilema R. Br. ex Salt., Abyss. Ap-
pend. 65. 1814, nom. nud.

Decaneurum DC., Arch. Bot. (Paris) 2:
516. 1833, nom. superfl. Type: as in
Gymnanthemum.

Monosis DC. in Wight, Contrib. Bot. Ind.
5. 1834. Type: Monosis wightiana DC.
in Wight [= Vernonia arborea Buch.-
Ham. |].

Vernonia sect. Strobocalyx Blume ex
DC., Prodr. 5:21. 1836. Type: Vernon-
ia arborea Buch.-Ham.

Plectreca Rafin., Fl. Tellur. 4:119. 1838
(“*1836’’). Type: Staehelina corymbo-
sa Thunb.

Keringa Rafin., Sylva Tellur. 144. 1838.
Type: Vernonia amygdalina Del.

Cheliusia Sch. Bip., Flora 24, 1. Intell.
26. 1841. Type: Cheliusia abyssinica
Sch. Bip. = Gymnanthemum amyg-
dalinum (Del.) Sch. Bip. ex Walp.

Strobocalyx (Bl. ex DC.) Spach, Hist.
Nat. Veg. Phan. 10:39. 1841.

Punduana Steetz in Peters, Reise Mos-
samb. Bot. 345. 1864. Type: P. vol-
kameriifolia (DC.) Steetz

VOLUME 112, NUMBER 1

Vernonia subsect. Urceolata S. B. Jones,
Rhodora 83:67. 1981. Type: Vernonia
sphaerocalyx O. Hoffm.

Shrubs or trees, moderately to densely
branching; hairs often forming felt, with
large often contorted cap cells basally or
nearly basally attached. Leaves alternate,
with short or winged petioles to rather long-
petiolate. Inflorescences terminal on stems
and branches, densely corymbiform. Heads
with involucres campanulate to ovoid;
bracts appressed, coriaceous to subcoria-
ceous, 25—35 in 4—5 series, inner bracts per-
sistent or easily deciduous; receptacle epa-
leaceous. Florets 1—50; corollas white to vi-
olet; tube cylindrical, throat longer than the-
cae or very deeply cut, lobes with glands
or spicules; anther base broadly tailed, often
long; apical anther appendages glabrous,
with rather thick-walled cells; style with or
without node. Achenes 5—10-costate, with
or without setulae or uniseriate hairs, ra-
phids short to elongate or lacking; pappus
of many rather persistent capillary bristles,
often broad-tipped, with short outer squa-
mellae. Pollen grains tricolporate, non-lo-
phate or rarely lophate, echinate, micro-
punctate. Chromosome numbers N = 7?,
10, 20, 2N = 30 (Jones 1979, 1982). Ses-
quiterpene lactones include elemanolides
(Bohlmann & Jakupovic 1990).

A broad interpretation of the Gymnanthe-
mum is accepted here. The type species lacks
a basal node on the style, but other closely
related species have such nodes. The number
of ribs on the achene varies from 5 to 10
within many subgroups. At least some of the
species with persistent inner involucral bracts
seem very closely related to species in which
they are deciduous. The leaves also seem to
vary in size, texture and margin. The corolla
throat is usually long, but a number of species
in Madagascar have sinuses cut nearly to the
base of the throat. The corolla lobes usually
show a characteristic long-triangular erect
form, but a few species have long narrow
lobes that are rolled back with age. The
sweeping hairs are usually blunt, but those of

239

G. amygdalinum are more pointed. The pol-
len is almost always non-lophate or Type A,
but rarely, as in the type of Punduana, Ver-
nonia volkameriaefolia, the pollen is lophate.
The genus is limited within the subtribe in
the present paper by only the most obvious
differences in inflorescence shape, involucral
bracts, leaf nervation, corolla symmetry and
color, and lack of pales on the receptacle.

The genus is credited here with the fol-
lowing 43 species:

Gymnanthemum amygdalinum (Del.) Sch.
Bip. ex Walp., Rep. 2:948. 1843.

Vernonia amygdalina Del., Cent. Pl. Afr.
Voy. Méroé 41. 1826.

Gymnanthemum abyssinicum Sch. Bip.
ex Walp., Rep. 2:948. 1843.

Vernonia vogeliana Benth. in Hook., Ni-
ger Fl. 427. 1849.

Vernonia condensata Baker., J. Bot. 8:
202. S75.

Vernonia eritreana Klatt,
Boiss. 4:826. 1896.

Vernonia randii S. Moore, J. Bot. 37:369.
1899.

Vernonia giorgii De Wild., Bull. Jard.
Bots Bruxs'5:92.11915.

Vernonia bahiensis Toledo, Arg. Bot. Es-
tado Sao Paulo, n.s. 1:52. 1939.

Vernonanthura condensata (Baker) H.
Rob., Phytologia 73:69. 1992.
Yemen and Ethiopia, South Uganda,
Kenya and Tanzania, Brazil, widely
cultivated.

Bull. Herb.

Gymnanthemum anceps (C. B. Clarke ex
Hook.f.) H. Rob., comb. nov. basionym:
Vernonia anceps C. B. Clarke ex Hook.
i, HA Brit, Ind. 53:233. 138k.

Sri Lanka.

Gymnanthemum andersonii (C. B. Clarke)
H. Rob., comb. nov. basionym: Vernonia
andersonii C. B. Clarke, Comp. Ind. 27.
1852.

Vernonia chevalieri Gagnap., Bull. Mus.
Hist. Nat. Paris 25:488. 1919, hom. il-
leg., not O. Hoffm., 1908.

Assam, Myanmar, China.

240

Gymnanthemum andrangovalense (Hum-
bert) H. Rob., comb. nov. basionym: Ver-
nonia andrangovalensis Humbert, Notul.
Syst. (Paris) 13:311. 1948.

Madagascar.

Gymnanthemum antanalus (Humbert) H.
Rob., comb. nov. basionym: Vernonia
antanala Humbert, Notul. Syst. (Paris)
13:314. 1948, in part., emend., Humbert,
Not. Syst. (Paris). 15:361. 1959.
Madagascar.

Gymnanthemum appendiculatum (Less.)
H. Rob., comb. nov. basionym: Vernonia
appendiculata Less., Linnaea 6:636.
1831.

Madagascar.

Gymnanthemum arboreum (Buch.-Ham.)
H. Rob., comb. nov. basionym: Vernonia
arborea Buch.-Ham., Trans. Linn. Soc.
London 14:218. 1824.

Monosis wightiana DC. in Wight, Con-
trib. Bot. Ind. 5. 1834, not Vernonia
wightiana Arn.

Vernonia celebrica DC., Prodr. 5:21.

1836.

Vernonia javanica DC., Prodr. 5:22.
1836.

Vernonia blumeana DC., Prodr. 5:22.
1836.

Strobocalyx arborea (Buch.-Ham.) Sch.
Bip., Jahres. Pollichia 18—19:171.
1861.

Vernonia monosis Benth. ex C. B.
Clarke, Comp. Ind. 24. 1852, nom.
nov. for Monosis wightiana DC. in
Wight, 1834.

Vernonia vaniotii Lévl., Repert. Spec.
Nov. Regni Veg. 12:531. 1913.

SE. Asia, Indonesia.
Gymnanthemum baronii (Baker) H. Rob.,

comb. nov. basionym: Vernonia baronii
Baker, J. Linn. Soc. Bot. 20:173. 1885.

Vernonia campenoni Drake, Bull. Soc.
Bot. France 46:241. 1899.
Madagascar.

Gymnanthemum bellinghamii (S. Moore)

H. Rob., comb. nov. basionym: Vernonia

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

bellinghamii S. Moore, J. Bot. 38:155.
May 1900.

Vernonia goetzeana O. Hoffm., Bot.
Jahrb. Syst. 28:503. July 1900.
Vernonia elizabethvilleana De Wild., Re-
pert. Spec. Nov. Regni Veg. 13:20.
1941.
Congo and Tanzania south to Moz-
ambique.

Gymnanthemum bockianum (Diels) H.
Rob., comb. nov. basionym: Vernonia
bockiana Diels, Bot. Jahrb. Syst. 29:608.
1900.

Pluchea rubicunda Schneid. in Sarg., pl.
Wis. 3:418. 1916. China.

Gymnanthemum chapelieri (Drake) H.
Rob., comb. nov. basionym: Vernonia
chapelieri Drake, Bull. Soc. Bot. France
46:234. 1899.

Madagascar.

Gymnanthemum coloratum (Willd.) H.
Rob. & B. Kahn, Proc. Biol. Soc. Wash.
99:501. 1986.

Eupatorium coloratum Willd., Sp. Pl. 3:
1768. 1803.

Baccharis senegalensis Pers., Syn. Pl. 2:
424. 1807.

Gymnanthemum cupulare Cass., Dict. Sc.
Nat., ed. 2, 20: 109. 1821.

Vernonia senegalensis (Pers.) Less., Lin-
naea 4:265. 1829.

Decaneurum grande DC., Prodr. 5:67.
1836.

Decaneurum senegalense (Pers.) DC.,
Prodr. 5:68. 1836.

Gymnanthemum grande (DC.) Sch. Bip.
ex Walp., Rep. 2:948. 1843.

Gymnanthemum senegalense (Pers.) Sch.
Bip. ex Walp., Rep. 2:948. 1843.

Gymnanthemum quercifolium Steetz in
Peters, Reise Mossamb. Bot. 334.
1864.

Vernonia oxyura O. Hoffm., Engl. Pflan-
zenw. Ost.-Afr. C. 403. 1895.

Vernonia polyura O. Hoffm., Bot. Jahrb.
Syst. 30:422. 1901.

VOLUME 112, NUMBER 1

Vernonia cirrhifera S. Moore, J. Linn.
Soc. Bot. 35:320. 1902.

Vernonia longipetiolata Muschl., Bot.
Jahrb. Syst. 46:74. 1911.

Vernonia aldabrensis Hemsl., J. Bot. 54,
suppl. 2:20. 1916.

Vernonia grandis (DC.) Humb., Fl. Mad-
ag. 189:44. 1960.
West tropical Africa east to Kenya,
south to Mozambique and Madagascar.

Gymnanthemum corymbosum (Thunb.) H.
Rob., comb. nov. basionym: Staehelina
corymbosa Thunb., Prodr. pl. Cap. 2:143.
1800.

Vernonia corymbosa (Thunb.) Less., Lin-
naea 6:647. 1831.

Plectreca corymbosa (Thunb.) Rafin., FI.
Tellur. 4:119. 1838 (“11836’’).
South Africa.

Gymnanthemum coursii (Humbert) H.
Rob., comb. nov. basionym: Vernonia
coursii Humbert, Notul. Syst. (Paris) 13:
310. 1948.

Madagascar.

Gymnanthemum crataegifolium (Hutch.)
H. Rob., comb. nov. basionym: Vernonia
crataegifolia Hutch., Bull. Misc. Inf.
Kew 7:330. 1912.

South Africa.

Gymnanthemum cumingianum (Benth. in
Hook.f.) H. Rob., comb. nov. basionym:
Vernonia cumingiana Benth., Hooker’s J.
Bot. Kew Gard. Misc. 4:232. 1852.
China, Philippines.

Gymnantheum cylindriceps (C. B. Clarke)
H. Rob., comb. nov. basionym: Vernonia
cylindriceps C. B. Clarke, J. Linn. Soc.
Bot. 25:35. 1880.

India, Nepal.

Gymnanthemum dissolutum (Baker) H.

Rob., comb. nov. basionym: Vernonia

dissoluta Baker, J. Linn. Soc. Bot. 20:
174. 1883.

Vernonia sparsiflora Baker, J. Linn. Soc.
Bot. 20:172. 1883.
Vernonia capreaefolia Baker, J. Linn.

241

Soc. Bot. 22:487. 1887.
Madagascar.

Gymnanthemum esculentum (Hemsl. ex FE
B. Forbes & Hemsl.) H. Rob., comb. nov.
basionym: Vernonia esculenta Hemsl. ex
E B. Forbes & Hemsl., J. Linn. Soc. Bot.
23:401. 1888.

Vernonia papillosa Franch., J. Bot. 10:
368. 1896.

Vernonia arbor Lévl., Repert. Spec. Nov.
Regni Veg. 11:304. 1912.
China.

Gymnanthemum exsertiflorum (Baker) H.
Rob., comb. nov. basionym: Vernonia
exsertiflora Baker, Bull. Misc. Inf. Kew
1898:147. July 1898.

Vernonia kreismannii Welw. ex Hiern,
Cat. Afrm PL Welw: 4:51. “Dec
1898.

Vernonia sphaerocalyx O. Hoffm., Bot.
Jahrb. Syst. 30:423. 1901.

Angola and Congo to Malawi and Tan-
zania.

Gymnanthemum exsertum (Baker) H.
Rob., comb. nov. basionvym: Vernonia
exserta Baker, J. Linn. Soc. Bot. 22:488.
1887.

Vernonia grisea Baker, J. Linn. Soc. Bot.
22:488. 1887, hom. illeg., not Baker,
1873.

Vernonia trichodesma Baker, J. Linn.
Soc. Bot. 25:325. 1890.
Vernonia lantziana Drake,
Bot. France 46:235. 1899.

Madagascar.

Bulk’ Soc.

Gymnanthemum glaberrimum (Welw. ex
O. Hoffm.) H. Rob., comb. nov. basio-
nym: Vernonia glaberrima Welw. ex O.
Hoffm., Bol. Soc. Brot. 13:15. Sept 1896.

Vernonia hensii Klatt, Bull. Herb. Boiss.
4:828. Dec. 1896.

Vernonia mashonica N. E. Br.,
Misc. Inf. Kew 1906:108. 1906.
West, central and south tropical Africa
north to Sudan.

Gymnanthemum hildebrandtii (Vatke) H.

Bull.

242

Rob., comb. nov. basionym: Vernonia
hildebrandtii Vatke, Osterr. Bot. Zeit. 25:
S23 LEIS:

Vernonia taylorii S. Moore, J. Bot. 38:
154. 1900.
Kenya, Somalia, Tanzania.

Gymnanthemum humblotii (Drake) H.
Rob., comb. nov. basionym: Vernonia
humblotii Drake, Bull. Soc. Bot. France
46:235. 1899.

Vernonia beforonensis Humbert, Notul.
Syst. (Paris) 13:320. 1948.
Madagascar.

Gymnanthemum louvelii (Humbert) H.
Rob., comb. nov. basionym: Vernonia
louvelii Humbert, Notul. Syst. (Paris) 15:
247. 1958.

Madagascar.

Gymnanthemum mespilifolium (Less.) H.
Rob., comb. nov. basionym: Vernonia
mespilifolia Less., Linnaea 6:641. 1831.
South Africa.

Gymnanthemum myrianthum (Hook.f.) H.
Rob., comb. nov. basionym: Vernonia
myriantha Hook.f., J. Linn. Soc. Bot. 7:
198. 1864.

Vernonia podocoma Sch. Bip. ex Vatke,
Linnaea 39:476. 1875.
Vernonia subuligera O. Hoffm., Engl.,
Pflanzenw. Ost.-Afr. C. 403. 1895.
Vernonia stipulacea Klatt, Bull. Herb.
Boiss. 4:457. 1896.

Vernonia lujae De Wild., Pl. Nov. Herb.
Hort. Then, 2:019st. 961.2900:

Vernonia ampla O. Hoffm., Bot. Jahrb.
Syst. 30:423. 1901.

Vernonia myrianthoides Muschl., Bot.
Jahrb. Syst. 46:84. 1911.

Vernonia uhligii Muschl., Bot. Jahrb.
Syst..46: 79.191

Vernonia oliveriana Pichi-Serm., Webbia
7:345. 1950, nom? itlesit superi. for
V. podocoma Sch. Bip. ex Vatke.

Vernonia chlarugii Pich-Serm., Miss.
Stud. Lago Tana 7, Ricerche Bot. 1:
195, 1 30/1951:
West Africa from Guinea and Sierra

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Leone to Cameroon, Congo, Sudan,
Ethiopia, Kenya, Tanzania, Uganda to
south tropical Africa.
Gymnanthemum obovatum Gaudich., Voy.
Uran. Bot. 471. 1830.

Vernonia cuneata Less., Linnaea 6:644.
1831.

Vernonia vagans DC., Prodr. 5:32. 1836.

Vernonia scandens DC., Prodr. 5:32.
1836.

Decaneurum obovatum (Gaudich.) DC.,
Prodr. 5:67. 1836.

Gymnanthemum scandens (DC.) Steetz in
Peters, Reise Mossamb. Bot. 338.
1864.

Gymnanthemum vagans (DC.) Steetz in
Peters, Reise Mossamb. Bot. 338.
1864.

Gymnanthemum affine Steetz in Peters,
Reise Mossamb. Bot. 338. 1864.

Vernonia obovata (Gaudich.) Boerl.,
Handl. Fl. Ned. Indié 2:1. 1891, not
Vernonia obovata Less., 1829.

India, Myanmar, Indonesia.
Gymnanthemum pectiniforme (DC. in

Wight) H. Rob., comb. nov. basionym:

Vernonia pectiniformis DC. in Wight,

Contr. Bot. Ind. 2. 1834.

Vernonia puncticulata DC., 7:264. 1838.
Lysistemma pectiniforme (DC. in Wight)
Steetz in Peters, Reise Mossamb. Bot.
343. 1864.
India, Sri Lanka.

Gymnanthemum pectorale (Baker) H.
Rob., comb. nov. basionym: Vernonia
pectoralis Baker, J. Bot. 20:139. 1882.
Madagascar.

Gymnanthemum pleistanthum (Humbert)
H. Rob., comb. nov. basionym: Vernonia
secundifolia subsp. pleistantha Humbert,
Bull. Soc. Bot. France 94:380. 1947.
Madagascar.

Gymnanthemum rueppellii (Sch. Bip. ex
Walp.) H. Rob., comb. nov. basionym:
Vernonia rueppellii Sch. Bip. ex Walp.,
Rep. 2:946. 1864.

Vernonia francavillana Oliv. & Hiern in

VOLUME 112, NUMBER 1

Oliv., Fl Trop. Afr. 3:296. 1877.
Ethiopia.

Gymnanthemum secundifolium (Boj. ex
DC.) H. Rob., comb. nov. basionym: Ver-
nonia secundifolia Boj. ex DC., Prodr. 5:
Ze 1836:

Vernonia quadriflora Baker, J. Linn. Soc.
Bot. 20:173. 1884.

Madagascar.

Gymnanthemum solanifolium (Benth.) H.
Rob., comb. nov. basionym: Vernonia so-
lanifolia Benth., Hooker’s J. Bot. Kew
Gard. Misc. 1:486. 1842.

China, Myanmar, Thailand, Vietnam.

Gymnanthemum subcrassulescens (Hum-
bert) H. Rob., comb. nov. basionym: Ver-
nonia subcrassulescens Humbert, Notul.
Syst. (Paris) 13:309. 1948.

Madagascar.

Gymnanthemum theophrastifolium
(Schweinf. ex Oliv. & Hiern) H. Rob.,
comb. nov. basionym: Vernonia theo-
Phrastifolia Schweinf. ex Oliv. & Hiern,
in Oliv., Fl. Trop. Afr. 3:294. 1877.

Vernonia myriocephala A. Rich., Tent.
Fl. Abyss. 1:374. 1848, hom. illeg.,
not DC., 1836.

Cacalia richardiana O. Kuntze, Rev.
Gen. Pl. 2:969. 1891, nom. nov. for V.
myriocephala A. Rich.

Vernonia seretii De Wild., Ann. Mus.
@oneo VV. fT:207- 1907.

Vernonia macrophylla Chiov., Ann. Bot.
Roma 9:70. 1911.

Vernonia richardiana (QO. Kuntze) Pichi-
Serm., Webbia 7:340. 1950.

Congo and Nigeria east to Uganda,
Kenya and Ethiopia.

Gymnanthemum thomsonianum (Oliv. &
Hiern in Oliv.) H. Rob., comb. nov. bas-
ionym: Vernonia thomsoniana Oliv. &
Hiern in Oliv., Trans. Linn. Soc. London
29°91, 1873.

Vernonia livingstoniana Oliv. & Hiern in
Oliv., Fl. Trop. Afr. 3:295. 1877.

Vernonia cruda Klatt, Bull. Herb. Boiss.
4:456. 1896.

243

Vernonia densicapitulata De Wild., Bull.
Jard. Bot, Brux:'5:92:'1915:.
West, central and south tropical Africa.
Gymnanthemum urticifolium (A. Rich.) H.
Rob., comb. nov. basionym: Vernonia ur-
ticifolia A. Rich., Tent. Fl. Abyss. 1:378.
1848.

Vernonia antinoriana Avetta, Nouv.
Giorn. Bot. Ital. 21:348. 1889.
Vernonia mellifera Muschl., Bot. Jahrb.
Syst. 46:90. 1911.
Congo, Ethiopia, Kenya.
Gymnanthemum vidalii (G. Mertr.) H. Rob.,
comb. nov. basionym: Vernonia vidalii
G. Merr., Philipp. Is., Bur. Gov. Lab.
Bull. 6:6. 1904 (*1903’’).
Philippines.
Gymnanthemum volkameriifolium (DC.)
H. Rob., comb. nov. basionym: Vernonia

volkameriaefolia DC., Prodr. 5:32. 1836.

Vernonia acuminata DC., Prodr. 5:32.
ES35C-

Vernonia punduana DC., Prodr. 5:32.
1836.

Vernonia esgirolii Lévl., Repert.
Nov. Regni Veg. 11:304. 1912.
China, India, Myanmar.

Gymnanthemum wightianum (Arn.) H.

Rob., comb. nov. basionym: Vernonia

wightiana Arn., Pugill. pl. Ind. Or. 27.

1836.

Sri Lanka.

Gymnanthemum zanzibarense (Less.) H.
Rob., comb. nov. basionym: Vernonia
zanzibarensis Less., Linnaea 6:637.
LSS.

Kenya, Tanzania.

Spec.

Gymnanthemum zeylanicum (L.) H. Rob.,
comb. nov. basionym: Eupatorium zey-
lantenm 4 Sp. pl. 837.1753.

Vernonia zeylanica (L.) Less., Linnaea 4:
344. 1829.

Sri Lanka.

A number of additional species have
been placed in the genus that are exclud-
ed here. These include G. congestum
Cass., = Critoniopsis triflosculosa (H. B.

244

K.) H. Rob. (Robinson 1993), G. frutes-
cens (Bl.) Sch. Bip. ex Walp., G. molle
(DC.) Sch. Bip. ex Walp., G. phyllolaen-
um (DC.) Sch. Bip. ex Walp. and G. re-
ticulatum (Wight) Sch. Bip. ex Walp.
which belong to Phyllocephalum (Kirk-
man 1981).

Brenandendron H. Rob., gen. nov. (Gym-
nantheminae).

Type: Vernonia titanophylla Brenan

Plantae arborescentes ad 8 m altae me-
diocriter ramosae; caules velutini, pilis fu-
siformibus vel leniter asymmetricis. Folia
alterna petiolata vel subsessilia, laminis
magnis interdum lobatis. Inflorescentiae
frondiformes in ramulis racemosae vel spi-
ciformes. Capitula sessilia vel breviter pe-
dunculata; involucra campanulata; bracteae
involucri usque ad 110 in seriebus 7—9 ap-
pressae interiores leniter deciduae subcori-
aceae vel coriaceae; receptacula epaleacea.
Flores 30-50 in capitulo; corollae pler-
umque tubiformes in faucibus elongatae,
lobis erectis vel interdum recurvatis; thecae
antherarum base longe obtuse caudatae; ap-
pendices apicales in parietibus cellularum
firmae; basi stylorum annulate noduliferi.
Achenia 8—10-costata glabra idioblastifera,
raphidis rhomboideis breviter vel mediocri-
ter elongatis; setae pappi interiores subplan-
ae subpersistentes exteriores squamiformes
vel nullae. Grana pollinis tricolporata
echinata non lophata. Numerus chromoso-
matum N = 9 Mangenot & Mangenot
(1957):

Germacranolides have been reported
from the genus (Bohlmann and Jakupovic
1990);

Brenandendron is closely related to Gym-
nanthemum, but it differs by the distinctive
frondiform inflorescence. Brenan (1953) pro-
vided a key to the three species. In addition
to the differences given by Brenan, the three
species differ in their corolla lobes. Brenan-
dendron donianum has erect lobes much
shorter than the corolla throat, B. frondosum
has narrow erect lobes, and B. titanophyllum
has strongly recurved narrow lobes.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

The three species are as follows:

Brenandendron donianum (DC.) H. Rob.,
comb. nov. basionym: Vernonia doniana
DC., Prodr. 5:23. 1836.

Vernonia conferta Benth. in Hook., Niger
Fl. 427. 1849.

West and central tropical Africa from An-
gola to Liberia and Sudan.

Brenandendron frondosum (Oliv. & Hiern
in Oliv.) H. Rob., comb. nov. basionym:
Vernonia frondosa Oliv. & Hiern in
Oliv., Fl. Trop. Afr. 3:294. 1877.

West Africa from Guinea to Cameroon.

Brenandendron titanophyllum (Brenan) H.
Rob., comb. nov. basionym: Vernonia ti-
tanophylla Brenan, Kew Bull. 1953:116.
1953.

Cameroon, Congo, Equatorial Guinea.

Myanmaria H. Rob., gen. nov. (Gymnan-
theminae).

Type: Vernonia calycina DC.

Plantae frutescentes ca. 1 m altae medi-
ocriter ramosae; caules puberuli, pilis sim-
plicibus uniseriatis multiseptatis. Folia al-
terna breviter petiolata, laminis oblongo-
Ovatis apice obtusis. Inflorescentiae cor-
ymbiformae. Capitula in ramis terminalia
pedunculata; bracteae involucri exteriores
ca. 20 ca. 2—3-seriatae late ovatae virides
foliiformes; bracteae interiores lineares acu-
tae; receptacula epaleacea. Flores 35—50 in
capitulo; corollae puberulae, tubis elonga-
tis, faucibus quoad lobis longioribus, lobis
2—3-plo longioribus quam latioribus extus
glanduliferis et pauce breviter pilosulis; the-
cae antherarum base distincte late caudatae;
appendices apicales antherarum distincte
induratae; basi stylorum non _ noduliferi;
rami stylorum obtuse papilliferi. Achenia
10-costata idioblastifera in costis setulifera,
raphidis subquadratis; setae pappi barbel-
latae numerosae 2—3-seriatae persistentes
exteriores breviores vel nullae. Grana pol-
linia ca. 50 wm in diametro tricolporata
echinata lophata micropunctata, lacunis po-
laribus triplicibus intercolpatis.

Myanmaria is endemic to Myanmar. The
genus is easily distinguished by the ca. 20

VOLUME 112, NUMBER 1

large, broad, green outer involucral bracts
that completely conceal the inner bracts.
The species is often described as herba-
ceous, and it apparently commonly turns
black when dry.

The single known species is as follows:

Myanmaria calycina (DC.) H. Rob., comb.
nov. basionym: Vernonia calycina DC.,
Prodr. 5:60. 1836.

Myanmar.

Lampropappus (QO. Hoffm.) H. Rob., stat.
nov. (Gymnantheminae). Vernonia section
Lampropappus O. Hoffm., Bol. Soc. Bro-
ter. 13:14. 1896. Type: Vernonia lampro-
pappus O. Hoffm. Vernonia subsect. Tur-
binella S. B. Jones, Rhodora 83:67. 1981.
Type: Vernonia lampropappus O. Hoffm.

Subshrubs to 0.6 m high; stems costate,
tomentose, hairs with basally or slightly ex-
centrically attached contorted capcells.
Leaves alternate, shortly petiolate to more
or less amplexicaul, blades densely tomen-
tose abaxially. Inflorescence densely cor-
ymbose, with many heads. Heads hemi-
spherical to turbinate; involucral bracts ca.
30, ca. 4—5-seriate, strongly gradate, outer
bracts ovate, apices acute to shortly apicu-
late; receptacle epaleaceous. Flowers ca. 20
in a head; corollas white with purple tips to
pale purple, actinomorphic or slightly zy-
gomorphic, with lobe or sinus centered on
outer side, longest lobe centered toward in-
side. Bases of anther thecae rounded, with-
out tails; apical appendage with somewhat
thickened and ornamented cell walls; base
of style with broadened node; sweeping
hairs of style branches with obtuse tips.
Achenes 5-costate, densely villous with un-
iseriate hairs, with few septae near base,
with few idioblasts; raphids subquadrate to
short-oblong; pappus setae persistent, 3-se-
riate, broad and flat, densely scabrid on
margins smooth on surfaces, outer seg-
ments shorter and narrower. Pollen grains
tricolporate, non-lophate, echinate.

Lampropappus, as a section, was origi-
nally compared with the American Stilp-
nopappus Mart. ex DC. and the Eastern

245

Hemisphere Vernonia sect. Stengelia (=

Baccharoides Moench) by Hoffmann

(1896) because of the broad flat pappus seg-

ments, but was distinguished at least from

the Vernonia sect. Stengelia by the exap-
pendiculate involucral bracts. Lampropappus
does not have the subulate tips on the pappus
elements seen in Stilpnopappus. The African
genus is further distinguished by the uniser-
iate rather than biseriate hairs densely cov-
ering the achene and by the non-lophate pol-
len. One species of the present genus, L. tur-
binella, proves to be distinct in the slightly
zygomorphic corollas that are rotated or re-
supinate by 1/10 so that a sinus is centered
to the outside and a lobe centered to the in-
side. This is not seen in the other two species
of the genus. This feature occurs elsewhere
in the Vernonieae in the American genus

Mattfeldanthus H. Rob. & R. M. King

(1979), which also has zygomorphic corollas.

The latter genus is related to Lepidaploa

(Cass.) Cass. with lophate pollen and seriate-

cymose inflorescences. Such resupination of

the corolla also seems to occur in the Gor-
teriinae of the Arctoteae.

The following three species are placed in
the genus:

Lampropappus eremanthifolia (O. Hoffm.)
H. Rob., comb. nov. basionym: Vernonia
eremanthifolia O. Hoffm., Bol. Soc. Bro-
ter, 03215. 1396.

Angola.

Lampropappus hoffmannii H. Rob., nom.
nov. basionym: Vernonia lampropappus
O. Hoffm., Bol. Soc. Broter. 13:14. 1896.
Angola.

Lampropappus turbinellus (S. Moore) H.

Rob., comb. nov. basionym: Vernonia
turbinella S. Moore, J. Linn. Soc. Bot.
7 2 6G V9O25.
Congo, Malawi, Zambia. Zygomorphy of
the corollas is not mentioned in the orig-
inal description of the species (Moore
L925).

Acknowledgments

The present study has been aided by loans
of material kindly sent by The Royal Botanic

246

Garden, Kew; British Museum (Natural His-
tory); Jardin Botanique de 1’Etat, Bruxelles;
and Muséum National d’Histoire Naturelle,
Laboratoire de Phanérogamie, Paris. Aspects
of this study have been assisted by Margorie
Knowles of the Department of Botany,
Smithsonian Institution. John Pruski of the
Smithsonian Department of Botany and Mi-
chael O. Dillon of the Field Museum of Nat-
ural History are thanked for critical reviews
of the manuscript.

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Pope, G. V. 1983. Cypselas and trichomes as a source
of taxonomic characters in the Erlangeoid gen-
era.—Kirkia 12:203-231.

VOLUME 112, NUMBER 1

. 1986. Vernonia chloropappa (Compositae)
and related species in tropical Africa——Kew
Bulletin 41:393—397.

Robinson, H. 1989. Acilepidopsis, a new genus of Ver-

nonieae from South America (Asteraceae).—
Phytologia 67:289-—292.

. 1990a. Six new combinations in Baccharoides
Moench and Cyanthillium Blume (Vernonieae:
Asteraceae).—Proceedings of the Biological
Society of Washington 103:248-—252.

. 1990b. New combinations in the Asteraceae
(Vernonieae, Heliantheae, Mutisieae).—Phyto-
logia 69:105—107.

. 1992a. Mesanthophora, a new genus of Ver-
nonieae (Asteraceae) from Paraguay.—Novon
2:169-172.

. 1992b. The Asteraceae of the Guianas, III:
Vernonieae and restoration of the genus Xiph-
ochaeta.—Rhodora 94:348-—361.

. 1993. A review of the genus Critoniopsis in
Central and South America (Vernonieae: Aster-
aceae).—Proceedings of the Biological Society
of Washington 106:606—627.

. 1996. 33. The status of generic and subtribal
revisions in the Vernonieae. Pp. 511—526 in D.
J. N. Hind and H. J. Beentje, eds., Compositae
Systematics. Proceedings of the International
Compositae Conference, Kew, 1994. D. J. N.
Hind, editor in chief, Royal Botanical Garden,
Kew 1:i—xiv, 1-784.

, & B. Kahn. 1986. Trinervate leaves, yellow
flowers, tailed anthers, and pollen variation in

247

Distephanus Cassini (Vernonieae: Astera-
ceae).—Proceedings of the Biological Society
of Washington 99:493-501.

, & R. M. King. 1979. Mattfeldanthus muti-
sioides gen. et spec. nov. (Asteraceae: Vernon-
ieae) from Bahia, Brazil.—Willdenowia 9:9-—12.
, & C. Marticorena. 1986. A palynological
study of the Liabeae (Asteraceae).—Smithson-
ian Contributions to Botany 64:i—iv, 1—50.

, E Bohlmann, & R. M. King. 1980. Chemo-
systematic notes on the Asteraceae. III. Natural
subdivisions of the Vernonieae.—Phytologia
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Scott, A. J. 1993. Distephanus. in D. J. N. Hind, C.

Jeffrey & A. J. Scott. Compositae. Pp. 19-21
in J. Bosser et al., eds., Flore des Mascareignes.
fasc. 109. Composées: Sugar Industry Research
Institute (Mauritius), ORSTOM (Paris) and
Royal Botanic Gardens, Kew.

Verdcourt, B. 1956. Notes from the East African Her-

barium: IV.—Kew Bulletin 1956:445—454.

Wild, H. 1977. The Compositae of the Flora Zambes-

jaca area, 7—Vernonieae (excluding Vernonia
Schreb.).—Kirkia 10:339-—384.

. 1978. The Compositae of the Flora Zambes-
jaca area 8—Vernonieae (Vernonia).—Kirkia
si 12 7.

, & G. V. Pope. 1978a. Brachythrix: a new ge-
nus of the tribe Vernonieae (Compositae) from
South Central Africa.—Kirkia 11:25—30.

WOE . 1978b. The Compositae of the Flo-
ra Zambesiaca area, 9—Vernonieae (Brachythrix
and Hoehnelia).—Kirkia 11:133-—142.

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
112(1):249—250. 1999.

INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE

Applications published in the Bulletin of Zoological Nomenclature

The following Applications were published on 30 September 1998 in Vol. 55,
Part 3 of the Bulletin of Zoological Nomenclature. Comment or advice on any of
these applications is invited for publication in the Bulletin and should be sent to the
Executive Secretary (I.C.Z.N.), c/o The Natural History Museum, Cromwell Road,
London SW7 5BD, U.K. (e-mail: iczn@nhm.ac.uk).

Case No.

3050 Pachylops Fieber, 1858 (Insecta, Heteroptera): proposed designation of Cap-
sus chloropterus Kirschbaum, 1856 (currently Orthotylus virescens
(Douglas & Scott, 1865)) as the type species.

3087 HAydrobia Hartmann, 1821 and Cyclostoma acutum Draparnaud, 1805 (cur-
rently Hydrobia acuta; Mollusca, Gastropoda): proposed conserva-
tion by replacement of the lectotype of H. acuta with a neotype;
Ventrosia Radoman, 1977: proposed designation of Turbo ventrosus
Montagu, 1803 as the type species; and HYDROBIINA Mulsant,
1844 (Insecta, Coleoptera): proposed emendation of spelling to HY-
DROBIUSINA, so removing the homonymy with HYDROBITIDAE
Troschel, 1857 (Mollusca).

Scarus chrysopterus Bloch & Schneider, 1801 (currently Sparisoma chrysop-
terum; Osteichthyes, Perciformes): proposed conservation of the
specific name and designation as the type species of Sparisoma
Swainson, 1839.

Osphronemus deissneri Bleeker, 1859 (currently Parosphromenus deissneri;
Osteichthyes, Perciformes): proposed replacement of holotype by a
neotype.

Cacatua Vieillot, 1817 and CACATUINAE Gray, 1840 (Aves, Psittacifor-
mes): proposed conservation.

LORISIDAE Gray, 1821 and GALAGIDAE Gray, 1825 (Mammalia, Primates):
proposed conservation as the correct original spellings.

250 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Opinions published in the Bulletin of Zoological Nomenclature

The following Opinions were published on 30 September 1998 in Vol. 55, Part 3
of the Bulletin of Zoological Nomenclature. Copies of these Opinions can be ob-
tained free of charge from the Executive Secretary, I.C.Z.N., c/o The Natural History
Museum, Cromwell Road, London SW7 5BD, U.K. (e-mail: iczn@nhm.ac.uk).

Opinion No.

1902. Anomalina d’Orbigny, 1826 (Foraminiferida): Anomalina ariminensis
d’Orbigny in Fornasini, 1902 designated as the type species.

1903. Umbellula Cuvier, [1797] (Cnidaria, Anthozoa): conserved as the correct
original spelling, and corrections made to the entries relating to Um-
bellularia Lamarck, 1801 on the Official Lists and Indexes of Names
in Zoology.

Aporcelaimus Thorne & Swanger, 1936 (Nematoda): Dorylaimus superbus
de Man, 1880 designated as the type species.

S.D. Kaicher (1973-1992), Card Catalogue of World-Wide Shells: not sup-
pressed for nomenclatural purposes.

Euchroeus Latreille, 1809 (Insecta, Hymenoptera): conserved; Chrysis pur-
purata Fabricius, 1787 (currently Euchroeus purpuratus): specific
name conserved; and Chrysis gloriosa Fabricius, 1793: specific
name suppressed.

Nothosaurus Minster, 1834 (Reptilia, Sauropterygia): given precedence over
Conchiosaurus Meyer, [1833]

Hemidactylus garnotii Duméril & Bibron, 1836 (Reptilia, Squamata): spe-
cific name conserved.

Holotropis herminieri Duméril & Bibron, 1837 (currently Leiocephalus her-
minieri), Proctotretus bibronii T. Bell, 1842 (currently Liolaemus
bibronii) (Reptilia, Squamata): specific names conserved, and Lio-
laemus bellii Gray, 1845 placed on the Official List.

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CONTENTS

Validation of Eleutherodactylus crepitans Bokermann, 1965, notes on the types and type lo-
cality of Telatrema heterodactylum Miranda-Ribeiro, 1937, and description of a new species
of Eleutherodactylus from Mato Grosso, Brazil (Amphibia: Anura: Leptodactylidae)

W. Ronald Heyer and Anna M. Munoz
\;new genus and species of frog from Bahia, Brazil (Amphibia: Anura: Leptodactylidae) with
“comments on the zoogeography of the Brazilian campos rupestres W. Ronald Heyer

: A mew species of cardinalfish (Perciformes: Apogonidae) from the Bay of Bengal, Indian Ocean

Thomas H. Fraser

er new species of deep-water skate, Rajella eisenhardti, (Chondrichthyes: Rajidae) from the

Galapagos Islands Douglas J. Long and John E. McCosker

; : A new genus of the subfamily Cubacubaninae (Insecta: Zygentoma: Nicoletiidae) from a Mex-

ican cave Luis Espinasa
Two new species of the genus Anelpistina (Insecta: Zygentoma: Nicoletiidae) from Mexican
caves, with redescription of the genus Luis Espinasa
Anacroneuria from northeastern South America (Insecta: Plecoptera: Perlidae) Bull P. Stark
A new species of Siamosquilla from Indonesia (Crustacea: Stomatopoda: Protosquillidae)
Mark V. Erdmann and Raymond B. Manning
The taxonomic status and zoogeography of Cambarus bartonii carinirostris Hay, 1914 (Crus-
tacea: Decapoda: Cambaridae) Roger F. Thoma and Raymond F. Jezerinac
Two new species of Aegla Leach (Crustacea: Decapoda: Anomura: Aeglidae) from southern
Chile Carlos G. Jara and Victor L. Palacios
The Hawaiian parthenopid crabs of the genera Garthambrus Ng, 1966, and Dairoides Stebbing,
1920 (Crustacea: Decapoda: Brachyura) Peter K. L. Ng and SSH Wan
Reinstatement and further description of Eualus subtilis Carvacho & Olson, and comparison
with E. lineatus Wicksten & Butler (Crustacea: Decapoda: Hippolytidae)
Gregory C. Jensen and Rachel C. Johnson
Deilocerus captabilis, a new species of cyclodorippid crab from southeastern Brazil (Crustacea:
Decapoda: Brachyura: Cyclodorippidae) Marcos Tavares
The Albuneidae (Decapoda: Anomura: Hippoidea) of the Hawaiian Islands, with description
of a new species Christopher B. Boyko
Two new species of Hansenium (Crustacea: Isopoda: Asellota) from Madang, Papua New
Guinea Kathrin S. Bolstad and Brian Kensley
Caecidotea simulator, a new subterranean isopod from the Ozark Springfield Plain (Crustacea:
Isopoda: Asellidae) Julian J. Lewis
Collocherides brychius, a new species (Copepoda: Siphonostomatoida: Asterocheridae) from a
deep-water hydrothermal site in the northeastern Pacific Arthur G. Humes
Cymbasoma californiense, a new monstrilloid (Crustacea: Copepoda: Monstrilloida) from Baja
California, Mexico E. Sudrez-Morales and R. Palomares-Garcia
Early development of the deep-sea ampharetid (Polychaeta: Puaphateudae) Decemunciger apa-
lea Zottoli Robert Zottoli
A new species of Spiochaetopterus (Chaetopteridae: Polychaeta) from a cold-seep site off
Hatsushima in Sagami Bay, central Japan
Eijiroh Nishi, Tomoyuki Miura, and Michel Bhaud
Two new subtribes, Stokesiinae and Pacourininae, of the Vernonieae (Asteraceae)
Harold Robinson
Revisions in paleotropical Vernonieae (Asteraceae) Harold Robinson
International Commission on Zoological Nomenclature
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
112(2):253-—302. 1999.

A revision of the Antarctic genus Chlanidota
(Gastropoda: Neogastropoda: Buccinulidae)

M. G. Harasewych and Yuri I. Kantor

(MGH) Department of Invertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560-0118, U.S.A.;
(YIK) Severtzov Institute, Russian Academy of Sciences, Leninski prospect 33,
Moscow 117071, Russia

Abstract.—The genus Chlanidota is revised to contain two subgenera,
Chlanidota sensu stricto and Chlanidota (Pfefferia). Chlanidota s.s. has a broad
distribution throughout the Antarctic region, and contains five species: Chlan-
idota (Chlanidota) vestita (Martens, 1881), C. (C.) densesculpta (Martens,
1885), C. (C.) paucispiralis Powell, 1951, C. (C.) pilosa Powell, 1951, and C.
(C.) signeyana Powell, 1951. The subgenus Pfefferia, which differs from
Chlanidota primarily in the morphology of its operculum, is endemic to South
Georgia Island, and is known from three species: Chlanidota (Pfefferia) chor-
data (Strebel, 1908), C. (P.) palliata (Strebel, 1908), C. (P.) invenusta, new
species. With its bulliform shell lacking a siphonal canal, the monotypic, cir-
cum-Antarctic genus Neobuccinum is conjectured to be the sister taxon of
Chlanidota, but is readily distinguished by its smoother, higher-spired shell,
radula in which the central cusp of the lateral teeth is medially placed, and a

penis lacking a papilla.

The genus Chlanidota Martens, 1878, is
one of the more diverse and wide-ranging
members of the antiboreal buccinoidean ra-
diation comprising the family Buccinulidae.
As is true of nearly all buccinoidean higher
taxa, this genus has been defined primarily
on the basis of shell morphology, occasion-
ally supplemented with observations of the
radula and operculum. Consequently, nei-
ther the limits of this genus nor its relation-
ships to other buccinoidean taxa have been
well established.

Martens (1878) first proposed Chlanidota
as a monotypic subgenus of Cominella, but
later (Martens 1881) transferred the subge-
nus to Buccinum Linné, 1758. Watson
(1886) included the type species of Chlan-
idota in the genus Neobuccinum Smith,
1877. Thiele (1912) erected the monotypic
Ficulina as a subgenus of Cominella. Later,
he discovered this taxon to be a junior syn-

onym of Ficulina Gray, 1867, and proposed
Notoficula Thiele, 1917, as a new name.
Still later, Thiele (1929:315) treated Chlan-
idota as a genus, and included Notoficula,
Pfefferia Strebel, 1908, and Bathydomus
Thiele, 1912, as subgenera. Powell (1951)
elevated each of these subgenera to generic
Status, but regarded them to be closely re-
lated. Oliver (1983) showed Notoficula to
be a neotenous eratoid rather than a bucci-
noidean.

Chlanidota has been treated most recent-
ly by Dell (1990), who included 13 species
(Table 1), some tentatively, and noted that
this genus has undergone an explosive ra-
diation in Antarctic waters. He commented
that the distributions, both geographic and
bathymetric, ““show puzzling patterns, es-
pecially through the Scotia Arc and adja-
cent regions of the Antarctic continent.”
The closely related taxon Pfefferia (see Ta-

254

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Listing of species attributed to Chlanidota and Pfefferia and their present taxonomic placement.
Taxa listed in bold are valid species referable to these subgenera of Chlanidota.

Chlanidota bisculpta Dell, 1990

Chlanidota elongata (Lamy, 1910)

Not a Chlanidota. Belongs in an as yet undescribed genus.
Chlanidota (Chlanidota) densesculpta (Martens, 1885)

Junior homonym of Cominella elongata Dunker, 1857; Chlani-

dota lamyi Dell, 1990 is a replacement name.

Chlanidota eltanini Dell, 1990
Chlanidota gaini Lamy, 1910
Chlanidota lamyi Dell, 1990
Chlanidota cf. lamyi Dell, 1990

Chlanidota (Chlanidota) paucispiralis Powell, 1951
Chlanidota (Chlanidota) pilosa Powell, 1951

Chlanidota polysperia Dell, 1990
Chlanidota pyriformis Dell, 1990
Chlanidota smithi Powell, 1958

Chlanidota (Chlanidota) signeyana Powell, 1951
Chlanidota (Chlanidota) vestita (Martens, 1881)
Chlanidota (Pfefferia) chordata (Strebel, 1908)

Pfefferia cingulata Strebel, 1908

Pfefferia elata Strebel, 1908
Chlanidota (Pfefferia) palliata (Strebel, 1908)
Chlanidota (Pfefferia) invenusta, new species.

ble 1), endemic to South Georgia Island,
was last reviewed by Powell (1951), who
selected a type species and added several
new records for two of the four species de-
scribed by Strebel (1908).

The present revision of the genus Chlan-
idota, which includes the subgenus Pfeffer-
ia, is based on a study of the large collec-
tions of these taxa sampled under the aus-
pices of the United States Antarctic Pro-
gram (USAP), as well as the type material
of all species described or subsequently as-
signed to these taxa. The objectives of this
study are: to assess the relationships among
nominal taxa of Chlanidota and Pfefferia
based on anatomical as well as concholog-
ical characters; to discern biogeographic
patterns in the distribution of these taxa;
and to begin to build a framework of ana-
tomical data that can be used to resolve
phylogenetic relationships of and among
the buccinoidean taxa of the Southern
Hemisphere.

Not a Chlanidota. Belongs in an as yet undescribed genus.
Belongs in the conoidean genus Belaturricula.

Synonym of Chlanidota signeyana Powell, 1951

Synonym of Chlanidota signeyana Powell, 1951

Not a Chlanidota. Belongs in an as yet undescribed genus.
Synonym of Chlanidota signeyana Powell, 1951
Synonym of Neobuccinum eatoni (Smith, 1875)

Synonym of Chlanidota (Pfefferia) palliata (Strebel, 1908)
Synonym of Chlanidota (Pfefferia) palliata (Strebel, 1908)

Materials and Methods

This report is based primarily on the ex-
tensive collections of Chlanidota and Pfef-
feria (including dry shells and alcohol pre-
served anatomical material) collected by the
United States Antarctic Program (USAP)
and housed at the National Museum of
Natural History, Smithsonian Institution
(USNM). Additional material, including
type specimens housed in the following in-
stitutions, were examined: BMNH—The
Natural History Museum, London; MNHN—
Muséum national d’ Histoire naturelle, Par-
is; MNH-U—Museum fiir Naturkunde der
Humboldt-Universitat zu Berlin; SAM—
The South Australian Museum, Adelaide;
SMNH—Swedish Museum of Natural His-
tory, Stockholm; ZMH—Zoologisches In-
stitut und Zoologisches Museum der Uni-
versitat Hamburg.

In the material examined sections, “‘spec-
imen’’ denotes alcohol preserved material,

VOLUME 112, NUMBER 2

239

Fie. 1:
operculum length; PE, posterior edge; SL, shell length; SW, shell width; 6, deflection of aperture from shell
axis.

while “‘shell’’ refers to records containing
only the empty shell.

Shell and operculum measurements were
obtained for representative specimens of
each species (n = 10, when available), as
detailed in Figure 1. Shell ultrastructure
was observed along freshly fractured sur-
faces parallel to the growing edge of the
shell. Anatomical descriptions are based on
gross dissections of preserved material. As
the apex of the shell was strongly eroded in
all specimens of Chlanidota, the aperture

Measurements of shell and operculum features. AL, aperture length; BWL, body whorl length; OL,

length (AL) rather than the shell length was
used as the reference when reporting the
relative size of anatomical features. Radulae
were removed by dissection, cleaned using
diluted bleach (NaClO), coated with carbon
and gold, and examined using a Hitachi
S570 Scanning Electron Microscope
(SEM).

Images were digitized using a Leaf Lu-
mina Digital Scanning Camera. Optical and
SEM images were processed using Pho-
toshop Version 4.01 (Adobe).

256

Systematics

Order Neogastropoda Wenz, 1938
Superfamily Buccinoidea
Rafinesque, 1815

The buccinoidean whelks comprise one
of the few uncontroversial, monophyletic
groups within the order Neogastropoda.
While buccinoideans are readily identified
on the basis of shell, radular, and alimentary
system morphology (e.g., weakly sculp-
tured shells lacking spines or columellar
folds, lateral teeth with =2 cusps, absence
of accessory salivary glands and rectal
gland), there is no consensus as to the rank
or inter-relationships of the group as a
whole, or its constituent higher taxa (e.g.,
Thiele 1929; Wenz 1938; Bouchet & Warén
1985; 1986; Ponder & Warén 1988; Ponder
& Lindberg 1996; Kantor 1996). As a re-
sult, most current classifications are of a ty-
pological rather than phylogenetic nature.

Based primarily on shell, radular, and
opercular characters, Powell (1929) divided
the Buccinoidea into the families Buccini-
dae, Neptuneidae, Buccinulidae, Cominel-
lidae and Fasciolariidae. In the same year,
Thiele (1929) published an alternative and
more widely accepted higher classification
of the Buccinoidea, comprising the families
Columbellidae, Buccinidae, Melongenidae,
Nassariidae, and Fasciolariidae. Powell
(1951) subsequently revised his classifica-
tion, regarding the Southern Hemisphere
Buccinulidae (now including the subfamily
Cominellinae) to be more closely related to
the boreal family Neptuneidae than to Buc-
cinidae. Ponder (1971) reviewed the New
Zealand species of Buccinulum and con-
cluded that anatomical differences did not
justify their segregation from Buccinidae.
Ponder (1974) and subsequently Ponder &
Warén (1988) reduced the Buccinoidea to a
family within Muricoidea, and also reduced
in rank or synonymized most of the previ-
ously recognized families and subfamilies.
Powell (1976) partially incorporated these
demotions in taxonomic rank into revisions
of his earlier work (e.g., Powell 1961), but

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(Powell 1979:192) clearly continued to re-
gard the Southern Hemisphere Buccinuli-
nae to be distantly related to the Northern
Hemisphere Buccininae. Other authors have
revised the higher systematics of the buc-
cinoidean fauna of the Northern Hemi-
sphere (Habe & Sato 1973) without treating
austral representatives.

Pending a phylogenetic revision of the
higher taxa within Buccinoidea, we provi-
sionally retain the taxon Buccinulidae and
its subdivisions, as understood by Powell
(1951), to encompass the austral radiation
of buccinoidean taxa.

Family Buccinulidae Finlay, 1928

Finlay (1928:250) proposed the family
Buccinulidae (with subfamilies Buccinuli-
nae and Siphonaliinae) and enumerated the
included genera, but provided no diagnosis
or differentiating characters. Powell (1929:
57) suggested that Finlay’s work was based
on protoconch and teleoconch characters,
and provided a modified classification
‘““‘based primarily on the dentition.’’ Powell
(1951:151) further refined his concept of
Buccinulidae, noting that the chief charac-
teristics of the group were tricuspid rachi-
dian teeth and an operculum with a terminal
or subterminal nucleus. He subdivided the
family into three subfamilies, the Cominel-
linae, Buccinulinae and Prosiphiinae based
on the number of cusps on the lateral teeth.

Subfamily Buccinulinae Finlay, 1928

As refined by Powell (1951:151) the
Buccinulinae are characterized by radulae
with tricuspid rachidian teeth and tricuspid
lateral teeth. The subfamily ranges from
Antarctica to New Zealand, Australia, and
along the eastern Pacific coast as far north
as California.

Genus Chlanidota Martens, 1878

Powell speculated that Chlanidota, Pfef-
feria and Neobuccinum may represent a
transitional stage between the short-ca-

VOLUME 112, NUMBER 2

nalled Cominellinae and the long-canalled
Buccinulinae, the latter including the Ant-
arctic genera Probuccinum, Cavineptunea,
and Bathydomus. He included Chlanidota,
Pfefferia, and Neobuccinum within Buccin-
ulinae primarily on the basis of their tricus-
pid lateral teeth, but noted that in these taxa
the intermediate cusp was likely the result
of a bifurcation of the inner cusp.

Subgenus Chlanidota Martens, 1878

Cominella (Chlanidota) Martens, 1878:
[Type (by monotypy): Cominella (Chlan-
idota) vestita Martens, 1878].

Chlanidota Thiele, 1929:314—315; Powell,
P9512 139: Dell, 1990:177.

Description—Shell medium-sized for
family, reaching 25.5 to 42.6 mm, depend-
ing on species. Shell usually thin, fragile,
ovate to elongate in outline, spire short to
very short. Shell covered with thin to thick,
smooth or hirsute periostracum. Spiral
sculpture of either thin to very thin threads
or of prominent cords. Aperture high, oval.
Operculum very small (0.18—0.37 AL),
ovate, coiled, with nucleus terminal, rotated
to left, nearly perpendicular to long axis of
operculum. Siphonal notch broad, dorsally
recurved, siphonal fasciole with ridges
along margins. Shell composed of 3 crys-
talline layers, outermost layer prismatic,
middle layer of crossed-lamellar crystals
with crystal faces comarginal (oriented par-
allel to growing edge), innermost layer of
crossed lamellar crystals radial (oriented
perpendicular to growing edge).

Digestive system generally typical of
buccinids. Proboscis of medium length or
long. Radular ribbon long, triserial. Rachi-
dian tooth with arched base, straight sides,
3 large, robust cusps, usually of equal
length. Lateral teeth with 3 cusps, outer
cusp longest, middle cusp shortest, closely
adjacent to innermost cusp. Salivary glands
small, fused or separate. Valve of Leiblein
large, well defined, lacking ciliary cone.
Gland of Leiblein long, tubular, convoluted
anteriorly, straightening and tapering pos-

257

teriorly. Oesophagus wide, muscular, with
posterior crop-like enlargement lined with
tall longitudinal folds. Stomach broadly U-
shaped, without caecum.

Penis long, dorsoventrally flattened, with
long, cylindrical terminal papilla, surround-
ed by deep circular fold at its base.

Remarks.—This subgenus is endemic to
the Antarctic region. The highest diversity
is in the Weddell Quadrant, especially in the
vicinity of South Georgia Island. There are
no records from the Ross Quadrant. Only
Chlanidota vestita, the type species, ex-
tends northward beyond the Antarctic Con-
vergence. The bathymetric range of Chlan-
idota sensu stricto is from 3 to 1116 m.

Chlanidota (Chlanidota) vestita
(Martens, 1878)
Figs. 2—3; Table 2

Cominella (Chlanidota) vestita Martens,
1878:23; Martens, 1904:63; Thiele, 1904:
168, Taf. IX, fig. 56 (radula); Lamy,
1O Lip he 3:

Buccinum (Chlanidota) vestitum Martens,
1881:43—44, Taf. 9, Fig. 3a—c.

Chlanidota vestita Tryon, 1881:201, pl. 79,
fe: 591: Smith, 1902-203; Powell, 1951:
1392 (Carcelles, 1953-191. pk 3. tig. 58;
Powell, 1957:133; Powell, 1960:150;
Dell, 1964:288; Gaillard, 1971:100; Can-
tera & Arnaud, 1984:68; Dell, 1990:184,
fies S11.

Neobuccinum vestitum Watson, 1886:216.

Description.—Shell (Fig. 2) small for ge-
nus (to 29 mm), thin, translucent, but
strong, ovate in outline, with low, rounded
spire. Protoconch unknown, upper teleo-
conch whorls heavily eroded. Teleoconch of
up to 3% evenly-rounded, convex whorls,
deeply eroded where periostracum damaged
or missing. Suture strongly impressed,
forming extremely narrow channel between
adapical spiral cord and previous whorl.
Spiral sculpture of prominent but unequal
cords, intervening spaces 2—4 times as
broad as cords. Lectotype with 23 cords on
body whorl, 8 on penultimate whorl. Para-

258

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Big.
D-F Paralectotype, MNH-U 25613-b. Both from Kerguelen Island. Scale bar = 1 cm.

lectotype with 22 cords on body whorl, 7
on penultimate whorl. Numerous fine, sin-
uated spiral threads between cords. Axial
sculpture limited to growth lines, producing
finely cancellate surface sculpture. Aperture
broadly ovate, deflected from shell axis by
14-18°. Outer lip thin, evenly rounded,
simple. Columella ~% AL, weakly con-
cave, with strong siphonal fold. Parietal re-
gion, siphonal fasciole overlain by thin,
grayish callus. Siphonal notch broad, dor-
sally recurved, with straight columellar,
rounded apertural margins forming borders
of fasciole. Ridge margin of fasciole runs
from apertural margin of siphonal notch.
Shell color chalky white, aperture weakly
glazed. Periostracum thin, straw-colored,

Chlanidota (Chlanidota) vestita (Martens, 1878). A-C. Lectotype, MNH-U 25613-a (here designated).

smooth, glossy. Operculum, as illustrated
by Dell (1990:fig. 311), very small (0.28
AL), ovate. Radula (Thiele 1904:fig. 56)
with tricuspid rachidian and lateral teeth, as
described for other species of Chlanidota
illustrated herein.

Type locality.—Kerguelen Island.

Type material.—Two lots of syntypes,
MNH-U 25613 (2 shells), MNH-U 25626
(3 shells). One specimen from MNH-U
25613, corresponding in size to the original
description, is here designated as the lec-
totype (Fig. 2A—C). The remaining four
Specimens are paralectotypes.

Material examined.—The lectotype
(MNH-U 25613-a) and one paralectotype
(MNH-U 25613-b).

VOLUME 112, NUMBER 2

Pig: 3.
= type locality, A = published records, and Chlanidota (Chlanidota) pilosa (Powell, 1951). * = type locality.
A. Bathymetric range of C. (C.) vestita (Martens, 1878). B. Bathymetric range of C. (C.) pilosa (Powell, 1951).
Antarctic quadrants follow terminology of Markham (1912). Dashed line indicates Antarctic Convergence.

Table 2.—Chlanidota (Chlanidota) vestita (Mar-
tens, 1878). Shell measurements of lectotype and par-
alectotype 1. Linear measurements in mm.

Paralec-
Character Lectotype totype |
Shell Length (SL) 2205 19.4
Body Whorl Length (BWL) 20.1 16.8
Aperture Length (AL) he aS.
Shell Width (SW) 17.1 14.5
BWL/SL 0.89 0.87
AL/SL 0.76 0.69
SW/SL 0.76 0.75
No. spiral cords on:
Penultimate whorl 8 f
Body whorl 23 22

259

100

150

100

Geographic distribution and bathymetric ranges of Chlanidota (Chlanidota) vestita (Martens, 1878),

Published records.—H.M.S. Challenger:
Sta. 149, Accessible Bay, Kerguelen,
49°08’S, 70°12'E, in 20-25 fm volcanic
mud; Sta. 149B, off Royal Sound, Kergue-
len, 49°28’S, 70°30’W, 25 fm, volcanic
mud; Sta. 149D, off Royal Sound, Kergue-
len, 49°28’S, 70°13’W, 28 fm, volcanic
mud: Sta. 151, off Heard Island,
52°59'30"S, 73°33'30"W, 75 fm, volcanic
mud (Watson, 1886). British, Australian
and New Zealand Antarctic Research Ex-
pedition (BANZARE): Sta. 12, Off Grave
Island, Island Harbor, Kerguelen, 5 m in red
and brown algae; Sta. 58, Hydrography
Channel, SE of Green Island, Kerguelen, in
50 m (Powell 1951). R/V Southern Cross:
Cape Adare, Ross Sea, 43-47 m, BMNH
(Smith 1902:203, see Dell 1990:fig. 311).

260

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 3.—Chlanidota (Chlanidota) densesculpta (Martens, 1885). Measurements of shell characters. Linear

measurements in mm (n = 10, holotype not included).

Character Mean

Shell Length (SL) 34.3
Body Whorl Length (BWL) 2938
Aperture Length (AL) 24.4
Shell Width (SW) Dae
Operculum Length (OL) 6.2
BWL/SL 0.87
AL/SL 0.71
SW/SL 0.68
OL/AL O25

Distribution:—Kerguelen, the Crouzets,
Heard Island and the Ross Sea, in 5—150 m
(Fig. 3).

Remarks.—Chlanidota vestita is readily
distinguished from all congeners by its
small, broadly shouldered shell, stepped
spire, high body whorl, and moderately
strong spiral cords.

In addition to the Kerguelen and Crozet
Plateaus, records include a single report
from Cape Adare on the Antarctic continent
(Smith 1902). Dell (1990:184, fig. 311) ex-
amined and figured the Cape Adare speci-
men [BMNH], and confirmed it to be C.
vestita, and not C. smithi as reported by
Powell (1958). Dell (1964:288) reported C.
vestita from Heard Island.

Chlanidota (Chlanidota) densesculpta
(Martens, 1885)
Figs. 4-7; Table 3

Cominella (Chlanidota) densesculpta Mar-
tens, 1885:91; Martens & Pfeffer, 1886:
71, pl. 1, fig. 3a-f.

Chlanidota densesculpta Thiele, 1912:263;
David, 1934:128; Powell, 1951:140, pl.
$,, fags..3 1-33; Carcelles, 1953192. pL,3.
fig, .60;.. Powell, 19602150; Dell, 71990
(partim):183, fig. 305, non fig. 306.

Description.—Shell (Fig. 4) large for ge-
nus (to 40.3 mm), thin, translucent, fragile,
ovate in outline, with low, rounded spire.
Protoconch unknown, early whorls heavily
eroded. Teleoconch of up to 6 evenly-
rounded convex whorls, deeply eroded

o Range Holotype
PHD 29.5—36.6 S12
1.90 25.9-32.3 27.8
2S 20.6—27.8 24.4
2.20 18.7—27.2 24.5
0.87 4.4-7.5 —
0.01 0.85—0.88 0.89
0.03 0.67—0.75 0.78
0.04 0.63—0.74 0.79
0.04 0.18-0.31 a

where periostracum damaged or missing.
Suture impressed, obscured by thick perios-
tracum. Spiral sculpture of numerous ex-
tremely fine spiral threads, broader than the
Spaces between them. Spiral threads sinu-
ated, equal in strength, closely spaced (4—5
per mm), ~40 on penultimate whorl, >80
on body whorl. Axial sculpture limited to
growth lines, producing finely cancellate
surface sculpture appearing in places more
like a rectangular array of fine pits. Aper-
ture broadly ovate, deflected from shell axis
by 9-—13°. Outer lip thin, evenly rounded,
slightly reflected. Columella <2 AL, weak-
ly concave, with strong siphonal fold. Cal-
lus consisting of thin, silver-edged glaze
overlying parietal region, siphonal fasciole.
Siphonal notch broad, dorsally recurved,
with straight columellar and rounded aper-
tural margins that form borders of fasciole.
Ridge margin of fasciole runs from aper-
tural margin of siphonal notch, may be ev-
ident beneath columellar callus in some
specimens. Shell color chalky white, aper-
ture weakly glazed. Periostracum (Fig. 6C)
thick, orange-tan, hirsute. Hair-like projec-
tions emanating from edges of lamellae at
intersections of spiral threads with axial
growth lines. Operculum (Fig. 4G—I, K)
small (0.18—0.31 AL), broadly ovate, weak-
ly coiled, with terminal nucleus (usually
abraded) rotated nearly perpendicular to
long axis. Attachment area nearly circular,
spanning % of inner surface, posterior, left
margins thickened, glazed, abraded.

VOLUME 112, NUMBER 2

Fig. 4. Chlanidota (Chlanidota) densesculpta (Martens, 1875). A—C. holotype, MNH-U 37478. South Geor-
gia Island. D-E, H, J. R/V Islas Orcadas, Sta. 20, Off South Georgia Island, 54°00'06’S, 37°40'36’W, in 68-—
80 m, USNM 906152. Periostracum of specimen in figure J removed with bleach to reveal spiral sculpture. E
G, I. R/V Eltanin, Sta. 1533, South Georgia Island, 54°00'S, 37°27’ W, in 3—6 m. EF USNM 870722, G, I. USNM
896097. K. Operculum of specimen in fig. D. Scale bar = 1 cm for AJ, 2 mm for K.

262 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Chlanidota (Chlanidota) densesculpta (Martens, 1875). R/V Islas Orcadas, Sta. 19, Off South Geor-
gia Island, 54°01'42”S, 37°40’00’W, in 46-70 m, USNM 901651. A. Left, and B. right lateral views of d animal
removed from shell. C. Ventral and D. left lateral views of anterior alimentary system. E. Posterior portion of
the proboscis, with salivary glands removed to show the valve of Leiblein. E Dorsal, and G. ventral views of
stomach. H. Stomach, opened mid-dorsally. I. Penis. Scale bars = 5 mm. Abbreviations: ao, anterior aorta; ct,
ctenidium; ctz, compacting zone of the stomach; ddg, duct of the digestive gland; dg, digestive gland; gL, gland

VOLUME 112, NUMBER 2

Ultrastructure.—(Fig. 6D) Outermost
layer prismatic (~90 wm), middle layer
comarginal crossed-lamellar (~140 wm),
inner layer radial crossed-lamellar (~32
Lm).

External anatomy.—(Fig. 5A—B). Soft
tissues comprise approximately 2% whorls.
Mantle cavity spans just under % whorl,
kidney %4 whorl, digestive gland and gonad
-1% whorls. Columellar muscle short, broad,
attached to shell at rear of mantle cavity.
Foot large, broadly rectangular (L/W = 1-
1.3). Body base color yellowish tan, dorsal
surfaces of the head, tentacles, siphon, foot
mottled with dark grayish black. Head
small, with broad, tapering tentacles, black
eyes.

Mantle cavity—Mantle edge smooth. Si-
phon of medium length (~% AL), muscular,
extending substantially beyond mantle
edge. Osphradium greenish, bipectinate,
with narrow axis, spanning about % mantle
cavity length. Ctenidium large, wide, span-
ning about % mantle cavity length. Hypo-
branchial gland formed of few, distinct,
widely spaced folds.

Alimentary system.—FProboscis smooth,
unpigmented, of moderate length when re-
tracted (~0.6 AL), but long in semiprotract-
ed position (~1.1 AL; Fig. 5C, D). Probos-
cis sheath very thin-walled, translucent.
Mouth opening forming triangular slit. Buc-
cal mass muscular, large, filling retracted
proboscis. Odontophoral cartilages paired,
fused anteriorly, spanning % of buccal
mass. Radular ribbon long, 10.9—13.2 mm
(0.47-0.52 AL), ~540 wm wide (~0.023
AL), triserial (Fig. 6A, B), consisting of
75-80 rows of teeth, posteriormost 5—7
teeth nascent. Rachidian teeth with 3 cusps
on central portion of broad, anteriorly
arched basal plate. Central cusp slightly

<_

263

longer, narrower than lateral cusps. Lateral
teeth with 3 cusps. Outer cusp stouter, near-
ly twice as long as inner cusp. Intermediate
cusp slightly shorter than, immediately ad-
jacent to inner cusp. Salivary glands small
(Fig. 5C, D; sg), fused, dorsal to nerve ring,
right salivary gland totally covering valve
of Leiblein. Salivary ducts pass loosely
along both sides of esophagus, becoming
embedded in esophageal walls near rear of
buccal mass. Valve of Leiblein well de-
fined, large (Fig. 5E; vL), pyriform, without
ciliary cone. Gland of Leiblein convoluted
anteriorly, tapering posteriorly (Fig. 5C, D;
gL), opening without constriction to mid-
esophagus, just anterior to, left of crop (Fig.
5C; poe). Esophagus wide, muscular, en-
larging posteriorly (Fig. 5C; poe) to form a
crop-like structure lined with tall longitu-
dinal folds. Stomach U-shaped, without
caecum (Fig. 5E H). Paired ducts of diges-
tive gland closely spaced (Fig. 5H; ddg).
Typhlosoles present (Fig. 5H; tph), poorly
defined. Digestive gland clearly divided
into 2 lobes. Zone of compaction well pro-
nounced (Fig. 5H; ctz). Rectum terminates
with well-defined anal papilla.

Female reproductive system.—Typically
buccinoidean. Oviduct opens into medium-
sized albumen gland. Ingesting gland sin-
gle. Capsule gland large, occupies ~% of
mantle cavity. Bursa copulatrix present,
simple, hemispherical.

Male reproductive system.—Seminal
vesicle of medium size, highly coiled. Pros-
tate narrow, running along posterior half of
mantle cavity wall. Penis dorsoventrally
compressed, long (0.8 X mantle cavity
length), with smooth walls, long, cylindri-
cal papilla surrounded by deep circular fold
around its base (Fig. 5I).

Type locality.—South Georgia Island.

of Leiblein; h, heart; ht, cephalic tentacles; ig, intestinal groove; mo, mouth; nep, nephridium; nr, circumoeso-
phageal nerve ring; oe, oesophagus; op, operculum; os, osphradium; p, penis; poe, posterior oesophagus; pr,
proboscis; prr, proboscis retractors; rhd, proboscis sheath; s, siphon; sg, salivary gland; st, stomach; t, testes;

vL, valve of Leiblein.

264

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6.
the central portion of the radular ribbon taken from animal in Fig. 5. C-D. R/V Eltanin, Sta. 1533, South Georgia
Island, 54°00'S, 37°27'W, in 3-6 m, USNM 896097. C. Periostracum. D. Shell ultrastructure, fracture surface
parallel to growing edge. Scale bar = 200 pm for A-C, 100 um for D.

Type material.—Holotype, MNH-U
37478 (fig. 4A-C).

Material examined.—Holotype. R/V EI-
tanin: Sta. 1533, South Georgia Island,
54°00'S, 37°27'W, in 3—6 m, 7 Feb 1966, 1
shell, USNM 870722, 6 specimens, USNM
896097. R/V Islas Orcadas: Sta. 18, Off
South Georgia Island, 54°02'30’S,
37°39'36"W, in 60-71 m, 15 May 1975, 4
specimens, USNM 901650; Sta. 19, Off
South Georgia Island, 54°01'42’S,
37°40'00"W, in 46-70 m, 15 May 1975, 10
specimens, USNM 901651 (anatomical de-

Chlanidota (Chlanidota) densesculpta (Martens, 1875). A. Dorsal, and B. left lateral (45°) views of

Scriptions are based on specimens from this
lot); Sta. 20, Off South Georgia Island,
54°00'06"S, 37°40'36"W, in 68-80 m, 15
May 1975, 7 specimens, USNM 901652;
Sta. 21, Off South Georgia Island,
53°57'30"S, 37°20'42”W, in 27—40 m, 16
May 1975, 1 specimen, USNM 901653;
Sta. 95, Off South Georgia Island,
54°11'48"S, 37°41'06’W, in 68-80 m, 9 Jun
1975, 3 specimens, USNM 901654. R/V
Prof. Siedlecki, Sta. 81, Off South Georgia
Island, 54°43’S, 35°13'W, in 300—306 m, 11
Dec 1986, 1 broken shell, USNM 897523.

VOLUME 112, NUMBER 2

Fis.’ 7.
1875). The type locality is ““South Georgia Island.”’

South Georgia, Bay of Islands, in 7.5 m, 3
specimens, USNM 252874.

Published records.—R/V_ Discovery II:
Sta. 45, 2.7 miles S 85°E of Jason Light,
South Georgia, in 238-270 m; Sta. 141,
East Cumberland Bay, South Georgia, 200
yards from shore under Mt. Duse, in 17—27
m; Sta. 145, Stromness Harbour, South
Georgia, between Grass Island and Tons-
berg Point, in 26-35 m; Sta. 1941, Leith
Harbour, South Georgia, in 55—22 m; Sta.
WS 62, Wilson Harbour, South Georgia, in
26—83 m; Sta. MS 6, East Cumberland Bay,
% mile south of Hope Point to 1% cables S
x E of King Edward Point Light, South
Georgia, in 24-30 m; Sta. MS 10, East
Cumberland Bay, % mile south of Hope
Point to 4 mile south of Government Flag-
staff, South Georgia, in 26-18 m (Powell
1951). SW of Snow Hill Island, 64°36’S,
57°42'W, in 125 m (Strebel 1908). Kergue-
len (22-345 m) and Crozet (22—505 m) Is-
lands (Cantera & Arnaud 1984).

Distribution.—All of the specimens that
we examined were from South Georgia Is-

265

No. of spec.x 5

Geographic distribution and bathymetric range of Chlanidota (Chlanidota) densesculpta (Martens,

land. There have been several published re-
ports of C. densesculpta occurring at other
localities. According to Powell (1951),
Martens (1903) record of C. densesculpta
from Bouvet Island is Notoficula bouveti
Giinele> 1912). Strebel 1908:33) neither
discussed nor figured the single specimen
of C. densesculpta he reported from off the
Antarctic Peninsula. Similarly, the recent
reports of this species from the Kerguelen
(22-345 m) and Crozet (22—505 m) Islands
(Cantera & Arnaud 1984:68) lack descrip-
tions or illustrations. Pending confirmation
of the identification of these records, we
provisionally regard Chlanidota dense-
sculpta to be endemic to South Georgia Is-
land (Fig. 7). Live-collected specimens
were all taken at depths ranging from 3 to
80 m. The only record outside this range is
a single dead and broken specimen (R/V
Prof. Siedlecki Sta. 81) taken in 300—306
m. Dell (1990) misidentified two lots of the
bathyal species Chlanidota (Pfefferia) in-
venusta (described below) as C. (C.) den-
sesculpta, which led to his incorrect report

266

Table 4.—Chlanidota (Chlanidota) pilosa Powell,
1951. Shell measurements of holotype. Linear mea-
surements in mm.

Character Holotype

Shell Length (SL) DIS
Body Whorl Length (BWL) DoS
Aperture Length (AL) PA\a
Shell Width (SW) 20.0
BWL/SL 0.92
AL/SL 0.83
SW/SL 0.79
No. spiral cords on:

Penultimate whorl 21

Body whorl 57

of a very broad bathymetric range (O—1400
m) for the latter species.

Remarks.—This species has a striking
conchological resemblance to Chlanidota
(Pfefferia) palliata Strebel, which is also
endemic to South Georgia Island, but can
be discriminated most easily on the basis of
its small operculum (18—0.31 AL, vs. 0.67—
0.83 AL in C. palliata), which lacks the
“‘feathered”’ posterior edge diagnostic of

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

the subgenus Pfefferia. Chlanidota dense-
sculpta is readily distinguished from other
species of Chlanidota s.s. by its extremely
fine and indistinct spiral sculpture as well
as by the presence of a thick, finely hirsute
periostracum.

Skeletal ossicles of holothurians were
found in the rectum of one of the specimens
dissected.

Chlanidota (Chlanidota) pilosa
Powell, 1951
Figs. 3, 8; Table 4

Chlanidota pilosa Powell, 1951:139, 194,
fig. L73, pl. 8, figs. 29-30; Kaicher 1990:
5807.

Description.—Shell (Fig. 8) small for ge-
nus (to 25.5 mm), very thin, translucent,
fragile, globose, with very low, rounded
spire. Protoconch unknown, early whorls
heavily eroded. Teleoconch of evenly
rounded, highly convex whorls. Spiral
sculpture of numerous fine spiral threads,
broader than intervening spaces. Spiral
threads sinuated, equal in strength, closely

Fig. 8.
456, 1 mile east of Bouvet Island, in 40—45 m. Scale bar = 1 cm.

Chlanidota (Chlanidota) pilosa Powell, 1951. Holotype, BMNH 1961500. R/V Discovery II, Sta.

VOLUME 112, NUMBER 2

267

Table 5.—Chlanidota (Chlanidota) signeyana (Powell, 1951). Measurements of shell characters. Linear mea-

surements in mm (n = 10, holotype not included).

Holotype
of C.
Holotype Holotype elongata
of C. of C. (from Lamy,
Character Mean o Range signeyana pyriformis 1911)
Shell Length (SL) 29.8 6.7 21.442.2 37.4 28.6 30.0
Body Whorl Length (BWL) 24.5 Se 17.8—33.4 31.9 24.4 26.5
Aperture Length (AL) 193 3D 14.1—25.7 255 20.1 20.7
Shell Width (SW) 18.2 3.2, 13.5—22.8 23.6 18.6 16.9
Operculum Length (OL) yo 1:5 4.5-9.4 6.4 a —
BWL/SL 0.83 0.02 0.79-0.86 0.86 0.85 0.87
AL/SL 0.65 0.04 0.58—0.70 0.68 0.70 0.69
SW/SL 0.61 0.06 0.54—0.71 0.64 0.65 0.56
OL/AL 0.30 0.04 0.25—0.37 O27 — a
No. spiral cords on:
Penultimate whorl 10.1 222 7-14 11
Body whorl 25.0 Gr. 15-35 26 21 2

spaced (2.4—3 per mm), 21 on penultimate
whorl, 57 on body whorl. Axial sculpture
limited to growth lines. Aperture broadly
ovate, deflected from shell axis by 15°. Out-
er lip very thin, evenly rounded, slightly re-
flected. Columella <% AL, weakly con-
cave, with strong siphonal fold. Callus con-
sisting of a thin glaze overlying parietal re-
gion and siphonal fasciole. Siphonal notch
broad, slightly dorsally recurved, with
Straight columellar, rounded apertural mar-
gins that mark borders of fasciole without
forming raised ridge margins. Shell color
white, aperture weakly glazed. Periostrac-
um thin, light yellowish-brown, hirsute.
Operculum small (0.3 AL), irregularly
ovate. Radula is similar to that C. (C.) dense-
sculpta except rachidian cusps are closer to-
gether, more broadly triangular, with the
central cusp broader and longer than flank-
ing cusps (Dell 1951:fig. L73).

Type locality.—1 mile east of Bouvet Is-
land, in 40—45 m (R/V Discovery II, Sta.
456).

Type material.—Holotype (live collect-
ed) BMNH 1961500 (Fig. 8), and paratype
(empty shell).

Material examined.—The type material.

Distribution.—Known only from Bouvet
Island.

Remarks.—This species is thus far
known only from the type material. Chlan-
idota pilosa is similar to low-spired, rotund
specimens of C. densesculpta (e.g., Fig.
4E), but differs in having fewer and more
pronounced spiral riblets on the penultimate
and body whorls, and by having a fasciole
that lacks a ridge along its margin. These
two species are also widely separated in
their distribution. Chlanidota pilosa differs
from C. vestita in having a lower spire, as
well as more numerous and finer spiral
threads.

Chlanidota (Chlanidota) signeyana
Powell, 1951
Figs. 9-11; Table 5

Chlanidota signeyana Powell, 1951:141,
fig. F 194, fig L74, 196, fig. N129, pl. 8,
figs. 34-35; Carcelles, 1953:191, pl. 3,
fig. 59; Powell, 1960:150; Kaicher, 1990:
5801; Dell, 1990:177, fig. 307.

Cominella (Chlanidota) vestita var. elon-
gata Lamy, 1910:319; Lamy, 1911:6, pl.
nis. ©:

Chlanidota elongata Powell, 1951:140,
194. tie 76; Carcelles, 19532191, ply 5,
fig. 56; Powell, 1960:150; Cernohorsky,
1977:110; Horikoshi et al., 1979: 22, fig.

268 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 9. Chlanidota (Chlanidota) signeyana Powell, 1951. A-C. Holotype, Chlanidota signeyana, BMNH
1961512, R/V Discovery IT, Sta. 167, Off Signy Island, South Orkney Islands, 60°50’30”S, 46°15’00”’W, in 244—
344 m. D. Holotype, Cominella (Chlanidota) vestita var. elongata Lamy, 1910 (from Lamy, 1911), King George
Island, South Shetland Islands, in 420 m. E, FE Holotype, Chlanidota pyriformis Dell, 1990, USNM 613075,

VOLUME 112, NUMBER 2

8; Hain, 1989:71, Taf. V/4, Taf. XXIII/1;

Numanami, 1996: 160-162, figs. 106— -:

107.

Chlanidota lamyi Dell, 1990:182, fig. 310,
new name for Cominella (Chlanidota)
elongata Lamy, 1910, not Cominella
elongata Dunker, 1857.

Chlanidota pyriformis Dell, 1990:182, fig.
309.

Chlanidota cf. lamyi Dell, 1990:182, fig.
315.

Description.—Shell (Fig. 9) large for ge-
nus (to 42.6 mm), thin, fragile, elongate to
ovate in outline, with medium to high,
rounded to turreted spire. Protoconch un-
known, early whorls heavily eroded. Teleo-
conch of up to 5+ evenly-rounded convex
whorls, deeply eroded where periostracum
damaged or missing. Suture deeply im-
pressed, sharply abutting. Spiral sculpture
typically of distinct, sharp, narrow cords
(7—14 on penultimate whorl, 15—35 on body
whorl) equal or subequal in prominence,
that may be reduced or nearly absent in
some specimens (Fig. 9E K). In specimens
with pronounced cords (Fig. 9H—I), includ-
ing the holotype (Fig. 9A—C), 4—5 adapical
cords on body and penultimate whorls usu-
ally more widely spaced, giving the illusion
that they are more pronounced. Spaces be-
tween cords equal to cord width on early
whorls, at least twice cord width on body
whorl. Axial sculpture limited to fine,
weakly recurved, prosocline growth lines.
Aperture elongate to broadly ovate, deflect-
ed from shell axis by 14—17°. Outer lip thin,
evenly rounded, fragile. Columella 1/3—1/2
AL, weakly convex, with strong siphonal
fold. Callus of thin, silver-edged or thick,
brownish glaze overlying parietal region,
siphonal fasciole. Siphonal notch broad,

—

269

dorsally recurved, with straight columellar
and rounded apertural margins that form
borders of fasciole. Ridge margin of fasci-
ole extends from apertural margin of si-
phonal notch. Shell color chalky-white to
brownish or orange-tan. Aperture weakly
glazed. Periostracum (Fig. 10E) thin to
moderately thick, orange-tan to greenish-
tan, smooth in weakly sculptured speci-
mens, forming low, axial lamellae with
short, hairlike projections on intersections
with spiral cords of strongly sculptured
specimens. Operculum (Fig. 9L) small
(0.25—0.37 AL), broadly ovate, weakly
coiled, with terminal nucleus rotated nearly
perpendicular to long axis. Operculum may
be thin, yellowish, translucent, but usually
Opaque, brownish to dark brown. Attach-
ment area oval, spanning about 7; of inner
surface, posterior and left margins thick-
ened, abraded.

Ultrastructure.—(Fig. 10F) Outermost
layer prismatic (~38 pum), middle layer
comarginal crossed-lamellar (~60 wm), in-
ner layer radial crossed-lamellar (~20 um).

Anatomy.—Gross anatomical features of
C. signeyana very similar to those of C.
densesculpta, but left and right salivary
glands of C. signeyana separate, not fused.
Radular ribbon long, 9.6—12.2 mm (0.48—
0.55 AL), 790-910 pm wide (0.04 AL),
triserial (Fig. l1OA—D), consisting of 65-75
rows of teeth, posteriormost 5—6 rows na-
scent. Rachidian teeth with arched base,
straight lateral sides, 3 large, robust cusps
of equal length. Lateral teeth with 3 cusps,
outer cusp longest, intermediate cusp short-
est, situated close to inner cusp.

Type locality—[Chlanidota signeyana]
Off Signy Island, South Orkneys,
60°50'30"S, 46°15’00"W, in 244-344 m

R/V Westwind, Sta. W-10, off South Shetland Islands, 63°00’S, 60°32’W, in 159 m. G. R/V Eltanin, Sta. 997,
Gibbs Island, Bransfield Strait, South Shetland Islands, in 769 m, USNM 881971. H-I. R/V Eltanin, Sta. 426,
Bransfield Strait, South Shetland Islands, in 809-1116 m, USNM 886204. J—K. R/V Islas Orcadas, Sta. 55,
Saunders Island, South Sandwich Islands, in 64-88 m, USNM 901664. L. Operculum of specimen in figs. J, K.

Scale bar = 1 cm for A—K, 1 mm for L.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 10. Chlanidota (Chlanidota) signeyana Powell, 1951. A. Dorsal and B. left lateral (45°) views of the
central portion of the radular ribbon taken from the animal in Fig. 9J, K. C. Dorsal and D. left lateral (45°)
views of the central portion of the radular ribbon taken from the animal in fig. 9H, I. E. Periostracum and EF
shell ultrastructure of the same specimen. Scale bars = 200 pm for A—-D, 500 pm for E, 50 pm for FE

VOLUME 112, NUMBER 2

Fig. 11.

Depth, m

1000

No. of spec.x 5

Geographic distribution and bathymetric range of Chlanidota (Chlanidota) signeyana Powell, 1951.

© = type locality, Chlanidota signeyana; + = type locality, Chlanidota lamyi; ¢ = type locality, Chlanidota
pyriformis. @ = examined material; _] = published records.

(R/V Discovery II, Sta. 167); [Cominella
(Chlanidota) vestita var. elongata Lamy,
1910] King George Island, South Shet-
lands, in 420 m; [Chlanidota pyriformis
Dell, 1990] Off South Shetland Islands,
63°00’S, 60°32’W, in 26 m (R/V Westwind:
Sta. W.10).

Type material.—{Chlanidota signeyana]
Holotype, BMNH 1961512 (fig. 9A—C),
and 2 paratypes; [Cominella (Chlanidota)
vestita var. elongata Lamy, 1910] The ho-
lotype (Fig. 9D) was not found in MNHN;
[Chlanidota pyriformis Dell, 1990] Holo-
type, USNM 613075 (Fig. 9E-F) paratypes
1-6, USNM 860181, Paratype 7, ME
47204.

Material examined.—R/V Eltanin: Sta.
426, South Shetland Islands, Bransfield
Strait, 62°27'S, 57°58’W, in 809-1116 m, 5
Jan 1963, 3 shell fragments, USNM

870290, 7 shells or fragments, USNM
870291, 4 specimens, USNM 886204; Sta.
428, South Shetland Islands, Bransfield
Strait, /62-41'S., 57° SW. 1n:662—1120 m5
Jan 1963, 1 shell fragment, USNM 870293;
Sta. 432, South Shetland Islands, 62°52’S,
59°27'W, in 884—935 m, 7 Jan 1963, frag-
ments of 2 shells, USNM 870303; Sta. 439,
Antarctic Peninsula, 63°51'S, 62°38’W, in
128-165 m, 9 Jan 1963, 1 shell, USNM
870313, 1 specimen, USNM 881919; Sta.
444, South Shetland Islands, 62°56’S,
62°02'W, in 750—732 m, 11 Jan 1963, 1
specimen, USNM 881923; Sta. 538, South
Orkney Islands, Bransfield Strait, 60°30’S,
47°34'W, in 616—662 m, 6 Mar 1963, frag-
ments of 2 shells, USNM 870330; Sta. 993,
South Shetland Islands, Elephant Island,
61°25'S, 56°30’W, in 300 m, 13 Mar 1964,
1 specimen, USNM 881970; Sta. 997,

242.

South Shetland Islands, Bransfield Strait,
Gibbs Island, 61°44.18'’S, 55°56.06’W, in
769 m, 14 Mar 1964, 1 specimen, USNM
881971; Sta. 1079, Scotia Ridge, 61°26’S,
41°55'W, in 593-598 m, 13 Apr 1964, frag-
ments of 3 shells, USNM 870627, 2 spec-
imens, USNM 881983; Sta. 1084, Scotia
Ridge, 60°22’S, 46°50'W, in 298—403 m, 15
Apr 1964, 3 shells, USNM 870641, 7 shells
and fragments, USNM 870642, 1 specimen,
USNM 881988; Sta. 1997, Ross Sea,
72°00’S, 172°28’E, in 530—549 m, 2 shells,
USNM 898205; Sta. 2124, Ross Sea,
71°38'S, 172°00’'E, in 606—622 m, 1 shell,
USNM 898202; Sta. 2127, Ross Sea,
71°23'S, 171°36’E, in 515-521 m, 1 shell,
USNM 898040.

R/V Eastwind: Sta. EW66-009, Palmer
Peninsula 62°43.1'S, 62°17.5'W, in 561 m,
31 Jan 1966, 1 specimen, USNM 678473;
EW66-012, Palmer Peninsula 63°23'S,
60°51’W, in 405 m, 3 Feb 1966, 2 speci-
mens, USNM 678475; EW66-021, off
South Orkney Islands, 60°21'S, 45°55’W, in
102 m, 9 Feb 1966, 1 specimen, USNM
678396; EW66-022, off South Orkney Is-
lands, 60°26.5'S, 45°53.3’W, in 168 m, 9
Feb 1966, 1 specimen, USNM 678476;
EW66-029, off South Orkney Islands,
61°06'S, 44°57'W, in 290 m, 15 Feb 1966,
1 specimen, USNM 678398; EW66-036,
off Elephant Island, 61°16’S, 54°45’W, in
300 m, 17 Feb 1966, 2 shells, USNM
678480; off beach wrack, Collins Pt., De-
ception Island, 4 Jan 1966, 4 specimens,
USNM 678378.

R/V Hero: Cruise 691, Sta. 23, off Ant-
arctic Peninsula, Palmer Archipelago, Bra-
bant Island, 64°12.06’S, 62°39.36’W, in 93—
95 m, 9 Feb 1969, 1 specimen, USNM
896260; Cruise 691, Sta. 31, South Shet-
land Islands, Deception Island, 62°58.25'S,
60°45.40'W, in 37-51 m, 13 Feb 1969, 7
specimens, USNM 897557; Cruise 721,
Sta. 751, off Antarctic Peninsula,
64°46'28"S, 64°04’20"W, in 33 m, 31 Dec
1971, 1 shell, USNM 901659; Cruise 721,
Sta. 765, off Antarctic Peninsula, 64°47.3'S
64°0O7.4'W, in 55 m, 3 Jan 1972, 1 shell,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

USNM 901671; Cruise 721, Sta. 1058, off
Antartic Peninsula, 62°19.0'S, 59°11.4'W,
in 44 m, 19 Dec 1971, 1 shell, USNM
901657; Cruise 731, Sta. 1806, 64°46'31”S,
64°04'52”W, in 47-75 m, 18 Feb 1973, 1
specimen, USNM 901686; Cruise 824, Sta.
32-1, Antarctic Peninsula, Anversa Island,
64°37'S, 62°50.48'W, in 640—670 m, 23
Mar 1982, 1 shell, USNM 881583; Cruise
824, Sta. 7-1, Antarctic Peninsula, Adelaide
Island, 66°40.40'S, 67°31.23'W, in 510—730
m, 17 Mar 1982, 1 specimen, _USNM
896309; Sta. 32-1, Antarctic Peninsula,
64°37'S, 62°50.80'W, in 640—670 m, 25
Mar 1982, 1 shell, USNM 901658.

R/V Islas Orcadas: Sta. 39, South Sand-
wich Islands, 57°01.2'S, 26°44.3'W, in 97—
100 m, 23 May 1975, 1 shell, USNM
901663; Sta. 40, South Sandwich Islands,
Kadlemas Island, 57°06.8’S, 26°43.36’W, in
15-33 m, 23 May 1975, 1 specimen,
USNM 901660; Sta. 42, South Sandwich
Islands, Kadlemas Island, 57°06.8’S,
26°43.6'W, in 22—44 m, 24 May 1975, 12
specimens, USNM 901669; Sta. 46, South
Sandwich Islands, Kadlemas _ Island,
57°06.2'S, 26°44.5'W, in 26-60 m, 25 May
1975, 2 specimens, USNM 901665; Sta. 48,
South Sandwich Islands, Kadlemas Island,
57°06.3’S, 26°44.2’W, in 27-62 m, 23 May
1975, 4 specimens, USNM 901670; Sta. 55,
South Sandwich Islands, Saunders Island,
57°47.12'S, 26°22.30'W, in 64—88 m, 27
May 1975, 1 specimen, USNM 901664;
Sta. 57, South Sandwich Islands, 27 May
1975, 5 specimens, USNM 901656; Sta. 80,
South Sandwich Islands, Zavodovski Is-
land, 56°20.0’S, 27°38.8'W, in 351—393 m,
4 Jun 1975, 13 specimens, USNM 901667;
Sta. 110, Inaccessible Island, 60°28.1’S,
46°27.2'W, in 115—132 m, 16 Feb 1976, 2
specimens, USNM 901666; Sta. 115, Inac-
cessible Island, 60°32.4’S, 47°22.7'W, in
567—671 m, 17 Feb 1976, 2 shells, USNM
901655; Sta. 118, off South Orkney Islands,
62°01.5'S, 43°06.2'W, in 759-857 m, 20
Feb 1976, 1 specimen, USNM 901661; Sta.
121, off South Orkney Islands, 61°47.00'S,

VOLUME 112, NUMBER 2

43°40.00’W, in 616—642 m, 21 Feb 1976, 1
specimen, USNM 901662.

R/V Westwind: Sta. W-10, off South
Shetland Islands, 63°00’S, 60°32’W, in 159
m, 26 Jan 1958, 6 shells, paratypes of C.
pyriformis Dell, 1991. USNM 860181, 4
shells, USNM 890897.

R/V Polar Duke: off Palmer Peninsula,
65°36'S, 67°21'W, in 200 m, 6 Sep 1985, 1
shell, USNM 846179.

R/V Deepfreeze IV: Sta. TD2-ED 14, off
Cape Norwegia, Weddel Sea, 71°50’S,
15°50’W, 1028-1122 m, 18 Jan 1959, 1
shell and fragments, USNM 638862.

Published records.-R/V Discovery I],
Sta. 162, Off Signy Island, South Orkneys
60°48'00"S, 46°08’00"W, in 320 m; Sta.
Menon -Cape: Bowles,..Clarence Is:,
Gie2a30'S, 60°28'00'W, in 342: m; Sta.
175, Bransfield Strait, South Shetlands,
GSoebi20S, 59°48'15"W, in 200: m;_ Sta:
1952, between Penguin Island and Lion’s
Rump, King George I, South Shetlands, in
367-383 m; Sta. 1957, off south side of
Clarence Island, South Shetlands, in 785—
810 m (Powell, 1951). R/V Eltanin: Sta.
426, off South Shetland Islands, 62°27’S,
57°58'W, in 1116—809 m; Sta. 439, west of
Antarctic Peninsula, 63°51’'S, 62°38'’W, in
128—165 m; Sta. 1084, north of South Ork-
ney Islands, 60°22’S, 46°50’W, in 298—403
m, R/V Hero: Sta. 23, Antarctic Peninsula,
64°12.1'S, 62°39.6’W, in 93-95 m; Arthur
Harbor, Anvers Island, Antarctic Peninsula,
30 m (Dell, 1990:180). PS ANT III/3: Sta.
345, Weddel Sea, 73°27’S, 21°37'W, in 617
m (Hain, 1989). Syowa Sta., Enderby Land,
in 98 m (Horikoshi et al., 1979). JARE Sta.
9, Breid Bay, Queen Maud Land,
we 15.7 S, 24°25.7'E; m’ 276289 m, Sta.
10, 25 (Numanami, 1996); NZOI: Sta.
A463, 72°20’S, 174°50’E, in 460 m; Sta.
A464, 72°20'S, 174°00’E, in 376 m (Dell,
1990, as C. cf. lamyi).

Distribution.—South Orkneys, South
Shetlands, Antarctic Peninsula, and Queen
Maud Land, and the eastern margins of the
Ross Sea. Living specimens were taken in
30-1116 m (Fig. 11).

2T3

We include records for this species re-
ported from the area of Queen Maud Land
(as C. elongata). Figured specimens from
this region (Numanami, 1996:fig. 1O6A—B,
E-—F). appear to have weaker, more numer-
ous spiral cords. One specimen from Syowa
station has rachidian teeth with narrower
cusps and additional serrations, features that
had not previously been observed in spec-
imens of C. signeyana. Powell (1951:142,
fig. L78) reported and illustrated similar ad-
ditional serrations in a specimen of Chlan-
idota (Pfefferia) elata.

We have examined and also include re-
cords from the eastern margin of the Ross
Sea, which had been listed as Chlanidota
cf. lamyi by Dell (1990:310). Dell (1990)
noted that these broken and worn speci-
mens closely resemble C. signeyana (as C.
lamyi) from the South Shetland Islands, but
differed only in having more numerous spi-
ral cords on their body whorl.

Remarks.—Lamy (1910) was the first to
recognize this species, proposing the taxon
Cominella (Chlanidota) vestita var. elon-
gata, without realizing that the name was
preoccupied by Cominella elongata Dun-
ker, 1857. Powell (1951:141) considered
Chlanidota elongata to be restricted to the
South Shetland Islands and erected the tax-
on C. signeyana based on material from the
South Orkney Islands. He recognized that
C. signeyana was closely related to C. elon-
gata, but distinguished it as being ‘“‘con-
stantly broader and of ovate rather than cy-
lindrical outline.’’ Cernohorsky (1977:110)
noted that Cominella (Chlanidota) vestita
var. elongata Lamy, 1910 was preoccupied,
but questioned whether this taxon was dis-
tinct from C. vestita. After examining a
broader range of specimens than were
available to Powell, Dell (1990:180) noted
that both ovate and cylindrical phenotypes
co-occurred in South Shetland and South
Orkney samples, but that specimens could
be sorted into one or the other phenotype
on the basis of the ratio of shell width (SW)
to shell height (SH). He regarded Chlani-
dota signeyana to be limited to the ovate

274

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 6.—Chlanidota (Chlanidota) paucispiralis Powell, 1951. Measurements of shell characters. Linear mea-

surements in mm (n = 10, holotype not included).

Character Mean

Shell Length (SL) 29.8
Body Whorl Length (BWL) 22.2
Aperture Length (AL) eS)
Shell Width (SW) 15
Operculum length (OL) 4.5
BWL/SL 0.85
AL/SL 0.67
SW/SL 0.65
OL/AL O27
No. spiral cords on:

Penultimate whorl 4.4

Body whorl 10.6

phenotype with SW/SH ratios ranging from
0.60 to 0.66 and proposed the new name
Chlanidota lamyi (for Cominella elongata
Lamy, 1910, non Dunker, 1857) for the
elongate phenotype, with SW/SH ratios be-
tween 0.55 and 0.60. He did, however, rec-
ognize the possibility that “these two forms
are but extremes of the same species.”’

Dell (1990) also described C. pyriformis
based on a single lot (11 specimens) col-
lected off the South Shetland Islands, which
he regarded to be a morphologically con-
sistent shallow-water form allied to both C.
signeyana and C. lamyi, but distinguished
by its tapering, conical spire.

We examined and measured a large series
of specimens from throughout the ranges of
the three nominal species, but were unable
to differentiate between them in any repro-
ducible manner. Larger samples from sev-
eral stations (e.g., USNM 901669, USNM
897557, USNM 860181) revealed that each
population spans the range of morphologies
of at least two, and sometimes all three of
these nominal species. Chlanidota signey-
ana is highly variable in shell outline and
spiral sculpture, and encompasses the phe-
notypes of Chlanidota lamyi and C. pyri-
formis.

There does appear to be a clinal gradient
in shell shape, with northern specimens
tending to be more inflated, while popula-
tions from the southern portion of the spe-

o Range Holotype
3.0 21.9-31.7 209
2.4 19.2—27.2 js
1.8 15.8-21.9 15.8
Zl 14.4—21.7 14.4
0.7 4.0-5.0 4.0
0.01 0.84—0.88 0.88
0.03 0.63—0.72 O72
0.02 0.62—0.68 0.66
0.02 0.25—0.28 0.25
0.5 4-5 4
1.6 8-13 10

cies’ range, including the Antarctic main-
land, have a higher proportion of narrower
and more cylindrical shells. Numanami
(1996:146) recognized a similar gradient in
shell morphology in the taxon Neobuccin-
um eatoni, noting that specimens from the
Kerguelen and South Shetland Islands tend-
ed to be more elongated, while the samples
from Enderby Land tended to be more in-
flated and shorter spired. However, his large
sample (n = 98) from Enderby Land
showed a wide variability in shell form, and
included examples from the entire range of
variation for the species.

Chlanidota signeyana is most similar to
C. (C.) paucispiralis Powell, 1951, but dif-
fers in having more numerous and weaker
spiral cords. Moreover, C. (C.) signeyana
has not been recorded off South Georgia,
while C. (C.) paucispiralis is endemic to
South Georgia Island.

Chlanidota (Chlanidota) paucispiralis
Powell, 1951
Figs. 12,13; 14)°15; Yable

Chlanidota paucispiralis Powell, 1951:141,
194, fig. L77, pl. 8, figs. 36-37; Carcel-
les, 1953:191,. pl. 3, fig. 57; -Beme
1960:150; Kaicher, 1990:5806; Dell,
1990:183—4, fig. 308.

Description—Shell (Fig. 12) large for
genus (to 40.3 mm), thin, strong, ovate to

VOLUME 112, NUMBER 2

elongate-ovate in outline, with moderately
high, conical spire. Protoconch unknown,
early whorls heavily eroded. Teleoconch of
up to 5% evenly-rounded convex whorls.
Suture strongly impressed, forming very
narrow channel between adapical spiral
cord and previous whorl. Spiral sculpture of
narrow, sharp, widely spaced cords, 4—5 on
penultimate whorl, 8—13 on body whorl.
Space between cords wide, evenly concave,
with O-8 very fine spiral threads. Two
adapical cords form narrowly tabulate
shoulder. Axial sculpture of fine, closely
spaced growth lines. Aperture ovate, de-
flected from shell axis by 9—15°. Outer lip
thin, evenly rounded, with serrated edge
formed by spiral cords. Columella <% AL,
weakly concave, with pronounced siphonal
fold. Parietal callus narrow, thick, overlying
spiral cords. Siphonal notch dorsally re-
curved, with straight columellar, rounded
apertural margins that define limits of fas-
ciole. Apertural margin of siphonal notch
dorsally deflected, giving rise to low, sharp
keel that forms “ridge margin”’ of fasciole.
Shell color chalky white, aperture thinly
glazed, occasionally with margins of glaze
discolored to gray or golden tan. Perios-
tracum (Fig. 13C, D) thin, smooth, yellow-
ish-tan, sometimes consisting of closely ad-
jacent, blade-like lamellae, which overlay
spiral sculpture without giving rise to hair-
like projections. Operculum (Fig. 12J)
small (0.25—0.28 xX AL), broadly ovate,
weakly coiled, with laterally terminal nu-
cleus (usually abraded) rotated nearly per-
pendicular to long axis. Attachment area
nearly circular, spanning *% of inner surface,
posterior and left margins thickened,
glazed.

Ultrastructure.—(Fig. 13D) Outermost
layer prismatic (~90 pm), middle layer
comarginal crossed-lamellar (~235 jm),
inner layer radial crossed-lamellar (~55
1m).

Anatomical features of C. (C.) paucispir-
alis are very similar to those of C. (C.) den-
sesculpta. Only a single female specimen

275

(SL = 27.7 mm) was available for dissec-
tion.

External anatomy.—Soft tissues com-
prise ~3% whorls. Mantle cavity just over
% whorl, kidney %4 whorl, digestive gland
2% whorls. Columellar muscle short, broad,
attaching to shell at rear of mantle cavity.
Foot small, rectangular (L/W = 1.4), with
long (0.2 X foot length) medial, ventral
pedal gland. Base color yellowish tan, mot-
tled with patches of grayish black on dorsal
surfaces of siphon, tentacles, foot. Head
small, with thin cylindrical tentacles, large
black eyes. Siphon long, free, muscular.

Alimentary system.—Radular ribbon
= 9.0 mm long (0.51 AL), 830 wm wide
(0.05 AL), triserial (Fig. 13A—B), of ~70
rows of teeth, of which 5—7 are nascent.
Rachidian teeth with arched base, straight
lateral sides, 3 large, robust cusps, central
cusp slightly longer that lateral cusps. Lat-
eral teeth with 3 cusps, outer cusp longest
and, intermediate cusp shortest, adjacent to
inner cusp. Anterior foregut very similar to
that of its congeners. Stomach (Fig. 14) dif-
fers in having very short caecum, which
may be homologous to caecum (posterior
mixing area) of Buccinidae.

Type locality—South Georgia, 53°52'30'S,
36°08'00’"W, in 160 m (R/V Discovery II,
Sta. 159):

Type _ material.—Holotype, BMNH
1961513 (fig. 12A—C), and 2 paratypes,
BMNH 1961513.

Material examined.—Holotype. R/V Is-
las Orcadas: Sta. 8, off South Georgia Is-
land, 53°35'48"S, 37°35'12”"W, in 254-366
m, 11 May 1975, 1 shell, USNM 881743;
Sta. 10, off South Georgia Island,
53°47'48"S, 37°26'42"W, in 165—234 m, 12
May 1975, 1 shell, USNM 881745; Sta. 12,
off South Georgia Island, 53°38'12’S,
37°54'42’W, in 130-137 m, 13 May 1975,
4 shells, USNM 881708; Sta. 13, off South
Georgia Island, 53°44'12”S, 37°59'30’W, in
128—137 m, 13 May 1975, 5 shells + frag-
ments, USNM 901672; Sta. 14, off South
Georgia Island, 53°41'48"S, 37°57'12”W, in
144-150 m, 14 May 1975, 1 shell, USNM

276 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 12. Chlanidota (Chlanidota) paucispiralis Powell, 1951. A-C. Holotype, BMNH 1961513, R/V Dis-
covery II, Sta. 159, South Georgia, 53°52'30"S, 36°08'00"W, in 160 m. D, E. R/V Prof. Siedlacki, Sta. 105,
South Georgia, 53°40’S, 36°48’W, in 161-192 m, USNM 897515. E R/V Prof. Siedlacki, Sta. 20, 53°58’S,
38°42'W, in 189-200 m, USNM 897583. G. R/V Prof. Siedlacki, Sta. 16, Off South Georgia Island, 53°44’S,

VOLUME 112, NUMBER 2

Pgh

Fig. 13.
the central portion of the radular ribbon taken from animal in fig. 12H, I. C. Periostracum and D. shell ultra-
structure. R/V Prof. Siedlacki, Sta. 24, 54°05'S, 38°25'W, in 197-207 m, USNM 897531. Scale bars = 200 wm.

881716; Sta. 15, off South Georgia Island,
53°37'42"S, 38°04'00"W, in 128-137 m, 14
May 1975, 1 shell, USNM 901673; Sta. 16,
off South Georgia Island, 53°38'12’S,
38°01'06"W, in 130-134 m, 14 May 1975,
1 shell, USNM 901675; Sta. 17, off South
Georgia Island, 53°36'00"S, 38°03’00"W, in
122-124 m, 14 May 1975, 1 specimen + 2
shells, USNM 901674 (anatomy studied);
Sta. 30, off South Georgia Island,

<_

Chlanidota (Chlanidota) paucispiralis Powell, 1951. A. Dorsal and B. left lateral (45°) views of

53°50'36"S, 36°18'36"W, in 185—205 m, 19
May 1975, 2 shells, USNM 887839.

R/V Prof. Siedlecki: off South Georgia
Island, Sta. 16, 53°44’S, 39°22’W, in 304—
342m, 02) Dec’ 1986, 4 shells, USNM
897553; Sta. 20, 53°58'S, 38°42’W, in 189—
200 m, 02 Dec 1986, 2 shells, USNM
897583; Sta. 24, 54°05’S, 38°25'W, in 197—
207 m, 03 Dec 1986, 2 shells, USNM
897531; Sta. 28, 54°30’S, 38°56’W, in 220—

39°22'W, in 304-342 m, USNM 897553. H, I. R/V Islas Orcadas, Sta. 17, Off South Georgia Island, 53°36'00"S,
38°03'00"W, in 122-124 m, USNM 901674, (anatomical data based on this E specimen). J. operculum of
specimen in figs. H, I. Scale bar = 1 cm for A-I, 2 mm for J.

See ee
seen,

Fig. 14.
view of the stomach. B. Stomach opened mid-ventrally. Scale bars = 2 mm. Abbreviations: ctz, compacting
zone of the stomach; dc, duct pouch; ddg, duct of the digestive gland; ig, intestinal groove; pm, posterior mixing
area, Or Caecum; poe, posterior oesophagus; sa, sorting area; tph, typhlosoles.

232m; 03> Dee 1986." 1 shell] “USNM
897484; Sta. 87, 54°18'S, 35°37'W, in 238—
247 mm, 11° Dec 1986; (2<shells, USNM
$907537: Sta. 101, 53°47'S2 36°34 Wonia
263-277 m, 1 shell, USNM 897457; Sta.
105, 53°40'S, 36°48'W, in 161—192 m, 14
Dec 1986, 1 shell, USNM 897515; Sta.
116, 53°43’S, 38°36’W, in 260-306 m, 16
Dec 1986, 1 shell, USNM 896993; Sta.
121,-58°57'S,.38°10!Wo in? 80-100 me 16
Dec 1986, 1 shell, USNM 896991.

R/V Eltanin: Sta. 671, off South Georgia
Island, 54°41’S, 38°38’W, in 220-320 m, 23
Aug 1963, 1 shell, USNM 870378; Sta.
678, off South Georgia Island, 54°49’S,
38°O1'W, in 732-814 m, 24 Aug 1963, 1

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

pm

poe

Chlanidota (Chlanidota) paucispiralis Powell, 1951. Stomach of specimen in fig. 12H I. A. Dorsal

juvenile specimen, USNM 881942; Sta.
1535, off South Georgia Island, 53°51’S,
37°38'W, in 97-101 m, 7 Feb 1966, 10
shells + fragments, USNM 898666.
Distribution.—South Georgia Island, in
97-814 m (Fig. 15). Live specimens were
collected in 122—814 m.
Remarks.—Unlike most of its congeners,
this geographically restricted species dis-
plays remarkably little variation in shell
morphology. It is readily distinguished from
all other Chlanidota sensu stricto by its
prominent, broadly spaced, spiral cords. It
most closely resembles Chlanidota (Pfeffer-
ia) chordata (Strebel, 1908), from which it
is readily distinguished by its small, smooth

VOLUME 112, NUMBER 2

Big, 15.
1951. & = type locality; @ = examined material.

operculum. Chlanidota (Pfefferia) chordata
(Strebel, 1908) has a thicker shell and a
thicker, hirsute periostracum.

Subgenus Chlanidota (Pfefferia)
Strebel, 1908

Pfefferia Strebel, 1908:33—4; Powell, 1951:
142.
Chlanidota (Pfefferia) Thiele, 1929:315.

Type species.—Pfefferia palliata Strebel,
1908, by subsequent designation, Powell,
1951:142.

Description.—Shell medium sized for
family, maximum adult size 34.5 to 38.3
mm, depending on species. Shell usually
moderately thick, ovate to elongate in out-
line, with short spire. Shell covered with
thick periostracum, either smooth or with
fine axial lamellae. Spiral sculpture of thin
threads or prominent cords. Aperture high,
oval. Operculum (Fig. 16) large (0.48—0.83
AL), leaf-shaped, tapered above, below,
with terminal nucleus rotated nearly per-
pendicular to long axis of operculum. Pos-
terior edge of operculum with tall ridge of

279

Depth, m

No. of spec.x 5

Geographic distribution and bathymetric range of Chlanidota (Chlanidota) paucispiralis Powell,

feathered lamellae, broadest just above mid-
length. Siphonal notch broad, dorsally re-
curved, siphonal fasciole with ridged mar-
gins. Shell ultrastructure and gross anatomy
of mantle cavity, alimentary and male re-
productive systems as in Chlanidota sensu
stricto.

Remarks.—The subgenus Chlanidota
(Pfefferia) is endemic to South Georgia Is-
land. In his description of Pfefferia, Strebel
(1908) noted many similarities with Chlan-
idota, especially in the morphology and
sculpture of the shell and periostracum. He
distinguished the two as genera on the basis
of their opercula, that of Chlanidota being
small, triangular, with rounded corners, and
with its apex and nucleus along the poste-
rior margin. The operculum of Pfefferia is
much larger, elongated, tapering above and
below, and has a characteristic, ‘““feathered’”’
posterior edge. Strebel further commented
that low and high-spired forms (without in-
termediates) were encountered at each of
the two stations from which Pfefferia had
been collected. Each station also contained
strongly and weakly sculpture forms. While

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 16. Chlanidota (Pfefferia) palliata (Strebel, 1908). A. Operculum showing terminal nucleus rotated
nearly perpendicular to long axis of operculum. Lines C-C’ and F-F’ correspond to the sections shown by Figs.
C and E B. Left lateral (45°) view of operculum. C. Section trough flat, central portion of operculum (along the
line C-C’ on Fig. A), nucleus at the bottom. D, E. Successive layers are closely appressed, adherent in this

VOLUME 112, NUMBER 2

expressing uncertainty as to whether these
differences warranted specific status, Stre-
bel nevertheless described four species,
commenting that additional material would
allow these questions to be better answered.

Examination and dissection of additional
material of three species of Chlanidota
(Pfefferia) revealed that Chlanidota (Pfef-
feria) cannot be distinguished from Chlan-
idota sensu stricto on the basis of gross
anatomy. Of special interest is the variable
operculum of the newly described C. (P.)
invenusta, which ranges from the typical
Chlanidota (Pfefferia) operculum to one
very similar to, but larger than, that of
Chlanidota (Chlanidota), and includes in-
termediate morphologies. Due to the pau-
city of distinguishing features, we consider
Chlanidota (Pfefferia) to warrant only sub-
generic distinction from Chlanidota sensu
stricto.

Chlanidota (Pfefferia) palliata
(Strebel, 1908)
Figs. 17, 18, 19, 20; Table 7

Pfefferia palliata Strebel, 1908:34-—5, pl. 3,
fiewooca—t Ehiele, 1912:pl. 16, fig. 20
(radula); Powell, 1951:143; Carcelles,
£9532 193.

Pfefferia elata Strebel, 1908:35, pl. 3, fig.
40; Powell, 1951:142, 194, fig. L78 (rad-
ula), 196, fig. N 128 (operculum).

Pfefferia cingulata Strebel, 1908:36, pl. 3,
fig. 42a—c; Powell, 1951:142-3, 194, fig.
L79 (radula); Carcelles, 1953: 193.

Description.—Shell (Fig. 18) large for
genus (to 34.5 mm, Powell, 1951:142),
thick, solid, ovate-rounded. Protoconch,
early teleoconch whorls eroded in all spec-
imens. Teleoconch with up to 4 evenly in-
flated whorls. Shoulder not pronounced.
Suture deep, adpressed, narrowly chan-

a

281

neled. Spiral sculpture of alternating broad,
low cords (10—17 on body whorl, 3—5 on
penultimate whorl) and fine threads (20—24
per 5 mm), spanning entire shell surface.
Cords more clearly visible on young spec-
imens with thinner periostracum. Axial
sculpture limited to indistinct growth lines.
Aperture tall, ovate, deflected from the shell
axis by 16—18°. Siphonal canal is not pro-
nounced. Outer lip evenly rounded, thick,
weakly reflected in adults, very thin in ju-
venile and some adult specimens. Columel-
la <% AL, weakly concave, with fine pus-
tules. Siphonal fold strong. Callus thick,
clearly demarcated, narrowly overlying pa-
rietal region, siphonal fasciole in mature
specimens. Siphonal notch broad, dorsally
reflected, columellar margin straight, aper-
tural margin rounded, reflected, forming
pronounced ridge margin along adapical
edge of fasciole. Shell color grayish white,
translucent. Periostracum very thick, tightly
adherent to shell surface, with evenly
spaced, axially reflected lamellae with
fringed edges evident on early whorls and
juvenile specimens (Fig. 19C). Operculum
(Fig. 16A, D) large, leaf-shaped, with api-
cal nucleus, massive, raised, lamellose bor-
der along posterior margin, 0.67—0.83 AL.
Attachment area spans nearly entire inner
surface, posterior, left margins thickened,
glazed.

Ultrastructure.—(fig. 19E) Outermost
layer prismatic (~50 pum), middle layer
comarginal crossed-lamellar (~175 pm),
inner layer radial crossed-lamellar (~95
1.1m). Inner surface of the outer aperture lip
with numerous, fine spherules (Fig. 17D).

External anatomy.—Body of 242 whorls
(Fig. 18A, B), mantle cavity spanning ~2
whorl, kidney ¥; whorl, digestive 134 whorl.
Foot of preserved specimens short (L/W
~1), with rounded posterior edge. Head

region of the operculum. F Section through “‘feathered”’ portion of operculum (along the line F-F’ on Fig. A),
nucleus at the bottom. G, H. Successive layers are unattached along apical portion of posterior edge of oper-
culum. I. new lamellae are deposited along the entire length of the operculum.

N
(o@)
i)

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ahd

Rhy Senile
Jats ens eet v
ue r r ai
&

|

ah

Fig. 17. Chlanidota (Pfeffeira) palliata (Strebel, 1908). A-C. Lectotype, SMNH (here designated), Sveska
Siidpolarexp., Sta. 22, South Georgia, 54°17’S, 36°28’W, in 75 m. D. Operculum of lectotype, enlarged.

VOLUME 112, NUMBER 2

283

Table 7.—Chlanidota (Pfefferia) palliata (Strebel, 1908). Measurements of shell characters. Linear measure-

ments in mm (n = 10, type material not included).

Lectotype Lectotype Holotype of

of C. of C. C. elata

Character Mean o Range palliata cingulata (broken)
Shell Length (SL) 272k 3.6 18.1-30.0 32.8 25.9 oa yf
Body Whorl Length (BWL) 24.6 3.4 16.4—27.1 292 Da Se
Aperture Length (AL) |e 2.0 14.4-20.4 2a 19.8 ae,
Shell Width (SW) 19.4 2.3 14.0—21.2 23.4 19.4 aa
Operculum Length (OL) 14.1 2.0 9.6—16.7 — 15.4 =

BWL/SL 0.91 0.02 0.86—0.95 0.89 0.92 ~0.86

AL/SL 0.72 0.04 0.67—0.80 O72 0.76 G2

SW/SL O72. 0.03 0.68—0.78 0.71 0.75 ~0.60
OL/AL 0.73 0.06 0.67—0.83 oa 0.78 —

narrow, with stout, conical tentacles, eyes
on small lobes. Body lacks pigmentation.

Mantle cavity.—Mantle edge smooth. Si-
phon medium to long, muscular, extending
substantially beyond mantle edge (~% AL).
Osphradium dark greenish-brown, bipectin-
ate, spans ~% mantle length. Hypobranchi-
al gland of few, distinct, widely spaced
folds. Ctenidium large, wide, spans ~%4
mantle length, lamellae tallest in posterior
% of ctenidium, becoming shorter anterior-
ly.

Alimentary system.—Proboscis (Fig.
19C, D, pr) thick (L/D = 3.5), of moderate
length (0.9 AL). Mouth opening, triangular
slit. Buccal mass muscular, large, nearly
equal to retracted proboscis in length.
Odontophoral cartilages paired, fused an-
teriorly, spanning ~% of proboscis length.
Radular ribbon (Fig. 18A, B) of moderate
length, 10.6 mm (0.66 AL), ~760 wm wide
(0.05 AL), triserial, consisting of 60-65
rows of teeth, posteriormost 5 nascent.
Rachidian teeth with arched base, nearly
straight lateral sides, 3 large, robust cusps
of equal length. Central cusp longer than
flanking cusps in one specimen (Fig. 18A,

—

B). Lateral teeth with 3 cusps, outer cusp
longest, intermediate cusp shortest, adjacent
to inner cusp. Intermediate cusp nearly
fused to inner cusp in one specimen (fig.
18A, arrow). Salivary glands small, fused
(Fig. 19G), situated above nerve ring. Right
salivary gland completely covering valve of
Leiblein. Salivary ducts run loosely along
both sides of oesophagus, entering esopha-
geal wall near posteriormost portion of re-
tracted proboscis. Valve of Leiblein (Fig.
18E, vL) well defined, large, without cili-
ated cone. Gland of Leiblein (Fig. 18C—E,
gL) long, tubular, coiled anteriorly, tapering
posteriorly. Oesophagus wide, muscular,
expanding posteriorly to form crop (Fig.
18I, poe) lined with tall longitudinal folds.
Stomach (Fig. 18H, I) U-shaped, without
caecum. Ducts of digestive gland paired,
closely spaced, transverse fold slightly
raised. Typhlosoles present, poorly defined.
Rectum terminating with well defined anal
papilla.

Female reproductive system.—Typically
buccinoidean. Oviduct opens into medium-
sized albumen gland. Ingesting gland sin-
gle. Capsule gland large, occupies ~'2 of

E. Paralectotype, SMNH, from the type locality. F-H. Lectotype of Pfefferia cingulata (Strebel, 1908) (here
designated), Sveska Siidpolarexp., Sta. 34, Cumberland Bay, South Georgia, 54°11'S, 36°18’W, in 252-310 m.
I. Holotype of Pfefferia elata (broken specimen), same station as types of C. (P.) palliata. J, K, L, both from
R/V Islas Orcadas, Sta. 33, 54°30.7'S, 35°35.9'W, in 261-267 m, USNM 901676. Scale bar = 1 cm for A-C,
E-L, 5 mm for D.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 18. Chlanidota (Pfefferia) palliata (Strebel, 1908). A. Dorsal, and B. left lateral (45°) views of the
central portion of the radular ribbon taken from animal in Fig. 17J. C. Periostracum. D. Edge of outer lip. E, F

Shell ultrastructure. E. Fracture surface parallel to outer lip. EK Fracture surface perpendicular to outer lip. Scale
bars = 200 pm for A, B, 100 wm for C—E

VOLUME 112, NUMBER 2

mantle cavity. Bursa copulatrix present,
simple, hemispherical.

Male reproductive system.—Very similar
to that of C. (C.) densesculpta, penis has
the same size, overall shape, terminal pa-
pilla.

Type locality.—|Pfefferia palliata & P.
elata| South Georgia, 54°17'S, 36°28’W, in
75 m (Sveska Siidpolarexp., Sta. 22); [Pfef-
feria cingulata] Cumberland Bay, South
Georgia, 54°11'S, 36°18'W, in 252-310 m.
(Sveska Siidpolarexp., Sta. 34).

Type material.—|Pfefferia palliata| Lec-
totype (here designated) SMNH (fig. 17A—
C), and 2 paralectotypes SMNH (fig. 17E);
[Pfefferia elata| Holotype, SMNH 3661
(Fig. 171), broken specimen; [Pfefferia cin-
gulata| Lectotype (here designated) (Fig.
16F—H) and juvenile paralectotype, SMNH.

Material examined.—Type material. R/V
Islas Orcadas: Sta. 31, 19 May 1975,
54°05.36'S, 36°30.48’W, 130-143 m, 1
Shell, USNM 887863; Sta. 32, 19 May
1975, 54°21.36’S, 35°58.42'W, 141-164 m,
1 shell, USNM 887872; 9 shells, USNM
887867; Sta. 33, 19 May 1975, 54°30.7'S,
35°35.9'W, 261-267 m, 27 specimens,
USNM 901676.

Published records.—R/V Discovery II,
Sta. 30, West Cumberland Bay, South
Georgia, 2.8 miles S, 24°W of Jason Light
[16 Mar 1926], 251 m (Powell, 1951:142).

Distribution.—Known only from off the
northeastern coast of South Georgia (Fig.
19). Live material was collected between 75
and 310 m.

Remarks.—The type specimens of
Chlanidota (Pfefferia) palliata and C. (P.)
elata were taken from the same dredge
haul. Strebel (1908) distinguished these
taxa primarily on the difference in their
spire height and the strength of their spiral
sculpture, but questioned whether these dif-
ferences merited specific recognition. We
were able to examine a larger sample than
was available to Strebel (USNM 901676, n
= 27) and found it to contain specimens
spanning a morphological continuum be-
tween these two forms. The type series of

285

C. (P.) cingulata was collected very near
the type locality of C. (P.) palliata, but at
slightly greater depth. The specimen select-
ed as lectotype of C. (P.) cingulata is, in
our opinion, conspecific with the specimen
selected as lectotype of C. (P.) palliata, as
both fall within the range of morphological
variation found in a single population
(USNM 901676). Since Powell (1951) des-
ignated P. palliata to be the type species of
Pfefferia, we retain this name for this spe-
cies, and synonymize the remaining nomi-
na.

Powell (1951:142) noted that the only
radula he examined was abnormal, with two
small, “‘incipient’’ cusps flanking the nor-
mally tricuspid rachidian teeth. The radulae
we examined were typical of Chlanidota.

Juvenile specimens are bicolored, with
the periostracum being dark chestnut brown
above the periphery, and a pale, olive green
below the periphery. The periostracum of
adult specimens is thicker and uniformly
chestnut brown in color. Conchologically
this species is similar to C. (C.) dense-
sculpta, but may be readily distinguished on
by the presence of weakly raised spiral
cords, as well as by its distinctive opercu-
lum.

Chlanidota (Pfefferia) chordata
(Strebel, 1908)
Pres: 2122, 23; Table 8

Pfefferia chordata Strebel, 1908:36-—7, pl.
3, fig. 4la—c; Powell, 1951:143; Carcel-
les, 1953:193.

Description.—Shell (Fig. 20) large for
genus (to 35.3 mm), thick, solid, ovate-
rounded, highly variable. Protoconch, early
teleoconch whorls eroded in all specimens.
Teleoconch with up to 4+ evenly rounded,
very convex whorls. Shoulder not pro-
nounced. Body whorl comprises 0.6—0.89
of the total shell length. Suture strongly im-
pressed, forming narrow channel between
adapical spiral cord and previous whorl.
Spiral sculpture variable, spiral cords (7—26
on body whorl, 4—11 on penultimate whorl)

286 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 19. Anatomy of Chlanidota (Pfefferia) palliata (Strebel, 1908). Specimen in fig. 17 J. A. Right and B.
left lateral views of animal removed from shell (SL = 29.7 mm). C. Ventral, D. left lateral, and E. right lateral
views of anterior alimentary system (salivary glands removed to show valve of Leiblein in E). E Juncture of
gland of Leiblein and esophagus. G. Ventral view of fused salivary glands. H. Dorsal view of stomach. I.
Stomach, opened mid-dorsally. Scale bars = 5 mm for A-E, H—I; 2 mm for E G. Abbreviations: ao, anterior

VOLUME 112, NUMBER 2

287

No. of spec.x 3

Fig. 20. Geographic distribution and bathymetric range of Chlanidota (Pfefferia) palliata (Strebel, 1908).

© = type locality; @ = examined material.

low, weak, and smooth to very tall and
sharply-defined, forming corresponding spi-
ral grooves on inner surface of outer lip.
Cords of weakly sculptured specimens (Fig.
20L, M) closely spaced, as wide or wider
than intervening spaces. Cords of moder-
ately sculptured specimens (e.g., Fig. 20A—
C, F-G) % to % width of intervening spac-
es. Cords of strongly sculptured specimens
(Fig. 20D—-E, K—J) more than % the width
of intervening spaces. Adapical cord of
strongly corded specimens may form nar-
rowly concave subsutural rim. Fine spiral
threads may be present on cords and in in-
terspaces. Axial sculpture limited to weakly
defined growth lines. Aperture tall (0.60—
0.84 SL), ovate, deflected from shell axis
by 16—19°. Outer lip thin to moderately
thick, evenly rounded or scalloped, fragile
to strong, sometimes slightly reflected. Col-

a

umella less than half aperture length, weak-
ly convex or straight, with strong siphonal
fold. Callus usually thick, narrow, overlying
parietal region, siphonal fasciole, grayish to
brownish in color. Siphonal notch broad,
dorsally recurved, with straight columellar
and rounded apertural margins that form
borders of fasciole. Ridge margin of fasci-
ole pronounced, originating at apertural
margin of siphonal notch, sometimes pro-
truding through callus. Shell color chalky-
white. Periostracum (Fig. 21) yellow-or-
ange, olive to dark brown, nearly black,
moderately to extremely thick, tightly ad-
herent to the shell surface. Periostracum
surface smooth or of closely adjacent,
blade-like lamellae that overlay spiral
sculpture without giving rise to hair-like
projections, or hairy, producing one or sev-
eral hairs at intersections of lamellae with

aorta; ct, ctenidium; ddg, duct of the digestive gland; dg, digestive gland; dgL, duct of gland of Leiblein; gL,
gland of Leiblein; h, heart; ht, cephalic tentacles; ig, intestinal groove; nep, nephridium; nr, circumoesophageal
nerve ring; oe, oesophagus; op, operculum; os, osphradium; ov, ovary; poe, posterior oesophagus; pr, proboscis;
pir, proboscis retractors; rhd, proboscis sheath; s, siphon; sd, salivary duct; sg, salivary gland; st, stomach; vL,

valve of Leiblein.

288

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 8.—Chlanidota (Pfefferia) chordata (Strebel, 1908). Measurements of shell characters. Linear mea-

surements in mm (n = 10, including holotype).

Character Mean

Shell Length (SL) 2e9
Body Whorl Length (BWL) 21.9
Aperture Length (AL) 197
Shell Width (SW) pd We
Operculum Length (OL) 12.0
BWL/SL 0.79
AL/SL 0.68
SW/SL 0.75
OL/AL 0.61
No. spiral cords on:

Penultimate whorl a)

Body whorl 13

spiral cords. Operculum large (0.46—0.73
AL), massive, typical of genus.

Ultrastructure.—(Fig. 22C, E) Outer-
most prismatic layer comprising spiral
cords, greatly variable in thickness, ranging
from 825 wm in strongly corded specimens
(Fig. C) to 58 pm in lightly corded shells
(Fig. E). Middle, comarginal layer ranges
from 60 to 210 wm, inner, radial layer
reaches ~60 wm in thickness.

Anatomy.—Gross anatomy of C. (P.)
chordata very similar to that of C. (P.) pal-
liata. Radular ribbon long (0.66 AL), ~720
wm wide (0.036 AL), triserial (Fig. 22A—
B), of ~65 rows of teeth, of which 5 are
nascent. Rachidian teeth with deeply arched
base, nearly straight lateral sides, 3 large,
robust cusps of equal-length. Lateral teeth
with 3 cusps, outermost cusp largest, inter-
mediate cusp thinnest and shortest, adjacent
to inner cusp. Stomach U-shaped without
obvious caecum, too poorly preserved to re-
veal internal details.

Type locality——Cumberland Bay, South
Georgia, 54°11’S, 36°18’W, in 252-310 m
(Sveska Siidpolarexp., Sta. 34).

Type material.—Holotype, SMNH 3660

Material examined.—Holotype. R/V Is-
las ‘Oreadas: South ‘Georgia "Sta. 9;
55.435 30'S, 37 30 06"W, 271-393. mer IZ
May 1975, 2 specimens, USNM 901677;
Sta. 18, 54°02'30"S, 37°39'36"W, 60-71 m,
15 May 1975, 3 specimens, USNM 901678;

o Range Holotype
SD) 23.2—34.4 34.7
Sth 16.7—29.7 29.0
age) 15.3—23.2 Zee
3.4 17.2—26.7 DBEG
33 7.7-16.2 —
On 0.60—0.89 0.83
0.05 0.60—0.74 0.62
0.06 0.68—0.86 0.69
0.09 0.46—0.73 —
2.4 4-11 5)
6.3 7-26 12

Sta. 19, 54°01'42”S, 37°40’00"W, 46—70 m,
15 May 1975, 3 specimens + 1 shell,
USNM 901679; Sta. 20, 54°00'06’S,
37°40'36"W, 68-80 m, 15 May 1975, 4
specimens, USNM 901680; Sta. 22,
54°02'48"S, 37°23'42”"W, 66-75 m, 16 May
1975, 3 specimens + 6 shells, USNM
881724; Sta. 25, 53°51'02’S, 36 49: 037;
199-247 m, 17 May 1975, 1 shell, USNM
901681; Sta. 26, 53°43'06"S, 36°49'18’"W,
183-192 m, 17 May 1975, 3 specimens +
3 ‘shells, “USNM “88175627 "Stamwesae
54°31'45"S, 36°48'42”"W, 150—154 m, 7 Jun
1975, 2 specimens + 1 shell, USNM
896287; Sta. 89, 54°44’12”S, 37°11'12’W,
225-265 m, 7 Jun 1975, 6 specimens + 7
shells, USNM 881760; Sta. 90, 54°50’48"S,
37°23" 48" W, 223-227 “m;. 7 Jono
shells, USNM 881762; Sta. 101,
54°14’10"S, 37°54’20"W, 164-183 m, 10
Jun 1975, 1 shell, USNM 901682. R/V
Prof. Siedlecki: South Georgia: Sta. 20,
53°58’S, 38°42’W, 189-200 m, 2 Dec 1986,
1 shell, USNM 901683; Sta. 37, 54°18’S,
37°54'W, 158-194 m, 5 Dec 1986, 1 shell,
USNM 897573; Sta. 83, 54°39'S, 35°49'W,
98-127 m, 11 Dec 1986, 2 specimens + 4
shells, USNM 897534; Sta. 84, 54°28’S,
35°39'W, 231-249 m, 11 Dec 1986, 1 spec-
imen, USNM 897536; Sta. 88, 54°15’S,
35°51’ W,.. 232-254 m, 11 Dec iSsemee
shells, USNM 897506; Sta. 90, 54°10’S,
35°15'W, 242-262 m, 12 Dec 1986, 1 shell,

VOLUME 112, NUMBER 2

USNM 896998; Sta. 91, 54°09’S, 35°55'W,
218-227 m, 12 Dec 1986, 3 specimens,
USNM 897450; Sta. 101, 53°47'S,
36°34'’W, 263-277 m, 2 shells, USNM
897457; Sta. 105, 53°40’S, 36°48’W, 161-
192 m, 14 Dec 1986, 2 specimens + 6
shells, USNM 897515; Sta. 106, 53°44’S,
36°51'W, 178-201 m, 14 Dec 1986, 5
shells, USNM 896985.
Distribution.—Off Southern Georgia
(Fig. 22). Living specimens collected at
depths from 46 to 313 m.
Remarks.—Chlanidota (Pfefferia) chor-
data is highly variable in shell morphology,
with the extremities of the morphological
range appearing very different (compare,
e.g., Fig. 21K, J to L, M). Incrementally
transitional forms ranging from nearly
smooth to highly corded individuals were
present in our samples (e.g., Fig.
2ZIMBLOFOAGDOHS J). Specimens
from depths (<100 m) (e.g., Fig. 21L, M)
tend to have smaller shells with more nu-
merous and more densely spaced spiral
cords than specimens from deeper water
(150 m) (e.¢., Fig. 21D, H, J). Interme-
diate specimens between these extremes oc-
cur within single dredge hauls at both shal-
low and deep stations. Specimens with in-
termediate shell morphology might be con-
fused with Chlanidota (Pfefferia) palliata
(compare Fig. 21F—G with Fig. 17F—G).

Chlanidota (Pfefferia) invenusta,
new species
Figs. 24—26; Table 9

Chlanidota densesculpta (partim)—Dell,
1990:183, fig. 306.

Description.—Shell (Fig. 23) thick, sol-
id, ovate-rounded. Protoconch, 1% upper te-
leoconch whorls eroded. Teleoconch with
~=4 strongly convex whorls. Shoulder
rounded. Suture adpressed, deep, wide,
nearly channeled. Subsutural rim well pro-
nounced, slightly convex. Spiral sculpture
of weak cords (25—40 on body whorl, 5—14
on penultimate whorl) constant in width
adapically, of variable width and promi-

289

nence below shoulder. Axial sculpture lim-
ited to fine, often indistinct growth lines.
Aperture of moderate height (0.60—0.72
SL), elongate-ovate, deflected from the
shell axis by 17—21°. Outer lip thin to thick,
evenly rounded, weakly reflected in thick-
lipped adults. Outer lip thin and evenly
rounded, slightly deflected. Columella short
(<¥% AL), straight, with strong, long si-
phonal fold, siphonal region slightly elon-
gated, inflected in some specimens. Callus
narrow, thin to thick overlying parietal re-
gion, siphonal fasciole, white when thin,
greenish-gray at margins when thick. Si-
phonal notch broad, dorsally recurved, with
straight columellar, rounded apertural mar-
gins forming borders of fasciole. Apertural
margin of siphonal notch gives rise to weak
ridge margin. Shell color grayish white.
Periostracum (Fig. 24D) light-olive, mod-
erately thick, tightly adherent to shell sur-
face, forming very low, densely spaced la-
mellae along the growth lines. Operculum
(Fig. 23D, H, J, L) large (0.47—0.62 AL),
oval, light to dark brown, apical end round-
ed to weakly tapering, nucleus, terminal, ro-
tated nearly perpendicular to long axis of
operculum, posterior edge smooth (Fig.
23d, J), or with weakly (Fig. 23H) to
strongly (Fig. 23L) feathered lamellae.
Ultrastructure.—(Fig. 25C) Outer pris-
matic layer comprises spiral cords, variable
in thickness, ranging from ~80 wm to
~=140 wm, depending on strength of cord.
Middle layer comarginal (~120 wm). Inner
layer radial, reaching ~50 um in thickness,
is the last layer to be deposited, may not be
evident in immature specimens (Fig. 25C).
Anatomy.—The gross anatomy of para-
type 1 was examined, and found to be sim-
ilar in all details to that of C. (P.) palliata
with the exception that the animal lacked
eyes. Radular ribbon (Fig. 24A—B) long,
13.6 mm (0.54 AL), ~760 wm wide (0.030
AL), triserial, consisting of ~80 rows, 6 of
which are nascent. Rachidian teeth with
arched base, straight lateral sides, 3 large,
robust cusps, central cusp slightly larger
that flanking cusps. Small intermediate ser-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 21. Chlanidota (Pfefferia) chordata (Strebel, 1908). A-C. Holotype, SMNH 3660, Sveska Siidpolarexp.,
Sta. 34, Cumberland Bay, South Georgia, 54°11'S, 36°18'W, in 252-310 m. D, E. R/V Islas Orcadas, Sta. 89,
South Georgia, 54°44'12”S, 37°11'12”W, in 225-265 m, USNM 881760. E G. R/V Islas Orcadas, Sta. 9, South
Georgia, 53°43'30"S, 37°30'06’W, in 271-313 m, USNM 901677. H, I. R/V Prof. Siedlecki, Sta. 105, South

VOLUME 112, NUMBER 2

ration present between central cusp and
right flanking cusp of one specimen. Lateral
teeth with 3 cusps, as in genus.

Type locality—South Georgia Island,
53°31'12"S, 37°50'54’W, in 1267-1599 m
(R/V Islas Orcadas, Sta. 7).

Type material.—Holotype, USNM
881782, paratypes 1-9, USNM 880280, all
from the type locality.

Material examined.—South Georgia:
Rayeisias’ Orcadas, Sta. 7, 53°31'12'S,
37°50'54"W, 1267-1599 m, 11 May 1975,
Holotype, USNM 881782, paratypes 1-9,
USNM 880280, 17 shells, USNM 901684;
STa. 27, 53°34.9'S, 36°47.8'W, 448-872 m,
17 May 1975, 1 specimen, USNM 901685;
R/V Eltanin: Sta. 731, 53°35’'S, 36°28'W,
796-824 m, 11 May 1975, 13 specimens,
USNM 896048; Sta. 734, 53°23’S,
37°11'W, 1299-1400 m, 11 May 1975, 4
shells, USNM 870389, 8 specimens,
USNM 896049.

Distribution.—Known only from the
northwestern coast of South Georgia (Fig.
25). Living material was collected at depths
ranging from 448 to 1599 m.

Etymology.—invenustus (Lat.)—unat-
tractive.

Remarks.—Chlanidota (Pfefferia) inven-
usta may be distinguished from the other
species in the subgenus Pfefferia by its
more inflated shell, with a stepped rather
than conical spire, more numerous and finer
spiral cords, a broader, more rounded ap-
erture, and an operculum in which the
feathering along the posterior edge is much
narrower to entirely absent. Some speci-
mens superficially resemble Chlanidota
densesculpta, but may be distinguished on
the basis of their larger operculum, thicker
Shell, more rounded and stepped spires, and
proportionally smaller, more rounded, and
more deflected apertures.

—

291

The nine paratypes collected with the ho-
lotype exhibit some variation in shell out-
line and the number of spiral cords. Smaller
specimens tend to have more rounded
shells, while the number of spiral cords
does not correlate with shell size. Opercula
of the smallest specimens are yellow and
translucent, and clearly show that the ter-
minal nucleus is coiled, as in Neobuccinum.

Species excluded from
Chlanidota sensu lato

Chlanidota smithi Powell, 1958
Fig. 27

Chlanidota smithi Powell, 1958:192, pl. 3,
heres Dell) 1L990s1 77.

Type locality.—Off Enderby Land,
65°50'S, 54°23’E, in 220 m (BANZARE,
Sta. 42).

Material examined.—Holotype, SAM
Dirs47 Liz. 27).

Published records.—BANZARE, Sta.
41, Off Enderby Land, 65°48’S, 53°16’E,
193 m (Powell, 1958:192).

Remarks.—Powell (1958) placed this
species in a group with Chlanidota denses-
culpta, C. vestita, and C. pilosa, but distin-
guished it from these taxa on the basis of
its “‘disproportionally large,’’ bulbous pro-
toconch and the almost smooth surface on
all whorls. Powell (1958) conjectured that
Smith’s (1902) record of C. vestita from
Cape Adare, 24—26 fm [45-48 m] might
represent C. smithi. However, Dell (1990:
fig. 311) illustrated the Cape Adare speci-
men and showed it to represent C. vestita.
Thus, records of C. smithi are restricted to
Enderby Land. This species differs from
typical Chlanidota in number of characters,
most notably its very large size (SL = 51.5
mm) and very large protoconch. Compari-

Georgia, 53°40’S, 36°48’W, in 161-192 m, USNM 897515. J, K. R/V Islas Orcadas, Sta. 26, South Georgia,
53°43'06"S, 36°49'18”W, in 183-192 m, USNM 881756. L, M. Both from R/V Islas Orcadas, Sta. 19, Off South
Georgia Island, 54°01'42”S, 37°40’00’W, in 46-70 m, USNM 901679. Scale bar = 1 cm.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 22. Chlanidota (Pfefferia) chordata (Strebel, 1908). A. Dorsal and B. left lateral (45°) views of the
central portion of the radular ribbon taken from specimen in fig. 21F—G. C. Shell ultrastructure and D. perios-
tracum of the strongly corded shell. R/V Prof. Siedlecki, Sta. 83, South Georgia, 54°39’S, 35°49’W, in 98-127
m, USNM 897534. E. Shell ultrastructure and FE periostracum of the weakly corded shell (fig. 20M). Scale bars
= 200 pm for A, B, E, EF 500 um for C, 1000 wm for D.

VOLUME 112, NUMBER 2

Table 9.—Measurements of shell characters of the type series of Chlanidota (Pfefferia) invenusta, new species.

Paratype Paratype Paratype Paratype Paratype Paratype Paratype Paratype Paratype
1 2 3 4 5) 6 7 8 9 Mean

Holotype

Character

Tie!

39.6 32.6 38.6 31.8 31.0 36.0 2015 2 19.0 31.6

38.3

Shell Length (SL)

Se) PS 30.7 25.8 26.3 30.3 22) 18.7 16.3 26.1 35.5
4.6

31.6

Body Whorl Length (BWL)

Shell Width (SW)

Dales 23.4 25.6 229) 22.0 25.6 17.4 16.0 14.4 Dpes|

26.4

25.0 21.9 DS) 19.0 20.1 22.0 18.2 15.6 13.7 20.3 Sei)

24.6

Aperture Length (AL)

2.0

0.02
0.04
0.03
0.06

8.0 (EP 10.3
0.84
0.70

0.72
0.51

8.8

0.84
0.69
0.67
0.48

ley 10.5

11.7

10.2

12

12.4

Operculum Length (OL)

BWL/SL
AL/SL
SW/SL

0.83
0.64
0.70
0.53

0.86
0.72
0.76
0.53

0.84
0.61
0.71

0.85
0.65
0.71
0.58

0.81
0.60
0.72
0.62

0.80
0.60
0.66

0.84
0.67
0.72
0.47

0.81
0.63
0.69
0.49

0.82
0.64
0.70
0.50

OL/AL

Number of spiral cords

4.5

12

11

12
29

24

19
40

16
DS

14

26

15
32

Penultimate whorl

Body whorl

38 27 29.9 53)

29

297

3++ 3+ 3++ 3+ 3+ 3+ 2++ 2++ 2+

3+

No. whorls

293

son of the holotype of C. smithi (Fig. 27)
with the specimens of Neobuccinum eatoni
Smith, 1875, clearly reveal a striking con-
chological similarity between these species.
Numanami (1996:146) noted that the ma-
jority of specimens of N. eatoni taken off
Syowa Station (Enderby Land) are obese,
with a large, well inflated body whorl.
Comparison of Numanami’s (1996:fig.
94A, B, D—G) illustrations of Neobuccinum
eatoni with the holotype of Chlanidota smi-
thi leave no doubt that these two taxa are
conspecific.

Chlanidota gaini (Lamy, 1910)

Sipho gaini Lamy, 1910:319; Lamy, 1911:
ie phe le hes: 7 —8.

Prosipho? gaini Thiele, 1912:262.

?Chlanidota gaini Powell, 1951:142.

Chlanidota gaini Carcelles, 1953:191; Dell,
1990:177.

Type locality.— Off King George Island,
South Shetlands, in 420 m.

Material examined.—Holotype, NMNH.

Remarks.—*‘Sipho”’ gaini was provi-
sionally attributed to Chlanidota by Powell
(1951) and by Carcelles (1953). Dell (1990)
considered Chlanidota gaini to be a species
of uncertain affinity, known only from its
holotype.

Examination of the holotype, which had
not been illustrated since its description,
clearly indicates that this species is refer-
able to the family Conidae (sensu Taylor et
al., 1993), and is, in fact, the senior syno-
nym of Belaturricula antarctica (Dell,
1990). The composition and relationships
of the genus Belaturricula are the subject
of a separate report (Kantor & Harasewych
1999). Hedley (1916) was the first to sur-
mise the conoidean affinities of Sipho gaini,
suggesting that it was closely related to
Pontiothauma ergata Hedley, 1916.

Chlanidota eltanini Dell, 1990

Chlanidota eltanini Dell, 1990:184-—5S, figs.
2901292)i 2975314:

Figs 23)
© = type locality; @ = examined material.

Type locality.—East of Falklands Islands
(Islas Malvinas), 51°58’S, 56°38'W, depth
845—646 m (R/V Eltanin, Sta. 558).

Type material.—Holotype, USNM
860124, 2 paratypes USNM 860125, 1 par-
atype NMNZ ME56613, from the type lo-
cality. R/V Eltanin: Sta. 557, East of Falk-
land Islands, 51°56’S, 56°39’W, 855-866 m,
2 paratypes USNM 860126; Sta. 1521,
South Atlantic Ocean, 54°09’S, 52°08'W,
419—483 m, 2 paratypes USNM 860127, 1
paratype NMNZ ME56614.

Remarks.—Dell (1990) described Chlan-
idota eltanini, C. bisculpta and C. polys-
peira from the Magellanic Province. While
these three species are likely congeneric,
they differ significantly from species as-
signed to the subgenera Chlanidota and
Pfefferia. Conchologically, these species
are readily distinguished by their small (15—
16 mm), slender, thick shell, very thin,
smooth periostracum, and large (0.6 AL),
coiled operculum. We were able to examine
the anatomy of Chlanidota bisculpta, as
well as of a closely related but as yet un-
described species, and found differences in

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

No. of spec.x 4

Geographic distribution and bathymetric range of Chlanidota (Pfefferia) chordata (Strebel, 1908).

radular morphology, stomach shape, size of
gland of Leiblein, and penis morphology.
We provisionally consider these species to
be more closely related to Neobuccinum,
and exclude them from Chlanidota.

A revision of this group will be pub-
lished separately.

Chlanidota bisculpta Dell, 1990

Chlanidota bisculpta Dell, 1990:185, figs.
DON, 294, 2955, sl

Type locality—Burdwood Bank, 53°08’S,
59°23'W, in 578—567 m (R/V Eltanin, Sta.
340).

Type material.—Holotype, USNM
860128; R/V Eltanin: Sta. 557, East of
Falkland Islands (Islas Malvinas), 51°56’S,
56°39'W, 855-866 m, 6 paratypes USNM
860129, 1 paratype NMNZ ME56615; Sta.
740, off Cape Horn, 56°06'S, 66°19'W,
384—494 m, 2 paratypes USNM 860130, 1
paratype NMNZ ME56616.

Remarks.—See remarks under Chlani-
dota eltanini Dell, 1990.

VOLUME 112, NUMBER 2 295

Fig. 24. Chlanidota (Pfefferia) invenusta new species. A—C. holotype, USNM 881782, R/V Islas Orcadas,
Sta. 7, South Georgia, 53°31'12”S, 37°50'54’W, in 1267-1599 m. D. Operculum of holotype, enlarged. E.
Paratype 1, (anatomy studied), and F paratype 5, USNM 880280, both from type locality. G, H. Specimen with
narrowly rimmed operculum. I, J, M. Specimens with smooth operculum. K, L. Specimen with broadly rimmed
operculum, all from R/V Eltanin, Sta. 734, South Georgia, 53°23’S, 37°11'W, in 1299-1400 m, USNM 896049.
Scale bar = 1 cm for shells, 5 mm for opercula.

296

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Pig.-25:
portion of the radular ribbon of paratype 1 (Fig. 24E). C. Shell ultrastructure, and D. periostracum of the paratype
5 (Fig. 24F). Scale bar = 200 um for A, B, 100 wm for C, 500 wm for D.

Chlanidota polyspeira Dell, 1990

Chlanidota polyspeira Dell, 1990:186, figs.
297 D093, S03,

Type _locality.—Patagonian Shelf,
54°04'S, 63°35’W, depth 247-293 m (R/V
Eltanin Sta. 369).

Type material—Holotype, USNM 860131,
3 paratypes, USNM 860132, 1 paratype
NMNZ ME56617, all from the type local-
ity.

Remarks.—See remarks to Chlanidota
eltanini Dell, 1990.

Chlanidota (Pfefferia) invenusta, new species. A. Dorsal and B. left lateral (45°) views of the central

Discussion

Buccinoidean classification has tradition-
ally been based on a combination of shell,
opercular, and radular characters, and these
structures are well documented for the ma-
jority of Antarctic genera (e.g., Thiele
1904, Powell 1951, Hain 1989, Dell 1990,
Numanami 1996). Ponder (1974) reported
that most organ systems were weakly dif-
ferentiated among buccinoidean higher
taxa, and that it was difficult to identify an-
atomical features that could be used to seg-

VOLUME 112, NUMBER 2 2ST

—
S
fo)

Depth, m
or
=)

No. of spec.x5

Fig. 26. Geographic distribution and bathymetric range of Chlanidota (Pfefferia) invenusta, new species. ©
= type locality, @ = examined material.

regate them reliably. More recently, Hara- gland, that vary among buccinoideans and
sewych (1990) identified several characters, may be phylogenetically informative. While
including the morphology of the gland of there have been a substantial number of an-
Leiblein and the presence of an ingesting atomical investigations of boreal (e.g., Da-

Fig. 27. Chlanidota smithi Powell, 1958. Holotype, SAM D15471, BANZARE Sta. 42, Off Enderby Land,
65°50’'S, 54°23’E, in 220 m. Scale bar = 1 cm.

298

kin 1912; Lus 1978, 1981, 1989; Kantor
1988, 1990), tropical (e.g., Marcus & Mar-
cus 1962, 1964) and deep-sea (e.g., Ponder
1968, Harasewych 1990) buccinoidean
taxa, the anatomy of Antarctic representa-
tives has not previously been studied in any
detail.

It is not surprising that Chlanidota shares
many anatomical features with the boreal
family Buccinidae, among them similar ar-
rangements of the mantle cavity organs, re-
productive system, and many characters of
the digestive system. Characters such as a
long, thick proboscis and fused salivary
glands occur in Chlanidota as well as in the
buccinid taxa Habevolutopsius (Kantor
1990) and Ancistrolepidinae (Kantor 1988).
The stomach of Chlanidota, however, dif-
fers from that of most Buccinidae in being
broadly U-shaped and in lacking a caecum.
The caecum, or posterior mixing area, is
usually well developed in boreal Buccini-
dae, and is sometimes very large (e.g., in
Volutopsius, subfamily Volutopsiinae),
making the stomach appear sac-like. The
stomach of Southern Hemisphere taxa, such
as Penion adustus (Philippi, 1845) (Ponder
1973) and Ratifusus mestayerae (Iredale
1915) (Ponder 1968), which had been as-
signed to Buccinulidae by Powell (1961),
also lack a caecum.

The northern Atlantic buccinid Colus
gracilis (da Costa 1778) is unusual in lack-
ing a gastric caecum (Smith 1967). How-
ever, its stomach differs from the stomach
of Chlanidota in having a cuticularized
shield and a distinct sphincter between the
oesophagus and stomach. The stomach ep-
ithelium of Colus gracilis is glandular and
darker than the esophageal epithelium,
while in Chlanidota the epithelial lining of
the esophagus is dark and that of the stom-
ach is lighter.

Unlike all other Chlanidota and Pfeffer-
ia, the stomach of Chlanidota (Chlanidota)
paucispiralis has a small enlargement (Fig.
14, pm) corresponding in position to the
caecum of Neptunea (Smith 1967), Buccin-
um (Medinskaya 1993) and other boreal

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

buccinids. It is still not clear, whether the
presence of a gastric caecum is an advanced
or plesiomorphic character, as the structure
occurs in many neogastropod families (e.g.,
Buccinidae, Volutomitridae, Muricidae, Mi-
tridae) but is absent in others (e.g., Col-
umbellidae, Volutidae, Cancellariidae and
all Conoidea).

The posterior oesophagus of Chlanidota
is greatly enlarged, thick-walled, and lined
with tall epithelial folds (e.g. Fig. 5H),
which are darker than the rest of esophageal
epithelium. The transition between the pos-
terior oesophagus and the stomach is clear-
ly evident on dissection, as it is marked by
a color change of the epithelium near the
posterior duct of the digestive gland.

The morphology of the radula and major
organ systems is remarkably invariant in all
of the species of Chlanidota and Pfefferia
that we examined. The only exceptions
were the presence of a small caecum in C.
(C.) paucispiralis and of separate rather
than fused salivary glands in C. (C.) Sig-
neyana. Preliminary dissections of other
Antarctic buccinulids generally regarded as
being closely related to Chlanidota (e.g.,
Neobuccinum eatoni, “‘Chlanidota” bis-
culpta, see above) revealed significant dif-
ferences in their alimentary and male repro-
ductive systems, providing support for re-
ducing Pfefferia to a subgenus of Chlani-
dota.

The morphology of the operculum of
Chlanidota (Pfefferia) is both striking and
unique within Neogastropoda. The mecha-
nism by which the characteristic “feath-
ered’’ margin is formed along the posterior
edge of the operculum is readily inferred.
Newly secreted layers of conchiolin are
broader adapically than abapically, and are
attached to the posterior edge of the oper-
culum only along their proximal margins.
While operculum morphology is constant in
C. (P.) palliata and C. (P.) chordata, it
varies considerably in the newly described
C. (P.) invenusta. Juveniles of C. (P.) in-
venusta have an operculum with a “‘feath-
ered’? margin. The width of this margin de-

VOLUME 112, NUMBER 2

creases with increasing shell size, the mar-
gin disappearing entirely in large specimens
(see Fig. 24L—H-J—D). The size of the
operculum of C. (P.) invenusta (0.47—0.62
AL) is larger than in any species of Chlan-
idota (Chlanidota) (0.18—0.37 AL), com-
parable to that of C. (P.) chordata (0.46—
0.73 AL), but smaller than that of C. (P.)
elata (0.68—0.78 AL). The large operculum
of Chlanidota (Pfefferia) allows it to seal
the aperture tightly when the animal with-
draws into its shell (e.g., Figs. 17F 21H),
while its “‘feathered”’ posterior margin pro-
vides a flexible, tight seal against the outer
lip. In contrast, the small size of the oper-
culum in Chlanidota s.s. precludes its util-
ity for sealing the aperture. The decrease in
relative size and “‘degree of feathering”’ of
the operculum in C. (P.) invenusta with in-
creasing shell size, suggests that the ability
to seal the aperture tightly diminishes in im-
portance as the animal grows.

While the anatomy and radular morphol-
ogy are largely invariant in Chlanidota and
Pfefferia, shell shape and the strength and
number of spiral cords and threads vary
considerably. The limited number of spec-
imens available to previous workers has, in
a few instances, led to the description of
species based on differences that now ap-
pear to fall within a broad continuum of
morphological variability revealed by larger
sample sizes spanning wider geographical
areas.

As restricted in this paper, the genus
Chlanidota is confined to the Antarctic re-
gion, with only the type species ranging
slightly beyond the Antarctic Convergence.
The present revision reduces the number of
recognized species within Chlanidota. Nev-
ertheless, it remains the second most di-
verse buccinoidean genus (following Pro-
sipho) in Antarctic waters.

Of the eight recognized species in the ge-
nus Chlanidota, only C. (C.) signeyana has
a broad circum-Antarctic distribution (al-
though records from the poorly sampled
Ross Quadrant are lacking). Except for a
single record of C. (C.) vestita from Cape

299

Adare, all remaining species of Chlanidota
are restricted to subAntarctic islands, with
two species of Chlanidota sensu stricto, and
all three species of Pfefferia endemic to
South Georgia. Of the species inhabiting
subAntarctic islands, most are sublittoral,
and only Chlanidota (Pfefferia) invenusta is
restricted to bathyal depths. In contrast, the
circum-Antarctic C. signeyana has a broad
bathymetric range (30-1116 m).

The protoconchs of all specimens of
Chlanidota were severely eroded, hamper-
ing inference of developmental mode.
However, most polar species, including
buccinoideans, develop directly without a
pelagic larval stage (Thorson 1946). Thus,
it seems likely that the island populations
of Chlanidota are vicariant isolates from a
more broadly ranging species.

Acknowledgments

We wish to express our thanks to Dr. An-
ders Warén for loan of the Strebel type ma-
terial, to Dr. Philippe Bouchet for access to
Lamy’s types, and to Kathie Way and Joan
Pickering for the loan of Powell’s types at
BMNH. Dr. W. Ziedler South Australian
Museum made available the type specimen
of C. smithi. We are grateful to Ellen Strong
for translation of the relevant portions of
Strebel (1908), and to Dr. Guido Pastorino
for his assistance at various stages of man-
uscript preparation, especially with digital
photography. We also thank Susann Braden
for her assistance with scanning electron
microscopy.

The junior author gratefully acknowledges
Mrs. Mabs Mango for her kind hospitality
during the preparation of this manuscript.

This research was supported by a grant
from the NSF-USAP United States Antarc-
tic Program [Contract No. OPP-9509761].

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(2):303—312. 1999.

Hydroid and medusa stages of the new species Ectopleura obypa
(Cnidaria: Hydrozoa: Tubulartidae) from Brazil

Alvaro E. Migotto and Antonio C. Marques

(AEM) Centro de Biologia Marinha, Universidade de Sao Paulo, Caixa Postal 83,
11600-970 Sao Sebastiao, SP, Brazil;
(ACM) Departamento de Biologia, Faculdade de Filosofia, Ciéncias e Letras de Ribeirao Preto,
Universidade de Sao Paulo, Av. Bandeirantes 3900, 14040-901 Ribeirao Preto, SP, Brazil

Abstract.—Ectopleura obypa, a new species referable to the family Tubu-
lariidae, is described from the southeastern coast of Brazil. Specimens were
collected on experimental panels and other artificial substrates at depths from
about 1 to 15 m. The hydroid stage is a remarkable green color when blasto-
styles are developed. A free medusa, with 2 opposed capitate marginal tentacles
and 8 meridional tracks of nematocysts on the exumbrella, is present in the

life cycle.

During a study of hydroid recruitment on
experimental plates in waters of southeast-
ern Brazil we discovered a species of Ec-
topleura, not yet described, remarkable for
its bright green color. Several other speci-
mens of the species were found at various
times later on other artificial substrates in-
cluding nylon ropes, buoys, and iron pipes.
Specimens kept in the laboratory and cul-
tured under controlled conditions liberated
free medusae.

The genus Ectopleura L. Agassiz, 1862,
a basal clade of the family Tubulariidae, has
20 valid species divided in two phyloge-
netic groups distinguishable basically by
their cnidomes and stolon growth pattern
(Petersen 1990:160). There is much confu-
sion between the genera Tubularia and Ec-
topleura, the first having sessile gonophores
and the latter traditionally considered as
having free medusae (e.g., Millard 1975:
31-32; Bouillon 1985:112). However, Pe-
tersen (1979:120; 1990:160) redefined the
tubulariid genera based on features of the
hydroid stage. His classification of tubular-
iid polyps included: the solitary Zyzzyzus,
with a thin perisarc and tuber-like attach-
ment of hydrocaulus; the solitary Tubularia
with firm perisarc and producing either eu-

medusoid or cryptomedusoid gonophores
of the symmetrical type; the solitary Hy-
bocodon, producing fixed asymmetrical
gonophores; the colonial or solitary Ecto-
pleura with firm perisarc and producing ei-
ther free medusae or fixed gonophores; the
solitary or colonial Ralpharia with firm
perisarc and producing reduced medusae
with an internal raised collar around bell
opening; and the solitary Bouillonia with
barrel-shaped hydranth, firm perisarc and
reduced medusae.

Material and Methods

Hydroids of Ectopleura obypa, new spe-
cies, were collected on a number of sub-
strates (experimental plates, ropes, buoys,
iron pipes) on the coast of Sao Sebastiao,
Sao Paulo State, Brazil. Type material,
studied here, was collected by skin and
SCUBA diving from a light buoy in the Sao
Sebastiao Channel. Part of the material was
anesthetized in a 1:1 solution of 7.5%
MgCl, solution and seawater, and preserved
in 4% formaldehyde solution in seawater;
part was kept alive in the laboratory. Live
hydranths were examined under a stereo-
microscope, and newly liberated medusae

304

were transferred to finger bowls and kept in
constant temperature chambers at 19, 24
and 26°C with a 12 h light/12 h dark pho-
toperiod. Water in the bowls was changed
daily, shortly after medusae were measured
and fed with Artemia nauplii.

Stems for scanning electron microscopy
(SEM) were post-fixed in 1% OsQO,, dehy-
drated in a graded series of ethanol, dried
in a critical-point drier, and sputter-coated
with gold. Nematocyst types and their dis-
tributions were determined using a light mi-
croscope with interference-contrast. Nema-
tocyst nomenclature used here is that of
Mariscal (1974) and Millard (1975). Only
capsules of undischarged nematocysts were
measured. Abbreviations are: AM (collec-
tion of A. Migotto); MZUSP (Museu de
Zoologia da Universidade de Sao Paulo,
Sao Paulo); USNM (United States National
Museum, Smithsonian Institution, Washing-
ton D.C.).

Results and Discussion

Genus Ectopleura L. Agassiz, 1862
Ectopleura obypa, new species
Figs. 1-3; Tables 1—2

Material.—Holotype, 1 hydroid colony,
2.0 cm high, Sao Sebastiao Channel, Sao
Sebastido, Sao Paulo State, Brazil
(23°49.86'S, 045°25.28'W), 4 Jan 1996, 5
m, colony with gonophores and several hy-
dranths growing epibiotically on barnacles
and mussels, on light buoy, collected by
hand, coll. A. E. Migotto (MZUSP-12.813);
paratypes (all from same locality as holo-
type), medusae liberated in the laboratory 6
Jan 1996 from hydroid colony collected 4
Jan 1996, 6 medusae ca. 2 h old (AM1325);
2 medusae, 6 d old (MZUSP-12.814); 4
medusae, 7 d old (AM1327); 26 medusae,
34 d old (AM1328); one hydroid colony 35
mm high, on mussels and ascidian, with
gonophores, 16 Jan 1996, 2 m, coll. A. C.
Morandini (AM1323); one hydroid colony
25 mm high, on mussel, with gonophores,
17 Jun 1996, 2-3 m, coll. A. E. Migotto
(AM1324; USNM 99449); 84 medusae ca.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

3 h old, liberated in the laboratory on the
same day of collection, from colony col-
lected 17 Jun 1996 (AM1329).

Description.—colony up to 35 mm high,
arising from branched stolons. Hydrocaulus
unbranched, increasing in width from base
to distal end, covered with a smooth light
brown perisarc without annuli. Neck region
long (Figs. 1C, 3A), flexible and contrac-
tile, covered by a filmy perisarc; extended
neck up to 3.5 times longer than contracted;
in live specimens, neck entirely white or
light pink, with white pigments concentrat-
ed in 2 narrow longitudinal bands. Hy-
dranth vasiform, pink-white and transpar-
ent, with one whorl of 16—30 aboral tenta-
cles and one whorl of 14—29 oral tentacles
(Figs. 1A, 2A, 3A). Aboral tentacles long,
filiform, laterally flattened (Fig. 1A), with
nematocysts concentrated along adoral sur-
face, and on tip. Oral tentacles short, cir-
cular in cross section, basal part adnate to
hypostome (Fig. 1D), with nematocysts
concentrated in a terminal capitulum and 1—
3 irregular swellings on the adoral surface
of the tentacle (Fig. 1B, D); free part of oral
tentacles slightly fused at base (Fig. 1B).
Medusa buds on 4—14 (usually 7—10) blas-
tostyles arising above aboral tentacles; old-
er medusa buds at ends of blastostyles (Fig.
3B). Blastostyles yellow-green, composed
of a main stem on which medusae arise ei-
ther directly or from irregular branches;
main stem may bifurcate. In large speci-
mens the developed blastostyles hang be-
tween aboral tentacles.

Newly-liberated medusa with thin umbrel-
la, dome-shaped and without a distinct apical
projection but with a small apical canal, with
8 slightly raised exumbrellar, meridional
nematocyst tracks; with 4 marginal bulbs, 2
of which with opposed tentacles; marginal
tentacles with a terminal knob and 1-3 ab-
axial nematocyst clusters (Fig. 2B); abaxial
nematocyst cluster grows and involves the
tentacle (Figs. 2D, 3C). Manubrium tubular,
without lips, with nematocysts around mouth,
and about *% the length of bell cavity. Apical

VOLUME 112, NUMBER 2

Fig. 1. Scanning electron micrographs of Ectopleura obypa, new species. A, fronto-lateral view of hydranth,
note incipient blastostyles just above aboral tentacles; B, oral view of hydranth; C, neck region; the arrow points
to the groove where filmy perisarc is secreted; D, lateral view of hypostome and oral tentacles. Scale bar, A, B

= 100 pm; C, D = 50 pum.

canal short. Color: umbrella transparent;
subumbrella green when illuminated side-
ways; manubrium pink-white; marginal bulbs
milky-white with small red dots.

The types and measurements of nema-
tocysts of hydranth and medusa are listed
below. Morphological character variation of
many colonies and newly released medusae
are in Tables 1 and 2, respectively.

Nematocysts.—(Qn ym) (Fig. 3D)
Hydranth
Oral tentacle:
Scho. 4 foo ee A
13.0 (abundant)
Srcnmtele: "Oa = SS Xx 6:0 — 75
(abundant)
Microbasic eurytele — 9.0 — 10.0 X
4.0 — 4.5

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Photographs of live specimens of Ectopleura obypa, new species. A, lateral view of a hydranth with
incipient blastostyles: B, lateral view of a newly-liberated medusa; C. lateral view of a 6-day-old medusa; D,
detail of a tentacle of a 7-day-old medusa. Scale bar, A = 1.0 mm; B, D = 100 pm; C = 200 um.

VOLUME 112, NUMBER 2 307

A B ioe

Fig. 3. Ectopleura obypa, new species. A, outline diagram of hydranth, traced from a whole mount prepa-
ration; the upper arrow points to the region where the film perisarc is secreted; the lower arrow indicates the
beginning of the firm perisarc; B, lateral view of a blastostyle with several well developed medusa buds; C.
sequence in the development of a medusa tentacle; C4 is the oldest stage, just before liberation from blastostyle;
D. nematocysts; D1, stenotele; D2, basitrichous isorhiza; D3, microbasic mastigophore; D4, desmonome. Scale
bar, A = 0.5 mm; B = 100 pm; C = 50 pm; D = 10 um.

308

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Morphological character variation of hydroids of Ectopleura obypa, new species.

Hydrocauli
Length (mm)
Distal diameter (mm)
Proximal diameter (mm)
Hydranth
Total length (mm)

Length from the base of aboral tentacles to the apex of hypostome (mm)

Diameter at the level of aboral tentacles (mm)
Tentacles
Aboral
Number
Length (mm)
Oral
Number
Length (mm)
Gonosome

Number
Length (mm)

Aboral tentacle:

Stenotele = 1975 7——- 1450 = On
13.0 (rare)

Stenofele = 6.0%— 785% 6.0 7-5
(abundant)

Microbasic eurytele — 9.0 — 10.0 X
4.0 — 4.5 (abundant; more at tip)
Desmoneme —-4.0"— 507355 40
(abundant)
Newly released medusa

Basitrichous isorhiza — 8.0 — 10.5 X
S.0:= 90

Stenotele —-3:0'>. 850670) 35)

Microbasic eurytele:— 9.0 —10. >" <
4.0 — 4.5

Desmoneme — 6.0 X 4.0

Etymology.—The species name is de-
rived from the word “‘obypa’’, in the lan-
guage of the Brazilian Tupi natives, mean-
ing green, the color of the blastostyles. It is
pronounced ‘‘6-bi-pa’’.

Development of medusa.—Medusae kept
in the laboratory, despite several different
combinations of temperature, salinity, type
of food and volume of culturing water, grew
slowly and did not change significantly in

Measurements
[average + SD (range)]

18.5 + 6.9 (0.87-30.3)
0.20 + 0.03 (0.13-0.26)
0.34 + 0.07 (0.22—0.60)

1.85 = 0.55 (118-3230)
1.41 = 0.42 (0.88—2.70)
1.09 = 0.30 (0.62—1.80)

23.8 + 2.9 (16-30)
2.50 + 0.59 (0.91—3.90)

20.8 = 3.25 (14-29)
0.79 + 0.16 (0.42—1.20)
8.55 + 2.10 (4-14)

0.91 + 0.55 (0.15—2.10)

morphology. Medusae usually did not cap-
ture food by themselves. Ingestion occurred
only when the offered nauplius was touched
to the manubrium; we never saw the trans-
fer process of the prey from the tentacles to
the mouth of the medusa. From the several
cultures started only one batch had medusae
with gonads, although even these were not
completely developed (medusae 25 days
old, from colony collected 17 Jun 1996,
kept at 19°C). We are aware that these un-
successful efforts probably indicate the cul-
tivated medusae do not represent the spe-
cies under natural conditions, but we decid-
ed to include our observations on growth
rate and morphology. Developing gonads
were first observed in 7 to 9 d old medusae.
These medusae were 1380—1440 pum high,
1050-1150 ym in maximum diameter and
had tentacles with 6 knobs, attaining ca.
800 ym in length (Fig. 2C, D). They lived
16 more days, maintaining approximately
the same size and general features except
for a slight growth of the gonads, which
bulged a little around the middle of the ma-
nubrium. Tentacles developed up to 7 nem-

VOLUME 112, NUMBER 2

309

Table 2.—Morphological character variation of newly-released medusae of Ectopleura obypa, new species,

from three different colonies.

Height (wm)

Maximum diameter (wm)
Diameter at base (wm)
Diameter of aperture (wm)
Length of manubrium (wm)

atocyst knobs. The number of tentacles did
not change in any specimens during
growth; nor was there any indication that
the two other perradial bulbs would give
rise to tentacles.

Remarks.—This description of Ectopleu-
ra obypa, new species, especially color, size
and general shape, is based mainly on live
material. Colors in fixed material disap-
peared. Some anesthetized and fixed hy-
dranths, although well preserved, did not
keep their natural shape: hydranths from
AM1324 are inflated and most have the up-
per part of the neck dilated. Fixed medusae
usually have longer manubria than live
specimens, and perradial and interradial
grooves appear in the umbrella (as also ob-
served for E. sacculifera, see Brinckmann-
Voss 1970:28).

The long, flexible and contractile neck
enables the hydranth to bend in any direc-
tion. In still water under laboratory condi-
tions, the hydranths performed regular cir-
cular movements interspersed with strong
contractions of the neck. We could not as-
certain the number of ridges in the endo-
derm of hydrocaulus, but we suppose the
two bands of white pigments present along
the neck indicate the existence of two of
these ridges in E. obypa. The blastostyles
were so vivid a green, even early in devel-
opment, that when fully developed the
whole hydranth appeared green to the na-
ked eye. The medusa started pulsating
hours before liberation, already capable of
catching and ingesting food, and the ma-
nubrium moved peristaltically and vermic-
ularly.

[average + SD (range)]

594.5 + 101.9 (460-780)

516.0 + 82.1 (396-660)

344.5 + 69.6 (240-440)
AZO

510.2 + 141.7 (276-780)

Besides E. obypa, new species, two other
species of Ectopleura were recorded from
the region of Sao Sebastiao: Ectopleura du-
mortieri (Van Beneden, 1844) and Ecto-
pleura warreni (Ewer, 1953) (see Migotto
& Silveira 1987:100—103, Migotto 1996:
24-25). The polyp of Ectopleura dumorti-
eri is solitary, and has free medusae with 4
marginal tentacles; E. warreni has fixed
gonophores.

Ectopleura obypa is referred to a large
group of species diagnosed by the presence
of free medusae with 8 meridional nema-
tocyst tracks in the umbrella, issuing in
pairs from tentacle bulbs (Schuchert 1996:
107). Complementary characters are: even-
ly rounded umbrella; four radial canals and
tentacle bulbs; manubrium short, not ex-
tending beyond bell margin; medusa tenta-
cles moniliform or with abaxial nematocyst
clusters (Schuchert 1996:107). The cni-
dome of species having free medusae is
characterized by microbasic euryteles (be-
sides other types such as stenoteles and des-
monemes), a type of nematocyst not present
in the species of Ectopleura with fixed gon-
ophores (see Petersen 1990:160; note that
the author interchanged the distribution of
anisorhiza and microbasic euryteles in fig-
ure 19, characters 2 and 6).

Among species of Ectopleura with free
medusae, two subgroups are clearly recog-
nized on the basis of the number of mar-
ginal tentacles in the medusa: 2 tentacles
and 4 tentacles.

Species described as having 4 perradial
tentacles are E. americana Petersen, 1990,
E. dumortieri (Van Beneden, 1844), E. be-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

310

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VOLUME 112, NUMBER 2

thae (Warren 1908) and E. indica Petersen,
1990.

Besides E. obypa there are six species of
Ectopleura having medusae with 2 opposed
tentacles: Ectopleura wrighti Petersen,
1979, E. pacifica Thornely, 1900, E. mayeri
Petersen, 1990, E. minerva Mayer, 1900, E.
sacculifera Kramp, 1957, and E. xiamenesis
Zhang & Lin, 1984 (see Table 3). The last
three of these are known only from the me-
dusa stage. Ectopleura sacculifera and E.
xiamenensis do not have an apical canal,
contrasting in this with E. minerva and E.
obypa. Moreover, Ectopleura sacculifera is
clearly distinct from the rest by having a
thick umbrella and pendent gonads on the
manubrium wall (Kramp 1957:7, plate 3,
figs. 1—3). Neither an apical canal nor an
apical projection were mentioned by
Brinckmann-Voss (1970:25—27) in the de-
Scription of young and adult medusae of E.
wrighti (as E. larynx), and we suppose they
are not present in this species. Nematocyst
clusters of the marginal tentacles in medu-
sae of E. obypa are clearly diffrent from
those of E. minerva Mayer, 1900 (Mayer
1900, fig. 125) and E. sacculifera (cf.
Brinckmann-Voss 1970:28, fig. 29.1),
which are exclusively abaxial and do not
involve the tentacle as in E. obypa (exclu-
sively abaxial clusters of nematocysts are
also present in an unidentified species of
Ectopleura with a two-tentacled medusa de-
scribed by Schuchert 1996:112, fig. 67).

It is difficult to link the species known
only by the medusoid stage with polypoid
stages already described in the literature,
due to the lack of life cycle studies. Except
for E. wrighti (see Brinckmann-Voss 1970:
25, as E. larynx) and E. obypa, data on me-
dusa stages are few and based on immature
specimens still attached to blastostyles. Ec-
topleura wrighti is the only species in
which medusae obtained from hydroid col-
onies were raised through maturity (Brinck-
mann-Voss 1970:25). Petersen (1990:166)
described Ectopleura mayeri (=E. pacifica
of Calder 1988:53-—55) as a “‘new species’’,
even though he acknowledged the possibil-

311

ity of its being the hydroid stage of E. mi-
nerva Mayer, 1900, because the medusa of
the first “has not been reared to a stage
where it can be identified with certainty”’.
Hydroid stages of other species with two-
tentacled medusae have many similar fea-
tures: general size, number of oral and ab-
oral tentacles, and number of blastostyles.
They all have 2 endodermal ridges in the
hydrocaulus, but this character is shared
with other species having different numbers
of tentacles. The aboral tentacles of E. oby-
pa and E. pacifica are slightly adnate to the
hypostome, in which they differ from E.
wrighti and E. mayeri. Ectopleura obypa
differs from E. pacifica in the morphology
of its oral tentacles (see Table 3), and is
unique in having flattened aboral tentacles.

Acknowledgements

We thank the staff of the Laboratério de
Microscopia Eletr6nica and of the Depar-
tamento de Zoologia, Instituto de Biocién-
cias, USP, for assistance with scanning elec-
tron microscopy. We also thank the two re-
viewers (Dr. Dale Calder and Dr. Paul E S.
Cornelius) whose comments improved the
manuscript. This work was supported by
the Fundacgao de Amparo a Pesquisa do Es-
tado de Sao Paulo (FAPESP grants n° 95/
3022-5; 96/10544-0 and 97/04572-4).

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112(2):313-—318. 1999.

Striatodoma dorothea (Cheilostomatida: Tessaradomidae), a new
genus and species of bryozoan from deep water off California

Judith E. Winston and Stace E. Beaulieu

(JEW) Virginia Museum of Natural History, Martinsville, Virginia 24112, U.S.A.;
(SEB) Marine Biology Research Division, Scripps Institution of Oceanography,
La Jolla, California 92093-0202, U.S.A.

(Current address) Applied Ocean Physics and Engineering Department,
Woods Hole Oceanographic Institution, Woods Hole, Massachusetts 02543, U.S.A.

Abstract—Striatodoma dorothea, a new genus and species of cheilostomate
bryozoan, is described from material found attached to hexactinellid sponges
and pogonophoran tubes at an abyssal station (4100 m depth) off central Cal-
ifornia. Members of this new genus can be distinguished from other members
of the family Tessaradomidae by the presence of biserial, rather than quadris-
erial branches, and a peristomial sinus, rather than an enclosed spiramen. Two
other Pacific species, Diplonotos striatum Canu & Bassler, 1930, and Tessar-
adoma bifax Cheetham, 1972, are transferred to Striatodoma.

Although bryozoans have been identified
from deep-sea stations down to 8300 m, re-
views by Schopf (1969) and Hayward
(1981) have shown that only a tiny portion
of the deep ocean floor has been sampled
for the group. Nothing is known of the
deep-sea bryozoan fauna of the eastern Pa-
cific with the exception of three species
found at two stations in the eastern Pacific
between Acapulco and Panama during the
Galathea Expedition (Hayward 1981). Re-
cently, as part of a study by Beaulieu
(1998) of hard substrate epifauna at abyssal
depths off California, an attempt was made
to identify all taxa attached to biogenic
structures that protruded from the soft sed-
iment of the sea floor. Individual stalks of
the hexactinellid sponge Hyalonema sp. and
individual tubes of the pogonophoran Uni-
brachium sp. were sampled in tube cores
using the submersible Alvin at 4100 m
depth. Two of the approximately 140 spe-
cies found attached to the sponge stalks and
pogonophoran tubes were bryozoans. One
is a ctenostome, Arachnidium hippothooi-
des Hincks, 1862. The other is an undes-
cribed genus and species of deep-sea chei-

lostome which we name and describe be-
low.

Tessaradomidae Jullien, 1903
Striatodoma, new genus

Diagnosis.—Tessaradomidae character-
ized by subcylindrical, proximally thick-
ened branches with two series of zooids,
longitudinally striated calcification, a spi-
ramen in close association with zooid peri-
stome, rows of marginal pores, some of
them replaced by oval avicularia, and a
subglobular imperforate ovicell. Striatodo-
ma differs from Tessaradoma in possessing
branches made up of two, rather than four
series of zooids, and in confluence of the
Spiramen with a peristomial sinus.

Type species.—Striatodoma dorothea,
new species, by present designation.

Additional species of Striatodoma.—Tes-
saradoma bifax Cheetham, 1972 and Dr-
plonotos striatum Canu & Bassler, 1930.

Etymology.—tThe first part of the genus
name is from the Latin, striatus = fur-
rowed, channeled, descriptive of the striated
appearance of colony walls. The second

314

part, doma, is derived from the Greek
dwpa, SwpaTol, house, roof, to parallel the
other genus name in the family, Tessara-
doma. Gender neuter.

Remarks.—During the last hundred years
seven living and fossil species of deep wa-
ter cheilostomes have been described and
placed in the family Tessaradomidae Jul-
lien, 1903, and genus Tessaradoma Nor-
man 1869, type species Onchopora borealis
Busk, 1860. Jullien defined the family as
having erect or encrusting colonies and zo-
oids, a tubular peristome with a tubuliform
Spiramen opening into its base, and with a
small, spherical, imperforate ovicell, also
opening into the peristome above the zo-
oidal operculum. The genus Tessaradoma
has a colony form consisting of erect, un-
jointed cylindrical branches arising from an
encrusting base, zooids with a projecting
peristome and prominent spiramen, imper-
forate ovicells obscured by increasing cal-
cification, and adventitious avicularia (Hay-
ward & Ryland 1979). The type species,
Tessaradoma boreale, has a Recent distri-
bution in both the North and South Atlantic
and is also known from Neogene fossil lo-
calities in western Europe and the Mediter-
ranean (Cheetham 1972, Lagaaij & Cook
1973). Its colonies are erect, rigid, and
spreading, with quadriserial branches con-
sisting of elongate oval zooids, arranged
back to back in alternating pairs. The zooid
primary orifice is concealed by a tubular
peristome with a spiramen tube near its
base, its opening projecting from the zooid
frontal surface about one third of the way
down the frontal wall. Wall calcification is
granular, becoming striated around the con-
spicuous marginal pores. Oval adventitious
avicularia are also developed in the mar-
gins, particularly lateral to the spiramen,
The ovicell is small, smooth, imperforate,
subspherical (slightly broader than long),
and though conspicuous in young zooids
becomes increasingly immersed in calcifi-
cation as zooids age (Hayward & Ryland
49770):

Hayward (1981) described Tessaradoma

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

brevissima from the Tasman Sea, but Gor-
don (1989), who recorded T. brevissima
from additional deep water localities off
New Zealand, placed the species in the ge-
nus Galeopsis (family Celleporidae) on the
basis of the following characters: paired av-
icularia in close association with peristome
and sinus, and an ovicell with a central tab-
ulate or fenestrate area and labellum. Two
South African species, Tessaradoma bispi-
ramina and Tessaradoma circella, de-
scribed by Hayward & Cook (1979), and
the Indonesian species Tessaradoma bipa-
tens, described by Harmer (1957) also seem
to belong in this group.

In contrast, the new Pacific species de-
scribed below, as well as two other Pacific
species, the Eocene Tessaradoma bifax
from the small western Pacific Island of
Tonga (Cheetham 1972), and the Recent
Tessaradoma striatum from the Galapagos
(Canu & Bassler 1930), differ from them in
the furrowed appearance of frontal wall cal-
cification, the subspherical imperforate ovi-
cell and the association of the spiramen
with the peristomial sinus. These Pacific
species appear to be related to Tessarado-
ma, but their shared features indicate they
should be grouped with the new species de-
scribed below in a new genus, Striatodoma.
The Eocene Tessaradoma bifax described
by Cheetham (1972) from Tonga, is very
similar in morphology to Striatodoma do-
rothea, with biserial branches, tubular peri-
stome, and with lateral pores and avicularia
making a sinuate double trail along the
sides of branches. In this species, however,
the sinus (at least in the fragmentary ma-
terial available) does not become calcified
into a tube. In addition, its ovicell is not as
prominent as that of S. dorothea, appearing
only as a slight enlargement of the peri-
stome of the maternal zooid, and a swelling
of the frontal shield of the distal zooid. The
Recent Diplonotos striatum collected from
the Galapagos at 1251 m depth (Canu &
Bassler 1930) also belongs in this group.
Based on his studies of syntype material,
Cheetham (1972) transferred that species to

VOLUME 112, NUMBER 2

315

Fig. 1.
attached to pogonophoran tube collected 4100 m depth on Alvin Dive 2828 at 34°42'N, 123°00'W, off California.
(A) Largest colony fragment showing colony form and branching pattern [scale = 2 cm]. (B) Zooid morphology
[from left to right] autozooids in side and front view; ovicelled zooids in front and side view [scale bar = 0.6
mm]. (C) Closeup of peristome, showing spiramen sinus [scale bar = 150 wm]. (D) Closeup of mature ovicelled
zooids, showing constricted peristome and completely enclosed sinus [scale bar = 0.270 mm]. (E) Closeup of
lateral avicularium, showing diagonally tilted rostrum and calcified pivotal hinges [scale bar = 100 pm].

Tessaradoma on the basis of its similarity
to T. bifax, remarking that, contrary to
Canu and Bassler’s original description, the
primary orifice does lie at the base of a
peristomial shaft, but that shaft is almost
completely immersed in the thickened fron-
tal shield. Its chief difference from the other
two species lies in the extreme development
of frontal wall calcification, in which stri-
ation becomes rugosity and external zooid
boundaries are lost. The positions of lateral
avicularia and pores may still be faintly dis-
cerned, however, and match the pattern in
the other two species. Like S. dorothea, the

Striatodoma dorothea new species. All illustrations from portions of holotype colony. CASIZ 113579,

sinus in S. striatum may become completely
enclosed to form a tube.

Striatodoma dorothea new species
Fig. 1

Holotype.—California Academy of Sci-
ences, CASIZ 113579, attached to pogo-
nophoran tube collected as stalk no. 3
(Beaulieu 1998) at 34°42’N, 123°00’W,
4100 m depth, 17 Sep 1994.

Paratypes.—Virginia Museum of Natural
History 567; from stalk 4, ~4100 m,
PULSE 22 Cruise, Chief Scientist Kenneth

316

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Measurements (in mm) of holotype and paratype specimens of Striatodoma dorothea.

Character Range

Zooid Length 0.364-0.501
Zooid Width 0.191—0.273
Orifice Length (autozooid) 0.073—0.100
Orifice Width (autozooid) 0.09 1—0.109
Orifice Length (ovicelled zooid) 0.073

Orifice Width (ovicelled zooid) 0.100

Ovicell Length 0.109-0.155
Ovicell Width 0.182—0.210
Avicularia Length 0.036—0.055
Avicularia Width 0.027—0.046
Branch Width (Distal end) 0.29 1—0.337
Branch Width (Basal end) 0.364—0.728

L. Smith, Jr., Scripps Institution of Ocean-
ography, Alvin Dive 2828, 17 Sep 1994.
Water temperature 1.2°C, 34°42'N,
123°00'W:) CASIZ - LISS SOe estate oi.
34°56'N, 123°07'W, 4100 m, PULSE 24
FVGR;°16 Feb 1995; CASTZ 1135815 stalk
33, 34°42’N, 123°00’W, 4100 m, PULSE
23, Dive 2920; 30. Ape) 1995e(CASIZ
113582, stalk 12, 34°42’N, 123°00’W, 4100
m, PULSE 22, Dive 2834, 23 Sep 1994;
collector for all, Stace Beaulieu.
Etymology.—dorothea, the Latinized
spelling of Dorothy, used as a noun in ap-
position. The species is named in honor of
Dorothy E Soule, in recognition of her el-
egant studies of Pacific bryozoans and of
her active stewardship of California marine
environments.
Diagnosis.—Characterized by subcy-
lindrical branches made up of two series of
rectangular zooids, with longitudinally stri-
ated calcification, rows of marginal pores,
with occasional pores replaced by oval av-
icularia, short peristome with spiramen en-
closed in proximal peristomial sinus, and
relatively prominent ovicell.
Description.—Colony erect, rigidly cal-
cified, unjointed, broadly branching in a
planar fashion, up to several cm in size, the
two biggest branch fragments of the largest
specimen found (the holotype), measuring
Scan h:X Tcmew and '5::cimm-hooovemiw:
respectively (Fig. 1A). Attached to stalks of
deep water glass sponges and pogonopho-

Mean SD n
0.444 0.038 12
0.231 0.027 12
0.085 0.009 12
0.102 0.008 12

— — 2
— — 2

0.134 0.016 6
0.196 0.011 6
0.044 0.007 11
0.036 0.004 11
0.301 0.025 12
0.558 0.103 9

ran tubes by an encrusting base. Zooids
elongate, rectangular, growing back to back
in two alternating longitudinal series (Fig.
1B). Frontal wall convex, with faint longi-
tudinal striations, most of its surface im-
perforate, but with a row of small oval
pores just inside zooid lateral margins. Pri-
mary orifice transversely oval, surrounded
by a short peristome (Fig. 1C). No out-
wardly visible spiramen; instead, the peri-
stome of young zooids has a proximal si-
nus, which is encircled by calcification as
the zooid ages, becoming increasingly tu-
bular and projecting. Ovicells are smoothly
calcified and globular, reaching the height
of the peristome opening, which becomes
slightly narrowed in fertile zooids (Fig.
1D). About one pore per zooid is replaced
by an oval adventitious avicularium, about
50 wm in length. In side view the sinuate
double track of pores and avicularia is dis-
tinctive (Fig. 1E, B). In basal regions of the
colony zooid openings are calcified over
and branches become thickened. Some of
the avicularia and pores are calcified over
also. Branch thickness (the depth of two
back to back zooids) averages 0.301 mm in
zooids near the growing edge of branches
and 0.558 mm in zooids near the colony
base. Zooid measurements are summarized
in Table 1.

Distribution and ecology.—The species
was observed and/or collected at 4060—
4100 m depths off California, between lat-

VOLUME 112, NUMBER 2 317,

itudes 34°38’ and 34°56’N and longitudes
122°59’ and 123°15’W. General area: 220
km west of Point Conception, CA. Beaulieu
(1998) collected a total of 35 tube core
samples at the abyssal station and found
seven colonies of S. dorothea (one colony
per substrate; Table 2). The species was at-
tached to pogonophoran tubes, the basal
spicules of Hyalonema sp., and to other or-
ganisms that were attached to the host sub-
strate. The branches of S. dorothea also
provided substrate on which other species
attached (Table 2).

In order to determine the abundance of
S. dorothea at the abyssal station, all colo-
nies large enough to identify with certainty
were enumerated in photographic transects
of the sea floor (procedures described in
Beaulieu 1998). We photographed approx-
imately 9 km of the sea floor (total of seven
transects) in which we encountered 55 col-
onies of S. dorothea. Of these, 53 were at-
tached to dead Hyalonema spicules, one to

Epizoites attached to S. dorothea
tulus), colonial hydroid, sabellid, serpulids (Bathyvermilia sp. and

Hyalopomatus mironovi), isopod (Arcturus sp.)
Agglutinated foraminiferan, calcareous foraminiferan (Cibicides lob-

atulus), colonial hydroid, serpulid (Hyalopomatus mironovi)

Overgrown by colonial hydroid

Agglutinated foraminifera, calcareous foraminiferan (Cibicides loba-
None

Overgrown by colonial hydroid

None
None

a pogonophoran tube, and one to an un- fi

identified structure. The encrusting bases of S

most of the colonies were attached at the 2 Q

middle of the host substrate, elevating the = Be =

colonies about 10 cm above the sea floor. : = a

Density estimates (no. colonies per unit 9 ty) Gaee S

area of sea floor) were calculated using the 3 bs 2 2 6 : 6

computer program DISTANCE (Laake et S aT Pas 5 = 8

al. 1994). Mean density estimates for the S| “| 22 & ae 3

individual transects ranged between 3 and iS a S rs : = =

8 colonies per 1000 m2. Only about 2% of : seu S38

the biogenic structures enumerated in pho- 3 e S ee = 5 =

tographic transects appeared to be colo- 8

nized by S. dorothea. However, 20% of the 5

structures collected from the sea floor had &

S. dorothea attached; therefore, the density &

estimates from the photographic transects E c c

may be an order of magnitude low. mM 2| oO ees
Discussion.—Tessaradomids belong to = 1 33| Scie S72) S18

the fauna of the outer continental shelf and 8 ; 6 6 ~ = = 3 3 5 z

slope. The 4100 m depth recorded for S. 2 a4 coy ely any : =

dorothea in this study is the deepest re- ta x 9

corded, but it is not much deeper than the | 23

3700 m recorded for T. boreale. However, wp hiaw Pu oe

T. boreale is found in much shallower wa- = [se] — oy MES Pes k

ter in the Arctic (70 m depth), and Cheet-

318

ham (1972) suggested that distribution of
such deep-sea species may be temperature
controlled, limited to water temperatures
between 2° and 13° C. This paper and Beau-
lieu (1998) provide a glimpse of how erect
bryozoans with an attached base may sur-
vive on the muddy deep sea floor. But as
yet, these species are known only from a
few broken fragments, collected at widely
scattered localities by various expeditions.
Better understanding of their ecology as
well as clarification of their relationships
with shallow water species must await the
kinds of collections that can provide mate-
rial adequate for the detailed anatomic
study necessary for phylogenetic analysis.

Acknowledgments

We would like to thank K. Smith and the
pilots of Alvin for making the collections
and in situ observations possible. S. Beau-
lieu was supported by a National Defense
and Engineering Grant and by NSF grant
OCE92-17334 to K. Smith. We are also in-
debted to Dr. W. C. Banta (American Uni-
versity) and Dr. A. H. Cheetham (National
Museum of Natural History) for their help-
ful reviews of the manuscript.

Literature Cited

Beaulieu, S. E. 1998. The ecology of glass sponge
communities in the abyssal N.E. Pacific. Un-
published Ph.D. thesis, University of California,
San Diego, 206 pp.

Busk, G. 1860. Zoophytology. Shetland Polyzoa col-
lected by Mr. Barlee.—Quarterly Journal of Mi-
croscopical Science 8:143—145.

Canu, R., & R. S. Bassler. 1930. The Bryozoan fauna

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

of the Galapagos Islands.—Proceedings of the
U.S. National Museum 76, Art. 13:1—78.

Cheetham, A. H. 1972. Cheilostome Bryozoa of Late
Eocene Age from Eua, Tonga.—U.S. Geologi-
cal Survey Professional Paper 640-E:1—26.

Gordon, D. P. 1989. The marine fauna of New Zea-
land: Bryozoa: Gymnolaemata (Cheilostomida
Ascophorina) from the western South Island
continental shelf and slope.—New Zealand
Oceanographic Institute Memoir 97:1—158.

Harmer, S. E 1957. The Polyzoa of the Siboga Expe-
dition, part 4, Cheilostomata Ascophora II.—
Leyden, Siboga Expedition Reports, 28d:641—
1147.

Hayward, P. J. 1981. The Cheilostomata (Bryozoa) of
the deep sea.—Galathea Report 15:21—68.

, & P. L. Cook 1979. The South African Mu-

seumis Meiring Naude Cruises, part 9. Bry-

ozoa.—Annals of the South African Museum

79:43—-130.

, & J. S. Ryland. 1979, British Ascophoran
Bryozoans. London, Academic Press, 312 pp.

Hincks, T. 1862. A catalogue of the zoophytes of south
Devon and south Cornwall.—Annals and Mag-
azine of Natural History, series 3, 9:467—475.

Jullien, J. 1903. Bryozoaires provenant des campagnes
de |’ Hirondelle (1886—88).—Résultats des cam-
pagnes scientifiques accompli par le Prince Al-
bert I, 23:1—120.

Laake, J. S., S. T. Buckland, D. R. Anderson, & K. P.
Burnham. 1994. DISTANCE User’s Guide V2.1
Colorado Cooperative Fish and Wildlife Re-
search Unit, Colorado State University, Fort
Collins, Colorado, 84 pp.

Lagaaij, R., & P. L. Cook. 1973. Some Tertiary to Re-
cent Bryozoa. Pp. 489-498 in A. Hallam, ed.,
Atlas of Palaeobiogeography. Elsevier Scientific
Publishing Company, Amsterdam.

Norman, A. M. 1869. Last report on dredging among
the Shetland Islands.—Report of the British As-
sociation for the Advancement of Science for
1868:303-312.

Schopf, T. J. M. 1969. Geographic and depth distri-
bution of the phylum Ectoprocta from 200 to
6000 m.—Proceedings of the American Philo-
sophical Society 113:464—474.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
112(2):319—326. 1999.

Two new species of Dentatisyllis and Branchiosyllis
(Polychaeta: Syllidae: Syllinae) from Venezuela

Guillermo San Martin and David Bone

(GSM) Laboratorio de Biologia Marina e Invertebrados, Departamento de Biologia,
Unidad de Zoologia, Universidad Aut6noma de Madrid, Canto Blanco,
E-28049 Madrid, Spain, e-mail: guillermo.sanmartin@uam.es;

(DB) Departamento de Biologia de Organismos, Instituto de Tecnologia y Ciencias Marinas,

Universidad Simon Bolivar, Caracas, 1080 Venezuela, A. P. 89000

Abstract.—A new species of the genus Dentatisyllis Perkins, 1981 and an-
other new species of the genus Branchiosyllis Ehlers, 1887 are described: Den-
tatisyllis morrocoyensis and Branchiosyllis lorenae. The specimens of both new
species were collected during a study of the polychaetes from Thalassia tes-
tudinum beds in Morrocoy Park, Venezuela. One specimen of B. lorenae was
previously collected and reported as Branchiosyllis sp., from Cuba by San
Martin (1991), so this species could be distributed throughout the Caribbean
area. Dentatisyllis morrocoyensis is distinguished from all other species of the
genus by the high number and shape of teeth in the trepan and shape of the
blades of the compound setae, unique in the genus. Branchiosyllis lorenae is
very similar to B. exilis but differs by having, on the anterior and midbody
parapodia, 1—2 compound setae, the bidentate blades of which are slightly
curved and bear a distal tooth somewhat shorter than the proximal one. This

latter characteristic is unique to the genus.

During a study on the ecology of the
polychaetes inhabiting Thalassia testudin-
um beds in the Morrocoy Park, Venezuela,
specimens of two undescribed species of
Syllidae were collected. A series on the tax-
onomy and ecology of several families of
polychaetes from this area and habitat has
been recently begun (Bone & Viéitez
1999). This paper deals with the description
of the two new species of Syllidae: Den-
tatisyllis morrocoyensis and Branchiosyllis
lorenae. The study of the syllids from Mor-
rocoy Park has been supported by the
Agreement between the Universities Simon
Bolivar (Venezuela) and Aut6noma de Ma-
drid (Spain).

Materials and Methods

All samples were collected in shallow
Thalassia testudinum seagrass beds in less

than 0.5 m depth. The samples were taken
using a 38 cm? corer which was pushed 25
cm into the sediment. Sediment samples
were preserved in 10% buffered formalin
and washed through a 1 mm mesh sieve.
All organisms were hand-picked under a
magnifying-lens from the remaining mate-
rial and separated for taxonomic identifi-
cations. Measurements are referred to the
holotype or largest specimen studied; width
is measured across the proventriculus and
excludes cirri, parapodia, and setae. Obser-
vations, drawings, and measurements were
made using a microscope with interference
contrast optics. Drawings were made with
the aid of a drawing tube. The SEM micro-
graphs were taken at the SIDI (Servicio In-
terdepartamental de Investigacién) of the
University Aut6noma of Madrid. Types are
deposited in the Museo Nacional de Cien-
cias Naturales de Madrid, Spain.

320

Results and Discussion

Family Syllidae Grube, 1850
Subfamily Syllinae Grube, 1850
Genus Dentatisyllis Perkins, 1981
Dentatisyllis morrocoyensis, new species
Fig. 1

Material examined.—Morrocoy Park
(Venezuela), Thalassia testudinum beds,
Holotype.

Description.—Body small, thin, cylindri-
cal, without color marking, incomplete,
about 4.2 mm long, 0.32 mm wide, 30 se-
tigers. Prostomium semicircular; four small
eyes in open trapezoidal arrangement and
two anterior eyespots. Only one lateral, bro-
ken antenna present on this specimen, with
10 articles, originating in front of anterior
eyes. Palps slightly longer than prostomi-
um, fused at bases. Tentacular segment dis-
tinct, somewhat shorter than following seg-
ments; dorsal tentacular cirri with about 13
articles; ventral tentacular cirri somewhat
shorter, with about 8 articles. Dorsal cirri of
first setiger long, with about 23 articles; re-
maining dorsal cirri alternating long and
short; long dorsal cirri similar in length to
body width, with about 17—18 articles, short
dorsal cirri shorter than body width, with
about 10 articles (Fig. 1A). Parapodia elon-
gate, each with distal anterior lobe, some-
what shorter than distal posterior lobe; ven-
tral cirri long, digitiform, extending past
tips of parapodial lobes, distally broad (Fig.
1D). Compound setae all heterogomph fal-
cigers, similar throughout; about 8—10 setae
on each parapodium. Blades bidentate, with
both teeth very close, similar in length.
Marked dorso-ventral gradation in shape
and length of blades; blades of dorsal-most
compound setae long, about 38 wm, with
short and coarse spines on margin; remain-
ing blades provided with long, coarse, up-
wards dressed, 3—4 spines on bases, espe-
cially on medium size ones, and shorter and
thinner spines distally; blades of ventral-
most compound setae about 20 wm length
(Fig. 1F). Simple dorsal and ventral setae
not seen. Parapodia each with two slender

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

aciculae, ending with fine tips (Fig. 1E).
Pharynx long, everted on this specimen
(Fig. 1A); anterior margin (Fig. 1A, B) sur-
rounded by a trepan of about 50 curved,
hooked teeth (Fig. 1B, C), and a crown of
about 20 soft papillae; midorsal pharyngeal
tooth rhomboidal, small, located subtermin-
ally to anterior margin. Proventriculus
shorter than pharynx, through 4 segments,
with about 27 muscle cell rows.

Remarks.—The genus Dentatisyllis was
erected by Perkins (1981) for species hav-
ing a cylindrical body, pharyngeal tooth
and a trepan on the anterior margin of the
pharynx; the genus has been recently re-
vised by Ding et al. (1998) who provided a
diagnosis and a key to all the known spe-
cies of the genus. Dentatisyllis morrocoy-
ensis N. sp., is the only species of the genus
provided with a very high number of mar-
ginal teeth on the trepan; all other species
have about 10, whereas D. morrocoyensis
has about 50. The blades of the compound
setae and the aciculae of D. morrocoyensis
n. sp. are very similar to those of Opistho-
syllis longidentata San Martin, 1991, but
the pharyngeal armature is completely dif-
ferent (San Martin 1991).

Etymology.—The species is named after
the type locality, Morrocoy Park (Venezue-
la).

Genus Branchiosyllis Ehlers, 1887
Branchiosyllis lorenae, new species
Figs. 2, 3, 4

Branchiosyllis sp. San Martin (1991):234,
fig. 1O-S.

Material examined.—Morrocoy Park
(Venezuela), Thalassia testudinum beds,
Holotype and 27 paratypes. Additional ma-
terial: 2 specimens used for SEM.

Description.—Body long, cylindrical
(Figs. 2A, 4A), holotype incomplete speci-
men, 5.3 mm long, 0.4 mm wide, 54 setig-
ers; longest complete paratype 7.2 mm
long, 0.48 mm wide, 50 setigers, a few
somewhat longer, incomplete paratypes.
Most anterior segments without color mark-

VOLUME 112, NUMBER 2 321

CWA

————— =

ae -coerrvranruuetitiecitnnrn

\
\ \

\ \

\
\

Fig. 1. Dentatisyllis morrocoyensis, n. sp. Holotype. A, anterior end, dorsal view; B, ventral view of the
anterior end of the pharynx; C, detail of the teeth of the trepan; D, midbody parapodium, anterior view; E,
aciculae, from midbody; EK compound setae, midbody. Scale.—A:0.11 mm. B, D:65 «wm. C:48 wm. E, F:20 wm.

ing, anterior and midbody segments provid- dorsal cirri with dark spots (Fig. 2A). Pro-
ed dorsally each with three ovate dark stomium oval, wider than long; four eyes in
spots, sometimes forming nearly a row; very open trapezoidal arrangement, nearly
some articles of antennae, tentacular and on line, and two small anterior eyespots.

322 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Pst

a

Se 2
SS

= — sat
ore Pest tiss
(<XE YES RAL Ii £2 a

Fig. 2.

Branchiosyllis lorenae, n. sp. Holotype. A, anterior end, dorsal view; B, parapodial lobe, dorsal

view. C, aciculae, anterior parapodium: D, aciculum, posterior parapodium. Scale-—A:0.11 mm. B:48 pm.

C, D:20 um.

Median antenna originating between pos-
terior pair of eyes, somewhat longer than
prostomium and palps together, with 14 ar-
ticles; lateral antennae shorter than median
antenna, originating between anterior pair
of eyes and eyespots, with 12-13 articles.
Palps broad, longer than prostomium, fused
at bases. Tentacular segment reduced, cov-
ered dorsally by prostomium and setiger 1;
dorsal tentacular cirri longer than median

antenna, with about 20 articles; ventral ten-
tacular cirri similar in length to lateral an-
tennae, with about 13 articles. Dorsal cir
of setiger 1 very long, with about 30—33
articles; remaining dorsal cirri alternating
long, somewhat longer than body width,
with 25—30 articles, and short cirri, shorter
than body width, with about 16—18 articles.
Parapodial lobes conical, provided with two
distal, triangular lobes (branchiae), anterior

VOLUME 112, NUMBER 2

Ey. 5.

323

Branchiosyllis lorenae, n. sp. Compound setae: A, dorsal from anterior parapodium; B, median and

ventral from anterior parapodium. C, from anterior midbody; D, from posterior midbody; E, from posterior
parapodium; FE claw-shaped setae, from posterior-most parapodia. Scale.—20 p.m.

lobe somewhat shorter than posterior lobe
(Figs. 2B, 4E). Ventral cirri digitiform,
elongate, reaching distal level of parapodial
lobes. Anterior parapodia each with 1-2,
sometimes 3, dorsal, compound setae pro-
vided with long, somewhat curved, biden-

tate blades, 50-55 wm, distal tooth short
and small proximal tooth somewhat larger,
more prominent (Figs. 3A, 4C); spines on
margin short, several rows of spines ob-
served by SEM, slightly longer than those
of the rest of blade, partially covering prox-

324 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Branchiosyllis lorenae, n. sp. SEM. A, anterior end, dorsal view; B, anterior rim of pharynx; C, long
bladed, dorsal compound seta, anterior parapodium; D, detail of the same, showing several rows of spines on
distal part. E, midbody posterior parapodium; FE dorsal compound seta from midbody; G, ventral compound
setae from midbody; H, claw-shaped compound setae, from posterior-most parapodia.

VOLUME 112, NUMBER 2

imal tooth (Fig. 4D), 7—10 median and ven-
tral compound setae with blades curved, bi-
dentate, both teeth similar on longer blades,
proximal tooth shorter than distal one on
shorter blades; gradation in length, 32 wm
above, 24 wm below (Fig. 3B). Progres-
sively decreasing number of compound se-
tae on each parapodium, blades shorter,
lacking compound setae with long blades;
midbody parapodium with a few bidentate
blades and several unidentate, smooth or
slightly bidentate, hooked blades (Figs. 3D,
4E G); posterior parapodia each with a few
modified compound setae with blades
turned 180°, smooth, strongly hooked,
claw-shaped (Fig. 4H); posterior-most para-
podia only with 2-3 claw-shaped com-
pound setae (Fig. 3F). Shafts of compound
setae distally provided with spines, which
are increasingly more numerous and longer
in dorsal and anterior setae. Solitary dorsal
and ventral simple setae absent. Anterior
parapodia each with 3-4 aciculae, dimin-
ishing progressively to only 1—2 on poste-
rior parapodia; aciculae with curved tip
(Fig. 2C, D). Pharynx broad, through 5 seg-
ments; anterior margin provided with a
crown of ten soft papillae and one conical
middorsal tooth (Figs. 2A, 4B). Proventric-
ulus somewhat longer than pharynx,
through 6 segments, with about 30 muscle
cell rows. Pygidium small, with two long,
articulated anal cirri.
Remarks.—Branchiosyllis is a genus
with only five described species: B. oculata
Ehlers, 1887 (Ehlers 1887), distributed in
the Caribbean Sea; B. exilis (Gravier, 1900)
(Gravier 1900, Westheide 1974, San Martin
1984), a circumtropical species, also pre-
sent in temperate waters; B. pacifica Rioja,
1941 (Rioja 1941), from the Pacific coasts
of Mexico; B. diazi Rioja, 1958 (Rioja
1958), from the Caribbean coast of Mexico;
and B. abranchiata Hartmann-Schréder,
1965 (Hartmann-Schroéder 1965) from Sa-
moa. A table summarizing the main char-
acters of all these species is provided in
Hartmann-Schroder (1978) and San Martin
(1984). Branchiosyllis lorenae, n. sp. is eas-

S25

ily distinguished from B. oculata, B. paci-
fica and B. abranchiata because these three
species have only claw-shaped seta, and
lack unmodified ones. Branchiosyllis lor-
enae differs from B. diazi in lacking a dor-
sal branchia on each parapodium and in the
shape of anterior compound setae. The most
similar species is B. exilis; however, B. lor-
enae differs in having compound setae on
anterior parapodia with long, curved,
blades, with the distal tooth small, shorter
than the proximal tooth; this kind of blade
is very unusual and B. lorenae, n. sp., is the
only member of this genus with this kind
of seta.

Etymology.—The species is named in
honor of Lorena Galindo, who collaborated
in the collection of the samples and sepa-
ration of specimens.

Literature Cited

Bone, D., & J. M. Viéitez, 1999. Spionidae from Mor-
rocoy Park (Venezuela).—Bulletin of Marine
Science (in press).

Ding, Z., E Licher, & W. Westheide. 1998. New and
newly assigned species of the genus Dentati-
syllis (Polychaeta, Syllidae, Syllinae), with
comments on the reproduction, together with a
key and a synoptic table of all species of the
genus.—Sarsia 83:29-—43.

Ehlers, E. 1887. Report on the annelids of the dredging
expedition of the U. S. coast survey steamer
‘“‘Blake’’.—Memoires of the Museum of Com-
parative Zoology of Harvard 15:335 pp.

Gravier, C. 1900. Contribution a l’étude des Annélides
Polychétes de la Mer Rouge.—Nouvelles Arch-
ives du Museum d’ Histoire naturelle de Paris 2:
137-282.

Hartmann-Schréder, G. 1965. Zur Kenntnis der Euli-
toralen Polychaetenfauna von Hawaii, Palmira
und Samoa.—Abhandlungen und Verhandlun-
gen des Naturwissensschaftlichen Vereins in
Hamburg 9:81—161.

. 1978. Einige Sylliden-Art (Polychaeta) von
Hawaii und aus dem Karibischen Meer.—Mit-
teilungen aus dem Hamburgischen Zoologisch-
en Museum und Institut 75:49-—61.

Perkins, T. H. 1981. Syllidae (Polychaeta), principally
from Florida, with descriptions of a new genus
and twenty-one new species.—Proceedings of
the Biological Society of Washington 93:1080—
Lie

Rioja, E. 1941. Estudios Anelidoldgicos. III. Datos para
el conocimiento de la fauna de poliquetos de las

326

costas del Pacifico de México.—Anales del Insti-

tuto de Biologia de México 12:669—746.

. 1958. Estudios Anelidolégicos. XXII. Datos
para el conocimiento de la fauna de Anélidos
Poliquetos de las costas orientales de México.—
Anales del Instituto de Biologia de México 29
(1/2):219-301.

San Martin, G. 1984. Estudio biogeografico, faunistico
y sistematico de los Poliquetos de la familia Sil-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

idos (Syllidae: Polychaeta) en Baleares.—Tesis

Doctoral, Ediciones de la Universidad Complu-

tense de Madrid, n° 187:529 pp.

199]. Syllinae (Polychaeta: Syllidae) from
Cuba and the Gulf of Mexico.—Bulletin of Ma-
rine Science 48:227—235.

Westheide, W. 1974. Interstitielle Fauna von Galapa-
gos. XI. Pisionidae, Hesionidae, Pilargidae, Syl-
lidae.—Mikrofauna Meeresbodens 44:195—338.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(2):327-—337. 1999.

Parvidrilus strayeri, a new genus and species, an enigmatic interstitial

clitellate from underground waters in Alabama

Christer Erséus

Department of Invertebrate Zoology, Swedish Museum of Natural History,
Box 50007, SE-104 05 Stockholm, Sweden

Abstract.—An exceedingly small oligochaete, Parvidrilus strayeri, a new
genus and species, is described from a streambed in northern Alabama, U.S.A.
The species is up to 1.4 mm long (with up to 33 segments) and has several
unusual features. In ovigerous specimens, a clitellum is developed as a pair of
lateral rows of a few, large, swollen epidermal cells in segments (IX—) X—XII
(—XIII, —XIV). Chaetae are absent from segment II. Otherwise, in the anterior
part of the body, there are long hair chaetae as well as small crotchet chaetae
in both dorsal and ventral bundles. Furthermore, the chaetal bundles are situated
posteriorly in each segment. It is unclear whether gonads are paired or un-
paired; testes (testis?) are in segment XI, ovaries (ovary?) in XII. The other
reproductive organs include a U-shaped muscular ‘genital body’ in segment
XII and a V-shaped ‘copulatory organ’ in segments XII—XIII; the exact nature
and function of these structures are unknown. The genus is proposed to be
classified as the single member of Parvidrilidae, new family. It appears to be
most closely related to two, largely Southern Hemisphere, aquatic clitellate
families, Capilloventridae and Phreodrilidae. A possible relationship to the

monotypic, South American taxon, Narapidae, is also discussed.

In the course of a study of the under-
ground fauna of a crystal clear, spring fed
stream in Alabama (Strayer et al. 1995),
specimens of an exceedingly small, undes-
cribed, oligochaete were encountered. Dr.
David L. Strayer (Institute of Ecosystem
Studies, Millbrook, New York, U.S.A.)
placed the material at the present author’s
disposal. It is described as the type species
of a new genus, Parvidrilus, in the present
paper. The phylogenetic affinities of the
new species, which appears to represent a
new higher level taxon of the Clitellata, are
also discussed.

Material and Methods

The worms were collected at a single site
in Hendrick Mill Branch, northern Ala-
bama, in October 1990 (Strayer et al. 1995).
They were fixed in buffered formalin,

stained with Rose Bengal, sorted under a
dissecting microscope, and stored in 70%
ethanol. The material was then sent to the
present author, who stained several individ-
uals in alcoholic paracarmine and mounted
them whole in Canada balsam on micro-
scope slides. Most of the measurements in
the description refer to this whole-mounted
material, examined under a light micro-
scope. Due to the small dimensions, how-
ever, the details of the chaetae could not be
observed even when using a high-resolution
100 oil immersion lens. Therefore, exter-
nal traits were also investigated by means
of scanning electron microscopy. Since the
animals were too small for the perforated
containers available for critical point dry-
ing, they were simply dehydrated in 99%
ethanol, transferred to butyl acetate, and air-
dried. This is a rough method, but thanks
to their sturdy cuticle and small size the

328

worms maintained the body shape accept-
ably, at least in a comparison with individ-
uals prepared for light microscopy. Three
specimens were mounted on double-sided
tape, coated with gold and examined with
SEM.

The holotype and some paratypes of the
new species, Parvidrilus strayeri, are de-
posited in the National Museum of Natural
History (USNM), Smithsonian Institution,
Washington, D.C., additional paratypes in
the Swedish Museum of Natural History
(SMNBH), Stockholm.

Parvidrilidae, new family

Etymology.—Family name (-idae) based
on Parvidrilus, new genus (type genus).

Diagnosis.—Until additional species are
found, the new family is characterized by
the features of the type species of the type
genus. They can be summarized as follows:

Meiofaunal freshwater oligochaetes,
about 1 mm long. Chaetal bundles situated
posteriorly in each segment, but completely
absent from segment II. In anterior part of
body, both dorsal and ventral bundles with
very long hair chaetae as well as short
crotchet chaetae; dorsal crotchets single-
pointed, ventral ones bifid. In posterior part
of body, dorsal bundles similar to anterior
dorsal ones, ventral bundles with bifid
crotchets only. In ovigerous specimens, cli-
tellum developed as a pair of lateral rows
of large, swollen, transparent, cells in seg-
ments ([X—) X—XII (-XIII, —XIV). Alimen-
tary canal simple, without diverticula. It is
unclear whether gonads are paired or un-
paired; testes (testis?) in segment XI, ova-
ries (ovary?) in XII. Reproductive organs
complex, including U-shaped muscular
‘genital body’ in XII and V-shaped ‘copu-
latory organ’ in XII—XIII.

Parvidrilus, new genus

Etymology.—A combination of parvus
(Latin for ‘small’) and drilus (Greek for
‘worm.’ ).

Diagnosis.—As for family.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Type species.—Parvidrilus strayeri, new
species.
Other species.—None.

Parvidrilus strayeri, new species
Figs 1-2

Oligochaeta n. sp.: Strayer et al. 1995:506.

Holotype.—USNM_ 185769, whole-
mounted specimen.

Type locality——A hyporheic site in a sand-
gravel bar, in Hendrick Mill Branch, a spring-
fed stream about 20 km NE of Pinson,
Blount Co., Alabama, U.S.A., 33°52’12’N,
86°33'57’W, 4 Oct 1990, coll. D. L. Strayer.
At this site, the stream is 3—6 m wide and
runs through a hardwood forest in a lime-
stone terrain. The bottom varies from sand
to exposed bedrock, with a predominance
of coarse sand, angular gravel and small
(10—15 cm) stones. Generally the alluvium
is thin, but a large deposit of sand and grav-
el is held in place by a dam about 3 m high.
At the time of collection, the emergent part
of this deposit was 3—4 m wide, about 30
m long, and reached about 0.5 m above the
stream water level. Because the dam leaked
from the base, water flowed down into and
through the bar. Subsamples were taken
from 16 wells in the bar, using a Bou-
Rouch pump (Bou 1974), all wells yielded
water freely. Dissolved oxygen was O-8.0
ppm, with most readings 2—6 ppm. Other
chemical characteristics (means of several
subsamples): Ca 22.2 ppm; Mg 11.6 ppm;
K 0.5 ppm; Na 0.7 ppm; NO, 0.9 ppm; SO,
1.8 ppm; Cl 1.2 ppm; Dissolved Organic
Carbon (DOC) 0.5 ppm (Strayer et al. 1995;
Strayer, pers. comm.).

Paratypes.—USNM 185770—-185776-
000000, seven whole-mounted specimens;
SMNH Type coll. 5085—5092, eight whole-
mounted specimens; SMNH Type coll.
5093, three specimens mounted on a SEM
stub; all from type locality.

Etymology.—Named for Dr. David L.
Strayer, who collected the material and was
the first to realize that the species was a

VOLUME 112, NUMBER 2

A

eM

~

Fig. 1. Parvidrilus strayeri, new genus and species; SEM micrographs. A, postclitellar part of body, showing
dorsal hair chaetae (hc) ventral bifid chaetae (in bottom part of figure), and intersegmental furrows (if); note the
posterior location of bundles in the segments; B, anterior end of body showing mouth (m), achaetous segment II (ID,
and dorsal (top of figure) and ventral chaetae (middle and bottom of figure) of segments III—V; C, bundle of ventral
chaetae of segment IX, showing hair chaeta (hc) and bifid crotchets (bc); D, bundle of dorsal chaetae of segment X,
showing hair chaeta (hc), and single-pointed crotchets (sc); note also thin outer part of hair chaetae of another segment;
E, bundle of ventral chaetae of postclitellar segment, showing two bifid crotchets; EK ventral view of genital bursa
(bu) in segment XII, showing lateral bulbous swellings (bs), posterior triangular lappet (tl), and ventral chaetae (vc).

330 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

50 um

(B-C)

= AE
<a
pe ZF SON
STB fi Cty
paral dys

gb Ds

Fig. 2. Parvidrilus strayeri, new genus and species. A, schematical, horizontal view of segments XII and
XIII, showing general outline of ‘genital body’ and ‘copulatory organ’; B, somewhat horizontal view of segments
XI-XIII of one paratype; C, somewhat lateral view of segments XI—-XIII of another paratype. Abbreviations: bs,
bulbous swelling (of body wall); bu, location of genital bursa (cf. Fig. 1F); c, chaeta; cc, clitellar cell; co,
copulatory organ; gb, genital body; ov, ovary; pe, penis/pseudopenis; ps, penial/pseudopenial sac; sf, sperm
funnel, t, testis.

VOLUME 112, NUMBER 2

new taxon of considerable systematic inter-
est.

Description.—Length (five complete
specimens) 0.7—1.4 mm, 18—33 segments.
Width at genital region 0.03—0.10 mm. Pro-
stomium rounded, somewhat narrower than
and well set off from peristomium; prosto-
mial epidermis with numerous (stained) cell
nuclei, probably indicating a glandular or
sensory function. Posterior part of body
(Fig. 1A) more clearly annulated (i.e., with
distinct intersegmental furrows; Fig. 1A, if)
than anterior part. Pygidium generally de-
marcated, somewhat narrower than last
(true) segment. Cuticle smooth, but foreign
particles often adhering to it here and there
along body of worm. Clitellum poorly de-
veloped in most individuals; but in oviger-
ous specimens, a row of large, swollen,
transparent, epidermal cells (Fig. 2B, cc)
present on each side of worm in segments
(IX—) X—XII (—XII], —XIV).

Chaetal bundles situated posteriorly in
each segment (Fig. 1A), but lacking com-
pletely in segment II (Fig. 1B). Anterior
dorsal bundles each with one to two (oc-
casionally three) long, whip-like, hair chae-
tae (Fig. 1D, hc), alternating with two or
three (occasionally four) single-pointed,
smaller crotchet chaetae (Fig. 1D, sc); hairs
occasionally up to about 200 wm long (ex-
act length difficult to measure due to strong
curvature of hairs and poorly resolved view
of tips), entally about 1 um wide, progres-
sively tapering and only about 0.1 um wide
at ectal end; crotchets 15-25 pm long,
about 0.5—0.6 wm thick, with somewhat
blunt tips. Postclitellar dorsal bundles each
with one or no hair chaeta (Fig. 1A, hc) and
two single-pointed crotchets. In anterior
segments (at least in III—VII, but generally
also including a few additional pre-genital
segments, and sometimes even as far back
as in XV), ventral bundles each with one
hair chaeta (Fig. 1C, hc), and three (occa-
sionally four) bifid crotchets (Fig. 1C, bc);
ventral hairs slightly shorter and thinner
than corresponding dorsal hairs; bifids
about 20—25 um long, about 0.9-1.4 pm

5a

thick, with upper tooth thinner and shorter
than lower tooth. Within these bundles, up-
permost (most lateral) chaeta bifid, fol-
lowed by a hair, and these two chaetae
clearly separated from two, more ventral,
bifids (Fig. 1C). Postclitellar ventral bun-
dles with two to three bifid crotchets (Fig.
1E), similar to anterior ventral bifids. Mod-
ified genital chaetae absent.

Ventrally in segment XII, body wall
forming a conspicuous, generally somewhat
**X’’-shaped genital bursa (Fig. 1F); im-
pression of an X facilitated by a pair of
large, bulbous, swellings (Fig. 1E bs), one
at each side of bursal opening, and an in-
distinct, mid-ventral, triangular lappet (Fig.
1E tl), located immediately posterior to bur-
sal opening.

Spermathecal openings not observed.

Brain located within, and with front end
reaching anterior coelomic lining of, peris-
tomium. Nerve cord ventral, of normal mi-
crodrile oligochaete appearance. Alimenta-
ry system of normal microdrile oligochaete
type too: simple pharynx with dorsal pha-
ryngeal bulb, somewhat sinuous esophagus
with a few pharyngeal (?) gland cells scat-
tered in segments V—VIII, followed by a
narrow, simple intestine; no particular en-
largements or diverticula on gut. Large, dif-
fuse cells of chloragogen tissue present
along most of gut. Free coelomocytes not
observed.

In three specimens, a small lump of cells
in anterior end of coelom of segment XI
probably an (unpaired?) testis (Fig. 2C, t?);
in another worm, a single sperm bundle
present in coelom of this segment; other-
wise male gonads or developing spermato-
zoa not observed. In nine specimens, an
ovary (apparently developed on one side
only) observed in anterior part of segment
XII (Fig. 2B, C, ov); three of these worms
ovigerous, i.e., with a large mature egg fill-
ing whole width of coelom, and extending
through one or two segments, in region of
XIII—XV. In two specimens (only), a sperm
funnel discernible at anterior side of septum

332

between XI and XII (Fig. 2B, C, sf), but its
continuation into a vas deferens not seen.
Other internal genitalia Gn XII—XIII)
complex (Fig. 2), but due to quality of fix-
ation and small dimensions difficult to in-
terpret using conventional oligochaete ter-
minology. Two conspicuous structures pres-
ent, here referred to as the ‘genital body’
and the ‘copulatory organ’, respectively.
Genital body (Fig. 2A—C, gb) a “U”’-
shaped structure incorporated in ventral
body wall in segment XII; the two arms
forcing body to bulge considerably, corre-
sponding to two bulbous swellings at lateral
sides of genital bursa (Fig. 1K bs). Arms of
genital body more or less oval, 20—35 pm
long, 15—23 pm wide, oriented along long
axis of worm, and united anteriorly. At ear-
ly developmental stages, lobes appear to
contain a lumen. At full maturity, wall of
genital body thick, heavily muscular and
folded in complex manner (Fig. 2B, C, gb),
with indistinct lumen (lumina?); sometimes
densely packed cilia (see Fig. 2B) and/or
spermatozoa (?) visible inside. At late stag-
es of development, lobes tend to break up
into a number of secondary, pear-to-spin-
dle-shaped, packages of muscle fibers (as in
Fig. 2B). Function of genital body un-
known; possibly having something to do
with storage and/or ejaculation of sperm, ei-
ther before (an atrium?) or after copulation
(a spermatheca?). Whether genital body has
any permanent or temporary opening to the
exterior or any connection with the sperm
funnels or copulatory organ unknown.
Copulatory organ (Fig. 2A, co) bilobed,
somewhat “‘V’’-shaped, consisting of a pair
of straight or somewhat curved sacs (Fig.
2B, C, ps), communicating with each other
mid-ventrally in most posterior part of seg-
ment XII. At point of union of the two sacs,
copulatory organ appears anchored in the
ventral body wall, in a position correspond-
ing to apex of the mid-ventral, triangular,
lappet located in posterior wall of genital
bursa (Fig. 1K tl; see above). Inner ends of
sacs extending obliquely and freely back-
wards into coelom of XIII. Each sac 25—60

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

wm long, 10—21 wm wide, with thick, mus-
cular, wall, and containing a coiled, slender
tube. In one damaged specimen, tubes
squashed out from sacs, appearing as long
penes (Fig. 2C, pe), but it appears as if tips
of these are normally located inside inner
ends of sacs rather than at mid-ventral at-
tachment point in body wall. It is thus un-
known whether tubes are protrusible (.e.,
penes) or eversible structures (1.e., pseudo-
penes).

Remarks.—Although there is a possibil-
ity that the genital body in segment XII is
used for storing sperm from a concopulant
(see above), typical spermathecae were not
observed.

Several points in this description need to
be clarified by future studies. Nevertheless,
Parvidrilus strayeri is easily separated from
all other known oligochaete taxa. See Dis-
cussion below.

Distribution and habitat——Known only
from the type locality in Alabama, USA.
Interstitial groundwater.

Discussion

Parvidrilus strayeri is clearly a clitellate.
It appears to be hermaphroditic (the geni-
talia of all specimens look the same) with
gonads in specific segments, and it has an
eversible, thickened, pharynx roof, a brain
located behind the prostomium, and large
glandular epidermal cells in the genital re-
gion suggesting the existence of a clitellum
(see Purschke et al. 1993). In a traditional
sense, the worm is a member of the Oli-
gochaeta; but there is now increasing evi-
dence that this taxon is paraphyletic unless
leeches and leech-like groups are included
(Purschke et al. 1993, Brinkhurst 1994, Sid-
dall & Burreson 1996, Ferraguti & Erséus
1999). A formal phylogenetic analysis to
establish the more specific systematic po-
sition of Parvidrilus within the Clitellata, is
not meaningful until some uncertainties in
the description have been clarified. As there
is yet no strong indication as to which cli-
tellate subgroup the new species belongs,

VOLUME 112, NUMBER 2

outgroups for such an analysis would need
to be selected from outside the Clitellata,
implying that most ingroup and outgroup
character states would not be comparable.

In some ways, however, the new species
is unusual and unique (see Table I). First, it
is one of the smallest clitellates known.
With a body length of about 1 mm it resem-
bles only the smallest Chaetogaster spp.
(Naididae) (see Sperber 1948), an undes-
cribed freshwater species tentatively as-
signed to Capilloventer (Capilloventridae)
from Western Australia (Pinder & Brink-
hurst 1997b); and a miniature terrestrial
species of Enchytraeidae from Italy, Mar-
ionina eleonorae Rota, 1995. Some marine
oligochaetes are just slightly longer. Ex-
amples within the 1.5—2.5 mm range are nu-
merous species of the subfamily Phallodri-
linae (Tubificidae; see, e.g., Erséus 1980,
1989, 1990, 1992), and Randiella caribaea
Erséus & Strehlow, 1986 and R. minuta Er-
séus & Strehlow, 1986 (Randiellidae; see
Erséus & Strehlow 1986, Erséus 1997). To
my knowledge, there is no freshwater tu-
bificid that is even close to the 0.7—1.4 mm
range that characterizes Parvidrilus.

Second, in Parvidrilus strayeri, the four
chaetal bundles of each segment are located
in a more posterior position within the seg-
ment than in other oligochaetes. Third, the
clitellar cells are large in relation to the
body diameter, few in number, and restrict-
ed to two lateral rows, one at each side,
through a few segments of the worm. These
features are likely to be autapomorphies of
Parvidrilus or of a group including as yet
unknown taxa.

Although complete male ducts have not
been observed in Parvidrilus, the position
of the genital bursa indicates that male
pores (or an unpaired male pore?) are pres-
ent in segment XII. In this respect the new
taxon bears resemblance to four other mi-
crodrile families, Enchytraeidae, Propappi-
dae, Capilloventridae and Phreodrilidae,
and a few representatives of the Lumbri-
culidae. Capilloventridae and Phreodrilidae
are regarded as endemic to the Southern

333

Hemisphere (although a few phreodrilids
occur north of the equator), but a further
comparison with them is pertinent here as
both (see Harman & Loden 1984; Erséus
1993; Pinder & Brinkhurst 1997a, 1997b)
are characterized by absence of chaetae in
segment II (for phreodrilids, at least with
regard to the dorsal chaetae; Pinder &
Brinkhurst 1997a), and presence of hair
chaetae in the other segments, i.e., two
striking similarities with the chaetal pattern
in Parvidrilus (Table I).

The additional feature of ventral hair
chaetae is, within the Clitellata, shared only
by Parvidrilus and Capilloventridae and
may be homologous. However, Capilloven-
tridae, as currently defined (Pinder &
Brinkhurst 1997b), is characterized by
modified hair-like genital chaetae, sperma-
thecae in segment VII, and a pair of blind
ventral sacs (salivary glands?) opening into
the mouth cavity (the two first features also
known from the Randiellidae; see Erséus
and Strehlow 1986). As none of these is
present in Parvidrilus, inclusion of the new
genus in Capilloventridae would imply re-
ductions of all three traits, which is a less
parsimonious hypothesis than to regard
Parvidrilus as a taxon outside Capilloven-
tridae. Moreover, in Capilloventridae but
not in Parvidrilus, the chaetal bundles of
each side are located close together and
widely separated from those of the other
side, somewhat like the distribution of
chaetae on parapodia of polychaetes (Er-
séus 1993). This could be a plesiomorphic
condition reflecting ancestry of the Clitel-
lata among the polychaetes (cf. Westheide
1997), and in which case the Capilloventri-
dae could be the most ancestral group of all
Clitellata; alternatively, it is an additional
autapomorphy of Capilloventridae.

The Phreodrilidae comprises a total of
about 60 species with great morphological
variation with regard to chaetae as well as
genital organs (Pinder & Brinkhurst 1997a).
In phreodrilids, the dorsal chaetal bundles
frequently consist of long hair-like chaetae
accompanied by short, single-pointed, “‘lat-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

334

juosqe juosqe juosqe yuasoid juosqe sovs [eoong jenUsA

x ul IIA Wl Tx ut IIA Ut juosqe svooyjeulods

IX ul IA Ul Ix ul Ix ul Ix ul sorod oe

Apoq sutpuno.ins Apoqg surpuno.zins Apoq surpuno.ins Apoqg surpunoiins SMO [eIO}e] SV S][99 IeTONTO
eTpunq

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(o[qvorjdde jou)

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juviognjoid
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yuosoid (o]qvorjdde jou) juasqe SOUITJOUIOS juosqe juasqe [J Ul ovjoryD

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VOLUME 112, NUMBER 2

eral support chaetae’’, an arrangement sim-
ilar to that found in Parvidrilus strayeri
(but also similar to the dorsal ‘hairs and
needles’ in species of Naididae and Opis-
tocystidae). The crucial point would then be
to what extent the curious genital body and
copulatory organ of the new species are ho-
mologous to phreodrilid genital structures.
They show some superficial resemblance to
the atria and eversible pseudopenes, respec-
tively, of the species of Phreodrilus Bed-
dard (see Pinder & Brinkhurst 1997a), al-
though in Parvidrilus the two kinds of
structures do not appear to be continuous
with each other as the atria and pseudope-
nes are in Phreodrilus. Moreover, the
phreodrilid atria are filiform invaginations,
lying freely in the coleomic cavity, whereas
the Parvidrilus genital body is in very close
contact with (merely as a pair of swellings
of) the body wall. There are also other dif-
ferences between Phreodrilidae and Parvi-
drilus (Table I). In phreodrilids, the general
body size is one order of magnitude larger
than that of Parvidrilus, the ventral chaetae
are consistently paired, there is a thin cli-
tellum of normal microdrile type and it is
generally restricted to segments XII—XIII,
there are spermathecae in segment XIII, and
if hairs are present, the lateral support chae-
tae do not usually project from the chaetal
sacs. Many of these features are shared with
other aquatic clitellate taxa, but the lateral
Support chaetae, the position of the sper-
mathecae (in segment XIII), and the well
developed tubular atria mentioned above,
are likely to be autapomorphies supporting
monophyly of the Phreodrilidae. Thus, it
does not appear appropriate to place Par-
vidrilus in the Phreodrilidae.

Narapa bonettoi Righi & Varela, 1983,
the single representative of the family Nar-
apidae and known only from the Parana
River in Argentina (Righi & Varela 1983,
Brinkhurst & Marchese 1989), is another
aquatic oligochaete taxon with an unclear
systematic position. It shows some curious
resemblance to Parvidrilus. Although lack-
ing chaetae completely and having its gen-

385

ital system in a more anterior position (Ta-
ble I), Narapa has a male gonoduct (in seg-
ment VI) (Righi & Varela 1983:figs 4—6)
with an outline comparable to one possible
interpretation of the reproductive system in
Parvidrilus. In Narapa, the sperm funnel is
followed by a short vas deferens (not yet
observed in Parvidrilus), leading to a ven-
tral glandular, tubular atrium (possibly cor-
responding to a lateral arm of the ‘genital
body’ in Parvidrilus), followed by a pos-
teriorly bent-over penial sac with a winding
lumen (possibly corresponding to a copu-
latory sac in Parvidrilus). It is also note-
worthy that the ovary (in VII) is reported
to be unpaired in Narapa; in none of the
specimens of Parvidrilus strayeri was more
than one ovary observed. It is an open
question whether these similarities are syn-
apomorphic or convergent, but Narapidae
and Parvidrilidae are both monotypic and
each possesses its own autapomorphies (Ta-
ble I). As currently defined, they can, there-
fore, at most be regarded as sister taxa.
However, relevant comparisons with oth-
er aquatic microdrile taxa have still not
been exhausted. For instance, it could be
suggested that the U-shaped genital body of
Parvidrilus strayeri is a derivative of a pair
of atria of the kind found in the family Tub-
ificidae, and that its position in segment XII
is merely an autapomorphic rearward shift
of a tubificid male system. Following this
line of reasoning, the copulatory organ of
Parvidrilus (Fig. 2C) could be interpreted
as an indication of a close relationship with,
e.g., Teneridrilus flexus Erséus & Hiltunen,
1990 (in Erséus et al. 1990). The latter is a
North American Great Lakes freshwater tu-
bificid (reported also by Stacey & Hubley
1994), with small, convoluted, tubular pe-
nial organs contained within muscular sacs
(Erséus et al. 1990:fig. 2D, E). However,
these similarities are probably coincident
and convergent. As noted above, the chaetal
pattern and the segmental position of the
genitalia strongly suggest that Parvidrilus is
more closely related to the Capilloventridae
and Phreodrilidae than to the Tubificidae

336

(Table I). Furthermore, the tubificid atria
extend freely into the coelom, rather than
being incorporated in the body wall as is
the case with the genital body of Parvidri-
lus.

To conclude, it seems justified to estab-
lish a higher level taxon, Parvidrilidae, new
family, for Parvidrilus on the basis of the
available morphological evidence. This
classification is not likely to render any of
the other clitellate families paraphyletic.
Several features of Parvidrilus/Parvidrili-
dae are probably autapomorphic: the ex-
treme miniaturization, the chaetal bundles
situated posteriorly in each segment, the cli-
tellum modified into two lateral rows of a
few large glandular cells, and the unique,
complex, genital organs surrounding a con-
spicuous mid-ventral bursa in segment XII.

In its area of distribution, Parvidrilus is
part of a particular association of inverte-
brates (‘stygobionts’) adapted to life in in-
terstitial groundwater, an association which
also includes the polychaete Troglochaetus
sp., aeolosomatids (Annelida, Aphanoneu-
ra), smaller lumbriculid (e.g., Stylodrilus
wahkeenensis Rodriguez & Coates, 1996)
and naidid oligochaetes, bathynellacean
crustaceans, microcerberid isopods and nu-
merous benthic cyclopoid copepods (Rod-
riguez & Coates 1996, Strayer et al. 1995,
Strayer & Reid in preparation, Reid et al.
in preparation). Although still greatly over-
looked, this rich stygobiont fauna appears
to characterize a vast area of unglaciated
ancient terrain in eastern North America.
Some authors (e.g., Wagele et al. 1995,
Reid 1998) have already pointed out that
many endemic interstitial microcrustaceans
of this region appear to represent an old
continental fauna, with closely related, but
anatomically distinct Eurasian counterparts.
It is possible that Parvidrilus is a northern
representative of a group of aquatic clitel-
lates, which also contains the Southern
Hemisphere families, Capilloventridae,
Phreodrilidae and, possibly also, Narapidae.

Parvidrilus was common at Hendrick
Mill (D. L. Strayer, pers. comm.) and is

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

probably not a rare animal, once the right
habitats are searched.

Parvidrilus strayeri deserves further at-
tention. New material needs to be studied
by methods that would enable a detailed
scrutiny of the true nature of its miniature
genital organs, which may throw additional
light on its systematic position. Molecular
systematic analyses of this enigmatic taxon
may contribute towards the same end.

Acknowledgments

I am indebted to Dr. David L. Strayer, for
generously placing the material at my dis-
posal, and for sharing published and un-
published information; to Dr. Anders Warén
(SMNH) for invaluable assistance with
scanning electron microscopy; to Ms. Bar-
bro L6fnertz (University of Géteborg), Mrs.
Anna Hedstrém and Ms. Christine Hammar
(both SMNH), for technical assistance; to
Dr. Ralph O. Brinkhurst (Hermitage, TN,
USA) and Dr. Kathryn A. Coates (Bermuda
Biological Station for Research, Inc), for
constructive criticism of the text; and the
Swedish Natural Science Research Council,
for financial support.

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, & M. Marchese. 1989. Guide to the fresh-
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Erséus, C. 1980. Taxonomic studies on the marine gen-
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atlanticus gen. et sp. n., a member of a new
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14:7-15.

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Stacey, D. F, & B. R. Hubley. 1994. New records of
a sludge worm Teneridrilus flexus Erséus &
Hiltunen (Oligochaeta: Tubificidae) from Lake
Huron.—Canadian Field. Naturalist 108:182—
185.

Strayer, D. L., S. E. May, P. Nielsen, W. Wollheim, &
S. Hausam. 1995. An endemic groundwater fau-
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Wagele, J. W., N. J. Voelz, & J. V. McArthur. 1995.
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America and erection of Coxicerberus, new ge-
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1-15.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(2):338-351. 1999.

On the entocytherid ostracods of the Brazos River basin and
adjacent coastal region of Texas

Daniel J. Peters and Jean E. Pugh

(DJP) New Horizons Governor’s School for Science & Technology, 520 Butler Farm Road,
Hampton, Virginia 23666, U.S.A.;
(JEP) Christopher Newport University, 6640 Fields Landing Road, Hayes, Virginia 23072, U.S.A.

Abstract.—New records of entocytherid ostracods infesting crayfishes are
recorded from the Brazos River basin in southeastern Texas, which extend the
ranges of Ankylocythere ancyla, A. sinuosa, Entocythere harrisi, E. reddelli,
and Uncinocythere simondsi. A review of the variations in the copulatory com-
plex of A. ancyla is provided and the synonymy of A. tiphophila with A.

sinuosa is proposed.

Upon the completion of his study of the
crayfishes of the Brazos River basin and
nearby areas, Douglas W. Albaugh (1973,
1975; Albaugh & Black 1973) sent sedi-
ments from the containers in which his
specimens were preserved to the Smithson-
ian Institution. This report is based on the
ostracods contained in these samples which
were made available to us through the kind-
ness of the late Horton H. Hobbs, Jr.

Five species of entocytherid ostracods
were retrieved from 118 samples collected
between 21 February and 8 March 1973.
Hosts, which were identified by Dr. AI-
baugh (the subgeneric assignments were
provided by Horton H. Hobbs, Jr.), include
the following: Cambarellus (Dirigicamba-
rus) shufeldtii (Faxon), C. (Pandicambarus)
ninae Hobbs, C. (P.) puer Hobbs, Fallicam-
barus (Creaserinus) fodiens (Cottle), Pro-
cambarus (Capillicambarus) brazoriensis
Albaugh, P. (C.) hinei (Ortmann), P. (C.)
incilis Penn, P. (Girardiella) simulans
(Faxon), P. (Ortmannicus) a. acutus (Gi-
rard), P. (O.) texanus Hobbs and P. (Sca-
pulicambarus) clarkii (Girard). Insofar as
we have been able to determine, all of the
collections were made from open water; no
crayfishes were taken from burrows. The
ostracods, locality data, host and entocy-
therid identifications are deposited at the

National Museum of Natural History,
Smithsonian Institution, Washington, D.C.

Ankylocythere ancyla Crawford
Figs. la—h, 2a—h, 4

Ankylocythere ancyla Crawford, 1965:148,
149, 152, 153, figs. 1-3, 6, 7 [Type lo-
cality: “‘... in the city limits of Greens-
boro, Guilford County, North Carolina.”
Types: holotype, allotype, morphotype,
and dissected male paratype USNM;; par-
atypes in the collection of E. A. Craw-
ford, Jr., and USNM. Host: Cambarus la-
timanus (LeConte) (=Cambarus (De-
pressicambarus) catagius Hobbs & Per-
kins, 1967)].

Ankylocythere species g Hobbs III, 1969:
32-34, figs. 4d—g, 1.

Ankylocythere species h Hobbs III, 1969:
34-36, figs. 4h—k, 1.

Except for the omission of the references
to Hobbs III (1969), Andolshek & Hobbs
(1986:10) included a compilation of all ref-
erences in the literature to this ostracod.

Diagnosis of Texas material.—Shell
length of male 336 to 378 (avg. 357) um;
shell height 196 to 224 (avg. 218) wm. Pen-
iferum truncate to tapering with acute an-
tero- and posteroventral angles. Clasping
apparatus L-shaped with vertical ramus lon-

VOLUME 112, NUMBER 2 359

——>

Fig. 1. Variation in penifera, lateral view, of Ankylocythere ancyla in Brazos River basin (a, h—Refugio
Co.; b, f—Washington Co.; c, g—Fort Bend Co.; d—Austin Co.; e—Brazoria Co.); scale 0.02 mm.

340

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2.

Variation in clasping apparatus, lateral view, of Ankylocythere ancyla in Brazos River basin (a,

h—Refugio Co.; b, e, g—Washington Co.; d—Fort Bend Co.; f—Austin Co.; c—Brazoria Co.); same scale

ASHE TE. by.

ger than horizontal ramus; latter with tooth
on preaxial border near midlength, and
postaxial border with long, curved talon
reaching to midway between preaxial tooth
and apex of ramus; extreme apical part of
talon sometimes strongly curved mesially.
Apex of clasping apparatus with 2 denti-
cles.

Range.—Andolshek & Hobbs (1986)
quoted the range of the species as cited by
Hobbs & Peters (1977) as extending “...

along the Atlantic and Gulf slopes from the
Mobile River drainage in Alabama and
Mississippi northeastward to the Potomac
drainage in Virginia and in the New River
Basin of North Carolina.”’ The locality re-
cords established herein extend the west-
ward limits some 750 kilometers to the
Brazos basin in Texas.

Southeastern Texas records (Fig. 4).—
Nineteen localities from the following
counties: Austin (2), Brazoria (2), Brazos

VOLUME 112, NUMBER 2

(2), Calhoun (3), Fort Bend (2), Matagorda
(1), Refugio (6), and Washington (1).

Remarks.—In his study of the entocy-
therids infesting burrowing crayfishes in the
coastal plain between extreme eastern Texas
and the Apalachicola Basin in Alabama and
Florida, Hobbs III (1969) recognized 12
species belonging to the genus Ankylocy-
there. Specimens of two of these, A. species
‘9’? and “‘h”’ were lent to us by Dr. Hobbs,
and, after comparing them with represen-
tatives of A. ancyla from throughout its
range, we are convinced that they are re-
ferable to this species. Thus there are few
gaps in its known range. Andolshek &
Hobbs (1986:14) reviewed available data
on the size of the shell of this ostracod and
found that the smallest individuals occurred
in southeastern Georgia and the largest in
Virginia. Those in North Carolina were, on
the average, intermediate in size. The range
in size of the material from along the coasts
of Florida, Alabama, Mississippi, Louisi-
ana, and Texas reported herein falls within
that cited for the species in southeastern
Georgia (length, 315 to 399 um; height 175
to 245 wm). Thus it appears that the shells
of southern populations of A. ancyla are
smaller than those occurring in the more
northern parts of the range.

A cursory examination of the ventral part
of the peniferum would suggest that a di-
morphic condition exists in this appendage,
one in which the anteroventral extremity is
produced in a subspiculiform prominence
(Fig. 1d), and in the other, subtruncate (Fig.
le). That the difference is more in the angle
from which the penifera are viewed rather
than due to morphological variation be-
comes apparent when those of a number of
specimens are compared. In this ostracod,
the basic structure of the ventral extremity
of the peniferum is more clearly observed
in specimens from the eastern part of the
range where a broad concavity exists be-
tween the acute cephloventral and rounded
posteroventral extremities (See fig. 4a in
Andolshek & Hobbs (1986)). In specimens
from the Brazos region, the anteroventral

341

angulate extension often appears to be more
strongly produced, and its base to bear a
thickened, sclerotized prominence, which
when viewed at some angles, seems to pro-
ject posteriorly or posteroventrally; also the
anteroventral apex of the posteroventral
prominence is procurved, diminishing the
maximum diameter of the concavity, and is
frequently rather strongly sclerotized. (For
variations compare Fig. 1b, c, e, g, h). Thus,
whereas the ventral extremity of the peni-
fera of the eastern and western members of
the species appear to be markedly different,
the contrast is less marked than seems ap-
parent when only a superficial comparison
is made.

Among other variations noted in the cop-
ulatory complex of this ostracod are the
thickness of the junction of the horizontal
and vertical rami of the clasping apparatus
and the curvature of the vertical ramus. In
specimens from Washington County, TX,
males were found that possess a sinuous
vertical ramus (Fig. 2e), and in the area of
the junction of the rami there occurs a con-
spicuous thickening (Fig. 2e). Even though
different, this variance must be considered
to be within the range of variation in the
species. For example, in one specimen one
of the pair of clasping apparati exhibits
such a thickening and the other resembles
the more frequently observed apparatus
(Fig. 2f, g). Considerable variation occurs
in the curvature of the vertical ramus and
in that of the talon (Fig. 2a—h).

Hosts.—In the Brazos Basin, A. ancyla
is known to infest F. (C.) fodiens, C. (P.)
ninae, and P. (S.) clarkii, and has been re-
trieved from collections containing all of
the crayfishes known to occur in the area
except C. (P.) texanus, P. (C.) hinei, and P.
(C.) brazoriensis; rarely, however, was it
found in collections containing representa-
tives of C. (P.) puer, C. (D.) shufeldtii, and
P: (C.) incilis.

Entocytherid associates.—In the 19 lo-
calities in which this ostracod was found, it
was the only one infesting the crayfishes in
nine sites. Its most frequent associate (in 9

342

localities) was A. sinuosa, and only rarely
was it found with E. reddelli (2), and U.
simondsi (1).

Ankylocythere sinuosa (Rioja)
Figs: 3.3

Entocythere cambaria.—Hobbs, 1941:4 [in
part].

Entocythere (Cytherites) heterodonta sin-
uosa Rioja, 1942a:203, 204, figs. 5, 6
[Type locality: La Cueva Chica, San Luis
Potosi, Mexico. Types: not extant. Host:
Procambarus acutus cuevachicae Hobbs
1941.]; 1953:287.

Entocythere (Cytherites) sinuosa.—Rioja,
1942b:688, 689, 695, 696, fig. 20; 1943a:
564; 1943b:576.

Entocythere sinuosa.—Hoff, 1944:330,
3322 SIG=-Rioja) 1949371 S22 Flin
partijehigs: 1Sy 1L4) 19st 7Os19532298,
292.—Tressler, 1954:138; 1959:731, fig.
28.190.—Hobbs, 1957:431.—Crawford,
LOSS -WAIBANTS:

Ankylocythere sinuosa.—Hart, 1962:127;
1964:246.—Crawford, 1965:149.—Red-
dell, 1965:156; 1970:395; 1971:18; 1981:
82.—Hobbs, 1966:70, fig. 18; 1971:34-—
5 Jsatlies022:=—Fersuson,y 1968-501
Hobbs & Walton, 1968:246.—Baker,
1969:293.—Reddell & Mitchell, 1969:6;
1971:142.—Young, 1971:399—409.
Hart & Harty 974) 2; 44, 21,.22,.29=
31, 34, pl. 3: figs. 11-13, pl. 41.—Hobbs
III, 1969:5, 14, 20—22, 27, 30, 32-35, 39,
41, 43, 46, 55, 65, 66, 71, 74, 78, 79, fig.
Sa—k; 1975:281, 290; 1978:506; 1982:
2.—Hobbs & Peters, 1977:13; 1991:66,
67.—Hobbs & McClure, 1983:773.

Entocythere tiphophila Crawford, 1959:
150, 1515. 173=17 82) 80phSie nies. Sil=37
[Type locality: roadside ditch 9.1 miles
(14.6 km) SE of University of South Car-
olina stadium, Richland County, South
Carolina, on St. Rte. 48. Types: USNM.
Hosts: Fallicambarus (C.) uhleri and
Procambarus (F.) troglodytes|.—Hart,
£96721235 128;

Ankylocythere tiphophila.—Hart, 1962:128;

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1964:245.—Crawford, 1965:149.—
Hobbs, 1966:71, fig. 16.—Ferguson,
1968:501.—Peters, 1974:74; 1975:ii1, 5—
8; 103:13,.20) 22; 23; 27, 45) figs: Zeek
14.—Hart & Hart, 1974:15, 20, 33, pl. 4:
figs. 6-8, pl. 45.—Hobbs & Peters, 1977:
iu, 3-7, 9, 12, 16, 19-22, 28, 40, 41, 43,
46, 49-54, 70, fig. 5, map 5.—Hobbs,
1981:140, 499, 501.—Hobbs & McClure,
1983:773.

Ankylocythere tiphophyla.—Hobbs, Holt, &
Walton, 1967:77 [erroneous spelling].

Diagnosis of Texas material.—Shell
length of male 329-378 (avg. 350) wm;
shell height 168—210 (avg. 191) wm. Peni-
ferum varying from deeply cleft to truncate
with tapering acute anteroventrally project-
ing prominence. Clasping apparatus L-
shaped with vertical ramus longer that hor-
izontal ramus; latter with truncate, almost
straight, anteroventrally projecting talon sit-
uated slightly proximal to midway between
preaxial tooth and apex of ramus; apex of
apparatus with 2 denticles.

Range.—On the Gulf of Mexico versant,
from the Cordillera volcanica Transversal
along the Gulf and Atlantic (lower pied-
mont and coastal plain) slope to the York
River Basin in Virginia. Hart and Hart
(1974:33) also reported it from two locali-
ties in Ohio, records that should be con-
firmed.

Southeastern Texas records (Fig. 5).—
This ostracod is the most widespread of the
entocytherids within the study area, occur-
ring in 94 of the 118 localities represented
among the collections examined.

Hosts.—In southeastern Texas this ostra-
cod was associated with three crayfishes: P.
(O.) a. acutus, P.. (S:) clarkit andsPa(G)
simulans, but has been found in collections
containing specimens of all of the other
species in the area. In the collections from
94 localities where the ostracod was found,
P. (O.) a. acutus was a potential host in 60
of them, P.-G:) clarkiiiin 52;'P2(Gies
mulans in 31, P. (C.) incilis in 17, and F.
(C.) fodiens in 14. All of the other crayfish-

VOLUME 112, NUMBER 2 343

Fig. 3. Variation in male copulatory complex, lateral view, of Ankylocythere sinuosa (a—Tamaulipas, Ace-
quia, Mexico; b—Burleson Co, TX; c—Robertson Co., Texas) and Ankylocythere tiphophila (d—Dorchester
Co., South Carolina; e—Newport News, Virginia); same scale as Fig. 1.

es were present in fewer than 10 of the col- host(s) with E. reddelli, in eight with A. an-

lections. cyla, and two each with E. harrisi and U.
Entocytherid associates.—In the 94 lo- simondsi.
calities where A. sinuosa was found, it was Remarks.—There is nothing remarkable

the only ostracod infesting the crayfish(es) concerning the size of the animals or in the
in 57 of them. In 26 localities it shared the structures employed in distinguishing this

344 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

@ A. ancyla

A JU. simondsi

Fig. 4. Distribution of Ankylocythere ancyla and Uncinocythere simondsi in the Brazos River drainage.

ostracod from its congeners. The length of in Mexico, cited similar ranges in size: 0.34
the shells of males ranges from 329 to 378 to 0.37 mm and 0.19 to 0.22 mm, respec-
wm and the height from 168 to 210 pm. tively.

Hobbs (1971), in reporting on this ostracod Perhaps because of the apparent discon-

VOLUME 112, NUMBER 2

@ A. sinuosa

345

Fig. 5. Distribution of Ankylocythere sinuosa in the Brazos River drainage.

tinuity between the ranges of A. sinuosa
and A. tiphophila and the seemingly con-
sistent presence of a cleft peniferum in the
range of the former and its absence in that

of the latter, there seemed to be no reason
to suspect that they represented two forms
of a single species. With the acquisition of
collections almost merging the ranges of

346

the two species, we were prompted to com-
pare specimens from Mexico to Virginia
(Fig. 3) and found that in the Gulf Coastal
area forms with a cleft peniferum, the only
feature that has served consistently to sep-
arate the two (Fig. 3a, c), occur in localities
in which some of the males were lacking a
cleft (Fig. 3b, d, e). Moreover, in specimens
from Bleckley County, Georgia (see An-
dolshek & Hobbs, 1986:fig. 12a), the pen-
iferum is distinctly excavate, tending to-
ward the cleft condition. In view of the dis-
covery of an almost continuous range and
no character that can be relied upon invari-
ably to separate the two, we propose that
A. tiphophila be considered a synonym of
Rioja’s Entocythere sinuosa.

Reinforcing this proposal is the seeming-
ly subparallel clinal distribution with re-
spect to the size of the shell. Andolshek &
Hobbs (1986:25) reviewed in tabular form
the shell size reported for A. tiphophila re-
vealing that the largest specimens occur in
North Carolina (410 pm) and Virginia (390
wm), smaller ones in South Carolina (346
im), and the smallest in southeastern Geor-
gia (321 pm). Perhaps significant are the
sizes reported by Hobbs III (1969:74) for
A. sinuosa occurring from eastern Texas to
the panhandle of Florida. The mean shell
length is 323 wm as compared with one of
321 wm in the material from southeastern
Georgia. The cline, however, seems to be
reversed between eastern Texas and Mexi-
co, for the mean shell length for specimens
from Mexico was reported by Hobbs (1971:
36) to be 350 pm.

Entocythere harrisi Peters
Fig. 6

Entocythere harrisi Peters, 1975:32-33,
figs. 5a, 6e, f, 7a [Type locality: Rocky
Creek 4.3 mi (6.9 km) E of U.S. Hwy 29
on U.S. Hwy 60, Amherst County, Vir-
ginia. Types: holotype and allotype,
USNM; paratypes, USNM, H.H. Hobbs
III, and DJP. Hosts: Cambarus (C.) bar-
tonii bartonii (Fabricius), C. (Hiaticam-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

barus) longulus (Girard), and C. (P.) ac-
uminatus Faxon].—Hobbs & Peters,
L977TaAvy SeOf 1214, 21, 29533, 362248
45, 47, 51, 52, 54, 55, 60, 61, 64, fig. 25;
1982:314; 1989:328.—Andolshek &
Hobbs, 1986:30. [The references cited
here constitute a complete bibliography
for the species.]

Diagnosis.—Shell length of male 441—
570 (avg. 477) wm; shell height 210—300
(avg. 244) um. Peniferum truncate distally.
Clasping apparatus “‘with postaxial border
[slightly] bowed into heellike prominence
at junction of horizontal and vertical rami,
junction thickened; mesial surface of area
of junction without flange; horizontal ramus
without oblique ridges on mesial surface”
(Hobbs & Peters 1977:51).

Range.—The most recent summary of
the range of this ostracod was that of Hobbs
& Peters (1977:52) who based their records,
except for the type locality, on female spec-
imens. In the present study we have become
convinced that the characters they used for
distinguishing between the females of this
species and those of their E. internotalis
(=E. elliptica Hoff, 1944) are not reliable.
This conclusion is based upon the obser-
vation that in two collections of E. harrisi
(one from Pike County, Arkansas, and an-
other from Angelina County, Texas) con-
taining several male and females, none of
the latter possess the type of genital appa-
ratus similar to those that were identified
with that species in North Carolina and Vir-
ginia. Instead, the genital apparatus of the
Arkansas and Texas females are indistin-
guishable from those of £. elliptica and E.
reddelli. Thus we believe this ostracod is
represented in collections only by the ho-
lotype (from Amherst County, Virginia)
and the specimens cited herein from the
Brazos River basin, Texas.

Remarks.—With the new records cited
herein for the males of this species, we are
inclined to propose a clinal distribution in
size with respect to shell height. As the
largest member is reported from Virginia

VOLUME 112, NUMBER 2

A. OH) harrisi

@ £. reddelli

347

Fig. 6. Distribution of Entocythere harrisi and Entocythere reddelli in the Brazos River drainage.

(300 wm), smaller members occur from
Pike Co., Arkansas (265 wm) and the Braz-
os River drainage (256 wm), with the small-
est (224 wm) appearing in Angelina, Texas.

However, when the shell length is com-
pared, the largest member remains in Vir-
ginia (570 wm) and the smallest (456 wm)
in Angelina, Texas, whereas members from

348

the Brazos River basin (490 wm) and Ar-
kansas (476 wm) appear reversed. This may
be due to the small number of specimens
available for examination.

Southeastern Texas records (Fig. 6).—A
male from each of the following localities:
4.0 mi (6.4 km) E of Marlin, Falls County,
and McLaughlin Creek, 12.5 mi (20 km) E
of Cameron on U.S. 90 at Branchville, Mil-
an County.

Hosts.—In both localities this ostracod
was found on P. (G.) simulans and P. (O.)
a. acutus.

Entocytherid associates.—A. sinuosa
was present in the two Brazos localities.

Entocythere reddelli Hobbs & Walton
Fig. 6

Entocythere reddelli Hobbs & Walton,
1968:243-246, fig. 2a—d [Type locality:
Golden Fawn Cave, 8 mi (12.8 km) NNE
of Boerne, Kendall County, Texas.
Types: holotype, allotype, and paratypes,
USNM;; paratypes, Academy of Natural
Sciences of Philadelphia and H.H. Hobbs
II. Host: P. (S.) clarkii (Girard)].—Red-
dell & Mitchell, 1969:6.—Reddell, 1970:
395.—Hobbs, 1971:40.—Hart & Hart,
1974:83, 92—93, pl. XXVII: figs. 11-15,
pl. LIl—Hobbs III, 1975:281.—Hobbs
& Peters, 1977:iv, 5—7, 9-10, 12, 14, 29,
30, 41,43. 45; 47, 52, 54,95, 73; 1982:
313.—Andolshek & Hobbs, 1986:30.
[These references constitute what we be-
lieve to be a complete bibliography for
the species. ]

Diagnosis.—Shell length of male 546—
581 (avg. 564) wm; shell height 280—294
(avg. 287) wm. Peniferum subtruncate ven-
trally and possessing rounded antero- and
posteroventral extremities. “‘Clasping ap-
paratus with postaxial border at junction of
horizontal and vertical rami produced in
heellike prominence and junction thick-
ened; mesial surface of area of junction
with angular flange, apex of angle reaching
level proximal to proximal tooth on preax-
ial margin of horizontal ramus, horizontal

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ramus lacking long oblique ridge extending
across mesial surface. Female genital ap-
paratus composed of ventrally directed sub-
spiculiform projection arising from bipartite
base, latter embedded in amorphous mass”’
(Hobbs & Peters 1977:55), second antenna
with appendix at base of terminal claws
conspicuously enlarged, pectinate, bearing
2 or 3 broad teeth or as many as 6 finely
divided denticles.

Range.—In addition to its presence in
Kendall County, Texas, Hart & Hart (1974:
93) reported its occurrence in Greene Coun-
ty, Arkansas, and Sumner County, Kansas.
Hobbs & Peters (1977:55) also reported it
from the Catawba, French Broad, and Hi-
wassee basins in North Carolina. Twenty-
five localities in the Brazos River basin of
southeastern Texas are cited here.

Southeastern Texas records (Fig. 6).—
Twenty-five localities from the following
counties: Brazoria (8), Brazos (1), Burleson
(1), Fort Bend (7), Matagorda (4), Milan
(1), Robertson (2), and Victoria (1).

Remarks.—Except for the distribution of
this ostracod in the Brazos River basin, its
range is poorly known. The total absence of
records in what appears to be the central
part of its range is disturbing, but there is
little reason to suspect that the type locality
lies on its western limit, and if and where
its range and that of E. cambaria Marshall
and E. illinoisensis Hoff meet or intersect
to the north have not been determined.

Hosts.—In the Brazos basin, E. reddelli
was found infesting P. (O.) a. acutus and
P. (S.) clarkii. It also occurred in collections
containing one or both of these crayfishes
along with one or more of the following:
C..(D.) shufeldti, C. (P.), ninaex GG
puer, C. (P.) texanus, F. (C.) fodiens, P.
(C.) brazoriensis, P. (C.) incilis, and Ea (G)
simulans.

Entocytherid associates.—In all 25 lo-
calities in which this ostracod was found, it
was associated with A. sinuosa, and in one
of them A. ancyla (hosts: P. (O.) a. acutus
and P. (S.) clarkii) was also present; in an-

VOLUME 112, NUMBER 2

other of these localities U. simondsi was in-
festing the same two host species.

Uncinocythere simondsi Hobbs & Walton
Fig. 4

Entocythere simondsi Hobbs & Walton,
1960:17, 20-21, figs. 1-10 [Type locali-
ty: Dunn Creek, 1.9 mi (3 km) west of
Fighting Town Creek on Hell’s Hollow
Road, Fannin County, Georgia. Hosts: C.
(C.) bartonii and C. sp. (=C. (D.) lati-
manus)}.

Uncinocythere simondsi.—Hart, 1962:138.

[A complete bibliography for the species
is presented by Andolshek & Hobbs,
1986:39.]
Diagnosis.—Shell length of male 329—
343 (avg. 336) wm; shell height 154-196
(avg. 175) pm. Copulatory complex of
peniferum terminating distally in bifid
tip. Clasping apparatus L-shaped with
preaxial border bearing 3 distinct teeth;
postaxial border entire, lacking any ex-
crescence; distal extremity with 3 denti-
cles.

Range.—From Illinois, Kentucky, and
North Carolina southward to Brazoria and
Washington counties, Texas, and northern
Florida, previously known no farther west
than Mississippi (Hart & Hart 1974; Hobbs
& Peters 1977, 1982; Andolshek & Hobbs
1986).

Southeastern Texas records (Fig. 4).—
This ostracod was found in the following
localities: 2 mi (3.2 km) N, 2 mi (3.2 km)
E of Brenham on St Rte 90, Washington
County; and 1.25 mi (2 km) E of Rosharon
on Farm Rd 1462, Brazoria County.

Hosts.—Procambarus (S.) clarkii was
one of the hosts in both of the Brazos col-
lections in which this ostracod was found.
In one of them, P. (O.) a. acutus was also
present, and in the other, P. (G.) simulans
was in the container from which the ostra-
cods were removed.

Entocytherid associates.—In both of the
localities in which this ostracod was found,
A. sinuosa and E. reddelli also were pre-

349

sent. In the collection from Washington
County, A. ancyla also was found with
them.

Acknowledgments

We extend our thanks to the late H. H.
Hobbs, Jr. who encouraged us in this en-
deavor and provided the materials, guid-
ance, and insight for this study. We are also
grateful to Horton Hobbs III for his criti-
cism of the manuscript and to Brian Ken-
sley, Ray Manning, and Karen Reed for
their assistance.

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567-585. Sons, New York, 1248 pp.

. 1949. Estudios carcinologicos. XXI. Contri- Young, W. 1971. Ecological studies of the Entocyther-
bucion al conocimiento de las especies del Ge- idae (Ostracoda).—American Midland Natural-

nero Entocythere de Mexico.—Anales del In- ist 85:399—409.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(2):352-—361. 1999.

A new genus, Neodoxomysis
(Crustacea: Mysidacea: Mysidae: Leptomysini), with
description of two new species

Masaaki Murano

Institute of Environmental Ecology, METOCEAN Co. Ltd., 1334-5 Riemon,
Ooigawa-cho, Shida-gun, Shizuoka 421-0212, Japan

Abstract.—A new genus, Neodoxomysis, is established for two new species,
N. elongata and N. sahulensis, collected from Sahul Shelf. The new genus is
closely related to Doxomysis, but is different from the latter genus in having
only a single strong, modified seta on the exopod of the fourth male pleopod,
as compared with two in Doxomysis. Doxomysis littoralis Tattersall, 1922, is

transferred to the new genus.

A single male specimen identified by li
(1964) as Doxomysis littoralis Tattersall,
1922, is quite different from the other Dox-
omysis species in having a single strong
modified seta on the exopod of the fourth
male pleopod. The exopod in all other Dox-
omysis species is armed with two strong
modified setae.

During a cruise to southeastern Asian
seas (KH-72-1) by the R/V Hakuho Maru
of the Ocean Research Institute, University
of Tokyo, two undescribed species, which
were similar to D. littoralis with respect to
the morphology of the fourth male pleopod,
were collected. This morphological char-
acter is distinct these species from those of
the genus Doxomysis. Therefore, a new ge-
nus is established to receive these three spe-
cies, and two new species are described.
The type specimens are deposited in the
National Science Museum, Tokyo (NSMT).

Neodoxomysis, new genus

Diagnosis.—Carapace produced anteri-
orly into triangular rostral plate with round-
ed apex. Antennal scale lanceolate with
rounded apex, armed with setae on whole
margins. Eye functionally normally devel-
oped, without papilliform process on eye-
stalk. Maxilla with second segment of en-

dopod wider than long, expanded distally,
with about 10 strong spines on distal mar-
gin. Labrum with rounded frontal margin.
Endopods of third to eighth thoracic limbs
with propodus divided into 2 subsegments.
Endopod of uropod with row of numerous
spines along inner margin. Telson with lat-
eral margin armed with spines throughout,
apical cleft deep, armed with marginal spi-
nules and pair of plumose setae arising
from anterior end. Exopod of fourth pleo-
pod of male modified, longer than endopod,
ultimate segment small, with 1 or 2 short
simple setae, penultimate segment with sin-
gle strong seta.

Type species.—Neodoxomysis elongata,
new genus, new species.

Etymology.—Derived from its relation-
ship to Doxomysis. It is feminine in gender.

Remarks.—The new genus Neodoxomy-
sis closely resembles Doxomysis Hansen,
1912, except for the fourth male pleopod.
In the new genus, the exopod of the fourth
male pleopod is armed with only a single,
strong, modified seta arising from the pen-
ultimate segment, while in Doxomysis as
well as eight related genera in the tribe Lep-
tomysini (Talbot 1997), the exopod is
armed with two strong modified setae, one
arising from the penultimate segment and
the other from the antepenultimate segment.

VOLUME 112, NUMBER 2

Among 11 Doxomysis species in which
the exopod of the fourth male pleopod has
been described, only D. littoralis bears a
single strong modified seta on the exopod.
This species was established on specimens
from the Andaman Islands by Tattersall
(1922); later the male was described from
the South China Sea by li (1964). Accord-
ingly, D. littoralis should be transferred to
the new genus. The new genus Neodoxo-
mysis comprises a total of three species.

Neodoxomysis elongata, new species
Figs: 1f)2

Type _ series.—Holotype (NSMIT-Cr
12487), adult male (5.3 mm), allotype
(NSMT-Cr 12488), adult female with em-
bryos (4.9 mm), paratypes (NSMT-Cr
12489), 6 adult males (6.1—6.8 mm) and 3
adult females (4.5—5.0 mm); Sahul Shelf,
Paes Ss. 4 024°33:9'E to 12°36.0'S;
124°36.4’E; 74-78 m; 25-26 June 1972;
plankton net installed in mouth of 3-m
beam trawl.

Other material.—8 adult males, 3 adult
females, 15 immature males and 19 imma-
ture females; collection data same as type
series.

Description.—Body somewhat slender.
Carapace produced anteriorly into triangu-
lar rostral plate with rounded apex extend-
ing beyond basal margin of antennular pe-
duncles, leaving whole eyes uncovered
(Fig. 1A, B); anterolateral corner rounded;
posterior margin slightly emarginate, leav-
ing last thoracic somite exposed.

Eye developed, relatively large; cornea
occupying more than half of whole organ,
spherical, wider than eyestalk; eyestalk his-
pid on anterior and posterior surfaces, with-
out papilliform process (Fig. 1A, B).

Antennular peduncle of male more robust
than that of female, first segment with sev-
eral setae at outer distal corner and 1 seta
at inner distal corner, second segment short,
narrower than preceding one; third segment

355

as long as first, wider than preceding two
segments, with 1 straight and 4 short
curved setae at inner distal corner; proces-
sus masculinus large, hirsute (Fig. 1A). An-
tennular peduncle of female slender; first
segment as long as succeeding 2 segments
together; second segment short, narrow,
with 1 seta at inner distal corner; third seg-
ment with 1 seta at distal third of inner mar-
gin and 5 setae on inner distal margin (Fig.
1B).

Antennal scale extending beyond distal
margin of third segment of antennular pe-
duncle for about % of its length (Fig. 1A,
B), lanceolate, 6.3 times as long as broad,
outer margin slightly concave, distal seg-
ment Y,, of length of scale (Fig. 1C). An-
tennal peduncle short, reaching proximal
third of antennal scale, 3-segmented, third
segment longest; sympod with thorn at out-
er distal corner (Fig. 1C).

Mandible with well developed mastica-
tory edge. Mandibular palp 3-jointed, sec-
ond segment elongated oval, third segment
0.52 of second in length, outer margin with
2 series of setae, proximal setae longer,
barbed on proximal half, distal setae barbed
on whole length (Fig. 1D). Maxillule: inner
lobe with 9 setae on inner margin, 5 setae
on outer margin, 3 setae on ventral surface
and 3 stout and 1 slender setae on distal
margin; outer lobe with about 10 spines on
apical margin and 3 setae on ventral surface
(Fig. 1E). Maxilla: second segment of en-
dopod expanded distally, wider than long,
distal margin slightly convex, 2.5 times as
wide as at base, armed with 10 stout spines,
which are rounded and flattened at tip, out-
ermost spine longer than others, inner 6
spines, especially 2 innermost ones armed
with short setae on margins; exopod rather
rectangular in shape, extending beyond dis-
tal margin of first segment of endopod (Fig.
1F). Labrum with frontal margin rounded
(Fig. 1G):

Endopod of first thoracic limb robust,
dactylus wider than long, with strong ter-
minal claw (Fig. 1H). Endopod of second

354 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

<a
C7 La
V7

Fig. 1. Neodoxomysis elongata, new species; A, CJ: holotype; B: allotype. A, anterior end of adult male;
B, anterior end of adult female; C, antenna; D, mandible and mandibular palp; E, maxillule; FE maxilla; G,
labrum; H, endopod of first thoracic limb; I, endopod of second thoracic limb; J, sixth thoracic limb.

VOLUME 112, NUMBER 2

va

va

a
>

Vi
)
i

we -

Fie \
= aa /
Sy,

—

—S,
Pee
ene

i
— —
es mes

SS

=
ZA
ca
——
—

Se
a
ee
——
2 —
Ewes

—,
a Sy .
a a)
—

=A
——

——
me

—

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=)

oe
Dp

EE

Hh

:

ws

- _—_—— oo a =

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———.

3 oe

ee

Fig. 2. Neodoxomysis elongata, new species; A—F: holotype; G: allotype. A, first male pleopod; B, fourth

male pleopod; C, distal part of fourth male pleopod; D, fifth male pleopod; E, endopod of uropod; EK uropod
and telson; G, distal part of telson.

356

thoracic limb slender, basis with inner lobe
developed, ischium with long setae on inner
margin, merus as long as carpopropodus
and dactylus combined, dactylus with ter-
minal claw and 6 barbed setae (Fig. 11).

Endopod of sixth thoracic limb slender,
carpus separated obliquely from 2-subseg-
mented propodus, distal propodal subseg-
ment longer than proximal, equal to carpus
in length; dactylus small, with slender ter-
minal claw (Fig. 1J). Exopods of thoracic
limbs with basal plate armed with small
acute process at outer distal corner, flagel-
liform part 8-segmented in first and eight
limbs, 9-segmented in second to seventh
limbs (Fig. 1J).

Abdomen consisting of 6 somites, first 5
somites subequal, sixth somite 1.7 times
longer than fifth.

Male pleopods well developed, natatory.
First pair with 7-segmented exopod and un-
segmented endopod (Fig. 2A). Second and
third pairs with 7-segmented exopod and 6-
segmented endopod extending to distal
margin of sixth segment of exopod. Fourth
pair: exopod 1.6 times longer than endopod,
11-segmented, seventh and eighth segments
unarmed with setae, ninth segment with 1
short seta at outer distal end, tenth segment
with 2 setae on distal end, longer one ex-
tremely long, 0.4 of exopod in length,
Straight, feathered in distal half, shorter one
simple, 0.2 as long as longer one, terminal
segment very small, % of preceding one in
length, with 1 short seta; endopod 6-seg-
mented, reaching distal end of sixth seg-
ment of exopod, without modified setae
(Fig. 2B, C). Fifth pair: exopod 7-segment-
ed, longer than endopod, endopod 6-seg-
mented, with triangular lobe tipped with
seta on outer margin of first segment (Fig.
2D).

Exopod of uropod slender, long, slightly
curved outwardly, extending beyond distal
end of telson for its distal half (Fig. 2F).
Endopod of uropod extending beyond distal
end of telson for distal third, armed along

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

inner margin from statocyst region to near
apex with about 42 spines which are made
up with longer and shorter ones arranged
alternately, except several spines in distal
and proximal portions (Fig. 2E, F).

Telson slightly longer than last abdomi-
nal somite, 1.7 times as long as broad at
base, abruptly narrowing near base, paral-
lel-sided in middle part, then gradually nar-
rowing towards distal end, cleft at apex for
% of telson length; lateral margin armed
along whole length with about 16 spines,
sparsely in proximal third and densely in
distal *%4; each apex of distal lobes rather
truncate, with 3 somewhat obtuse spines,
outermost one longest in male and middle
one in female; cleft with small notch at an-
terior end, each side with about 16—18 spi-
nules along whole length except for poste-
rior short distance naked, pair of plumose
setae arising from anterior end of cleft (Fig.
2E G).

Etymology.—The name elongata refers
to the slender body.

Remarks.—Neodoxomysis elongata is
considerably different from the other two
species of the genus, N. sahurensis and N.
littoralis, in the following aspects: The an-
tennal scale is 6.3 times as long as broad in
this species, while it is about 5 times as
long in the other two species; the exopod
of the fourth male pleopod is 11-jointed in
this species as compared with 7 in the other
two species; and, spines on the second en-
dopod segment of the maxilla are rounded
and flattened distally in this species, while
these are sharply or obtusely pointed in the
other two species.

Neodoxomysis sahulensis, new species
Figs. 3, 4

Type series.—Holotype (NSMT-Cr
12490), adult male (5.0 mm); allotype
(NSMT-Cr 12491), adult female (4.0 mm);
paratypes (NSMT-Cr 12492), 2 adult fe-
males (3.7, 4.0 mm), 1 adult male (divided
into two parts); Sahul Shelf, 12°17.3’S,

VOLUME 112, NUMBER 2 307

Fig. 3. Neodoxomysis sahulensis, new species; A, C—G: holotype; B: allotype. A, anterior end of adult male;
B, anterior end of adult female; C, antenna; D, mandible and mandibular palp; E, maxillule; EK maxilla; G,
labrum; H, endopod of first thoracic limb; I, second thoracic limb.

358 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

a Via

‘
REESE
WANS AN

SSH p~>p3s~
ZZ

<a

Fig. 4. Neodoxomysis sahulensis, new species; A: allotype; B—F: holotype. A, endopod of one of posterior
thoracic limbs; B, endopod of eighth thoracic limb; C, fourth male pleopod; D, distal part of exopod of fourth
male pleopod; E, fifth male pleopod; E uropod and telson.

VOLUME 112, NUMBER 2

129°40.9’E to 12°17.2'S, 129°41.8’E; 49-
52 m; 24 June 1972; plankton net installed
in mouth of 3-m beam trawl.

Description.—Carapace with anterior
margin produced in triangular rostrum with
narrowly rounded apex and almost straight
lateral margins, leaving whole eyes exposed
(Fig. 3A, B); anterolateral corner rounded;
posterior margin somewhat emarginate,
leaving last thoracic somite uncovered.

Eye moderately developed; cornea oc-
cupying about half of whole eye, slightly
wider than stalk; stalk hispid on anterior
and posterior surfaces (Fig. 3A, B).

Antennular peduncle of male more robust
than that of female, first segment as long as
third, with anterolateral corner slightly pro-
duced anteriorly, armed with several setae,
second segment short and narrow, third seg-
ment wider than preceding segments, with
1 short seta at distal third of inner margin
and 2 setae at anteromedian corner; proces-
sus masculinus large (Fig. 3A). In female,
first segment longer than and as wide as
third, second segment with 1 seta at anter-
omedian corner, third segment with 1 seta
at distal third of inner margin and 4 setae
at anteromedian corner (Fig. 3B).

Antennal scale setose on whole margins,
overreaching distal end of antennular pe-
duncle for % of its length (Fig. 3A, B),
more than 5 times as long as broad, outer
margin nearly straight, inner margin convex
in proximal half, distal segment % of length
of scale (Fig. 3C). Antennal peduncle short,
barely extending to middle of antennal
scale, composed of 3 segments, distal 2
equal in length, longer than proximal one;
antennal sympod with long, acute process
at outer distal corner (Fig. 3C).

Mandible with well developed masticatory
edge; mandibular palp with second segment
elongated oval, third segment 0.6 as long as
second, outer margin with 2 series of setae,
proximal setae longer, barbed on proximal
half, distal one barbed along whole length
(Fig. 3D). Maxillule: inner lobe with 7 setae
on inner margin, 4 setae on outer margin and

359)

3 stout and 1 slender setae on apical margin;
outer lobe with about 10 spines on apical
margin, about 10 spinules on middle part of
outer margin and 3 setae on ventral surface
(Fig. 3E). Maxilla: distal segment of endopod
expanded distally, wider than long, distal
margin slightly arched, 2.5 times as wide as
at base, armed with 7 strong spines of which
outer one is longer than others; exopod ex-
tending to distal end of first endopod segment
(Fig. 3H). Labrum with frontal margin round-
ed (Fig. 3G).

First thoracic endopod robust, ischium
wider than long, with developed inner lobe,
merus relatively slender, dactylus wider
than long, with stout, straight claw (Fig.
3H). Second thoracic endopod rather slen-
der, basis with developed inner lobe, ischi-
um with about 8 long setae on inner margin,
merus as long as carpopropodus and dac-
tylus together, dactylus longer than wide,
with 1 barbed claw and 6 barbed setae (Fig.
31). Third to eighth thoracic endopods be-
coming more slender towards posterior
pairs, carpus articulated obliquely with pro-
podus but in eighth limb articulation is
nearly transverse, propodus divided into 2
subsegments, in eight limb distal subseg-
ment 1.6 times longer than proximal (Fig.
4A, B). Exopods with flagelliform part 9-
segmented in middle pairs; basal plate with
outer distal corner pointed (Fig. 31).

Abdomen with sixth somite longest, 1.3
times as long as fifth.

Male pleopods well developed, bira-
mous. First pleopod with 7-segmented ex-
opod and unsegmented endopod. Second
and third pleopods with 7-segmented exo-
pod longer than 6-segmented endopod. Ex-
opod of fourth pleopod modified, 7-seg-
mented, nearly 1.5 times longer than en-
dopod, terminal segment very small, % of
preceding segment in length, armed with 1
tiny seta at distal end, penultimate segment
with 1 strong spinose seta which is nearly
4 times longer than segment supporting it,
antepenultimate segment armed with 1
short feeble seta at distal end of outer mar-
gin; endopod 6-segmented, normal, without

360

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Morphological differences between Neodoxomysis sahulensis and N. littoralis.

N. sahulensis

Second segment of endopod

of maxilla
Exopod of 4th pleopod of male
Endopod of 4th pleopod of

male of exopod

Endopod of 5th pleopod of Extending to distal end of 5th
male segment of exopod

Telson 1.5 times as long as broad, lateral

margin with 16—17 spines; cleft 4
as long as telson, with 11—12
spines on each side

modified setae (Fig. 4C, D). Fifth pleopod
with exopod 7-segmented, 1.2 times longer
than 6-segmented endopod; endopod with
first segment with digitiform projection
tipped with 1 seta on outer margin in ad-
dition to usual side lobe (Fig. 4E).

Endopod of uropod extending beyond
distal end of telson for % of its length, inner
margin from statocyst region to near apex
with 34 spines, which are composed of lon-
ger and shorter spines arranged alternately
except for several spines on proximal and
distal portions, distal 4 spines long and
acute, other spines obtuse (Fig. 4F). Exopod
of uropod overreaching endopod for ¥; of
its length (Fig. 4P).

Telson as long as last abdominal somite,
1.5 times as long as broad at base, deeply
cleft at apex; lateral margin furnished with
16-17 rather long spines along whole
length, distal 10 spines gradually increasing
in length towards apex; each apex of distal
lobes armed with 3 spines, outer one of
which is slightly longer than others; apical
cleft %4 as long as telson, furnished with 11—
12 spinules on each side and pair of plu-
mose setae arising from anterior end (Fig.
4F). Spination on telson in female similar
to that of male.

Etymology.—The name sahulensis refers
to the locality in which the specimens were
collected.

Remarks.—Neodoxomysis sahulensis is
closely similar to N. littoralis (Tattersall,

With 7 acute, simple spines

7-jointed, with 1 seta at distal end
Extending to middle of 4th segment

N. littoralis

With 9 slender, bluntly pointed
spines

7-jointed, with 2 setae at distal end

Extending to distal end of 5th
segment of exopod

Extending to distal end of 6th
segment of exopod

1.75 times as long as broad, lateral
margin with 16 spines; cleft % as
long as telson, with 16—19 spines
on each side

1922) in many respects, but distinguished
from that species as shown in Table 1.

In the new species the outer lobe of the
maxillule is furnished with a row of spi-
nules on the middle part of the outer mar-
gin. Such a character has been observed in
Doxomysis quadrispinosa by Pillai (1973)
and D. australiensis by Tattersall (1940).

Literature Cited

Hansen, H. J. 1912. Report on the scientific results of
the expedition to the eastern tropical Pacific, in
charge of Alexander Agassiz, by the U.S. Fish
Commission Steamer “Albatross’’, from Au-
gust, 1899, to March, 1900, Commander Jeffer-
son E Moser, U. S. N., commanding. XVI. Re-
ports on the scientific results of the expedition
to the eastern tropical Pacific, in charge of Al-
exander Agassiz, by the U.S. Fish Commission
Steamer ‘‘Albatross’’, from October, 1904 to
March, 1905. Lieut.-Commander L. M. Garrett,
U. S. N., commanding. XX VII. The Schizopo-
da.—Memoirs of the Museum of Comparative
Zoology at Harvard College 35:175—296, pls.
1-12.

li, N. 1964. Fauna Japonica, Mysidae (Crustacea). Bio-
geographical Society of Japan, Tokyo, 610 pp.

Pillai, N. K. 1973. Mysidacea of the Indian Ocean.
Papers on the zooplankton collection of the
ITOE.—Handbook to the International Zooplank-
ton Collections 4:1—125.

Talbot, M. S. 1997. Doxomysis acanthina, a new lep-
tomysinid (Crustacea: Mysidacea) from the
northern Great Barrier Reef, Australia, with ex-
tensions to the known distributions of D. aus-
traliensis W. M. Tattersall, 1940 and D. spinata
Murano, 1990, and a key to the genus Doxo-

VOLUME 112, NUMBER 2

mysis.—Proceedings of the Biological Society
of Washington 110:426—438.

Tattersall, W. M. 1922. Indian Mysidacea.—Records
of the Indian Museum 24:445—504.

361

. 1940. Report on a small collection of Mysi-
dacea from the coastal waters of New South
Wales.—Records of the Australian Museum,
20:327—340.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
112(2):362—367. 1999.

First record of the family Gynodiastylidae Stebbing, 1912

(Crustacea: Malacostraca: Cumacea) from Antarctic waters with the

description of Gynodiastylis jazdzewskii, a new species

Magdalena Btlazewicz and Richard W. Heard

(MB) University of L6dz, Department of Invertebrate Zoology and Hydrobiology,

Laboratory of Polar Biology and Oceanobiology, 90-237 L6dz, S. Banacha str. 12/16, Poland;

(RWH) University of Southern Mississippi Institute of Marine Sciences,
Gulf Coast Research Laboratory, PO. Box 7000, Ocean Springs, Mississippi 39566-7000

Abstract.—An adult male of an undescribed gynodiastylid cumacean was
collected in a Menzies trawl sample taken at a depth of between 388 and 399
m in the Ross Sea (76°01.5’—76°01.0'S, 179°49.9’—179°52.3’E) during February
1972. The specimen, which had lost its third maxillipeds and the last four
articles of the first pereopods, is placed in the genus Gynodiastylis Calman,
1911. Gynodiastylis jazdzewskii, new species, the first member of its family
reported from Antarctic waters, can be distinguished from the 59 previously
described species of Gynodiastylis and members of the other five gynodiastylid
genera by a combination of characters, including the finer, more irregularly
wavy, longitudinal ridges, and the length and setation of the telson, on the
carapace, the shape and terminal setation of the telson, the length of telson in
relation to that of the last abdominal somite and uropodal peduncles, and the

setation and relative length of the uropodal rami.

An examination of the Cumacea collect-
ed by the R/V Eltanin in the Ross Sea, re-
vealed a single adult male specimen of an
undescribed gynodiastylid. The specimen
was damaged (third maxillipeds and the
four distal articles of the first pereopods
missing), but can be reliably assigned to the
genus Gynodiastylis Calman, 1911 and rep-
resents the first record of the family Gyno-
diastylidae from the Antarctic region.

Gynodiastylis jazdzewskii, new species
Figs. 1—2

Material examined.—Holotype: adult
male (USNM 243765); Ross Sea, between
76°01.5'S—179°49.9’E and 76°01.0’S—
179°52.3'E; depth 388-399 m; 08 February
1972; Menzies trawl; R/V Eltanin, Cruise
51, Sta. 5761; Coll., Smithsonian Oceano-
graphic Sorting Center.

Diagnosis —Gynodiastylidae. Adult male.

Carapace with numerous (approximately
17) fine, irregular, wavy, longitudinal ridg-
es, many coalescing (especially anteriorly).
Exopods on pereopods 1—4. Telson lingui-
form, distinctly longer than sixth abdominal
somite, extending to distal %4 of uropodal
peduncle, armed with 2 distinct, closely set,
terminal spine-setae. Uropods relatively
long, attenuated; peduncle with 4 spine-se-
tae along inner distal margin; endopod and
exopod with 2 articles, endopod (excluding
terminal setae) slightly longer than exopod,
exopod with 2 long terminal setae, inner-
most about as long as rest of exopod, outer
seta about *% length of inner one.
Description.—adult male holotype (Fig.
1), carapace length 0.9 mm, total length 3.2
mm. Carapace nearly % of total length; hav-
ing numerous (approximately 17) fine, ir-
regular, wavy, longitudinal ridges (or stria-
tions), many coalescing anteriorly; ocular
lobe well-developed, eyes obscure, unpig-

VOLUME 112, NUMBER 2

363

Fig: (1.

mented; pseudorostrum strongly developed,
acutely pointed, slightly decurved, extend-
ing well beyond ocular lobe. Antennal
notch broad, not defined ventrally by sharp
tooth. Thoracic somites 3-5 as illustrated
(Fig. 1). Abdomen (Fig. 1) subequal in
length to carapace; somites of similar
length. Telson (Fig. 2J) linguiform, distinct-
ly longer than sixth abdominal (telsonic)
somite, extending to distal %4 of uropodal
peduncle, armed with 2 distinct, closely set,
terminal spine-setae. First and second an-
tennae (not dissected), appearing similar to
those of other described members of genus.
Mandible (Fig. 2A): with well developed
incisor and molar. First maxilla (Fig. 2B):

Gynodiastylis jazdzewskii, n. sp. Lateral view of male holotype. Scale = 0.2 mm.

outer endite with 14 terminal or subtermi-
nal, stout, spine-setae. Second maxilla (fig.
2E): characterized by well-developed, stout
comb setae. Maxilliped 1 (Fig. 2D): bran-
chial lobe (not illustrated) lacking branchial
processes; siphon long, twisted distally, ter-
minating in bent acute tip; endopod with
well-developed specialized setae and spine-
setae on inner plate, coxa, and propodus
(Fig. 2D, enlargements). Second maxilliped
(Fig. 2C): carpus nearly twice as long as
propodus; propodus more than twice as
long as dactyl; dactyl terminating in 3 well
developed, simple spine-setae. Third max-
illipeds: lost. Pereopods 1—4: bearing exo-
pods, exopods decreasing in size posteriorly

364 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

WY
RQ\\hie
IY \)

WAN

Z
“4
Y

Fig. 2. Gynodiastylis jazdzewskii, n. sp. A, left mandible: incisor (on left), all spine teeth, except distal most

broken off; molar (on right); B, first maxilla; C, second maxilliped (terminal articles); D, first maxilliped (ex-
cluding damaged branchial lobe) showing enlargement of setal types; E, second maxillia EK second pereopod
(exopod not shown); G, third pereopod; H, fourth pereopod (exopod not shown); I, fifth pereopod; J, sixth

abdominal (telsonic) somite, uropods, and telson. Scale a = 0.1 mm for E G, H, I; Scale b = 0.3 mm for A,
B, C and 0.1 mm for J.

VOLUME 112, NUMBER 2

(Fig. 1). First pereopod (Fig. 1) damaged,
only basis present; basis strongly devel-
oped, extending anteriorly past midlength
of carapace, nearly as long as entire second
pereopod. Second pereopod (Fig. 2F), basis
subequal to combined length of other arti-
cles; ischium distinguishable; carpus longer
than merus; propodus subequal in length to
dactyl. Third pereopod (Fig. 2G), merus
well developed, subequal in length to basis;
carpus approximately as long as propodus,
nearly as wide as long with 3 well devel-
oped distal setae, 1 extending to tip of dac-
tyl and 2 extending well beyond dactyl;
dactyl about as long as propodus. Fourth
pereopod (Fig. 2H) similar to third, but hav-
ing more slender carpus. Fifth pereopod
(Fig. 21), basis relatively short, subequal in
length to combined lengths of ischium,
merus, and carpus; merus broad, slightly
longer than carpus; carpus longer than pro-
podus; propodus with long, stout, distal seta
extending well past dactyl; dactyl shorter
than propodus. Uropods (Fig. 2J): relatively
long, approximately equal to combined
length of abdominal somites 5 & 6, atten-
uated; peduncle with 4 spine-setae along in-
ner distal margin; inner ramus with 2 arti-
cles, longer than outer ramus, proximal ar-
ticle with inner margin bearing 3 spine-se-
tae, distal article longer than proximal,
inner margin with 3 spine-setae, 2 terminal
setae present, outer seta distinctly longer
than distal article and over 4 times longer
than inner seta; outer ramus with 2 articles,
distal article with one relatively short sub-
terminal seta on inner margin and 2 long
terminal setae, innermost approximately as
long as rest of exopod, outer seta about %
length of inner.

Etymology.—This species is named in
honor of Professor Krzysztof Jazdzewski
(University of L6dz, Poland) in recognition
of his many significant contributions to
Antarctic biological research.

Comparisons.—Gynodiastylis jazdzew-
skii can be distinguished from the previ-
ously described species of the genus by the
fine, irregularly wavy, longitudinal ridges

365

on the carapace, the shape and terminal se-
tation of the telson, the relative length of
the telson in relation to that of the last ab-
dominal somite and the uropodal peduncles;
and by the setation and relative length of
the uropodal rami.

Of the known species of Gynodiastylis,
four (G. costata Calman, 1911; G. turgida
Hale, 1928; G. lata Hale, 1946; G. lineata
Day, 1980) have carapaces with longitudi-
nal ridges or striations. Of these, G. ja-
zdzewskii appears to be most closely related
to G. costata, the only other member of the
genus having numerous, fine, irregular stri-
ations on its carapace. Gynodiastylis cos-
tata was originally described from Gulf of
Siam at depths ranging from 9 to 37 m. It
was later reported from “night surface
plankton’? in Japanese waters by Gam6é
(1968).

Gynodiastylis jazdzewskii is readily dis-
tinguishable from the comparable males of
G. costata sensu Calman (1911) and Gam6é
(1962) by having: (1) a longer and more
acutely tipped rostrum, (2) a distinctly lon-
ger telson (approximately *%4 the length of
the uropodal peduncle), and (3) two long
terminal setae on the tip of uropodal exo-
pod. Based on Calman’s (1911) and Gam6’s
(1962) descriptions and illustrations (plate
36, figs. 1-10 and fig. 40, respectively) the
telson of male G. costata is half or less the
length of the uropodal peduncle. There is
only a single long inner terminal seta on the
uropodal exopod of G. costata, the outer
terminal seta being weakly developed (less
than Y; the length of the inner).

The variously modified peduncular arti-
cles of the first antennae of Allodiastylis
Hale, 1936 and Sheardia Hale, 1946 distin-
guish G. jazdzewskii from the species of
these two genera. The presence of uropodal
endopods with two articles distinguishes G.
Jazdzewskii from the species of the genus
Dicoides Hale, 1946, which are character-
ized by uropodal endopods with three arti-
cles. The members of the remaining two
gynodiastylid genera, Halina Day, 1980,
and Zimmeriana Hale, 1946, all lack the

366

fine, longitudinal, carapace ridges that char-
acterize G. jazdzewskii.

Discussion.—In her revision and rein-
statement of the family Gynodiastylidae,
Day (1980) recognized six genera, Gyno-
diastylis, Allodiastylis, Dicoides, Haliana,
Sheardia, and Zimmeriana. Based on the
summary works by Day (1980) and Baces-
cu (1992), records for all previously de-
scribed gynodiastylids appear to be con-
fined to the Indo-West Pacific region be-
tween 40°N (Japan) and 43°S (New Zea-
land).

Because present day gynodiastylids have
predominantly tropical and temperate dis-
tributions, G. jazdzewskii might represent a
relic species whose ancestral stock was
present in cold Antarctic waters before the
break up of Pangaea over 180 million yr
ago. Based on its similarity to other warmer
water species like G. costata, G. jazdzews-
kii could have split off from the same stock
that gave rise to the rich present day gy-
nodiastylid fauna. Following this reasoning,
present day gynodiastylids may have
evolved from a small group of cold water,
Antarctic progenitors and then underwent
rapid speciation and dispersal as Australia
and South Africa slowly drifted northward
into the warmer latitudes. There, however,
are some problems with this hypothesis. For
example, if the origin of gynodiastylids was
the cold water of the Antarctic Ocean, their
apparent absence from continental waters of
South America is puzzling. Recently, how-
ever, Les Watling (pers. comm) informed us
that he has collected gynodiastylids from
waters off Chile.

There is the additional possibility that,
due to their small size and specialized hab-
itat requirements, the lack of previous rec-
ords for the family in Antarctic waters also
may be due to an artifact of sampling. How-
ever, if populations of gynodiastylids are
common in the shelf waters of Antarctica,
at least some additional specimens should
have been collected in studies conducted in
the waters adjacent to the Antarctic Penin-
sula. There has been considerable benthic

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

sampling in this region, and the cumaceans
from the resulting collections have been
identified (Lowry 1975, Richardson &
Hedgpeth 1977, Ledoyer 1993, Btazewicz
& Jazdzewski 1995, R. Heard, personal ob-
servations).

A third plausible option is that gynodias-
tylids originated in the warm temperate wa-
ters of the Indo-Pacific (Day 1980). If so G.
jJazdzewskii might simply be a pioneer spe-
cies, representing a relatively recent and
isolated dispersal event for the family in
Antarctic waters.

The occurrence of G. jazdzewskii at a
depth between 388 and 399 m represents
the second deepest record for the genus and
family. A southern African species, G. pro-
fondus Day, 1980, was reported from
depths of 68 to 680 m by Day (1980).

The description of any new species,
based on a single damaged specimen, is
tenuous. Notwithstanding, G. jazdzewskii is
readily distinguishable from all previously
described gynodiastylids and represents an
important zoogeographic record for the
family in Antarctic waters.

Acknowledgments

We are grateful to Dr. G. Hendler and
Ms. B. Landrum for making material and
station data from the R/V Eltanin available.
The senior author thanks Dr. Robert T. van
Aller (former Director of the Gulf Coast
Research Laboratory, Ocean Springs, MS)
and Prof. Krzysztof Jazdzewski (Director of
the Laboratory of Polar Biology & Ocean-
obiology, University of L6dz, Poland) for
their help in arranging financial assistance
to work in the United States. Ms. Sara
LeCroy graciously helped with the prepa-
ration and reading of the manuscript. This
research was supported in part by the U.S.
Antarctic Research Program of the National
Science Foundation, through a _ contract
with the Smithsonian Oceanographic Sort-
ing Center (Grant DPP-8214878).

VOLUME 112, NUMBER 2

Literature Cited

Bacescu, M. 1992. Cumacea II (Fam. Nannastacidae,
Diastylidae, Pseudocumatidae, Gynodiastylidae,
et Ceratocumatidae)—Crustaceorum Catalogus,
Pars 8, H.-E. Gruner and L. B. Holthuis, eds.,
SPD Academic Publishing, The Hague. pp. i1—
iv, 175—468.

Blazewicz, M., & K. Jazdzewski. 1995. Cumacea
(Crustacea, Malacostraca) of Admiralty Bay
(King George Island, South Shetlands): a pre-
liminary note.—Polish Polar Research 16(1—2):
71-85.

Calman, W. T. 1911. On new or rare Crustacea of the
order Cumacea from the collection of the Co-
penhagen Museum. II. The families Nannasta-
cidae and Diastylidae.—Transactions of the
Zoological Society of London 18:341-398.

Day, J. 1980. South African Cumacea. Part 4. Families
Gynodiastylidae and Diastylidae.—Annals of
the South African Museum 82(6):187—292.

Gam6o, S. 1962. On the Cumacean Crustacea from Tan-
abe Bay, Kii Peninsula.—Publications of the
Seto Marine biological Laboratory 10(2):10—
66.

. 1968. Studies on the Cumacea (Crustacea,

Malacostraca) of Japan, part II.—Publications

367

of the Seto Marine biological Laboratory 16(3):
147-192.

Hale, H. M. 1928. Australian Cumacea.—Transactions
of the Royal Society of South Australia 52:31-—
5

1936. Cumacea from a South Australian
reef.—Records of the South Australian Museum
5(4): 404-438.

. 1945. Australian Cumacea. No. 11. The fam-

ily Diastylidae, part 1.—Transactions of the

Royal Society of South Australia 69:173—211.

. 1946. Australian Cumacea. No. 12. The fam-
ily Diastylidae, part 2 Gynodiastylis and related
genera.—Records of the South Australian Mu-
seum 8:357—444.

Lowry, J. K. 1975. Soft bottom macrobenthic com-
munity of Arthur Harbor, Antarctica.—Antarc-
tic Research Series 23:1-19.

Richardson, M., & J. W. Hedgpeth 1977. Antarctic
soft-bottom macrobenthic community adapta-
tions to cold stable, highly productive, glacially
affected environment. pp. 181-196 In G. A.
Liano, ed., Adaptations within Antarctic eco-
system.—Proceedings SCAR Symposium on
Antarctic Biology, Smithsonian Institution,
Washington.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(2):368—380. 1999.

A new genus and species of sphaeromatid isopod (Crustacea) from
Atlantic Costa Rica

Regina Wetzer and Niel L. Bruce

(RW) Department of Biological Sciences, University of South Carolina,
Columbia, South Carolina 29208 U.S.A.; rwetzer@sc.edu;
(NLB) 138 Carmody Road, St. Lucia, Queensland 4067, Australia; nbruce@ globec.com.au

Abstract.—Beatricesphaera ruthae, a new genus and species of sphaero-
matid isopod inhabiting shallow water along the Atlantic Costa Rican coast is
described. This small species is placed into a new genus as the combination
of characters which describe it precludes inclusion into any known sphaero-
matid genus. It is best recognized by these character states: short epistome,
flattened antenna 1 peduncle articles 1 and 2, robust setose pereopods, lamellar
uropods, simple pleopods 4 and 5, short and unfused penes and an elongate
basally attached appendix masculina. It is not readily allied to any genus or
group of genera within the Sphaeromatidae.

Costa Rican shores boast every major
tropical coastal marine habitat, from rocky
shores and sandy beaches, to mangrove la-
goons and coral reefs. In a joint project
between Universidad de Costa Rica and
the Los Angeles County Natural History
Museum, the Cahuita Reef (9°18'N,
82°7'W) at Parque Nacional on the Carib-
bean coast of Costa Rica was surveyed and
monitored between April 1986 and August
1987 for habitat degradation resulting from
upland deforestation. This national park in-
cludes a small 240 hectare barrier reef
(Wellington 1974; Risk et al. 1980; Cortés
et al. 1984; Guzman & Cortés 1984a,
1984b; Murillo & Cortés 1984; Cortés &
Risk 1985).

Knowledge of the Pacific Costa Rican
isopod fauna rests with Brusca & Iverson’s
1985 summary. The greater Caribbean re-
gion is covered by the publications of Kens-
ley (1984), the field guide of Kensley &
Schotte (1989) and miscellaneous other
contributions, notably by Miller (e.g.,
1993a, 1993b). This new genus and species
of marine sphaeromatid isopod brings the
total number of sphaeromatid genera known
from the Caribbean to 17.

Bruce (1993, 1995, 1997) discusses some
of the taxonomic difficulties of the Sphae-
romatidae (e.g., high levels of homoplasy
and the certain lack of monophyly of many
larger genera) when compared to other fla-
belliferan families. The Sphaeromatidae are
remarkable for their morphological diver-
sity and are highly speciose. To date almost
100 genera have been described (Bruce
1995), and the latest count reports over 600
described species (see Kensley & Schotte,
USNM Isopod List on the WWW at: (URL:
http://nmnhwww-.si.edu/gopher—menus/Iso-
pods.html)). For the Caribbean, Kensley &
Schotte (1989) reported 32 species in 13
genera, and White (1996 unpublished) re-
ports 7 described sphaeromatids from the
Cahuita reef and two undescribed species.

Methods and terminology.—Coral rubble
and algal samples from the intertidal to 12 m
were collected and rinsed in fresh water and
sieved through a 240 pm mesh screen. All
material was fixed and preserved in 95%
ethyl alcohol. Material was examined with
dissecting, compound, and scanning elec-
tron microscopes. Appendages were drawn
with the aid of a camera lucida. Setal and
spine nomenclature as well as the nomen-

VOLUME 112, NUMBER 2

clature of appendage orientation follows
Brusca et al. 1995. Abbreviations used are:
LACM—Los Angeles County Museum of
Natural History, Los Angeles, California;
MOV—Museum of Victoria, Melbourne,
Australia; SDNHM-—San Diego Natural
History Museum, San Diego, California;
UCR—Universidad de Costa Rica, San
Jose, Costa Rica; USNM—National Muse-
um of Natural History, Smithsonian Insti-
tution, Washington, D.C.; PMS, plumose
marginal setae; SEM, scanning electron mi-
croscope.

Systematics

Order Isopoda Latreille, 1817
Suborder Flabellifera Sars, 1882
Family Sphaeromatidae
Milne Edwards, 1840
Beatricesphaera, new genus

Type species.—Beatricesphaera ruthae,
new species, here designated.

Diagnosis of male.—Body vaulted, about
twice as long as wide, widest at pereonites
5 and 6; body with translucent, flexible
membrane-like covering; body margins
with setose fringe, membrana cingula (see
Buss & Iverson 1981). Cephalon lacking
rostral point, lateral margins not expanded;
eyes lateral, distinctly faceted. Pereonites
2—6 of about equal length, pereonites 1 and
7 longest; pereonite 7 not extending to lat-
eral margins, laterally overlapped by per-
eonite 6; coxae without coxal keys (see
Bruce 1994a, Figs. 2G, 22D), sutures ab-
sent. Pleon consisting of 1 segment. Pleo-
telson anterolateral margins extending to
full body width, posterior margin narrowed,
with distinct dorsal foramen, posterior mar-
gins of which contact posteriorly.

Antenna 1 peduncle articles 1 and 2
strongly flattened, anteriorly expanded; ar-
ticle 2 with anterodistal lobe; anterior mar-
gins of article 1 and 2 contiguous; article 3
slender, not expanded; flagellum short,
slightly longer than article 3. Antenna 2 pe-
duncle article 1 shortest, article 5 longest,

369

articles 2—4 progressively increasing in
length.

Epistome short, not laterally narrowed or
constricted, with blunt median point; not
separating antennule bases; lateral margins
narrow, encompassing labrum. Right man-
dible incisor blade-like, narrow, unicuspid;
without spine row and lacinia mobilis; left
mandible incisor similar to right, but with
lacinia mobilis and spine row of 2 spines;
both mandibles with robust molar provided
with proximal marginal teeth, gnathal sur-
face smooth. Maxilla 1 outer lobe with 10
spines, outer 5 smooth, inner 5 weakly ser-
rate, inner lobe with 4 feathered spines and
2 short simple spines. Maxilla 2 with 3 sub-
equal lobes, provided with setae. Maxilli-
ped palp articles 2 and 3 with short distinct
distomedial lobes, article 4 and 5 quadrate;
endite flat, distal margin subtruncate, with
acute and blunt spines.

Pereopods robust, ambulatory, accessory
unguis of dactylus simple. Pereopod 2
markedly slender in comparison to pereo-
pod 1, pereopods 3—7 more robust than 1
and 2, posterior margins of pereopods with
fringe of simple and plumose setae on ven-
tral margins of merus, ischium, and pro-
podus.

Paired short flattened penes present on
posterior of sternite 7, not extending to ple-
opod rami.

Pleopods 1—3 rami with PMS, 4—5 with-
out PMS. Pleopod 1 rami almost subequal
in length, neither indurate nor operculate.
Pleopod 2 endopod about 1.5 times as long
as exopod; appendix masculina on basal
lobe, nearly twice (1.8) as long as endopod,
apically acute. Pleopod 3 endopod without
suture; pleopods 4 and 5 without ridges or
folds; pleopod 5 exopod with 3 scale patch-
es. Uropods lamellar, extending just beyond
posterior of pleotelson, not meeting at mid-
line; exopod prominent, about half as long
as endopod.

Female.—Mouthparts not metamor-
phosed; pleotelson posterior margin entire,
without exit channel; oostegites absent;

370

eggs apparently held within pereon; other-
wise similar to male.

Remarks.—This small species of sphae-
romatid presents a combination of charac-
ters that prevents its inclusion in any known
sphaeromatid genus, and furthermore, can-
not readily be allied to any group of genera
within the Sphaeromatidae. We therefore
regarded it as incertae sedis. Superficially it
is perhaps most similar to the “‘Cassidini-
dea group” discussed by Bruce (1994b),
but it lacks the large dorsally visible epi-
stome, fused penes, laterally extended ceph-
alon, and also has short rather than elongate
pleopod 1 rami. In common with that
group, it has an elongate appendix mascu-
lina which is set on a basal lobe, the mar-
supium is composed of anterior and poste-
rior pockets (see Harrison 1984, Kensley &
Schotte 1989: 205 for terminology), the
pleon consists of a single segment entirely
without sutures, and pleopods 4 and 5 are
similar to those of Cassidinidea (all of these
characters mentioned might be considered
possible apomorphies). The significance of
the shared characters is uncertain as all of
these states are known to occur in other
sphaeromatid genera. In common with the
‘‘Leptosphaeroma group’”’ (sensu Bruce
1994b) are the short epistome and short pe-
nes, but the genera of this group are oth-
erwise distinct. What is remarkable, if the
affinities of this new genus do lie with Cas-
sidinidea and related genera, is that this is
the first instance of a perforate pleotelson
in that group of genera.

There is, as mentioned earlier, a superfi-
cial resemblance to the genus Cassidinidea
Hansen, 1905 (reviewed in Bruce 1994b).
That genus is immediately distinguished by
the dorsally visible epistome, narrow anten-
nule peduncle articles, expanded lateral
margins of the head, and fusion of the pe-
nial processes (Kensley & Schotte’s 1989
statement that the penes are “elongate, sep-
arate’’ is in error, applying only to “‘Cas-
sidinidea’ mosaica Kensley & Schotte,
1987, a species incorrectly placed in the ge-
nus). In addition both male and female Cas-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

sidinidea have the pleotelson posterior mar-
gin entire, whereas it is perforate in the
male of Beatricesphaera new genus. A
*“‘gel-layer’’ occurs in Cassidinella (Bruce
1994a) as well as other genera, but seems
easily lost, and is usually not recorded, so
its significance is entirely uncertain. It was
not observed in Beatricesphaera. :

Lack of thickened ridges or folds on ple-
opods 4 and 5 has repeatedly been shown
to be a character of dubious phylogenetic
significance (most recently Bruce 1995,
1997) that can vary within genera, and the
flattened antennule peduncle article(s) can
also vary within genera (e.g., Cassidinella,
Bruce 1994a). Another character state that
is not widely distributed in the family is the
marked slenderness and elongation of the
second leg in comparison to the first. This
is also known to occur in some genera and
species of the Jschyromene-group (Bruce
1995) as well as some species of Dyname-
nella (e.g., D. nuevitas Kensley et al. 1997).
Lamellar uropods, which occur in most
genera formerly placed in the subfamily
Cassidininae, are also to be found in other
genera such as Exosphaeroma for example.
Without a full phylogenetic analysis of the
genera with the associated identification of
informative characters and resolution of the
evidently high level of homoplasy in the
family (Bruce 1993, 1995, 1997), problems
such as the one that this genus presents will
continue to plague taxonomists working on
the Sphaeromatidae.

Of the other sphaeromatid genera known
from the Caribbean region, none is partic-
ularly similar, and the genus can best be
identified by the characters of short epi-
stome, flattened antenna 1 peduncle articles
1 and 2, robust setose pereopods, lamellar
uropods, simple pleopods 4 and 5, short and
unfused penes and an elongate basally at-
tached appendix masculina. Females can be
identified by the antenna 1, epistome and
pereopod characters. The prominent bosses
on pereonite 7, while not included as a ge-
neric character, further serve to identify the
genus and species.

VOLUME 112, NUMBER 2

Etymology.—During her tenure at the
SDNHM, RW had the great pleasure of
working with and benefiting from two tre-
mendously wise women (Beatrice Koep-
nick and Ruth Kantor), whose efficacy, in-
sight, and hard-earned wisdom left an in-
delible impression on all they met. It is the
energy that these two women gave to so
many scientific projects that this new genus
and species honors. The new genus honors
Bea who passed away April 1995 and un-
fortunately before this manuscript was com-
pleted. The generic name is feminine, as the
second element, sphaera, is a feminine Lat-
in noun.

Beatricesphaera ruthae, new species
Figs. 1-6

Material examined.—Male holotype
(LACM 86-196.4), Atlantic, Costa Rica, Li-
mon Province, Parque Nacional Cahuita,
Punta Cahuita Reef, 9°44.18’N, 82°48.7'W,
inner reef flat, brown algae wash, 1-3 m
depth. 29 Oct 1986. Coll. R. C. Brusca and
P. M. Delaney, LACM 86-196. Ovigerous
female allotype (LACM 86-202.3) Parque
Nacional Cahuita, Punta Cahuita Reef,
9°40.3'N, 82°45’'W, outer reef crest, algae
and algae holdfast washes, 1—8 m depth. 30
Oct 1986. Coll. R. C. Brusca and P. M. De-
laney, LACM 86-202. Paratypes (2 mature
males, 2 juvenile males) (LACM 86-196.7),
Parque Nacional Cahuita, Punta Cahuita
Reef, 9°44.18’N, 82°48.7'W, inner reef flat,
brown algae wash, 1-3 m depth. 29 Oct
1986. Coll. R. C. Brusca and P. M. Delaney,
LACM 86-196. Paratypes (1 male, 1 fe-
male, 1 juvenile) (LACM 86-202.4), Parque
Nacional Cahuita, Punta Cahuita Reef,
9°40.3'N, 82°45'W, outer reef crest, algae
and algae holdfast washes, 1-8 m depth. 30
Oct 1986. Coll. R. C. Brusca and P. M. De-
laney, LACM 86-202. Paratypes (2 males,
3 females) (LACM 87-3.2), Parque Nacion-
al Cahuita, Punta Vargas, 9°44.18'N,
82°48.7'W, inner reef lagoon, algae washes,
1—3 m depth. 8 Jan 1987. Coll. R. C. Brusca
and R. Wetzer, LACM 87-3. Paratype (1

STi

male, dissected and figured appendages)
(LACM 86-122.4), Limon Province, north
of Puerto Viejo, 9°40.8'N, 82°45.53’W, cor-
alline algae rock wash, offshore algae ridge,
1-3 m depth, CRA-86-13. 27 Jul 1986.
Coll. G. L. Hendler, LACM 86-122. Para-
types) (Seanales; 5 Ofemales) (USNM
286886), Parque Nacional Cahuita, Punta
Vargas, 9°44.18'N, 82°48.7'W, inner reef la-
goon, algae washes, 1-3 m depth. 8 Jan
1987. Coll. R. C. Brusca and R. Wetzer,
LACM 87-3. Paratypes (4 males, 4 fe-
males) (MOV J45491 through J45498),
Parque Nacional Cahuita, Punta Cahuita
Reef, 9°40.3’N, 82°45’W, outer reef crest,
algae and algae holdfast washes, 1-8 m
depth. 30 Oct 1986. Coll. R. C. Brusca and
P. M. Delaney, LACM 86-202. Paratypes (2
males, 3 females) (UCR-2248), Parque Na-
cional Cahuita, Punta Cahuita Reef,
9°40.3'N, 82°45’W, outer reef crest, algae
and algae holdfast washes, 1—8 m depth. 30
Oct 1986. Coll. R. C. Brusca and P. M. De-
laney, LACM 86-202.

Other material examined: Atlantic, Costa
Rica, specimens: Limon Province, Parque
Nacional Cahuita, Punta Cahuita Reef,
9°44.35’N, 82°48.7'W, approximately 500 m
from shore, algae and coral rubble washes,
3—4 m depth. 3 Apr 1986. Coll. R. C. Brus-
ca and P M. Delaney, LACM 86-98, 2
mancas. Parque Nacional Cahuita, Punta
Cahuita Reef, 9°44.35’N, 82°48.7'W, ap-
proximately 500 m from shore, brown algae
washes, 3—4 m depth. 4 Apr 1986. Coll. R.
C. Brusca, P. M. Delaney, and R. Wetzer,
LACM 86-100, 9 specimens. Limon Prov-
ince, north of Puerto Viejo, 9°40.8'N,
82°45.53'W, coralline algae rock wash, off-
shore algae ridge, 1-3 m depth, CRA-86-
Say 27) | SulesbOSGn Colke Goal sw Hendler
LACM 86-122, 24 specimens. Limon Prov-
ince, north of Puerto Viejo, 9°40.8'N,
82°45.53'W, outer reef crest, coralline algae
on Sargassum reef, 6-11 m depth. 27 Jul
1986. Coll. R. Wetzer, LACM 86-128, 2
specimens each attached to a walking leg
of brachyuran crab: Acanthonyx petiverii.
One specimen attached to left 3rd walking

372

leg; second specimen attached to left 2nd
walking leg. Limon Province, Parque Na-
cional Cahuita, Punta Cahuita Reef,
9°44.18'N, 82°48.7’W, inner reef flat,
brown algae wash, 1—3 m depth. 29 Oct
1986. Coll. R. C. Brusca and P. M. Delaney,
LACM 86-196, 40+ specimens. Limon
Province, Parque Nacional Cahuita, Punta
Cahuita Reef, 9°40.3’N, 82°45'W, outer reef
crest, algae and algae holdfast washes, 1-8 m
depth. 30 Oct 1986. Coll. R. C. Brusca and
P. M. Delaney, LACM 86-202, 64+ speci-
mens. Limon Province, Parque Nacional
Cahuita, Punta Vargas, 9°44.18'N,
82°48.7'W, inner reef lagoon, algae washes,
1—3 m depth. 8 Jan 1987. Coll. R. C. Brusca
and R. Wetzer, LACM 87-3, 70+ speci-
mens. Limon Province, Parque Nacional
Cahuita, Punta Vargas, 9°44.18'N, 82°48.7'W,
inner reef lagoon, coral rubble washes, 1—
12 m depth. 8 Jan 1987. Coll. R. C. Brusca
and R. Wetzer, LACM 87-4, 2 females. Li-
mon Province, Parque Nacional Cahuita,
Punta Vargas, 9°44.18'’N, 82°48.7'W, inner
reef lagoon, rock, coral rubble, and algae
washes, 1—13 m depth. 8 Jan 1987. Coll. R.
C. Brusca and R. Wetzer, LACM 87-5, 2
mancas. Limon Province, Parque Nacional
Cahuita, Punta Vargas, 9°44.5'N, 82°48.37'W,
inner reef lagoon, skimming Thallassia bed
off point, intertidal to 1 m depth. 8 Jan
1987. Coll. R. C. Brusca and R. Wetzer,
LACM 87-7, 1 male. Limon Province,
Parque Nacional Cahuita, Punta Vargas,
9°44.18'N, 82°48.7'W, inner reef lagoon,
broken coral and rubble washes, 1—3 m
depth. 8 Jan 1987. Coll. R. C. Brusca and
R. Wetzer. LACM 87-8, 1 molt. Limon
Province, Puerto Viejo, 9°40.3'N, 82°45’W,
forereef crest and along sand bottom junc-
tion, algae and gorgonian washes, 1—6 m
depth. 9 Jan 1987. Coll. R. C. Brusca and
R. Wetzer, LACM 87-9, 34 specimens. L1i-
mon Province, Parque Nacional Cahuita,
Punta Cahuita Reef, 9°44.37'N, 82°48.7'W,
rock, coral, and algae holdfast washes, 0.9—
3 m depth. 19 Aug 1987. Coll. R. C. Brusca
and R. Wetzer, LACM 87-158, 21 speci-
mens. Limon Province, Parque Nacional

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Cahuita, Punta Cahuita Reef, 9°44.52'N,
82°48.37'W, rock, coral, and algae holdfast
washes, 0.5-10 m depth. 20 Aug 1987.
Coll. R. C. Brusca and R. Wetzer, LACM
87-165, 64 specimens.
Description.—Adult male: Body strongly
vaulted, limpet-like, about 2.3 times as long
as wide (Figs. 1A, B, 2A). Antennular pe-
duncle articles, lateral body margins, uro-
pods, and pleotelson with conspicuous setal
fringe, membrana cingula (Figs. 1A, B, C,
3A). Frontal margin of cephalon without
rostral process (Fig. 1A). Dorsum of per-
eonite 7 elevated into large dome-like con-
ical tubercle, with lateral notches on pos-
terior margin. Single free pleonite also with
tubercle, somewhat smaller than tubercle on
pereonite 7; with lateral projections on pos-
terior margin. Pleotelson acute; pleotelsonic
foramen dorsally directed, triangular, entire-
ly enclosed (Figs. 1A, B, 2B). A thin mem-
brane visible with SEM overlie the dorsal
cuticle of entire body including antennular
peduncle articles and appears finely granu-
lar with small, numerous, evenly spaced
pits; each pit with 1 to 3 simple or palmate
setae (Fig. 2A, B, C). Coxal sutures indis-
tinct dorsally and ventrally. Eyes round,
well developed, slightly elevated, with fac-
ets and pigmentation, deeply immersed in
pereonite 1. Pereonite 1 longest, pereonites
2—6 subequal, pereonite 7 longer than pre-
ceding pereonites. First pleonite subequal in
length to pereonites 2—6 (Figs. 1A, B, 2A).
Antennae 1 short, extending to posterior
margin of pereonite 2 (Fig. 3A). Peduncular
articles 1—2 shovel-shaped, dorsoventrally
compressed and greatly expanded. Article 1
rectangular, almost twice as long as wide.
Last (third) peduncular article narrow, not
expanded; about twice as long as wide. Fla-
gellum 6-articulate, article 1 short, article 2
longer, article 3 longest, terminal articles
gradually tapering distally. One long, “ar-
ticulated’’ aesthetasc on lateral, distal fla-
gellar articles 2, 3, and 4 (Fig. 3A, B).
Antennae 2 extending to anterior margin
of pereonite 3; usually not visible in dorsal
aspect. Peduncle composed of 4 articles, 1

VOLUME 112, NUMBER 2

( ATT mm / ¢

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—
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S

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ANTS

Beatricesphaera ruthae, new species. A, dorsal view, adult male, holotype LACM 86-196; B, lateral

view, adult male, holotype LACM 86-196; C, uropod (right), adult male, paratype LACM 86-122.

and 2 subequal in length, article 3 1.6 times

as long as 2 and somewhat expanded, arti-
cle 4 slender and about twice as long as
article 1. Simple and palmate setae present
on peduncular articles as figured. Flagellum
8-articulate, each article with 2-6 simple
setae on margin (Fig. 3C).

Frontal lamina wide, crescent shaped, en-
compassing clypeus and part of labrum. La-
brum with dense short simple setae on me-
dial margin, setae become less dense and
longer on lateral margins (Figs. 3D, 4E, F).
Clypeus and labrum fused, together form-
ing a subovate structure. Mandibular inci-

sors distally narrow, flattened, unicuspid;
lacinia mobilis composed of 2 large bifid

spines; spine row composed of 2 comb
spines; molar process with proximal mar-
ginal teeth; gnathal surface almost smooth;
palp 3-articulate, article 2 with 3 stout bi-
serrate setae, article 3 with 6 biserrate ro-
bust setae (Figs. 3E, 4A, B, C). Maxilla 1
outer lobe nearly twice as wide as inner
lobe, with 5 flattened, smooth robust spines
laterally, 4—5 large, serrate spines medially;
inner lobe with 4 slender, plumose robust
setae (Fig. 3F). Maxilla 2 with 3 subequal
lobes; outer lobe with 4 long robust setae;
middle lobe with 4 long robust setae; inner
lobe with simple, plumose and comb setae,
as figured; numerous short, simple setae
dispersed along medial margin; endite basis

373

374 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Beatricesphaera ruthae, new species. Scanning electron micrographs. A, adult male LACM 86-122,
lateral view; B, adult male LACM 86-122, pleotelson; C, male LACM 87-7 palmate setae on pleotelson, note
torn “‘cuticle;” i.e., membrana cingula, scale bar = 20 wm; D, gravid female LACM 86-122 ventral brood pouch;
E, adult male LACM 86-7 penes, scale bar = 200 pm.

VOLUME 112, NUMBER 2 375

Fig. 3. Beatricesphaera ruthae, new species. Adult male paratype LACM 86-122. A, antenna 1 (right); B,
flagellular articles antenna 1 (right); C, antenna 2 (right); D, frontal lamina, clypeus, labrum; E, mandible (left);
FE maxilla | (right); G, maxilla 2 (right); H, maxilliped (right).

376 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Beatricesphaera ruthae, new species. Scanning electron micrographs. A, male LACM 87-7, mouth-
field with right and left molar processes, left incisor, lacinia mobilis, and spine row, scale bar = 100 wm; B,
same as A, scale bar = 50 wm; C, male LACM 86-122, mouthfield, frontal view, scale bar = 100 wm; D, male
LACM 87-7, palmate setae on pleotelson, membrana cingula torn, discs are diatom symbionts, scale bar = 20
wm; E, male LACM 87-7, peduncular articles of antennules, frontal lamina, scale bar = 200 wm; K male LACM
87-3, mouthfield with clypeus, labrum, left and right maxillipeds, scale bar = 200 wm.

VOLUME 112, NUMBER 2

377

Fig. 5. Beatricesphaera ruthae, new species. Adult male paratype LACM 86-122. All appendages from left
side of body. A, pereopod 1 posterior surface; B, pereopod 2 anterior surface; C, pereopod 3 posterior surface;
D, pereopod 4 anterior surface; E, pereopod 5 posterior surface; EK pereopod 6 posterior surface; G, pereopod 7

posterior surface.

with long setae on dorsal surface, excite ba-
Sis with very short simple setae on lateral
margin (Fig. 3G). Maxillipedal endite ex-
tended to middle of palp article 2 (carpus);
endite with 4 short, stout, setose robust se-
tae, and 4 acute plumose setae on distal

margin; 1 coupling hook present on each
maxilliped (Figs. 3H, 4F).

All pereopods with secondary unguis.
Ratios of lengths of pereopods 1—7 respec-
tively (relative to pereopod 1): 1.0:1.6:1.3:
1.3:1.3:1.4:1.5 (Fig. SA—G). Pereopod 1

378

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LS
SSS

a=
a

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AYN
\
‘

i
AK \
CVV A
AAG
Mya

‘

\

5

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Beatricesphaera ruthae, new species. Adult male paratype LACM 86-122. All appendages from left
side of body. A, pleopod 1; B, pleopod 2; C, pleopod 3; D, pleopod 4; E, pleopod 5.

propodus inferior margin with 2 stout ser-
rate setae and one simple distal robust seta;
merus and carpus inferior margin each with
single simple stout seta; merus anterior su-
perior margin with stout serrate spine (Fig.
5A). Pereopod 2 propodus 2.5 times as long
as carpus (Fig. 5B). Pereopods 4—6 with
small tubercles on propodal inferior margin;
tubercles more prominent on pereopod 5
compared to 4, largest on pereopod 6. Pe-
reopods 3-7 with cuticular scale spines on
inferior dactyl surface; with setose fringe
on inferior margin of merus, carpus, and
propodus; fringe becomes longer, more
complex, and denser on posterior pereopods
(i.e., fringe composed of variable length
simple and plumose setae); simple and plu-

mose setae on superior margins of merus,
carpus, and propodus as figured, becoming
more abundant on posterior pereopods (Fig.
5C-G).

Penes 2.5 times as long as basal width,
conical, separate at base (Fig. 2E).

Pleopods 1—3 endopods and exopods
with long plumose marginal setae. Pleopod
1 endopod weakly subtriangular, exopod
ovoid, 0.8 times length of exopod; peduncle
with 4 coupling spines (Fig. 6A). Pleopod
2 endopod long, 2.4 times as long as wide,
1.5 times as long as exopod; appendix mas-
culina arising basally from endopod and ta-
pering to acute tip, distal % separated from
basal % by distinct cuticular “‘interruption,”’
1.8 times as long as endopod, ornamented

VOLUME 112, NUMBER 2

with serrations and minute spines; peduncle
with 2 coupling spines (Fig. 6B). Pleopod
3 endopod 1.5 times as long as exopod, ex-
opod weakly indurate, peduncle with 2 cou-
pling spines (Fig. 6C). Pleopods 4 and 5
rami without plumose marginal setae (Fig.
6D, E). Uropodal endopod 1.3 times as long
as exopod, endopod reaching posterior mar-
gin of pleotelson, both rami with setal
fringe composed of short and long setae,
the latter extending beyond membrana cin-
gula and feathered distally (Fig. 1C).

Adult female: Similar to male, except
dorsal tubercles on pereonite 7 and first
pleonite less pronounced than in adult
males; pleotelson entire, not perforated
(pleotelson also entire in juveniles and sub-
adult males). Without oostegites, internal
morphology of brood pouches and brood
pouch openings not discernible, ventrum of
ovigerous females appears entire (see Har-
rison 1984, fig. 2D).

Remarks.—The thin membrane overlying
the cuticle is easily damaged during pres-
ervation and subsequent handling and with
light microscopy has the appearance of
shattered glass, which SEM reveals is torn
membrane (Figs. 2C, 4D). In preparing
specimens for SEM this membrane cover-
ing has a tendency to distort and curl es-
pecially on appendage margins (Fig. 2D,
evident on distal coxal margins).

Color.—Live color a uniform pink, lost
Shortly after preservation in ethanol.

Size.—Maximum length to 3.2 mm.

Variation.—Cephalon of some speci-
mens with anterior row of 5 long simple
setae and one pair of medial setae on per-
eonites 2—7. Absent in holotype.

Distribution and ecology.—Habitats
from which specimens were collected in-
clude seagrass beds, coral rubble, and algal
holdfasts in Limon Province, Parque Na-
cional Cahuita and south to Puerto Viejo.
Two specimens were each attached to a
walking leg of brachyuran crab Acanthonyx
petiverii.

Etymology.—tThe specific epithet honors
Ruth Kantor. The name of the new genus is

B79

feminine and the specific name of the type
species also is feminine because of its der-
ivation; ruthae is the Latin feminine geni-
tive singular of a modern feminine noun.

Acknowledgments

We thank Bea Koepnick and Ruth Kantor
for their help with processing and sorting
samples and Manuel Murillo for assistance
with field logistics. Thanks go to artist Chip
Griffin for his illustrations, Dave White for
help with dissections, Todd Haney and
Shawn Vose for assistance with SEM, and
Clint Cook for his assistance with Adobe
Photoshop. The field work was in part sup-
ported by the Weiler Foundation, and this
project benefited from National Science
Foundation grant (DEB-9701524) to RW.

_Marilyn Schotte, Angelika Brandt, N. Dean

Pentcheff, Rick Brusca and two anonymous
reviewers are thanked for their helpful com-
ments on the manuscript.

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(2):381-—395. 1999.

Icelopagurus tuberculosus, a distinct new hermit crab species
(Crustacea: Decapoda: Paguridae) from Japan

Akira Asakura

Natural History Museum and Institute, Chiba, 955-2, Aoba-cho, Chuo-ku, Chiba 260-8682, Japan

Abstract.—A new species of pagurid hermit crab of the genus Jcelopagurus
McLaughlin, J. tuberculosus, is described and illustrated. It is the second spe-
cies in the genus and is distinct in having a large well-calcified, immovable
plate that is partially fused with the shield’s posterior and posterolateral mar-
gins, a transverse row of four prominent protuberances on the dorsal surface
of the shield, a pair of similar protuberances on each anterolateral margin of
the shield, and a somewhat reduced abdomen. The new species is compared
with the only other congeneric species, /. crosnieri McLaughlin. Morphological
similarities and differences of [celopagurus with selected, specialized pagurid

genera are discussed.

In the course of studies of the decapod
crustacean fauna from the Ryukyu Archi-
pelago (Okinawa) by Keiichi Nomura of
the Kushimoto Marine Park and his co-
workers, numerous specimens of hermit
crabs were collected and given to the author
for identification. A partial account of these
hermit crabs was published together with
data on other decapod crustaceans (Nomura
et al. 1996). However, a number of speci-
mens still remained unidentified, some of
which resemble species of Catapagurus A.
Milne Edwards, 1880 with sabre blade-like
dactyls on the ambulatory legs, although
the males differ in having a short sexual
tube.

McLaughlin (1997) recently established
the monotypic genus Jcelopagurus for I.
crosnieri McLaughlin, 1994, based on spec-
imens collected from Indonesian waters.
Icelopagurus is superficially similar to Ca-
tapagurus but differs in the shortness of the
sexual tube in males, which does not curve
up over the dorsal surface of the body, and
by the distinct development of the telson.
The specimens from the Ryukyu Archipel-
ago belong to Jcelopagurus and represent a
new species described herein.

Material and Methods

Collections were made by SCUBA div-
ing on coral reefs of the Kumejima Island,
northern part of the Ryukyu Archipelago
(Okinawa), Japan. Specimens were fixed in
10% formalin solution and then transferred
to 75% alcohol solution for preservation.
Specimens were stained with a 5% Meth-
ylene Blue solution and examined under a
stereoscopic microscope OLYMPUS SZH
and using a NIKON UW for high magni-
fication. Drawings were made using a cam-
era lucida attachment. All specimens are
deposited in the Natural History Museum
and Institute, Chiba, with the registration
code CBM-ZC.

Terms used in this study follow Mc-
Laughlin (1974) in general, and Lemaitre
(1995) for the cephalothorax.

Shield length (SL), measured from the tip
of the rostrum to posterior end, is used as
indicator of size. The length of segments in
chelipeds and cephalic appendages is mea-
sured along the dorsomedian line. The
length of segments in the second and third
pereopods is measured along the laterome-
dian line.

To evaluate the affinities of the new spe-

382

cies, material of the following species was
examined: Jcelopagurus crosnieri, paraty-
pes, 1,9. Sk = Ac ng one Se,
3.90 mm, 884-891 m, Station CP91l,
8°44’S—131°05’E, Tanimbar Islands, Indo-
nesia, 5 Nov 1991, Karubar Cruise, Musé-
um national d’Histoire naturelle, Paris,
MNHN-Pg 5279; Catapagurus ensifer Hen-
derson, 1893, syntypes, 2 6, SL = 2.40,
2.35 mm.-? SL 1240 may 2-ov 2 SE
= 2.15, 2.45 mm, Gulf of Martabian, Bur-
ma, The Natural History Museum, London,
1888.34; Catapagurus granulatus Edmond-
son, 1951, holotype, ¢, SL = 2.58 mm, 58—
79 m, off Bird Island, Hawaii, Bernice P.
Bishop Museum, Hawaii, BPBM 5446, par-
atypes, 2 6, SL = 2.05, 2.00 mm, 36—49
m, off the south coast of Oahu, Hawaii,
BPBM No. 5514; Catapagurus japonicus
Yokoya, 1933, syntypes, 3 6, SL = 2.10,
2.60, 2.95 mm, st. 324, Bungo Strait, 106
m, 21 Jul 1928, Kitakyushu Museum of
Natural History; 1 ¢, SL = 2.20 mm, 65
m, Maruyamadashi, Amadaiba, Sagami
Bay, 27 Jul 1958, Biological Laboratory,
Imperial Household, Showa Memorial In-
stitute, Tsukuba, BLIH 1474: 1 6, SL =
2.65 mm, 84-85 m, Aoyamadashi-Maruy-
amadashi, Amadaiba, Sagami Bay, 24 Jan
1968, BLIH 3551; 1 ovi. 2, SL = 3.10 mm,
80—85 m, 2.41 km western southwest off
Jyougashima Island, Sagami Bay, 25 Jul
1959, BLIH 1615.

Icelopagurus tuberculosus, new species
Figs. 1A—H, K—M, 2-4

Holotype.—sé, SL = 5.70 mm, CBM ZC
4700, outer side of coral reefs, northern
coast of Kumejima (26°20’N, 126°45’E),
Shimajiri-gun, Okinawa-ken, Ryukyu Ar-
chipelago, SCUBA diving, coll. Yuisho
Sakamoto, 16 Aug 1993.

Paratypes.—1 ¢, SL = 4.90 mm, CBM
ZC 4701; 1 2, SL = 5.10 mm, CBM ZC
4702; same data as holotype.

Type locality—Kumejima, Ryukyu Ar-
chipelago.

Description.—Eleven pairs of phyllo-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

branchiate gills: 2 pairs of arthrobranchiae
on either side of coxo-thoracic articulations
of third maxilliped and first to fourth pe-
reopods, plus 1 pair of pleurobranchiae on
either side of pleural plate of seventh tho-
racic somite (above fourth pereopod).

Shield (Fig. 1A, B, D) slightly broader
than long, 1.12:1.00; anterior margin be-
tween rostrum and lateral projections dis-
tinctly concave and terraced; anterolateral
margins each bearing prominent protuber-
ance armed with numerous often spiniform
tubercles on middle portion; lateral margins
Straight and armed with numerous spini-
form tubercles; posterolateral margins ir-
regular; posterior margin truncate. Dorsal
surface of shield vaulted and rugose, cov-
ered with numerous often spiniform tuber-
cles and scattered with short setae: anterior
region with a transverse row of 4 prominent
protuberances armed with numerous often
spiniform tubercles; posterolateral regions
each with 3 distinct incisions. Rostrum
broadly rounded, overreached by lateral
projections. Lateral projections distinctly
produced, triangular, upturned. Anterior
faces of shield between rostrum and lateral
projections each with 3—5 distinct blunt-
tipped spines on ventral margin laterally.
Very broad, flat, well-calcified plate sur-
rounding margin of posterior half of shield
(Fig. 1A), unarmed, with scattered long se-
tae; partially fused with shield, immovable;
lateral and posterior margins straight and
edged.

Posterior carapace (Fig. 1A, B) with pair
of partially calcified broad bands in areas
between cardiac sulci and sulci cardiobran-
chialis, broader anteriorly; partially calci-
fied, small plate present between cardiac
sulci, abutting to medial portion of posterior
margin of accessory portion of shield; linea
anomurica extending from anterior extrem-
ity of accessory portion of shield to sub-
proximal portion of ventrolateral margin of
posterior carapace. Branchiostegites (Fig.
1B) not calcified except for somewhat cal-
cified anterodorsal margin; anterodorsal and
anterior margins unarmed.

VOLUME 112, NUMBER 2

Ocular peduncles (Fig. 1A, D) short, dis-
tinctly shorter than shield, 0.30:1.00; each
with constriction near base of cornea;
strongly tuberculate dorsally and dorsome-
sially; corneae dilated; basal margins of
corneae each fringed with thick, long setae
dorsally and mesially. Ocular acicles dis-
tinctly shorter than ocular peduncles, 0.40:
1.00; elongated triangular, subacute; sepa-
rated basally by width of rostrum; mesial
margins fringed with few, thick setae; lat-
eral margins with few short, fine setae. In-
terocular plate rectangular, calcified; sepa-
rated from surrounding region by soft mem-
brane.

Antennular peduncles (Fig. 1A, E, F)
very long, with penultimate segment ex-
ceeding ocular peduncle (excluding cornea)
by approximately 0.80 own length, when
fully extended. Ultimate segment unarmed,
with few long setae dorsodistally. Penulti-
mate segment unarmed, with few short se-
tae on dorsal face subdistally. Basal seg-
ment slightly produced distomesially, with
1 acute spinule at ventrodistal mesial angle.

Antennal peduncles (Fig. 1A, B, G, H)
moderately long, with distal margin of
fourth segment reaching beyond base of
corneae but not to distal margin, when fully
extended; with supernumerary segmenta-
tion. Fifth and fourth segments with few
scattered long setae. Third segment with
ventrodistal angle produced. Second seg-
ment with dorsolateral distal angle pro-
duced, terminating in strong spine accom-
panied ventrally by short accessory spine,
dorsolateral face with several tubercles,
dorsomesial distal angle with acute spine.
First segment with strong, hook—shaped
spine laterally and strong spine at ventro-
distal margin mesially. Antennal acicle
moderately short, overreaching ocular pe-
duncles; straight, blunt-tipped, with scat-
tered setae; mesial face slightly tuberculate.
Flagella very long, 6.2 times length of
shield, scarcely setose.

Mandible (Fig. 2A) without distinguish-
ing characters. Maxillule (Fig. 2B, C) with
external endopodal lobe obsolete; internal

383

lobe with 1 apical bristle. Maxilla (Fig. 2D)
with moderately broad scaphognathite. First
and second maxillipeds (Fig. 2E, F) without
distinguishing characters. Third maxilliped
(Fig. 2G—1) with 3 acute corneous teeth on
basis; ischium with crista dentata well de-
veloped, composed of 15—16 corneous teeth
and with 1 strong accessory tooth; merus
with acute dorsodistal spine.

Sternite of 3rd maxillipeds unarmed,
with shallow median cleft.

Right cheliped of male (Fig. 3A—D) larg-
er and stouter than that of left; compressed
dorsoventrally. Dactyl distinctly shorter
than palm, 0.32:1.00; forming weak hiatus
with fixed finger; terminating in strong,
curved calcareous claw; dorsal face convex,
tuberculate mesially, with tufts of long se-
tae; cutting edge with prominent blunt-
tipped calcareous tooth medially. Fixed fin-
ger with dorsal surface tuberculate, with
tufts of long setae; cutting edge with minute
calcareous teeth on distal half. Palm very
long, with dorsal surface covered with nu-
merous spiniform tubercles, with 2 faint
longitudinal sulci on proximal *%4, continu-
ing to proximal very deep depression, dor-
somesial and dorsolateral surfaces covered
with dense tubercles, with proximal angles
protruded and strongly armed with spini-
form tubercles; ventral surface covered with
dense minute tubercles and scattered long
setae. Carpus distinctly shorter than palm,
0.56:1.00; swollen distally; dorsal surface
flat, with scattered spiniform tubercles, dor-
solateral and dorsomesial margins forming
distinct ridges, each armed with strong spi-
niform tubercles; lateral, mesial and ventral
surfaces glabrous. Merus with dorsal mar-
gin irregular, covered with strong spiniform
tubercles and protuberances, and scattered
long setae, dorsodistal angle bearing slight-
ly upturned large spine; lateral and mesial
faces tuberculate, ventromesial and ventro-
lateral faces tuberculate with distal angles
only protruded. Ischium with very strong,
acute spine on ventrodistal margin laterally.
Coxa with tubercles on ventromesial distal
angle.

384 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Jcelopagurus tuberculosus, new species: holotype male, SL = 5.70 mm, Okinawa, Japan (CBM ZC 4700);
A-H, cephalothorax and cephalic appendages: A, whole (dorsal view); B, whole (right lateral view); C, tip of sexual
tube arising from coxa of night fifth pereopod; D, ocular peduncle and anterolateral portion of shield (right, dorsal
view); E, antennule (right, lateral view); E same, distal portion of basal segment (mesial view); G, antenna (night,
lateral view); H, same (ventral view); K—M, proportion of cephalothorax and abdomen: K, holotype male; L, paratype
male, SL = 4.90 mm (CBM-ZC 4701); M, paratype female, SL = 5.10 mm (CBM-ZC 4702). Icelopagurus crosnieri

VOLUME 112, NUMBER 2

Left cheliped of male (Fig. 3E—G) slight-
ly shorter than that of right, 0.92:1.00; con-
siderably more slender than right, 0.65:1.00
in widest portion of palm and 0.32:1.00 in
widest portion of carpus, respectively. Dac-
tyl strongly curved distally, forming large
hiatus with fixed finger; approximately
equaling palm in length, with tufts of long
setae; terminating in strong corneous claw;
cutting edge with numerous tiny corneous
teeth. Fixed finger tuberculate laterally,
with tufts of long setae; terminating in
strong recurved corneous claw; cutting
edge with numerous tiny corneous teeth.
Palm with dorsal surface tuberculate, with
very broad faint longitudinal sulcus on
proximal half continuing to proximal de-
pression; mesial, lateral and ventral surfaces
covered with dense blunt or spiniform tu-
bercles. Carpus moderately long, approxi-
mately equaling palm in length; dorsal sur-
face flat, with numerous blunt or spiniform
tubercles, dorsolateral and dorsomesial
Margins distinct, each armed with strong,
spiniform tubercles; mesial and lateral faces
granular or tuberculate. Merus with dorsal
margin strongly tuberculate, dorsodistal an-
gle bearing very large spine; lateral face
granular; ventrolateral margin tuberculate,
with distal angle bearing blunt-tipped spine,
ventromesial distal angle only produced and
unarmed. Ischium with strong spine at ven-
trolateral distal angle.

Second (Fig. 2J—O) and third (Fig. 2P—
R) pereopods morphologically similar; third
Slightly longer than second, 1.09:1.00, es-
pecially dactyl of third longer than second,
1.15:1.00, but broader in second than in
third (ratio of width/length = 0.17 in sec-
ond but 0.12 in third). Dactyls slightly
shorter than propodi (second), 0.91:1.00, or
as long as propodi (third); broad, sabre
blade-shaped; each terminating in strong
corneous claw; mesial faces concave lon-

—

385

gitudinally, with scattered long setae; dor-
somesial and ventromesial margins each
with row of corneous spinules; lateral faces
slightly convex longitudinally. Propodi with
entire surfaces minutely tuberculate; dorsal
margins each with 2 to 5 thick, spine-like
setae, dorsodistal margins each with pair of
thick, spine-like setae; ventral margins each
with 1 or 2 thick, spine—like setae. Carpi
each with tiny spinule at dorsodistal angle;
dorsal, lateral and mesial surfaces covered
with dense tubercles or granules. Meri with
dorsal, lateral and mesial surfaces tubercu-
late; dorsal face tuberculate, dorsodistal an-
gles each only slightly protruded (second)
(with tiny spinule on left second of paratype
male) or bearing very strong spine (third);
dorso-subdistal portion with 1 strong spine
(second, but 2 spines in right second of par-
atype male), or 2 strong spines (third); ven-
trolateral distal angles each bearing large
spine (second) or slightly protruded (third),
ventromesial distal angles unarmed; distal
half of ventrolateral margins slightly (sec-
ond) or strongly (third) expanded ventrally,
but ventromesial margins not. Female with
coxae of third pereopods each with gono-
pore (Fig. 4N).

Fourth pereopods (Fig. 4A—C) semiche-
late. Dactyl moderately long, terminating in
strong corneous claw; prominent preungual
process present, covered with dense short
setae; cutting edge with several corneous
teeth. Propodus with rasp composed of 1
row of strong corneous scales along 0.80
length of ventral margin; dorsodistal angle
unarmed. Carpus with blunt-tipped spine at
dorsodistal angle.

Fifth pereopods chelate (Fig. 4D). Dac-
tyls and propodi with well developed rasps.
Male (Fig. 1B, C) with right coxa bearing
moderately-short sexual tube directed to ex-
terior; sexual tube with blunt-tipped apex,
not filamentous, covered with scattered

McLaughlin, 1997: paratype female, SL = 3.15 mm, Tanimbar Island, Indonesia (MNHN-Pg 5279): I, carapace,
dorsal view; J, same, lateral view; N, proportion of cephalothorax and abdomen. Scales = 1 mm.

386 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

E Py
LN AS Me
0h

a Rete
. ae '
is

Ct

cn

ae
oth
ee

Fig. 2. Icelopagurus tuberculosus, new species: holotype male, SL = 5.70 mm, Okinawa, Japan (CBM ZC
4700); A-—I, mouthparts (right): A, mandible (internal view); B, maxillule (external view); C, same, endopod;
D, maxilla (external view); E, first maxilliped (external view); E second maxilliped (external view); G, third
maxilliped (external view); H, same, ischium and basis (external view); I, same, ischium and basis (internal
view); J—O, second pereopod (left): J, whole (lateral view); K, dactyl (mesial view); L, same, ventral portion of
mesial face; M, same (lateral view); N, propodus, schematic diagram indicating condition of spine-like setae

VOLUME 112, NUMBER 2

short setae apically; gonopore of left coxa
(Fig. 4E) with vas deferens only slightly
protruded, obscured by dense short setae
posteriorly and dense medium-length setae
anteriorly. Female with nearly symmetrical
coxae (Fig. 40).

Sternite of second pereopods (Fig. 4F)
broad, subrectangular; divided into 2 lobes
by longitudinal shallow median groove; an-
terior margin of each lobe round, unarmed,
with long setae. Sternite of third pereopods
(Fig. 4E G) with perpendicular anterior
lobe bearing pair of small projections
armed with minute tubercles and fringed
anteriorly with setae; horizontal posterior
plate very broad, with setae anteriorly; sub-
divided into 2 lobes by longitudinal shallow
median groove. Sternite of fourth pereo-
pods (Fig. 4F) as transverse rod, accompa-
nied by setae anteriorly. Sternite of fifth pe-
reopods (Fig. 4E, F) reduced to very narrow
transverse rod with pair of round projec-
tions with setae; widely separated from pre-
ceding sternal plates.

Abdomen (Fig. 1K—M) small in holotype
male, comparatively reduced in paratype
male, reduced in paratype female; membra-
nous, dextrally twisted. Tergite of first ab-
dominal somite small and chitinous. Ter-
gites of second to fifth abdominal somites
not clearly delineated, membranous. Tergite
of sixth abdominal somite calcified, sub-
rectangular, unarmed; divided into anterior
and posterior lobes by shallow, transverse
groove. Male with 3 unpaired left unira-
mous pleopods (Fig. 4H—J) fringed with
long setae, arising from third to fifth ab-
dominal somites. Female with no paired
first pleopods modified as gonopods but
with 3 unpaired left biramous pleopods
(Fig. 4P—R) arising from second to fourth
abdominal somites; existence of fourth ple-
opod uncertain because of damaged abdo-

—

387

men of single female specimen. Uropods
(Fig. 4K, L) strongly asymmetrical, left dis-
tinctly larger than right; rasps of exopods
and endopods well developed; protopods
with some tubercles on posterodistal face
(right) or unarmed (left). Telson (Fig. 4M)
with lateral constrictions; posterior lobes
separated by very broad median cleft; ter-
minal margins of left and right lobes round,
with short setae mesially.

Variations.—A marked sexual dimor-
phism is seen in chelipeds. Right cheliped
of female (Fig. 3H—L) distinctly more slen-
der and more compressed dorsoventrally
than male; dactyl shorter than palm, 0.48:
1.00, cutting edge with 2 broad but weak
calcareous teeth on proximal half and many
small corneous teeth on distal half; fixed
finger not forming hiatus with dactyl, cut-
ting edge with minute calcareous teeth on
distal half; palm long, with dorsal face flat
and weakly granular; carpus long and slen-
der, approximately equaling palm in length,
dorsal surface flat, with dorsolateral and
dorsomesial margins each forming distinct
ridge bearing row of large tubercles; merus
with row of large tubercles and long setae
on dorsal margin, dorsodistal angle bearing
strong spine, ventromesial and ventrolateral
distal angles each with blunt-tipped spine.
Left cheliped of female (Fig. 2M-—P) more
slender than male; slightly shorter than
right, 0.90:1.00; more slender than right,
0.75:1.00 in widest portion of palm and
0.79:1.00 in widest portion of carpus; dac-
tyl and fixed finger straight, not forming hi-
atus; palm slender, with dorsal surface mi-
nutely tuberculate, median portion flat; car-
pus longer than palm, 1.46:1.00, slender,
dorsal surface flat, dorsolateral and dor-
somesial margins forming ridges, each
bearing distinct row of granules; merus
with dorsal face tuberculate, dorsodistal an-

(mesial view); O, carpus and distal portion of merus (lateral view); P—R, third pereopod (left): P, whole (lateral
view); Q proximal portion of carpus and distal portion of merus (lateral view); R, merus (mesial view). Scales

= | mm.

388 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ae \ i f
Zt +
(| So aR
Vag: a Se W ae ZA
wy LAS Sys
y ERS
IN 3
i — a Mr
oe aes
Seay 3

~s
Its.
ws

ny yp
a 7 ~_ ~

Wwe

c

e cue Sc

Sor we Seice
segs Lote

wee te pie ret
aed eee
—

Fig. 3. Icelopagurus tuberculosus, new species. A—D, male right cheliped: A, whole (dorsal view); B, whole
(lateral view); C, dactyl and fixed finger (dorsal view); D, merus and ischium (ventral view); E—G, male left
cheliped: E, whole (lateral view); E whole (dorsal view); G, merus and ischium (ventral view); H—L, female
right cheliped: H, whole (lateral view); I, whole (dorsal view); J, dactyl and fixed finger (dorsal view); K, distal
portion of merus (lateral view); L, merus and ischium (ventral view); M—P, female left cheliped: M, whole
(dorsal view); N, whole (lateral view); O, merus and ischium (ventral view); P, dactyl and fixed finger (ventral

VOLUME 112, NUMBER 2

gle bearing very large spine, with distal an-
gles each bearing blunt-tipped spine.

The size of the abdomen is subject to
variation. The abdomen is small in the ho-
lotype male and comparatively reduced in
male and female paratypes, although no dis-
tinct morphological variations are seen in
the uropods and telson. Neither the number
or shape of the pleopods differ between the
holotype and paratype males.

Coloration.—Unknown.

Habitat.—10-—20 m,
Housing unknown.

Distribution.—Known only from the
type locality, Kumejima, Okinawa.

Etymology.—The name tuberculosus is
from the Latin, tuberculum, diminutive of
tuber, a swelling or lump, and the suffix,
-osus, alluding to the tuberculate surfaces
of the shield and pereopods characteristics
of this new species.

Remarks.—Because the posterior portion
of the abdomen of the paratype female is
damaged, it cannot be confirmed whether or
not it had a fourth pleopod.

Comparison with Icelopagurus crosni-
eri—The new species agrees with all of the
diagnostic characters of Icelopagurus pro-
posed by McLaughlin (1997) but is unique
in having a large well-calcified, immovable
plate that is partially fused with the poste-
rior and posterolateral margins of the
shield. This plate is most probably derived
from the right and left accessory portions
of the shield (sensu Lemaitre 1995) plus the
linea transversalis (sensu Boas 1926, Pil-
grim 1973). Generally in hermit crabs (ex-
cluding Pylochelidae and Lithodidae), the
accessory portions of the shield consists of
small, elongated movable, partially-calci-
fied plates on either side of the posterolat-
eral margin of the shield. The linea trans-
versalis is a narrow, uncalcified hinge sep-
arating the gastric and cardiac regions of

sandy bottom.

——

389

the carapace. However, as discussed below,
well-developed and strongly-calcified ac-
cessory portions and linea transversalis oc-
cur in five genera which comprise a highly
specialized group within pagurids (Poupin
& McLaughlin 1996, McLaughlin 1997).

McLaughlin (1997) did not mention ac-
cessory portions of shield and linea trans-
versalis in Jcelopagurus crosnieri. Two par-
atype specimens of that species were ex-
amined (Fig. 1I, J, N), and no such struc-
tures were found; instead, there is a pair of
broad, partially-calcified accessory portions
of the shield and a narrow, uncalcified linea
transversalis along the posterior margin of
the shield. These structures are clearly sep-
arated from the shield by a soft membrane
(Fig: AL Sp

The morphology of the dorsal surface of
the shield in the new species, J. tuberculo-
sus, iS also distinct. Anteriorly there is a
transverse row of four prominent protuber-
ances and a pair of similar protuberances
on the anterolateral margins, all strongly
armed with minute tubercles. The ocular
peduncles are also distinct, armed with
many tubercles. None of these characters
occur in J. crosnieri.

The size of the abdomen of J. tubercu-
losus is small or somewhat reduced (Fig.
1K—M). The abdomen in J. crosnieri is nor-
mal, similar to that of typical pagurid crabs
(Fig. 1N). Other morphological differences
between the two species are given in Ta-
ble 1.

Discussion

When establishing Jcelopagurus, Mc-
Laughlin (1997) stated that it was superfi-
cially very similar to Catapagurus, both
genera being characterized by a well-devel-
oped right sexual tube in males, females
lacking specialized secondary sexual char-

view). A—G, holotype male, SL = 5.70 mm, Okinawa, Japan (CBM ZC 4700); H-P, paratype female (CBM ZC

4702), SL = 5.10 mm, Okinawa, Japan. Scales = 1 mm.

390 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1 mm

K, L, M

1 mm

1 mm

Y piv WN Lith foray Ce
Yf WSS OZ Yi

N, P,Q,R

O 1 mm

Fig. 4. Icelopagurus tuberculosus, new species. A—C, fourth pereopod (left): A, whole (lateral view); B,
dactyl (lateral view); C, terminal claw and preungual process of dactyl (mesial view); D, fifth pereopod (right);
E, coxae and sternite of fifth pereopods of male (ventro-posterior view); EK coxae and sternites of second to fifth
pereopods of male (ventral view); G, anterior plate of sternite of third pereopods (ventral view); H—J, first to
third pleopods of male (arising from third to fifth abdominal somites); K, left uropod; L, right uropod; M, telson
(dorsal view); N, coxae and sternites of second to fifth pereopods of female (ventral view); O, coxae and sternite
of fifth pereopods of female (ventral view); P—R, first to third pleopods of female (arising from second to fourth
abdominal somites). A-M, holotype male, SL = 5.70 mm, Okinawa, Japan (CBM ZC 4700); N-R, paratype
female, SL = 5.10 mm, Okinawa, Japan (CBM ZC 4702). Scales = 1 mm.

VOLUME 112, NUMBER 2

391

Table 1.—Comparison of morphology of ambulatory pereopods and telson in species of /celopagurus.

I. tuberculosus,
new species

I. crosnieri

Second and third pereopods

Dactyl: mesial face
dorsomesial margin

concave

cormneous spines
ventromesial margin
corneous spines

Propodus: dorsal face granular

spine-like setae on distal half

Carpus: dorsal face
Merus: dorsal face

granular

ventrolateral face granular
Posterior lobes of telson

posterolateral margin unarmed

mesial margin unarmed

row of numerous short

row of numerous short

1 or 2 very strong spines

flat or slightly convex proximally

row of short setae on proximal half and
row of every long setae on distal half

very few setae

few irregular rows of spines
spine-like setae entirely

few irregular rows of spines
rugose

numerous spines

4 long corneous spine-like setae
few calcareous spines

acters, a rudimentary or vestigial external
endopodal lobe on the maxillule, more or
less reduced crista dentata with one acces-
sory tooth, and a distinct tubular preungual
process on the dactyl of the fourth pereo-
pod. However, Jcelopagurus is readily sep-
arated from Catapagurus by the shortness
of the sexual tube and the distinct devel-
opment of the telson.

Although the new species, J. tuberculo-
sus, differs from both I. crosnieri and Ca-
tapagurus species in having short ocular
acicles, the morphology of the ambulatory
pereopods of I. tuberculosis is more similar
to those of Catapagurus species which have
dactyls shaped like sabre blades, such as C.
granulatus Edmondson, 1951 and C. ensi-
fer Henderson, 1893, than that of I. cros-
nieri. These characters include the ambu-
latory pereopods ending in sabre blade-
shaped dactyls with dorsal and ventral rows
of numerous corneous spine-like setae on
the mesial face, and the meri having one or
two very strong, subdistal dorsal spines.
The ambulatory dactyls of I. crosnieri are
not sabre blade-shaped, their mesial faces
having a dorsal row of short setae on the
proximal half, very long setae on the distal
half, and ventrally only a few setae; the
meri of the ambulatory pereopods of J.
crosnieri lack a distinct dorsal spine.

During the course of this study, similar
patterns of calcification were found in the
posterior carapace in both /celopagurus and
Catapagurus species examined (C. granu-
latus, C. ensifer, C. japonicus, and three un-
described species collected). There are a
pair of partially calcified bands in the areas
between the cardiac sulci and sulci cardio-
branchialis in J. tuberculosus (Fig. 1A) and
in I. crosnieri (Fig. 11). When McLaughlin
& Lemaitre (1997) determined character
states of calcification of the posterior cara-
pace of hermit crabs, i.e., entirely calcified;
posterolateral and/or posteromedial calcifi-
cation; and entirely membranous or with
only scattered small areas of calcification,
they included /celopagurus in the latter cat-
egory. Although I agree with their classifi-
cation because of small areas of those par-
tially calcified bands, I think that future de-
tailed studies on patterns of calcification of
posterior carapace may provide useful in-
formation in hermit crab phylogeny.

McLaughlin (1997) also stated that Ice-
lopagurus was ostensibly quite similar to
Pagurodes Henderson, 1888 sensu stricto,
which is currently represented only by Pa-
gurodes inarmatus Henderson, 1888. How-
ever, P. inarmatus differs from both J. cros-
nieri and I. tuberculosus in having tricho-
branchiate gills, fourth pereopod dactyls

392

without a preungual process, and telson
with acutely triangular posterior lobes.

The new species, I. tuberculosus, shares
several characters with taxa considered
among the most highly specialized pagurid
genera by Poupin & McLaughlin (1996)
and McLaughlin (1997), namely, Alainop-
agurus Lemaitre & McLaughlin, 1995,
Alainopaguroides McLaughlin, 1997, Soli-
tariopagurus Tirkay, 1986, Porcellanopa-
gurus Filhol, 1885, and Ostraconotus A.
Milne Edwards, 1880. These five genera are
characterized by a well-calcified shield, dis-
tinct reduction of the abdomen, symmetri-
cal uropods and telson, very broad sternites
on the pereopods, absence of paired or un-
paired pleopods in males, and lack of left
fifth pleopod in females. Because of these
morphological similarities, McLaughlin
(1997) stated that these genera might con-
ceivably be considered as sister taxa. Since
I. tuberculosus also has a vaulted, well-cal-
cified shield, a more or less reduced abdo-
men and elongated, slender chelipeds and
ambulatory legs with very broad sternites,
general impression of similarity is gained
between J. tuberculosus and these genera.

As previously mentioned, the large, well-
calcified and immovable plate on the cara-
pace of J. tuberculosus is derived from the
well-developed and calcified accessory por-
tions of the shield and linea transversalis.
The well-calcified condition of the linea
transversalis is also seen in the five men-
tioned specialized genera (Table 2), and in
at least two genera, Solitariopagurus and
Alainopagurus, it is partially fused with the
shield (Poupin & McLaughlin 1996, Le-
maitre & McLaughlin 1995, McLaughlin
1997). The development and strong calci-
fication of shield accessory portions are
also known in these genera (Lemaitre &
McLaughlin 1995, Poupin & McLaughlin
1996, McLaughlin 1997) (Table 2). In all
of these genera, the linea transversalis and
accessory portions are not completely fused
together, being at best partially fused in So-
litariopagurus and Porcellanopagurus.

In overall morphology, the two species

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

of Icelopagurus are quite similar to Alain-
opaguroides, a monotypic genus represent-
ed by A. lemaitrei McLaughlin, 1997, in the
morphology of the shield, cephalic append-
ages, mouthparts, pereopods and secondary
sexual characters in the both sexes (Table
2). Icelopagurus tuberculosus is more sim-
ilar to Alainopaguroides than I. crosnieri in
the short ocular acicles, ambulatory legs
with more or less blade-shaped dactyls
bearing dorsal and ventral rows of regular-
ly-spaced setae on the mesial face, and a
more or less reduced abdomen. Alainopa-
guroides is distinguished from Icelopagu-
rus in lacking unpaired or paired pleopods
in males, having vestigial propodal rasps on
the fourth pereopods, and symmetrical uro-
pods and telson (Table 2). In addition, J.
tuberculosus differs from Alainopaguroides
in having large protuberances anteriorly on
the shield.

Icelopagurus shares some characters
with Solitariopagurus, currently recognized
as having three representatives, S. profun-
dus Tiirkay, 1986, S. triprobolus Poupin &
McLaughlin, 1996, and S. tuerkayi Mc-
Laughlin, 1997. Both genera have a vault-
ed, well-calcified shield, obsolete or no ex-
ternal endopodal lobe of the maxillule,
elongated, slender chelipeds and ambula-
tory legs, and a moderately long right sex-
ual tube in males. Further characters shared
between J. tuberculosus and Solitariopa-
gurus, but not with I. crosnieri, include four
large protrusions on the antero-dorsal por-
tion of the shield (except S. profundus) with
a well-developed, calcified accessory por-
tion (but not fused as one in Solitariopa-
gurus), and a more or less reduced abdo-
men. In Solitariopagurus, two or three
wing-like expansions (except lateral projec-
tions of the shield) occur on each lateral
margin of the shield. Also, J. tuberculosus
has a projection on each anterolateral mar-
gin of the shield, and distinct anterolateral
angles armed with tubercles. Solitariopa-
gurus and Icelopagurus are immediately
distinguished by a number of gills and
shapes of a rostrum, lateral projections, oc-

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394

ular facicles and morphology of the abdo-
men and its appendages (Table 2).
Porcellanopagurus also has two or three
wing-like expansions on each lateral margin
of the shield (Borradaile 1916, Forest 1951,
Takeda 1981, 1985; Suzuki & Takeda 1987,
McLaughlin 1997, McLaughlin & Hogarth
1998). However, it differs from [celopagu-
rus in morphology of the abdomen and its
appendages, and male sexual tube (Table 2).
Ostraconotus (based on Tiirkay 1986, Pou-
pin & McLaughlin 1996, McLaughlin
1997, McLaughlin & Lemaitre 1997) also
has the shield with a transverse row of four
somewhat large tubercles on the anterodor-
sal portion, and a right sexual tube in males,
but differs from Jcelopagurus in number of
gills and structure of the abdomen and its
appendages (Table 2). Furthermore, in the
Paguridae, Ostraconotus is the only genus
in which the posterior carapace is complete-
ly calcified (McLaughlin & Lemaitre 1997).

Acknowledgments

I am grateful to K. Nomura (Kushimoto
Marine Park, Wakayama-kKen) and Y. Sak-
amoto (Kumejima, Okinawa) for the op-
portunity to examine the specimens de-
scribed here and to N. Ngoc-Ho (Muséum
national d’Histoire naturelle, Paris), P.
Clark and M. Lowe (The Natural History
Museum, London), L. G. Eldredge (Bernice
P. Bishop Museum, Hawaii), H. Namikawa
and K. Hasegawa (Biological Laboratory,
Imperial Household, Showa Memorial In-
stitute, Tsukuba) and Y. Yabumoto (Kita-
kyushu Museum of Natural History) for the
loans of specimens. Mark J. Grygier (Lake
Biwa Museum) kindly commented on an
earlier draft of the manuscript. I am deeply
indebted to Dr. P. A. McLaughlin (Shannon
Point Marine Center, Western Washington
University) for her kind advice, generous
help and elaborate review of a draft of the
manuscript. Dr. R. Lemaitre and two anon-
ymous referees greatly improved the man-
uscript.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Literature Cited

Boas, J. E. V. 1926. Zur Kenntnis Symmetrische Pa-
guriden.—Det Kongelige Danske Videnskaber-
nes Selskabs Biologiske Meddelelser 6:1—52.

Borradaile, M. A. 1916. Crustacea, part II. Porcellan-
opagurus: an instance of carcinization.—British
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1913.—Natural History Reports, Zoology 3:
111-126.

Edmondson, C. H. 1951. Some central Pacific crusta-
ceans.—Occasional Papers of the Bernice Pau-
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Filhol, H. 1885. Description d’un nouveau genre de
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Forest, J. 1951. Contribution a |’étude du genre Por-
cellanopagurus Filhol (Paguridae). I. Descrip-
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lected by H. M. S. Challenger during the years
1873—76.—Report on the scientific results of
the exploratory voyage of H.M.S. Challenger
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. 1893. A contribution to Indian carcinology.—
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genus Xylopagurus A. Milne Edwards, 1880
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, & P. A. McLaughlin. 1995. Alainopagurus
crosnieri, gen. et sp. nov. (Decapoda, Anomura,
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du Muséum national d’Histoire naturelle, Paris
4e série 17:273-—282.

McLaughlin, P. A. 1974. The hermit crabs (Crustacea,
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. 1997. Crustacea Decapoda: Hermit crabs of

the family Paguridae from the KARUBAR

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séum national d’ Histoire naturelle, Paris 172.

, & R. Lemaitre 1997. Carcinization in the An-

omura—fact or fiction? I. Evidence from adult

morphology.—Contributions to Zoology 67:

79-123.

, & P. J. Hogarth 1997. Hermit crabs (Deca-

poda: Anomura: Paguridae) from the Sey-

chelles.—Zoologische Verhandlingen 318:1—

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Milne Edwards, A. 1880. Reports on the results of
dredging, under the supervision of Alexander
Agassiz, in the Gulf of Mexico, and in the Ca-
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Nomura, K., S. Nagai, A. Asakura, & T. Komai. 1996.
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Kingdom 53:363-—396.

Poupin, J.. & P. A. McLaughlin 1996. A new species
of Solitariopagurus Tiirkay (Decapoda, Ano-
mura, Paguridae) from French Polynesia.—Bul-

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Takeda, M. 1981. A new hermit crab of the genus Por-
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Takeda, M. 1985. Occurrence of a new hermit crab of
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pan.—Memoirs of the National Science Muse-
um, Tokyo 18:141-144.

Tiirkay, M. 1986. Crustacea Decapoda Reptantia der
Tiefsee des Roten Meeres.—Senckenbergiana
maritima 18:123—185.

Yokoya, Y. 1933. On the distribution of Decapod crus-
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1930.—Journal of the College of Agriculture,
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(2):396—404. 1999.

A new species of crayfish of the genus Procambarus
(Crustacea: Decapoda: Cambaridae) from Veracruz, Mexico

Yolanda Rojas, Fernando Alvarez, and José Luis Villalobos

Colecci6n Nacional de Crustaceos, Instituto de Biologia,
Universidad Nacional Autonoma de México, Apartado Postal 70-153,
México 04510 D.F, México

Abstract.—Procambarus (Austrocambarus) citlaltepetl, new species, is de-
scribed from Ciudad Mendoza, Veracruz, Mexico. The new species can be
placed in the mexicanus Group, and is most similar to P. (A.) mexicanus, from
which it can be distinguished by the absence of branchiostegal spines, gonopod
with blade-like mesial process curved over the apex and thicker lateral portion,
and annulus ventralis with complete preannular plate. SEM micrographs of the
male and female genitalia of the new species, as well of those of P. (A.)
mexicanus, are provided to facilitate comparisons, and to clarify the identity

of the latter.

Confusion concerning the identity of the
crayfish Procambarus (Austrocambarus)
mexicanus (Erichson, 1846) has existed for
more than a century, primarily because of
the incomplete original description lacking
figures, loss of the holotype, and the high
degree of morphological variation found
among the many populations assigned to
this taxon (Villalobos 1954). Several efforts
have been made to clarify the status of P.
(A.) mexicanus. Villalobos (1954) described
four new species, one subspecies, and re-
described six species of Procambarus, in-
cluding P. (A.) mexicanus, most from cen-
tral and southern Veracruz, and proposed
the “‘mexicanus’’ Group for eight of these
species. Hobbs (1987) synonymized Cam-
barus aztecus Saussure, 1857 with P. (A.)
mexicanus, which he redescribed, listing 23
collection sites; he also described P. (A.)
olmecorum Hobbs, 1987 an placed it in the
‘““mexicanus’’ Group.

In spite of these efforts, the distribution
range of P. (A.) mexicanus as reviewed by
Hobbs (1987), still included a vast area in
the states of Veracruz, Puebla, and Oaxaca
(Fig. 1), and included a wide array of mor-
phological forms. Presumably, the northern

limit of the distribution range is in the en-
virons of Jalapa, Veracruz; the southern
limit occurs in Tuxtepec, Oaxaca; to the
west it occurs along the slope of the Mex-
ican central plateau; and to the east it reach-
es the Sierra de Los Tuxtlas, Veracruz (Fig.
i):

After a detailed examination of speci-
mens from Rincon de la Doncella, Ciudad
Mendoza, it became clear that they repre-
sented a new species, different but morpho-
logically similar to P. (A.) mexicanus. In
this study we describe this new species,
Procambarus (Austrocambarus) citlalte-
petl, and in order to clearly separate the
new species from P. (A.) mexicanus, we
provide scanning electron microscope
(SEM) micrographs of the gonpod, annulus
ventralis, and epistome of both species. The
SEM micrographs of P. (A.) mexicanus
were taken from specimens from the type
locality in order to help in the recognition
of other new forms which may exist within
this Group.

All specimens used are deposited in the
Coleccién Nacional de Crustaceos, Instituto
de Biologia, Universidad Nacional Aut6on-
oma de México (CNCR). Other abbrevia-

VOLUME 112, NUMBER 2 S97

Mexico

Jals
Coate pec»
Teocelo

rac ©

@® Huatusco
eCoscomatepec
in de las Flores

udad Mendozé e8 Orizaba

6
santiago

Puebla @® Suxtepec

Veracruz

Fig. 1. Localities where Procambarus (Austrocambarus) mexicanus has been collected in the states of Pueb-
la, Oaxaca, and Veracruz.

398

tions used are: coll, collector; RL, rostrum
length; TCL, total carapace length; USNM,
National Museum of Natural History,
Smithsonian Institution, Washington, D.C.

Procambarus (Austrocambarus)
citlaltepetl, new species
Figs. 2A—C, 3A-C, 4

Diagnosis.—Body pigmented, eyes nor-
mally developed. Rostrum devoid of mar-
ginal spines. Acumen length ranging from
0.07% to 0.14% of RL (X = 0.10%). Car-
apace with or without small cervical tuber-
cle. Areola ranging from 7.1 to 16.8 (X =
10.9) times as long as wide, 30.8% to
39.5% (KX = 34.9%) of TCL, with 1-3
punctations across narrowest part. Subor-
bital angle weak, infraorbital spine absent.
Branchiostegal spine absent. Postorbital
ridges well marked, slightly sinuous in dor-
sal view, converging anteriorly, ending in
spine anteriorly. Antennal scale varying
from 1.61 to 1.96 (X = 1.79) times as long
as wide; maximum width at midlength.
Chelipeds subequal, shorter than TCL, par-
tially covered with squamous tubercles;
chela moderately elongated, not pubescent,
fingers shorter than palm, palm about twice
as long as broad; movable finger with squa-
mous tubercles along dorsal margin, oppos-
able margin devoid of tubercles distally,
finely serrate; fixed finger punctate, devoid
of tubercles except for inferior margin, op-
posable margin finely serrate distally with
5 tubercles proximally. First pair of pereio-
pods shorter than total length of body, cov-
ered with small squamous tubercles from
distal half of merus to proximal half of fin-
gers. Ischium of third pereiopod armed with
single, acute hook, reaching beyond artic-
ulation with basipodite. First pair of pleo-
pods of male form I symmetrical, devoid of
subterminal setae, reaching coxae of third
pair of pereiopods; shoulder on cephalic
surface slightly concave, forming rounded
angle with cephalic border (Figs. 2A, 3A);
mesial process blade-like, becoming nar-
rower distally, laterodistally oriented (Fig.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

2B), curved over apex in cephalic view
(Fig. 2C); cephalic process absent; central
projection triangular, cephalodistally orient-
ed. Female with hinged annulus ventralis;
preannular plate strong, complete, oriented
perpendicular to longitudinal axis of body;
annulus oval-shaped, divided by deep and
narrow mesial groove, seminal receptacle
opening on caudal portion as sinuous
groove; postannular sclerite strong, oval-
shaped, wider than annulus, cephalic mar-
gin subacute, caudal margin almost straight
(Fig. 3B). Sternal plates adjacent to fifth
pair of pereiopods wide, rounded, extend-
ing mesially, reaching postannular sclerite.
Coxae of fourth and fifth pair of pereiopods
separated.

Measurements of types.—See Table 1.

Holotypic male, form I.—Body pigment-
ed. Cephalothorax subcylindrical, becoming
thicker in branchial region, 0.91 times
length of abdomen. Areola 15.6 times as
long as wide, 34.9% of TCL (Fig. 4A).
Dorsal surface of carapace densely punctate
along branchial and cardiac regions. Lateral
surfaces finely granulate along hepatic and
branchial regions. Rostrum devoid of mar-
ginal spines; dorsal surface slightly con-
cave, densely punctate posteriorly. Rostrum
with lateral margins convergent, anterior
width 2.1 mm, posterior width 3.6 mm,
ending anteriorly in acute angle; subrostral
ridges not evident in dorsal view, oriented
ventrally. Acumen length 0.11% of RL,
reaching anterior portion of third article of
anntenular peduncle. Postorbital ridge mod-
erately strong, ending anteriorly in trian-
gular spine oriented laterally. Suborbital an-
gle weak, infraorbital spine absent. Bran-
chiostegal spine absent. Cervical groove
with blunt tubercle in hepatic region (Fig.
4B).

Abdomen narrower than carapace. So-
mites with tergal region finely punctate,
pleural region densely punctate. Dorsal sur-
face of telson covered with short setae, ce-
phalic section with 3 spines in posterolat-
eral angles, caudal section with posterior
margin rounded. Epistome triangular, ante-

VOLUME 112, NUMBER 2 Sly)

- ou
ant

: - i | V4 . :
Nah: CRE
Fig. 2. Lateral (A, D), caudal (B, E), and cephalic views (C, F) of left gonopod. A-C, Procambarus (Aus-

trocambarus) citlaltepetl, new species, holotype (CNCR 17317); D-EK Procambarus (Austrocambarus) mexican-
us (CNCR 2339).

rior vertex triangular, posterior angles with second, larger depression (Fig. 3C).
rounded, ventral surface slightly concave; Antennal scale 1.72 times as long as wide,
cephalic lobe elongated, oval-shaped; main maximum width at midlength, lateral mar-
body with well defined fovea, anteriorly gin terminating in spine (Fig. 4C). Antennal

400 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Mesial view of left gonopod (A, D), annulus ventralis (B, E), and epistome (C, F). A-C, Procambarus
(Austrocambarus) citlaltepetl, new species holotype (CNCR 17317); D—E Procambarus (Austrocambarus) mex-
icanus (CNCR 2339).

peduncle with basis ending in short disto- cle; midventral surface of ischium covered
lateral spine, ischium with lateral blunt tu- with fine, simple, plumose setae, laterally
bercle. Third maxilliped reaching distal with tufts of short setae.

portion of third segment of antennal pedun- Chelipeds shorter than TCL, tuberculate

VOLUME 112, NUMBER 2

401

Fig. 4. Procambarus (Austrocambarus) citlaltepetl, new species (all figures of holotype (CNCR 17317),
except F of female allotype (CNCR 17318)): A, carapace, dorsal view; B, carapace, lateral view; C, antennal
scale; D, distal podomeres of cheliped; E, basal podomeres of left second, third, and fourth pereiopods; F distal

podomeres of cheliped. Scale bars represent 1 mm.

from distal half of merus to midlength of
fingers. Right chela moderately elongated,
2.9 times as long as broad, not pubescent,
fingers slightly shorter than palm; uniform-
ly covered with squamous tubercles except
on fingers. Palm about 1.8 times long as

broad. Movable finger with squamous tu-
bercles along dorsal margin, lateral surfaces
punctate, covered with regularly spaced
tufts of setae, opposable margin finely ser-
rate, devoid of tubercles distally, with 4—5
small tubercles proximally. Surface of fixed

402

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Table 1.—Measurements (mm) of type specimens of Procambarus (Austrocambarus) citlaltepetl, new species.

Holotypic male,

form I
Total length 52.6
Carapace
Total length 2592
Postorbital length 20.6
Width 12.6
Areola
Length 8.8
Width 0.7
Rostrum
Length 6.8
Anterior width BSI
Posterior width 3.6
Acumen length 0.7
Cheliped
Length of mesial margin
of palm OZ
Width of palm 6.2
Length of lateral margin 16.7
Length of dactyl 8.7
Abdomen
Width 10.8
Length OHSS

finger punctate, devoid of tubercles except
for difuse row along inferior margin; lateral
surfaces covered with regularly spaced tufts
of setae; opposable margin finely serrate
with 5 well defined tubercles proximally
(Fig. 4D).

Carpus short, approximately conical in
shape, dorsal and lateral surfaces covered
with subsquamous tubercles, lateral surface
with shallow longitudinal groove, inferior
surface with small, scattered subsquamous
tubercles. Distal third of merus with minute
tubercles along mesial and lateral surfaces;
inferior border with 2 rows of blunt tuber-
cles, inner row with small tubercles increas-
ing in size distally, outer row with larger
tubercles ending distally in large spiny tu-
bercle.

Ischium of third pair of pereiopods with
simple acute hook, overreaching basio-
ischial articulation (Fig. 4E).

First pleopods as described in Diagnosis.

Proximal lobes of uropods with short,

Morphotypic male,

Allotypic female form II
675 61.8
S25) 30.5
26.4 24.5
16.0 14.4
WA 10.5

1.0 OW
de® 7.8
DS 2.4
510) 4.7
10 1.0
6.6 8.7
6.2 65
163 18.6
Onl 10.5
14.3 10.4
34.9 3129

acute spines. Endopodite with distomedial
spine located premarginally, and well de-
veloped distolateral spine. Exopodite with
3 spines on distolateral angle, external and
internal ones fixed, medial one largest, ar-
ticulated.

Allotypic female.—Similar to holotype,
differing in following characters: acumen
length 0.12% of RL; areola 12.2 times as
long as wide, 37.3% of TCL; chelae more
robust, 2.7 times as long as broad, fingers
longer than palm, palm about 1.5 times as
long as broad (Fig. 4F); sternal thoracic
plate tapering anteriorly, wide between
third pair of pereiopods, becoming ridge-
like by third maxillipeds. Annulus ventralis
as described in Diagnosis.

Paratypic male, form IT.—Differing from
holotype in following characters: cardiac
and gastric regions of carapace less densely
punctate, acumen length 0.12% of RL, are-
ola 15 times as long as wide; rostrum with
scattered punctations, rostral borders con-

VOLUME 112, NUMBER 2

verging anteriorly; ischium of third pair of
pereiopods with small blunt projection, not
reaching basioischial articulation.

Apical structures of first pair of pleopods
poorly developed, non corneous; mesial
process strong and rounded apically, ante-
rior margin of central projection rounded,
central projection divided into cephalic and
caudal portions by groove.

Type locality.—Rinc6n de la Doncella
Park (altitude 1400 m), Ciudad Mendoza,
Municipio de Camerino Z. Mendoza, Ve-
racruz (18°48'N, 97°10’W). Rinc6én de la
Doncella is a spring fed lake that carries
water all year.

Disposition of types.—Holotypic male
form I CNCR 17317, allotypic female
CNCR 17318, and morphotypic male form
II CNCR 17319. Paratypic males form I (8)
CNCR 2326.

Material examined.—All from Rinc6én de
la Doncella Park, Ciudad Mendoza, 1400 m
of altitude, Veracruz, México: 23 ¢ form I,
17 3 form II, 25 2 (CNCR 500), 18 May
1956, coll. A. Villalobos; 3 6 form I, II 3
form 219 (ENCR 508), 1.6 form I
(eN@iat7317), 12 (CNCR. 17318),.1.d
form II (CNCR 17319), 28 Aug 1948, coll.
A. Villalobos; 2 ¢ form I, 2 6 form II, 11
2, 20 juv. (CNCR 17021), 1 Nov 1995,
colls. J. L. Villalobos, F Alvarez, R. Ro-
bles, J. Calderén.

Etymology.—The specific epithet citlal-
tepetl is taken from “‘Citlaltepetl’’, in na-
huatl “mountain of the star’, or “Pico de
Orizaba’’, the highest mountain in Mexico,
from which the springs feeding Rinc6én de
la Doncella originate. The name is treated
as a noun in apposition.

Remarks.—Procambarus (Austrocamba-
rus) citlaltepetl, new species, is morpholog-
ically similar to P. (A.) mexicanus. The new
species can be separated from P. (A.) mex-
icanus through the following characters:
branchiostegal spine absent; gonopod with
mesial process blade-like, curved over lat-
eral portion of apex; in cephalic view, lat-
eral portion of gonopod thicker than mesial
portion; preannular plate of annulus ven-

403

tralis straight, undivided, perpendicular to
longitudinal axis of body; annulus ventralis
more rounded, mesial groove narrower and
deeper, seminal receptacle opening shorter,
postannular sclerite of annulus ventralis
oval-shaped; and epistome with well de-
fined fovea and an anterior excavation, an-
terior lobe triangular, cephalic lobe elon-
gated.

The new species is also similar to P. (A.)
veracruzanus Villalobos, 1954, P. (A.) ruth-
vent (Pearse, 1911), and P. (A.) zapoapensis
Villalobos, 1954, in the general shape of the
gonopod and annulus ventralis. However,
marked differences exist in the carapace
and chelipeds among these species.

Procambarus (Austrocambarus) mexi-
canus, as defined by Hobbs (1987), still
represents a variety of closely related forms
distributed over a distance of more than 185
km from Jalapa, Veracruz to Tuxtepec, Oa-
xaca, from the rim of the central plateau at
an altitude of 1440 m to areas in the coastal
plains near the Gulf of Mexico. A detailed
study of the populations assigned to this
species will probably result in the discovery
of additional new species. Rojas (1998) de-
scribed the high degree of morphological
variation found among these populations.

Acknowledgments

We wish to thank Sara Fuentes for taking
the SEM micrographs, Rolando Mendoza
for producing the drawings, and Citlali
Pérez and Carmen Loyola for preparing the
final prints of the SEM micrographs.

Literature Cited

Erichson, W. FE. 1846. Uebersicht der Arten der Gattung
Astacus.—Archiv fiir Naturgesichichte, Berlin
12(1):83-—106.

Hobbs, H. H., Jr. 1987. On the identity of Astacus
(Cambarus) mexicanus Erichson (1846) (Deca-
poda: Cambaridae) with the description of Pro-
cambarus olmecorum, new species, from Vera-
cruz, Mexico.—Proceedings of the Biological
Society of Washington 100:198—215.

Pearse, A. S. 1911. Report on the Crustacea collected
by the University of Michigan-Walker Expedi-
tion in the state of Veracruz, Mexico.—13" An-

404 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

nual Report of the Michigan Academy of Sci- cés nouveaux de I’ Amérique tropicale-—Revue
ences 13:108—113. et Magasin de Zoologie Pure et Appliquée, se-
Rojas, Y. 1998. Revisi6n taxonémica de ocho es- ries 2, 9:501—505.
pecies del género Procambarus (Crustacea: Villalobos, A. 1954. Estudios de los cambarinos mex-
Decapoda: Cambaridae) del centro de Vera- icanos. XII, parte 1. Revisidn de las especies
cruz, México. Tésis Professional, Facultad de afines a Procambarus mexicanus (Erichson),
Ciencias, Universidad Nacional Aut6noma de con descripcidn de nuevas formas.—Anales del
México, 158 p. Instituto de Biologia, Universidad Nacional Au-

Saussure, H. de. 1857. Diagnoses de quelques crusta- tonoma de México 25(1—2):299-379.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(2):405—409. 1999.

A new species of freshwater crab of the genus Strengeriana
Pretzmann, 1971, from Colombia
(Crustacea: Decapoda: Pseudothelphusidae)

Martha R. Campos

Universidad Nacional de Colombia, Instituto de Ciencias Naturales, Apartado Aéreo 103698,
Santa Fé de Bogota, D.C., Colombia, South America

Abstract.—A new species of the genus Strengeriana Pretzmann, 1971, is
described. With the addition of the new species, this genus now includes 15
species, distributed in the Sierra Nevada de Santa Marta, and the Western,
Central and Eastern Cordilleras of the Colombian Andes, at altitudes ranging
from 700 to 1800 m above sea level. The species are distinguished by mor-
phological characteristics of the first male gonopod.

The genus Strengeriana Pretzmann,
1971, comprises a group of small pseudo-
thelphusid crabs that inhabit mountain
streams. The discovery of a new species of
Strengeriana, described herein, raises the
number of species in the genus to 15, all of
which are distributed in the Sierra Nevada
de Santa Marta, and the Western, Central
and Eastern Cordilleras of the Colombian
Andes, at altitudes ranging from 700 to
1800 m above sea level. The systematics
and biogeography of the genus were re-
viewed by Rodriguez & Campos (1989),
Campos & Rodriguez (1993), and Campos
(1995). The new species was discovered in
the collections of Museo La Salle, Santa Fé
de Bogota, Colombia (MLS). The abbrevi-
ations cb and cl stand for carapace breadth
and carapace length, respectively.

Family Pseudothelphusidae Rathbun, 1893
Tribe Strengerianini Rodriguez, 1982
Genus Strengeriana Pretzmann, 1971

Strengeriana casallasi new species
Figs. 1-3

Holotype.—Vereda San Jer6nimo, Mun-
icipio Casabianca, Tolima Department, Co-
lombia, 1500 m alt., 1 Jun 1997, leg. R.
Casallas: 1 male cl 14.6 mm, cb 23.5 mm
(MSL 278).

Paratypes.—Same locality data as holo-
type:: | male.) cl-13.0 Gum,-cb 19.7 mm
(ICN-MHN-CR 1711), 2 males, cl 12.3,
Pt-8 mm, cb 18.5, 17:6 mm, 3 females, cl
range 11.2—12.0 mm, cb range 17.4—18.0
mm (MLS 279).

Diagnosis.—First male gonopod short,
stout; mesial lobe forming crest, with 2
spines adjacent to mesial process; mesial
process wide, lanceolate, with 4 strong
spines on mesial border, bearing 5 less
prominent spines on lateral border, and 1
bifid spine distally; cephalic lobe bearing
strong conical process, a subtriangular, al-
most rounded, process, directed latero-cau-
dally; with proximal, cephalic acute spine
fused to mesial process, but with free tip.

Description of holotype.—Carapace (Fig.
la) with wide, deep cervical groove curv-
ing posteriorly and not reaching lateral mar-
gin. Anterolateral margin with deep depres-
sion behind anteroexternal orbital angle,
followed by papillae, and second shallow
depression at level of cervical groove; rest
of margin with papillae. Postfrontal lobes
small, oval, delimited anteriorly by trans-
verse depression; median groove absent.
Surface of carapace in front of postfrontal
lobes flat, inclined anteriorly. Upper border
of front convex in dorsal view, marked with

406

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fis.

Strengeriana casallasi, new species, holotype (MLS 278), a, dorsal view of carapace and pereiopods;

b, aperture of efferent channel; c, left third maxilliped external view.

row of tubercles; lower margin visible in
dorsal view, strongly sinuous in frontal
view, and with tubercles. Surface of front
between upper and lower borders high and
vertical. Upper and lower orbital margins
each with row of tubercles. Dorsal surface
of carapace covered with small papillae;
limits between regions well demarcated.
Merus of endognath of third maxilliped reg-
ularly recurved, with shallow depression on
distal part of external margin; exognath
overreaching lateral margin of ischium of
third maxilliped (Fig. 1c). Orifice of effer-

ent branchial channel partially closed by
Spine at jugal angle and by extension of lat-
eral lobe of epistome (Fig. 1b).

First pereiopods heterochelous, right che-
liped larger than left. Merus with 3 crests:
upper crest with rows of tubercles, internal
lower crest with rows of teeth, and external
lower crest with few tubercles. Carpus with
blunt distal spine. Palms of both chelipeds
smooth, swollen. Fingers of chelae gaping
when closed, tips crossing, and with rows
of tubercles (Fig. 1a) Walking legs (second
to fifth pereiopods) slender, but not unusu-

VOLUME 112, NUMBER 2 407

Fig. 2. Strengeriana casallasi, new species, holotype (MLS 278), left first gonopod: a, whole gonopod,
caudal view; b, whole gonopod, cephalic view; c, whole gonopod, mesial view; d, whole gonopod, lateral view;
e, apex, distal view. Left second gonopod: f, whole gonopod, caudal view; g, apex distal view.

408

Icp

mep

csp

Ris. 3:

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

b

mel
Icp

cel

mcp

__ Amep

Strengeriana casallasi, new species, holotype (MLS 278), left first gonopod: A, apex distal view;

Strengeriana maniformis Campos and Rodriguez, 1993, holotype (ICN-MHN-CR 0938), left first gonopod: B,
apex distal view. Abbreviations: cel, cephalic lobe; csp, cephalic spine; Icp, lateral-cephalic process; Isp, lateral
spine; mcp, mesial-cephalic process; mel, mesial lobe; mep, mesial process.

ally elongated, those of second to fifth pe-
reiopods each with 5 rows of large spines
diminishing in size proximally, arrange-
ment of spines on dactylus of left third pe-
reiopod as follows: anterolateral and anter-
oventral rows with 5 spines plus 3 interca-
lated papillae, external row with 5 spines
plus 2 proximal papillae, posteroventral and
posterolateral rows with 4 spines.

First male gonopod short, stout; mesial
lobe forming crest, with 2 spines adjacent
to mesial process (Fig. 2c, d); mesial pro-
cess wide, lanceolate, with 4 strong spines
on mesial border, bearing 5 less prominent
spines on lateral border, and 1 bifid spine
distally (Fig. 2a, b, c, d, e); marginal lobe
simple, with long ridge on lateral surface;
mesial lobe forming long slit with cephalic
lobe where spermatic channel is located;
cephalic lobe bearing strong conical pro-
cess, (Fig. 2a, b, c, e), a subtriangular, al-
most rounded process directed latero-cau-
dally (Fig. 2a, b, e), and proximal cephalic
acute spine fused to mesial process, but

with free tip (Fig. 2b, c, d, e). Spermatic
channel with rows of dark spines, and spi-
nules on distal border of cephalic lobe.
Caudal setae strong. Cephalic distal surface
with rows of strong dark spines. Second
male gonopod with spinules on distal por-
tion, tip cup-shape (Fig. 2f, g).

Color (nomenclature after Smithe
1975).—Specimens preserved in alcohol
are dark brown (near 223A, Mars Brown),
with olive-brown specks on the dorsal side
of carapace. The walking legs are paler
brown (Verona Brown, 223B) dorsally, and
reddish-brown (Tawny, 38) ventrally. The
chelae are paler brown (Verona Brown,
223B) dorsally, and buffy-brown (Sayal
Brown, 223C) ventrally. The ventral sur-
face of the carapace is buffy-brown (Sayal
Brown, 223C).

Etymology.—The species is named in
honor of Hermano Roque Casallas, who
collected the specimens.

Remarks.—This species is similar to
Strengeriana maniformis Campos & Rod-

VOLUME 112, NUMBER 2

riguez, 1993. The two can be distinguished
by features of the first male gonopod. The
mesial process (Fig. 3a, mep) in S. casallasi
is wide, lanceolate, with four strong spines
on the mesial border, five less prominent
spines on the lateral border, one bifid spine
distally, and shows a torsion of 90° relative
to S. maniformis. The mesial process (Fig.
3b, mep) in S. maniformis is longer, oblong,
with a strong distal spine, followed proxi-
mally by a series of spines, diminishing in
size, and a slender spine (Fig. 3b, Isp) lo-
cated near the middle of the lateral surface
of the process (the last absent in S. casal-
lasi). The cephalic lobe (Fig. 3a, cel) in S.
casallasi shows a subtriangular, almost
rounded process (Fig. 3a, lcp) directed la-
tero-caudally, and a proximal acute cephalic
spine (Fig. 3a, csp) fused to the mesial pro-
cess. In S. maniformis, the subtriangular
process (Fig. 3b, lcp) on the lateral side of
the gonopod is vestigial, and the proximal
cephalic spine (Fig. 3b, csp) is well sepa-
rated from the mesial process.

Acknowledgments

I am very grateful to Dr. Rafael Lemaitre
and Dr. John Lynch for their most valuable

409

comments to this article. The illustrations
have been prepared by C. Sarmiento.

Literature Cited

Campos, M. R. 1995. A new species of freshwater crab
of the genus Strengeriana from Colombia
(Crustacea: Decapoda: Pseudothelphusidae).—
Proceedings of the Biological Society of Wash-
ington 108:98-101.

, & G. Rodriguez. 1993. Three new species of
Strengeriana from Colombia (Crustacea: De-
capoda: Pseudothelphusidae).—Proceedings of
the Biological Society of Washington 106:508—
SVL:

Pretzmann, G. 1971. Fortschritte in der Klassifizierung
der Pseudothelphusidae.—Anzeiger der Mathe-
matisch Naturwissenschaftliche Klasse der Os-
terreichischen Akademie der Wissenschaften (1)
179(1—4):14—24.

Rathbun, M. 1893. Descriptions of new species of
American freshwater crabs.—Proceedings of
the United States National Museum 16(959):
649-661, pl. 73-77.

Rodriewez, Go») 1982 6. Leswcrabésr d' cau douce
d’Amérique. Famille des Pseudothelphusi-
dae.—Faune Tropicale 22:1—223.

, & M. R. Campos. 1989. Cladistic relation-
ships of fresh-water 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.

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
112(2):410—432. 1999.

A new genus and species of South American fishes

(Teleostei: Characidae: Cheirodontinae) with a derived caudal fin,

including comments about inseminating cheirodontines

Luiz R. Malabarba and Stanley H. Weitzman

(LRM) Laborat6rio de Ictiologia, Museu de Ciéncias e Tecnologia, Pontificia Universidade

Catdélica do Rio Grande do Sul, Av. Ipiranga, 6681, CEP 90619-900, Porto Alegre, RS, Brazil,

and Departamento de Zoologia, IB, Universidade Federal do Rio Grande do Sul,
Porto Alegre, RS, Brazil;

(SHW) Division of Fishes, Department of Vertebrate Zoology, MRC-159, Smithsonian Institution,

Washington, D.C. 20560-0159, U.S.A.

Abstract.—Acinocheirodon melanogramma from tributaries of the rio Sao
Francisco and the rio Jequitinhonha, Minas Gerais, Brazil, is described as a
new genus and species of inseminating cheirodontines. The adult males are
distinguished by possession of modified thirteenth to fifteenth, ventral caudal-
fin rays bearing dorsally directed hooks surrounded by fleshy, possibly glan-
dular, structures. These structures appear similar to, but are here hypothesized
as convergent with some of the derived caudal glandular organs of certain
glandulocaudine characids and the species of the incertae sedis characid genus
Brittanichthys. Putative relationships of these taxa to the new genus are ex-
plored and rejected. The new species is referred to the Cheirodontinae on the
basis of a cladistic diagnosis of this subfamily. Hypotheses of relationships
with other inseminating cheirodontines bearing modified caudal fins are ex-
plored.

Resumo.—Acinocheirodon melanogramma, dos afluentes do rio Sao Fran-
cisco e rio Jequitinhonha, Minas Gerais, Brasil, é descrito como um novo
género e nova espécie de Cheirodontinae com insemina¢ao. Os machos adultos
sao caracterizados por possuirem o décimo terceiro ao décimo quinto raios da
nadadeira caudal modificados, curvados ventralmente, portando ganchos e te-
cidos hipertrofiados semelhantes, mas consideradas convergentes com os 6r-
gaos caudais glandulares de certos Glandulocaudinae e de Brittanichthys, um
género incertae sedis em Characidae. Relacoes hipotéticas entre estes taxons e
O novo género sao exploradas e rejeitadas. A nova espécie é referida a Chei-
rodontinae com base em uma nova diagnose cladistica da subfamilia. Sao dis-
cutidas as hipéteses de relagdes com outros Cheirodontinae inseminados e por-
tando nadadeiras caudais modificadas.

While searching for cheirodontine char-
acids in the collections of the Museu Na-
cional de Rio de Janeiro, one of us (LRM)
found seven specimens of an undescribed
small characid species from the rio Jequitai,
rio Sao Francisco drainage, Minas Gerais,
Brazil. Another specimen from the middle
portion of the rio Jequitinhonha, Minas

Gerais, Brazil was found (by SHW) in the
Museu de Zoologia da Universidade de Sao
Paulo while searching for glandulocaudine
characids. The rio Jequitinhonha is inde-
pendent of the rio Sao Francisco basin, but
is an adjacent coastal plain river of eastern
Brazil. Subsequent joint collections of Mu-
seu de Ciéncias e Tecnologia, Pontificia

VOLUME 112, NUMBER 2

Universidade Catdlica do Rio Grande do
Sul, Porto Alegre, the Academy of Natural
Sciences, Philadelphia, and the Universi-
dade Federal de Sao Carlos, Sao Paulo re-
sulted in additional specimens from both
rivers.

Some of the caudal-fin structures of this
species appear convergent with those of
certain of the glandulocaudine characids,
especially those of the tribe Corynopomini
of eastern Brazil and the rio Paraguay ba-
sin. Also, males in the new genus bear de-
curved caudal-fin rays in the ventral caudal-
fin lobe, suggesting some of the caudal-fin
modifications present in the glandulocau-
dine tribe Glandulocaudini. Our collabora-
tion regarding the description of the new
species grew out of our respective and joint
interests in the cheirodontine and glandu-
locaudine characids and the primary and
secondary sexual features found in both
groups such as their gonad modifications
associated with insemination and their cau-
dal-fin modifications associated with court-
ship.

The new species has hooks on some of
the caudal-fin rays and hypertrophied cau-
dal-skin tissue surrounding the hooks that
may be glandular. The caudal-fin hooks in
males of the new genus (Figs. 3—6) are de-
rived compared to other inseminating chei-
rodontines that have caudal-fin hooks in
that not only are they directed dorsoanter-
iorly, but the hooks are longer and more
slender and are borne by thickened ray seg-
ments. Compare these hooks with those
shown for “‘Odontostilbe”’ mitoptera (Fink
& Weitzman, 1974:fig. 3). Note that Mala-
barba et al. in Malabarba (1998) included
“O.” mitoptera in their new tribe Comp-
surini, but so far the genera of this group
have not been fully analyzed cladistically
and a meaningful generic assignment for
this species is not available. We therefore
use the most recent generic allocation from
Malabarba (1998). Because the new genus
and species is included in the cheirodontine
tribe Compsurini by several synapomor-
phies at several nodes and is therefore ex-

411

cluded from the Glandulocaudinae as well
as other characid groups bearing caudal-fin
hooks, its caudal hooks may be considered
derived independently from those excluded
taxa. The posteriorly placed caudal-fin
hooks of the new genus and species might
suggest a relationship to the species in the
characid genus Brittanichthys Géry, 1965b:
13, but this is rejected in more detail below.

Four synapomorphies were provided by
Malabarba (1994, 1998), also briefly dis-
cussed by Weitzman & Malabarba (1999),
to diagnose a monophyletic Cheirodontinae
in a more restricted and rigorous sense than
that defined by Eigenmann (1915) and dis-
cussed by Géry (1977). The new genus and
species shares all four of these synapomor-
phies and we do not consider it to be close-
ly related to the glandulocaudines, or to
Brittanichthys.

After initiating our study of this new spe-
cies, it was discovered that those cheiro-
dontine characids bearing modified hyper-
trophied caudal tissue and/or hooks on
some of the caudal-fin rays are inseminat-
ing (Burns et al. 1997) as are all glandu-
locaudines (Burns et al. 1995), and Brittan-
ichthys (see Weitzman et al. 1994, Weitz-
man & Menezes 1998). However, Burns et
al. (1998) found that the testis morphology
and the fine structure of the sperm cell nu-
clei of the glandulocaudines are uniquely
derived. Inseminating cheirodontines also
have derived sperm nuclei, but these are
differently modified from those of the glan-
dulocaudines as well as those of certain oth-
er inseminating characids (Burns et al.
1998). This suggest a hypothesis of inde-
pendent acquisition of insemination of chei-
rodontines and glandulocaudines, but the
two subfamilies could be sister taxa, simply
having their sperm cell bodies derived in
different directions from an inseminating
common ancestor. However, Malabarba
(1988) offered evidence that the Compsur-
ini which includes the new genus, is a sister
group to a taxon derived from within the
non-inseminating cheirodontine groups and
therefore the tribe is not a sister taxon to

412

the Glandulocaudinae or other inseminating
characid groups. The possible relationships
of the new genus to other inseminating
cheirodontines are briefly explored below.

Methods

Our comments on the phylogeny of the
Cheirodontinae and other characids are
based on the concepts of phylogenetic sys-
tematics as reviewed and discussed by Wi-
ley (1981). The term diagnostic character is
used to designate one or a series of syna-
pomorphies or autapomorphies that are
characteristic of a particular taxon. The
term distinguishing character is used for
one or a series of characters that help to
distinguish one taxon from others, but no
phylogenetic information is implied; see
also Weitzman et al. (1994:48). Distin-
guishing characters may or may not be apo-
morphic. No phylogenetic information is
implied because the potential comparative
derived nature of distinguishing characters
are not analyzed. The two terms, diagnostic
character(s) and distinguishing character(s),
are so defined in order to differentiate phy-
logenetic (cladistic) treatments from those
bearing no phylogenetic information.

Counts and measurements are the same
as those described by Fink & Weitzman
(1974:1—2) and Menezes & Weitzman
(1990:382—383). For counts recorded in the
descriptions of the new species, the holo-
type is listed separately and first. This is
followed in parentheses by the range, mean
(X) and total number of specimens having
that count. When the condition of the spec-
imens were such that certain characters
could be taken from only a few specimens
and reasonable descriptive statistics were
impossible, only the extremes and a state-
ment of the modal count are given, but
there will be no mode when all counts are
the same. In the Table nearly all measure-
ments other than standard length (SL) are
expressed as a percentage of SL, except
subunits of the head that are recorded as a
percentage of the head length (HL). Verte-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

bral counts include the Weberian apparatus.
The terminal half centrum, hypural bones
and associated vertebral elements, together
usually designated as PU, + U,, but not
necessarily consisting only of those verte-
bral elements, were counted as one verte-
bra. Vertebral counts and usually fin-ray
counts were taken from radiographs and
from C&S (cleared and stained) prepara-
tions prepared according to Taylor & van
Dyke (1985).

As in Menezes & Weitzman (1990:383),
geographic entities (such as rivers) and
place names (except those of countries) are
in the language of the country of origin.
Countries are in English because English
names are available for these. The word rio
(rio in Portuguese, rio in Spanish) is not
capitalized when referring to a river (thus,
for example, rio Baria in Venezuela and rio
Ita in Brazil). However, rio would be capi-
talized when referring a place name such as
a town or city. We do this in an attempt to
avoid the transferal of English style and
grammar onto foreign place names and geo-
graphical entities. Coordinates are used
whenever possible, but are used only when
we have confidence in their approximate
accuracy.

The following acronyms are used for in-
stitutions and collections: FMNH, Field
Museum of Natural History, Chicago; MCP,
Museu de Ciéncias e Tecnologia, Pontificia
Universidade Catdlica do Rio Grande do
Sul, Porto Alegre; MNRJ, Museu Nacional
do Rio de Janeiro, Rio de Janeiro; MZUSP,
Museu de Zoologia da Universidade de Sao
Paulo, Sao Paulo; UFSCar, Universidade
Federal de Sao Carlos, Sao Carlos; USNM,
former United States National Museum,
now the National Museum of Natural His-
tory, Smithsonian Institution, Washington
D.C.

In addition to HL and SL the following
abbreviations are used in the text or figures:
C&S = alizarin red s and alcian blue
stained specimens cleared with trypsin;
spm(s) = specimen(s).

VOLUME 112, NUMBER 2

Acinocheirodon, new genus

Type species.—Acinocheirodon melano-
gramma, new species.

Diagnosis.—The penultimate and ante-
penultimate ventral procurrent caudal-fin
rays of males have their tips ventrally ex-
panded, forming a small keel in the ventral
profile of the caudal fin (Figs. 1, 3-4). The
cheirodontines Serrapinnus piaba (Liitken,
1874), S. calliura (Boulenger, 1900), S.
kriegi (Schindler, 1937), and S. notomelas
(Eigenmann, 1915) also bear three ventrally
expanded posteriormost ventral procurrent
caudal-fin rays, forming a small keel in the
ventral profile of the caudal fin. Although
the same procurrent rays are involved in the
keels of those cheirodontine species re-
ferred to Serrapinnus Malabarba (1998),
and Acinocheirodon, these differ in shape
and involve different portions of the fin rays
in the two genera. The four Serrapinnus
species have an expansion along the pos-
terior half of the ray rather than at the tip
of the last three ventral procurrent caudal-
fin rays and are more closely related to
Cheirodon Girard (1855) with which they
share several synapomorphies involving the
anteriormost ventral procurrent caudal-fin
rays, and specifically to Spintherobolus Ei-
genmann (1911) by sharing several syna-
pomorphies associated with the anal fin of
males, (Malabarba 1998, Weitzman & Mal-
abarba, 1999).

The thirteenth, through the fifteenth prin-
cipal caudal-fin rays of males are ventrally
bowed, leaving wider spaces between prin-
cipal caudal-fin rays twelve to fifteen than
found between the other caudal-fin rays of
this species (Figs. 1, 3—6).

The thirteenth and fourteenth principal
caudal-fin rays of fully mature males bear
thickened ventrally expanded segments.

The fourteenth principal caudal-fin ray of
males has a dorsally extending flap along
the proximal two thirds of its length. This
does not overlap the thirteenth principal
caudal-fin ray. The tissue between the four-
teenth and fifteenth principal caudal-fin rays

413

of males has a large dorsally extending flap
that overlaps the fourteenth ray along al-
most its entire length. There is a ventral
smaller skin flap along the posterior half of
this structure that does not overlap the fif-
teenth principal caudal-fin ray ventrally
(Figs. 3, 6). Hypertropied soft tissue sur-
rounds the caudal-fin ray hooks and extends
anteriorly in diminishing amounts along fin
rays thirteen to fourteen to their bases (Figs.
3, 6). This tissue may be glandular, but this
needs histological confirmation.

The presence of anal-fin hooks posi-
tioned along the posterolateral border of the
anal-fin rays and directed to the anal-fin
base; the anal fin not pigmented along its
distal margin; the five cusped dentary teeth,
with three central cusps larger and nearly
equal in size (Fig. 7B); and the complete
lateral line; were all found ambiguous re-
garding relationships, but also distinguished
Acinocheirodon from the other members of
the Compsurini in a global parsimony anal-
ysis (Malabarba 1998).

Etymology.—tThe first component of the
name Acinocheirodon is from the Greek,
akaina = thorn or spike and refers to the
spines on the caudal-fin rays. The remain-
der of the word refers to the characid genus
Cheirodon in reference to our referral of the
new genus to the Cheirodontinae.

Acinocheirodon melanogramma,
new species
Figs. 1—7

Note: This species and genus equals “‘un-
described genus and species B”’ in Burns et
al. (1997) and “‘new genus and species B”’
in Malabarba (1998).

Type specimens.—Brazil, Minas Gerais,
rio Sao Francisco drainage.

Holotype-MNRJ 16455, male, 31.8 mm
SL, municipio de Bocaitiva, cérrego Cacho-
eira, tributary to rio Jequitai, km 413 of
highway BR 135, approximately 17°28’S,
44°02'W (Reference map: WAC—Carta
Aeronautica Mundial 3190, Goiania, 1976),
24-30 Sep 1990, D. E Moraes Jr.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Acinocheirodon melanogramma, new species, holotype, MNRJ 16455, male, 31.8 mm SL; Brazil,
Minas Gerais, municipio Bocaitiva, cérrego Cachoeira, tributary to rio Jequitai, rio Sao Francisco basin, 24—30
September 1990.

Paratypes—MNRJ 13435, 4 spms., 2 described genus and species B in Burns et

males, 27.4—28.0 mm SL, 1 female, 27.4 al. 1997); MCP 18598, 1 female, 26.3 mm
mm SL, 1 male C&S, 28.6 mm SL; MCP SL (listed as undescribed genus and species
18596, 1 male, 27.6 mm SL (listed as un-_ B in Burns et al. 1997), all collected with

Fig. 2. Acinocheirodon melanogramma, new species, paratype, MNRJ 13435, female, 27.4 mm SL; Same
locality data as in Fig. 1.

VOLUME 112, NUMBER 2

415

Fig. 3.

Acinocheirodon melanogramma, new species, paratype, MNRJ 13435, male, 28.0 mm SL; same

locality data as in Fig. 1. Caudal peduncle and fin (lateral view, left side, rays 13 and 14 bear fin hooks).

the holotype. MCP 16472, 6 spms, 2 males,
27.0—28.1 mm SL, 4 females, 26.2—30.0
mm SL; ANSP 176238, 7 spms, 3 males,
24.5—26.2 mm SL, 4 females, 21.9—25.6
mm SL, both lots collected from highway
BR 153, S of Montes Claros, municipio de
Bocaitiva, approximately 17°65'31"S,
43°49'48"W, 20 Jul 1993, R. E. Reis, S. A.
Schaefer, J. P. Silva, & E. L. Pereira.
Non-type specimens from rio Jequiti-
nhonha drainage:—MCP 19142, 16 spms,
9 males, 31.6-34.4 mm SL, 7 females,
29.1—37.3 mm SL; ANSP 173799, 16 spms,
14 males, 31.4—36.5 mm SL, 2 females,
29.6—-31.4 mm SL, both lots from rio
Jequitinhonha at Sao Pedro do Jequitinhon-
ha, Itaobim, approximately 16°30'35’S,
41°20'02’W, 20 Jan 1995, R. E. Reis, W. G.
saul; S. Schaefer & J. E P. da Silva.
MZUSP 5132, 1 spm., C&S, male, 34.4
mm SL, municipio de Medina, rio Jequiti-
nhonha at Itaobim, approximately 16°33’S,
041°30'W (Reference map: WAC—Carta

Aeronautica Mundial 3189, Belo Horizonte,
1976), 25 Jun 1966, H. Britski and N. Me-
nezes. USNM 318486, 59 spms, 4 young
males, 23.3—29.7 mm SL, 55 juveniles and
females, 17.3—24.9 mm SL, tributary of rio
Jequitinhonha known as rio ribeirao, ap-
proximately 4—5 km ESE of municipio de
Jordanic, about 15°54’S, 40°10’W, 12 Jul
1991, R. M. C. Castro & S. Jewett. MCP
18597, 2 spms, females, 30.5—37.7 mm SL
(listed as undescribed genus and species B
in Burns et al. 1997); MCP 19238, 8 spms,
4 males, 28.4—32.2 mm SL, 1 male, C&S,
33.3 mm SL, 3 females, 26.4—35.1 mm SL;
MZUSP 52434, 8 spms, 4 males, 28.3—35.7
mm SL, 4 females, 29.9—30.6 mm SL, all
three lots from rio Jequitinhonha below
mouth of rio Araguai, municipio de Itira, 10
Jan 1989, J. C. Garavello, A. S. Soares, A.
I. Alves, & J. C. Soares.

Diagnosis.—The same as for the genus.
All unambiguous characters listed in the di-
agnosis above are secondary sexual features

416

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Acinocheirodon melanogramma, new species, paratype, MNRJ 13435, cleared and stained male, 28.6
mm SL; same locality data as in Fig. 1. Caudal fin base, procurrent caudal-fin rays and hook-bearing portion of

caudal fin.

that allow only the identification of males
of A. melanogramma among cheirodonti-
nes.

Distinguishing characters.—The follow-
ing distinguishing characters allow the rec-
ognition of both male and female speci-
mens: A small, but distinctive black band
occurs along the distal tip of the first un-
branched dorsal-fin ray (Figs. 1-2) and dis-
tinguishes A. melanogramma from other
cheirodontines, except Macropsobrycon
uruguayanae Eigenmann (1915) and
Compsura heterura Eigenmann (1915). The
absence of modified caudal-fin scales and
the presence of multicuspid instead of con-
ical teeth, respectively distinguish A. me-
lanogramma from C. heterura and M. uru-
guayanae.

Description.—Morphometrics given in
Table 1. Body moderately elongate and

compressed, greatest depth at dorsal-fin or-
igin or somewhat anterior to this position if
intestinal canal full of food or female grav-
id. Predorsal profile convex to snout tip.
Profile of body from base of posterior dor-
sal-fin ray to adipose-fin origin straight or
slightly convex, slightly concave posterior
to adipose-fin origin. Ventral profile of
body convex from tip of lower jaw to pel-
vic-fin origin and moderately concave to
straight from there to anal-fin origin. Profile
along male’s anal-fin base somewhat con-
vex in anterior half and slightly concave
posteriorly. In female entire anal-fin base
relatively straight. Dorsal and ventral pro-
files of caudal peduncle somewhat concave
in both sexes. Caudal peduncle slightly lon-
ger than deep. Head small, snout short and
rounded. Mouth terminal, maxilla short and
positioned at an angle of approximately 45

VOLUME 112, NUMBER 2

417

Fig. 5. Acinocheirodon melanogramma, new species, paratype, MNRJ 13435, cleared and stained male, 28.6
mm SL; same locality data as in Fig. 1. Caudal-fin portion bearing hooks, right side, anterior is to right.

degrees relative to long axis of body. Pos-
terior end of maxilla reaching a vertical
passing through anterior border of iris.
Premaxilla (see Fig. 7A) with 6—7 large
more or less symmetrical teeth bearing 3—5
cusps. Center cusp largest, producing an
arched tooth profile. Three main cusps are
always large. When more than three cusps
present, lateral cusps very small. Small
cusps usually absent on most medial tooth
and on fifth or sixth large tooth posteriorly.
Teeth of premaxilla somewhat compressed
with teeth and all cusps in a single row.

Premaxillary teeth not compressed as much
as those of dentaries or maxillae. Maxilla
with 2, rarely 1 or 3 teeth, each usually
bearing 5 cusps. Lateral cusps, when pres-
ent, always smaller than central ones. Max-
illary teeth compressed, bearing 3 major
more or less flattened conical cusps of ap-
proximately equal size; these form a single
row on tooth’s cutting edge. Dentary teeth
7-9 in a single row, each tooth with 5
cusps. Each tooth with 3 large central
cusps: each of these compressed, in a single
row and forming sharp cutting edge. One

418

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

re sa
ln

Fig. 6. Acinocheirodon cf. melanogramma, new species, MCP 19142, male, 34.2 mm SL, Brazil, Minas
Gerais, Itaobim, rio Jequitinhonha at SAo Pedro do Jequitinhonha, 20 January 1995. Caudal-fin rays twelve (up)
through fifteen (below), other lower caudal-fin rays partially drawn. Figure shows the distribution of the hyper-
trophied soft tissue (also present in some specimens from the rio Sao Francisco) surrounding the hooks and
extended anteriorly in diminishing amounts along fin rays thirteen to fourteen to their bases (darkened areas);
the large dorsally directed slightly recurved hooks dorsal to the thirteenth and fourteenth principal caudal-fin
ray; and the large dorsally extending flap from the fourteenth and fifteenth principal caudal-fin rays of males.

very small cusp at base of larger lateral
cusps on each side of each tooth. Median
tooth largest and next 3—4 teeth gradually
diminish in size, fifth or sixth tooth abruptly
smaller and with only 3 cusps. Next pos-
terior teeth much smaller, bearing 2 cusps
and last tooth small, unicuspid and conical.

Dorsal-fin rays ii, 9 in all specimens (n
= 19). First unbranched ray about half
length of second. Dorsal-fin origin approx-
imately at midlength of body. Adipose-fin
origin a little anterior to posterior anal-fin
insertion.

Anal-fin rays iv, 19 (iii-v, 16-19, X =
17.7, n = 19; mostly 17—18; 1 specimen 16,
and 3 specimen 19). Female anal fin with
anterior 6—7 branched rays very long, form-
ing prominent anterior lobe. Rays posterior
to anterior lobe abruptly short. Distal border
of anal fin concave and then straight pos-
terior to anterior lobe. Distal border of male
anal fin decreasing in length more gradually
than in females so that lobe not as sharply
distinct from remainder of fin as in females,
but anterior anal-fin lobe of adult males
proportionally longer than in females or ju-
veniles. Anal-fin rays of males with small,

elongate, retrorse hooks present on longest
unbranched ray, and on first 7-13 (usually
8—10) branched rays. Hooks present only
on posterior branches of rays and on distal
parts of main ray, never on its proximal
portion. One pair of bony hooks per ray
segment, two pairs rarely occur on first
branched ray and posterior unbranched ray-
bearing hooks.

Pectoral-fin rays i, 10 G, 10-12, = 10.8,
n = 20). Distal ends of longest rays not
extending to pelvic-fin origin. Pelvic-fin
rays i, 7 (n = 20). Pelvic-fin origin anterior
to a vertical passing through dorsal-fin or-
igin. In females longest ray does not reach
anal-fin origin; in males longest ray reaches
just to anal-fin origin. Pelvic fins with uni-
lateral, ventromedian antrorse bony hooks;
unbranched and branched rays bearing 1, 2,
or rarely 3 slender hooks per segment along
most of their lengths. Number of segments
bearing hooks varies among males, those
males bearing more hooks appearing more
mature.

Principal caudal-fin rays 10/9. Dorsal
procurrent caudal-fin rays 13 (7-13, X =
10.1, n = 20; 7 in one specimen). Ventral

VOLUME 112, NUMBER 2 419

Fig. 7. Acinocheirodon melanogramma, new species, paratype, MNRJ 13435, cleared and stained male, 28.6

mm SL; same locality data as in Fig. 1. Jaws and teeth of right side. Anterior is to right. Top = maxilla and
premaxilla. Bottom = mandible. Note: replacement teeth visible because bone is translucent.

procurrent caudal-fin rays 11 (7-11, X = forming small ventral keel seen in ventral
8.8, n = 20). Distal tip of penultimate and profile of caudal fin; Figs. 1, 3—4. Other
antepenultimate ventral procurrent caudal- ventral procurrent caudal-fin rays not mod-
fin rays of males ventrally expanded and ified. Males with modified 13th through

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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VOLUME 112, NUMBER 2

15th principal caudal-fin rays curved, hav-
ing dorsally concave profile; rays 13th and
14th thickened and bearing dorsally (on
each side) 8 to 11 elongated and strong
hooks in 1 row; Figs. 4—6. Young or im-
mature males have 1 to 8 hooks on each of
these rays; number of hooks increases ac-
cording to size of specimen.

Scales cycloid. Lateral line usually not
complete, with perforated scales 37 (31-37,
X = 34.9, n = 4; 31 in one specimen, other
specimens 34-37), last one to three scales
of this longitudinal row of scales not per-
forated. Total number of scales in lateral
line row 37 (34-37, X = 36.5, n = 14).
Scale rows between dorsal-fin origin and
lateral line 6 (5-6, X = 5.8, n = 18), scale
rows between lateral line and pelvic-fin or-
igin 4 (3-4, X = 3.9, n = 18). Predorsal
scales 12 (11-13, X = 11.7, n = 18), usu-
ally in 1 regular series. Scale rows around
caudal peduncle 14. Males and females
with a scale sheath on anal-fin base con-
sisting of one row with 4 or 5 small scales
covering bases of unbranched rays and first
4—5 branched rays.

Supraneurals 5 (4 or 5). Precaudal ver-
tebrae, including Weberian apparatus, 17
(17-18, X = 17.1, n = 12); caudal verte-
brae, including posterior half centrum, 17
(17-19, X = 17.8, n = 12); counts taken
from x-ray negatives and one cleared and
Stained specimen.

Color in alcohol.—Both sexes with ap-
proximately same preserved color pattern;
Figs. 1-2. Body pale brownish yellow in
Specimens preserved in formalin long
enough to destroy guanine pigment. Lateral
body stripe broad and black posteriorly, be-
coming pale and narrow anteriorly and dis-
appearing anterior to dorsal-fin origin. Hu-
meral spot absent. What appears to be a hu-
meral spot in Figs. 1—2 is black interior wall
of swim bladder seen through membranous
tympanum plus a few extra dark chromato-
phores on borders of scales just anterior to
tympanum. Dark body stripe continues
broadly onto middle caudal-fin rays, but not
extending to termination of these rays. Oth-

421

erwise caudal fin white or pale brownish
yellow. Exposed borders of scales delineat-
ed by dark chromatophores, especially on
dorsal body surface where the pigment is
darker. Midline of dorsal body surface from
nape to caudal-fin origin very dark. All fins
hyaline except dorsal fin with 1 small dark
bar between first and second rays from
about distal tip of first unbranched ray and
to about midpoint of second unbranched
ray; Figs. 1-2. Head black to gray dorsally,
especially dark near nape. Sides of head
and opercles silvery where guanine pigment
not destroyed by formalin, sides of head
otherwise whitish yellow. Some dark chro-
matophores surround nares and border of
upper jaw bones. Lower jaw white, without
dark pigment except at symphisis where
few dark chromatophores border its con-
tours.

Sexual dimorphism.—The females lack
hooks in the pelvic, anal, and caudal fins,
while the mature, sexually active males
bear numerous hooks on these fins. Caudal-
fin rays thirteen and fourteen (rarely fifteen)
of males bearing hooks and hypertrophied
tissue. Large dorsal skin flaps occur along
the connective tissue between the four-
teenth and fifteenth caudal-fin rays and
along thirteenth caudal-fin ray. Females
have dorsal skin flaps along the proximal
length of fourteenth and fifteenth caudal-fin
rays, but these are small and almost imper-
ceptible. The penultimate and antepenulti-
mate ventral procurrent caudal-fin rays of
males are ventrally expanded forming a
small ventral keel on the caudal fin. These
keels are absent in females.

Distribution.—Known from the basin of
the rio S40 Francisco in Minas Gerais, Bra-
zil. A second population found in the basin
of the rio Jequitinhonha, Minas Gerais, Bra-
zil is tentatively assigned to A. melano-
gramma (see Remarks below).

Etymology.—The name melanogramma
is from the Greek, melas, = black +
gramme = line. The name refers to the dis-
tinctive black bar on the large anterior un-
branched dorsal-fin ray.

422

Remarks.—Besides the type specimens
from the rio Sao Francisco basin, on which
we based the above description, we ob-
tained additional lots of what appear to be
A. melanogramma from four localities
along the rio Jequitinhonha. Although these
specimens have the same counts and most
other characters as those from the rio Sao
Francisco, mature males of the population
samples from these two river systems can
be differentiated by their caudal peduncle
depth relative to caudal peduncle length.
Figure 8 illustrates these differences. Spec-
imens available from the rio Sao Francisco
basin seem to be fully mature, even though
they are of smaller size than the fully ma-
ture specimens from the rio Jequitinhonha,
but the sample sizes are small. We suggest
that the differences we found in caudal pe-
duncle length versus its depth might pos-
sibly be due to small sample size of fully
mature adult males and prefer to not assign
a new name for the rio Jequitinhonha pop-
ulation until further comparative material is
available. However, it seems best to refer to
the rio Jequitinhonha population sample as
Acinocheirodon cf. melanogramma. Other
data from rio Jequitinhonha samples are:
Analstntrays aii 16-2 1k = 18 Ane
49; mostly 17-19; 1 specimen 16, 2 speci-
mens 20, and 1 specimen 21. Pectoral-fin
rays 1, 9=11, X = 10.4, n = 33. Pelvie=-tins
i, 7-8, X = 7, n = 33; i, 8 in one specimen.
Dorsal procurrent caudal-fin rays 8-13, X
= 10.4, n = 46. Ventral procurrent caudal-
fin rays 7-11, X = 9.6, n = 46. Lateral line
with 34—36 perforated scales, X = 35.5, n
= 13. Total number of scales in lateral line
row 35-38, X = 36.8, n = 13. Scale rows
between dorsal-fin origin and lateral line 5—
6, X = 5.8, n = 16, scale rows between
lateral line and pelvic-fin origin 3—4, O=
3.9, n = 16. Predorsal scales 11-13, X =
11.8, n = 16, usually in regular series. Scale
rows around caudal peduncle 14. Supra-
neurals 4 or 5. Precaudal vertebrae, includ-
ing Weberian apparatus, 16-18, X = 17, n
= 66; caudal vertebrae, including posterior
half centrum, 16-19) °X%¢=> 19:5; n7?="66)-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

counts taken from x-ray negatives and two
cleared and stained specimens. For body
and head ratios expressed as percentages;
see Table 1. One male (MCP 19238, 31.9
mm SL) unusual in that 15th principal cau-
dal-fin ray has two hooks dorsally.

Discussion of possible relationships.—
The new species is placed in the Cheiro-
dontinae on the basis of a new diagnosis of
this characid subfamily provided by Mala-
barba (1994, 1998) and briefly discussed in
Weitzman & Malabarba (1999). The new
species shares all four synapomorphies pro-
posed to diagnose cheirodontines: Presence
of a large, nearly triangular hiatus of mus-
cles covering the anterior swimbladder be-
tween the first and second pleural ribs
(pseudotympanum), limited dorsally by the
lateralis superficialis muscle, posteriorly by
a naked anterior face of the second pleural
rib, postero-ventrally by the obliquus infer-
ioris muscle, and antero-ventrally by the
obliquus superioris muscle; Humeral spot
absent (Fig. 1—2); Teeth pedunculate, great-
ly expanded and compressed distally (Fig.
7); Only one tooth series present in the pre-
maxilla (Fig. 7).

The new species also displays some, we
suggest convergent, similarities with certain
members of the characid subfamily Glan-
dulocaudinae and the characid genus Brit-
tanichthys. Burns et al. (1997) found that
some cheirodontines are inseminating, as is
the case for all examined glandulocaudines
(Burns et al. 1995). We hypothesize, how-
ever, that the similarities of A. melanogram-
ma to the species of Brittanichthys and
glandulocaudines are homoplastic because
Brittanichtys lacks the synapomorphies di-
agnosing the Cheirodontinae.

Mimagoniates microlepis (Steindachner,
1876) and Mimagoniates rheocharis Me-
nezes & Weitzman (1990:389 fig. 5, and p.
401, fig. 24, p. 408 + pp. 419) also have
hook-bearing, ventrally bowed caudal-fin
rays. Menezes & Weitzman (1990) hypoth-
esized a phylogeny of the glandulocaudine
tribe Glandulocaudini that would exclude
A. melanogramma from close relationship

VOLUME 112, NUMBER 2 423

Squares, rio Jequitinhonha, males

n=20, Y = -0.0432 + 1.0048X; r? = 0.8780
Circles, rio Sao Francisco basin, males

n= 8, Y=0.3974 + 0.7201 X; r? = 0.7757

Caudal Peduncle Depth (mm)

3.0 3.5 4.0 4.5 5.0
Caudal Peduncle Length (mm)

Fig. 8. Graph of caudal peduncle depth as a function of its length for Acinocheirodon melanogramma from
rio Sao Francisco basin (circles) and of Acinocheirodon cf. melanogramma from the rio Jequitinhonha (squares).
Note that the 95% confidence intervals do not overlap with the means of the compared population samples
indicating a statistically significant difference. These differences are discussed in the text.

with the Glandulocaudini and especially phied caudal scale or scales associated with
with the derived glandulocaudin species, M. glandular tissue at the base of the tail that
microlepis and M. rheocharis. Acinochei-. is apparently present in all glandulocau-
rodon melanogramma lacks a hypertro- dines except Landonia latidens Eigenmann

424

& Henn in Eigenmann et al. (1914). Acino-
cheirodon melanogramma also lacks the
synapomorphies of the tribe Glandulocau-
dini and of the subfamily Glandulocaudinae
(Weitzman & Menezes 1998). There are
some soft structures associated with the
base of the caudal fin in A. melanogramma
that might be assumed to be an indication
of glandulocaudine relationships. These are
described as skin flaps and hypertrophied
tissues between the thirteenth to fifteenth
principal caudal-fin rays. Acinocheirodon
melanogramma apparently acquired its de-
rived caudal tissues independently of those
of the glandulocaudines because it lacks
most of the features diagnosing the Glan-
dulocaudinae and has those diagnosing the
Cheirodontinae of Malabarba (1994, 1998).

Acinocheirodon melanogramma, as well
as certain other cheirodontines (see Burns
et al. 1997), are inseminating characids as
are glandulocaudines (Burns et al. 1995).
We note, however, that glandulocaudines
have a large area of the posterior testis
modified for sperm storage, but this was not
observed among examined inseminating
cheirodontines, tribe Compsurini, including
Acinocheirodon, see Malabarba (1998).
This further suggests, among other charac-
ters respectively diagnosing the glandulo-
caudine tribes and the Cheirodontinae, that
insemination arose independently within
the Cheirodontinae and in the Glandulocau-
dinae.

The male specimens of A. melanogram-
ma have ventrally bowed caudal-fin rays
bearing dorsoposteriorly recurved hooks in
the ventral caudal-fin lobe as do males of
Brittanichthys axelrodi Géry (1965b:fig. 9—
10). The question of a relationship among
A. melanogramma and the species of Brit-
tanichthys is as problematic as the sugges-
tion of a relationship of A. melanogramma
to members of the Glandulocaudinae. Un-
fortunately the relationships of Brittani-
chthys with other characids remain uncer-
tain. Brittanichthys was referred to a sub-
tribe, the Aphyoditeini, of his tribe Chei-
rodontidi by Géry (1965a:14, 18, 20—22)

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

and to a tentative subgroup of the Cheiro-
dontinae named the Aphyoditeina by Géry
(1977:543) who spent a considerable effort
in a precladistic attempt to discern the re-
lationships of Brittanichthys based on cer-
tain of its anatomical resemblance’s to other
characid genera. Géry justifiably had many
reservations about his results, but suggested
that Brittanichthys might be related to Lep-
tobrycon Eigenmann (1915) and Thrissob-
rycon Bohlke (1953). We do not argue with
that opinion and suggest that the prelimi-
nary osteological evidence we have seen
may confirm this, but, as Géry, we believe
that the matter needs much further investi-
gation. Géry (1977:591, 594) essentially
did not alter his estimate of the relation-
ships of Brittanichthys. Géry’s Aphyoditei-
na has never been phylogenetically diag-
nosed but its included genera (Paracheiro-
don Géry (1960a), Atopomesus Myers
(1927), Oligobrycon Eigenmann (1915),
Aphyocharacidium Géry (1960b), Oxybry-
con Géry (1964), Axelrodia Géry (1965a),
Prodontocharax Eigenmann & Pearson in
Pearson (1924), Microschemobrycon Ei-
genmann (1915), Parecbasis Eigenmann
(1914), Macropsobrycon Eigenmann
(1915), Aphyodite Eigenmann (1912), Brit-
tanichthys, Leptobrycon and Thrissobry-
con) were placed in the Cheirodontinae by
Géry (1977:586—587, 590-591, 594-595)
apparently because its species have a single
row of premaxillary teeth. We note here
that Paracheirodon simulans Géry (1963)
is an exception and has two rows of pre-
maxillary teeth; see Weitzman & Fink
(1983). We cannot treat the putative
Aphyoditeini or Aphyoditeina here except
to note that only two of the genera (Ma-
cropsobrycon and Prodontocharax) placed
in this subtribe by Géry were considered
Cheirodontinae in the more restrictive sense
of Malabarba (1994). See Malabarba (1998)
for further comments on the Aphyoditeini
and the Aphyoditeina and a historical re-
view of the various treatments by Géry of
the genera he ultimately included in his
Cheirodontinae, Géry (1977).

VOLUME 112, NUMBER 2

425

Big? 9.

Brittanichthys axelrodi, USNM 221992, cleared and stained male, 22.7 mm SL; Brazil, Amazonas,

rio Negro, near mouth of rio Urubaxi. Caudal-fin base, procurrent caudal-fin rays and hook-bearing portion of

caudal fin. Anterior to right.

Géry (1965:14) also considered the pos-
sibility that Brittanichthys might be a glan-
dulocaudine. His opinion was that if it does
belong among that group (which he sug-
gested is polyphyletic) its relationships
could be remotely with the Xenurobrycon-
ini. Weitzman & Fink (1985:13, 21) noted
that although Brittanichthys shared fused
hypurals and parhypural with the xenurob-
ryconin glandulocaudines, its lack of both
a caudal gland and the modified xenurob-
ryconin arrangement of the caudal squa-
mation excluded any relationship based on
a cladistic analysis of the character distri-
butions of the taxa they studied. According
to Weitzman et al. (1994:53), however, the

fact that Brittanichthys is an inseminating
characin with a sperm storage area in the
testis raises the question again. See also
Weitzman & Menezes (1998).

Although A. melanogramma and Brittan-
ichthys axelrodi share recurved hooks on
the caudal fin and share curved ventral cau-
dal-fin rays, these structures are derived in
different ways in these two genera (com-
pare Figs. 3—6 and 9-10). Acinocheirodon
melanogramma has several soft caudal
structures that are derived states apparently
absent in Brittanichthys. The derived
curved fin ray of Brittanichthys is the
twelfth principal caudal-fin ray (Fig. 9—10)
while in A. melanogramma it is the thir-

426

~~

:_

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 10. Brittanichthys axelrodi, USNM 221992, cleared and stained male, 22.7 mm SL; same locality data
as in Fig. 9. Hook-bearing portion of caudal fin, right side, anterior to right.

teenth through the fifteenth principal cau-
dal-fin rays (Figs. 4-6). As noted in Weitz-
man & Fink (1985:13, 21) and Weitzman
et al. (1994) the parhypural and hypural are
fused in Brittanichthys. In our cleared and
stained specimens of Acinocheirodon these
bones are separate (one C&S specimen of
A. melanogramma has them so closely
placed that we can only distinguish the two
isolated elements by pressing them with a
needle where they are in contact). This re-
moves a source of possible synapomorphic
support for a relationship between these

taxa. The fact that A. melanogramma has
the cheirodontine synapomorphies and that
these are absent in Brittanichthys corrobo-
rates a hypothesis that A. melanogramma is
not closely related to Brittanichthys axel-
rodi.

Among cheirodontine genera, in the
more restricted sense of Malabarba (1994),
A. melanogramma (called “‘undescribed ge-
nus and species B’—MCP 18596, MCP
18597 and MCP 18598—by Burns et al.
1997 and ‘‘new genus and species B”’ by
Malabarba, 1998), Saccoderma melanostig-

VOLUME 112, NUMBER 2

ma Schultz (1944), Compsura heterura Ei-
genmann, Compsura gorgonae (Evermann
& Goldsborough, 1909), Macropsobrycon
uruguayanae, some species currently re-
ferred to the polyphyletic “‘Odontostilbe,”
“O.” dialeptura (Fink & Weitzman, 1974)
and ‘“‘O.” mitoptera, and an undescribed
species (called “undescribed genus and
species A’ by Burns et al., 1997 and “new
genus and species A’ by Malabarba, 1998)
were found to be inseminating. Also, all
these species have hypertrophied caudal tis-
sues and/or hooks on the caudal-fin rays,
suggesting they may represent a monophy-
letic group and were united in the cheiro-
dontine subtribe Compsurini by Malabarba
et al. in Malabarba (1998).

Homologies of the specialized structures
and putative apomorphies in the caudal fin
among inseminating cheirodontines, need
further investigation. Each of the three gen-
era listed above have apomorphies that al-
low their recognition as natural assemblag-
es among the Cheirodontinae, but possible
hypotheses of possible relationships among
them and with A. melanogramma need phy-
logenetic study.

Acinocheirodon melanogramma has
hooks in the ventral lobe caudal-fin rays as
do certain other inseminating cheirodonti-
nes. Acinocheirodon melanogramma has
large recurved hooks along the dorsolateral
surface of the dorsal branches of caudal-fin
rays 13-14, and rarely and less-developed
hooks also on ray 15. Caudal-fin hooks are
also present in Saccoderma hastata (Eigen-
mann, 1913) and are distributed on the thir-
teenth to eighteenth (usually the fourteenth
to the seventeenth) principal caudal-fin
rays. In “‘Odontostilbe”’ dialeptura caudal-
fin hooks occur on the twelfth to sixteenth,
and in Macropsobrycon uruguayanae from
the twelfth to fourteenth caudal-fin rays
(plus several spinelets along the proximal
half of the caudal-fin rays 14 to 18). See
Fink & Weitzman (1974:8, fig. 3 and 34,
fig. 26) respectively for figures of the cau-
dal fins of “O.” dialeptura and S. hastata.
These two species have one hook for each

427

ray segment except at the distal end of each
ray where the hooks are small and there
may be as many as three hooks per seg-
ment, especially in S. hastata. Macropsob-
rycon uruguayanae has one hook for each
ray segment along the distal one-third of
caudal-fin rays twelve to fourteen. The
hooks on the caudal-fin rays of all these
species are on the dorsolateral surfaces of
the rays. In Saccoderma hastata and O. di-
aleptura the hooks occur on at least two of
the branches of the larger rays (13-15). The
hooks are not large and organized like those
of A. melanogramma on thickened rays. In-
stead the rays of all examined cheirodontine
Species except A. melanogramma remain
slender in their hook-bearing areas. Further,
the hooks of the other inseminating chei-
rodontines are relatively small compared to
the hooks on the anal fin of A. melano-
gramma whose caudal-fin hooks are con-
siderably enlarged. Although the hooks are
present on the ventral lobe of the caudal fin
in all the species discussed, they do not all
occur on the same caudal-fin rays and are
of different shapes. Caudal-fin ray hooks
may be informative regarding relationships
among inseminating cheirodontines and de-
serve further investigation, but the presence
of hooks in the lower lobe of the caudal fin
itself as a synapomorphy uniting A. melan-
ogramma to those other inseminating chei-
rodontines bearing hooks is not congruent
with a cladogram (Malabarba 1998) of in-
seminating cheirodontines that results from
other characters discussed below.

Species of Saccoderma, have an enlarged
caudal-pouch scale in the ventral lobe of the
caudal fin, mainly represented by the pos-
terior ultimate scale of the scale row im-
mediately ventral to the lateral-line scale
row. This scale has a free posterior border
and covers a pouch open to the surrounding
water. While bearing hooks on the lower
lobe of the caudal fin, neither A. melano-
gramma nor Macropsobrycon uruguayanae
have modified caudal-fin scales, as de-
scribed for Saccoderma. A pouch scale, not
as well-developed as in species of Saccod-

428

erma, is also found in the lower caudal-fin
lobe of Compsura heterura. This species
bears no caudal-fin hooks, but is insemi-
nating. Compsura gorgonae is like Comp-
sura heterura, but has a less modified cau-
dal scale. Compsura heterura, Compsura
gorgonae, and Saccoderma species have
the enlarged scale connected dorsally to lig-
aments arising from the twelfth and thir-
teenth principal caudal-fin rays, and ven-
trally to a skin flap connected to the nine-
teenth principal caudal-fin ray. This sug-
gests that the derived caudal fin, its scales,
and its ligaments are homologous among
these cheirodontines. Odontostilbe dialep-
tura also has some enlarged scales on the
ventral caudal-fin lobe, but these are not as
developed as those found in the species of
Compsura and Saccoderma and are at-
tached to the fourteenth and nineteenth
principal caudal-fin rays.

Males of A. melanogramma have a large
dermal flap or fold that extends posteriorly
from between the interradialis muscles.
This flap originates between the fourteenth
and fifteenth principal caudal-fin rays and
mostly overlaps the fourteenth ray along al-
most its entire length (Fig. 6). This flap
forms the ventrolateral wall of a shallow,
elongate dorsally open pocket or trough that
lies between the flap and the body of the
fourteenth ray. There is a smaller ventral
flap, along the posterior half of this struc-
ture that does not overlap the fifteenth prin-
cipal caudal-fin ray ventrally. The four-
teenth principal caudal-fin ray also has a
dorsally directed skin flap along the proxi-
mal two-thirds of its length. This flap does
not overlap the thirteenth principal caudal-
fin ray. On the dorsal caudal-fin lobe there
are two additional flaps. These are folded
or directed ventrally. These flaps extend
about half the proximal length of the sixth
and seventh rays. Females also have dermal
flaps extending ventrally from the sixth and
seventh caudal-fin rays and dorsally from
the fourteenth and fifteenth rays, but these
are always smaller than that flaps described
for the males. These flaps do not show in

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2, being obscured by other caudal-fin
tissues. These flaps are very similar in ap-
pearance to lateral ray skin flaps sometimes
found on the dorsal, anal and also caudal-
fin rays of fishes that live in fast water
streams and fold their fins to reduce friction
of the fin while swimming. Many characids,
as for example Brycon pesu Miiller and
Troschel, have such flaps, although they are
smaller in relative size than those of A. me-
lanogramma. The large, derived flaps pre-
sent in males of Acinocheirodon constitute
one of the apomorphies of the genus (see
the Diagnosis of Acinocheirodon). None of
the other inseminating cheirodontines has
derived skin flaps as described for A. me-
lanogramma. Species of Saccoderma and
Compsura heterura also have small, slight-
ly circular skin flaps along the midlength of
the dorsal border of the principal caudal-fin
rays 16-19 and 15-19, respectively. These
flaps occur near the opening of the caudal
pouch medial to the enlarged pouch scale.
These flaps are found on different caudal-
fin rays than in Acinocheirodon, and have
distinctly different shapes. They are puta-
tively considered non homologous with the
similar structures in Acinocheirodon be-
cause they occur on different rays.

In addition, A. melanogramma has hy-
pertrophied soft tissue, especially around
the hooks of the thirteenth and fourteenth
fin rays (Fig. 3). This hypertrophied tissue
extends anteriorly in continuously reduced
amounts along these fin rays to their prox-
imal ends and also occurs distributed along
the fifteenth principal caudal-fin ray that
has no hooks.

Weitzman & Fink (1985:96—99) rather
extensively discussed caudal-fin structures
and their possible function among glandu-
locaudines, and proposed that many of the
modified structures of the male xenurobry-
conin caudal fin and the pouch scales of
other glandulocaudines may serve as a
pumping mechanism increasing the rate of
dissemination of a pheromone or phero-
mones into the surrounding water from
glandular tissue situated in and around the

VOLUME 112, NUMBER 2

opening of the pouch or sac. Almost all
glandulocaudines have a caudal pumping
mechanism, but most have a “‘passive
pump” in that there is no derived direct
caudal muscle involvement. In the Xenu-
robryconini there is direct, derived muscle
involvement, especially in Xenurobrycon
Myers & Miranda-Ribeiro (1945), Tytto-
charax Fowler (1913), and Scopaeocharax
Weitzman & Fink (1985). Although consid-
ered non homologous structures with regard
to those in glandulocaudines, the relatively
large scale structure and flaps of the insem-
inating cheirodontines could have a similar
function to that in the species of the glan-
dulocaudines. This could be true for the
various species of Saccoderma and for C.
heterura, which have developed skin flaps
near the opening of the caudal pouch scales.
However, A. melanogramma does not have
any pumping mechanism involving caudal-
fin scales. Instead, it has two large skin
flaps between the thirteenth and fifteenth
principal caudal-fin rays, just ventral to and
surrounding the hypertrophied soft tissue
found around the hooks of the thirteenth
and fourteenth fin rays. This tissue may be
glandular in nature, but we have no histo-
logical preparations of it at this time. The
courtship behavior of this species has not
been observed, but we suggest that the
large and specialized skin flaps of the cau-
dal fin in A. melanogramma males may also
be part of a mechanism for increasing the
rate of pheromone dissemination in the sur-
rounding water during courtship. Mecha-
nisms for the dissemination of pheromones
may be important for inseminating chara-
cids, both cheirodontines and glandulocau-
dines.

Caudal fin structures aside, tooth mor-
phology suggests that A. melanogramma
could be related to the genera Aphyochei-
rodon Eigenmann (1915) and Cheirodon-
tops Schultz (1944). The teeth of Acinoch-
eirodon usually have five cusps and are dif-
ferentiated according to their distribution on
the jaws (Fig. 7). The five-cuspid premax-
illary teeth are symmetrical and elongate

429

with the central cusp the largest. The an-
teromedial dentary teeth are the largest in
the lower jaw and are usually five-cuspid.
Each of the dentary teeth has the central
three cusps largest and of equal size. The
lateral cusps of each tooth are tiny and are
situated more ventrally than the three me-
dial cusps. The remaining more laterally-
placed dentary teeth, are posteriorly graded
and reduced in size, robustness, and cusp
number with the most posterior tooth often
unicuspid. These teeth are similar to those
found in Aphyocheirodon hemigrammus E1i-
genmann (1915) and Cheirodontops geayi
Schultz (1944), but similar teeth are also
found in Serrapinnus microdon (Eigen-
mann, 1915), Serrapinnus heterodon (Ei-
genmann, 1915), and Holoshesthes pequira
(Steindachner, 1882).

Serrapinnus microdon and Serrapinnus
heterodon share several synapomorphies
with Cheirodon and Spintherobolus as brief-
ly discussed by Malabarba (1998). Species
in these genera are not inseminating and
thus are not considered as closely related to
Acinocheirodon melanogramma or the oth-
er inseminating cheirodontines. See Mala-
barba (1998) for a more complete discus-
sion. Holoshesthes pequira appears more
closely related to Odontostilbe fugitiva
Cope than to any other described Cheiro-
dontinae (as hypothesized by Malabarba,
1994, 1998), and is also not inseminating
(Burns et al. 1997).

Although tooth morphology suggests
Acinocheirodon may be related to the gen-
era Aphyocheirodon and Cheirodontops, in-
semination is unknown for both later gen-
era. However, no mature specimens of these
last two species were available for exami-
nation. We are not confident about a hy-
pothesis of close relationship between Acin-
ocheirodon and the genera Aphyocheirodon
and Cheirodontops because tooth morphol-
ogy is highly variable among other chara-
cids and also among cheirodontines. The
specialized dentary teeth, with the three
central cusps about equally developed and
forming the cutting edge may have evolved

430

several times in the Cheirodontinae, for ex-
ample in H. pequira, S. microdon plus S.
heterodon, Acinocheirodon, Aphyocheiro-
don and Cheirodontops, as supported by
Malabarba’s (1998) hypothesis of relation-
ships among cheirodontines. Histological
studies of gonads and morphological stud-
ies of the caudal fin of adult and mature
specimens of both genera may provide new
and better insights about the relationships
of these cheirodontine genera.

Comparative material.—Brittanichthys
axelrodi: USNM 221992, 1 C&S spm,
male, 22.7 mm SL, Brazil, Amazonas, rio
Negro, near mouth of rio Urubaxi, Ama-
zonas, Brazil. Compsura gorgonae: MCP
11988, 3 C&S spms, MCP 11989, 7 spms,
rio Caimito at Chorrera falls, in backwater
above dam, Panama Prov., Panama. Comp-
sura heterura: MZUSP 39177, 9 spms, ri-
beirao do Gado, tributary of rio Sao Fran-
cisco, Minas Gerais, Brazil. Macropsobry-
con uruguayanae. MCP 11996, 2 C&S
spms, arroio Dom Marcos, on the road BR
290, Rio Grande do Sul, Brazil and MCP
16169, 13 spms, Antioquia, Uraba, tribu-
tary of rio Guap, rio Leon drainage, Colom-
bia. Odontostilbe dialeptura: MCP 11992,
5 C&S spms and MCP 11989, 15 spms,
creek about 4 mi up Pese Road from junc-
tion with Chitre—Divisa Road, Herrera
Province, Panama. Odontostilbe mitoptera:
MCP 14713, 2 C&S spms, rio Claro-Golfito
road, just east of Bar Roded, Costa Rica.
Saccoderma hastata: FMNH 56383 (holo-
type), Soplaviento, Colombia. Saccoderma
melanostigma: USNM 228325, 7 speci-
mens, Zulia, rio Yasa MCPO. Libertad, Es-
tado Zulia, Venezuela. Saccoderma robus-
ta: USNM 175308, 16 spms, Chibogado,
Sinu Cordoba, Colombia.

Acknowledgments

Financial aid for this research and asso-
ciated travel for museum study and field
work in Brazil were supplied for SHW and
to LRM for museum study in the United
States in part by the International Environ-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

mental Science Program of the Smithsonian
Institution, Neotropical Lowland Research
Program. This project was under the con-
secutive direction of W.R. Heyer and R.P.
Vari during the duration of this research.
Further similar support was obtained from
the following Brazilian agencies, institu-
tions and museums: Conselho Nacional de
Desenvolvimento Cientifico e Tecnolégico
(CNPq)—Proc. 451459/96-2—Brazilian
government grants; and Fundacao de Am-
paro a Pesquisa do Estado do Rio Grande
do Sul (FAPERGS)—96/0653.5—state
government grant for Malabarba’s travels to
the US; Museu de Zoologia da Universi-
dade de Sao Paulo (N.A. Menezes and PE.
Vanzolini); Laboratério de Ictiologia, Mu-
seu de Ciéncias e Tecnologia, Pontificia
Universidade Catdlica do Rio Grande do
Sul (J. Bertoletti, Director). Osvaldo Oyak-
awa (MZUSP), Paulo A. Buckup (MNRJ),
Roberto Reis (MCP), and William G. Saul
(ANSP), loaned specimens in their care or
provided cataloging services. Lisa Palmer
(USNM) and José E P. Silva provided cat-
aloging and other technical services. This
paper was improved by the comments of
several reviewers, Richard P. Vari, Jonathan
W. Armbruster, Antony S. Harold and John
G. Lundberg.

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
112(2):433—442. 1999.

Psilotris amblyrhynchus, a new seven-spined goby
(Teleostei: Gobiidae) from Belize, with notes on
settlement-stage larvae

David G. Smith and Carole C. Baldwin

Department of Vertebrate Zoology, MRC 159, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560-0159, U.S.A..,
email: smith.davidg@nmnh.si.edu; baldwin.carole@nmnh.si.edu

Abstract.—Psilotris amblyrhynchus is described from one adult and one ju-
venile collected in shallow waters in the vicinity of Carrie Bow Cay, Belize,
Central America. Settlement-stage larvae of the new species were collected in
a stationary plankton net and are used to augment the description; two of these
were reared through metamorphosis. Psilotris amblyrhynchus is distinguished
from other western Atlantic seven-spined gobies on the basis of meristic fea-
tures and pigment pattern. Psilotris is one of 15 genera assigned by Birdsong
et al. (1988) to the ““Gobiosoma group,’’ but the genus is not defined on the
basis of derived characters. Assignment of the new species to Psilotris is based
on its having separate pelvic fins, seven spines in the first dorsal fin, no scales,
and no head pores. None of these characters by itself is unique within the
Gobiosoma group to Psilotris, and the configuration of the pelvic fins is highly
variable among gobies. Our inability to diagnose Psilotris cladistically, along
with our discovery of resemblances between larvae of Psilotris amblyrhynchus
and those of another western Atlantic member of the Gobiosoma group, Nes
longus, suggest that generic concepts within the Gobiosoma group should be

reassessed.

The gobiid genus Psilotris is distin-
guished from all other western Atlantic
genera by the following combination of
characters: seven spines in the first dorsal
fin; pelvic fins separate; scales absent; head
pores absent. These characters, while of
questionable phylogenetic significance,
conveniently characterize the included spe-
cies. Five species are currently recognized:
the type species, Psilotris alepis Ginsburg,
1953; P. batrachodes Bohlke, 1963; P. cel-
sus Bohlke, 1963; P. kaufmani Greenfield
et al., 1993; and P. boehlkei Greenfield,
1993. They are distinguished from one an-
other by color pattern, fin-ray counts (Table
2), and a few proportional measurements.
The specimens described here possess the
four distinguishing features of Psilotris, but

they cannot be assigned to any of the five
recognized species.

Methods

Counts and measurements follow Hubbs
& Lagler (1958). Institutional abbreviations
follow Leviton et al. (1985). The dorsal-fin
formula was proposed by Birdsong et al.
(1988) to describe the relationship between
the dorsal-fin pterygiophores and the neural
interspaces of the vertebral column. The
first digit represents the neural interspace
into which the anteriormost pterygiophore
inserts. The digits following the hyphen de-
note the number of pterygiophores that in-
sert into each subsequent interspace. The
holotype was x-rayed to obtain vertebral

434

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1.

and fin-ray counts. For larvae, these counts
were obtained by clearing several speci-
mens in trypsin and counterstaining them
with alizarin red and alcian blue. Adult and
juvenile specimens of the new species were
collected with rotenone or quinaldine. Lar-
vae were collected in a 1.0 by 0.5-m rect-
angular plankton net with 505 ym mesh,
fished passively over the reef flat by sus-
pending it from a pier on the seaward side
of Carrie Bow Cay, Belize. Selected cap-
tured larvae were anesthetized with MS-
222, examined under a dissecting micro-
scope, then reared in small, nylon-mesh
breeder traps set in a raceway in a flow-
through seawater system. Reared larvae
were fed cultured brine shrimp.

Psilotris amblyrhynchus new species
Figs. 1, 2, 3A, B

Holotype-—USNM 321019 (34.0 mm
SL), Belize, western Caribbean, between
Carrie Bow Cay and Twin Cays, lagoon—
‘shot hole,’ 15—20 ft (4.6—6.1 m), 17 Sep
1990, G. D. Johnson, et al.

Other specimens.—USNM 347250 (1,

Psilotris amblyrhynchus, holotype, USNM 321019, 34.0 mm SL.

14.2), Caribbean, Belize, shallow spur and
groove just east of Carrie Bow Cay, ca. 20
ft (6.1 m), 9 Sep 1997, C. C. Baldwin and
D. G. Smith. Larvae: all collected in a sta-
tionary plankton net on seaward side of
Carrie Bow Cay, Belize; USNM 350086 (1,
1f75), 10 Jun 1993, 2130 hrvsnmy
350087 (@; 18:5), 13 Jun 1193, Zi3Geae
cleared and stained; USNM 350088 (1,
13.0), 14 Jun 1993, 2330 he VSN
350089 (1, 12.5), 20 Jun 1993, 2100—2330
hr; USNM 350090 (1, 11.5), 20 Jun 1993,
2130 hr; USNM 350091 (1, 11.0), 21 Jun
1993, 2300 hr; USNM 350092 (1, 14.0),
19—22 Jun 1993, reared; USNM 350093 (1,
16.8), 28 Jul 1994, reared; USNM 350094
(2, 10.5—11.5), 8 Aug 1994, cleared and
stained.

Comparative material.—Larvae of Nes
longus, all collected in a stationary plank-
ton net on seaward side of Carrie Bow Cay,
Belize: USNM 350189 (1, 11.0), 23 Jul
1992, 2230-2300 hr; USNM 350190 (,
10.0), 27 Jul 1992, 2000—2030 hr; USNM
350191 (1, 11.0), 27 Jul 1992, 2230n2300
hr; USNM 350192 (3, 9.5), 16 Jun 1993,

WA —— a Ss
i Ez

——

=] SSS
— —

Fig. 2. Psilotris amblyrhynchus, larva, USNM 350088, 13.0 mm SL.

VOLUME 112, NUMBER 2 435

Fig. 3. Pelvic fins: development from larva to juvenile. A. Psilotris amblyrhynchus, USNM 350088, larva,
13.0 mm SL. B. Psilotris amblyrhynchus, USNM 347250, juvenile, 14.2 mm SL. C. Nes longus, USNM 350194,
larva, 10.5 mm SL. D. Nes longus, USNM 350197, reared juvenile, 16.5 mm SL.

436

2030 hr; USNM 350193 (4, 9.0—11.0), 20
Jun 1993, 2000—2030 hr, three specimens
cleared and stained; USNM 350194 (1,
10.5), 2 Sep 1996; USNM 350195 (1, 10.5),
5 Sep 1996; USNM 350196 (7, 9.5—17.0),
28 Jul 1994, reared; USNM 350197 (5,
15.0—19.3), Sep 1997, reared: USNM
590198 Coy Ss) 12 7han 1993.5 2100-hr,
cleared and stained. Psilotris alepis: USNM
123231 (holotype, 13 mm SL), St. Croix,
Virgin Islands, 8 Apr 1937. Psilotris ba-
trachodes: USNM 274946 (1, 13.0), Carrie
Bow Cay, Belize, depth 12—25 ft (6-8 m),
10 Jun 1981; USNM 317025 (1, 17.8), Ca-
ribbean, Tobago, Sisters Rocks, depth to 60
ft (18 m), 14 Sep 1990; USNM 317026 (3,
7.0—8.5), Caribbean, Tobago, windward
side of Little Tobago Is., depth 60 ft (18
m), 7 Sep 1990; USNM 317480 (1; 15-5);
patch reef south of Carrie Bow Cay, Belize,
depth 20-—30 ft (6—9 m), 26 Mar 1987. Psi-
lotris boehlkei: ANSP 124619 (3 paratypes,
17.5—18.2), St. Barthelmy, West Indies, Port
de Gustavia, off rocky cliffs just S. of Anse
Galet, depth 60—65 ft (10.3—19.8 m), 14 Jul
1965. Psilotris celsus: ANSP 133274 (4,
20.0—26.0), Bermuda, between Sinky and
Cross Bays, off Sonesta Beach, 9.1—10.6 m,
30 Jul 1975. Psilotris kaufmani: ANSP
131712 (2 paratypes, 15.0—16.5), Jamaica,
Discovery Bay, depth 130 ft (40 m), 16 Jun
1974. Gobulus myersi: USNM 347348 (3,
22-25), Belize, Pelican Cays, Manatee Cay,
depth 1-3 ft (0.3—1 m), 19 Oct 1997).

Diagnosis.—A _ species of Psilotris, as
currently construed, with 11—12 elements in
the second dorsal fin; 9—11 anal-fin rays;
pale ground color with a series of dusky
blotches on side of body; small, distinct,
dark spots on dorsal-fin rays; three narrow,
oblique bars on caudal fin; first dorsal spine
elongate in adults.

Description of holotype (Fig. 1).—A
moderately elongate goby without scales or
head pores. Snout short and blunt, anterior
profile of head steep, no rostral frenum. Eye
large, its diameter greater than snout length;
interorbital space very narrow. Both nostrils
tubular, the anterior one longer. Posterior

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

end of jaw below posterior half of eye.
Teeth slender, conical, slightly recurved,
widely spaced, uniserial. Gill opening re-
stricted, not extending past base of pectoral
fin. Second dorsal and anal fins relatively
long, the base of each longer than caudal
peduncle. Anteriormost spine of first dorsal
fin greatly elongate, reaching middle of
base of second dorsal fin when appressed.
Caudal fin elliptical, middle rays much lon-
ger than those above and below. Pelvic fins
almost completely separate, only a small
basal membrane connecting the inner rays
of the two sides (Fig. 3B).

Measurements in mm, with proportions
in parentheses: SL 34.0, preanal length 19.7
(58% SL), head length 10.0 29% SL),
snout length 2.2 (22% head length), eye di-
ameter 2.5 (25% head length), interorbital
width 0.3 (3% SL), upper-jaw length 4.8
(48% head length), depth at dorsal-fin ori-
gin 5.1 (15% SL), depth at anus 4.5 (13%
SL), least depth of caudal peduncle 3.5
(10% SL), length of base of second dorsal
fin 8.6 (15% SL), posterior end of dorsal
fin to origin of caudal fin 3.4% 10% SL,
base of anal fin 6.7 (20% SL), pectoral-fin
length 8.6 (25% SL), caudal-fin length 9.7
(29% SL), pelvic-fin length 5.8 (17% SL).
Meristic features: dorsal-fin rays VII+12,
anal-fin rays 11, pectoral-fin rays 17, bran-
chiostegal rays 5, vertebrae 11 + 16 = 27,
dorsal-fin formula 3-221110.

Color in alcohol: Ground color pale yel-
lowish brown, with dusky markings as fol-
lows: diffuse dark spot on cheek below eye;
dark oblique streak from just behind oper-
cle to dorsum directly above pectoral base
and before dorsal origin; a diffuse spot on
pectoral base; approximately 16 diffuse
dark saddles, irregular and uneven in inten-
sity, from nape to base of caudal fin; ap-
proximately seven diffuse midlateral
blotches on body between pectoral fin and
base of caudal fin, with smaller blotches in
between; discrete, small, dark spots on dor-
sal-fin spines and rays; three oblique dark
bars on caudal fin, oriented anterodorsal to
posteroventral.

VOLUME 112, NUMBER 2

437

Table 1.—Meristic characters for specimens of Psilotris amblyrhynchus. * = cleared and stained. * = x-rayed.

USNM SL in mm Dorsal rays Anal rays Pectoral rays Vertebrae
321019* 34.0 WAL 12 LA 17 11 + 16 = 27
347250 14.2 Viti it 10 — —
350092 14.0 WEE 9 17 —
350093 16.8 MIF "it 10 -- =
350087° cD Vike 12 11 19 i 16327
350087° Piss WEE + h2 ial 19 11 +16 = 27
350094° 11.0 Vii ila 18 11 + 16 = 27
350094° 115 Vik +, 12 10 18 Lies 16i—.27

'Rleven rays but 12 pterygiophores.

Other specimens.—Only one adult spec-
imen, the holotype, is known, and the de-
scription is based on it. We have also col-
lected 12 immature specimens, including a
small, field-caught juvenile and 11 larvae,
two of which were reared through transfor-
mation. None of these specimens has the
full complement of adult characters, and we
therefore do not designate them as para-
types. Nevertheless, they do provide addi-
tional meristic data that help characterize
the species. Four of the larvae were cleared
and stained, and their counts are presented
in Table 1, along with counts from the field-
caught juvenile and the two transformed
larvae. Of the eight specimens for which
counts are available, all have seven spines
in the first dorsal fin. Four specimens have
12 elements in the second dorsal fin, and
four have 11. In one of the latter, however,
the penultimate pterygiophore does not sup-
port a ray; the specimen thus has 12 pter-
ygiophores but only 11 rays. Fin rays can
be difficult to count in small specimens, es-
pecially when the fin is depressed, as it is
in most of the specimens. The only way to
obtain unambiguous counts is to x-ray the
specimen (if it is large enough) or to clear
and stain it. Of the five specimens subjected
to one of these techniques, four have 12
second-dorsal elements, and the fifth has 11
rays but 12 pterygiophores. Four of these
five also have 11 anal-fin rays. The three
specimens that were simply counted by
gross examination have 11 second dorsal-
fin elements and 9-10 anal rays. The pec-

toral-ray counts obtained were also higher
in the cleared and stained specimens (18—
19) than in the alcohol specimens (17), sug-
gesting to us that we may be undercounting
the fin rays in gross examination.

The field-caught juvenile (USNM
347250, 14.2 mm SL) was photographed
shortly after capture and had the following
color pattern: a few melanophores on side
of opercle and on base of pectoral fin; one
melanophore immediately above opercle on
right side; one or two melanophores behind
eye and on top of head above brain; a single
small melanophore on ventral midline just
behind anal fin; a small melanophore on
fifth dorsal spine; a few small melano-
phores on snout and upper jaw; three indis-
tinct yellow stripes on head, originating
from upper, middle, and lower margin of
eye and extending approximately to poste-
rior end of opercle; a few indistinct yellow
spots along base of dorsal fin.

Development.—The settlement-stage lar-
vae (Fig. 2) are characterized by a large,
very conspicuous, expanded melanophore
midventrally over the posterior end of the
anal fin. There is very little other pigment,
only some internal melanophores on the
dorsal surface of the gas bladder and at the
posterior end of the intestine, and a dark
midventral streak immediately before the
pelvic fin. The caudal fin is truncate rather
than elliptical. The pelvic fins are largely
united, the rays of the two sides nearly par-
allel rather than sharply diverging (Fig.
3A). The membrane connecting the inner-

438

Table 2.—Meristic characters in species of Psilotris.

Species Dorsal rays
P. alepis Vit 10
P. batrachodes VII + 9-10
P. boehlkei VII + 10-11
P. celsus VII + 10-11
P. kaufmani Vit + 10-11
P. amblyrhynchus Vil + 11-12

most rays of the two sides extends approx-
imately % to % of the way from the base to
the tip. There is no frenum connecting the
spines, however. The largest of the larvae
collected in the plankton net are about 11.5
mm SL. The 14.0-mm reared specimen still
has the conspicuous ventral melanophore,
but it is smaller relative to the size of the
body. The pelvic fins appear more diver-
gent, and the membrane connecting the me-
dial rays is greatly reduced, extending not
more than about an eighth of the distance
from the base of the rays to their tips. The
caudal fin is more elongate, but the middle
rays are not greatly prolonged. The 16.8-
mm reared specimen retains the midventral
melanophore, but it is even less conspicu-
ous than in the 14.0-mm specimen. A few
melanophores are present along the base of
the dorsal fin, but no other pigment is vis-
ible. The pelvic fins are almost completely
separate, and they diverge strongly. The
membrane connecting the medial rays is re-
stricted to the base of the rays. The caudal
fin is rounded, although the elongation of
the middle rays is still not extreme. The first
dorsal spine is not elongate. The field-
caught juvenile, although smaller (14.2 mm
SL) than the larger of the two reared spec-
imens, is slightly more developed. The ven-
tral melanophore is still present but is small
and contracted. Definitive pigment as de-
scribed above is beginning to appear. The
pelvic fins are separate and diverge strongly
(Fig. 3B). The caudal fin has still not at-
tained the elongation seen in the holotype,
and the first dorsal spine is not elongate.
Distribution and habitat.—Psilotris am-
blyrhynchus is currently known only from

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Anal rays Pectoral rays
9 LS
7-8 16
10 16-18
9-11 16-17
10-11 16-19
9-11] 17-19

the vicinity of Carrie Bow Cay on the bar-
rier reef in Belize, western Caribbean
(16°48'N, 88°05'’W). The holotype was col-
lected between Carrie Bow Cay and Twin
Cays in one of a series of “‘shot holes”’ cre-
ated by seismic sounding. The habitat was
described as an open sand patch in beds of
turtle grass (Thalassia testudineum) at a
depth of 15—20 feet (4.5—6.1 m). The field-
caught juvenile was taken from the shallow
spur and groove reef complex just seaward
of Carrie Bow Cay at a depth of approxi-
mately 20 feet (6.1 m).

Etymology.—From the Greek amblys,
blunt, obtuse; and rhynchos, snout, muzzle.
In reference to the blunt anterior profile of
the head. To be treated as a noun in appo-
sition.

Key to the species of Psilotris
(modified from Greenfield, 1993)

la. Pectoral fin with dark brown to
black bar running dorsoventrally at
a posterior angle across fin; dorsal
and caudal fins also with dark bars;
anal-fin elements 7 or 8 (usually 7);
second dorsal-fin elements 9-10
(nearly always 9) .... P. batrachodes
Pectoral fins lacking bar running
dorsoventrally at a posterior angle
across fin, fin is bicolored (upper
half black) or unpigmented; anal-

1b.

fin elements’ 8=1'1 "422 3) eee 2
2a. Pectoral-fin rays 15; anal-fin ele-

ments: 89.) Aes eee P. alepis
2b. Pectoral-fin rays 16—19; anal-fin el-

ements'9=110. S722.) eae 3

3a. Caudal fin with 3 oblique, dark

VOLUME 112, NUMBER 2

bars; snout very short and blunt,
with steep anterior profile; second
dorsal-fin elements 11—12
a ns de oy ogi P. amblyrhynchus
3b. Caudal fin without 3 oblique, dark
bars; snout more acute, with flatter
anterior profile; second dorsal-fin
elements 10—11
4a. Pectoral fin bicolored, dark brown
to black on upper 9-11 rays and
membranes and white below; anal-
fin elements 10—11 (usually 11) ..
P. kaufmani
4b. Pectoral fin not bicolored; anal-fin
elements 9—11 (usually 10)
5a. Posterior end of jaw extending past
posterior margin of pupil; caudal
peduncle slender (80-89 thou-
sandths of SL); snout short (44—55
thousandths of SL) P. boehlkei
5b. Posterior end of jaw not extending
past posterior margin of pupil; cau-
dal peduncle deeper (greater than
92 thousandths of SL); snout lon-
ger (greater than 55 thousandths of
SL) P. celsus

Comparison and Relationships

‘“Anyone who has attempted to identify
the naked or partially scaled Atlantic gobies
in and around the genera Gobiosoma and
Garmannia will appreciate the difficulty ex-
perienced in placing even generically any
specimen.’ With those words, Bohlke &
Robins (1968:47) began their study of New
World seven-spined gobies, which remains
the most comprehensive treatment of the
group. They included 17 genera in their key
(Bohlke & Robins 1968:51): Psilotris, Var-
icus, Chriolepis, Gobulus, Nes, Pycnomma,
Gymneleotris, Eleotricus, Risor, Parrella,
Barbulifer, Enypnias, Bollmannia, Aruma,
Microgobius, Ginsburgellus, and Gobioso-
ma. Four other genera were subsumed as
subgenera under an expanded concept of
Gobiosoma: Austrogobius, Elacatinus,
Garmannia, and Tigrigobius. Bohlke &
Robins (1969) added two more genera, Ev-

439

ermannichthys and Pariah. They excluded
Parrella, Microgobius, and Bollmannia
from their analysis of relationships because
*‘Parrella is a composite and the other two
are only distantly related’’ (BOhlke & Rob-
ins 1968:146). The resulting diagram (Fig.
4) expressed Bohlke & Robins’s concept of
relationships among the 16 genera and five
subgenera. This is not a classification based
on cladistic methods, which had not yet
come into wide use, and the genus Gobio-
soma as construed by Bohlke & Robins is
a classic paraphyletic group. Nevertheless,
by resurrecting Austrogobius, Elacatinus,
Garmannia, and Tigrigobius, one can ex-
tract a generic phylogeny from the diagram,
even though the characters on which it is
based are not shown, and this phylogeny
can serve as a testable hypothesis.
Birdsong (1975:180), although still not
using phylogenetic methods, considered the
American seven-spined gobies to be a “‘nat-
ural assemblage,’ based largely on verte-
bral number (11 + 16—17) and dorsal-fin
formula (3-221110), characters that he ob-
served to be “‘extremely stable’. . . with
many groups of gobioids.’’ He placed the
17 genera from BohlKe & Robins’s (1968:
51) key, plus Evermannichthys, Pariah, and
Palatogobius, in the tribe Gobiosomini
(Birdsong 1975:182, properly spelled Go-
biosomatini). He noted that the “‘Gobioso-
ma group” is united by a distinctive char-
acter, the fusion of hypurals 1—2 with hy-
purals 3—4 and the urostyle, but he did not
provide a list of genera in that group. Bird-
song stated only that he found this character
state in four subgenera of Gobiosoma as
well as in nine genera presumably derived
from Gobiosoma. He further noted that Mi-
crogobius, Bollmannia, Parrella, and Pal-
atogobius do not show this fused caudal
condition and thus are excluded from the
Gobiosoma lineage. Birdsong et al. (1988:
189) treated the ““Gobiosoma group” as
comprising 17 genera: Birdsong’s (1975:
182) Gobiosomatini minus the “‘Microgo-
bius group,’ and with the questionable ad-

440

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

CHRIOLEPIS
VARICUS
EVERMANNICHTHYS \ PSILOTRIS
\. PARIAH pS GOBULUS
\ RISOR
\ Be. BARBULIFER
keeoay, | a ENYPNIAS
eae ius
GINSBURGELLUS \ 2 eee ST
we Tigrigobius EN ae a GYMNELEOTRIS
oe \ d ‘fini PY CNOMRIN
; 7.
be \ \ Austrogobius Zo am ELEOTRICA
/ An Yj \
J NG : | Vi j ———— ARUMA
/ 3 wg
/ a SX Gobiosoma Yo :
i 3 \ / Wi \
Ww pp ;
: Elacatinus NN \ V f Li ;
: \ \ I ZZ
\ - i / SF |
\ \ / | Uf |
ee) FO ~ GOBIOSOMA
A
Y/ a, ' \\

Fig. 4. Diagram of relationships (from Bohlke & Robins 1969).

dition of Ophiogobius, an eastern Pacific
genus.

Psilotris is a member of the Gobioso-
matini of Birdsong (1975) based on the
presence of seven dorsal spines, 11 + 16—
17 vertebrae, and a dorsal-fin formula of 3-
221110. It further belongs to the “‘Gobio-
soma group”’ by virtue of its fused hypural
elements. In BoOhlke & Robins’s (1968:51)
key to genera, Psilotris falls out in the first
couplet along with Varicus, Chriolepis,
Gobulus, and Nes, all characterized by the
absence of head pores. Within this group,
the genera are separated by the presence or
absence of scales and by the form of the
pelvic fins, either united or separate. Bohlke
& Robins made no claim that their key is
anything more than a convenient way to
identify specimens. Nevertheless, it is in-
triguing that four of the five genera are ad-
jacent on Bohlke & Robins’s tree (Fig. 4).
The lone outlier is Nes, which arises from
near the base of another branch. Nes, how-
ever, is of special interest to us, as we have
collected larvae of Nes longus and reared
them through transformation. Larvae of Nes
longus and Psilotris amblyrhynchus are

strikingly similar, both having a very large
melanophore on the ventral midline at the
posterior base of the anal fin. Indeed, at first
we did not separate larvae of the two spe-
cies. It was only after our reared specimens
transformed into two distinct species that
Wwe reexamined the larvae and recognized
the difference. Larvae of Nes longus have
a second midventral melanophore at the an-
terior base of the anal fin, in addition to the
large one at the posterior end.

The most obvious difference between
adults of Nes longus and Psilotris ambly-
rhynchus is the form of the pelvic fins.
Throughout the ontogeny of Nes, the pelvic
fins of the two sides are completely fused,
and there is a well developed frenum be-
tween the spines (Fig. 3C, D). In juvenile
and adult Psilotris amblyrhynchus the pel-
vic fins are almost entirely separate, with
only a rudimentary membrane between the
bases of the inner rays (Fig. 3B). As dis-
tinctive as this character may seem, it is
subject to great variation among gobies,
even among closely related species. Within
the genus Coryphopterus, for example, the
pelvic-fin condition spans the complete

VOLUME 112, NUMBER 2

spectrum from fully fused to separate. In
larvae of Psilotris amblyrhynchus, the pel-
vic fins are largely united (Fig. 3A), the in-
terradial membrane retreating during trans-
formation (Fig. 3B); this suggests that the
separate pelvic fins are secondarily derived.
Although the character is often treated di-
chotomously (united vs. separate) in keys
such as Bohlke & Robins’s, the condition
is not so clear-cut. Gobulus, for instance, is
keyed out by choosing “‘ventral fin united,”
yet Gobulus has no interspinal frenum, and
the membrane between the inner rays of the
two sides extends no more than a third of
the way from the base of the ray to its tip.
This condition is actually closer to that in
Psilotris than in Nes. With only a slightly
greater reduction of the interradial mem-
brane in Gobulus, the distinction between it
and Psilotris would disappear.

The other distinctions between Nes lon-
gus and Psilotris amblyrhynchus are the
greater elongation of the body with age in
Nes and the presence of an extra vertebra
(28 rather than 27). Resemblances between
the two species are the steep anterior profile
of the head, with its large eye and short
snout, and the elongate first dorsal spine in
adults.

There is no unique feature that distin-
guishes Psilotris from the other members of
the ““Gobiosoma group.”’ It is characterized
only by the combination of three character
states: no scales, no head pores, and sepa-
rate pelvic fins. Each of these characters
shows a wide range of conditions. For ex-
ample, some genera are completely scaled
from head to tail, some are partially scaled,
some have only two small scales at the base
of the caudal fin, and some have no scales
at all. The number of pores also varies;
sometimes within a single species certain
pores can be present or absent. The vari-
ability in pelvic-fin condition has already
been discussed. The trends within the evo-
lution of these characters cut in different
ways across genera. Greenfield (1993:773)
correctly pointed out that Psilotris is not de-
fined by any shared derived characters, and

44]

the species are not very similar in general
appearance. Psilotris alepis (the type spe-
cies) and P. batrachodes seem most differ-
ent from P. amblyrhynchus. They are both
small, relatively stubby gobies with low fin-
ray counts. Neither has an elongate dorsal
spine, and the anterior head profile is dif-
ferent. Psilotris celsus has two elongate
dorsal spines rather than one as in P. am-
blyrhynchus. This species is also relatively
stubby, and the eye and mouth are smaller
than those of P. amblyrhynchus. The teeth
of P. alepis, P. batrachodes, and P. celsus
are described as being in several to many
rows on the jaws, as opposed to the widely
spaced, uniserial pattern found in P. am-
blyrhynchus. Psilotris kaufmani also has the
first two dorsal spines elongate (based on
fig. 1 in Greenfield et al. 1993, though it
was not mentioned in the text). The color
pattern of P. kaufmani is quite different
from that of P. amblyrhynchus, it has a
much shallower snout profile, it has fewer
second-dorsal-fin elements (10—11 vs. 11—
12), and it inhabits deeper areas of reefs and
dropoffs. Psilotris boehlkei lacks elongate
dorsal spines (again based on the figure in
Greenfield, 1993, as the character was not
mentioned in the text), and it has fewer sec-
ond dorsal-fin elements (10-11).

Our assignment of the new species to
Psilotris must be considered tentative pend-
ing further study of generic concepts among
seven-spined gobies. Although larval mor-
phology suggests a possible relationship be-
tween Psilotris amblyrhynchus and Nes lon-
gus, little comparative information is avail-
able on larval morphology of other species
of the ““Gobiosoma group.”

Acknowledgments

Our studies at Carrie Bow Cay were sup-
ported by grants from the Caribbean Coral
Reef Ecosystems program, K. Ruetzler, di-
recton..M.'R: Carpenter, KK. CColevand.c.
E. Thacker assisted in various aspects of
our field work. B. Pfeiffer and M. Lara as-
sisted us in rearing larvae. Finally, we wish

442

to thank G. D. Johnson for his support and
encouragement of our work at Carrie Bow
Cay. This is Contribution No. 561 from the
Caribbean Coral Reef Ecosystems program.

Literature Cited

Birdsong, R. S. 1975. The osteology of Microgobius
signatus Poey (Pisces: Gobiidae), with com-
ments on other gobiid fishes.—Bulletin of the
Florida State Museum Biological Sciences 19:
135-187.

, E. O. Murdy, & FE L. Pezold. 1988. A study
of the vertebral column and median fin osteol-
ogy in gobioid fishes with comments on gobioid
relationships.—Bulletin of Marine Science 42:
174-214.

Bohlke, J. E. 1963. The species of the west Atlantic
gobioid fish genus Psilotris.—Notulae Naturae
(362):1—-10.

, & C. R. Robins. 1968. Western Atlantic sev-

en-spined gobies, with descriptions of ten new

species and a new genus, and comments on Pa-
cific relatives—Proceedings of the Academy of

Natural Sciences of Philadelphia, 120:45—-174.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

44 1969. Western Atlantic sponge-
dwelling gobies of the genus Evermannichthys:
their taxonomy, habits and relationships.—Pro-
ceedings of the Academy of Natural Sciences
of Philadelphia 121:1—24.

Ginsburg, I. 1953. Ten new gobioid fishes in the Unit-
ed States National Museum, including additions
to a revision of Gobionellus—Journal of the
Washington Academy of Science 43:18—26.

Greenfield, D. W. 1993. New goby, Psilotris boehlkei
(Pisces: Gobiidae), from the western Atlantic,
with a key to the species —Copeia 1993:771—
Tafa):

, L. T. Findley, & R. K. Johnson. 1993. Psilo-
tris kaufmani n. sp. (Pisces: Gobiidae), a fourth
western Atlantic species of Psilotris——Copeia,
1993:183-186.

Hubbs, C. L., & K. EF Lagler. 1958. Fishes of the Great
Lakes Region.—Bulletin of the Cranbrook In-
stitute of Science 26:213 pp.

Leviton, A. E., R: H. Gibbs, Jn, Es Heal é21@E
Dawson. 1985. Standards in herpetology and
ichthyology, part I. Standard symbolic codes for
institutional resource collections in herpetology
and ichthyology.—Copeia 1985:802—832.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(2):443—450. 1999.

Diagnoses of hybrid hummingbirds (Aves: Trochilidae).
7. Probable parentage of Calliphlox iridescens Gould, 1860

Gary R. Graves

Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560, U.S.A.

Abstract.—Calliphlox iridescens Gould, 1860 is hypothesized to be a hybrid
between Calliphlox amethystina and Chlorostilbon aureoventris. The hybrid,
collected at Nova Friburgo, Rio de Janeiro, Brazil, exhibits a blended mosaic
of plumage characters of the presumed parental species. External measurements
of the hybrid fall between the character means of the parental species and
approach the values expected from least squares regression of parental mea-

surements.

The miniature woodstar, Calliphlox iri-
descens Gould, 1860, was described from a
unique specimen collected at Nova Fribur-
go, about 100 km northeast of Rio de Ja-
neiro, Brazil. Gould (1860:310) observed.

“If, as I believe, I am right in referring this little
bird to the genus Calliphlox, it is one of the most
remarkable Humming-birds that it has fallen to my
lot to describe. In its size and form it is very similar
to C. amethystina, but in colouring it is like a Chlo-
rostilbon.”’

The singular appearance of the specimen
prompted Gould (1861:plate 359) to make
it the type of a new genus, Smaragdochry-
sis, which was adopted by Elliot (1878) and
Salvin (1892). The taxonomic validity of ir-
idescens was not questioned until Butler
(1931:347) remarked in a brief note:

“May I regard my belief that the little Humming-
bird which Gould described (P.Z.S. 1860, p. 310) as
Calliphlox? iridescens . . . is really a hybrid be-
tween Calliphlox amethystina (Gm.) and Chlorostil-
bon [aureoventris] prasinus (Less.)? . . . I have ex-
amined it repeatedly, and to my eye its external
characters are entirely a mixture of those of these
two species.”

Subsequent authorities listed Calliphlox
iridescens as a hybrid (e.g., Berlioz 1932,
1938; Peters 1945; Gray 1958; Wolters
1976) or omitted it altogether (e.g., Morony
et al. 1975, Sibley & Monroe 1990). The

taxonomic status of C. iridescens is still un-
certain, however, because the accounts of
Butler (1931) and Berlioz (1932, 1938) did
not adequately review the morphological
characters of the specimen in question and
those of its putative parental species. In this
paper, I confirm the hybrid origin of Calli-
phlox iridescens employing the methods
outlined in Graves (1990) and Graves &
Zusi (1990).

Material and Methods

The type of Calliphlox iridescens
(BMNH 1888.7.25.102 in The Natural His-
tory Museum, formerly British Museum of
Natural History) appears to be an adult
male in definitive plumage. This opinion is
based upon the absence of striations on the
maxillary ramphotheca (Ortiz-Crespo
1972), the presence of an iridescent gorget,
and moderately elongated outer rectrices
which lack terminal spots or markings.

I compared the specimen with series of
all species in the subfamily Trochilinae, the
typical hummingbirds (Zusi & Bentz 1982,
Sibley & Monroe 1990, Bleiweiss et al.
1997), in the collection of The Natural His-
tory Museum. Color transparencies and
videotape of the specimen were also com-
pared with the collections of the National

444 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

x 2
Museum of Natural History, Smithsonian = €
: : . ° Q
Institution. A second specimen of Calli- Sy
ss ; S
phlox iridescens, reported by Ruschi (1951) 0 g g
and deposited in the Museu Nacional, Rio ‘< 2 B28 aANMoanOm
: ee : S SP2ArTPFatrraAnesao
de Janeiro (M. N. 18275; “‘Brasil), was not S = Tas f MAA ANAA MH
2 > : Dn ee
examined. For the purposes of hybrid di- 2
agnosis (Graves 1990), I considered all 2 =
hummingbirds (Trochilinae) that occur in 8 a
the state of Rio de Janeiro as potential pa- Son
: : SA
rental species (Appendix 1). gc See ane
Measurements of wing chord, bill length S 09 i 2 ir y > 2 ‘
(from anterior extension of feathers), and S . ~| omaotaom
rectrix length (from point of insertion ofthe 22 oe Soaaada
central rectrices to the tip of each rectrix) E 2 Ss ot ete ee
° SD . a ct 4! 3b (On
were taken with digital calipers and round- BS ep Oh ena
\O AANDMNNAN WO
ed to the nearest 0.1 mm (Table 1). Color < 0 Kt nme ew
“ 5 2 a 7. Toe naaadand
descriptions were made under natural light. 2
I considered four alternatives—the spec- ‘3 8
° é 2)
imen represents an unrecognized color oe
morph of a species listed in Appendix 1, a 2 §
5) eS 2
chemically-altered artifact, a hybrid, or a aS
: es Ae oe Reonagngr
valid species. Because Calliphlox irides ie ee aS
cens differs significantly in size and shape oS Pere eee
1 i I ili- = Siew Se gen Oa €8
from all species in the subfamily Trochili 2s Silaats ee =
nae, it does not represent a previously un- a S esp CuTccre ce
é el gif aESrtoOrWg :
discovered color morph or chemically-al- Se Sspaawr ais s
tered artifact. As hybrids have no standing 2: id hele ee ©
i wo) = . aay . . ° ° : =
in zoological nomenclature, the burden of eS we SSeS Ss
oes 5
proof rests on the systematist to refute the ae e
Love sre 2 on} ue)
possibility of hybridization before bestow- oar 6
ing species status on a unique specimen. I — Vv aM
: : OX IPS
was unable to reject the hypothesis of hy- = Il &
bridity and thus refer to the specimen asa = 8& ah ee alee
ies = maeeoanacl ots
hybrid in the remainder of the paper. Bs CL ot) Tor ee
. . ° . iol
The diagnosis was approached hierarchi- -¥ & | Pee SS
i a 85 gS] 6 oT Slee
cally. The pool of potential parental species z . z 0 RS) ee
(a maximum of 37 = 351 pairwise combi- 3 a Sif haAnomte one |
nations, Appendix 1) was narrowed by the SHS aS m = a a a a & Looe
comparative analysis of plumage and soft Hes es ean =o ‘sg
¥ wm Gam Se ee © |l
part colors and feather shape. The restric- 225 c Sent meee
° . . (=|
tive hypothesis then was tested with an 2 FS E S : 5
analysis of size and external proportions. In 3 a ES cs
: : . = = he
previous papers I used bivariate plots of nabs J came) s
(Oe=) aS
mensural characters and least squares re- og pe ak ae
> - - 3 : ae = onmN &
gression lines (Wilkinson 1989) projected ‘ SE DRS ese
through parental measurements to illustrate us = ce Oe & SS ererae
. . . e = c= LY ~ oH Dien!
the relationship of hybrids to their hypoth- 2 2 & Se op al el knee eee
; oss whe eee) Ss aioe
esized parental species (e.g., Graves & a8 iS Sips es esis sus | [less 22) 62) 22
< SS 0 0 0 0 O s 2 o BD
Newfield 1996, Graves 1998a, 1998b). sO Smeets

VOLUME 112, NUMBER 2 445

Fig. 1. Ventral and dorsal views of adult male Chlorostilbon aureoventris (top), Calliphlox amethystina

(bottom), and their putative hybrid, C. aureoventris X C. amethystina, (=Calliphlox iridescens Gould, 1860:
BMNH 1888.7.25.102).

446 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

55 55
50 50 P
=45
= 45 &
© @
S bs
e S
= 40 = 40
35 35
30
6 20 30 25 30 35 40
Length Rectrix 1 Length Rectrix 5
18 35
17
30
16
ve = 25
® 15 5
®
=) a
=o £
= 2 20
=|
13
15 4
12
"4 10
30 35 40 45 50 55 25 30 35 40
ing Cengin Length Rectrix 5

Fig. 2. Bivariate plots of selected measurements (see Table 1) of adult male Chlorostilbon aureoventris (A),
Calliphlox amethystina (@), and their putative hybrid (#), C. aureoventris X C. amethystina, (=Calliphlox
iridescens Gould, 1860; BMNH 1888.7.25.102). Least squares regression lines are illustrated for comparison.

Table 2.—The percent difference between measurements (mm) of the hybrid (=Calliphlox iridescens Gould,
1860; BMNH 1888.7.25.102) and the mensural midpoints (average of character means from Table 1) of species
combinations.

Chlorestes notatus & Chlorostilbon aureoventris &
Calliphlox amethystina Hybrid Calliphlox amethystina Hybrid
Parental Percent Parental Percent
Midpoint Difference Midpoint Difference
Wing chord 40.4 oe, oe} 0.4
Bill Length 14.4 0.5 1BK7 3.8
Rl 2ACS 24.0 iL (AF pL 8)
R2 232 10.2 107 G7
R3 26.1 9.2 Daa 0.9
R4 29.2 pH) 28.3 1.0

R5 30.5 0.8 S1e5 BS)

VOLUME 112, NUMBER 2

Close proximity of hybrids and regression
lines (for all pairwise combinations of var-
iables) was interpreted as evidence consis-
tent with the specified hybrid hypothesis,
assuming polygenic inheritance of external
morphology. Concordance of results from
plumage and size analyses is regarded as
strong support for the hypothesis (Graves
1990, Graves & Zusi 1990).

Results and Discussion

Plumage characters.—The hybrid pos-
sesses several characters that facilitate the
identification of its parental species: (a)
brilliant silvery-green gorget; (b) moderate-
ly forked tail (fork depth = 43% of tail
length); and (c) mandibular ramphotheca
yellowish-brown (Fig. 1). Perhaps as infor-
mative, the hybrid lacks several conspicu-
ous traits that are present among source
pool species (Appendix 1): (a) contrasting
rump band; (b) brilliant frontlet or coronal
patch; (c) rufous or chestnut pigmentation
on rectrices; (d) pronounced blue or violet
iridescence on body plumage; (e) white rec-
tricial spots; (f) white bases or margins of
gorget feathers; (g) thickened primary ra-
chises; and (h) racket-tipped or attenuated
rectrix tips.

This association of characters can be de-
rived from only two of the possible pair-
wise combinations of species (Appendix 1):
Chlorestes notatus X Calliphlox amethys-
tina and Chlorostilbon aureoventris X Cal-
liphlox amethystina. Other combinations of
species can be eliminated from consider-
ation because they either lack characters ex-
hibited by the hybrid, or possess one or
more distinctive characters that are not ex-
pressed, even subtly, in the hybrid. The
geographic ranges of C. aureoventris and
C. amethystina overlap extensively in Bra-
zil, and both are found in the vicinity of
Nova Friburgo. C. notatus appears to reach
its southern limit on the Atlantic coastal
plain near the city of Rio de Janeiro and is
not known to occur in the uplands near
Nova Friburgo. However, 19th century col-

447

lections of birds from Nova Friburgo often
contained species from nearby lowlands
(fide J. E Pacheco, pers. comm.). C. notatus
and C. aureoventris are similar in size and
plumage color, differing most noticeably in
tail shape—square or slightly rounded in C.
notatus, shallowly forked in C. aureoventris
(Table 1).

External measurements.—I evaluated the
two parental hypotheses by inspecting raw
data, bivariate plots, and least squares re-
gressions of measurements. Measurements
of the hybrid fell within the character
means of both possible parental combina-
tions, Chlorestes notatus X Calliphlox ame-
thystina and Chlorostilbon aureoventris X
Calliphlox amethystina. External measure-
ments of the hybrid most closely approxi-
mate the values expected from least squares
regression of measurements of Chlorostil-
bon aureoventris X Calliphlox amethystina
(Fig. 2, Appendix 2). Hybrid characters dif-
fer from the parental midpoints (average of
parental character means, Table 2) of C. au-
reoventris X C. amethystina by 0.4—6.7%,
and from C. notatus X C. amethystina by
0.5—24%. Measurements of the hybrid are
closer to the parental midpoint of C. au-
reoventris X C. amethystina for 5 of the 7
characters.

In summary, both plumage and external
morphology are consistent with the hypoth-
esis that Calliphlox iridescens represents a
hybrid between Chlorostilbon aureoventris
and Calliphlox amethystina. For taxonomic
purposes, Calliphlox iridescens Gould is
available only for the purpose of homony-
my.

Acknowledgments

I thank Robert Prfys-Jones, Michael Wal-
ters, and Mark Adams of The Natural His-
tory Museum, Tring, for permitting me to
study the type of Calliphlox iridescens on
two different occasions, and Jose Fernando
Pacheco (Rio de Janeiro) for providing the
list of species in Appendix 1 and distribu-
tional data for Chlorestes notatus. The man-

448

uscript was reviewed by Richard Banks,
Jose Fernando Pacheco, Robert Pr¥ys-Jones,
Michael Walters, and Richard Zusi, whom
I thank for their comments. Leslie Over-
street helped with literature. The Smithson-
ian photographic services prepared the
prints. Travel was supported by the Alex-
ander Wetmore Fund and the Research Op-
portunities Fund, Smithsonian Institution.

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. 1998a. Diagnoses of hybrid hummingbirds

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Appendix 1

Species of trochiline hummingbirds that occur in the
state of Rio de Janeiro, Brazil (fide J. E Pacheco, pers.
comm.). Vagrant species (less than three records in Rio
de Janeiro) are marked by an asterisk. Parentheses en-
close a representative list of characters or traits that
would probably be expressed in hybrid progeny of
these species, but that do not occur in Calliphlox iri-
descens Gould, 1860 (BMNH 1888.7.25.102). Taxon-
omy follows Sibley & Monroe (1990): Eupetomena
macroura (violet-blue head and breast, thickened pri-
mary rachises); Melanotrochilus fuscus (black body
plumage, white outer rectrices); Colibri serrirostris
(purple auricular tufts, subterminal band on rectrices);
Anthracothorax nigricollis (black ventral plumage, ru-
fous pigmentation on rectrices); *Chrysolampis mos-
quitus (brilliant coronal patch, rufous pigmentation on
rectrices); Stephanoxis lalandi (brilliant coronal patch,
elongated crest plumes, blue ventral plumage); Lo-
phornis magnificus (rufous crest, contrasting rump
band); Lophornis chalybeus (white-tipped gorget
feathers, contrasting rump band); Popelairia langs-
dorffi (contrasting rump band, attenuated rectrices);
*Discosura longicauda (contrasting rump band, rack-
et-tipped rectrices), Chlorestes notatus, Chlorostilbon

VOLUME 112, NUMBER 2

aureoventris, *Thalurania furcata (violet breast and
belly); Thalurania glaucopis (brilliant coronal patch),
Hylocharis sapphirina (rufous chin and rectrices, vio-
let head and breast); Hylocharis cyanus (white chin,
violet head and upper breast); *Hylocharis chrysura
(cinnamomeus chin, golden-bronze tail); Leucochloris
albicollis (white throat, white-tipped rectrices); Polyt-
mus guainumbi (white-tipped rectrices); Amazilia ver-
sicolor (white throat, dark subterminal band on outer
rectrices); Amazilia fimbriata (white-margined throat
feathers); Amazilia lactea (violet-blue throat and upper
breast); Aphantochroa cirrochloris (dull plumage,
large size); Clytolaema rubricauda (rufous rectrices);
Heliothryx aurita (brilliant coronal patch, white outer
rectrices); Heliomaster squamosus (brilliant coronal
patch, white malar mark, white medial stripe from up-
per breast to vent); Calliphlox amethystina.

Appendix 2

General comparative description of definitive plum-
ages of male Chlorostilbon aureoventris, Calliphlox
amethystina, and the hybrid, C. aureoventris X C.
amethystina (=Calliphlox iridescens Gould, 1860;
BMNH 1888.7.25.102). Descriptions of structural col-
ors are unusually subjective, as color seen by the ob-
server varies according to the angle of inspection and
direction of light. For this reason I use general color
descriptions.

The dorsal plumage in amethystina, from crown to
uppertail coverts, is weakly iridescent and dull green
to pale bronzy-green in coloration; the iridescence is
brighter from a “‘tail-on”’ view, as opposed to a “‘head-
on”’ view. The crown is dull dark green viewed head-
on. The dorsum of aureoventris is significantly more
iridescent than that of amethystina, appearing golden-
green to bluish-green, depending on the angle of ob-
servation. The crown is brilliant golden-green, viewed
head-on, with coppery reflections on the periphery.

The quality and brightness of dorsal iridescence in
iridescens is intermediate to those of the parental spe-
cies, but closer in overall appearance to amethystina.
The crown reflects a pale, but variable, bluish-green
iridescence when viewed head-on.

The brilliant rosy-red to purplish-red gorget of ame-
thystina, which extends from the chin laterally to the
eye and posterior to the upper throat, is bordered pos-
teriorly by a white or grayish-white pectoral band that
blends posteriorly into dull green on the sides. Gorget
feathers are of moderate length (6.1—7.0 mm), medium
gray basally bordered distally by a narrow transitional
band of gray glossed with green and tipped with a
rosy-red terminal disk (from posterior margin of gor-
get: 2.1-2.4 mm deep, 1.7—2.9 mm wide). Feathers of
the lower breast, sides, and flanks are dark gray ba-
sally, tipped subterminally with a weakly-iridescent
green disk, and fringed (heavily along the midline)
with grayish-buff or buff. Vent feathers are white. Tib-

449

ial plumes, which extend past the base of the hallux,
are dark gray, broadly tipped with buffy-white. Un-
dertail coverts are grayish-buff fading to white or pale
buffy-white at the margins (subterminally glossed with
green in some individuals).

With the exception of white vent plumes, the ventral
plumage of aureoventris exhibits brilliant iridescence
when viewed head-on. Although there is considerable
color variation among individuals, iridescence is pre-
dominately bluish-green on the throat, upper breast,
and undertail coverts, tending toward golden-green on
the lower breast, sides and belly. Throat feathers are
medium gray basally, becoming dark gray distally, and
abruptly tipped with a bluish-green disk (from lower
throat, 1.6—2.0 mm deep, 3.0—3.3 mm wide). Gorget
feathers (5.1—-5.8 mm) are relatively shorter than in
amethystina. Tibial feathers are dark gray and reach
but do not exceed the base of the hallux.

The gorget of iridescens, similar in shape to that of
amethystina, exhibits a peculiar pattern of iridescence,
predominately pale silvery-green viewed head-on, but
irregularly marked with a coppery hue, especially on
the sides of the throat. Closer inspection reveals this
is due to coppery or bronze iridescence emanating
from barb tips of otherwise silvery-green disks (or sil-
very-blue in certain lights). Lateral gorget feathers
(5.9-6.0 mm long) are dark gray basally, broadly
tipped with a silvery-green disk (2.1—2.2 mm deep,
2.7—2.8 mm wide). The depth (usually <1.0 mm) and
intensity of coppery disk margins increase laterally, a
few gorget feathers lacking coppery iridescence are
juxtaposed among margined feathers in the center of
the throat. The breast and sides of iridescens are dark
green (dark gray with only a hint of green iridescence
viewed head-on); feather bases are dark gray and gray-
ish feather margins are largely restricted to the lower
midline above the vent. Evidence of the white pectoral
band of amethystina is limited in iridescens to a scat-
tering of white and pale gray basal feather barbs. Tibial
plumes, which are dark brownish-gray lightly tipped
pale buffy-gray, are intermediate in length between
those of amethystina and aureoventris, narrowly pass-
ing the base of the hallux. Undertail coverts are buffy
gray with a weakly-defined subterminal green spot of
variable size and extensively margined with pale buf-
fy-gray.

The tail of amethystina is moderately forked. The
outer rectrices (R2—R5) are narrow (3.3—3.6 mm wide)
and dull purplish-black in coloration. The outer vane
is faintly (R3) or moderately (R2) glossed with green.
R3-RS5 are faintly tipped with green in some individ-
uals (unstriated ramphotheca). Both vanes of R1 are
extensively glossed with dark green. Rachises are dark
brown on both surfaces. The shallowly forked tail of
aureoventris, which is shining steel-blue on both sur-
faces, contrasts highly with the brilliant bluish-green
tail coverts. Outer rectrices are 5.4—6.8 mm wide.

The color and shape of the hybrid’s tail are inter-

450

mediate between those of amethystina and aureoven-
tris. The right outer rectrix (R5) is 4.0 mm wide. The
outer vane of R2 and both vanes of RI are glossed
with green.

Remiges of amethystina are dark purplish-brown,
whereas those of aureoventris are bluish-black and sig-
nificantly glossier. Neither species shows unusual
notching or emargination of the primaries and second-
aries. The remiges of the hybrid are intermediate in
color between those of the hypothesized parental spe-
cies.

The maxillary ramphotheca in amethystina is black,
the mandibular ramphotheca is brownish-black distal-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ly, medium brown at the base of the bill. Feathering
on the maxillary ramphotheca extends to the anterior
edge of the nasal operculum but does not obscure it.
The mandibular ramphotheca and the proximal % of
the maxillary ramphotheca of aureoventris is light yel-
lowish-brown (red in life). Feathering does not reach
the anterior edge of the nasal operculum, which is fully
exposed. The bill of the hybrid is almost perfectly in-
termediate in color. The maxillary ramphotheca is dark
brown proximally becoming black distally. The man-
dibular ramphotheca is pale yellowish-brown, gradu-
ally darkening to brownish-black on the distal fifth.
Feathering extends to the anterior edge of the nasal
operculum, which is slightly inflated.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
112(2):451—452. 1999.

INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE

Applications published in the Bulletin of Zoological Nomenclature

The following Applications were published on 31 March 1999 in Vol. 56, Part 1
of the Bulletin of Zoological Nomenclature. Comment or advice on any of these
applications is invited for publication in the Bulletin and should be sent to the
Executive Secretary (I.C.Z.N.), c/o The Natural History Museum, Cromwell Road,
London SW7 5BD, U.K. (e-mail: iczn@nhm.ac.uk).

Case No.

3074 Eudendrium arbuscula Wright, 1859 (Cnidaria, Hydrozoa): proposed con-
servation of the specific name.

3048 NYMPHULINAE Duponchel, [1845] (Insecta, Lepidoptera): proposed precedence
Over ACENTROPINAE Stephens, 1835.

3054 AUGOCHLORINI Moure, 1943 (Insecta, Hymenoptera): proposed precedence
Over OXYSTOGLOSSINI Schrottky, 1909.

3069 Solenopsis invicta Buren, 1972 (Insecta, Hymenoptera): proposed conserva-
tion of the specific name.

3064 Strongylogaster Dahlbom, 1835 (Insecta, Hymenoptera): proposed conser-
vation by the designation of Tenthredo multifasciata Geoffroy in
Fourcroy, 1785 as the type species.

Hemibagrus Bleeker, 1862 (Osteichthyes, Siluriformes): proposed stability of
nomenclature by the designation of a single neotype for both Bagrus
nemurus Valenciennes, 1840 and B. sieboldii Bleeker, 1846, and the
designation of the lectotype of B. planiceps Valenciennes, 1840 as
the neotype of B. flavus Bleeker, 1846.

Megalotragus Van Hoepen, 1932 (Mammalia, Artiodactyla): proposed con-
servation, and Alcelaphus kattwinkeli Schwarz, 1932 (currently Me-
galotragus kattwinkeli): proposed conservation of the specific name.

452 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Opinions published in the Bulletin of Zoological Nomenclature

The following Opinions were published on 31 March 1999 in Vol. 56, Part 1 of
the Bulletin of Zoological Nomenclature. Copies of these Opinions can be obtained
free of charge from the Executive Secretary, I.C.Z.N., c/o The Natural History Mu-
seum, Cromwell Road, London SW7 5BD, U.K. (e-mail: iczn@nhm.ac.uk).

Opinion No.

1913 Pila R6ding, 1798 and Pomacea Perry, 1810 (Mollusca, Gastropoda): placed
on the Official List, and AMPULLARIIDAE Gray, 1824: confirmed as
the nomenclaturally valid synonym of PILIDAE Preston, 1915.

1914 Belemnotheutis Pearce, 1842, Geopeltis Regteren Altena, 1949, Geoteuthis
Miinster, 1843, Jeletzkyteuthis Doyle, 1990, Loligosepia Quenstedt,
1839, Parabelopeltis Naef, 1921, Paraplesioteuthis Naef, 1921
(Mollusca, Coleoidea): conserved, and the specific name of Belem-
noteuthis (sic) montefiorei Buckman, 1880: conserved.

Suchonella Spizharsky, 1937 (Crustacea, Ostracoda): Suchonella typica Spi-
zharsky, 1939 designated as the type species.

BRACHYPTERINAE Zwick, 1973 (Insecta, Plecoptera): spelling emended to BRA-
CHYPTERAINAE, SO removing the homonymy with BRACHYPTERINAE
Erichson, [1845] (Insecta, Coleoptera); KATERETIDAE Erichson in Ag-
assiz, [1846]: given precedence over BRACHYPTERINAE Erichson.

Papilio camillus Fabricius, 1781 (currently Cyrestis camillus) and Limenitis
reducta Staudinger, 1901 (Insecta, Lepidoptera): specific names con-
served.

MELOIDAE Gyllenhal, 1810 and NEMOGNATHINAE Castelnau, 1840 (Insecta, Co-
leoptera): given precedence over HORIDAE Latreille, 1802.
Polyrhachis Smith, 1857 (Insecta, Hymenoptera): given precedence over

Myrma Billberg, 1820.

Strongylopus Tschudi, 1838 (Amphibia, Anura): Rana fasciata Smith, 1849
designated as the type species.

PETROPEDETINAE Noble, 1931, CACOSTERNINAE Noble, 1931 and PHRYNOBA-
TRACHINAE Laurent, 1941 (Amphibia, Anura): given precedence over
HEMIMANTIDAE Hoffman, 1878, and PHRYNOBATRACHINAE: not given
precedence over PETROPEDETINAE.

Loris E. Geoffroy Saint-Hilaire, 1796 (Mammalia, Primates): conserved, and
correction made to the entry for Choloepus Illiger, 1811 (Xenarthra)
on the Official List.

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CONTENTS

A revision of the Antarctic genus Chlanidota (Gastropoda: Neogastropoda: Buccinulidae)
M. G. Harasewych and Yuri I. Kantor
Hydroid and medusa stages of the new species Ectopleura obypa (Cnidaria: Hydrozoa: Tub-

ulariidae) from Brazil Alvaro E. Migotto and Antonio C. Marques
Striatodoma dorothea (Cheilostomatida: Tessaradomidae), a new genus and species of bryo-
zoan from deep water off California Judith E. Winston and Stace E. Beaulieu
Two new species of Dentatisyilis and Branchiosyllis (Polychaeta: Syllidae: Syllinae) from
Venezuela Guillermo San Martin and David Bone
Parvidrilus strayeri, a new genus and species, an enigmatic interstitial clitellate from under-
ground waters in Alabama Christer Erséus

On the entocytherid ostracods of the Brazos River basin and adjacent coastal region of Texas
Daniel J. Peters and Jean E. Pugh
A new genus, Neodoxomysis (Crustacea: Mysidacea: Mysidae: Leptomysini), with description
of two new species Masaaki Murano
First record of the family Gynodiastylidae Stebbing, 1912 (Crustacea: Malacostraca: Cumacea)
from Antarctic waters with the description of Gynodiastylis jazdzewsku, a new species
Magdalena Blazewicz and Richard W. Heard
A new genus and species of sphaeromatid isopod (Crustacea) from Atlantic Costa Rica
Regina Wetzer and Niel L. Bruce
Icelopagurus tuberculosus, a distinct new hermit crab species (Crustacea: Decapoda: Paguridae)
from Japan Akira Asakura
A new species of crayfish of the genus Procambarus (Crustacea: Decapoda: Cambaridae) from
Veracruz, Mexico Yolanda Rojas, Fernando Alvarez, and José Luis Villalobos
A new species of freshwater crab of the genus Strengeriana Pretzmann, 1971, from Colombia
(Crustacea: Decapoda: Pseudothelphusidae) Martha R. Campos
A new genus and species of South American fishes (Teleostei: Characidae: Cheirodontinae)
with a derived caudal fin, including comments about inseminating cheirodontines
Luiz R. Malabarba and Stanley H. Weitzman
Psilotris amblyrhynchus, a new seven-spined goby (Teleostei: Gobiidae) from Belize, with

notes on settlement-stage larvae David G. Smith and Carole C. Baldwin
Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 7. Probable parentage of Calliphlox
iridescens Gould, 1860 Gary R. Graves

International Commission on Zoological Nomenclature

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303
313
319
327
338

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362
368
381
396

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410
433

443
451

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— D
,,
bs,

ys 112(3):453-469. 1999.
Cytogeogr phy SePhi ine bats (Mammalia: Chiroptera)
i

(EAR, JAM) Utah Museum of
U.S.A.; (LRH) Department of Zoology, Field Museum of Natural History,
Chicago, Illinois 60605, U.S.A.

Abstract.—Standard karyotypes of 17 species of Philippine bats representing
12 genera and 5 families are described. Karyotypes of five pteropodid species
(Alionycteris paucidentata, Otopteropus cartilagonodus, Pteropus pumilus, P.
leucopterus, and Eonycteris robusta), four rhinolophids (Rhinolophus arcuatus,
R. inops, R. rufus, and Hipposideros obscurus), and two vespertilionids (Myotis
macrotarsus and Murina cyclotis) are reported for the first time. Available data
reveal a pattern of distinctive karyotypes among many Philippine bats. In sev-
eral independent groups, karyotypes are unique at different taxonomic levels.
Pteropodid and rhinolophid species endemic to the Philippines have karyotypes
that represent major new arrangements for their families or genera. Some spe-
cies widespread in Asia are chromosomally polymorphic, with major differ-
ences between populations from within and outside of the Philippines. The
large proportion of unique karyotypes seen among bats from the oceanic Phil-
ippines reflects the isolation of the archipelago from mainland southeast Asia.

PRGO EDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

al History, University of Utah, Salt Lake City, Utah 84112,

The bat fauna of the Philippine Islands is
large and diverse, consisting of 73 species
belonging to 36 genera and 6 families. It
also is relatively rich in endemic species,
with 25 (Heaney et al. 1998). High levels of
species richness and endemism are charac-
teristic of the Philippine terrestrial mammal
fauna as a whole, reflecting a history of rel-
ative isolation in an ancient tropical island
archipelago (Heaney 1986, Heaney & Rick-
art 1990). Studies of non-volant mammals
have revealed a pattern of colonization fol-
lowed by repeated speciation (Heaney 1986,
Musser & Heaney 1992, Rickart & Musser
1993, Heaney & Ruedi 1994). These pro-
cesses have shaped patterns of distribution
and species richness among Philippine fruit
bats (Heaney 1991), and similar patterns are
likely to exist among other groups of bats,
although underlying rates of colonization
and speciation are expected to vary among
them as a function of dispersal ability (Hea-
ney 1986, 1991).

Cytogenetic studies of mammals can pro-

vide insight into phylogenetic and biogeo-
graphic relationships. However, previous
information on cytogenetics of Philippine
bats is limited to a single study on members
of one family (Rickart et al. 1989). In many
other respects bats constitute one of the
most poorly known groups of Philippine
mammals (Ingle & Heaney 1992). In this
paper, we describe the standard karyotypes
(non-differentially stained chromosome
complements) of 17 species of Philippine
bats representing 12 genera and 5 families,
and discuss the significance of chromosom-
al data with respect to systematics and bio-

geography.

Methods

Specimens were obtained during field
studies conducted from 1987 to 1993. All
specimens were collected directly from nat-
ural populations and euthanized with sodi-
um pentobarbital or chloroform within 24 h
of capture. Karyotypes were prepared from

454

bone marrow and/or spleen cells following
standard in vivo methodology (Patton 1967,
as modified by Rickart et al. 1989). Fixed
cell suspensions were frozen in liquid nitro-
gen in the field, and standard karyotypes
prepared in the laboratory after storage at
0°C to —70°C for several years. Determi-
nations of diploid number were based on
chromosome counts from a minimum of 10
mitotic spreads per individual. As used
herein, fundamental number (FN) refers to
the total number of arms in the autosomal
complement. Due to the presence of minute
chromosomes, small chromosome arms, or
inability to distinguish sex chromosomes,
FN values are provisional for some species,
as indicated by question marks. Species no-
menclature follows Heaney et al. (1998).
Voucher specimens were prepared as
skins with partial skeletons, complete skel-
etons, or preserved in fluid and are depos-
ited in the Field Museum of Natural His-
tory, Chicago (FMNH), the National Mu-
seum of Natural History, Smithsonian In-
stitution, Washington (USNM), the
Philippine National Museum, Manila
(PNM), and the Utah Museum of Natural
History, University of Utah, Salt Lake City
(UMNH). Microscope slides and photomi-
crograph negatives derived from all speci-
mens examined are housed at UMNH.

Specimens Examined

Alionycteris paucidentata Kock, 1969a.—
Mindanao Island, Bukidnon Province, Mt.
Kitanglad Range, 16.5 km S, 4 km E Camp
Phillips, elev. 1900 m, 8°10'30'N,
124°51/EB;) 2ofemales ((PMNHO14782 It:
FMNH 1478272).

Otopteropus cartilagonodus Kock,
1969b.—Luzon Island, Camarines Sur
Province, Mt. Isarog, 5 km N, 20 km E
Naga, elev. 900 m, 13°40'N, 123°21’E, 2
females (USNM 573692, USNM 573695).

Pteropus leucopterus Temminck,
1853.—Catanduanes Island, 1 km N, 8.5
km W Gigmoto, elev. 200 m, 13°48’N,
124°19’E, 1 male (USNM 573263).

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Pteropus pumilus Miller, 1910.—Negros
Island, Negros Oriental Province, 3 km N,
14 km W Dumaguete, elev. 950 m, 9°22’N,
123°11’'E, 1 female (USNM 573466), 9 km
N, 14 km W Dumaguete, elev. 600 m,
9°23 'N, '123°I1'E,“?!’ female “(shim
573749).

Eonycteris robusta Miller, 1913.—Catan-
duanes Island, 1 km N, 8.5 km W Gigmoto,
elev. 200 m, 13°48’N, 124°19’E, 2 females
(USNM 573210, USNM 573211).

Taphozous melanopogon Temminck,
1841.—Negros Island, Negros Oriental
Province, 4 km N Manjuyod, elev. 20 m,
9°43'N, 123°10'E, 1 female (seme
459344).

Megaderma spasma (Linnaeus, 1758).—
Catanduanes Island, 1 km N, 8.5 km W
Gigmoto, elev. 200 m, 13°48’N, 124°19’E,
1 male (USNM 573267); Biliran Island, 5
km N, 10 km E Naval, elev. 850 m,
11°36'N, 124°29’E, 1 female (PNM speci-
men, EAR field number 1445); Leyte Is-
land, Leyte Province, 10.2 km N, 2.2 km E
Baybay, elev. 320 m, 10°46’N, 124°49’E, 1
male (USNM 459392), 10.5 km N, 4 km E
Baybay, elev. 700 m, 10°47’N, 124°50’E, 1
female (USNM 459393).

Rhinolophus arcuatus Peters, 1871.—Bi-
liran Island, 5 km N, 10 km E Naval, elev.
850 m, 11°36’N, 124°29’E, 2 males, small
morph (USNM 459445, USNM 459446);
Luzon Island, Camarines Sur Province, Mt.
Isarog, 4 km N, 18 km E Naga, elev. 475
m, 13°40’N, 123°20’E, 1 femalepoiare
morph (USNM 573757), 5 km N, 20 km E
Naga, elev. 900 m, 13°40’N, 123°21’E, 1
male, large morph (USNM 573758).

Rhinolophus inops K. Andersen, 1905.—
Leyte Island, Leyte Province, 8.5 km N, 2.5
km E Baybay, elev. 500 m, 10°46’N,
124°49’E, 1 male (USNM 458606).

Rhinolophus rufus Eydoux & Gervais,
1839.—Catanduanes Island, 1 km N, 8.5
km W Gigmoto, elev. 250 m, 13°47’N,
124°19’E, 1 male (USNM 573588).

Hipposideros ater Tempieton, 1848.—
Catanduanes Island, 4 km W Virac, elev. 50

VOLUME 112, NUMBER 3

455

ARK OW ORR AK KA FY

“a WS az

AA aA

am re

ns ee | e ¢

10 um

HE CC OY cr re

O§ AA

1g
At

Pa SB

Fig. 1.

a @

Karyotypes of: A, Alionycteris paucidentata 2 (FMNH 147821), 2N = 36, FN = 58(?); B, Otop-

10 um

C GA SB ha ba
Ao OR

fA @8 f2

teropus cartilagonodus 2 (USNM 573695), 2N = 48, FN = 62(?).

m, 13°35'N, 124°11’E, 1 female (USNM
5373274).

Hipposideros diadema (E. Geoffroy,
1813).—Leyte Island, Leyte Province, 7 km
N, 1.5 km E Baybay, elev. 50 m, 10°45’N,
124°48'’E, 2 males (USNM 458637,
459418).

Hipposideros obscurus (Peters, 1861).—

Catanduanes Island, 1 km N, 8.5 km W
Gigmoto, elev. 250 m, 13°47'N, 124°19’E,
1 female (USNM 573280), 1 km S, 4 km
W Gigmoto, elev. 600 m, 13°46’N,
124°21’E, 1 male (UMNH 29005); Negros
Island, Negros Oriental Province, 3 km N,
14 km W Dumaguete, elev. 950 m, 9°22’N,
123°11’E, 1 female (USNM 459435).

456

Myotis macrotarsus (Waterhouse,
1845).—Negros Island, Negros Oriental
Province, 4 km N Manjuyod, elev. 20 m,
9°43'N, 123°10’'E, 2 males (USNM 459739,
USNM 459740).

Pipistrellus javanicus (Gray, 1838).—
Luzon Island, Camarines Sur Province, Mt.
Isarog, 4 km N, 21.5 km E Naga, elev.
1,550 m, 1340'N, 123°22'E,.2 males
(USNM 573822, USNM 573829).

Scotophilus kuhlii Leach, 1821.—Leyte
Island, Leyte Province, 7 km N Baybay,
elev. 10 m, 10°45'N, 124°47’E, 1 male, 1
female (USNM 458697, USNM 458699);
Negros Island, Negros Oriental Province, 2
km E Siaton, elev. 5 m, 9°3’N, 123°3’E, 1
male (USNM 458719).

Murina cyclotis Dobson, 1872.—Luzon
Island, Camarines Sur Province, Mt. Isarog,
4 km N, 18 km E Naga, elev. 475 m,
13°40'N, 123°20’E, 1 female (USNM
573776), 4.5 km N, 20.5 km E Naga, elev.
1 1235..meel3-405N,.. 123°21'E,. 1 female
(USNM 573777).

Results

Standard karyotypes for specimens of 14
species are presented in Figs. 1—7 and brief-
ly described below. Karyotypes of 3 addi-
tional species are described but not illus-
trated.

Family Pteropodidae
Subfamily Pteropodinae

Alionycteris paucidentata. 2N = 36, FN
= 58(?), Fig. 1A.—This is the first report
on the chromosomes of this species, be-
longing to a monotypic genus endemic to
Mindanao Island. The Karyotype consists of
13 pairs of metacentric or submetacentric,
and 5 pairs of acrocentric chromosomes.
The sex chromosomes cannot be deter-
mined because females only were exam-
ined, but the X chromosomes presumably
are metacentric or submetacentric as in oth-
er Philippine cynopterines (Rickart et al.
1989).

Otopteropus cartilagonodus. 2N = 48,

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FN = 62(?), Fig. 1B.—The karyotype of
this species, belonging to a monotypic ge-
nus endemic to Luzon Island, is reported
for the first time. It includes 6 pairs of meta-
centric or submetacentric chromosomes, 3
pairs of subtelocentric chromosomes, and
15 pairs which appear to be acrocentric.
Male specimens were not examined, so the
sex chromosomes cannot be determined.
However, the X chromosomes presumably
are metacentric or submetacentric as in oth-
er cynopterines (Rickart et al. 1989).

Pteropus pumilus. 2N = 38, FN = 72,
Fig. 2A.—The karyotype of this Philippine
endemic is reported for the first time. It is
composed entirely of biarmed elements, in-
cluding 10 metacentric or submetacentric,
and 9 subtelocentric pairs. Only females
were examined, so the sex chromosomes
cannot be determined. However, the X
chromosomes presumably are subtelocen-
tric as in congeners sharing this same basic
karyotype (Hood et al. 1988, RicKart et al.
1989).

Pteropus leucopterus. 2N = 38, FN = ?,
Fig. 2B.—A single male specimen yielded
the first available karyotype for this species
endemic to the Philippines. Although the
preparation quality is poor, basic features of
the karyotype are clear: it includes 1 chro-
mosome pair that is submetacentric, and 18
pairs that are either acrocentric or subtelo-
centric. The sex chromosomes cannot be
differentiated and the FN value cannot be
determined.

Subfamily Macroglossinae

Eonycteris robusta. 2N = 36, FN =
66(?), Fig. 3A.—The standard karyotype of
the endemic Philippine nectar bat is report-
ed for the first time. It includes 17 pairs of
small to large-sized biarmed chromosomes,
and one pair of small acrocentric elements.
The sex chromosomes cannot be distin-
guished since females only were examined,
but the X chromosomes presumably are
submetacentric as in E. spelaea (Rickart et
al. 1989).

VOLUME 112, NUMBER 3

457

“CE AD RC GS a8 ve

by a:

oi

na

H§ GE AG Oh Gh se

A4 Sh *

10 um

Pam Ad 4B 4% 4A wa

@a 2e

ese se eo = *
an 22 j“88@
a | | 10 um
Fig. 2. Karyotypes of: A, Pteropus pumilus 2 (USNM 573749), 2N = 38, FN = 72; B, Pteropus leucopterus

3 (USNM 573263), 2N = 38.

Family Emballonuridae

Taphozous melanopogon. 2N = 42, FN
= 64(?), not figured.—The karyotype of a
single female specimen of this widespread
species is consistent with those reported for
specimens from India (Ray-Chaudhuri et al.
1971) and Thailand (Hood & Baker 1986).
It includes 13 pairs of metacentric or sub-

metacentric, and eight pairs of acrocentric.
chromosomes. The X chromosomes are
presumably submetacentric as reported pre-
viously for this species.

Family Megadermatidae

Megaderma spasma. 2N = 46, FN = 64,
Fig. 3B.—Philippine specimens of this

458

widespread species have a karyotype that
includes 10 pairs of biarmed autosomes and
12 pairs of acrocentric autosomes. The sub-
telocentric X chromosome is the largest el-
ement in the karyotype, and the Y is a small
acrocentric chromosome.

Family Rhinolophidae
Subfamily Rhinolophinae

Rhinolophus arcuatus. 2N = 58, FN =
60, Fig. 4A.—Horseshoe bats included un-
der this name are widely distributed among
the islands of Southeast Asia (Koopman
1993), and have not been karyotyped pre-
viously. Philippine specimens from Biliran
and Luzon islands have a uniform standard
karyotype consisting of 2 pairs of large
metacentric and 26 pairs of small to large
acrocentric autosomes, a medium-sized
subtelocentric X and an acrocentric Y chro-
mosome. One pair of medium-sized acro-
centric autosomes have a secondary con-
striction.

Rhinolophus inops. 2N = 58, FN = 60,
Fig. 4B.—Bats included under this name
are endemic to the Philippines and have not
been karyotyped previously. A single male
specimen from Leyte Island has a standard
karyotype that is identical to those seen for
specimens of R. arcuatus (Fig. 4A).

Rhinolophus rufus. 2N = 40, FN = 60,
Fig. 5A.—The karyotype of a single male
specimen is the first reported for this en-
demic Philippine horseshoe bat. It includes
11 pairs of biarmed autosomes, 8 pairs of
acrocentric autosomes, a subtelocentric X,
and a acrocentric Y chromosome.

Subfamily Hipposiderinae

Hipposideros ater. 2N = 32, FN = 60,
Fig. 5B.—The karyotype of a single female
specimen of this widespread species closely
resembles those reported previously for
specimens from India (Ray-Chaudhuni et al.
1971, Sreepada et al. 1993). It consists of
14 pairs of metacentric or submetacentric,
and 3 pairs of subtelocentric elements. The
X chromosomes cannot be distinguished

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

from the autosomes, but presumably are
medium-sized and metacentric as reported
previously for this species.

Hipposideros diadema. 2N = 32, FN =
60, not figured.—Standard karyotypes from
Philippine specimens of this widespread
Asian species consist entirely of biarmed
chromosomes, and closely resemble the
karyotype reported for a specimen from
Borneo (Harada & Kobayashi 1980).

Hipposideros obscurus. 2N = 24, FN =
44, Fig. 5|C.—The karyotype of this endem-
ic Philippine roundleaf bat is reported for
the first time. It includes 10 pairs of meta-
centric or submetacentric, and 2 pairs of
subtelocentric chromosomes. The sex chro-
mosomes cannot be distinguished with cer-
tainty, but the X is presumably medium-
sized and either metacentric or submetacen-
tric, as in congeners.

Family Vespertilionidae
Subfamily Vespertilioninae

Myotis macrotarsus. 2N = 44, FN = 50,
Fig. 6A.—The karyotype of this species is
reported for the first time. It has the ar-
rangement typical for members of the genus
Myotis (Bickham 1979a, Volleth & Heller
1994), consisting of 3 large metacentric, 1
small submetacentric, and 17 acrocentric
autosomal pairs, a submetacentric X and an
acrocentric Y chromosome.

Pipistrellus javanicus. 2N = 38, FN =
48, Fig. 6B.—Philippine specimens of this
widespread east Asian species have a kKar-
yotype consisting of 6 pairs of large biarm-
ed autosomes, 12 pairs of acrocentric au-
tosomes, and acrocentric X and Y chro-
mosomes.

Scotophilus kuhlii. 2N = 36, FN = 48,
not figured.—Philippine specimens of the
widespread Asian house bat have a standard
karyotype that includes seven pairs of bi-
armed and ten pairs of acrocentric auto-
somes, a submetacentric X chromosome,
and an acrocentric Y chromosome.

VOLUME 112, NUMBER 3

“OH MH OG

459

BR SS ag

10 um

ae Hae ww &©w eh ee

Ra “AM aa
aann ne

ae not na

Fig. 3.
spasma 2? (EAR 1445), 2N = 46, FN = 64.

Subfamily Murininae

Murina cyclotis. 2N = 44, FN = 50(?),
Fig. 7.—The karyotype of this widespread
Asian species is reported for the first time.
The autosomes include three pairs of large
metacentric and one pair of small submeta-
centric elements, and seventeen pairs of
small to medium-sized elements that appear
to be acrocentric. Females only were ex-
amined, so the sex chromosomes cannot be

10 um

L

aanan a0

a eo sé

Karyotypes of: A, Eonycteris robusta 2 (USNM 573210), 2N = 36, FN = 66: B, Megaderma

determined with certainty. Based on com-
parisons with karyotypes of congeners
(McBee et al. 1986, Harada et al. 1987),
two medium-sized submetacentric elements
are tentatively identified as the X chromo-
somes.

Discussion

Several studies have demonstrated that
non-differentially stained chromosomes

“RR M8
@f GA
aa Ae
oe ne

=> * a: &

= >

po ee
nh ac
ap an

~~ Ge &
=2* £8

a ee a .

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10 um Ae
xX Y
oP

aa ae fo ef
u&® C® AA @A

~a at Aa*® a ®8

10 um 4
X

FA AN AA G0
Fa ADB aan a

ar #62 2 &@& “*

= & & %& = -“

Fig. 4. Karyotypes of: A, Rhinolophus arcuatus (small morph) 6 (USNM 459445), 2N = 58, FN = 60; B,
Rhinolophus inops 3 (USNM 458606), 2N = 58, FN = 60.

VOLUME 112, NUMBER 3

461

“3B OR Rk BR SD eg

RN gw A

as sa fle
Y

X
aa 2@e aa one ef +@-
10 um

PES AX Re ome xe xs

XR KX 5% 44 as

a A 10 um

mr jt fy (8 ass

| See wee & sty
Zi a4 10 um

Fig. 5. Karyotypes of: A, Rhinolophus rufus 3 (USNM 573588), 2N = 40, FN = 60: B, Hipposideros ater
(USNM 573274), 2N = 32, FN = 60: C, Hipposideros obscurus 2 (USNM 459435), 2N = 24, FN = 44,

462

WN AK KG em

AB AN ag

46> @) nA

A

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x

aN ANH NE

Ef S38 22 &8 4.
fe a) m=

@§6 an ad

ee ah

Fig. 6.
Javanicus 6 (USNM 573822), 2N = 38, FN = 48.

ce @

may greatly underestimate the magnitude of
karyotypic evolution (Haiduk et al. 1981,
and references therein). In our analysis of
results we therefore consider any observ-
able differences between standard karyo-
types to be minimal reflections of actual
differences. More importantly, we avoid
drawing conclusions based solely on appar-
ent similarities.

With 25 species, the Pteropodidae is one
the most diverse mammalian families in the

a @ ® a

oe -* e@ « es °@

Karyotypes of: A, Myotis macrotarsus 6 (USNM 459740), 2N = 44, FN = 52; B, Pipistrellus

Philippines. It also displays a high level of
endemism; 15 species and 4 genera are re-
stricted to the archipelago (Heaney et al.
1998). All five species of pteropodids ex-
amined in this study are Philippine endem-
ics, and two belong to monotypic genera.
Alionycteris and Otopteropus are part of
the diverse cynopterine clade that has un-
dergone substantial radiation in southeast
Asia (Rickart et al. 1989:528, fig. 5). Kar-
yotypes of the two genera differ substan-

VOLUME 112, NUMBER 3

ig. 7:

tially, providing no support for a previously
hypothesized sister-group relationship
(Rickart et al. 1989, Heaney 1991). The
standard karyotype of Alionycteris (2N =
36; Fig. 1A) differs from those of other
pteropodid genera that have been examined
thus far. It most closely resembles karyo-
types of Cynopterus that have 2N = 34 and
FN = 58 (Ando et al. 1980a, Harada & Ko-
bayashi 1980, Rickart et al. 1989), the only
apparent difference involving a single Rob-
ertsonian translocation. The karyotype of
Otopteropus (2N = 48; Fig. 1B) also may
represent a novel arrangement for the fam-
ily. It resembles those of Ptenochirus (2N
= 44-46), another cynopterine genus en-
demic to the Philippines, and thought to be
closely related (Rickart et al. 1989); both
genera possess three pairs of large-sized
biarmed chromosomes and several smaller
subtelocentric and acrocentric elements.
The east Asian cynopterine Penthetor, con-
sidered a more distant relative of Otopter-
opus (Rickart et al. 1989), also reportedly
has 2N = 48 (Yong & Dhaliwal 1976), but
the karyotype has not been published so
comparisons are not possible.

Previous studies on the genus Pteropus

463

X X
?

06 AA ay

q4@4@ ee” @ «64

10 um

aA

Karyotypes of Murina cyclotis 2° (USNM 573777), 2N = 44, FN = 50(?).

have characterized it as chromosomally
conservative: most species, including P.
pumilus (Fig. 2A), have karyotypes of 2N
= 38 and FN = 72 (Hood et al. 1988, Rick-
art et al. 1989). The karyotype of the one
specimen of Pteropus leucopterus (Fig. 2B)
represents a major departure from the pre-
dominant arrangement for the genus. The
absence of large biarmed autosomes and the
predominance of acrocentric or subtelocen-
tric elements distinguish this Philippine en-
demic from all other species of Pteropus
that have been examined thus far.

The standard karyotype of the Philippine
nectar bat, Eonycteris robusta (2N = 36;
Fig. 3A), is indistinguishable from that of
the widespread E. spelaea (Harada et al.
1982, Rickart et al. 1989). From the limited
perspective of standard karyotypes, there
appears to be little chromosomal variation
among the few members of the subfamily
Macroglossinae that have been examined to
date (Haiduk et al. 1980, Rickart et al.
1989).

Our results demonstrate that the wide-
spread Asian ghost bat, Megaderma spas-
ma, 1s chromosomally polytypic. Hood et
al. (1988) reported a karyotype of 2N = 38

464

for specimens from Thailand. Philippine
specimens from three islands representing
two distinct faunal regions (Heaney 1986)
all have 2N = 46 (Fig. 3B). Philippine and
Thai specimens have similar sex chromo-
somes and share FN = 64, indicating that
Robertsonian translocations among the au-
tosomes may account for all or most of the
observed differences between the karyo-
types. Megaderma lyra, the only other me-
gadermatid that has been karyotyped, has a
very different arrangement: specimens from
both India and Thailand have 2N = 54, FN
= 104 (Ray-Chaudhuri et al. 1971, Naidu
& Gururaj 1985, Hood et al. 1988). The
two species also differ substantially in
amount and distribution of C-positive het-
erochromatin and in patterns of G-banding
(Hood et al. 1988).

Karyotype differences of the magnitude
seen in Megaderma spasma suggest that
more than one species may be included un-
der this name at present. Although chro-
mosomal polymorphism alone does not pre-
clude possible gene flow between cytotypes
(e.g., as occurs in Uroderma bilobatum;
Baker 1981, Greenbaum 1981), in M. spas-
ma it is correlated with geographic isolation
by extensive water barriers. Additional data
from both mainland and island populations
are needed, and a systematic review of the
group is warranted.

The genus Rhinolophus is one of the
most diverse yet poorly known mammalian
groups in the Philippines (Heaney et al.
1998). The status of two of the taxa ex-
amined herein, R. arcuatus and R. inops, is
uncertain. Both of these taxa have been
considered members of the same species
group (Andersen 1905, Tate & Archbold
1939). Philippine specimens currently in-
cluded within R. arcuatus probably repre-
sent more than one species. Two body size
classes are present on most islands, and ad-
ditional geographic variation is apparent be-
tween different islands. Specimens referred
to R. inops also exhibit geographic varia-
tion, and may represent more than one spe-
cies (Heaney et al. 1991, Ingle & Heaney

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1992, Heaney et al. 1998). Data presented
here do not help resolve either of these is-
sues. Specimens of both size morphs of R.
arcuatus and a single example of R. inops
have the same standard karyotype of 2N =
58, FN = 60 (Fig. 4), which is a common
arrangement in the genus (Qumsiyeh et al.
1988, Zima et al. 1992b).

Among the more than 30 species of
Rhinolophus that have been karyotyped, the
limited range in FN values (60 to 64) in-
dicates that Robertsonian rearrangements,
that do not alter FN, may account for much
of the karyotypic variation (Qumsiyeh et al.
1988, Zima et al. 1992b). A 2N = 62, FN
= 60 karyotype has been hypothesized as
primitive for the genus. That arrangement
predominates among east Asian taxa,
whereas karyotypes of 2N = 58, FN = 60-—
62 predominate in Africa, Europe, and
northern Asia (Bogdanowicz 1992, Zima et
al. 1992b). With this geographic perspec-
tive, the 2N = 58 karyotypes of R. arcuatus
and R. inops are unusual among southeast
Asian members of the genus, and may rep-
resent novel developments within the oce-
anic Philippines. Support for this is seen in
the phenetic analysis of Bogdanowicz
(1992) which places R. arcuatus within a
group of southeast Asian species that in-
cludes several taxa with 2N = 62 karyo-
types (Zima et al. 1992b).

The endemic Philippine species Rhino-
lophus rufus (2N = 40, FN = 60; Fig. 5A)
has one of the lowest diploid numbers
known for the genus, surpassed only by R.
luctus (2N = 32) and R. rouxi (2N = 36)
from mainland Asia (Zima et al. 1992b). In
a recent phenetic analysis (Bogdanowicz
1982:221, fig. 3), R. rufus tentatively clus-
tered with R. pearsonii (2N = 42) and R.
yunanensis (2N = 60), two widespread spe-
cies from mainland Asia that also have
unique karyotypes (Zima et al. 1992b). If
reduction in diploid number through centric
fusion has been the primary direction of
karyotypic evolution in Rhinolophus, as hy-
pothesized by Zima et al. (1992b), then the
unique arrangement of R. rufus may repre-

VOLUME 112, NUMBER 3

sent a novel derivation within the oceanic
Philippines.

Previous work has characterized the ge-
nus Hipposideros as chromosomally con-
servative: 14 species, including both H.
diadema and H. ater (Fig. 5B) have stan-
dard karyotypes of 2N = 32, FN = 60 with
only very minor interspecific variation in
arm length ratios (Ray-Chaudhuri et al.
1971, Peterson & Nagorsen 1975, Ando et
al. 1980b, Handa & Kaur 1980, Harada &
Kobayashi 1980, Harada et al. 1982, Hood
et al. 1988, Sreepada et al. 1993). The kar-
yotype of the Philippine endemic H. obscu-
rus (2N = 24, FN = 44; Fig. 5C) represents
a radical departure from this common ar-
rangement. The low diploid and fundamen-
tal numbers of this species distinguish it not
only from congeners, but apparently from
all other rhinolophids karyotyped thus far
which have 2N = 30-62 and FN = 58-64
(Baker et al. 1974, Ando et al. 1980b, Han-
da & Kaur 1980, Zima et al. 1992b). Given
the extent of this FN pattern and the pre-
vailing view on the primary mode and di-
rection of karyotypic evolution in the fam-
ily (Zima et al. 1992b, Sreepada et al.
1993), the chromosome arrangement of H.
obscurus may be viewed as another unique
derivation within the oceanic Philippines.

All members of the genus Myotis kar-
yotyped thus far, including M. macrotarsus
(Fig. 6A), exhibit the same basic karyotype
of 2N = 44, FN = 50-52 that is widely
regarded as the primitive arrangement for
the family Vespertilionidae (Bickham
1979b, Volleth & Heller 1994). Members of
the genus Murina that have been examined
have karyotypes similar to Myotis with 2N
= 44 and FN = 50-58 (McBee et al. 1986,
Harada et al. 1987, Volleth and Heller 1994,
this study, Fig. 7). Very small secondary
arms present in some species of Murina
cannot be discerned in the material avail-
able for M. cyclotis.

Species of Scotophilus examined to date
have 2N = 36, and FN = 48—54, with in-
terspecific differences in the size and num-
ber of arms on some autosomes (Schlitter

465

et al. 1980, McBee et al. 1986, Ruedas et
al. 1990). Scotophilus kuhlii (= S. tem-
mincki of some authors) is chromosomally
polytypic. Specimens from the Philippines
examined here have the same standard kar-
yotype of 2N = 36, FN = 48 seen in spec-
imens from Borneo and Thailand (Harada
& Kobayashi 1980:91, fig. 5, Harada et al.
1982:274, fig. 2), whereas specimens from
India have 2N = 36, FN = 52 (Pathak &
Sharma 1969:43, fig. 10). At a minimum,
these karyotypes differ by two non-Robert-
sonian events involving additional arms on
the two smallest autosomal pairs. These dif-
ferences suggest that more than one species
may be included under this name at present.
Multiple cytotypes are seen elsewhere in
Scotophilus: the widespread African spe-
cies, S. dinganii and S. viridis, show vari-
ation comparable to that of S. kuhlii (Schlit-
ter et al. 1980, Ruedas et al. 1990).

Our results indicate chromosomal poly-
morphism within another widespread east
Asian vespertilionid, Pipistrellus javanicus.
Specimens from the Philippines have 2N =
38, FN = 48 (Fig. 6B), whereas those from
Malaysia have 2N = 34, FN = 46 (Zima
et al. 1992a). These differences suggest the
involvement of both Robertsonian and non-
Robertsonian events. The autosomal ar-
rangement of Philippine specimens of P. ja-
vanicus is similar to that reported for Indian
P. mimus (2N = 38, FN = 48) by Pathak
& Sharma (1969:37, fig. 2). However, the
acrocentric KX chromosome seen in both
Philippine and Malaysian P. javanicus is
distinctive and apparently derived (Volleth
& Heller 1994). Acrocentric X chromo-
somes occur also in P. abramus (2N = 26,
FN = 44; McBee et al. 1986) and P. endoi
(2N = 36, FN = 60; Ando et al. 1977).
These taxa are considered close relatives of
P. javanicus (Hill & Harrison 1987), and P.
abramus has been arranged as a synonym
by Koopman (1993). The substantial kar-
yotypic variation seen among these species,
and within P. javanicus, indicates the need
for further study of this group.

Of the 73 species of bats known to occur

466

in the Philippines (Heaney et al. 1998), ba-
sic information on standard kKaryology is
now available for 27 (Rickart et al. 1989,
this study). These data reveal a geographic
pattern of variation at different taxonomic
levels within several independent groups. It
involves endemic genera with unique kar-
yotypes within the family Pteropodidae
(Alionycteris, Haplonycteris, Otopteropus,
Ptenochirus), endemic species with unique
karyotypes within widespread genera that
are otherwise chromosomally conservative
(Pteropus leucopterus, Rhinolophus rufus,
Hipposideros obscurus), and widespread
species that are chromosomally polytypic,
with unique karyotypes observed for Phil-
ippine populations (Megaderma spasma,
Pipistrellus javanicus).

The historical development of the Phil-
ippine Islands and their mammal fauna pro-
vides the context for understanding these
patterns. One part of the Philippines (as a
political entity) is the Palawan island group
that lies immediately to the north of Bor-
neo. This is the only portion of the Philip-
pines that had dry land connections to the
Asian continent during Pleistocene periods
of low sea level, and on biogeographic
grounds it is considered part of the Sunda
Shelf faunal region rather than part of the
main body of the Philippines (Heaney
1985, 1986). The remaining Philippine is-
lands are oceanic in origin. Most originated
as parts of island arcs far to the southeast
of their current position, although a few
represent fragments of continental material
that dipped below the ocean surface and re-
emerged subsequently as effective oceanic
islands (Hall 1996, Hamilton 1979, Heaney
1986). The available geological evidence
therefore suggests that the fauna of the
main portion of the Philippines originated
through over-water colonization, and pat-
terns of distribution of the fruit bats and
murid rodents are consistent with this (Hea-
ney 1986, 1991; Heaney & Rickart 1991,
Musser & Heaney 1992).

Notably, all of the species (or current
subspecies) of bats that show distinctive

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

and/or unusual karyotypes occur in oceanic
portions of the Philippines. In contrast, spe-
cies in groups exhibiting little or no kar-
yotypic variation occur mainly in continen-
tal Asia (including islands of the continen-
tal shelf), although some chromosomally
conservative taxa also occur in oceanic ar-
eas. Our documentation of unique chro-
mosomal arrangements in 10 of the 27 Phil-
ippine bat species that have been karyo-
typed (37%) implies that the exceptionally
high level of endemism among Philippine
mammals (34% of the bat species, and
about 97% of the rodents; Heaney et al.
1998) resulted from speciation events that
included major chromosomal rearrange-
ments as a common feature. Chromosomal
rearrangements are thought to arise most
often in small, isolated populations (Patton
& Sherwood 1983); precisely the condi-
tions that would occur with the rare colo-
nization of oceanic islands.

The presence and consistency of this pat-
tern can be tested readily. We predict that a
significant portion of the remaining 46 spe-
cies of Philippine bats will exhibit standard
karyotypes as distinctive as those docu-
mented thus far. Banded karyotypes should
exhibit additional distinctive arrangements,
even among groups showing little or no
variation in standard karyotypes. Finally,
we predict that studies on population ge-
netics of species with unusual karyotypes
will provide evidence of population bottle-
necks that are associated with rapid and
substantial genetic change.

Acknowledgments

We thank S. Goodman, P. Heideman, M.
Lepiten, D. Samson, B. Tabaranza, L. Tag-
at, and R. Utzurrum, for assistance in col-
lecting specimens. Reviews by J. Bickham
and L. Ruedas led to significant improve-
ments in the manuscript. E. King assisted
with photomicrographs. Permits and logis-
tical support were provided by the Protect-
ed Areas and Wildlife Bureau, Philippine
Department of Environment and Natural

VOLUME 112, NUMBER 3

Resources. Various aspects of this study
were supported by the National Science
Foundation (grant numbers BSR 8514423
and BSR 8912415), the Smithsonian Insti-
tution, the John D. & Catherine T. MacAr-
thur Foundation, and the Ellen Thorne
Smith Fund of the Field Museum.

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On the valid name of the lesser New Zealand short-tailed bat
(Mammalia: Chiroptera)

Gregory C. Mayer, John A. W. Kirsch, James M. Hutcheon, Frangois-Joseph Lapointe,
and Jacinthe Gingras

(GCM) Department of Biological Sciences, University of Wisconsin-Parkside, Kenosha,
Wisconsin 53141, U.S.A.; (GCM, JAWK, JMH) The University of Wisconsin Zoological
Museum, 250 North Mills St., Madison, Wisconsin 53706, U.S.A.; (FJL, JG) Départment de
sciences biologiques, Université de Montréal, Montréal, Québec H3C 3J7

Abstract.—The lesser New Zealand short-tailed bat, sometimes known as
Mystacina tuberculata Gray, 1843 was, for 14 years after its description, con-
founded with the New Zealand long-tailed bat, Vespertilio (now Chalinolobus)
tuberculatus. This confusion can be traced to Gray’s account, in which he used
the name Vespertilio tuberculatus, attributing it to G. Forster, but also proposed
the new genus Mystacina to receive the species. Although Gray did not make
available two species-group names, two interpretations of his actions are pos-
sible, depending on who is considered to be the author of the name tubercu-
latus. If the author is Gray, Mystacina tuberculata Gray is composite, in which
case the name must be settled on a part of that composite. If the author is G.
Forster, Gray’s generic name Mystacina is based on a misidentified type spe-
cies, and there is no ““Mystacina tuberculata Gray’’. In that case, the specific
name must be replaced by the first available name, which is velutina Hutton,
1872, and the current usage of Mystacina may be preserved by choosing the
zoological species before Gray as its type species. We recommend this course
of action because G. Forster is properly the author of Vespertilio tuberculatus,
and because thereby familiar usage of Mystacina and Mystacinidae, and current
subspecific classification of Mystacina, are preserved. The valid name of the
lesser New Zealand short-tailed bat is therefore Mystacina velutina Hutton,
1872, and we select Hutton’s specimen from the Hutt Valley (near Wellington,
North Island, New Zealand) as its lectotype.

Résumé.—Durant les 14 années qui ont suivi sa description en 1843, la petite
chauve-souris a queue courte de Nouvelle-Zélande, Mystacina tuberculata
Gray, a été confondue avec la chauve-souris a queue longue de Nouvelle-
Zélande Vespertilio (maintenant Chalinolobus) tuberculatus, ce dernier nom lui
ayant été attribué soit par Gray, G. Forster ou J. R. Forster. La confusion
remonte au compte-rendu initial de Gray dans lequel il utilise le nom Vesper-
tilio tuberculatus qu’il attribue 4 G. Forster. Dans ce méme compte-rendu, Gray
propose le nouveau genre Mystacina pour y classer l’espéce V. tuberculatus.
Si les travaux de Gray ne rendent pas deux noms de groupes-espéces dispon-
ibles, deux autres interprétations sont possibles selon l’identité de l’auteur du
nom tuberculatus. Si ’ auteur est Gray, Mystacina tuberculata Gray représente
un mélange d’espéces (englobant Chalinolobus tuberculatus et Mystacina tub-
erculata). Par contre, si l’auteur est plutét G. Forster, le genre Mystacina créé
par Gray est basé sur une espéce-type mal identifiée et ““Mystacina tuberculata
Gray”’ n’existe pas. Dans ce cas, l’épithéte spécifique doit étre remplacée par
le prochain nom disponible, soit velutina Hutton, 1872 (dans le cas présent, il
s’agit du premier nom disponible). Cette option permet de préserver Il’ usage

VOLUME 112, NUMBER 3

471

courant du genre Mystacina en établissant que l’espéce-zoologique examiné
par Gray représente l’espéce-type. Nous recommandons de choisir cette option
pour que G. Forster soit a juste titre considéré comme |’ auteur de Vespertilio
tuberculatus, tel que stipulé dans l’article 50 du Code International de No-
menclature Zoologique. Par ailleurs, cette option permet de sélectionner un
spécimen type d’un lieu connu pour Mystacina velutina et de préserver la
classification infraspécifique actuelle de Mystacina ainsi que Il’ utilisation ha-
bituelle du genre Mystacina et de la famille Mystacinidae. Le seul nom valide
de la petite chauve-souris a queue courte de Nouvelle-Zélande est alors Mys-
tacina velutina Hutton, 1872, et nous désignons comme lectotype le spécimen
de Hutton, récolté a Hutt Valley (prés de Wellington, Ile du Nord, en Nouvelle-

Zélande).

New Zealand is home to but two endemic
land mammals, both of which are bats. Ac-
cording to current taxonomy (e.g., Koop-
man 1993) they are the vespertilionid Chal-
inolobus tuberculatus (Forster, 1844) and
mystacinid Mystacina tuberculata Gray,
1843. (A second Mystacina species, M. ro-
busta Dwyer, 1962, first described as a sub-
species of M. tuberculata, is thought to be
extinct—Daniel 1990.) Mystacina is of es-
pecial interest because of its broad ecolog-
ical niche, probable nearest relationship
with the American noctilionoids, and long
history of residence, if not also origin, in
New Zealand (Pierson et al. 1986, Hand et
al. 1998, Kirsch et al. 1998). In contrast,
Chalinolobus tuberculatus is clearly a close
relative of Australasian species belonging
to the same genus and subgenus (Daniel
1979, 1990; Koopman 1993, 1994).

The respective families of the two New
Zealand bats are widely separated in current
chiropteran classifications (Koopman 1994,
Kirsch et al. 1998, Simmons & Geisler
1998), yet for many years the two species
were confounded (Tomes 1857). While the
history of the specific classification of Mys-
tacina tuberculata has several times been
recounted, most recently by Hill & Daniel
(1985), we believe that the nomenclatural
conclusions drawn by most previous au-
thors are incomplete, if not in error. In fact,
the correct application of the name Mysta-
cina tuberculata is unclear, and it is thus

also unclear what the valid name is for the
lesser New Zealand short-tailed bat (here-
inafter, “the mystacine’’). Here we attempt
to clarify the history of the species’ no-
menclature, and to arrive at a resolution of
the difficulties encountered that will best
promote the stability and universality of no-
menclature. We will conclude that the spe-
cific name velutina Hutton, 1872 must be
adopted for the mystacine, and select the
zoological species before Gray as the type
species of Mystacina, so as to preserve the
genus and family names in their current
and, historically, sole applications.

An Historical Sketch

The first descriptions of the chalino-
lobe.—In May of 1773, at Queen Char-
lotte’s Sound, South Island, New Zealand,
Johann Reinhold Forster, naturalist on
Cook’s second voyage, shot and subse-
quently described, and his accompanying
son George drew (‘‘a me descriptus et a fi-
lio delineatus’’—Forster 1844:63), an ex-
emplar of the bat now called Chalinolobus
tuberculatus (hereinafter, ‘“‘the chalino-
lobe’’). In his journal, under the date of 21
May, Forster writes, “‘I shot a a new Shag,
never before described’’, and that, ‘‘[i]n the
morning I described the new Shag... &
George drew the Shag & a new Bat, which
we had got” (Hoare 1982:283); as Forster
used ship’s time (i.e., noon of the previous

472

day to noon of the date of entry) in dating
his entries, the drawings were made on the
morning of 21 May. There appears to be
some discrepancy concerning the dates of
collection and description of the shag:
Hoare (1982) identifies it as Pelecanus (=
Phalacrocorax) carunculatus, but Forster
(1844) proposed that name under the date
of 20 May, the apparent date of the shag’s
collection; neither Lysaght (1959) nor
Hoare (1982) question the date in Forster
(1844). However, the dating of events re-
lating to the bat is consistent among For-
Ster’s writings.

Despite an Admiralty ban on publication
urged by Cook (Quammen 1996), both
Forsters published early accounts of their
travels, in which allusions to the bat are
presented; neither Forster, however, men-
tions the bat by scientific name in their gen-
eral accounts of the 1772—1775 voyage. J.
R. Forster notes the resemblance of the
New Zealand bat to Pennant’s “‘New- York
bat”? (Forster 1778:130 in the 1996 reprint),
while George Forster (1777) is even more
casual in his Voyage, stating only that the
bat is one of five mammals found in New
Zealand.

It was not until nearly 50 years after his
death that J. R. Forster’s account of the zo-
ology of Cook’s second voyage, edited by
Lichtenstein, was published (Forster 1844).
In it the bat is described as Vespertilio tub-
erculatus under the entry for 22 May. As
we know from this description that the bat
lived two days after being shot in the wing,
20 May seems probable as the date of its
collection. Forster again refers to Pennant,
comparing the new species to “‘V. novebor-
acensi [s]”’, a synonym of Lasiurus boreal-
is. That the creature was regarded as a new
species of Vespertilio is unsurprising: at the
time of collection only two genera of bats
were recognized—WNoctilio Linnaeus, 1766
(placed by Linnaeus among Glires) and the
vastly heterogeneous Vespertilio Linnaeus,
1758 (then and until Erxleben [1777] in-
cluding even pteropodids, and placed with-
in Primates by Linnaeus). In a footnote on

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

page 62 of Forster (1844), added by the ed-
itor, comparison of Vespertilio tuberculatus
is made to species of Scotophilos [sic] and
Miniopterus. According to Whitehead
(1969), no surviving mammal specimens
can be traced from any of Cook’s voyages,
and so George’s drawing, now in the British
Museum (see below), is the only extant
non-written evidence concerning the first
bat collected in New Zealand.

While J. R. Forster’s description of the
bat was not published until 1844, George’s
illustration, with the name Vespertilio tub-
erculatus written on it, was known to Brit-
ish zoologists prior to posthumous publi-
cation of his father’s work, as, probably,
were some of both of the Forsters’ manu-
scripts (Whitehead 1978). On the basis of
this knowledge, a brief description of the
chalinolobe appeared in 1843 in an appen-
dix prepared by John Edward Gray for
Dieffenbach’s Travels in New Zealand. On
page 181 of volume two of Dieffenbach
(1843), the Forsters’ bat is characterized
thus:

Fam. VESPERTILIONID-.

1. Vespertilio tuberculatus. G. Forster. Icon. ined.,
1 em 8

Yellowish brown; ears small, rounded.
Inhab. Dusky Bay, New Zealand. G. Forster.

Hill & Daniel (1985), like Thomas (1905)
before them, were of the opinion that this
short characterization is sufficient to make
the name available, and we agree (Interna-
tional Commission on Zoological Nomen-
clature [ICZN] 1985:Art. 12a). As will be
seen, parts of this description are inaccu-
rate; however, a description need not be ac-
curate in order to make a name available.
Taken on its face, this passage seems to in-
dicate that G. Forster is the author; this 1m-
pression is reinforced when note is taken of
further passages in Dieffenbach in which
Gray clearly attributes to himself names he
(Gray) therein proposes. Hill & Daniel
(1985), however, attribute authorship to
Gray (in Dieffenbach 1843:181). Thomas

VOLUME 112, NUMBER 3

(1905:423) was a bit more equivocal in his
attribution, giving the full citation as “‘Gray
(ex Forst.)’’. Gray incontestably attributes
responsibility for the name and _ locality
(which, as we know from J. R. Forster
[1844], is incorrect) to George Forster. As
will become evident, the critical nomencla-
tural question is whether the whole of the
description is attributable to George, thus
establishing him as the one responsible both
for the name and for satisfying the criteria
that make it available (ICZN 1985:Art.
50a), or whether some part is attributable to
Gray.

George Forster’s drawing of the New
Zealand bat.—No illustrations, by George
or otherwise, appear in Forster (1844), so
that Hill & Daniel’s (1985) statements that
George Forster’s drawing was published in
1844, if intended to refer to the Lichtenstein
edition, are apparently in error. George’s
drawing of the bat has, to our knowledge,
been published only once, and then incom-
pletely, including just one of the two figures
on the drawing (Andrews 1986:28).
Through the courtesy of the authorities of
the British Museum (Natural History), we
have been able to examine the original in
London, and here publish it fully for, we
believe, the first time, as Fig. 1.

The drawing of the bat is number 1 of a
series of 271 zoological drawings from
Cook’s second voyage bound in two vol-
umes (Whitehead 1978). Although a few of
these drawings are by other artists, it is well
attested that the drawing of the bat is by
George (Forster 1844, Whitehead 1978,
Hoare 1982). In a catalogue of these draw-
ings (‘Catalogue B”’ of Whitehead 1978)
by one of Joseph Bank’s associates, prob-
ably Dryander (A. Wheeler, pers. comm.;
Whitehead [1969] agreed, but later [1978]
thought Solander was its author, although
this seems unlikely given Diment & Wheel-
er’s [1984] failure to discuss this catalogue),
the notation “‘Vespertilio tuberculatus, N.
Zel. Charlotte Sound” is made in reference
to the drawing. The recto of the drawing
has two views of the bat, one ventral, the

473

other lateral with wings folded. The draw-
ing is in pencil, and the bats have been col-
ored. The body of the upper bat is colored
dark brown and a lighter shade of brown.
The wings are gray. The lower bat is a more
uniform dark brown, and appears to be lit
from behind, with a shadow falling towards
the viewer. Figure 2 is a reproduction of a
sketch made from Forster’s drawing which
includes details not obvious in the repro-
duction in Fig. 1.

Below the bats, also in pencil, is written
‘““Vespertilio tuberculatus’”; in the upper
right is the number “1.’’, and in the lower
left “‘Geo Forster’, both of the latter in ink.
According to Whitehead (1978; see also
Lysaght 1959), scientific names were writ-
ten on the drawings by George or J. R. For-
ster, while George Forster’s name was add-
ed later by Dryander, it being uncertain by
whom and when the drawings were num-
bered (although certainly prior to 1843). On
the verso, ‘“‘New Zealand Charlotte’s
Sound” is written in pencil. Apparently dif-
ferent hands are responsible for the scien-
tific name, artist’s name, and locality.
Whitehead (1978) failed to record any writ-
ing on the verso of the bat drawing, but also
did not note that two figures are present, nor
the notation of locality in “Catalogue B”’.
It is possible that the verso locality was
added later, perhaps even after the first pub-
lication of the correct locality by J. R. For-
ster in 1844; Whitehead (1978) notes that a
number of later annotations were made to
various of the drawings, but we consider it
more: likely that Whitehead inadvertently
omitted this information than that the verso
locality was added after 1978. Wheeler
(pers. comm.) believes that the verso local-
ity may be in the hand of J. R. Forster; if
Whitehead’s (1978; see also Lysaght 1959)
attribution of the scientific names is to be
credited, then the binominal is in the hand
of George Forster.

The first description of the mystacine.—
The account of Vespertilio tuberculatus on
page 181 is not the only passage in Dief-
fenbach (1843) concerned with bats. In an

474 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

7 o, “2 oF
Cifrcs Sed at

. ’ © be - .
Ye. ida , t.S$eir, ive ane

Fig. 1. George Forster’s colored drawing of the bat obtained at Queen Charlotte’s Sound, South Island, New
Zealand, on or about 20 May 1773. Reproduced from a transparency, by permission of the authorities of the
British Museum (Natural History), a.k.a. The Natural History Museum, London. The insets are enlarged and
computer-enhanced to show more clearly some of the writing on the drawing.

VOLUME 112, NUMBER 3

475

Fig. 2. Pencil sketch of Forster’s drawing by JG showing details not evident in the reproduction in Fig. 1.
The shadowing in the lateral view of the bat has been omitted, and the distance between the two views reduced.

Scale line is 2 cm.

appended note on page 296, prepared after
earlier pages were already printed, Gray re-
turns to a consideration of the Chiroptera of
New Zealand, where he provides the first
description of a mystacine:

Vespertilio tuberculatus, p. 181.—I have just re-
ceived two specimens of this bat: it is a new genus,
differing from Embalonura [sic], Kuhl, and Uro-
cryptus, Temm., in having only two large cutting
teeth in the middle of the upper jaw; the fur is close,
erect, dark brown, with minute white tips to the
hair; the under surface is paler; the face has a series
of short, rigid, black bristles round the base of the
muzzle, the wings near the body and bones of the
limbs are thickened and tranversely grooved; the
tragus is elongate, subulate. It may be called Mys-
tacina tuberculata.—J. E. GRAY.

While this account makes it clear that the
two specimens before Gray were mysta-
cines (although, significantly, it does not
distinguish between the living species and
the then-extant M. robusta), it also dem-
onstrates that Gray believed he was describ-

ing further specimens of a bat already
known: the one described for the first time
on page 181. His intent on page 296 was
not to name a second species, but to point
out this previously known species’ generic
distinctness from Vespertilio, proposing for
it the new genus Mystacina. (Such a sepa-
ration from Vespertilio here is not auda-
cious: by 1843 over 40 genera of bats had
been described, many simply representing
subdivisions of the once-comprehensive
Vespertilio.)

There is only the slightest hint that Gray
in 1843 might have thought there to be
more than a single species of bat in New
Zealand. On page 182 of Dieffenbach, fol-
lowing the account of Vespertilio tubercu-
latus, Gray presents two quotations with
parenthetical and interpolated comments:

“The Pekdpekad, or Bats, and various small batlets,
are very common in the Island, but none of the
Vampire species. (Pteropus? or Glossophaga?) They

476

are among the smallest of the Australian spe-
cies.”’-—FPolack, i. 304. I am not aware that any of
these animals have reached Europe; they would be
interesting, and doubtless new. “There is, apparent-
ly, only one species; probably the one figured by
Forster.’ —Dieffenbach.

Hill & Daniel (1985) reproduce only the
second of the remarks quoted by Gray, as
evidence that he did not (then) believe in
the existence of additional New Zealand

species, yet Gray’s inclusion of the quota- .

tion from J. S. Polack (1838:304; impre-
cisely and incompletely transcribed by
Gray) might suggest otherwise. But in any
case, we reiterate there can be no doubt that
Gray considered his two new specimens to
be exemplars of the Forsters’ bat. Thus
Gray confounded two very different spe-
cies, with consequences to be explored be-
low.

Dieffenbach’s work, with the zoological
appendix entitled “‘Fauna of New Zea-
land’, was published in January 1843
(Sherborn 1932); later in 1843, Gray pub-
lished a List of the Specimens of Mammalia
in the Collection of the British Museum,
wherein he reiterated his belief in the iden-
tity of Mystacina tuberculata and Vesper-
tilio tuberculatus (Gray 1843a; the intro-
duction is dated “‘May-Day, 1843’, and a
copy in the library of the Field Museum of
Natural History bears the manuscript an-
notation ““May 13’, without a year).

Once more in 1843, Gray returned to
consider, at least briefly, the mystacine, in
his contribution to The Zoology of the Voy-
age of H.M.S. Sulphur (Gray 1843b). While
sometimes cited as the first description of
Mystacina (e.g., Dobson 1878, Miller
1907), this work undoubtedly appeared af-
ter Dieffenbach (1843) and Gray (1843a),
although, contrary to Hill & Daniel (1985),
apparently not as late as 1844. The Zoology
of the Sulphur was issued in parts from
1843 to 1845 (Zimmer 1926). A copy of
the third part (on birds) in original wrappers
in the library of the Field Museum bears
the date October 1843, while the fourth part
(also on birds), dated January 1844, has on

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

the wrappers an advertisement stating that
parts 1 and 2 on mammals and part 3 on
birds have already been published. A bound
copy of the first two parts (including the
‘““Summary of the Voyage’’ in part 1, as
well as the complete mammals) in the same
library has a ““Temporary Title’? page with
the date 1843. The section on mammals is
also cited as 1843 by earlier authors (e.g.,
Tomes 1857, Dobson 1878, Miller 1907).
The British Museum Catalogue’s (1903-—
1915) date of 1844, cited by Hill & Daniel
(1985), is simply in error; and there seems
to be no confusion of the Zoology of the
Sulphur and Captain Belcher’s general ac-
count, as suggested by Hill & Daniel
(1985). While thus appearing before 1844,
parts 1 and 2 of the Sulphur nonetheless
seem to have appeared after Gray’s List
(1843a; and thus also after Dieffenbach
1843), as in the main text of the List Gray
cites the Sulphur by unnumbered plates,
rather than by precise page and plate num-
bers as he does for his other works, and
states (page vii) that the Zoology of the Sul-
phur is “‘now in the course of publication.”
He does give precise plate references to the
Sulphur in a page (196) of ‘“‘Corrections”’
likely added to the List shortly before it was
published; in the Sulphur, by contrast, he
gives precise page references to his List
throughout. These facts indicate that the
Sulphur was in preparation but not yet com-
plete or published at the time of the writing
of Gray’s List (1843a).

Gray’s remarks in the Sulphur also give
no indication that they are intended to name
a taxon. In his account (page 22), Gray ex-
plicitly states that he is only mentioning
Mystacina and related genera “‘[flor the
purpose of showing the natural relations,
and the distinctive characters of the two
new genera [Mosia and Centurio] figured’’.
The entire account of Mystacina is but three
lines long, and read in context is clearly
synoptic rather than descriptive: ““MysTA-
CINA, Gray.—Nose rather produced, sur-
rounded at the base with a series of short
rigid bristles. Interfemoral membrane trun-

VOLUME 112, NUMBER 3

cated. Cutting teeth %, upper large. M. tub-
erculata, New Zealand.”’ Further, as Hill &
Daniel (1985) remark, the Sulphur did not
visit New Zealand, and no specimens of
Mystacina were described in the Zoology.

Subsequent recognition of two species.—
Gray (1843a), in his List, had restated his
belief in the identity of Mystacina tuber-
culata and Vespertilio tuberculatus. The
confounding of these two very different
species was not recognized until Tomes’
(1857) treatment of New Zealand bats,
which was based on specimens in the Col-
lege of Surgeons, British Museum, and Lei-
den collections. Therein, Tomes also reas-
signed Vespertilio tuberculatus to Scoto-
philus. Both New Zealand bat species were
figured by G. H. Ford for Tomes. In col-
oring, Ford’s plate of Vespertilio tubercu-
latus corresponds more closely to George’s
1773 drawing and the characterization in
Forster (1844), which Tomes accepts as the
first published description of the chalino-
lobe, than to the description given in Dief-
fenbach (1843:181). Ford’s lithograph is
clearly not a copy of Forster’s drawing, and,
contrary to Andrews (1986:64), there is no
indication that it is of the Forsters’ speci-
men: as noted earlier, all of the mammal
specimens from Cook’s voyages appear to
have been lost (Whitehead 1969).

While lamenting that the specific names
of Scotophilus tuberculatus and Mystacina
tuberculata were so similar, Tomes (1857:
135) realized that this was an irrelevant
consideration, as the bats were undoubtedly
different:

As the above-mentioned zoologists have certainly
been the first describers of two distinct animals, the
names imposed by them will of course be retained;
but it is much to be regretted that their specific
names are similar; and the more so, as the one most
recently given was clearly intended as a reference
to the earlier known species.

Thus Tomes accepted Gray’s description
and naming of the mystacine in Dieffen-
bach (1843) as valid, but either discounted
or (less probably) was unaware of the char-
acterization of the chalinolobe on page 181

477

of the same work, while also recognizing
that knowledge of the chalinolobe dates
from much earlier.

Hill & Daniel (1985) note the presence
of both species, properly labeled as such,
among material registered by Gray in 1844
(and sent in 1843 by Dr. FE Knox from Port
Nicholson [= Wellington]), which suggests
that Gray had come to recognize their dis-
tinctness on the basis of his own compari-
sons. Nonetheless, much later Gray (1875:
12b), in the Zoology of the Voyage of the
H.M.S. Erebus & Terror, acknowledged
Tomes’ analysis, and gave no indication
that he, Gray, had distinguished the bats by
himself:

I at first thought this was the little Bat named and
figured as Vespertilio tuberculata [sic] by Forster,
collected during Cook’s voyages, the drawings of
which are in the Banksian Library, British Museum,
and of which Lichtenstein published Forster’s MS.
descriptions and notes in 1844, but Mr. Tomes, who
has found in the British Museum two kinds of Bats
from New Zealand, believes that the one which is
a Scotophilus is the one which Forster described, on
account of the number of incisors he indicates, and
describes it as Scotophilus tuberculatus, P.Z.S..,
1857, 154 [sic: 135], pl. 43 [sic: 53], and he also
describes and figures the one that I have described
as Mystacina tuberculata, P.Z.S., 1857, 138, pl. 44
[sic: 54].

Gray’s comments here are accompanied by
a plate in which the mystacine is figured
(Gray 1875:plate 22, fig. 1; probably by
Waterhouse Hawkins, who did others in the
series). Although not published till 1875,
this plate was prepared much earlier (prob-
ably in 1844—-see Giinther [1875] and Hill
& Daniel [1985] on the publication history
of the Erebus & Terror), and had been seen
by Tomes (1857), but is unlikely to be of
the specimen figured in Ford’s drawing in
Tomes. The Erebus & Terror mystacine is
shown apparently in flight, and from the
dorsal aspect (presumably to show the em-
ballonurid-like penetration of the uropata-
gium by the tail), “‘flying’’ from the upper
left toward the lower right of the page, and
with its head thrown backwards—ceither be-
cause that was the case in the specimen or

478

in order better to display the face. However,
details of the patagia (as well as the position
of the head) are different from those in the
more hieratic dorsal view in Ford’s figure.

Later considerations of the chalino-
lobe.—Although the later nomenclatural
history of the chalinolobe is not entirely
without incident (Dobson [1878] confound-
ing it with the Australian species morio
Gray [Thomas 1889, 1905]), the details of
this history need not detain us. From the
point of view of our inquiries, the important
action is Peters’ (1867; as noted by a re-
viewer, though often cited as 1866, pages
657 ff. of this volume were published in
1867) proposal of the genus Chalinolobus,
type by monotypy Vespertilio tuberculatus.
Peters credits Forster with this binominal,
but neither specifies which Forster nor
gives any bibliographic citation. Fifteen
species are currently included in Chalino-
lobus, six in the nominate subgenus (Koop-
man 1993), and the stability of their names
is potentially affected by resolution of the
proper application of the species-group
name tuberculatus.

Later considerations of the mystacine.—
Subsequent authors have accepted the ex-
istence of two New Zealand bat species,
and have been more concerned with the pe-
culiar ecology and broader phylogenetic af-
finities of the mystacine (reviewed by Dan-
iel 1979, 1990; Kirsch et al. 1998) than
with nomenclature. Important exceptions,
however, are Hutton (1872), Thomas
(1905), and Hill & Daniel (1985).

Hutton (1872) briefly reviewed New
Zealand’s bats, and, like Tomes, regretting
the similarity of the specific names of the
chalinolobe and mystacine, proposed the
Species-group name velutina for the mys-
tacine; his comments were based on two
specimens of the mystacine then in the Co-
lonial Museum. Thomas (1905), having had
his attention called to the two passages on
bats in Dieffenbach (1843), which he had
previously overlooked, concluded that the
brief description of the chalinolobe under
the name V. tuberculatus on page 181 pre-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

empted Gray’s page 296 usage of the spe-
cific name for the mystacine, and therefore
Thomas adopted Hutton’s name velutina for
the mystacine. Thomas was likely influ-
enced by the fact that, although published
in the same work, and thus simultaneously,
the account on page 181 was clearly written
before that on page 296, and so held a sort
of temporal, as well as page, priority. (Odd-
ly, while Thomas credits Miller and Palmer
with alterting him to Dieffenbach, Miller
[1907] does not himself cite that publica-
tion in his account of the mystacine, and
uses the specific name tuberculatus for it.)

Both Thomas and Miller used the generic
name Mystacops Lydekker for the mysta-
cine. Mystacops had been proposed as a re-
placement name by Lydekker (in Flower &
Lydekker, 1891) in the belief that Mystaci-
na was preoccupied by Mystacina Boie,
1822 (a genus of birds). However, as point-
ed out by Simpson (1945) and Dwyer
(1962), Boie’s (1822) name was Mystaci-
nus, and the difference in ending is suffi-
cient to prevent homonymy (ICZN 1985:
Art. 56b).

Hill & Daniel (1985), in a study of geo-
graphic variation in the mystacine which in-
cluded a careful study of extant specimens
at the British Museum and a review of the
species’ nomenclature, differed with Thom-
as (1905). While accepting his conclusion
that the chalinolobe had been properly
named in Dieffenbach, they rejected his
conclusion that this necessitated the adop-
tion of velutina for the mystacine. They re-
jected this on two grounds: first, on the ir-
relevant basis that Thomas’ proposal had
not been generally adopted; and, second, on
the ground that no provisions of the Code
required such a change. As we shall see,
this latter point is problematic. The essence
of their analysis is that, in Dieffenbach,
Gray proposed two different species-group
names for two different zoological species.
But, as we have already seen in this histor-
ical review, and will explore further below,
such was certainly not the case.

VOLUME 112, NUMBER 3

Analysis

Two species-group names or one?—The
two bat descriptions in Dieffenbach—that
on page 181 relating to George Forster’s
drawing, and the one on page 296 referring
to the two mystacine specimens—create a
nomenclatural problem, as was first appre-
ciated by Thomas (1905). Are two species-
group names proposed in Dieffenbach or
only one? The preceding historical sketch
makes it certain that Gray regarded both the
bat illustrated by George Forster and the
two specimens Gray later received as mem-
bers of the same species. Thomas (1905:
423) also saw this clearly, noting that when
Gray discussed the mystacine on page 296
he did so “distinctly stating his opinion that
it was the same bat’’ as was mentioned on
page 181. On page 296, Gray had no inten-
tion of naming a second species, and in fact
did not do so. We can thus state categori-
cally that only one specific name was pro-
posed in Dieffenbach (1843).

Given that but one name is proposed,
much depends on who we suppose the au-
thor of that name to be. If it be Gray, as
Hill & Daniel (1985), and, somewhat less
emphatically, Thomas (1905) concluded,
then Mystacina tuberculata is composite
(the syntypical series including a chalino-
lobe and two mystacines), and the name
must be fixed on one part or the other. If,
on the other hand, the author is George For-
ster, as Sherborn (1931), definitely, and
Dwyer (1962), apparently, concluded, then
the name refers unequivocally to the cha-
linolobe, and the mystacine requires a
name. A further consequence of G. For-
ster’s authorship would be that the genus
Mystacina would then be based on a mis-
identified type species (ICZN 1985:Art.
70b), as Gray mistakenly believed his two
mystacines to be conspecific with the chal-
inolobe in George’s drawing.

The question of authorship hinges upon
who provided the name and the conditions
that make it available (ICZN 1985:Art.
50a), and this in turn depends on the inter-

479

pretation of the passage on page 181 of
Dieffenbach. In order for authorship of a
name to be attributed to someone other than
the author of the work in which it appears,
the Code requires that the evidence for this
attribution be “‘clear from the contents of
the publication” (ICZN 1985:Art. 50a). A
straightforward reading of page 181 seems
to indicate that this requirement is met:
Gray is attributing the name and the con-
ditions that make it available to G. Forster.
It is undeniable that Gray attributes both the
name and the locality to George, and the
name appears on George’s drawing, likely
in his own hand, and thus George surely has
provided at least one of the two elements
that the Code requires; but, it might be con-
tended that Gray has provided the descrip-
tion that makes the name available. The
question is, in short, who wrote the five
words of description on page 181?

Whose words are they ?—That Gray cites
G. Forster on two of the lines on page 181
could mean that the intervening line con-
taining the five words was his (i.e., Gray’s);
but it could equally plausibly mean that
Gray merely wanted to make clear the
source of the name, as distinct from the oth-
er information, which he also attributed to
George. Gray need not have put Forster’s
name at the end of every line in order to
make an attribution to George. Indeed, in
his account in Dieffenbach (1843) of the
‘““Sea Bear’ (Arctocephalus ursinus [Lin-
naeus, 1758], now Arctocephalus forsteri),
Gray gives information which comes di-
rectly from the Forsters (which we know
because the Forsters’ notes survive on
George’s drawing or in “Catalogue B’’—
Whitehead 1978) without immediate attri-
bution, showing that he did not always put
Forster’s name after every line derived from
him. In the account of the bat, Gray was, if
anything, more careful and explicit in citing
George twice, perhaps because a newly pro-
posed name was involved, whereas the
‘Sea Bear’ had been named previously.

Gray does himself name and describe
several new species in Dieffenbach, but

480

when doing so follows the new species’
name with “‘Gray, n.s.”’, then the locality,
and finally the description; this is not the
format he follows on page 181, in which
the words of description are enclosed with-
in attributions to G. Forster. Since the only
plausible interpretation of Gray as the au-
thor would have him describing George’s
drawing, it is significant that one of the new
species names proposed by Gray (Balaena
antipod[ar|um, now Eubalaena australis) is
based on the description of a drawing, and
that Gray is quite explicit about that basis
(page 184): ‘“‘The above short description of
this species is taken from a very good
drawing ...’’. No such statement of deri-
vation accompanies the account of the bat.

The description might credibly be attri-
buted to Gray if it consisted of a character-
ization of George’s drawing. The fact that
the locality given on George’s drawing does
not match that cited as from G. Forster on
page 181 suggests that Gray was working
from some written notes by George, and not
just the drawing. Perhaps even more per-
suasive evidence that Gray was not working
from the drawing alone is that its color does
not correspond to that given in Dieffenbach
(“Yellowish brown’’; cf. Fig. 1). These
notes could not have been J. R. Forster’s
manuscript descriptions (eventually pub-
lished in 1844), because the latter give a
locality (“in estuario reginae Charlottae”’
[Queen Charlotte’s Sound]—Forster 1844:
63) and color description (“‘Alae . . . fuscae,
.... Vellus ubique ... fusco-ferrugineum”’
[Wings.. .;..dusky,.......3 Pur:,dark-rusty
brown all over]|—Forster 1844:64) different
from those on page 181 of Dieffenbach,
and, in any event, Forster’s manuscript had
long been in Berlin by the time Gray was
writing. Unless he was simply making
things up, Gray must have been working
from written material by George, because
the source is clearly neither the drawing nor
J. R.’s manuscript. That some of the Fors-
ters’ manuscripts have been lost is undeni-
able: Whitehead (1978) records that the
Forsters furnished information to Joseph

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Banks, the documents of which no longer
survive, and George’s extant journal ends
at 11 May, eight days before the Resolution
reached Queen Charlotte’s Sound (Hoare
1982).

How might George have been the source
of erroneous locality and color information?
Dusky Bay was visited by the Resolution,
and it figures prominently in George’s ex-
tant journal (Whitehead 1978); it is also, as
H. Spencer (in litt.) has pointed out to us,
the locality of the “‘Sea Bear’’, the subject
of the second of George Forster’s drawings,
on the verso of which the locality is written,
and Gray (in Dieffenbach 1843:182) did
cite this drawing in his account. Gray may
just have made a hasty inference or tran-
scription error. As regards the color, notes
from the Forsters in “‘Catalogue B’’, repro-
duced in Whitehead (1978), make it clear
that their remarks on coloring often con-
sisted of critiques of the drawings; thus
“Yellowish brown’? could be an admoni-
tion for modulation of the applied color,
rather than a description. Gray, never hav-
ing seen the bat, could not have known the
import of such a note.

Absent written evidence, we cannot now
know for sure whether the five words of
description are Gray’s or George’s but giv-
en the citation of G. Forster, both following
the name and at the account’s conclusion,
it was clearly Gray’s intent to attribute the
name and description, and thus authorship
of Vespertilio tuberculatus, to the younger
Forster. This conclusion is strengthened by
looking at Gray’s List published a few
months later (1843a:34), which reads ““The
MYSTACINE. Mystacina tuberculata, Gray,
Dieffenb. Jour. App. 296. Vespertilio tub-
erculatus, G. Forster, Icon. ined. in Brit.
Mus. t. 1.” As is clear from the form of
citation Gray uses in his List, this passage
indicates that Gray attributes to himself the
combination Mystacina tuberculata, while
he attributes the specific name tuberculatus
to G. Forster (which also demonstrates, as
we believe is patent from Gray in Dieffen-
bach as well, and as Sherborn [1931] also

VOLUME 112, NUMBER 3

indicated, that Gray did not consider him-
self to have proposed a new species tuber-
culata in the genus Mystacina, but rather to
have placed Forster’s species V. tubercula-
tus in the new genus Mystacina). It is clear
that the name came from G. Forster, that all
knowledge Gray had of the species came
from Forster’s work (the drawing and per-
haps manuscripts), and that Gray attributed
the name and the description to Forster. We
should not cast aside these certainties as to
the source of the name, knowledge of the
species, and Gray’s own attribution.

Thus, it is our judgment that the descrip-
tion is more likely to be George’s than
Gray’s—and therefore that George not only
named the bat, but also satisfied the con-
ditions that make that name available. All
the internal evidence of the contents of the
publication—of Gray’s attributions, cita-
tions, and format—indicates that, under Ar-
ticle 50a, authorship should be attributed to
George Forster. The external evidence—the
lack of correspondence between the draw-
ing and the published account, Gray’s later
publications—even if it were admissible
under Article 50a (which it is not), also fails
to support the contention that Gray is the
author. The name is thus properly cited as
Vespertilio tuberculatus G. Forster in Dief-
fenbach, 1843:181.

The consequences of G. Forster as au-
thor.—On the interpretation that the de-
scription as well as the name on page 181
are George Forster’s, and therefore that he
and not Gray made that name available, an
inescapable conclusion is that there is no
such name as ‘“‘Mystacina tuberculata Gray
in Dieffenbach, 1843’’; there is only Mys-
tacina tuberculata (G. Forster in Dieffen-
bach, 1843) sensu Gray, 1843 (ICZN 1985:
Art. 51b(i)). Gray confused the two speci-
mens before him, and thus also the type
species of Mystacina, with Forster’s Ves-
pertilio tuberculatus. This leaves the bats
before Gray without a species-group name,
and the genus Mystacina with a misidenti-
fied type species. We consider these two is-
sues in turn.

481

The species-group name of the mysta-
cine.—**Mystacina tuberculata Gray in
Dieffenbach, 1843:296’’, being a misiden-
tified type species, rather than the proposal
of a new species, cannot possibly be the
name for this bat. Gray did use this com-
bination in several other publications (e.g.,
1843a, 1843b), and it might be argued that
these accounts make the name tuberculata
available in Mystacina as of these publica-
tions. The Code, however, explicitly pro-
hibits this: Article 49 states that “‘[a] pre-
viously established species-group name
wrongly used to denote a species-group tax-
on because of misidentification cannot be
used for that taxon even if it and the taxon
to which the name correctly applies are in,
or are later assigned to, different genera

The valid species-group name of the
mystacine must thus be the next available
name, which, in this case, is actually the
first available name (ICZN 1985:Art. 23e):
velutina Hutton, 1872. This name was in-
troduced by Hutton to replace “‘tuberculata
Gray’’. It is not, however, strictly speaking,
a replacement name in the sense of the
Code. A replacement name is proposed to
replace an available name, but “‘tuberculata
Gray” is not an available name, and was
not intended to be one by Gray. Thus Ar-
ticle 72e, on the typification of replacement
names, does not apply. Article 72b(ii) and
(iv), on the type series of names based in
whole or part on previously published mis-
identifications or unavailable names, does
apply, and thus the type series of Mystacina
velutina consists of all specimens before
Hutton, plus Gray’s two specimens.

Gray’s specimens cannot now be traced
(see below), and, given the recognized geo-
graphic variation in the lesser mystacine,
and the existence in the 1840s of a second
species of mystacine (M. robusta) in the en-
virons of the South Island, their lack of spe-
cific locality data makes them a poor choice
to be name bearers. The current where-
abouts of the two specimens examined by
Hutton in 1871 are also not known. They

482

are not in the National Museum in Welling-
ton; the earliest extant bats currently in that
collection were presented in 1877, virtually
all the earlier collection having been de-
stroyed by damp and mold in the 1880s—
1890s (C. Paulin, in litt.). It was the practice
of the Colonial Museum to send type spec-
imens from New Zealand to the British Mu-
seum, but such specimens were often not
clearly labeled as such, and may have been
accessioned into the British Museum col-
lection under some other name, and without
their nomenclatural importance being noted
(C. Paulin, in litt.). Dobson (1878) records
two specimens in the British Museum, “‘e”’
and “‘f’’ of his list, from “‘“Wellington, New
Zealand”’ received from the Colonial Mu-
seum. Hill & Daniel (1985) note that only
one of these two was registered, in 1876:
BM(NH) 76.4.8.1. It seems quite possible
that Dobson’s “e” and “f’’? (ncluding
BM(NH) 76.4.8.1) were those before Hut-
ton from the Colonial Museum (‘“‘Welling-
ton’’ recording from where they were sent,
rather than the exact locality), but we can-
not now be certain of this.

Although they cannot be traced with cer-
tainty, Hutton’s specimens do have exact lo-
calities: “the Hutt Valley, near Wellington’”’
and “Milford Sound, on the southwest
coast of the South Island.”’ This is fortu-
nate, as Hill & Daniel (1985:290) have
based their subspecific arrangement on a
conception of the nominate form as occur-
ring near Wellington. We thus ratify and
preserve the nomenclatural stability of their
arrangement by selecting as lectotype of M.
velutina Hutton, 1872 the specimen from
the Hutt Valley, near Wellington (ICZN
1985:Art. 74c). Being on the North Island,
this locality also excludes the possibility of
confusion with the extinct M. robusta, as
Wellington is outside its historic range
(Daniel 1990).

It is worthwhile noting here that the use
of velutina is in no way in violation of the
presumptive statute of limitation which
gives the benefit of the doubt to stability
when it conflicts with priority ICZN 1985:

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Art. 79c). This provision applies to the pro-
posed substitution of an unused senior syn-
onym for a junior synonym, not the substi-
tution of an available name for an unavail-
able name. Nor does it violate the stronger
injunction against the displacement of
names in the fourth edition of the Code
(ICZN 1999), which again does not apply
to unavailable names; and in any event, ve-
lutina has been used as a valid specific
name after 1899 (Minelli & Ride 1997).
The generic name of the mystacines.—In
the following, we apply the provisions of
the fourth edition of the Code (ICZN 1999),
on the recommendation of Dr. P. Tubbs, Ex-
ecutive Secretary of the ICZN, to use these
provisions during the interregnum between
publication of the new edition and its ef-
fectuation, as during this time action by the
Commission under Article 70 of the third
edition of the Code is no longer possible or,
indeed, necessary under the new edition.
Explicit provision is made for cases where
an author erects a new genus on the basis
of specimens to which he has applied the
wrong specific name, the applicable section
in the fourth edition being that on misiden-
tified type species in Article 70. The article
lays out two courses of action which an au-
thor discovering such a case may take: ei-
ther to let the species mentioned by the au-
thor of the genus, regardless of misidenti-
fication, remain as type species of the genus
(i.e., Vespertilio tuberculatus G. Forster
would be the type species of Mystacina);
or, to designate the species actually before
the author of the new genus as the type spe-
cies. The principle that we believe should
guide action in this case is that types of taxa
are zoological objects, not names (Mayr
1969). Thus we believe that the species ac-
tually before Gray should be the type of
Mystacina. We have already noted that
Gray’s description, while diagnostic to ge-
nus, does not exclude the possibility that he
may have had a specimen of M. robusta,
and the locality, ‘““New Zealand’’, ascribed
to Gray’s specimens in his List does not
allow us to exclude M. robusta on geo-

VOLUME 112, NUMBER 3

graphic grounds, either. However, in histor-
ic times the lesser mystacine (M. tubercu-
lata auctorum) has been more widespread
and abundant than M. robusta, and no spec-
imens of the latter are certainly known to
have been collected in the 19th century. In
addition, the specimens provided by Sir
George Grey, which we believe may have
been Gray’s specimens (see below), are of
the lesser mystacine. We are thus confident
that at least one of the two specimens be-
fore Gray was of the lesser mystacine, and
this species, of which we have shown ve-
lutina Hutton, 1872 to be the valid species-
group name, should be the type species of
Mystacina.

This action would also promote the sta-
bility and universality of nomenclature.
Were the misidentified species to remain the
type species of the genus, it would disas-
sociate Mystacina Gray, 1843 from all zoo-
logical species with which it has ever been
associated; make Mystacina Gray, 1843, by
virtue of its becoming a senior objective
synonym of Chalinolobus Peters, 1867, the
valid generic name for the chalinolobe and
its fourteen congeners, forcing a change in
the names of these species; make Mystacin-
idae a family name in a section of the Mi-
crochiroptera distant from its usual appli-
cation in or near the Noctilionoidea; and
leave the mystacines without a generic or
family name, leading to further nomencla-
tural changes. (Mystacops Lydekker, 1891,
and the family name based on it, Mysta-
copidae Miller, 1907, do not help here, in-
asmuch as Mystacops, being a replacement
name sensu stricto, takes the same type spe-
cies as Mystacina, and would thus also ap-
ply to the chalinolobe—ICZN 1985:Art.
67h.)

In contrast, having the species actually
before Gray as type species of the genus
would allow the name Mystacina to contin-
ue to be the valid generic name for the en-
demic noctilionoid bats of New Zealand, as
it has been assumed to be since 1843; allow
Mystacinidae to continue in its familiar ap-
plication; and lead to no changes in the no-

483

menclature of the chalinolobe and its con-
geners. We thus hereby designate the zoo-
logical species we believe to have been be-
fore Gray, the valid specific name of which
is velutina Hutton, 1872, as the type species
of Mystacina Gray, 1843. Should it later be
shown that both of Gray’s specimens were
M. robusta—an eventuality we consider
highly improbable—then M. robusta
Dwyer, 1962, as the only zoological species
before Gray, would be the type species of
Mystacina, and familiar usages of the ge-
neric and familial names would again be
preserved.

What if Gray were the author?—Though
we believe that a preponderance of the ev-
idence, including, most trenchantly, the
contents of Dieffenbach (1843), leads to the
conclusion that George Forster is the author
of the description of the chalinolobe, we ac-
knowledge the possibility that it might be
Gray. It is therefore useful to explore the
implications of this assignment of author-
ship. We believe that these implications,
though not admissible in deciding author-
ship, offer additional justification for our
choice in favor of George.

On the interpretation that Gray is the au-
thor, Mystacina tuberculata is a composite
species because Gray considered the Fors-
ters’ bat and the two later-received speci-
mens to be conspecific, thus forming a
compound syntypical series (ICZN 1985:
Art. 72b); although the two mystacines
were seen later by Gray, publication on the
three specimens (i.e., the mystacines and
the Forsters’ bat), because in the same
work, was simultaneous. In this case, Mys-
tacina tuberculata must be settled on one
or the other part of Gray’s composite series.

We imagine that most zoologists would
agree that the (or a) mystacine would be the
logical choice to bear the name, as this was
the zoological species before Gray. But be-
fore we can decide this issue, we should
consider the implications of the two alter-
natives, and whether a previous author may
have already settled the issue. There are two
candidates: Dobson (1878), who, apparent-

484

ly inadvertently, attempted to fix the name
tuberculata on the mystacine; and Thomas
(1905), who deliberately chose the chali-
nolobe.

Dobson’s view.—Dobson (1878:445), al-
though not at all appreciating the nomen-
clatural conundrum (he in fact seems to
have been unaware of Dieffenbach [1843]),
comes near to settling the issue by referring
to specimen “‘a”’ in his list of specimens of
Mystacina tuberculata as the “‘type’’. Were
this one of the two specimens which Gray
had mentioned in Dieffenbach (presumably
“a” and “b” in Gray’s 1843 List), this
would be a valid designation of a lectotype
(ICZN 1985:Art. 74a), and the name would
be fixed on the mystacine. It is unlikely,
however, that Dobson’s ‘‘a’”’ was one of
Gray’s specimens. Dobson further states
that ‘‘a’’ is the specimen figured in The Zo-
ology of the Erebus & Terror. In the text
accompanying this figure, Gray (1875:12b;
see quotation above) does not identify the
specimen figured, and certainly does not in-
dicate that it is part of his original series.
Because much, if not all, of Gray’s mysta-
cine material had been turned over to
Tomes for the latter’s use in writing his
1857 paper, and was seemingly not returned
to the British Museum until 1905 (Hill &
Daniel 1985), it may be that Dobson never
viewed all of the British Museum speci-
mens; and Gray was dead by the time Dob-
son’s catalogue appeared. Thus it is dis-
tinctly possible that Dobson’s statements
are merely conjectural.

Hill & Daniel (1985) have looked care-
fully into the question of the identity and
location of Gray’s specimens, examining
the accession registers and specimen labels,
but unfortunately they have not been able
to ascertain with certainty which, if any, of
the specimens currently present in the Brit-
ish Museum are Gray’s two bats, nor the
identity of Dobson’s specimen ‘a’. They
did, however, uncover evidence which ap-
pears to rule out Dobson’s specimen “‘a”’
from being one of Gray’s specimens, thus
invalidating it as a lectotype. Thomas an-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ee 99

notated specimen ‘a’? in a copy of Dob-
son’s Catalogue at the British Museum with
the following note: ““Dr. E J. Knox(P).—
see Trans. N.Z. Inst. IV, p. 186, 1871” (Hill
& Daniel 1985:286). If Thomas’s annota-
tion is correct, then specimen “a’’ cannot
be the type, as Knox (1872) states that he
sent his shipment to the British Museum in
or following July 1843, after Gray had al-
ready published in Dieffenbach. Perhaps
Knox, writing in 1871, misrecalled what he
had done almost 30 years earlier, in which
case his specimen could have been before
Gray in 1842, but there is no independent
evidence of this, and Knox in his 1872 ar-
ticle refers to notes he had written in 1843
or earlier. Hill & Daniel (1985) have sug-
gested that BM(NH) 44.10.29.7, sent by
Knox, may be Dobson’s specimen “‘a’’, but
the date of accession (29 October 1844)
lends no support to the notion that it might
have been before Gray in 1842. The only
thing about which we can be certain is that
for Dobson’s “‘a’”’ to be the type, one or
more authorities must be mistaken about
the identity of the specimen and/or the cir-
cumstances surrounding it.

Given that a number of errors in speci-
men identification and labeling have been
discovered in Dobson’s Catalogue (Thomas
1905), we are reluctant to assume that Dob-
son was correct in this case, as no support-
ing evidence in Gray’s publications, the ac-
cession registers, specimen labels, or
Thomas’s subsequent inquiries, is forth-
coming; and, if Thomas’s annotation is cor-
rect, specimen ‘“‘a’’ cannot be the type. Hill
& Daniel (1985) consider other possibilities
for the types, but do not seriously entertain
that a specimen (‘‘c’? of Dobson’s Cata-
logue) from 1842 provided by Sir George
Grey might be one of the type series (even
though Gray [1843a] specifically mentions
Grey’s collection), perhaps because it is not
an adult and Hill & Daniel’s subspecies-lev-
el taxonomy depends heavily on size dif-
ferences. The accession number of Grey’s
specimen indicates that it had been at the
Museum since at least February, although

VOLUME 112, NUMBER 3

Gray was unaware of it when writing the
main body of his contribution to Dieffen-
bach (1843), which is dated “‘15th August,
1842” on page 181. We do not now know
if Gray became aware of the specimen be-
tween this date and the publication of Dief-
fenbach, so that Grey’s specimen could
have been one of the two specimens re-
ferred to on page 296. Sir George Grey also
provided a second still-extant specimen (la-
beled “‘c*’’—Hill & Daniel 1985), cata-
logued in 1849; if this specimen was a du-
plicate from 1842 catalogued later, Grey
may well have provided both of Gray’s
bats.

W. D. L. Ride (in litt.) has suggested that
Dobson’s (1878) specimen “‘b’’, attributed
by Dobson to Captain Belcher of the Sul-
phur, may have been one of Gray’s speci-
mens. Although the Sulphur did not call at
New Zealand, and Gray (1843b) does not
refer to any mystacine specimens, Ride
notes that Belcher was an avid collector,
and could have received a specimen from
his assistant surgeon Andrew Sinclair. Sin-
clair left the Sulphur voyage early to return
to England via New Zealand, and collected
at the Bay of Islands in 1841, where he may
have obtained mystacine specimens. That
Knox, Grey, and Belcher (through Sinclair)
can all be argued to have provided Gray’s
bats shows how difficult it is to identify
their source or possible current where-
abouts.

Hill & Daniel (1985) ultimately conclud-
ed that they could not identify the types,
and based their conception of the nominate
form on BM(NH) 44.10.29.7, a specimen
which, if Knox is to be believed, could not
possibly be a type.

A further problem with all of the candi-
dates is that none has exact locality data.
This is a serious shortcoming, because Hill
& Daniel (1985) base their subspecific tax-
onomy of M. tuberculata on the supposition
that the nominate form occurs in the Wel-
lington area. As noted earlier, it is even con-
ceivable that Gray’s specimens might have
included M. robusta. At least some early

485

specimens, including perhaps Gray’s, came
from the South Island and vicinity, and thus
within or near the historic range of the now-
extinct M. robusta; both Hutton and Knox
mention a specimen (probably the same
one) from as far south as Milford Sound,
bats at least occasionally being caught in
the rigging of ships plying the South Island
coast. Ascribing authorship to Gray and fix-
ing the name on the mystacine thus results
not only in the name having a vague local-
ity, but also injects some uncertainty into
its specific identity.

Fixing tuberculata on the mystacine
would give authorship of Vespertilio tub-
erculatus for the chalinolobe to J. R. Forster
(1844), although there is some question as
to whether tuberculatus might be preoccu-
pied in Vespertilio by Gray’s usage of it in
Dieffenbach (1843). The Code is not ex-
plicit on this point, but the implication of
Article 51c(ii) seems to be that since the
proposal of Mystacina was not conditional,
then tuberculatus is not preoccupied. J. R.
Forster is cited as the author (“‘Forster,
1844’’) in Koopman’s authoritative list of
1993 (but not in Koopman’s 1994 mono-
graph, prepared earlier than the 1993 list,
where he attributes the names of both New
Zealand bats to Gray, 1843, apparently ac-
cepting the judgments of Hill & Daniel
[1985]).

In short, accepting Gray as the author
and settling the name of the resulting com-
posite on the mystacine leaves us without a
type specimen or type locality, and thus
also even without unequivocal correspon-
dence of the name with the extant mysta-
cine species, though it has a superficial ap-
peal in that it would preserve one version
of current usage and attribution for both the
mystacine and chalinolobe.

Thomas’s view.—Of all authors who
have considered the mystacine problem,
Thomas (1905) came closest to understand-
ing the situation. He realized that both New
Zealand bats had been discussed in Dief-
fenbach (1843), but that only one species-
group name had been proposed, so that the

486

name could apply to only one or the other.
He apparently regarded the description of
Vespertilio tuberculatus on page 181 of
Dieffenbach as originating with Gray, and
therefore that the same author’s later de-
scription of Mystacina tuberculata on page
296 was invalidated by page priority. He
thus settled what he took to be Gray’s spe-
cific name on the chalinolobe. Thomas’s
claim to have acted as first reviser and set-
tled the question is strengthened by his hav-
ing clearly been the first to realize that a
choice between two courses of action was
necessary, and to have made one; and his
mention of Forster’s figure, and citation of
Gray’s name as “‘ex. Forst.’’, comes close
to (although, because he did not use the
word “‘type’’, not quite) designating a lec-
totype. It is clear that he intended to restrict
application of tuberculatus to the chalino-
lobe, and may have done so according to
the nomenclatural standards then prevail-
ing, even if his actions are not conclusive
under today’s Code.

While appreciating Thomas’s insight, we
hasten to add that, unless we are willing to
consider that an author (in this case Gray)
may have misidentified a type species
which he himself has authored in the very
same publication (a possibility whose illog-
ic makes us loath to contemplate it), Thom-
as’s restriction leads necessarily to all of the
unfortunate consequences enumerated
above which flow from having the chali-
nolobe as the type species of Mystacina.
Thomas overcame these difficulties, in part,
by using Mystacops for the mystacine,
which may have sufficed under nomencla-
tural rules as then understood; but, as we
have already shown, under the current
Code, Mystacops must always be an objec-
tive synonym of Mystacina, and thus the
unpleasant consequences still follow.

What is to be done?

We believe the weight of the evidence
indicates that the description of the chali-
nolobe on page 181 of Dieffenbach (1843),

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

and hence authorship of Vespertilio tuber-
culatus, should be attributed to George For-
ster. It is clear from the contents and format
of Dieffenbach (1843) that Gray was attri-
buting the species’ name and description to
another, and thus that “‘some other person
is alone responsible both for the name and
for satisfying the criteria of availability”
(ICZN 1985:Art. 50a)—that other person
being George Forster. Gray never subse-
quently explicitly attributed the name to
himself. Attempts to attribute the conditions
of availability to Gray by appealing to the
contention that he was describing George’s
drawing fail on the ground that the descrip-
tion does not correspond to the drawing,
and also run afoul of Article 50a’s specific
admonition that such questions of author-
ship are to be determined by the “‘contents
of the publication’’, and not by reference to
external evidence. We are not entirely alone
in honoring George Forster: Sherborn
(1931:6670) cites the name as “G. Forster
in J. E. Gray in Dieffenbach, Trav. N. Z. II.
1843, 181.’ Dwyer (1962) also appears to
accept George as author, for he attributes
the name to G. Forster in his first mention
of the chalinolobe (and elsewhere just to
‘‘Forster’’). Having once settled authorship
upon George Forster, all else follows un-
ambiguously: all species-group names have
a firm basis in either extant types (robusta,
aupourica, rhyacobia) or well-established
type localities (tuberculatus, velutina)
which are in harmony with the current sub-
specific and specific arrangement, insuring
the nomenclatural stability of these names
into the indefinite future; the names Mys-
tacina and Mystacinidae continue in their
familiar applications; and no changes of
name are required in the genus Chalinolo-
bus. The only drawback of this decision is
that it requires use of the specific name ve-
lutina Hutton for the lesser mystacine,
which has not been used since earlier in this
century.

The other route, supposing Gray to be the
author, makes us choose between Dobson
and Thomas: it leads to a chain of uncer-
tainties and difficulties, concerning the

VOLUME 112, NUMBER 3

specimens before Gray, and whether any of
these are still extant; whether Dobson’s sup-
posed type was one of these specimens, and
thus if he had fixed the specific name on
the mystacine; whether Thomas’s annota-
tions to Dobson’s catalogue are accurate;
whether Knox in 1871 correctly recalled the
circumstances of his shipment of specimens
to the British Museum almost 30 years ear-
lier; and whether Thomas validly settled
tuberculatus on the chalinolobe. Although
by judicious resolution of these several un-
certainties we might retain tuberculatus(a)
as the specific name of both extant New
Zealand bats, we might also lose some type
localities; upset the subspecific classifica-
tion of Mystacina, even introducing uncer-
tainty into its specific classification; make
Chalinolobus an objective synonym of
Mystacina; transfer the latter name to a seg-
ment of the Chiroptera phylogenetically
distant from that to which the name has
long applied; leave the mystacines without
generic or family names, thus requiring the
proposal of new names; and, unlike our
proposed course of action, might well re-
quire exercise of the plenary power by the
Commission. And, whatever resolution we
might come to, it would be based on a se-
ries of doubtful inferences: a compounding
of uncertainties, a multiplication of improb-
abilities. We thus prefer to resolve the ini-
tial uncertainty in favor of G. Forster, as we
believe that this is not only correct and his-
torically true, but will do the most to pro-
mote the stability and universality of no-
menclature.

Synonymy.—Adopting the course here
recommended leads to the following syn-
onymies of the names concerned. For the
chalinolobe, only the specific synonymy is
given, as the generic synonymy would lead
us far astray from New Zealand. For the
mystacines, familial, generic, specific, and
subspecific synonymies are given. They are
complete, we believe, as regards synonyms
and combinations, but only a few of the
more salient citations to each name or com-
bination are given. Further citations may be
found in the references in Hill & Daniel

487

(1985), Daniel (1979, 1990), Hand et al.
(1998), and Kirsch et al. (1998).

Chalinolobus tuberculatus (G. Forster)

Vespertilio tuberculatus G. Forster in Dief-
fenbach, 1843:181. “Dusky Bay, New
Zealand’’, in error; correctly given as “‘in
estuario reginae Charlottae’? (= Queen
Charlotte’s Sound, South Island, New
Zealand) by J. R. Forster, 1844:63. Type
lost (Whitehead 1969); drawing of type
by George Forster in British Museum
(Natural History).

Mystacina tuberculata.—Gray, 1843a:34
(part).

Scotophilus tuberculatus.—Tomes, 1857:
135; Hutton, 1872.

Chalinolobus tuberculatus.—Peters, 1867:
680; Dobson, 1878 (part); Thomas, 1905;
Dwyer, 1962; Hill & Daniel, 1985.

Miniopteris [sic] morio.—Gray, 1875:12a
(?part). Although Gray states ‘“Inhab.
Australia’, Hill & Daniel (1985:288),
following unpublished notes by Thomas,
suggest that the specimen illustrated (as
Scotophilus morio; see next entry) was
from New Zealand.

Scotophilus morio.—Gray, 1875:plate 19,
fig. 2 (?part). Placed in the genus Min-
iopteris [sic] in the text; supposed by Hill
& Daniel (1985) to be of a specimen
from New Zealand (see previous entry).
The plate may have been prepared as ear-
ly as 1844 (Tomes 1857), and thus rep-
resents Gray’s earlier opinion concerning
the species’ generic position.

Chalinolobus morio.—TYhomas, 1889:462
(part); Flower & Lydekker, 1891 (part).

Chalinolobus tumorio Flower & Lydekker,
1891:671 (part; apparently a lapsus for
Chalinolobus morio, but perhaps arising
from an incomplete striking of “‘tuber-
culatus’’, which was overwritten by
“‘morio’’; not an available name).

Mystacinidae Dobson

Mystacinae Dobson, 1875:349. Type Mys-
tacina Gray. Proposed as a “Group” (=

488

tribe, and thus a family-group name
[ICZN 1985:Art. 35a)).

Mystacopidae Miller, 1907:239. Type Mys-
tacops Lydekker (a replacement name for
Mystacina Gray).

Mystacinidae.—Simpson, 1945:60. First
use with correct suffix (ICZN 1985:Art.
Pit).

Mystacina Gray

Mystacina Gray in Dieffenbach, 1843:296.
Type Vespertilio tuberculatus G. Forster
in Dieffenbach, 1843:181 sensu Gray in
Dieffenbach, 1843:296 = Mystacina ve-
lutina Hutton, 1872.

Mystacops Lydekker in Flower & Lydek-
ker, 1891:671. Replacement name for
Mystacina Gray, erroneously thought by
Lydekker to be homonymous with Mys-
tacinus Boie, 1822. Type Vespertilio tub-
erculatus G. Forster in Dieffenbach,
1843:181 sensu Gray in Dieffenbach,
1843:296 = Mystacina velutina Hutton,
1872.

Mystacina velutina Hutton

Mystacina tuberculata.—Gray in Dieffen-
bach, 1843:296; Gray, 1843a (part),
1843b, 1875; Tomes, 1857; Dobson,
1878; Hill & Daniel, 1985.

Mystacina velutina Hutton, 1872:186.
“(T]he Hutt Valley, near Wellington’,
North Island, and ‘‘Milford Sound, on the
southwest coast of the South Island’’,
New Zealand. Lectotype herein designat-
ed to be the Colonial Museum specimen
from the Hutt Valley (see text for the fate
of this specimen).

Mystacops velutinus.—Thomas, 1905:423.

Mystacops tuberculatus.—Miller, 1907:
241.

Mystacina_ tuberculata
Dwyer, 1962:3.

tuberculata.—

Mystacina velutina velutina Hutton

Mystacina velutina velutina, New combi-
nation.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Mystacina velutina aupourica Hill &
Daniel

Mystacina tuberculata aupourica Hill &
Daniel, 1985:294. ‘““Omahuta Kauri Sanc-
tuary, Northland, North Island, New Zea-
land”’. Type Auckland Institute and Mu-
seum (AIM) M309.

Mystacina velutina aupourica, New com-
bination.

Mystacina velutina rhyacobia Hill &
Daniel

Mystacina tuberculata rhyacobia Hill &
Daniel, 1985:295. *““Te Rimu area, upper
Waimarino River, Kaimanawa Forest
Park, SE of Lake Taupo, central North
Island, New Zealand’’. Type AIM M304.

Mystacina velutina rhyacobia, New com-
bination.

Mystacina robusta Dwyer

Mystacina tuberculata robusta Dwyer,
1962:3. ““Big South Cape Island’’, Stew-
art Island region, New Zealand. Type Do-
minion Museum 1083. Daniel, 1979.

Mystacina robusta.—Hill & Daniel, 1985:
297; Daniel, 1990.

Acknowledgments

We wish to express our thanks to the au-
thorities of the British Museum (Natural
History) for facilitating inspection of
George Forster’s drawing, granting permis-
sion to reproduce it, and discussing For-
ster’s work, especially to A. Datta, V. Skeet,
and A. Wheeler; B. Williams (Field Muse-
um of Natural History) for providing access
to earlier literature and assistance in dating
it; C. Paulin (National Museum, Welling-
ton) for information about specimens in the
Colonial Museum; W. D. L Ride and P.
Tubbs (ICZN) for advice concerning the
fourth edition of the Code; D. Lee and H.
Spencer for reminding us of Andrews
(1986); T. Griffiths for providing copies of
literature; W. Feeny and D. Lintner for re-
producing the figures; M. Clayton for nau-

VOLUME 112, NUMBER 3

tical advice; T. Griffiths and L. Ruedas for
comments on the manuscript; and especial-
ly to W. D. L. Ride for extensive discus-
sions.

Late in the preparation of the manuscript
we became aware that H. Spencer and D.
Lee (Otago) were pursuing a parallel inves-
tigation of the nomenclature of New Zea-
land’s bats. We are most grateful to them
for sharing their views with us, and for
much discussion of the issues involved. We
are encouraged that our independent analy-
ses agree on many points, but regret that
certain points of difference remain.

Although as yet unpublished, Spencer
and Lee are making an application to the
International Commission on Zoological
Nomenclature (Case No. 3095 “‘Mystacina
Gray, 1843, Chalinolobus Peters, 1866, M.
tuberculata Gray, 1843 [currently C. tub-
erculatus] [Mammalia, Chiroptera]: pro-
posed conservation of usage.’’ BZN 55:
205). Note that Case No. 3095 does not
support the arguments and resolution here
proposed. Also, an application means (un-
der Article 80) that current usage is to be
maintained until a ruling is made.

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Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 8. A
provisional hypothesis for the hybrid origin of Zodalia glyceria
(Gould, 1858)

Gary R. Graves

Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution,

Washington, D.C. 20560, U.S.A.

Abstract.—Zodalia glyceria (Gould, 1858), supposedly from the vicinity of
Popayan, Colombia, is hypothesized to be a hybrid between Lesbia victoriae
and Chalcostigma herrani, which are sympatric in shrublands and timberline
ecotones in the Andes from southern Colombia to northern Peru. Those por-
tions of the capital, spinal, and ventral feather tracts that exhibit green irides-
cence in the parental species are greenish-blue to purple in the hybrid, de-
pending upon the angle of observation. For example, the dominant wavelength
reflected from back plumage is much shorter in the hybrid (505 nm) than in
either of the parental species (561-576 nm). This color shift is thought to have
been caused by a developmental aberrancy, possibly associated with hybrid-
ization, which affected melanin granules that produce iridescence in feather
keratins. Rectricial measurements of the hybrid fall between the character

means for L. victoriae and C. herrani whose tails differ markedly in shape.

Among the many puzzling species of
hummingbirds described by John Gould,
Cometes glyceria stands apart in taxonomic
obscurity (Gould 1858: opposite plate 176):

‘During the many years that I have given atten-
tion to the Trochilidae, I have not met with a bird
which has caused me more thought, and I may say
perplexity, .. . it is intimately allied to the members
of the genera Lesbia, Cometes and Cynanthus, par-
taking as it does, either in form or colouring, of
characters pertaining to each of those genera. Some-
times it has occurred to me that it might be a hybrid
between either two of them, but I am perfectly at a
loss to say which two species would be likely to
produce such a cross. Such an idea has entered my
mind, but when I have again and again reconsidered
the matter, it has appeared to me that it is a distinct
species, and that it may ultimately prove to be the
female or young male of some gorgeous bird with
which we are at present unacquainted.”’

As a consequence of evolving generic
definitions, glyceria was placed first in
Sparganura (Cabanis & Heine 1860, Scla-
ter & Salvin 1873) and later transferred to
Zodalia (Mulsant & Verreaux 1876, Elliot

1878, Salvin 1892, Boucard 1893, Sharpe
1900, Oberholser 1902, Cory 1918). Simon
(1921) considered Zodalia glyceria to be a
senior synonym and immature plumage of
Z. ortoni (Lawrence 1869), a position em-
braced by Peters (1945). The taxonomic va-
lidity of Zodalia glyceria was not formally
challenged until Meyer de Schauensee
(1947) declared both Z. glyceria and Z. or-
toni to be hybrids of Lesbia victoriae and
Ramphomicron microrhynchum. This con-
clusion was followed by subsequent authors
(e.g., Morony et al. 1975, Greenway 1978,
Fjeldsa & Krabbe 1990, Sibley & Monroe
1990, Collar et al. 1992). Meyer de
Schauensee’s (1947) evaluation of Z. ortoni
appears to be valid (Graves 1997); however,
the holotype of Zodalia glyceria seems to
represent a different, and previously unre-
ported, intergeneric hybrid. Here I provide
a detailed hybrid diagnosis employing the
methods and assumptions outlined in
Graves (1990) and Graves & Zusi (1990).
In recognition of its 120-year association

492

with glyceria, I use Zodalia as a generic
identifer instead of Cometes.

Materials and Methods

The holotype of Zodalia glyceria (The
Natural History Museum; formerly British
Museum of Natural History, BMNH
1888.7.25.184) was procured by M. Mossa
before 1853 in the vicinity of Popayan, Co-
lombia (Gould 1858). Although Salvin
(1892) considered Z. glyceria a juvenile and
Simon (1921) believed it to be an immature
of Z. ortoni, the holotypes of neither of
these species exhibit striations on the max-
illary ramphotheca typical of juvenile hum-
mingbirds (see Ortiz-Crespo 1972). Instead,
both specimens appear to be males in de-
finitive plumage based on the presence of
elongated rectrices and a brilliant, narrow
gorget that extends from the chin to the up-
per breast. The unique appearance of Z. gly-
ceria (Figs. 1—3) cannot be attributed to a
developmental or geographic variation of
any known taxon. Two other possibilities
exist—it represents a valid species or a hy-
brid. Because hybrids lack formal standing
in zoological nomenclature, hybridity must
be ruled out before species status is granted.
As this paper demonstrates, I was unable to
reject the plausibility of a hybrid origin for
Z. glyceria.

Assuming a hybrid origin, the pool of po-
tential parental species (Appendix 1) in-
cludes all species of trochiline humming-
birds (subfamily Trochilinae: Zusi & Bentz
1982, Sibley & Monroe 1990, Bleiweiss et
al. 1997) that occur in southern Colombia
and northern Ecuador (Chapman 1917, Hil-
ty & Brown 1986, Fjeldsa & Krabbe 1990,
Schuchmann & Heindl 1997, Krabbe et al.
1998). Notes, photographs, and videotape
of Zodalia glyceria were compared with
similar material for the holotypes of Zoda-
lia ortoni (=Lesbia ortoni; American Mu-
seum of Natural History, AMNH 156651;
Graves 1997), Zodalia thaumasta (National
Museum of Natural History, Smithsonian
Institution, USNM 173911), Chalcostigma

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

purpureicauda (AMNH 483931), Aeronym-
pha prosantis (Field Museum of Natural
History, FMNH 11852), and Heliangelus
zusii (Academy of Natural Sciences of Phil-
adelphia, ANSP 159261; Graves 1993).
The type of Z. glyceria was compared with
specimens of all trochiline species deposit-
ed in the Natural History Museum.

Measurements of the aforementioned type
specimens and of selected species were tak-
en with digital calipers and rounded to the
nearest 0.1 mm: wing chord; bill length
(from anterior extension of feathers); rectrix
Jength (from point of insertion of the central
rectrices to the tip of each rectrix), and rec-
trix width (at widest point). Pairs of rectrices
are numbered from the innermost (R1) to the
outermost (R5). Lengths of crown and gor-
get feathers were measured by sliding a slip
of paper to the feather base and then mark-
ing the position of the feather tip on the pa-
per (Appendix 2). Data judged to be most
relevant to the hybrid diagnosis are present-
ed in Table 1. Untransformed measurements
and least squares regression lines were pro-
jected on bivariate plots to illustrate size dif-
ferences (Fig. 4) (Wilkinson 1989).

I evaluated the color of the dorsal plum-
age (center of back) with a calibrated col-
orimeter (CR-221 Chroma Meter, Minolta
Corporation) equipped with a 3.0 mm ap-
erture. The measuring head of the CR-221
uses 45° circumferential illumination. Light
from the pulsed xenon arc lamp (C illumi-
nant, 2° observer) is projected onto the
Specimen surface by optical fibers arranged
in a circle around the measurement axis to
provide 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 plumage
surface without depressing it. The default

VOLUME 112, NUMBER 3

493

Fig. 1. Dorsal and ventral views of male Lesbia victoriae (top), Chalcostigma herrani (bottom), and a
probable hybrid, L. victoriae X C. herrani (=Zodalia glyceria [Gould, 1858]; BMNH 1888.7.25.184).

setting for the CR-221 Chroma Meter dis-
plays mean values derived from three se-
quential, in situ measurements. I repeated
this procedure three times for each area of

plumage, removing the aperture between
trials. Each datum summarized in Table 2
thus represents the mean of three indepen-
dent measurements.

494

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fis: 2.

Lateral view of male Lesbia victoriae (top), Chalcostigma herrani (bottom), and a probable hybrid,

L. victoriae X C. herrani (=Zodalia glyceria (Gould, 1858]; BMNH 1888.7.25.184).

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 (tinie>= Rheretore - reaness “and
‘““greenness’’ 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 O 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 LZ value will be. Vi-
sual systems in hummingbirds (e.g., Gold-
smith & Goldsmith 1979) differ signifi-
cantly from those of humans. The relevance

of opponent color coordinates to colors per-
ceived by hummingbirds is unknown. Gen-
eral color descriptions presented in Appen-
dix 2 were made under natural light.

The hybrid diagnosis was approached in
a hierarchical manner. A hypothesis of par-
entage was first derived from comparison of
plumage pattern, color and feather shape.
As a second step, the restrictive hypothesis
was tested with an analysis of size and ex-
ternal proportions. Concordance of results
is regarded as strong support for the hy-
pothesis (Graves 1990, Graves & Zusi
1990).

Results and Discussion

Characters of Zodalia glyceria that per-
mit its parental species to be identified in-
clude: (a) moderately lengthened crown
feathers broadly margined with rufous; (b)
partially white outer vane of outermost rec-
trices (R5); (c) deeply forked tail (fork

VOLUME 112, NUMBER 3

495

Fig. 3. Dorsal (top) and ventral (bottom) view of rectrices of Zodalia glyceria [Gould, 1858]; BMNH

1888.7.25.184.

depth = 51.6 mm); (d) broad rectrices (all
>10 mm wide); (e) narrow brilliant gorget;
and (f) short tibial plumes. This suite of
characters (Figs. 1-3, Appendix 2) can be
recovered from a particular pairwise com-
bination of species, but not from any single
Species.

The pool of potential parental species (a
maximum of (8°) = 3,655 possible pairwise
combinations) may be quickly narrowed by
focusing on the elongated rufous-margined
crown feathers of Z. glyceria. Meyer de
Schauensee’s (1947) hypothesis, Lesbia vic-
toriae X Ramphomicron microrhynchum, is
unlikely because neither of those species
possesses a rufous or chestnut crown patch.

Only two species in the geographic source
pool (Appendix 1) have rufous or chestnut
crowns: Chalcostigma herrani and C. rufi-
ceps. The latter species can be ruled out as
a parent of Z. glyceria on the basis of ex-
ternal measurements (see below).

The partially white outer vanes of the
outermost rectrices (R5) of Z. glyceria were
contributed by the other parental species.
Five species in the geographic source pool
(Ocreatus underwoodii, Lesbia victoriae, L.
nuna, Aglaiocercus kingi, A. coelestis) have
deeply forked tails. Ocreatus can be elimi-
nated from consideration because Z. glycer-
ia shows no evidence of racket-tipped rec-
trices or elongated tibial plumes. Neither

496 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Ranges (mean + standard deviation) of measurements (mm) of adult males of Lesbia victoriae,
Chalcostigma herrani, and a probable hybrid, L. victoriae <* C. herrani (=Zodalia glyceria (Gould, 1858);
BMNH 1888.7.25.184). Measurements of adult male Lesbia nuna are included for comparison.

Hybrid
victoriae nuna herrani BMNH
(n = 12) (n = 12) (n = 15) 1888.7.25.184

Wing chord 58.7—62.2 50.3-52.8 66.7—72.7 65.9
(60 Sas rl) (51.8 += 0.6) (9l8 = we7)

Bill length 13.5-15.3 7.5—9.0 9.8—-12.2 10.7
(14.5 + 0.6) (8.2 + 0.4) (Citas) 2es (0)5))

Rectrix | 22.1—24.9 19.8—22.4 43.8-47.9 33.8
(23.5 = 0.9) (21.0 = 0.7) (45.8 + 1.4)

Rectrix 2 26.2-31.1 25.9—28.1 48.5—52.5 41.9
C828) 12) @7- O05) (49.9 = 1.4)

Rectrix 3 39.2-44.4 36.0-40.4 49.9-56.3 51.0
42 aay) (38.0 = 1.4) C6257 =snIk@)

Rectrix 4 62.1—68.5 51.3-57.6 50.3—56.4 63.3
(65.2723) (54.3 + 1.9) (50.4 + 1.6)

Rectrix 5 149.0-189.0 94.1-109.0 47.1—56.2 85.4
(ChB y7/ 2 110) )) (99.9 + 4.1) Gs 23276)

species of Aglaiocercus possesses white
outer rectricial vanes in male definitive
plumage. A species of Lesbia is thus im-
plicated, although I doubt that the specific
identity of the “‘trainbearer’’ parent can be
determined from plumage color or pattern
alone. In brief, details of plumage pattern
and color are consistent with the hypothesis
that Zodalia glyceria is a hybrid between a
rufous-crowned species of Chalcostigma
and a species of Lesbia (Appendix 2).
Two peculiar aspects of plumage color
need to be addressed in greater detail. The
greenish-blue iridescence of Z. glyceria un-
doubtedly influenced Meyer de Schauensee
(1947) to propose the purple-backed thorn-
bill (Ramphomicron microrhynchum) as
one of its parental species. However, both
R. microrhynchum and Lesbia victoriae, as
well as a robustly documented hybrid, L.
victoriae X R. microrhynchum (AMNH
156651), exhibit green (instead of greenish-
blue or purple) iridescence on the breast
and sides (Graves 1997). It is usually as-
sumed that hybridization in hummingbirds
produces no traits characteristic of genera
or species other than those involved in the
particular cross (Banks & Johnson 1961).
In a recently published exception to this
general rule, plumage iridescence of a hy-

brid specimen, Aglaiocercus kingi X Me-
tallura tyrianthina (AMNH 146645), was
found to be significantly bluer (dominant
wavelength, 511 nm) than that exhibited by
the parental species (cumulative range,
553-571 nm) (Graves 1998b). The broader
implications of this finding are unclear, but
the shift in dominant wavelength may result
from a developmental aberrancy or muta-
tion, directly related to hybridization, that
affects melanin granules that produce iri-
descence in feather keratins.

The holotype of Zodalia glyceria appears
to represent another case of the “blueing”’
phenomenon associated with intergeneric
hybridization (Graves 1998b). The dorsal
plumage is significantly bluer (dominant
wavelength, 505 nm) than that of Lesbia
victoriae or Chalcostigma herrani (cumu-
lative range, 561-576 nm) (Table 2). How-
ever, the pattern of greenish-blue irides-
cence (violet to purple at certain angles) in
Z. glyceria coincides precisely with the dis-
tribution of green iridescence in those spe-
cies, suggesting a single developmental or
mutational event affecting iridescence of
the entire plumage (Fig. 5).

A second puzzlement is the absence of
terminal white spots on the outermost rec-
trices (R4—R5) of Z. glyceria. Presuming

VOLUME 112, NUMBER 3 497

50 80

S,)
(oe)

Length Rectrix 1
3 5
Wing Length
& 3

2
: 10) 50 100 150 200

Le)
oO

30 40 50
Length Rectrix 5 Length Rectrix 1

80 80

70

60

50
20 30 40 50 60

Wing Length
Wing Length

oO

ie)

35 40 45 50 55 60
Length Rectrix 2 Length Rectrix 3
80 80
70 70
= : =
Ei e
= =
60 60
50 50
60 55 60 65 70 8) 50 100 150 200
Length Rectrix 4 Length Rectrix 5
Fig. 4.

Bivariate plots of mensural characters of males in definitive plumage: Lesbia victoriae (@); Chal-
costigma herrani (@); a probable hybrid (W), L. victoriae X C. herrani (=Zodalia glyceria [Gould, 1858];
BMNH 1888.7.25.184). Least squares regression lines are illustrated for comparison.

498

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Ranges and means (+ standard deviation) of opponent color coordinates (L, a, b) and dominant
wavelength (nm) reflected from dorsal plumage (center of back) in male Chalcostigma herrani, Lesbia victoriae,
and their probable hybrid, L. victoriae < C. herrani (=Zodalia glyceria [Gould, 1858]; BMNH 1888.7.25.184).

C. herrani
Variables n= 15
Back plumage

L (Lightness) 20.1—23.3
Dh 3S FOS

a (Red [+]/Green [—]) —8.8—-(4+2.5)
ah, Fe SE
b (Yellow [+]/Blue [—]) 9.6—15.2
bie oF
Dominant wavelength (nm) 561-576
56910" S53

Chalcostigma herrani as a parental species,
this observation apparently conflicts with
assumptions I made in previous papers
about the heritability of unpigmented tail
spots in trochiline hybrids (Graves 1990,
1998a; Graves & Zusi 1990). This discrep-
ancy seemed to be resolved by a closer ex-
amination of Z. glyceria’s rectrices, which
suggests that the white terminal spots in C.
herrani were phenotypically expressed as
conspicuously white rachises in R4 and R5
(see Appendix 2). Although the basal third
of the dorsal rachial surface of R5 is dull
white or cream-colored in some female Les-
bia victoriae, the rachis of R4 is brown in

Dominant Wavelength (nm) of Back Piumage

20 25 30 35 40
Lightness (L) of Back Plumage

Fig. 5. Bivariate plots of mensural characters of
males in definitive plumage: Lesbia victoriae (A);
Chalcostigma herrani (@); a probable hybrid, L. vic-
toriae X C. herrani (@) (=Zodalia glyceria [Gould,
1858]; BMNH 1888.7.25.184).

L. victoriae hybrid
n= 21 1888.7.25.184
30.5—36.0 25-5
5310) 22 ok
—16.8-(—5.4) =
=A B26
19.2 —29.6 1.0
26.4 + 2.8
563-570 505
564.8 + 1.7

all specimens examined (50+). Conse-
quently the white rachises of R4 and R5 in
Z. glyceria cannot be attributed to female
or immature characters of Lesbia victoriae.
In summary, details of plumage color and
pattern are sufficient to narrow the pool of
possible parental combinations: Lesbia (vic-
toriae or nuna) X Chalcostigma (herrani or
ruficeps).

External measurements.—Several straight-
forward comparisons permit the identifica-
tion of the parental species of Zodalia
glyceria: (a) length of R3 (Chalcostigma
herrani > Z. glyceria > Lesbia victoriae >
C. ruficeps > L. nuna); (b) length of R4 (ZL.
victoriae > Z. glyceria > L. nuna > C. her-
rani > C. ruficeps); (c) width of R5 (C.
herrani > Z. glyceria > C. ruficeps > L.
victoriae > L. nuna) (Table 1, unpublished
data). Crown and gorget feathers of C. her-
rani and Z. glyceria are also significantly
longer than those of the other taxa. Because
morphological luxuriance (where hybrids
are larger than their parental species) has
never been observed in trochilines, these
data indicate L. victoriae and C. herrani as
the most probable parental species of Z.
glyceria.

Male Lesbia victoriae and Chalcostigma
herrani in definitive plumage differ mark-
edly in tail shape and rectricial measure-
ments are non-overlapping. The deeply
forked tail (fork depth = 83-88% of tail
length) of male L. victoriae is among the

VOLUME 112, NUMBER 3

longest in the Trochilidae (319 species ac-
cording to Sibley and Monroe 1990), with
tail/wing ratios ranging from 2.4 to 3.1 in
Ecuadorian populations. In contrast, the tail
of C. herrani is relatively short (tail/wing
= 0.7-0.8) and shallowly forked (fork
depth = 5-16% of tail length). Bivariate
plots of wing length and rectricial measure-
ments of L. victoriae and C. herrani exhibit
both positive (R1, R2, R3) and negative
(R4, R5) allometry (Fig. 4). Rectricial mea-
surements of Z. glyceria fall between the
character means of L. victoriae and C. her-
rani, and, in most cases, approximate the
values predicted by least squares regression.

In summary, evidence derived from size
and shape characters, as well as plumage
pattern and color, strongly suggest that Z.
glyceria represents an intergeneric hybrid
between Lesbia victoriae and Chalcostigma
herrani. As such, Cometes (=Zodalia) gly-
ceria Gould is available only for the pur-
poses of homonymy in taxonomy.

Geographic overlap.—The geographic
and elevational ranges of the parental spe-
cies, Lesbia victoriae and Chalcostigma
herrani, overlap broadly in the Andes
(Fjeldsa & Krabbe 1990). Lesbia victoriae
inhabits forest edge and brushy slopes at
2600—4000 m elevation, whereas C. her-
rani prefers well-drained rocky slopes, for-
est edge and Polylepis woodland at 2700—
3600 m.

Acknowledgments

I thank Richard Banks, Robert Bleiweiss,
Richard Prum, and Robert Pr¥ys-Jones for
comments on the manuscript. I am grateful
to Robert Pr¥s-Jones, Michael Walters,
Mark Adams, and Don Smith of the Natural
History Museum, Tring, for permission to
study the type of Zodalia glyceria, and the
curators and staff of the Field Museum of
Natural History, American Museum of Nat-
ural History, and the Academy of Natural
Sciences of Philadelphia for access to col-
lections. Photographic prints were prepared
by the Smithsonian photographic services.

499

Leslie Overstreet (Smithsonian Institution
Libraries) assisted with bibliographic ma-
terial. Travel was supported by the Re-
search Opportunities Fund, the Alexander
Wetmore Fund, and the Department of Ver-
tebrate Zoology, Smithsonian Institution.

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SYSTAT, Inc., Evanston, Illinois, 822 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.

Appendix 1

Species of trochiline hummingbirds that occur in
southern Colombia (Cauca, Narino, Putamayo) and
northern Ecuador (Carchi, Esmeraldas, Imbabura): An-
drodon aequatorialis, Doryfera johannae, D. ludovi-
ciae, Campylopterus largipennis, C. falcatus, Colibri
delphinae, C. thalassinus, C. coruscans, Anthracoth-
orax nigricollis, Klais guimeti, Lophornis chalybeus,
Popelairia popelairii, P. conversii, Chlorostilbon mel-
lisugus, Thalurania colombica, T. furcata, Damophila
julie, Hylocharis grayi, Chrysuronia oenone, Amazilia
fimbriata, A. amabilis, A. rosenbergi, A. franciae, A.
cyanifrons, A. saucerrottei, A. tzacatl, Chalybura buf-
fonii, C. urochrysia, Adelomyia melanogenys, Phlo-
gophilus hemileucurus, Heliodoxa imperatrix, H. gu-
laris, H. schreibersii, H. aurescens, H. rubinoides, H.
jacula, H. leadbeateri, Topaza pyra, Oreotrochilus
chimborazo, Urochroa bougueri, Patagona gigas,
Aglaeactis cupripennis, Lafresnaya lafresnayi, Ptero-
phanes cyanopterus, Coeligena coeligena, C. wilsoni,
C. torquata, C. lutetiae, Ensifera ensifera, Boisson-
neaua flavescens, B. matthewsii, B. jardini, Heliange-
lus strophianus, H. exortis, Eriocnemis vestitus, E. lu-
ciani, E. mosquera, E. mirabilis, E. alinae, E. derbyi,
Haplophaedia aureliae, H. lugens, Urosticte benjami-
ni, U. ruficrissa, Ocreatus underwoodii, Lesbia victo-
riae, L. nuna, Ramphomicron microrhynchum, Metal-
lura williami, M. tyrianthina, Chalcostigma ruficeps,
C. stanleyi, C. herrani, Opisthoprora euryptera,
Aglaiocercus kingi, A. coelestis, Schistes geoffroyi, He-

VOLUME 112, NUMBER 3

liothryx barroti, H. aurita, Heliomaster longirostris,
Philodice mitchellii, Myrtis fanny, Acestrura mulsant,
A. bombus, A. heliodor, A. berlepschi.

Appendix 2

General comparative description of adult male Les-
bia victoriae, Chalcostigma herrani, and a probable
hybrid, Lesbia victoriae X Chalcostigma herrani (=
Zodalia _ glyceria [Gould, 1858]; BMNH
1888.7.25.184). Descriptions of structural colors are
unusually subjective, as color seen by the observer
varies according to the angle of inspection and direc-
tion of light. For this reason I use general color de-
scriptions.

Dorsal feathering (capital and spinal tracts) of vic-
toriae posterior to the upper tail coverts is medium dull
green; feather bases are gray and some lateral barbs
are narrowly fringed with buff. Crown feathers are un-
modified (length, 6.1-7.1 mm, X = 6.6 + 0.3; n =
IS):

The crown of herrani is dark saturated green (matte
black when viewed head-on), bisected by a medial
stripe of deep lustrous rufous, which begins at the base
of the bill, reaches its widest point on the forecrown
(where the barbs have a plush-like texture), and nar-
rows (one or two feathers wide) to a thin stripe on the
hindcrown. The bases of rufous-tipped crown feathers
(length, 8.1-10.6 mm, X = 9.6 + 0.9; n = 16) are
grayish-black. In males in subdefinitive plumage, the
feather barbs are less plush-like, and the rufous crown
stripe is considerably wider, often extending laterally
to the superciliary region. The back of herrani is dark
green (sooty green from head-on) becoming dark
bronzy-green with reddish (coppery) highlights on the
rump and uppertail coverts.

Dorsally, glyceria differs considerably in appear-
ance from victoriae and herrani, or for that matter, all
other species of hummingbirds. First, an oval crown
patch extends from the base of the bill to the nape.
Feathers in this weakly defined patch (~9.2 mm long)
are broadly tipped with rufous and possess a shining
central spot (imparting a spotted appearance) that, de-
pending upon the angle of inspection, reflects greenish-
blue or purple iridescence. The superciliary, nape,
back, scapulars, rump, and uppertail coverts are shin-
ing greenish-blue to purple; the distal ends of barbs
are pale rufous or buff, and not easily seen without
magnification.

In victoriae, a brilliant, medium-green gorget ex-
tends from the chin to the upper breast; the posterior
end of the gorget is broadly lanceolate. Feathers at the
gorget point are of moderate length (8.4—9.8 mm, X
= 9.1 + 0.4; n = 15) and width (~2.5—3.2 mm). From
base to tip they are grayish-buff proximally, bordered
distally by a broad band of pale buff, a narrow tran-
sitional band (<0.4 mm) of bronzy green, and broadly
tipped with medium green (terminal iridescent tip:

501

length ~2.8—3.0 mm, width ~2.5—3.2). The lores, au-
riculars, sides of neck, breast, and flanks are green,
new feathers are finely margined (10 magnification)
with buff. The basal third of feathers on the breast,
sides, and belly is dark gray; feathers are narrowly
fringed with buff, extensively so along the midline be-
low the gorget and on the abdomen. Vent plumes are
white; undertail coverts (13—16 mm long) are buff with
a muted and elongated grayish spot along the rachis.
Tibial plumes, which extend approximately half way
to the toes, are dark grayish-brown, broadly tipped
with buff; tibial plumes are buff.

In definitive plumage, herrani has a narrow and
highly iridescent gorget (“‘beard’’), extending from the
chin to the upper breast, that changes color in a step-
wise fashion from blue (chin) to red (gorget tail). Al-
though a large fraction of the visible spectrum is re-
flected in ~20—25 rows of gorget feathers, the exposed
portion of individual feathers appears to the eye
(viewed head-on) to be rather uniform in color, reflect-
ing a truncated range of wavelengths. The posterior %
of the gorget tail is frequently 1—2 feathers wide.
Feathers in gorget tail are relatively long (length, 12.0—
15.3 mm, 13.6 + 0.9; n = 16) and narrow (~1.7-2.5
mm), the iridescent red tip is similarly elongated
(ength ~5.0—6.0 mm, width ~2.0—2.3 mm). These
feathers, from base to tip, are sooty gray or black,
gradually becoming black glossed with green, and then
changing abruptly in an optically smooth gradient
(~0.7—1.2 mm) from greenish-gold, to coppery-gold
and coppery-orange, and finally to red (depending on
the angle of inspection, the terminal portion of the
feather can vary from reddish-orange to reddish-ma-
genta when viewed head-on). The auriculars and sides
of the throat are dusky green (matte black from head-
on), contrasting with the brilliant gorget. The breast,
sides, and belly are dusky green becoming buffy to-
ward the midline. In general, ventral feathers are gray-
ish-buff, marked with a diffuse dull green disc, which
becomes progressively smaller toward the midline.
The vent plumes are white. Undertail coverts are mod-
erately long (20-25 mm), pale buffy-white to buff,
marked with grayish smudges, or in some individuals,
by an indistinct grayish stripe of variable width. Tibial
plumes extend about % of the way to the toes, and are
dark brownish-gray with a scattering of buffy barbs.

The gorget of glyceria is uniformly silvery-green
(viewed head-on), and is composed of ~20—21 rows
of iridescent feathers that extend from the chin to the
upper breast. At midpoint, the gorget is about 5 feath-
ers in width. Chin feathers have intermingled buff and
greenish barbs. Feathers from the gorget tail most
closely resemble those of victoriae in color, exhibiting
an indistinct lanceolate spot (pale grayish-buff) cen-
tered at the rachis near the base of the feather, bordered
distally by a broad subterminal band of pale buff, and
terminated with a highly iridescent silvery-green tip
(length ~3.8 mm, width ~2.0; total feather length

502

~10.1 mm). Gorget feathers of glyceria are interme-
diate in shape between those of victoriae and herrani.
The color pattern of the ventral plumage of glyceria
most closely resembles that of victoriae. Ventral feath-
ers are dark gray basally, broadly tipped with buff, and
marked with an iridescent spot (greenish-blue to violet
and purple, depending on the angle of inspection), the
size and distinctiveness of which decreases toward the
midline. The tibial plumes, which extend approximate-
ly half way down the tarsus, are dark grayish-brown,
broadly tipped with pale buff and grayish-buff barbs
(approximately intermediate in appearance between
victoriae and herrani). The vent feathers are white.
Undertail coverts are pale buff, marked with a subter-
minal lanceolate spot (greenish-purple on largest co-
verts) whose darkness, color intensity and size increas-
es with covert size.

In victoriae, the rectrices (dorsally) are black with
brownish-purple reflections in bright light, conspicu-
ously (R1—R4) or inconspicuously (R5) tipped with
dark green. The proximal % of the lateral vane of R5
is gray (dorsally) and grayish-white (ventrally). Rec-
trices are relatively narrow (R3—R5 < 7.0 mm wide).
The dorsal surface of rachises is dark brown (the prox-
imal 10—25 mm of R5 in some specimens is cream-
colored). Ventrally, rachises are dark brown, gradually
shading to pale brown at the base, except for R5 in
which the proximal third of the rachis is grayish-white
(the pigmentation of R4 is intermediate). Tips of R5
are slightly subspatulate and “bowed” in cross-sec-
tion.

The dorsal and ventral surfaces of herrani rectrices
reflect a metallic iridescence that varies from dark blu-
ish-purple on the outer rectrices (R5) to dark purple
on the innermost (R11). The two outer rectrices are
tipped with large white spots (length, measured along
the rachis), 13.8-16.7 mm (RS5) and 10.0—11.7 mm

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(R4). R3 exhibits a faint white mark near the tip along
the rachis. Rachises are dark brownish black, becom-
ing white in the unpigmented spots on both surfaces.
Rectrices are relatively wide: (R1) 10.3-13.7 mm;
(R2) 11.1-15.8 mm; (R3) 10.9-14.6 mm; (R4) 11.2—
13.7 mm; (R5) 10.0—-12.8 mm. In cross-section, the
rectrices are nearly flat.

The rectrices of glyceria are nearly flat and relative-
ly wide: (R1) width = 10.5 mm; (R2) 11.0 mm; (R3)
11.6 mm; (R4) 10.4 mm; (R5) 10.1 mm. Rectrix shape
in glyceria is approximately intermediate between that
of victoriae and herrani. Dorsally, R1 is dark metallic
bronzy-purple gradually shading toward purple about
12 mm from the tip and finally to violet-purple (ter-
minal 3 mm). R2—R4 are similarly colored although
the basal bronzy-purple portions are largely obscured
in the folded tail. The central part (~24—62 mm from
feather tip) of the lateral vane of R5 is dull white on
both ventral and dorsal surfaces (dilute pigmentation
can be observed at 10X magnification). No traces of
white tipping or terminal spotting could be observed
on any of the rectrices of Z. glyceria under magnifi-
cation (10). Dorsally, the rachises of R1—R3 are dark
brown becoming light brown proximally. Rachises of
R4 and R5 are conspicuously white starting approxi-
mately 14 mm and 16 mm, respectively, from the
feather tip. All rachises are slighter darker on ventral
surfaces.

Remiges are dull dark brown and similar in shape
in victoriae, herrani, and in glyceria. Greater wing co-
verts and primary coverts are the same color as back
plumage in the respective specimens. The bill of gly-
ceria is intermediate in shape between that of victoriae
(slightly decurved in lateral profile, smoothly tapered
in dorsal profile) and herrani (straight in lateral profile,
abruptly tapered in dorsal profile). The ramphothecas
and feet of victoriae, herrani, and glyceria are black.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(3):503—509. 1999.

A new species of pelican (Aves: Pelecanidae) from the Lower
Pliocene of North Carolina and Florida

Storrs L. Olson

Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560-0116, U.S.A.

Abstract.—A new species of pelican, Pelecanus schreiberi, is described from
rare fossils from the Lower Pliocene Yorktown Formation of North Carolina.
Additional material from the nearly contemporaneous Bone Valley Formation
in Central Florida is tentatively referred to the same species, which was much
larger than any existing New World pelican and so distinctive in some features
that it appears to represent an extinct lineage with no living descendents. The
holotype is the distal portion of a femur with dense medullary bone indicating
a laying female, so the species presumably bred near the type locality.

Among the tens of thousands of bird fos-
sils recovered from early Pliocene marine
deposits excavated in a phosphate mining
operation in North Carolina (Olson & Ras-
mussen 2000), are a few specimens belong-
ing to an extremely large species of pelican
(Pelecanidae: Pelecaniformes). These are so
distinctive as to leave little doubt that they
constitute a previously unrecognized ele-
ment in the late Tertiary avifauna of North
America. With the discovery of additional
contemporaneous specimens from Florida
that may represent the same species, it was
decided that formal description was war-
ranted, despite the paucity of material.

Order Pelecaniformes
Family Pelecanidae

The femora are referable to the Pelecan-
idae by the combination of the very long,
broad anterior surface of the external con-
dyle; the long, squared outline of the fibular
condyle in posterior view; the very long,
distinct scar for attachment of M. gastroc-
nemius lateralis; and the distinct, deep de-
pression for the ligamentous loop of M. il-
iofibularis. The overall aspect of the distal
end of the femur in anterior view is some-
what similar in the Cathartidae but the gas-

trocnemius scar is small and round, the de-
pression for the loop of the iliofibularis is
indistinct, and the external condyle is rela-
tively shorter. The Pelagornithidae differ
markedly in having the popliteal fossa in-
distinct (versus very deep), the shaft broad-
er and not constricting above the condyles,
and the external condyle much less poste-
riorly produced. From the illustrations of
the femur of Teratornis merriami in Miller
(1925, plate 4c, d), the Teratornithidae dif-
fer in having the popliteal fossa shallower,
the fibular condyle much less produced lat-
erally, the gastrocnemius scar smaller, and
the intercondylar groove much shallower.
Other taxa of large birds in the Lee Creek
avifauna such as swans, storks, and cranes
(Anatidae, Ciconiidae, and Gruidae) differ
even more substantially from the Pelecani-
dae than the above families.

Pelecanus schreiberi, new species
(Figs. 1-2)

Holotype.—Distal third of right femur of
an egg-laying adult female (lumen filled
with dense medullary bone), paleontologi-
cal collections of the National Museum of
Natural History, Smithsonian Instution,
UNSM 192077; collected about 1972 by

504

Gerard R. Case; collector’s number NC
543.

Type locality.—Texasgulf Inc. Lee Creek
Mine, south side of Pamlico River, near Au-
rora, Beaufort County, North Carolina
(35°23'22" .N:.76°47'06" W).

Horizon.—Basal part of the Yorktown
Formation, Lower Pliocene. This determi-
nation was made from matrix removed
from paratypical femur USNM 263567 and
is based on the presence of secondary phos-
phate, abundant echinoid spines, and the fo-
raminiferan Nonionella labradorium
(Thomas G. Gibson, USGS, pers. comm.,
1999). Most of the avian fossils recovered
at Lee Creek are from the basal part of the
Yorktown Formation and it is assumed that
the other specimens cited here come from
that unit as well.

Measurements (mm) of holotype.—Distal
width 37.2, depth through external condyle
28.3, depth through internal condyle 22.3,
length from proximal edge of scar for M.
gastrocnemius lateralis to distal extent of
external condyle 34.2, width and depth of
shaft at proximal edge of scar of M. gas-
trocnemius lateralis 23.6 X 15.8.

Paratypes.—The following are topotyp-
ical paratypes from the Lee Creek Mine:
distal fourth of left femur, USNM 263567:
pedal phalanx 1 of digit III, USNM 446506;
pedal phalanx 2 of digit Ill, USNM 421948.

Mainly on the basis of their very large
size, the following specimens from the
Bone Valley Formation in Polk County,
central Florida, are also referred to P.
schreiberi: right quadrate lacking orbital
process, UF 125031 (Seminole phosphate
mine); axis vertebra lacking dorsal spine,
UF 65677 (Gardinier mine).

Measurements.—See Table 1.

Etymology.—To my late friend and col-
league Ralph W. Schreiber (6 July 1942-29
March 1988), former curator of birds at the
Natural History Museum of Los Angeles
County, authority on and devoted student of
pelicans, whose career was ended much too
soon (see Woolfenden 1989).

Diagnosis.—Larger than all modern pel-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

icans except the largest individuals of the
two largest species (Pelecanus crispus
Bruch and P. onocrotalus Linnaeus). Distal
end of femur in distal view with rotular
groove narrower and much deeper, anterior
crest of external condyle extending anteri-
orly well beyond the level of that of the
internal condyle; in posterior view fibular
condyle much more expanded laterally. The
pedal phalanges are much more robust than
the comparable elements in modern peli-
cans.

Discussion.—This very large species ex-
ceeds in size either of the living North
American pelicans (American White Peli-
can P. erythrorhynchos Gmelin and the
smaller Brown Pelican P. occidentalis Lin-
naeus). Because medullary bone functions
as a calcium reserve, forming 10 to 14 days
prior to egg-laying and being quickly re-
sorbed afterwards (see references cited in
Ballmann 1979, Mourer-Chauviré et al.
1999), the holotype is certainly from a fe-
male. The paratypical femur is essentially
the same size as the holotype and thus
could be from a non-breeding female. This
is turn suggests that males (the larger sex)
of the fossil species may have exceeded in
size any individuals of modern pelicans.

Pelecanus schreiberi must have been
breeding somewhere near the site of fossil
deposition in North Carolina, making it a
previously unknown member of the resi-
dent avifauna of eastern North America, de-
Spite its apparent rarity. Size alone pre-
cludes its inclusion in any existing species
of North American pelican and the quali-
tative differences in the femur are so great
as almost to suggest generic distinction.

Pelican remains are also represented in
the nearly contemporaneous Bone Valley
Formation of central Florida (Becker 1987),
but are likewise very scarce. A single axis
vertebra (UF 65677) is too large for Pele-
canus erythrorhynchos (Table 1) and is here
tentatively referred to P. schreiberi, al-
though otherwise I could detect no consis-
tent differences from P. erythrorhynchos or
Old World pelicans. Of three quadrates

VOLUME 112, NUMBER 3

Fig. 1. Femora of Pelecanus. A—-D, holotype of Pelecanus schreiberi, new species (USNM 192007): A,
proximal view; B, posterior view; C, distal view; D, anterior view. E—G, paratype of P. schreiberi, new species
(USNM 263567): E, proximal view; FE posterior view; G, anterior view. Unlettered elements are comparative
views of the modern species P. crispus (USNM 557493). Note the dense medullary bone in the lumen of the
femur of the holotype (A), indicating an egg-laying female, versus the hollow trabeculated lumen of the paratype
(E). Scale bar = 2 cm.

506

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Measurements of bones of Pelecanus. P. schreiberi n = 1 or 2; P. crispus n = 2 (1 unsexed, probably
female; 1 male); P. erythrorhynchos all are males (the larger sex); n = 8 except quadrate and axis n = 7.

P. schreiberi
range

P. crispus
range (mean)

P. onocrotalus
range (mean)n

P. erythrorhynchos
range (mean)

Quadrate
Depth 33.0
Length mandibular articulation 25.0
Axis
Length 38.5
Posterior width he
Femur
Distal width 37.2-38.2
Depth external condyle 27.3—27.9
Phalanx | Digit II
Length 33:6
Proximal width 132
Distal width 8.2
Phalanx 1 digit III
Length 37.8
Proximal width 9.6
Distal width Gee

from the Bone Valley Formation, one (UF
65699) is smaller than in males of P. ery-
throrhynchos but probably within the range
of females of the species, one (UF 125031)
is larger than in P. erythrorhynchos in one
measurement and within the range in an-
other, whereas the third (UF 125030) ex-
ceeds that species in both measurements
(Table 1) and is therefore referred to P.
schreiberi. The measurements of three dis-
tal ends of tarsometatarsi from Bone Valley
(UF 12344, UF 29738, UF 123868) all fall
within the range of variation of P. erythro-
rhynchos. The quadrates and axis in the
Brown Pelican (P. occidentalis) differ
markedly in morphology from those of oth-
er pelicans, and are unlike those elements
found at Bone Valley. If two species of pel-
ican are represented there, one would be re-
ferred to P. schreiberi and the other would
presumably be in the P. erythrorhynchos
lineage.

The fossil record of Pelecanidae else-
where in North America is also very mea-
ger. Miller (1944) tentatively assigned three
poorly preserved fragments of femora from

34.8—37.0 (35.9)
23.9-27.3 (25.6)

34.5—39.9 (37.2)
23.4—-25.8 (24.6)

35.3-37.4 (36.3)
25.2—27.9 (26.5)

48.1—57.3 (52.7)
10.5—11.9 (11.2)
7.1-8.9 (8.0)

37.2-44.5 (40.8)

30.4—33.6 (32.0) 2 28.7—29.8 (29.2)
25.0—29.4 (27.8) 4 23.0-23.9 (23.4)

35.8-43.7 (39.5) 7
23.5-28.7 262) 7

33.6—35.9 (35.0)
211-233-227)

33.5-37.6' G59y7
25.1—27.6 (26.7) 7

30.3-—33.2 (31.8)
21.5—24.0 (22.5)

47.5—57.8 (53.5) 5
10.6—12.6 (11.9) 5
7.6-9.5 (8.4) 5

40.746.5 (44.5)
8.7—11.0 (10.1)
6.6-75. GI)

36.9-45.0 (41.4) 5
8.4—-10.1 (9.0) 5
6.0-8.3 (7.0) 5

30.1-35.9 (34.1)
7.0-8.5 (7.9)
5.0-5.8 (5.6)

from the Pliocene of Oregon to P. erythro-
rhynchos. A new species, P. halieus, was
erected by Wetmore (1933) for the proximal
portion of a radius from the late Pliocene
Hagerman deposits of Idaho said to be sim-
ilar to P. erythrorhynchos but smaller. The
radius is usually not considered a very di-
agnostic element, but when I re-examined
the holotype of P. halieus (USNM 12233)
I could find nothing that it resembled more
than a pelican and it is indeed smaller than
in P. erythrorhynchos. Becker (1986) as-
signed additional material from Idaho to
this species.

Pelicans are much better represented in
the fossil record of Europe, where Pelecan-
us gracilis Milne-Edwards is known from
fairly abundant remains from the early Mio-
cene of France. Cheneval (1984) made this
the type of a new genus, Miopelecanus, and
also suggested that Pelecanus intermedius
Fraas, from the Middle Miocene of Ger-
many might be conspecific with Miopele-
canus gracilis, although he did not use the
former in a new combination, contra Mli-
kovsky (1992:435). These pelicans were

VOLUME 112, NUMBER 3

507

Pig, 2.

Pedal phalanges of Pelecanus crispus (USNM 557493) on the left in each pair compared with

paratypes of Pelecanus schreiberi, new species, on the right in each pair. A, pedal phalanx 1, digit Il (USNM
446506), lateral view; C, same, dorsal view; B, pedal phalanx 2, digit II] (USNM 421948), lateral view; D,

same, dorsal view. Scale bar = 2 cm.

much smaller and more gracile than P.
schreiberi, and the rotular groove of the fe-
mur is figured (Cheneval 1984: pl. 5-2) as
quite broad, not narrow and deep as in P.
schreiberi.

With one exception, discussed below,
other named pelicans from Europe, Asia,
and Africa would all have been smaller than
P. schreiberi (see Lydekker 1891, Harrison
& Walker 1976). The various fossil pelicans
from Australasia, reviewed by Rich & van

Tets (1981), likewise were smaller than P.
schreiberi, with the exception of Pelecanus
conspicillatus novaezealandiae Scarlett,
which Rich & van Tets elevated to full spe-
cies rank. The width of the femur in this
species, from the late Holocene of New
Zealand, was the size of that of P. schrei-
beri, although on temporal and geographic
grounds it would be highly unlikely for the
two to be closely related. Furthermore,
Scarlett’s (1966: fig. 4) illustration shows

508

the rotular groove of the femur of the New
Zealand bird to be very wide and shallow,
unlike that of P. schreiberi.

Perhaps of greater potential relevancy in
the present connection is the almost ethereal
taxon Pelecanus odessanus Widhalm
(1886), which was proposed in an obscure
publication that seems to have eluded many
researchers. Mlikovsky (1996), for exam-
ple, was unable to consult it, and one won-
ders if Brodkorb (1963) may have taken his
information about it from Lambrecht
(1933). Beginning at least with Lambrecht
(1933), the author’s name has most often
been incorrectly rendered as ‘“Wildhalm,”’
when in fact it was Germanicized as J.
Widhalm (from Ighnatii Martuinovich
Vidghal’m).

Widhalm’s paper treats a few fossils of
waterbirds (the pelican, a toe bone of “‘Co-
lymbus,”’ which at that time probably meant
a loon [Gavia], and an assortment of bones
of cormorants) from Tertiary deposits at
Novaja Slobodka, near Odessa, Ukraine,
previously assigned to the Lower Pliocene
but now to the Upper Miocene (MN 11-13,
Mlikovsky 1996:749). This was for its time
(or even now) an exemplary paper. In the
case of the pelican, for example, Widhalm
took into account recent views on the com-
plexities of nomenclature of modern spe-
cies, gave measurements of the fossil and
the two relevant modern species, along with
fairly detailed comparisons, and provided a
superb lithographic plate.

The problem arises with what has been
interpreted as Widhalm’s apparent failure
to conform to binomial nomenclature, but
in my opinion this is an unfair appraisal
probably arising from other authors’ lack
of access to the original publication. Brod-
korb (1963), for example, considered Wid-
halm to be nonbinomial and therefore at-
tributed the name Pelecanus odessanus to
Lambrecht (1933). Mlikovsky (1996) was
of the opinion that Widhalm’s apparent
treatment of cormorants from the site
called into question the validity of Wid-
halm’s nomenclature. Widhalm’s exact ty-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

pography where the new pelican was pro-
posed (p. 6) was as follows: “‘Pelecanus
odessanus. fossilis Widhalm.’’. It is evi-
dent that Widhalm did not intend the term
‘‘fossilis” to be part of the scientific name,
but merely an indication that his pelican
was a fossil, as opposed to a modern taxon.
The situation with cormorants is a little
more complex but again I believe does not
involve anything that can be considered
formal nomenclature as on page 8 they are
referred to as: ‘“‘Haliaeus fossilis, var.
Odessana major, medius, und minor. Wid-
halm.”’, which I interpret as simply a pro-
visional way of saying that there were
three sizes of fossil cormorants from Odes-
sa (Halieaus Illiger, 1811, is a synonym of
Phalacrocorax Brisson, 1760).

Regardless of its authorship, Pelecanus
odessanus was a very large pelican, the
length of the tarsometatarsus being given as
150 mm (that in the largest specimen of P.
crispus that I examined was 131 mm, and
in the two largest P. onocrotalus it was 138
and 145 mm). This species was temporally
quite close to P. schreiberi and was prob-
ably of similar size. Unfortunately it is not
known what became of Widhalm’s speci-
mens (Lambrecht 1933, Mlikovsky 1996).
Absent comparable elements it would not
be possible to determine the distinctness of
P. odessanus from P. schreiberi in any
case.

Acknowledgments

I am grateful to Mark Frank and David
W. Steadman, Florida Museum of Natural
History, Gainesville, for information, ar-
rangements, access to fossil and modern
collections, and for lending specimens,
and to Elizabeth Ann Schreiber for assis-
tance and encouragement. Mark Florence
assisted with USNM specimens. I am
grateful for useful comments on the man-
uscript by Steven D. Emslie and Kenneth
E. Campbell. The photographs are by
John Steiner, Smithsonian Photographic
Services.

VOLUME 112, NUMBER 3

Literature Cited

Ballmann, P. 1979. Fossile Glareolidae aus dem Mioz-
ain des Nordlinger Ries (Aves: Charadriifor-
mes).—Bonner Zoologische Beitrage 30:51—
101.

Becker, J. J. 1986. Fossil birds of the Oreana Local
Fauna (Blancan), Owyhee County, Idaho.—
Great Basin Naturalist 46:763—768.

1987. Neogene avian localities of North
America. Smithsonian Institution Press, Wash-
ineton, D.C., 171 pp.

Brodkorb, P. 1963. Catalogue of fossil birds: Part 1
(Archaeopterygiformes through Ardeifor-
mes).—Bulletin of the Florida State Museum,
Biological Sciences 7:179—293.

Cheneval, J. 1984. Les oiseaux aquatiques (Gaviifor-
mes a Ansériformes) du gisement Aquitanien de
Saint-Gérand-le-Puy (Allier, France): Révision
systématique.—Palaeovertebrata 14:33—115.

Harrison, C. J. O., & C. A. Walker. 1976. A new fossil
pelican from Olduvai.—Bulletin of the British
Museum Natural History (Geology) 27:315-
320.

Lambrecht, K. 1933. Handbuch der Palaeornitologie.
Gebriider Borntraeger, Berlin, 1024 pp.
Lydekker, R. 1891. Catalogue of the fossil birds in the
British Museum (Natural History). British Mu-

seum, London, 364 pp.

Miller, L. 1925. The birds of Rancho La Brea.—Car-
negie Institution of Washington Publication
349:1—106, 6 plates.

. 1944. Some Pliocene birds from Oregon and
Idaho.—Condor 46:25-32.

Mlikovsky, J. 1992. The present state of knowledge of
the Tertiary birds of central Europe.—Science

509

Series Natural History Museum of Los Angeles

County 36:433—458.

. 1996. Tertiary avian localities of Ukraine. Pp.
743-757 in J. Mlikovsky, ed., Tertiary avian lo-
calities of Europe.—Acta Universitatis Caroli-
nae Geologica 39 (for 1995):519—846.

Mourer-Chauviré, C., R. Bour, S. Ribes, & F Moutou.
1999. The avifauna of Réunion Island (Mascar-
ene Islands) at the time of the arrival of the first
Europeans. /n Storrs L. Olson, ed., Avian pa-
leontology at the close of the 20th century. Pro-
ceedings of the 4th International Meeting of the
Society of Avian Paleontology and Evolution,
Washington, D.C., 4—7 June 1996.—Smithson-
ian Contributions to Paleobiology 89 (in press).

Olson, S. L., & P. C. Rasmussen. 2000. Miocene and
Pliocene birds from the Lee Creek Mine, North
Carolina. In C. E. Ray and D. J. Bohaska, eds.,
Geology and paleontology of the Lee Creek
Mine, North Carolina, I1J].—Smithsonian Con-
tributions to Paleobiology (in press).

Rich, P. V., & G. E van Tets. 1981. The fossil pelicans
of Australasia——Records of the South Austra-
lian Museum 18:235—264.

Scarlett, R. J. 1966. A pelican in New Zealand.—No-
tornis 13:204—217.

Wetmore, A. 1933. Pliocene bird remains from Ida-
ho.—Smithsonian Miscellaneous Collections
87(20):1-12.

Widhalm, J. 1886. Die fossilen Vogel-Knochen der
Odessaer-Steppen-Kalk-Steinbritiche an der
Neuen Slobodka bei Odessa.—Schriften der
Neurussische Gesellschaft der Naturforscher zu
Odessa [Odesskoe obshchestvo estestvoispyta-
telei zapiski] 10:3—9, plate 5.

Woolfenden, G. E. 1989. In memoriam: Ralph W.
Schreiber, 1942—1988.—Auk 106:137—140.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(¢3):510-514. 1999.

A new species of Platymantis (Amphibia: Ranidae) from the Sierra
Madre Mountains, Luzon Island, Philippines

W. C. Brown, A. C. Alcala, P. S. Ong, and A. C. Diesmos

(WCB) Department of Herpetology, California Academy of Sciences, San Francisco, California
94118, U.S.A.; (ACA) Commission on Higher Education, DAP Bldg., San Miguel Avenue, Pasig
City, Philippines; (PSO) Institute of Biology, University of the Philippines, Diliman, Quezon
City, Philippines; (ACD) College of Arts and Sciences, University of the Philippines Los Bafios
College, Laguna, Philippines

Abstract.—A new species, Platymantis sierramadrensis, from the Sierra Ma-
dre Mountains in the provinces of Aurora and Isabela, Luzon Island, Philip-
pines, is described. This species is distinguished from other Philippines species
of Platymantis (hazelae group) by its pale color, smooth skin, advertisement
call, and other characters given in the diagnosis.

Recent field work in the Sierra Madre
Mountains, northeastern Luzon Island, con-
ducted separately by us in 1996 and 1997
resulted in the collection of 22 specimens
(16 adults and 6 juveniles) of this distinc-
tive frog. These specimens are platymantine
frogs based on structure of the pectoral gir-
dle (Brown 1952) and are referred to the
hazelae group of Platymantis based on dig-
ital structures (Brown et al. 1997). This spe-
cies is an addition to the list of eight other
species occurring on Luzon and the Central
and Western Visayas and currently assigned
to this group. The specimens are deposited
in the California Academy of Sciences
(CAS) and Philippine National Museum
(NMPH).

Materials and Methods

Materials examined included holotypes
of all other species of the hazelae group of
Platymantis from the Philippines. These ho-
lotypes with the exception of P. panayensis
are in the CAS collections. The holotype of
P. panayensis is in the NMPH.

Morphometric characters.—Snout—vent
length (SVL), head length (HL), head
breadth (HW), snout length (SnL), diameter
of eye (ED), diameter of tympanum (TD),

tibia length (TiL), third finger length from
proximal edge of basal tubercle (3FL), di-
ameter of third finger disk (3FD), and di-
ameter of third toe disk (3ToD).

Advertisement calls (Fig. 1) were record-
ed and analyzed, using a Kay Electrics
SonaGraph (Model #550) and SIGNAL
Sound Analysis System software.

Platymantis sierramadrensis, new species
Fig. 2

Holotype.-—NMPH 6465, an adult fe-
male, collected by Marisol Pedregosa in
disturbed lowland forest at Sitio Mapidjas,
Barangay Umiray, Municipality of General
Nakar, Quezon Province, Luzon Island on
29 May 1996.

Paratypes.—CAS 204738, 204742—45
and NMPH 5980, 6461-63, 6466-67,
CMNH 05678-79, collected in disturbed
lowland forest in Sitio Mapidjas, Barangay
Umiray, Municipality of Dingalan, Aurora
Province, Luzon Island at the southern end
of the Sierra Madre mountains.

Referred specimens.—CAS 204739-—41
and NMPH 6464, 6470—74 from Palanan,
northern Sierra Madre Mountains, Isabela
Prov., Luzon Island. These specimens are
in close agreement morphologically and in

fet Spal are
aa he ae
5 STERR at: :

~ >

)

FREQUENCY (KHZ

TIME (sec)

Fig. 1. Audiospectrograms of advertisement calls of (A) Platymantis sierramadrensis (NMPH 6472), (B) P.
subterrestris (CAS 204322), (C) P. montanus (CAS 201705).

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Rigs.
Luzon Island. Color (in life), pale yellowish without dark markings on either body or limbs, is shown in this
photograph of a male clinging to a branch of a forest shrub.

advertisement call with the type and para-
types. This population at the northern end
of the Sierra Madre needs further investi-
gation. It may prove to be a valid subspe-
cies.

Diagnosis.—Platymantis sierramadren-
sis differs from other members of the ha-
zelae group in having a pale creamy color
without dark brown or blackish markings
on body or limbs. In preserved specimens,
faint, scattered brownish flecks are evident
on dorsal and lateral surfaces. Other char-
acters are: smooth skin, terminal phalanges
rounded, and finger disks broader than
those on toes, SVL 22.7—25.2 for mature
males and 25.7 for one mature female; HL
(39% of SVL) about equal to HW (40% of
SVL) for 10 males and one female.

Description of holotype.—A gravid fe-

Platymantis sierramadrensis (NMPH 6464) from Sierra Madre National Park, Isabella Province,

male; dorsum uniformly off white; dorsum
and venter smooth; abdomen translucent,
some internal organs visible; a conical tu-
bercle on heel; measurements in mm: SVL
25.7, HL 10.0, HW 9.9, SnL 4.6, ED 3.6,
TD 1.5, TiL 13.3, 3FL. 3-7, 3FD 12Siteb
0.9.

Description.—SVL 22.7—25.2 for 10 ma-
ture males and 25.7 for one mature female:
HL 37-42% (mean 39%) of SVL; HW 37-—
42% (mean 40%) of SVL; SnL 17-21%
(mean 19%) of SVL; ED 12-15% (mean
14%) of SVL; TD 5-8% (mean 6%) of
SVL; TiL 49-56% (mean 52%) of SVL;
3FL 14-17% (mean 15%) of SVL; 3FD 5—
7% (mean 6%) of SVL; 3ToD 3—5% (mean
4%) of SVL; tympanum exposed; lore mod-
erately oblique, concave; vomerine teeth
only slightly protruding, patches widely

VOLUME 112, NUMBER 3

separated; fingers with minute webs at base;
fingers except first with broad disks; fingers
with shallow circum-marginal grooves; first
finger shortest, third finger longest, and sec-
ond and fourth about equal in length; su-
barticular tubercles large, round, low; a row
of low, inconspicuous supernumerary tu-
bercles on palm; inner, middle and outer
metacarpal tubercles oval, vague, inner and
middle about equal in size; larger than out-
er; hind limb long; toes webbed to distal
edge of tubercle on first and second, to dis-
tal edge of basal tubercle on third, and to
midway between tubercles on fifth; disks of
toes narrower than those of fingers, subar-
ticular tubercles rounded, low; plantar area
smooth; inner metatarsal tubercle elongat-
ed, outer vague; dorsum smooth, without
tubercles; venter generally smooth, coarsely
granular in some specimens.

Variations.—Based on the limited sam-
ple available to us, there are a few small
differences in the body proportions between
the northern (Isabela Province) and the
southern (Aurora Province) populations.
However, it is not possible to say whether
these differences are significant because of
the small samples. The northern population
has barely overlapping values for HL/SVL
(40—42 versus 37-40%) and TD/SVL (5-6
versus 6—8%) and non-overlapping values
for 3FD/SVL (16-17 versus 14-15%). The
two general collecting sites are far apart (ca.
200 km) and the forest environment is no
longer continuous. These populations may
not interact in any way.

Color (in life).—Pale cream without dark
bars or other marks on limbs or body and
without areolations.

Color (in preservative).—Pale cream
without dark bars or other marks. Small
brownish pigment flecks are scattered on
dorsal and lateral surfaces. The specimens
from the northern Sierra Madre (Isabela
Province) appear slightly darker.

Reproduction.—The only mature female
in the collections has about 10 ovarian eggs
of various sizes, the largest being 2.8 mm
in diameter. The eggs are yellow in color

513

and are devoid of dark pigments, suggesting
that the species undergoes direct develop-
ment like other members of Platymantis.
This female was collected in May, 1996 in
Aurora Province. However, rainfall occurs
throughout the year, and breeding activities
may also be non-seasonal.

Advertisement call.—The call of this frog
sounds like “‘pek-pek-pek’’ produced in a
forceful manner. Each note ranges from
2500 to 3250 Hz with a duration of 0.05 to
0.06 of a second. The time interval between
notes is about 0.53 to 0.64 seconds (Fig. 1).

Ecological notes.—Specimens of this
species were collected in virgin and dis-
turbed lowland forest at altitudes of 55 to
550 m. They were observed on leaves of
low shrubby plants and screw pines from
0.5 to 2.5m above the ground. In Palanan,
Isabela (northern part of Sierra Madre),
adult males were observed and heard call-
ing in March—April, 1997 while sitting on
leaves of shrubs.

Etymology.—The species name is de-
rived from Sierra Madre Mountains, the
type locality.

Comparisons.—Platymantis sierramad-
rensis, based on digital characters, belongs
in the hazelae group. This species differs
from the other species of the group primar-
ily in the distinctive color pattern. In life,
the color appears cream to creamy yellow
(Fig. 1) without the dark markings and/or
areolations on the body and limbs that char-
acterize other species of this group (Brown
et al. 1997). This species also differs in the
appearance of the ventral body wall. It per-
mits a partial view of underlying organs,
especially eggs in gravid females.

In terms of size (SVL) at maturity, based
on those species with samples of fine or
more specimens, this species is among
those with ranges from about 20 to 28 mm
for males and with ranges widely overlap-
ping when species are compared (Brown et
al. 1997).

The species of the hazelae group of
Platymantis are known only from the cen-
tral and northern islands of the Philippines

514

in contrast to the ranges exhibited by the
guentheri group (throughout the Philip-
pines), and the dorsalis group (Philip-
pines, Palau Islands, Melanesia, and New
Guinea). Three of previously recognized
species of the hazelae group are known
from the central islands and five species
are known from mountain ranges or iso-
lated mountains on the northern island of
Luzon. Platymantis sierramadrensis does
not change this zoogeographic concept. It
simply adds a sixth species of the group
for Luzon Island, a species apparently iso-
lated in a previously little known moun-
tain range, the Sierra Madre. Platymantis
sierramadrensis is not the first new spe-
cies of this genus to be described from the
Sierra Madres. It is preceded by Platy-
mantis pygmaeus (dorsalis group, Brown
et al. 1998).

Advertisement calls have been recorded
for three species of the hazelae group (sier-
ramadrensis, montanus, and subterrestris).
The note structure is similar for these three
species, but the frequencies and intervals
between notes differ (Fig. 1).

Acknowledgments

We wish to thank Robert C. Drewes for
clearing and staining a specimen, Jens Vin-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

dum for assistance in the laboratory and
Andrea Jesse for producing the audiospec-
trograms. We thank Danilo S. Balete, Ron-
ald Allan Altamirano, Arturo Manamtam,
Renato Fernandez, Genevieve Gee, Cristi
Nozawa, Arne Jensen, Finn Danielsen, and
Morten Heegard (NORDECO) for allowing
us to study the specimens from Isabela
Province. We also thank Roy Quiber and
Marisol Pedregosa for collecting the speci-
mens from Aurora Province. We also thank
W. Pollisco, Director of the Department of
Environment and Natural Resources-Pro-
tected Areas and Wildlife Bureau, for his
assistance.

Literature Cited

Brown, W. C. 1952. The amphibians of the Solomon
Islands.—Bulletin of the Museum of Compar-
ative Zoology 107(1):1—64.

, R. M. Brown, & A. C. Alcala. 1997. Species

of the hazelae Group of Platymantis (Amphibia:

Ranidae) from the Philippines, with descriptions

of two new species.—Proceedings of the Cali-

fornia Academy of Sciences 49(11):405—421.

, A. C. Alcala, & A. C. Diesmos. 1998. Two

new species of the genus Platymantis (Amphib-

ia: Ranidae) from Luzon Island, Philippines.—

Proceedings of the California Academy of Sci-

ences 50(17):381—-388.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
112@G):515-922. 1999.

Two new species of the Eleutherodactylus rugulosus group
(Amphibia: Anura: Leptodactylidae) from Honduras

James R. McCranie and Larry David Wilson

(JRM) 10770 SW 164th Street, Miami, Florida 33157-2933, U.S.A.; (LDW) Department of
Biology, Miami-Dade Community College, Kendall Campus, Miami, Florida 33176, U.S.A.

Abstract.—Two new species of Eleutherodactylus, E. pechorum and E. olan-
chano, of the E. rugulosus group are described from Honduras. The following
combination of characters will distinguish E. pechorum from the remaining E.
rugulosus group members: male vocal slits present; male nuptial thumb pad
absent; finger II < I; female loreal length/eye length ratio =1.00; keels present
on unwebbed portions of toes; toes with basal webbing; canthus rounded; pos-
terior surface of thighs marked with numerous pale spots and blotches; antero-
ventral and posteroventral surfaces of tibial segments barred; ventral surfaces
pale yellow; and size moderate. Eleutherodactylus olanchano is distinguished
from all other E. rugulosus group species by the following combination of
characters: male vocal slits and nuptial thumb pads absent; finger II > I or II
= I; tympanum length/eye length ratio 21.00; and male size very small (fe-

males unknown).

Recent fieldwork in Honduras has result-
ed in the discovery of two new species of
the Eleutherodactylus rugulosus group.
One of these species is a streamside dwell-
er, as are most members of the group. How-
ever, the other species occurs on forested
hillsides, well above streams that are inhab-
ited by E. aurilegulus, another rugulosus
group member. The forest dwelling species
represents only the third known member of
the rugulosus group occurring in that type
of habitat (Campbell et al. 1994, Johnson
& Savage 1995).

Materials and Methods

All morphological and color pattern traits
discussed herein follow the character states
and terminology defined by Savage (1975).
Comparative data for E. azueroensis, E.
taurus, and E. vocalis were taken from Sav-
age (1975). Comparative material examined
is listed in Appendix I. The name rugulosus
is in quotes in this appendix because work
in progress by J. A. Campbell and J. M.

Savage will likely severely restrict the geo-
graphic distribution of FE. rugulosus. Thus,
the specific identity of the Honduran and
Nicaraguan material of E. “‘rugulosus’’ ex-
amined is unresolved at this time. Abbre-
viations used in each species description are
EL (eye length), E—N (anterior border of
eye to posterior edge of nostril; equals lo-
real length of Savage 1975), HL (head
length; tip of snout to angle of jaw), HW
(greatest width of head), LL (leg length),
SL (snout length; anterior border of eye to
tip of snout), SVL (snout—vent length), TL
(tibia length), and TM (tympanum length).
Measurements were made to the nearest 0.1
millimeter under a dissecting microscope.
Museum acronyms follow those of Leviton
et al. (1985) and color codes are those of
Smithe (1975-1981).

Systematics
Eleutherodactylus pechorum, new species
Fig. 1

Holotype.—National Museum of Natural
History (USNM) 530000, an adult male,

516

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

tgs

4 fied se |

Figs:

from near a small dam along a small trib-
utary of the Quebrada de Las Marias
(15°17.64'N, 85°21.29'W), about 12 airline
km NNE La Colonia, Departamento de
Olancho, Honduras, 680 m elev., collected
2 Aug 1998 by J. R. McCranie, K. L. Wil-
liams, and L. D. Wilson. Original number
LADW2 11375.

Paratype.—USNM 529999, and adult fe-
male with the same locality data as the ho-
lotype, except collected 1 Aug 1998.

Referred specimen.—BMNH 1985.1454,
an adult female from along the Rio Cuya-
mel, Departamento de Colon, Honduras.

Diagnosis.—The combination of the
presence of vocal slits and the absence of
nuptial thumb pads in males will distin-
guish E. pechorum from all other described
members of the E. rugulosus group, except
for E. anciano, E. azueroensis, E. taurus,
and E. vocalis. Eleutherodactylus pechorum
is distinguished from E. anciano by having
finger IJ shorter than finger I (finger II lon-
ger than or equal in length to finger I in E.
anciano), toe keels on toe IV of females
(weakly infolded fringes on at least one side
of toe IV in all four females), shorter fe-
male snouts (E—-N/EL = 1.000 versus

Adult male holotype of Eleutherodactylus pechorum (USNM 530000), SVL 33.5 mm.

=1.000, n = 4), and larger female size (to
59 mm SVL versus to 41 mm, n = 4). In
addition, E. anciano is known only from
remnants of broadleaf cloud forest (Lower
Montane Moist Forest formation of Hold-
ridge 1967) at 1770 to 1840 m elevation in
southwestern Honduras. The new species
differs from E. azueroensis of the Peninsula
de Azuero, Panama, in having the posterior
surface of the thighs marked with numerous
pale spots and blotches (a few small pale
spots in E. azueroensis), the anteroventral
and posteroventral surfaces of the tibial
segments barred (uniformly brown), the
canthus rounded (moderately sharp), and
basal toe webbing (moderately webbed;
webbing formula IJ 2-3 1/2 III 3—4 1/3 IV
4 1/3-3- V in E. pechorum versus I 2-—3
1/4 Ill 2 2/3—4 IV 4-2 1/2 V in E. azue-
roensis). Eleutherodactylus pechorum dif-
fers from E. taurus of southwestern Costa
Rica and adjacent Panama in having toe
keels on unwebbed portions of toes (well
developed fleshy toe fringes in E. taurus),
four and one-third segments on toe IV free
of webbing (<4), definite toe discs (barely
expanded), and females to 59 mm SVL (fe-
males to 80 mm). The new species can be

VOLUME 112, NUMBER 3

distinguished from EF. vocalis of western
Mexico by having finger II shorter than fin-
ger I (finger II longer than or equal in
length to finger I in E. vocalis), ventral sur-
faces pale yellow (white), and the antero-
ventral and posteroventral surfaces of the
tibial segments barred (suffused).
Description of holotype (alcohol after
formalin).—An adult male with the follow-
ing measurements (percentages of SVL in
parentheses): SVL 33.5 mm; HL 13.1 mm
G91); HW 12.5 mm (37.3); EL 3.9 mm
(11.6); SL 5.6 mm (16.7); E-N 3.4 mm
Cet EM 3.5 mm (10.4); LL 56.5 mm
(168.7); and TL 19.6 mm (58.5). Dorsum
smooth with several enlarged tubercles on
upper eyelids, short postorbital ridges also
present; snout nearly rounded in dorsal as-
pect, rounded in profile; canthus rostralis
rounded; tympanum prominent; first finger
longer than second; finger discs definite, a
little over twice width of digit just proximal
to discs on fingers III-IV; well developed
inner tarsal fold; heels rugose; plantar tu-
bercles absent; inner metatarsal tubercle
elongated, elevated, visible from above;
outer metatarsal tubercle small, round, bare-
ly elevated; toe discs definite, that on toe
IV about 1.6 times width of digit just prox-
imal to disc; toes with marginal ridge; toes
basally webbed, webbing formula I 2—2 1/
2 I 2-3 1/2 Il 3-4 1/3 IV 4 1/3-3° V;
vomerine tooth patches on elevated, nearly
triangular-shaped ridges located postero-
medially to ovoid choanae, tooth patches
separated by distance less than width of ei-
ther patch; paired vocal slits present; nuptial
pads absent. Belly cream, lightly flecked
with dark brown; throat and chest cream,
moderately flecked with dark brown; dor-
sum brown with indistinct dark brown mot-
tling, several white spots scattered on back;
postorbital ridges slightly darker brown
than adjacent dorsum; dark line absent
along canthal ridges; loreal region brown;
upper and lower lips with dark brown bars
separated by white lines, dark lower lip bars
not extending onto chin; top of head brown
with slightly paler interorbital bar anterior

517

to median portion of orbit; supratympanic
fold slightly darker brown than adjacent
area; groin mottled with dark brown and
white; posterior surface of thighs dark
brown with numerous cream spots and
blotches; anterior surface of thighs pale
brown with dark brown vertical bars; un-
derside of thighs and tibial segments cream
with very sparse dark brown flecks, barred
with dark brown along margins.

Color in life (based on a Kodachrome®
slide, except for ventral coloration, which
was recorded in life): dorsal surfaces of
head and body Verona Brown (223B), with
slightly darker brown mottling and slightly
paler brown, scattered spots; postorbital
ridges mottled Verona Brown (223B) and
slightly darker brown; dorsal surfaces of
limbs Raw Umber (23), with darker brown
crossbands; interorbital region mottled pale
brown, medium brown, and dark brown,
suggestive of pale interorbital bar outlined
with darker brown; dark brown upper lip
bars separated by pale brown; tympanum
intermediate between Verona Brown
(223B) and darker brown mottling; iris cop-
pery brown, reticulated with black; belly
and ventral surfaces of thighs pale yellow.

Variation in paratype (alcohol after for-
malin).—Color is described as follows: bel-
ly, chest, and throat cream, lightly flecked
with grayish-brown; dorsum grayish-brown
with distinct dark brown mottling and elon-
gated, narrow spots; several dirty white
small spots also present on lower back;
postorbital ridges distinctly darker brown
than adjacent dorsum; dark line absent
along canthal ridges, although several dark
flecks present just anterior to eye; loreal re-
gion pale brown; upper and lower lips with
medium brown bars separated by dirty
white lines, dark lower lip bars not extend-
ing onto chin; top of head grayish-brown
with slightly paler brown interorbital bar,
pale bar bordered anteriorly and posteriorly
by dark brown; supratympanic fold darker
brown than adjacent area; groin mottled
with grayish-brown and pale brown, several
dark brown spots also present; posterior

518

surface of thighs grayish-brown with nu-
merous cream spots and blotches; anterior
surface of thighs pale brown with grayish-
brown vertical bars; underside of thighs and
tibial segments cream, barred with grayish-
brown along margins.

The following measurements (percent-
ages of SVL in parentheses) were recorded:
SVL 41.2 mm; HL 17.0 mm (41.3); HW
16.0 mm (38.8); LL 70.7 mm (171.6); and
TL 24.8 mm (60.2).

Comments on referred specimen.—A sin-
gle adult female (BMNH 1985.1454; SVL
59.1 mm) from about 50 airline km NNE
of the type locality of E. pechorum is re-
ferred to this species. This specimen is very
similar in color pattern and morphology to
the female paratype of E. pechorum. Also,
although the type locality for E. pechorum
is in the headwaters of the Rio Wampu, the
locality is only some 15 km from the head-
waters of the Rio Paulaya. The locality for
the BMNH specimen is along a tributary of
the latter river. Additionally, E. epochthi-
dius, another streamside Eleutherodactylus
(E. milesi group), is known from both the
E. pechorum type locality and from a lo-
cality very near that for BMNH 1985.1454.

Natural history notes.—Both type spec-
imens were collected at night alongside a
small stream flowing through nearly pris-
tine forest at 680 m elevation. However, at
660 m elevation, the former forest around
this stream has been cleared, as have most
of the hillsides surrounding the forest on
both sides of the stream. The forest along-
side this portion of the stream remains in-
tact only because water from that portion of
the stream is piped to several villages below
the stream. This extensive deforestation is
taking place even though the region is part
of the Rio Platano Biosphere Reserve, es-
tablished as a World Heritage Site in 1980.
The type locality is in the Premontane Wet
Forest formation of Holdridge (1967).

Etymology.—The name pechorum means
‘belonging to or pertaining to”’ the Pech,
in reference to this frog inhabiting an area

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

long, but sparsely populated by the indig-
enous Pech tribe.

Eleutherodactylus olanchano, new species
Fig: 2

Holotype.—National Museum of Natural
History (USNM) 529998, an adult male,
from a hillside above the Quebrada El Pinol
(15°07'N, 86°44’W), Parque Nacional La
Muralla, Departamento de Olancho, Hon-
duras, 1200 m elev., collected 14 Aug 1994
by J. R. McCranie and L. D. Wilson. Orig-
inal number LDW 10319.

Paratypes.—USNM 529991-97, all
adult males, all from the type locality,
1180—1200 m elev., collected 21 Jul 1993
(USNM 529996-97) or 12-14 Aug 1994
(the remaining specimens).

Referred specimens.—USNM 529989—
90, both adult males, from near Rio de En-
medio (14°52'N, 86°48’W), Montafia El Ar-
mado, Departamento de Olancho, Hondu-
ras, 1350 m elev., collected 13 Jun 1993.

Diagnosis.—The combination of small
adult male size (SVL 18.7—29.8, K = 24.8;
females unknown), large male tympanum
(TM/EL 1.00—1.14, X = 1.06), finger II
longer than or equal in length to finger I,
and males lacking vocal slits and nuptial
thumb pads will distinguish E. olanchano
from all other described species of the E.
rugulosus group. Eleutherodactylus olan-
chano is the smallest known member of the
E. rugulosus group. Males of all other spe-
cies reach at least 33 mm SVL (the single
known adult males of E. anciano and E.
pechorum), 38 mm SVL (E. vocalis), or 40
mm or longer (the remaining E. rugulosus
group species: Savage 1975, Savage et al.
1988, Campbell et al. 1994, Johnson &
Savage 1995).

Description of holotype (alcohol after
formalin).—An adult male with the follow-
ing measurements (percentages of SVL in
parentheses): SVL 29.8 mm; HL 13.1 mm
(44.0); HW 11.7 mm (39.3); EL 3.5 mm
(11.7); SL 4.8 mm (16.1); E-N 3.0 mm
(10.1); TM 4.0 mm (13.4); LL 55.6 mm

VOLUME 112, NUMBER 3

S19

Bis.<2.

(186.6); and TL 17.6 mm (59.1). Dorsum
smooth to weakly granular, with several
distinct tubercles on lower back and upper
eyelids; postorbital ridges absent; canthus
rostralis rounded; tympanum prominent;
first finger equal in length to second; finger
discs definite, about 1.8 times width of digit
just proximal to discs on fingers [J-IV;
well developed inner tarsal fold; toe discs
definite, that on toe IV about 2.0 times
width of digit; toes with marginal ridge;
toes moderately webbed, webbing formula
I 2—2 1/2 If 2-3 1/3 Il 3-4 IV 4-2 3/4 V;
vocal slits absent; nuptial pads absent. Belly
cream, very lightly flecked with dark
brown; chest cream, heavily flecked with
dark brown; throat heavily flecked with
dark brown; dorsum dark brown with sev-
eral very dark brown markings in postocu-
lar region; dark line absent along canthal
ridges; loreal region mottled pale and dark
brown; upper and lower lips with dark

Adult male holotype of Eleutherodactylus olanchano (USNM 529998), SVL 29.8 mm.

brown bars separated by cream lines, dark
lower lip bars not extending onto chin; top
of head pale brown with a few dark brown
small spots in interocular region near mid
level of orbit; supratympanic fold very dark
brown; groin cream, heavily flecked with
dark brown; posterior surface of thighs dark
brown, with indistinct pale brown blotches;
anterior surface of thighs pale brown, with
slightly darker, incomplete vertical bars; un-
derside of thighs and tibial segments cream,
with very sparse dark brown flecks, barred
with dark brown along margins.

Color in life (based on a Kodachrome®
slide, except for ventral coloration, which
was recorded in life): dorsal surfaces of
head and body Cinnamon-Rufous (40), with
several dark brown spots in postorbital and
interorbital regions and slightly darker
brown mottling on lower back; dorsal sur-
faces of limbs Cinnamon-Rufous (40), with
darker brown, somewhat indistinct, cross-

520

bands; dark brown upper lip bars separated
by pale brown; supratympanic folds dark
brown; tympanum pale brown; iris reddish-
brown, reticulated with dark brown on up-
per half, grayish-brown, reticulated with
dark brown on lower half, both halves not
separated by dark band; belly and ventral
surfaces of thighs pale yellow.

Variation in type series (alcohol after
formalin).—Belly coloration of all para-
types is similar to that of holotype; most
specimens have less brown flecking on
chest and throat than does holotype; how-
ever, one specimen (USNM 529992) has
throat and chest more intensely punctated
with dark brown; dorsal surfaces of head
and body vary from grayish-brown to dark
brown, with varying amounts of dark
brown spots or lines present in postorbital
and interocular regions, these dark marks
indistinct in darker specimens; one speci-
men (USNM 529992) has a rather broad,
pale brown middorsal stripe extending from
tip of snout to just above vent; most spec-
imens have indistinct pale brown blotches
on posterior surface of thighs; however,
these blotches can be rather distinct in a few
specimens (e.g., USNM 529991-92); un-
derside of thigh and tibial segments similar
to that of holotype in all specimens, except
that one specimen (USNM 529992) has
slightly more flecking medially on thighs
near knees than does the holotype.

Morphological measurements of the en-
tire type series include SVL given in mil-
limeters and other measurements as _ per-
centages of SVL, range followed by mean
in parentheses: SVL 18.7—29.8 (24.8); HL
40.3—44.6 (43.5); HW 37.1—40.1 (38.6); LL
168.8—208.6 (191.0); and TL 56.4—62.9
(59.4). Additionally TM/EL ranges and
means are 1.00—1.14 (1.06) in the entire
type series, and finger II varies from being
longer than finger I to being equal in length
to finger I. Examination of the testes of two
specimens (USNM 529993-94; 28.1 and
23.1 mm SVL, respectively) confirmed that
they are adults.

Comments on referred specimens.—Two

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

adult males (USNM 52998990; 26.5 and
24.0 mm SVL, respectively) from about 30
airline km SSW of the type locality agree
in all diagnostic features with those of the
type series. These specimens were collected
by M. R. Espinal. His collection from the
Rio de Enmedio region also contains two
adult females of E. aurilegulus (USNM
529987-88; 53.5 and 53.6 mm SVL, re-
spectively). Espinal did not record any hab-
itat information for this collection, but it
seems likely that the E. olanchano were
taken from forested hillsides above the
streamside locality where E. aurilegulus
would be expected to occur.

Natural history notes.—The type series
was collected both during the day and at
night while active among leaves on the for-
est floor on a single hillside above the Que-
brada El Pinol. Specimens were collected
in July and August. Two species of stream-
side Eleutherodactylus (E. aurilegulus and
E. stadelmani; E. rugulosus and E. milesi
groups, respectively) were common along-
side the Quebrada El Pinol at the base of
the hillside, while E. lauraster (E. rhodopis
group) was found in the same habitat as the
E. olanchano. This locality is in the Pre-
montane Wet Forest formation of Holdridge
(1967), as is the locality for the referred
specimens. The known elevational range
for this species is 1180 to 1350 m.

Etymology.—The name olanchano is
Spanish, meaning “‘native of Olancho,”’ and
is used in reference to this Honduran de-
partment, in which this species is apparent-
ly endemic. The name is a noun used in
apposition to the generic name.

Discussion

Members of the Eleutherodactylus ru-
gulosus group occur from southern San
Luis Potosi, Mexico on the Atlantic versant
and from northern Sinaloa, Mexico on the
Pacific versant southward to central Panama
(Savage 1975). The two new species de-
scribed herein appear to have relatively
small geographical distributions as do many

VOLUME 112, NUMBER 3

other Central American members of this
group (Savage, pers. comm.).

Among the described species of the E.
rugulosus group, E. pechorum appears to be
most similar morphologically to E. azue-
roensis of low and moderate elevations
(60—940 m) of the Peninsula de Azuero,
Panama. However, the intervening lowlands
(with the exception of the Golfo Dulce area
of the Pacific versant of southern Costa
Rica and adjacent Panama where E. taurus
occurs) are inhabited by one or more mod-
erately webbed species of the E. rugulosus
group in which the males lack vocal slits
and nuptial thumb pads. No specimens from
the intervening territory resemble either E.
pechorum or E. azueroensis in the subtle
features that distinguish species in this
group. The localities for E. pechorum lie in
excess of 900 km NNW of the nearest
known locality for E. azueroensis.

The second new species described herein
(E. olanchano) possesses a combination of
male characters (very small size, very large
tympanum, finger II longer than or equal in
length to finger I, and the absence of vocal
slits and nuptial thumb pads) not shared
with any other described member of the

group.

Acknowledgments

Collecting and exportation permits were
provided by M. Acosta, A. Barahona, E.
Munoz, G. Rodriguez, and C. Romero of
COHDEFOR, Tegucigalpa. Field assistance
was provided by D. Almendarez, M. R. Es-
pinal, G. A. Flores, and K. L. Williams. Es-
pinal also donated the specimens from his
Rio de Enmedio collection and provided
valuable help in obtaining the COHDEFOR
permits. Comparative material was provid-
ed by C. J. Cole (AMNHB), B. T. Clarke
(BMNH), W. W. Tanner (BYU), E. J. Cen-
sky and J. J. Wiens (CM), A. Resetar
(FMNH), J. E. Cadle and J. Rosado (MCZ),
A. Holman (MSUM), R. W. Murphy
(ROM), G. Kohler (SMF), K. R. Vaughan
(TCWC), D. L. Auth (UF), A. G. Kluge and

eval

G. Schneider (UMMZ), S. W. Gotte and R.
W. McDiarmid (USNM), and R. Giinther
(ZMB). An early draft of this manuscript
was reviewed by J. M. Savage, who also
provided many helpful comments concern-
ing the systematics of the E. rugulosus

group.

Literature Cited

Campbell, J. A., J. M. Savage, & J. R. Meyer. 1994.
A new species of Eleutherodactylus (Anura:
Leptodactylidae) of the rugulosus group from
Guatemala and Belize.—Herpetologica 50:412—
419.

Holdridge, L. R. 1967. Life zone ecology. Revised edi-
tion. Tropical Science Center, San José, Costa
Rica, 206 pp.

Johnson, J. D., & J. M. Savage. 1995. A new species
of the Eleutherodactylus rugulosus group (Lep-
todactylidae) from Chiapas, Mexico.—Journal
of Herpetology 29:501—506.

Leviton, A. Be Ror. Gibbs, Jr, E. Heal, & C. E.
Dawson. 1985. Standards in herpetology and
ichthyology: part I. Standard symbolic codes for
institutional resource collections in herpetology
and ichthyology.—Copeia 1985:802-832.

Savage, J. M. 1975. Systematics and distribution of the
Mexican and Central American stream frogs re-
lated to Eleutherodactylus rugulosus.—Copeia
1975:254-—306.

, J. R. McCranie, & L. D. Wilson. 1988. New
upland stream frogs of the Eleutherodactylus
rugulosus group (Amphibia: Anura: Leptodac-
tylidae) from Honduras.—Bulletin Southern
California Academy of Sciences 87:50—56.

Smithe, E B. 1975-1981. Naturalist’s color guide, part
I. Color guide. The American Museum of Nat-
ural History, New York, 182 color swatches.

Appendix I
Comparative material examined

Eleutherodactylus anciano. WHonduras—Ocote-
peque: El Chagiiit6n, KU 208999—9001, ROM 18076—
80.

Eleutherodactylus aurilegulus. Honduras—Atlanti-
da: Lancetilla, AMNH 54792—96, MCZ 16184-88,
16280, 16284—85, 17448—50, 21270; 2.0 km SE Lan-
cetilla, TCWC 30129; mountains S of Lancetilla, MCZ
16191; 7 km NW Las Mangas, UF 90216; Quebrada
La Muralla, SMF 77636—39; Quebrada de Oro, KU
209002-—32, 209033 (3), LACM 137286—97; Tela,
UMMZ 70329 (7). Colén: Balfate, AMNH 45696—
703, 45718; Trujillo, CM 63920-21. Olancho: Que-
brada La Calentura, USNM 343704; Quebrada Las
Cantinas, USNM 343710-—14; between El Dictamo and

522

Parque Nacional La Muralla Centro de Visitantes,
USNM 343625-36; Rio de Enmedio, USNM 529987-—
88; Quebrada La Habana, USNM 343705—09; Quebra-
da de Las Mesetas, USNM 343695-—702; Quebrada del
Monte Escondido, USNM 343703; near Parque Na-
cional La Muralla Centro de Visitantes, USNM
343622-—24; Quebrada El Pinol, USNM 343657-—94;
confluence of quebradas El Pinol and Las Cantinas,
USNM 343638—56; near Los Planes, USNM 343637;
Quebrada Salitre Lajas, USNM 343715-—16; Sendero
El Pizote, USNM 343717-19. Yoro: Monte Mataderos,
MCZ 21287; Portillo Grande, FMNH 21859-60,
34696-97, MCZ 21273-75, 21289, UMMZ 77852 (3);
Santa Marta, FMNH 21857-58, MCZ 21271-72,
21288, UMMZ 77851 (3); Subirana Valley, FMNH
21782, MCZ 21283-86; 6.6 km S Yoro, MVZ
171375-76, USNM 217582-83; ca. 3 km W Yoro,
TCWC 23619; ca. 32 km W Yoro, MVZ 175800,
USNM 217584-85.
Eleutherodactylus “‘rugulosus.

3°

Honduras—Co-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

mayagua: between La Mision and Trincheras, AMNH
54752; Siguatepeque, MCZ 26421. Cortés: Agua Azul,
AMNH 54951; Rio Guayabal, AMNH 54751; Haci-
enda Santa Ana, FMNH 4661 (3), 4663, 4665—69,
4691-93, 4695-97, MCZ 17433-34, 21282, SMF
29865: Canon Santa Ana, FMNH 4671, 4674-75:
Lago de Yojoa, MSUM 4540, 4637. El Paraiso: ca. 6
km E Danli, TCWC 23811; Danli, BYU 18209; Mon-
serrat, AMNH 54814; Montana del Volcan, MCZ
26433-35. Francisco Morazan: Agua Amarilla,
AMNH 54883; San Francisco, AMNH 54750; Cerro
Uyuca, MCZ 26440; Rio Yeguare Valley, MCZ
25954-56, UMMZ 94055 (6); El Zamorano, MCZ
26376—77, 26469. Intibuca: 5 km NE Jestis de Otoro,
TCWC 23810. Santa Barbara: Montafia de Santa Bar-
bara, AMNH 55309-12. ““HONDURAS,” ZMB
13202 (holotype of Hylodes laevissimus Werner). Ni-
caragua—Atlantico Sur: Rio Chiquito, SMF 77825.
Granada: Volcan Mombacho, SMF 78222—28. Mata-
galpa: Selva Negra, SMF 78207-21.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
112(3):523—530. 1999.

Taxonomic status and geographic distribution of
Bryconamericus eigenmanni Evermann & Kendall, 1906
(Characiformes: Characidae)

Amalia M. Miquelarena and Adriana E. Aquino

(AM) Museo de La Plata, Paseo del Bosque s/n, La Plata 1900, Argentina;
(AM, AA) Instituto de Limnologia “‘Dr. Raul A. Ringuelet’’, C.C.
712, La Plata (1900), Argentina;

(AA) Department of Ichthyology, American Museum of Natural History,
Central Park West at 79th Street, New York, NY 10024, U.S.A.

Abstract.—Bryconamericus eigenmanni Evermann & Kendall (1906) is re-
described on the basis of additional material collected at the type locality, Rio
Primero, Province of Cérdoba, central Argentina. Evidence supporting the va-
lidity of B. eigenmanni based on morphometric, meristic, and osteological char-
acters is discussed. The geographic distribution of B. eigenmanni is analyzed
and the species is compared with Bryconamericus theringi, which some authors
suggested was conspecific. Bryconamericus eigenmanni can be distinguished
from B. iheringi by the number of maxillary teeth, premaxilla shape, body
depth, caudal peduncle length, orbital diameter, and the secondary sexual di-
morphism of the pelvic fin of males. The known range of B. eigenmanni sug-
gests that it is endemic to endorrheic drainage basins of central Argentina.

Resumen.—Se redescribe Bryconamericus eigenmanni en base a material
adicional colectado en la cuenca de la localidad tipo, Rio Primero, Provincia
de Cérdoba, Argentina central. Se discute evidencia apoyando la validez de B.
eigenmanni en base a caracteres morfométricos, meristicos y osteolégicos. As-
imismo, se analiza la distribuci6n geografica y se compara con Bryconamericus
itheringi, la cual fue sugerida como coespecifica por algunos autores. Brycon-
americus eigenmanni puede distinguirse por el nimero de dientes maxilares,
forma del premaxilar, diametro orbitario, altura del cuerpo, longitud del pe-
dunculo caudal, y el dimorfismo sexual secundario en la aleta pélvica del ma-
cho. El rango de distribuci6n conocido de B. eigenmanni sugiere que es en-
démica de cuencas endorreicas de la regién central de Argentina.

The characid Bryconamericus eigenman-
ni (Evermann & Kendall 1906:83) was
originally described on the basis of two
specimens from the Rio Primero, the main
tributary of an endorrheic drainage basin of
the Province of Cérdoba, in central Argen-
tina. The taxonomic status of this species
has been regarded as unclear by a number
of authors primarily because of its close
similarity with B. iheringi, a species wide-
spread through the Plata basin. This simi-
larity was noted by Eigenmann (1927:379),

although he preferred to recognize the spe-
cies as distinct, a practice followed by sub-
sequent authors (e.g., Ringuelet et al. 1967:
116, Géry 1977:390). The lack of addition-
al collections of B. eigenmanni prevented
further analysis of the question. Malabarba
& Kindel (1995:684) explicitly noted the
need for a statistical comparison of popu-
lation samples from within the areas of dis-
tribution of both nominal species. However,
these authors added uncertainty to the iden-
tity of B. eigenmanni by erroneously citing

524

its type locality as being within the Rio Pa-
rana system, when it is actually a separate
drainage.

Bryconamericus eigenmanni, first as-
signed to Astyanax by Evermann & Kendall
(1906:83), was transferred to Bryconamer-
icus by Eigenmann (1910:434), who had
previously proposed the genus (in Eigen-
mann et al. 1907:139). The current defini-
tion of the genus follows Eigenmann
(1927), and was summarized by Vari & Sie-
bert (1990:516) and Malabarba & Kindel
(1995:679). These authors, among others,
pointed out the need for a reconsideration
of the monophyly of Bryconamericus, in-
cluding an evaluation of the taxonomic sta-
tus of the nominal species assigned to the
genus. Given those questions, we have fo-
cused on the taxonomic identity of the spe-
cies B. eigenmanni as part of a comprehen-
Sive revision of the species of the genus
Bryconamericus in Argentina. We present
evidence supporting the distinctiveness of
B. eigenmanni, redescribe the species, com-
pare it with B. theringi, and discuss its geo-
graphic distribution.

Material and Methods

Most of the examined material was col-
lected by members of the Laboratory of
Ichthyology of the Institute of Limnology
“Dr. Rat A. Ringuelet’’? (ILPLA) during
field trips to the Province of Cordoba, Ar-
gentina, using nets and the ichthyocide
Pronoxfish. Measurements were taken using
calipers to the nearest 0.05 mm. Caudal-pe-
duncle length was measured from the base
of the last anal-fin ray to the last perforated
scale of the lateral line. Osteological prep-
arations were made following Dingerkus &
Uhler (1977). Meristic data are presented as
ranges; the mean is indicated between pa-
rentheses when necessary. Counts of gill-
rakers on the lower portion of the arch in-
cludes the element at the corner between
inferior and superior arches. The anal-fin
ray count includes the last ray divided to its
base as a single element. The lateral-line

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

count includes the last pored scale. The fol-
lowing abbreviatures are used: C & S,
cleared and stained; EK females; HL, head
length; M, males; n, number of specimens;
and SL, standard length. Institutional ab-
breviations used in the listing of material
examined follow Leviton et al. (1985). The
drainage basins follow the terminology of
Mazza (1962).

Bryconamericus eigenmanni
(Evermann & Kendall, 1906)
Fig. la, b

Astyanax eigenmanni Evermann & Kendall,
1906: 83, fig. 1.

Bryconamericus eigenmanni.—Eigenmann,
1910: 434; 1927: 379, pl. 68, fig. 1, pl.
69, fig. 13.—Pozzi, 1945: 255.—Ringue-
let & Aramburu, 1962: 29.—Ringuelet et
al., 1967: 116.—Ringuelet, 1975: 82.—
Géry, 1977: 390.—Malabarba & Kindel,
1995: 684, fig. 3—-Miquelarena & Aqui-
no, 1995: 560.—Lé6pez et al., 1996: 6.

Material examined (all localities in Ar-
gentina unless noted otherwise).—Brycon-
americus eigenmanni: USNM 55570 (ho-
lotype of Astyanax eigenmanni Evermann
& Kendall, 1906), Rio Primero, Cérdoba,
1903-1904. J. W. Titcomb. ILPLA 970 (8
ex.), MLP 6-VII-83-22 (8 ex.), second Ar-
royo Mallin-Tanti, Cérdoba. ILPLA 971
(11 ex. + 8 c&s), MLP 6-VII-83-15 (55
ex.), Arroyo Cachimayo, near Taninga,
Cordoba. ILPLA 972 (17 ex.), Bialet Mas-
sé, Rio Cosquin, Cérdoba. ILPLA 973 (11
ex. + 4 c&s), MLP 6-VII-83-16 (54 ex.),
Arroyo Nifanquil, Cérdoba. ILPLA 974 (5
ex.), MLP 6-VII-83-20 (26 ex.), stream at
camping ground Villa Giardino, Cordoba.

Bryconamericus iheringi: ILPLA 297
(50 ex.), ILPLA 298 (7 ex.), ILPLA 595
(27 ex.), Laguna Chascomius, Buenos Aires.
ILPLA 975 (18 ex.), Laguna Cochic6,
Buenos Aires. MCP 11481 (3 ex.), Arroio
do Ouro, along road between Feliz and Ca-
xias do Sul (RS 452), almost 100 m above
bridge, Jacui drainage, Rio Grande do Sul,
Brazil. USNM 310959, Arroyo Pelotas at

VOLUME 112, NUMBER 3

525

Bis. 1.

Bryconamericus eigenmanni, ILPLA 970. Left lateral view. a, female, SL = 61.0 mm; b, male, SL

= 50.7 mm. Second Arroyo Mallin-Tanti, Cordoba, Argentina.

bridge crossing, along road between Pelotas
and Porto Alegre, near Pelotas, Rio Grande
do Sul, Brazil. USNM 310947, Rio Grande
do Sul, Brazil. USNM 310979, Arroyo Sar-
andi, Rio Grande do Sul, Brazil.

Description

Body elongate. Dorsal and ventral pro-
files slightly convex. Head short and robust;
snout deep and rounded, blunt. Caudal pe-
duncle deep and moderately long. Anal-fin
origin along vertical line through base of
last dorsal-fin ray. Mouth slightly inferior.
Maxilla approaching or reaching vertical
line through anterior border of eye. Infraor-
bitals well developed, 6, third largest. Ven-
tral and posterior margins of infraorbitals
reaching horizontal and vertical limbs of
preopercle.

Posterior border of dorsal, pectoral, and
caudal fins gently rounded. Pelvic-fin tip
rounded in males, pointed in females. Dor-
sal-fin rays ii, 7—9, typically 8. Second un-
branched and first branched dorsal-fin rays
almost equal in length. Tip of adpressed
dorsal fin not reaching adipose fin. Adipose
fin small. Pectoral-fin rays i, 10—12; these
occasionally followed by one or two un-
branched rays. Ventral-fin rays i-ii, 6-8,

typically u, 7. Principal caudal-fin rays
17+2; dorsal caudal-fin procurrent rays 10—
12, and ventral procurrent rays 8—10. Anal-
fin rays iv, 15-17.

Cycloid scales regularly distributed on
body. Single row of scales on base of 9 to
12 anterior branched anal-fin rays. Scales
present on caudal-fin base. Lateral line
complete, 38 to 39 perforated scales. Rows
of scales from dorsal-fin origin to lateral
line 5—6, and 4.5—5.5 from lateral line to
anal-fin origin. Vertebrae 36—38, typically
37. Supraneurals 4—6, typically 5. Gill rak-
ers 7—9+10-—12. Morphometric characters
presented in Table I.

Teeth.—Maxilla elongate, with 3 to 6
teeth along ventral margin (Fig. 2a). Each
tooth typically tricuspidate, though teeth
with 1 or 4 cusps also occur. Premaxilla as-
cending process short and slightly curved
(Fig. 2b). Premaxilla with an outer row of
4 or 5 tricuspidate teeth, and an inner row
of 4 teeth, with 4 or 5 cusps each. Dentary
with 8 to 12 teeth: 3 or 4 larger anterior
teeth with 4 or 5 cusps, followed by a series
of smaller teeth, usually with 1 or 3 cusps
(Fig. 2c).

Color in alcohol.—Ground color light
tan. Body and head finely dotted, more con-

526

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Morphometric data of Bryconamericus eigenmanni presented as percent of standard length (3-11),
distance between origins of pectoral and pelvic fin (12), and head length (13-15) (LPLA 970; ILPLA 971;

ILPLA 973; TE PEA 974)°G3 ML 72):

Character Range xX SD n

1 Standard length 46.0—73.0 59.6 5.8 20

2 Total length 58.0—88.0 72.4 6.9 20

3 Body depth 28.7—33.0 Sus sles 20

4 Head length 24.1-27.2 D0) 0.8 20

5 Caudal peduncle length 11.4—15.4 13e2 0.8 20

6 Caudal peduncle length 17.9-23.2 20.2 a 19

7 Predorsal distance 47.1—53.1 5108) 2.0 20

8 Prepelvic distance 42.5-49.1 46.4 1.8 20

9 Preanal distance 58.8—68.0 63.2 2.1 20

10 Pectoral-pelvic distance 19.5—26.8 23.6 1.6 20
11 Pelvic-anal distance 15.7—25.8 18.9 2.0 20
12 Pectoral length 73.8-125.2 103.9 Wei 17
13 Orbital diameter 21.9—30.7 2S) 1.3) 20
14 Interorbital width 30.3-39.6 55:9 2.4 20
15 Snout 16.2—27.4 2226 2.6 20

centrated along posterior scale margins
forming overall reticulated pattern more ev-
ident above trunk midlateral line. Dark pig-
mentation on head, opercle, supraorbitals,
maxilla, and along body dorsal of midline
relatively more intense. Vertically elongate
dark brown humeral mark between second
and fourth lateral-line scales. Midlateral
band extends from near vertical line
through dorsal-fin origin, along second
scale row dorsal to lateral line. Band faint
anteriorly and becoming darker posteriorly;
covering one-third of caudal peduncle depth
below adipose fin, and terminating as tri-
angular spot on caudal-fin base. Paired fins
light brown. Dorsal and anal fins dark
brown, with distal portions of first rays
lighter. Middle rays of caudal fin dark.
Sexual dimorphism.—First to fourth or
fifth branched pelvic-fin rays of males
curved to form a basket-like structure (Fig.
3a). In females ventral fins almost flat (Fig.
3b). Pectoral and pelvic fins larger in males,
where tip of pectoral fin reaches or slightly
surpasses pelvic-fin origin. Pelvic fin reach-
es anal-fin origin. In females tip of both
pectoral and pelvic fins separated from pel-
vic- and anal-fin origins, respectively, by
two or more scale rows. Margin of anal fin
almost straight in males and slightly con-

cave in females, in which first branched
rays relatively longer. Males with bony
hooks on pelvic and anal fins. Anal-fin
hooks short, curved (Fig. 4a), forming
smaller angle relative to ray axis than one
formed by longer and more pointed pelvic-
fin hooks (Fig. 4b).

Distribution.—Known only from the
Province of Cordoba in central Argentina
(31°00’S; 65°02'’W). It was collected in the
endorrheic drainage basins of the Rio Pri-
mero and the Rio Pichanas, to the east and
west respectively of the Sierras Grandes.

Ecology.—Collected in shallow creeks
(ca. 0.5 m deep) over sandy to rocky bot-
tom, and in deeper pools (up to 1.5 m deep)
of the same streams. Water in these envi-
ronments has the anion CO,H™ and the cat-
ions Ca** and Na* as the most abundant
elements (Menni et al. 1984). Following
Cabrera’s (1976) phytogeographical scheme,
the distribution range of B. eigenmanni is
the “‘Chaquefio-serrana’”’ area of the ““Cha-
queno”’ dominion.

Discussion and Conclusions

The original description provided by Ev-
ermann & Kendall (1906) does not permit
an unequivocal discrimination of B. eigen-

VOLUME 112, NUMBER 3

Pig. 2.

manni from congenerics. Eigenmann
(1927) noted its close similarity to B. iher-
ingi, however, indicating that B. eigenmanni
differed in having less convex dorsal and
ventral body profiles, longer pectoral fins,
and wider naked area below the infraorbi-

527

Eee ee Oe enero Orta eee |

Jaws of Bryconamericus eigenmanni, ILPLA 973. a, maxilla; b, premaxilla; c, dentary. Scale bar: 1 mm.

tals. Ringuelet et al. (1967:116) and Mala-
barba & Kindel (1995:684) pointed out that
neither the meristic nor morphometric val-
ues of B. eigenmanni clearly differed from
those of B. iheringi. Malabarba & Kindel
(1995) erroneously considered the type lo-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

BigS.
bar: 5 mm.

cality of B. eigenmanni to be within the rio
Parana basin, which implied overlapping
geographical ranges between the two spe-
cies. In actuality the type locality of B. ei-
genmanni is in an endorrheic drainage basin
of central Argentina, separate from the Pa-
rana/Plata basin.

Sexual dimorphism in ventral-fin shape of Bryconamericus eigenmanni. a, male; b, female. Scale

The two type specimens of B. eigenman-
ni had been the only source of information
on the species, leading to questions about
its identity. The examination of more ex-
tensive samples of specimens from the type
drainage of B. eigenmanni allows us to con-
firm the distinctiveness of the species and

VOLUME 112, NUMBER 3

Des. i one

Fig. 4. Bony branched fin ray hooks in males of
Bryconamericus eigenmanni. Left side, lateral view,
tip of ray to the right. a, anal fin; b, ventral fin. Scale
bar: 0.5 mm.

its restricted geographic distribution in cen-
tral Argentina.

Bryconamericus eigenmanni differs from
B. theringi by the following combination of
characters: higher number of maxilla teeth
(3-6 vs. 2—4 in B. iheringi) (Fig. 2a) (Mi-
quelarena & Aquino, 1995:567); premaxilla
ascending process pointed and slightly
curved (Fig. 2b); in B. itheringi, it is blunt
and strongly curved (Miquelarena 1986:
30-32, Miquelarena & Aquino 1995:567);
shallower body depth (28.7—33.0 (31.1) %
Sis 2 — 20 vs. 33.7-38.3 (35.9) % SL; n
= 19); shorter orbital diameter (21.9—30.7
(27.9) % HL; n = 20; 31.6—37.9 (34.9); n
= 19); longer caudal-peduncle length
(17.9—23.2 (20) % SL; n = 19 vs. 14.2-—
18.2 (15.9) % SL; n = 19); in males of B.
eigenmanni, the fins are distinctly second-
arily sexually dimorphic, with the first to
fourth or fifth lateral branched rays curved
to form a basket-like structure (Fig. 3a). In
males of B. iheringi, the pelvic fins are al-
most flat, the same condition in females of
both species.

According to the ichthyogeographical
scheme proposed by Ringuelet (1975), the
range of B. eigenmanni falls in the contact
zone between the Andean and Paranensean
domains, encompassing the so-called Sier-
ras Grandes, which represents the main oro-

529

graphic system of the region. Based on the
similarity in the composition of fish faunas
to both slopes of the Sierras Grandes, Men-
ni et al. (1984:28-—29) proposed a demar-
cation between those domains west to these
hills, a hypothesis which is also supported
by the distribution range of B. eigenmanni.

Acknowledgments

We would like to thank S. Jewett, R.
Vari, and S. Weitzman (USNM) for their
help in making available for examination
the material under their care, and C. Lucena
(MCP) for the loan of material. We are in-
debted to L. Protogino and H. Lopez for
their support and comments on the earlier
versions of the manuscript, and to S. Schae-
fer for his critical reading and checking of
the English. We also thank the two anony-
mous reviewers whose comments greatly
improved the manuscript.

Literature Cited

Cabrera, A. L. 1976. Regiones fitogeogrdaficas de la
Republica Argentina.—Enciclopedia Argentina
de Agricultura y Jardineria, 2nd edition, 2(1):
1-85.

Dingerkus, G., & L. D. Uhler. 1977. Enzyme clearing
of alcian blue stained whole small vertebrates
for demonstration of cartilage —Stain Technol-
ogy 52(4):229—2372.

Eigenmann, C. H. 1910. Catalogue of the fresh-water
fishes of tropical and south temperate Ameri-
ca.—Reports, Princeton University Expedition
to Patagonia 1896-1899, 3(2), Zoology:375—
S10,

. 1927. The American Characidae.—Memoirs

of the Museum of Comparative Zoology of Har-

vard 43(4):311—428.

, W. L. McAtee, & D. P. Ward. 1907. On further
collections of fishes from Paraguay.—Annals of
the Carnegie Museum 4(2):110—157.

Evermann, B. W., & W. C. Kendall. 1906. Notes on
collection of fishes from Argentina, South
America, with descriptions of 3 new species.—
Proceedings of the United States National Mu-
seum 31:67—108.

Géry, J. 1977. Characoids of the world. T.EH. Publi-
cations, Neptune City, New Jersey, 672 pp.

Leviton, A. E., R..H.. Gibbs, Jn, E. Heal. &'C. .E.
Dawson. 1985. Standards in ichthyology and
herpetology: Part I. Standard symbolic codes
for institutional resource collections in herpe-

530

tology and ichthyology.—Copeia 1985:802—
835.

Lopez, H. L., L. C. Protogino, & A. E. Aquino. 1996.
Panorama de la Ictiofauna Continental de la Ar-
gentina: Santiago del Estero, Catamarca, Cor-
doba, San Luis, La Pampa y Buenos Aires.—
Aquatec 3:1—14.

Malabarba, L. R., & A. Kindel. 1995. A new species
of the genus Bryconamericus Eigenmann, 1907
from southern Brazil (Ostariophysi: Characi-
dae).—Proceedings of the Biological Society of
Washington 108(4):679—86.

Mazza, G. 1962. Recursos hidraulicos superficiales.—
Serie Evaluacion de los Recursos Naturales de
la Argentina (Primera etapa) 4(1):1—459.

Menni, R. C., H. L. Lépez, J. R. Casciotta, & A. M.
Miquelarena. 1984. Ictiologia de areas serranas
de Cordoba y San Luis (Argentina).—Biologia
Acuatica 5:1—63.

Miquelarena, A. M. 1986. Estudio de la denticién en
peces caracoideos de la Republica Argentina.—
Biologia Acuatica 8:1—60.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

, & A. E. Aquino. 1995. Situaci6n taxonomica
y geografica de Bryconamericus thomasi Fowl-
er, 1940 (Teleostei, Characidae).—Revista Bras-
ilera de Biologia 55(4):559—569.

Pozzi, D. J. 1945. Sistematica y distribuci6n de los
peces de agua dulce de la Republica Argentina.
Anales de la Sociedad Argentina de Estudios
Geograficos.—_GAEA 7(2):239—292,

Ringuelet, R. A. 1975. Zoogeografia y ecologia de los
peces de aguas continentales de la Argentina y
consideraciones sobre las areas ictiologicas de
América del Sur—tEcosur 2(3):1—122.

, & R. H. Aramburu. 1962. Peces argentinos de

agua dulce.—Agro 3(7):1—98.

; , & A. Alonso de Aramburu. 1967.
Los peces argentinos de agua dulce.—Comision
de Investigaciones Cientificas de la Provincia de
Buenos Aires, La Plata, 602 pp.

Vari, R. P, & D. J. Siebert. 1990. A new, unusually
sexually dimorphic species of Bryconamericus
(Pisces: Ostariophysi: Characidae).—Proceed-
ings of the Biological Society of Washington
103:517-524.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112G6)73531-535: 1999.

Caridina clinata, a new species of freshwater shrimp
(Crustacea: Decapoda: Atyidae) from northern Vietnam

Yixiong Cai, Nguyen Xuan Quynh, and Peter K. L. Ng

(YC, PKLN) Department of Biological Sciences, National University of Singapore,
Lower Kent Ridge Road, Singapore 119260, Singapore; (NXQ)
Department of Invertebrate Zoology,

National University of Hanoi, 90, Nguyen Trai Road, Thanh Xuan, Hanoi, Vietnam

Abstract.—Caridina clinata, a new species of atyid shrimp, is described from
northern Vietnam. The new species is characterised by a short, sloping rostrum
which reaches to or slightly exceeds the distal margin of the basal antennular
segment, the shape of the sexual appendages, and its large egg size.

Eight species and subspecies of fresh-
water shrimps of the family Atyidae have
been previously reported from Vietnam:
Caridina vietnamensis Dang, 1967, C. sub-
nilotica Dang, 1975, C. acuticaudata Dang,
1975, C. flavilineata Dang, 1975, C. serrata
serrata Stimpson, 1860, C. serrata cuc-
phuongensis Dang, 1980, C. tonkinensis
Bouvier, 1919, and C. cantonensis Yu, 1938
(Dang 1967, 1975, 1980). Cai (1996) sub-
sequently synonymized Caridina vietna-
mensis with Neocaridina palmata palmata
(Shen, 1948). Recently, a revision of the
Caridina serrata species group (Cai & Ng
1999) indicated that the specimens illus-
trated as “C. serrata serrata’ by Dang
(1980) probably represent an undescribed
species and C. serrata cucphuongensis
should be elevated to specific rank. Cari-
dina tonkinensis and C. cantonensis have
been shown to have a wide Chinese, Indo-
Chinese and/or Southeast Asian distribution
(Johnson 1961; Ng & Choy 1990a, 1990b;
Cai & Ng, 1999). As a result of those stud-
ies, only four species: C. subnilotica, C.
acuticaudata, C. flavilineata and C. cuc-
phuongensis are known to be endemic to
Vietnam.

Recently, we had an opportunity to ex-
amine several lots of specimens of Caridina
collected from northern Vietnam. These
specimens proved to belong to an unde-

scribed species. Specimens are deposited in
the Zoological Reference Collection of the
Raffles Museum, National University of
Singapore (ZRC); Zoological Museum of
Hanoi University, Vietnam (ZMHU); Insti-
tute of Zoology, Academia Sinica, Beijing,
China (IZAS); National Museum of Natural
History, Leiden, The Netherlands (RMNH);
National Museum of Natural History,
Smithsonian Institution, Washington, D.C.,
USA (USNM); and Muséum National
d’Historie Naturelle, Paris, France
(MNHN). The abbreviation cl is used for
carapace length, measured from the post-
orbital margin to the posterior margin of the
carapace.

Caridina clinata, new species
(Figs. 1, 2)

Material examined.—Holotype: male, cl
3:3 aume(Z RC) (998.5560), \diutchpCusc
Phuong National Park, Ninh Binh Province,
northern Vietnam, 16 Sep 1997. Paratypes:
3 males, cl 3.0—3.4 mm, 7 females, cl 3.7—
4.2 mm (ZRC 1998.551—560), 2 males, cl
3.0—3.2 mm, 2 females, cl 4.0—4.1 mm
(IZAS), 1 male, cl 3.4 mm, 2 females, cl
3.4—3.6 mm (ZMHU), 1 male, cl 2.8 mm,
2 females, cl 3.2—4.0 mm (RMNH), 1 male,
cl 2.8 mm, 2 females, cl 3.8—3.9 mm
(USNM), same data as holotype. None-

532

Biers ly
1998.551); B, anterior portion of cephalothorax, lateral view, female, cl 4.0 mm (ZRC 1998.552); C, anterior
portion of cephalothorax, lateral view, female, cl 4.2 mm (ZRC 1998.553); D, right antennular peduncle; E,
right scaphocerite; E right third maxilliped; G, right first pereiopod of male; H, right first pereiopod of female;
I, right second pereiopod; J, preanal carina, lateral view; K, uropodal diaeresis. Scales: A, B, C = 1 mm; D, E,
E G, H, I, J = 0.5 mm. K = 0.1 mm.

types: 1 ovigerous female, cl 4.2 mm
(ZRC), approximately 8.7 km from Nho
Quan on Phu Ly to Cucphuong Road, Ba
Dien district, Ninh Binh Province, northern
Vietnam, coll. jH., Ha.Ne dD. €.\J. Yeo
(from fisherman), 16 Sep 1997.
Description.—Rostrum (Fig. 1A—C)
short, reaching to or slightly exceeding dis-
tal margin of basal antennular segment, not
reaching as far as middle of second seg-
ment, sloping ventrad anteriorly; armed
dorsally with 13—21 (mode 14-16) teeth,
including 3—5 (mode 4) teeth on carapace
posterior to orbital margin; armed ventrally
with 2—5 (mode 2-3) teeth. Suborbital an-
gle acute, completely fused with antennal
spine; pterygostomian margin rounded.
Abdomen with sixth somite 0.4 times as
long as carapace, 1.5 times as long as fifth

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Caridina clinata, new species, paratypes. A, cephalothorax; lateral view, male, cl 3.2 mm (ZRC

somite, slightly shorter than telson. Telson
(Fig. 2A) (not including marginal spines)
slightly longer than sixth somite, tapering
posteriorly, lacking posteromedian projec-
tion; 1 pair of dorso-lateral spines near dis-
tal end, 5 pairs of spiniform setae on distal
margin, lateral pair subequal to or slightly
shorter than intermediate pairs; preanal an-
gle rounded, symmetrical lacking spine.
Eyes well developed. Atennular peduncle
(Fig. 1D) 0.6 times length of carapace; bas-
al segment slightly shorter than half length
of peduncle; second segment distinctively
longer than third. Stylocerite not reaching
distal margin of of basal antennular seg-
ment. Scaphocerite (Fig. 1E) ovate, reach-
ing beyond distal end of antennular pedun-
cle; 3.4 times as long as broad. Branchial
formula as for genus. Epipods present on

VOLUME 112, NUMBER 3

first 4 pereiopods. Third maxilliped (Fig.
1F) reaching slightly beyond distal end of
antennular peduncle, ending in single ter-
minal claw; exopod reaching to fourth of
penultimate segment; ultimate segment as
long as penultimate.

First pereiopod (Fig. 1G, H) short, ro-
bust, reaching slightly beyond distal margin
of basal antennular segment; chela about
2.3 times as long as broad in male (Fig.
1G), 2.0 times in female (Fig. 1H); fingers
as long as palm in male, 0.7 times as long
as palm in female; carpus 1.5 times as long
as high in male, 1.2 times as long as palm
in female; merus slightly shorter than palm;
merus 2.3 times in female and 1.3 times in
male as long as broad. Second pereiopod
(Fig. 11) long, slender, reaching end of an-
tennular peduncle; chela about 2.5 times as
long as broad, fingers about 1.6 times as
long as palm; carpus slightly longer than
chela, about 4.7 times as long as high. Third
pereiopod (Fig. 2B) reaching slightly be-
yond end of antennular peduncle; dactylus
(Fig. 2C) terminating in 2 spines, bearing 5
spines on posterior margin; propodus 9
times as long as broad, about 3.6 times as
long as dactylus, with numerous spinules on
posterior margin. Fifth pereiopod (Fig. 1D)
reaching slightly beyond distal margin of
second antennular segment; dactylus (Fig.
1E) ending in curved claw, with row of 45—
49 closely spaced spinules; propodus slen-
der, 10 times as long as broad, about 3.3
times as long as dactylus (including termi-
nal claw), with numerous spinules on pos-
terior margin.

Endopod of male first pleopod (Fig. 2F)
2.8 times as long as broad, half as long as
exopod, rounded distally; with appendix
interna exceeding terminal margin of en-
dopod by 0.3 its length; endopod with long
plumose setae on outer and distal margins,
with short simple setae on inner margin.

Appendix masculina of male second ple-
opod (Fig. 2G) extending to proximal 0.6
length of endopod, with some short spi-
nules on outer surface and some long spi-

533

nules on distal surface; appendix interna
about 0.8 length of appendix masculina.

Uropodal diaeresis with 14-17 spinules.

Eggs large, ranging in dimensions from
0.60—0.75 to 1.10—1.15 mm.

Habitat.—This new species was found in
a ditch with a sandy substratum and clear
flowing water from the forest. Other deca-
pods found in the ditch are the potamid
crabs Potamiscus cuphuongense Dang,
1975 and Potamiscus kimboiense (Dang,
1975) eDs Gii> Yeo; pers. comm.). The lat-
ter species was originally placed in the ge-
nus Ranguna Bott, 1966, but is now re-
garded as a junior subjective synonym of
Potamiscus Alcock, 1909 (Ng & Naiyanetr
1993):

Color (from a color photograph taken
several hours after preservation in 10% for-
malin).—Body yellowish to grey and dark
grey. There is one transverse black stripe
which is irregularly broken at the lateral
posterior margin of the carapace. Some ir-
regular black spots are present on the ven-
trolateral parts of the carapace. The ventro-
lateral portion of the abdominal tergal pleu-
ra are mottled with black spots adjacent to
the articular knobs at the posterior bases of
the pleura. Black transverse stripes are pre-
sent at the posteroventral ends of the first
four pleura. The dorsum of the third ab-
dominal somite has a black transverse
stripe. The antenna, antennule and telson
are yellowish to orange, and the pereiopods
are translucent to yellowish.

Etymology.—The new species is named
as ‘clinata’, Latin, meaning “‘sloping’’, al-
luding to the shape of rostrum.

Remarks.—Caridina clinata new species,
is most similar to C. flavilineata from Nam-
ha, northern Vietnam, in the rostral formula
and egg size. It differs from the latter, how-
ever, by the form of the rostrum which is
short and sloping (vs. long and straight); the
dactylus of fifth pereiopod has 45 to 49
denticulate spinules (vs. 20—30); the endo-
pod of the male first pleopod reaching to
half the length of the exopod (vs. 1/3 in C.
flavilineata); the appendix masculina of the

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fis. 2.
right third pereiopod; C, dactylus of right third pereiopod; D, right fifth pereiopod; E, dactylus of right fifth pereiopod;
E left male first pleopod; G, left male second pleopod. Scales. A, C, E, E G = 0.2 mm, B, D = 0.5 mm.

male second pleopod reaching to 2/3 the
length of the endopod (vs. reaching to half
the length); and the appendix interna of the
male second pleopod is 2/5 as long as the
appendix masculina (vs. more than half the
length) (cf. Dang 1975:70, fig. 5, Dang
1980:412, fig. 235). Dang (1980) had de-
scribed C. cucphuongensis from the same
area as C. clinata, although we did not
manage to obtain fresh specimens from
there. Caridina clinata, can easily be sep-
arated from C. cucphuongensis by the form
of the rostrum and rostral formula [(2—
5)13-—21/2—5 vs. (1—3)5—-9/0-—2 in C. cuc-
phuongensis]; the short stylocerite which
does not reach the end of the basal segment
of the antennular peduncle (vs. reaches be-
yond); and the symmetrical shape of the
preanal carina (vs. asymmetrical) (cf. Dang
1980).

Caridina clinata, new species, paratype, male, cl 3.2 mm (ZRC 1998.551). A, distal portion of telson; B,

Acknowledgments

Thanks are due to Prof. Dang Ngoc
Thanh (Vietnam National Centre for Natu-
ral Science and Technology) for his support
of the present study, and for his hospitality
when one of us (PKLN) visited Vietnam; as
well as Mr. H. H. Ng and D. C. J. Yeo for
their kind help. This project is partially sup-
ported by a research grant (RP950362) to
the third author, and this paper is contri-
bution No. 98/30 from the Systematics &
Ecology Laboratory, Department of Biolog-
ical Sciences, National University of Sin-
gapore.

Literature Cited

Alcock, A. 1909. Diagnoses of new species and vari-
eties of freshwater crabs. Nos. 1—4.—Records
of Indian Museum 3:243—252, 375-381.

VOLUME 112, NUMBER 3

Bott, R., 1966. Potamiden aus Asien (Potamon Savig-
ny und Potamiscus Alcock) (Crustacea, Deca-
poda).—Senckenbergiana biologica, Frankfurt
47:469—S509, pls. 16-21.

Bouvier, E. L. 1919. Quelques espéces nouvelles de
Caridines.—Bulletin du Muséum National
d’Histoire Naturelle, 1919:330-—336.

Cai, Y. 1996. A revision of the genus Neocaridina
(Crustacea: Decapoda: Atyidae).—Acta Zootax-
onomica Sinica 21:129—160. (in Chinese with
English abstract)

Cai, Y., & N. K. Ng. 1999. A revision of the Caridina
serrata species group, with descriptions of five
new species (Crustacea: Decapoda: Caridea:
Atyidae).—Journal of Natural History (in
press).

Dang, N. T. 1967. Cac Loai moi va giong moi tim thay
trong khu he dong vat khong xuong song nuoc
ngot va nuoc lo mien Bac. Vietnam. Tap san
Sinh Vat-Dia Hoc (New species and genera in
the freshwater and brackish water Invertebrates
of North Vietnam.)—Journal of Biology-Geol-
ogy 6(3—4):155—165. [in Vietnamese with
French summary]

. 1975. Phan loai tom cua nuoc ngot mien bac

vietnam. Tap san Sinh Vat-Dia Hoc (The iden-

tities of North Vietnamese freshwater shrimps

and crabs.)—Journal of Biology-Geology 13(3):

65-78. figs. 1-8. [in Vietnamese with French

summary]

. 1980. Dinh loai dong khong xuong song nuoc

noot bac Viet Nam. Decapoda (The identities of

freshwater invertebrates of North Vietnam). p.

339

380-418, figs. 219-236. Hanoi. Vietnam. [in
Vietnamese]

Johnson, D. S. 1961. Notes on the freshwater crustacea
of Malaya. I. The Atyidae.—Bulletin of Raffles
Museum, Singapore 26:120—153.

Ng, P. K. L., & S. Choy. 1990a. The Caridean Prawns
(Palaemonidae and Atyidae) from the Endau-
Kompin area, Johore-Pahang, Peninsular Ma-
laysia.—Malayan Nature Journal 43(4):302—
5A?

, & . 1990b. Notes on some freshwater

Caridean Prawns (Palaemonidae and Atyidae)

from the Endau-Kompin area, Johore-Pahang,

Peninsular Malaysia.—Raffles Bulletin of Zo-

ology 38(1):11-—20.

, & P. Naiyanetr. 1993. New and recently de-
scribed freshwater crabs (Crustacea: Decapoda:
Brachyura: Potamidae, Gecarcinucidae and Par-
athelphusidae) from Thailand.—Zoologische
Verhandelingen 284:1—117.

Shen, C. J. 1948. On three new species of Caridina
(Crustacea Macrura) from south-west China.—
Contributions of Institute of Zoology, National
Academy of Peiping. 4(3):119-—126.

Stimpson, W. 1860. Prodromus descriptionis animal-
ium evertebratorum, quae in Expeditione ad
Oceanum Pacificum Septemtrionalem, a Repub-
lica Federata missa, C. Ringgold et J. Rodgers,
observavit et descriptsit—Proceedings of Na-
tional Academy of Sciences, Philadelphia 22—
47(97-116).

Yu, S. C. 1938. Studies on Chinese Caridina with de-
scriptions of five new species.—Bulletin of the
Fan Memorial Institute of Biology, Zoology Se-
ries 8(3):271—310; figs. 1-16.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(3):536—540. 1999.

Inclusion of the austral species Pinnotheres politus (Smith, 1869) and
Pinnotheres garthi Fenucci, 1975 within the genus Calyptraeotheres
Campos, 1990 (Crustacea: Brachyura: Pinnotheridae)

Ernesto Campos

Facultad de Ciencias, Universidad Aut6noma de Baja California, Apartado Postal 2300,
Ensenada, Baja California, México.

Abstract.—The limpet crabs Pinnotheres politus (Smith, 1869) known from
Pert to Chile, and P. garthi Fenucci, 1975 known from Brazil to Argentina,
are transferred to the genus Calyptraeotheres Campos, 1990. As result, the
diagnosis of the genus is emended in order to account for the presence or
absence of the minute dactylus of the third maxilliped. Additional characters
that remain diagnostics for Calyptraeotheres are, caparace with two longitu-
dinal cervical depressions, anteriorly arcuate and sharp edged; walking legs 1—
3 similar in shape, walking leg 4 slender and its dactylus longer than other
walking legs; third maxilliped with a robust carpus, larger than the propodus;
abdomen in both sexes with six abdominal somites and telson well separated.
Species of Calyptraeotheres are obligatory symbionts of slipper shells (family

Calyptraeidae).

The study of specimens and published
descriptions and figures indicates that the
austral limpet crabs Pinnotheres garthi Fen-
ucci, 1975 and P. politus (Smith, 1869)
Should be included in the genus Calyp-
traeotheres Campos, 1990. As a result the
original diagnosis of this genus needs to be
emended. However, most of the characters
provided by Campos (1990) to distinguish
Calyptraeotheres remain valid. The follow-
ing Institutions provided specimens for the
present study: National Museum of Natural
History, Smithsonian Institution (USNM);
Museo Argentino de Ciencias Naturales
‘““Bernardino Rivadavia’’ Washington D.C.
(MACN); Laboratorio de Invertebrados, Fa-
cultad de Ciencias, Universidad Aut6noma
de Baja California, Ensenada (UABC). Oth-
er abbreviations used are: MXP3, third
maxilliped; WL, walking legs.

Family Pinnotheridae de Haan, 1833
Calyptraeotheres Campos, 1990

Emended diagnosis (emendations under-
lined).—Female. Carapace arcuate anteri-

orly, sharp-edged; regions ill-defined, with
2 longitudinal cervical depressions from or-
bits to middle of carapace; front slightly
projecting: MXP3 obliquely placed in buc-
cal cavity; ischium and merus fused; palp
2- or 3-segmented, carpus larger than pro-
podus, dactylus, when present, minute, in-
serted distoventrally on propodus; exopod
with thin, unsegmented flagellum. WL1—3
similar in shape; WL4 more slender than
others; propodus of WL1-—2 with tuft of
short stiff setae on distoventral margin; dac-
tyli of WL1-3 similar in shape, acute and
curved at tip, those of WL4 longest and
sword-shaped. Abdomen covering sternum,
with 6 abdominal somites and telson well
separated.

Male: Carapace subpentagonal or subor-
bicular, regions ill-defined, dorsal region
even, with short spaced setae; anterolateral
margin with fringe of simple setae. MXP3
similar to that of female. Abdomen with 6
somites and telson well separated, widest at
third somite, narrowing toward telson.

Distribution.—Northeast Pacific: Mexi-

VOLUME 112, NUMBER 3

co, throughout Gulf of California, and west
coast of Baja California Sur at Bahia Mag-
dalena (C. granti). Southeast Pacific: Bahia
Ancon, Perti to Castro, Isla Chilé6e, Chile
(C. politus). Southwest Atlantic: Brazil, Rio
Grande do Sul; Argentina, Mar del Plata,
Necochea, Golfo San Matias (C. garthi).

Type species.—By original designation,
Fabia granti Glassell, 1933; host, Cruci-
bulum spinosum (Sowerby).

Calyptraeotheres garthi (Fenucci, 1975),
new combination
Figs. 1 E-K 2B

Pinnotheres politus (not Smith, 1869).—
Fenucci, 1971:355—367.

Pinnotheres garthi Fenucci, 1975:167,
169-171, 178, fig. 1A—B, 3D, I; Campos,
1990:365; Martins & D’Incao, 1996:11—
13, fig. 7, 8 14E

Material examined.—2 ovig. females
(MACN 26315, 26316), S. SE Puerto Que-
quén, 8 Feb 1962, in Crepidula sp. on My-
tilus sp., colls. M. Birabén & E. Martinez-
Fontes; 3 females (MACN 29265, 26700),
Necochea, Provincia de Buenos Aires, Apr
1978, coll. I. Pollites.

Distribution.—Southwest Atlantic: Bra-
zil, Rio Grande do Sul; Argentina, Mar del
Plata, Necochea, Golfo de San Matias.

Hosts.—Gastropoda: Calyptraeidae; in
Crepidula unguiformis Lamark, C. protea
Orbigny, and Crepidula sp.

Calyptraeotheres politus (Smith, 1869),
new combination
Figs. 1C—D, 2C

Restricted synonymy:

Pinnotheres politus.—Schmitt et al., 1973:
81-82; Fenucci, 1975:166; Saelzer &
Hapette, 1986:63-60; Campos, 1990:
365: Marquez & Pohle, 1995:349.

Material examined.—1 ovig. female, Ba-
hia Ancon, Peri, coll. R. E. Coker (USNM
40448).

Distribution.—Southeast Pacific: Bahia
Ancon, Pert to Castro, Isla Chil6e, Chile.

537

Hosts.—Gastropoda: Calyptraeidae: Cre-
pidula fecunda Gallardo [= C. dilatata (not
Lamark)] and Calyptraea sp (see Schmitt et
al. 1973). Gallardo (1979) split C. dilatata
Lamark in two species, C. dilatata from Af-
rica and C. fecunda Gallardo from Chile
(see Gallardo 1979, Hoagland 1983).

Taxonomic Remarks on the Genus
Calyptraeotheres

Several characters support the inclusion
of Pinnotheres granti and P. politus in the
genus Calyptraeotheres. These include the
presence of two longitudinal depressions on
the carapace, front little projected, WL1-3
of similar shape, WL4 slender and its dactyl
longer than the others, MXP3 with a carpus
robust, larger than the propodus, and ab-
domen in both sexes of six somites and tel-
son well separated. These features clearly
distinguish this genus from others currently
in the Pinnotheridae (Campos 1990). The
species of Calyptraeotheres are obligatory
symbionts of slipper shells (family Calyp-
traeidae). No other member of the Pinno-
theridae is known to be symbiotic with slip-
per shells (Geiger & Martin 1999).

Marques & Pohle (1995) included of C.
politus within the genus Tumidotheres
Campos, 1989 based on presumably shared
characters between C. politus and Tumi-
dotheres spp; namely the narrowly spatulate
dactylus of MXP3 that is inserted at an an-
gular notch on the middle of the ventral
margin of the propodus. As previously not-
ed (Campos 1989), these characters are di-
agnostics for Tumidotheres (Fig. 2D); how-
ever, MXP3 of C. politus does not exhibit
such characters. Calyptraeotheres politus
and C. garthi, have the dactylus of the
MXP3 rounded, minute and inserted sub-
distally on the ventral margin of the pro-
podus (Fig. 2B—C). The genus Tumidothe-
res can be separated from Calyptraeotheres
by the tumid and setose carapace, and by
the above noted shape and insertion point
of articles of the MXP3. In addition, species
of Tumidotheres inhabit the mantle cavity

538

Fig. tf.
E-E C. garthi (Fenucci). A-B from Campos, 1990; C modified from Retamal, 1981; D from Garth, 1957; E—F
from Fenucci, 19735. Scale in mm; A= 2.5: B= F:3; C = 2.6; D:-—= 1.35 BE = 34. EF = ES

of Bivalvia species, while species in Calyp-
traeotheres live between the shell and the
head of slipper shells (family Calyptraei-
dae). Their different mode of life can be
indicative of divergence and supports the
separation of these genera.

Acknowledgments

I am indebted to Juan L6pez Gappa and
Alejandro Tablado for the loan of C. garthi
deposited in the Museo Argentino de Cien-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Adult female and male respectively, A—B, Calyptraeotheres granti Glassell; C—D, C. politus (Smith);

cias Naturales ‘“‘Bernardino Rivadabia’’; to
my wife Alma-Rosa for her fine artistic
work; to Daniel L. Geiger, R. Lemaitre, G.
Pohle and E D’Incao for reviewing this ar-
ticle with great care and supplying pertinent
literature; and R. Lemaitre and R. B. Man-
ning for their continuous support of my
Pinnotherid crab studies. This work was
partially financed by the project UABC-
0134 “Crustaceos simbiontes de Baja Cal-
ifornia’’.

VOLUME 112, NUMBER 3 539

D

Fig. 2. Third maxilliped. A, Calyptraeotheres granti (Glassell); B, C. garthi; C, C. politus (Smith); D,
Tumidotheres margarita (Smith). Scale in mm, A = 0.95; B = 0.43: C = 0.39; D = 0.70.

540

Literature Cited

Campos, E. 1989. Tumidotheres, a new genus for Pin-
notheres margarita Smith, 1869 and Pinnothe-
res maculatus Say, 1818 (Brachyura, Pinnothe-
ridae).—Journal of Crustacean Biology 9:672—
679.

. 1990. Calyptraeotheres, a new genus of Pin-
notheridae for the limpet crab Fabia granti
Glassell, 1933 (Crustacea, Brachyura).—Pro-
ceedings of the Biological Society of Washing-
ton 103:364—371.

Fenucci, J. L. 1971. Notas sobre las dos especies de
pinotéridos mas comtunes del litoral bonaerense
(Decapoda, Brachyura, Pinnotheridae).—Phy-
sis, serie A 30(81):355—367.

. 1975. Los cangrejos de la familia Pinnother-
idae del litoral argentino (Crustacea, Decapoda,
Brachyura).—Physis, seccion A 34(88):165—
184.

Gallardo, C. S. 1979. Especies gemelas del género
Crepidula en la costa de Chile; una redescrip-
ci6n de C. dilatata Lamark y descripcién de C.
fecunda n. sp.—Studies on Neotropical Fauna
and Environment 14(4):216—227.

Garth, J. S. 1957. The Crustacea Decapoda Brachyura
of Chile—Reports of the Lund University
Chile Expedition, 1948—49 29:1—127.

Geiger, D. C., & J. W. Martin. 1999. The pea crab
Orthotheres haliothidis new species (Decapoda:
Brachyura: Pinnotheridae) in the australian ab-
alone Haliotis asinina Linnaeus, 1758, and Hal-
iotis squamata Reeve, 1846 (Gastropoda: Veti-
gastropoda: Haliotidae).—Bulletin of Marine
Science 64:269-—280.

Glassell, S. A. 1933. Description of five new species
of Brachyura collected on the west Coast of
Mexico.—Transactions of the San Diego Soci-
ety of Natural History 7(28):331—334.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

de Haan, W. 1833-1850. Crustacea. In P. E von Sie-
bold, Fauna Japonica sive Descriptio Animal-
ium, quae in Itinere per Japoniam, Jussu et Aus-
piciis Superiorum, qui Summum in India Bata-
va Imperium Tenent. Suscepto, Annis 1823—
1830 Collegit, Notis, Observationibus et
Adumbrationibus I[lustravit. i-xvii, i-xxxi, ix-
xvi, 1-243, plates A-J, L-Q 1-55, circ. tab. 2
Lugduni-Batavorum [Leiden].

Hoagland, K. E. 1983. Notes on type specimens of
Crepidula (Posobranchia: Calyptraeidae) in the
Muséum National d’ Histoire Naturelle, Paris.—
Proceedings of the Academy of Natural Scienc-
es of Philadelphia 135:1-8.

Marques, F, & G. Pohle. 1995. Phylogenetic analysis
of the Pinnotheridae (Crustacea, Brachyura)
based on larval morpholgy, with emphasis on
the Dissodactylus complex.—Zoologica Scripta
24:347-364.

Martins, S. T. S., & E D’Incao. 1996. Os Pinnotheridae
de Santa Catarina e Rio Grande do Sul, Brasil
(Decapoda, Brachyura).—Revista Brasileira de
Zoologia 13:1—26.

Retamal, M. A. 1981. Catalogo ilustrado de los crus-
taceos decapodos de Chile.—Gayana, Zoologia
(Universidad de Cancepcién) 44:1—110.

Saelzer, H. E., & Hapette, A. M. 1986. Desarrollo lar-
vario de Pinnotheres politus (Smith, 1870)
(Brachyura: Pinnotheridae) en condiciones de
laboratorio.—Gayana, Zoologia 50:63—70.

Schmitt, W. L., J. C. McCain, & E. S. Davidson. 1973.
Decapoda I, Brachyura I. Family Pinnotheridae.
In H.-E. Gruner & L. B. Holthuis, eds., Crus-
taceorum Catalogus 3, W. Junk, Den Haag 1—
160.

Smith, S. I. 1869. Pinnotheres margarita Smith, sp.
nov. Jn A. E. Verrill. On the parasitic habits of
Crustacea. American Naturalist 3(5):245.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(3):541-552. 1999.

Description of Synalpheus williamsi, a new species of sponge-dwelling
shrimp (Crustacea: Decapoda: Alpheidae), with remarks on its first

larval stage

Rubén Rios and J. Emmett Duffy

School of Marine Science and Virginia Institute of Marine Science,
The College of William and Mary, Gloucester Point, Virginia 23062, U.S.A.

Abstract.—A new species of Synalpheus is described based on specimens
collected from sponges in Belize and the Atlantic coast of Panama. The new
species, S. williamsi, is most similar morphologically to S. goodei Coutiére,
but the two species are consistently different in several morphological and
larval features, and occupy distinct species of sponges. The shape of the major
chela and of the uropodal exopods are the most reliable morphological char-
acters that distinguish the two species. The first larval stage, a zoea I, was
obtained from an ovigerous female of the new species. The zoea I is similar
to that of S. neomeris (De Man), S. triunguiculatus (Paulson), S. tumidomanus
(Paulson), and S. scaphoceris Coutiére, in lacking pleopods and chelae, but
can be distinguished by the presence of an acute projection on the pterygos-
tomian corner.

Resumen.—Se describe una nueva especie de Synalpheus en base a especi-
menes recolectados dentro de esponjas en Belice y la costa Atlantica de Pan-
ama. La nueva especie, Synalpheus williamsi, es muy parecida a S. goodei
Coutiére, pero posee distintas caracteristicas ecologicas, morfoldgicas y lar-
varias. Estas dos especies de Synalpheus habitan distintas especies de esponjas.
Los caracteres morfol6gicos mas confiables para distinguirlas son la forma de
la quela mayor y de los ex6podos uropodales. De una hembra ovigera se obtuvo
la primera fase larvaria, una zoea I desprovista de pleépodos y de quelas. La
zoea I de la nueva especie es muy similar a la de S. neomeris (De Man), S.
triunguiculatus (Paulson), S. tumidomanus (Paulson), y S. scaphoceris Couti-
ére, pero se distingue de ellas por la presencia de una proyeccion pterigosto-
miana aguda en el caparazon.

During the course of collections made
Over several years in Belize and on the At-
lantic coast of Panam4, we obtained a num-
ber of shrimps of a sponge-dwelling Syn-
alpheus species morphologically similar to
S. goodei Coutiére, 1909. Specimens were
collected from living sponges taken by
SCUBA from various depths (1-15 m) on
the outer reef ridge at the Smithsonian In-
stitution’s field station on Carrie Bow Cay,
Belize (16°48'N, 88°05’W), and from Ulag-
sukun, Pico Feo, and Mamitupo reefs near
the Smithsonian Tropical Research Insti-

tute’s (STRI) field station in the San Blas
Islands, on the Atlantic coast of Panama
(9°34'N, 78°58'W). Specimens from both
areas were collected from the internal ca-
nals of the midnight-blue sponge Hymen-
iacidon caerulea Pulitzer-Finali, 1986, al-
though one was associated with an uniden-
tified orange encrusting sponge of tubular
shape. Four specimens were collected free
from any host, and had probably left their
host sponges during sample handling. Usu-
ally, a heterosexual pair or a single adult
was found in a same sponge. Here we de-

542

scribe these specimens. We also include a
diagnosis of the first zoeal stage of this spe-
cies hatched in the laboratory. Material is
deposited in the National Museum of Nat-
ural History, Smithsonian Institution,
Washington, D.C. (USNM); Coleccién Na-
cional de Crustaceos from the Instituto de
Biologia UNAM, México (CNCR), the Mu-
séum National d’Histoire Naturelle, Paris
(MNHN), and in the Virginia Institute of
Marine Science (VIMS). Measurements in-
dicated are of carapace length including
rostrum.

Synalpheus williamsi, new species
Figs. 1-6

Type specimens.—Holotype 6, 3.6 mm
(USNM 276158), allotype ovigerous ¢, 4.5
mm (USNM-276159), Carrie Bow Cay, Be-
lize, 13 Jun 1996, from canals of same
specimen of midnight-blue sponge Hymen-
iacidon caerulea, 18 m.—Paratypes: 6, 4.4
mm, 2 ovigerous 2 ¢, 4.3, 4.8 mm (USNM-
276160), Carrie Bow Cay, Belize, 4 Apr
1993, from canals of same specimen of
midnight-blue sponge H. caerulea, 13 m;
Paratype 3, 2.3 mm (CNCR 17987) Carrie
Bow Cay, Belize, 12 Jun 1996, from canals
of midnight-blue sponge H. caerulea, 18 m;
Paratype 6, 3.7 mm (MNHN-Na 13561)
Ulagsukun Reef, Panama, 17 Jan 1991,
from canals of midnight-blue sponge H. ca-
erulea.

Additional specimens examined (non
paratypes ).—Belize: Carrie Bow Cay, 5
Apr 1993, 2 66, 2.9, 3.8 mm (VIMS); 18
Aug 1994, 6, 3.5 mm (VIMS); 26 Aug
1994, 3, 4.7 mm (VIMS), zoea larvae
(USNM-276161); in midnight-blue sponge
H. caerulea. Panama: Ulagsukun Reef, 18
Jan 1991 ovigerous °, 4.2 mm (VIMS); 10
Nov 1992, 3, 3.7 mm (VIMS), in midnight-
blue sponge H. caerulea, 1 m; Pico Feo
Reef, 18 Jan 1991, 6, 3.4 mm, ovigerous
2, 4.2 mm (VIMS), in midnight-blue
sponge H. caerulea; San Blas Islands:
1991, 6, 4.3 mm (VIMS); Mamitupo Reef:

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

19 Jan 1991, 3, 4.1 mm (VIMS), in orange
tubular sponge.

Description of holotype.—Body form
subcylindrical. Carapace smooth, with
sparse short simple setae. Rostrum (Fig. 1a,
d, e) slender, distally upturned, slightly
overreaching orbital teeth, not reaching to
distal margin of first segment of antennular
peduncle; dorsally carinated; ventrally pro-
duced into small longitudinal keel, but
without true orbitorostral process. Ocular
hoods (Fig. la, d, e) dorsally convex, sep-
arated from rostral carina by broad depres-
sions; triangular ocular teeth and rostrum
with few apical setae; adrostral notches
broadly rounded. Pterygostomian corner
(Fig. 1b) produced into bluntly acute angle.
Posterior margin (Fig. 2a, d) with distinct
cardiac notch.

Abdomen with sparse simple setae; pleu-
ron of first somite (Fig. 2d) with anterior
corner almost in right angle, ventral margin
sinuous and posteriorly bearing a blunt
strongly hooked tooth directed anteriorly;
second pleura broadly rounded anteriorly,
with ventral margin slightly concave and
posterior corner rounded, but with widely
obtuse projection behind 3/4 of length; third
to fifth pleura anteriorly and posteriorly ob-
tuse, ventral margin triangular with apex
progressively displaced posteriorly; sixth
abdominal segment (Fig. 2a) with bluntly
acute projection on posteroventral margin,
shallow curved inferior concavity, triangu-
lar lateral lobe with convex sides, shallow
obtuse upper emargination, and convex dor-
sal margin slightly overhanging telson. Tel-
son (Fig. lc) subtriangular, truncate, proxi-
mal margin slightly more than twice as
wide as distal margin; lateral margins
slightly convex just as far as distal pair of
dorsal spines, then slightly concave; dorsal
surface with mesial shallow depression and
2 pairs of dorsal spines, anterior pair situ-
ated slightly anterior to proximal third of
length of telson, posterior pair of dorsal
spines at second third of length of telson;
distal margin arcuate, posterolateral corners
not projected, 2 pairs of flanking distal

VOLUME 112, NUMBER 3

C

Fig. 1.

543

Synalpheus williamsi, new species. Paratype male 4.4 mm (USNM 276160): a, anterior region of

carapace and cephalic appendages, dorsal view; b, same, lateral view; c, telson and uropods, dorsal view.
Paratype ovigerous female 4.3 mm (USNM 276160): d, anterior region of carapace and cephalic appendages,
dorsal view; e, anterior region of carapace, lateroventral view. Scale bar = 1 mm for a, b, d, e and 0.72 mm

for c.

movable spines, lateral pair shorter than in-
nermost; mesial gap wider than combined
bases of both pairs of spines, with 2 tufts
of 3 erect setae on dorsal surface mesially
adjacent to inner spines, and single row of
8 more conspicuous, posteriorly directed

plumose setae of similar size, emerging
from under mesial distal lobe.

Stylocerite (Fig. la, d) slender, with me-
sial side slightly concave, barely overreach-
ing distal margin of basal article of anten-
nular peduncle; this latter with lateral fan of

544

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VOLUME 112, NUMBER 3

setae proximally on dorsal face; second and
third segments of antennular peduncle de-
creasing in length. Upper flagellum (Fig.
la, b) biramous, rami fused for 5 segments;
lower ramus shorter, with 3 groups of aes-
thetascs on ventral face, and 4 and 5 addi-
tional groups posteriorly, upper ramus with
12 and 15 segments (left and right side, re-
spectively). Lower flagellum normal, longer
than upper.

Basicerite (Fig. la, b, d) with acute dor-
solateral projection, and longer acute ven-
trolateral spine overreaching tip of stylo-
cerite; scaphocerite with slightly reduced
blade reaching to distal edge of antennular
peduncle, narrower at base than _ lateral
spine, which is robust, acute, with slightly
concave lateral margin, and clearly over-
reaching antennular peduncle. Carpocerite
(Fig. 1b, d) about 7 times as long as wide
when viewed laterally, overreaching tip of
lateral spine of scaphocerite.

Exopod of third maxilliped (Fig. 3f) not
overreaching antepenultimate article, tip of
ultimate segment (Fig. 3g) with circle of 7
strong blunt spines finely denticulate on
proximal half of inner face. Remaining
mouthparts as figured from paratypes (Fig.
3 a-e).

Major first pereiopod (Fig. 4a, b, c) mas-
sive, subcylindrical, with few sparse setae;
palm about 1.75 times longer than wide,
dorsodistal margin with blunt prominence
overhanging insertion of dactyl, and bear-
ing subterminal, ventrally directed, acute
spine. Dactyl 0.3 times as long as palm,
strongly compressed, dorsal margin broadly
semicircular in lateral view (Fig. 4b), cut-
ting edge almost flat with discrete blunt tip,
internal face opposing outer face of pollex
when closed. Pollex 0.7 times as long as
dactyl, deeply excavated proximally to re-

a

Fig. 2.

545

ceive dactylar molar process, in ventral
view (Fig. 4g), thick, triangular, with ac-
cessory subdistal obtuse projection on outer
face, flanking outer face of dactyl when
closed. Carpus cup-shaped, short and broad.
Merus, about 0.8 times as long as height of
palm, nearly triangular in cross section, in-
ner edges slightly concave, outer convex,
lower side flattened, with lamellar triangu-
lar projection on distal outer corner, upper
side clearly convex.

Minor first pereiopod (Fig. 4d) slightly
compressed. Palm about 1.7 times longer
than high. Dactyl 0.7 times as long as palm,
with several tufts of regular setae; extensor
margin convex, with 2 longitudinal series
of transverse parallel rows of setae distally
curved towards tip of dactyl, inner series
composed of about 12 rows, first row at
start of second third of dactyl, last row at
end of fourth fifth, shorter outer series, with
first row opposing fifth row of inner series,
setae similar in shape, length, and orienta-
tion, but slightly thinner; dactyl flexor mar-
gin excavated (Fig. 4e), tip with strong
tooth flanked by two accessory blunt pro-
jections. Pollex with sinuous lower margin,
sparse tufts of setae, strong apical tooth
(Fig. 4f) continued backward into oblique
widely convex blade. Carpus cup-shaped,
about 0.5 times as long as palm; 1.2 times
higher distally than mesial length in lateral
view; upper and lower margins distally ex-
panded over proximal portions of palm.
Merus 1.5 times as long as palm, 2.5 times
longer than maximum width; almost trian-
gular in cross section; distal outer corner
flared, without true spine.

Second pereiopod (Fig. 4h) more slender
than all others. Fingers with strong tufts of
setae, slightly more than 1.3 times as long
as palm, latter 1.4 times longer than high.

Synalpheus williamsi, new species. Paratype ovigerous female 4.8 mm (USNM 276160): a, abdomen

with one egg, lateral view; b, first pleopod; c, second pleopod. Paratype male 4.4 mm (USNM 276160): d, first
5 segments of abdomen, lateral view; e, first pleopod; f, same, detail of endopod; g, second pleopod. Scale bar
= 2.2 mm for a, d, 1 mm for b, c, e, g, and 0.31 mm for f.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

546

‘\
)

mt ing

NN

\

VOLUME 112, NUMBER 3

Carpus composed of 5 articles with ratio
4.5:1:1:1:2, second through fourth equally
as long as high. Merus about 5.3 times lon-
ger than wide, and 0.9 times as long as car-
pus. Ischium four times longer than wide,
and (.7 times as long as merus.

Third pereiopod (Fig. 5a), strongest of
posterior legs. Dactyl (Fig. 5d) biunguicu-
late, slightly longer than width of propodus,
flexor margin concave. Propodus 6.3 times
longer than wide, flexor margin with a lon-
gitudinal series of 7 strong movable spines
and one terminal pair of movable spines
flanking base of dactyl. Carpus 2.6 times
longer than wide, extensor distal margin
projected over propodus, strong movable
spine on distal flexor margin. Merus 4.4
times longer than wide, unarmed. Ischium
unarmed.

Fourth pereiopod (Fig. 5b, e) very simi-
lar to third, but slightly weaker.

Fifth pereiopod (Fig. 5c, f) weaker than
fourth, with following slight differences,
propodus with only two or three spines on
flexor margin besides distal pair, and about
five combs of stout setae transversely ar-
ranged on distal half of posterior face; car-
pus without spine.

Endopod of first pleopod (Fig. 2e) not as
long as width of exopod, with few apical

—

Fig. 3.

547

setae (Fig. 2f). Second pleopod (Fig. 2g)
without appendix masculina. Pleopods 2—5
with appendix interna.

Uropodal exopod (Fig. 1c) with 5 strong
distolateral teeth, longer movable spine and
acute mesial tooth. Diaeresis present, dis-
tinct.

Color.—Live specimens were translucent
faint gold to golden brown, with tip of che-
lae (fingers and distal part of palm includ-
ed) of first pereiopods bright orange to red,
second legs golden, third to fifth transpar-
ent, and traces of blue in the gut and in the
branchiae. These traces were the same dark
blue as the host sponge, and they still can
be seen in some of the preserved speci-
mens.

Variations.—The few discrepancies be-
tween the illustrations, from a male para-
type, and the holotype are most probably
due to the larger size of the figured para-
type. The greater development of the en-
dopod on the first pleopod (Fig. 2f) could
be related to different stages of sexual ma-
turity. Regarding sexual dimorphism, the
most striking female characters are the
broadly rounded shape of the first to fourth
pleurae (Fig. 2a), the larger endopod of the
first pleopod (Fig. 2b), and the insertion of

Synalpheus williamsi, new species. Mouthparts. Paratype male 4.4 mm (USNM 276160): a, mandible;

b, first maxilla; c, second maxilla; d, first maxilliped; e, second maxilliped. Paratype ovigerous female 4.3 mm
(USNM 276160): f, third maxilliped; g, same, detail of tip. Scale bar = 0.5 mm for a, b, c, d, e, 1 mm for f,

and 0.25 mm for g.

a

Fig. 4. Synalpheus williamsi, new species. Paratype male 4.4 mm (USNM 276160): a, major first pereiopod
in lateral view; c, chela of same, dorsal view; g, same, tip of pollex, ventral view; h, second pereiopod. Paratype
ovigerous female 4.3 mm (USNM 276160): b, anterior portion of major first pereiopod, external face, lateral
view; d, minor first pereiopod, lateral view; e, distal portion of same, lateroventral view; f, same, detail of tip,
frontolateral view. Scale bar = 2.5 mm for a, 2.2 mm for b, c, 1 mm for d, 1.4 mm for e, g, 0.2 mm for f, and
2 mm for h.

Fig. 5. Synalpheus williamsi, new species. Paratype male 4.4 mm (USNM 276160): a, third pereiopod; b,
fourth pereiopod; c, fifth pereiopod; d, detail of distal portion of third pereiopod; e, detail of distal portion of
fourth pereiopod; f, detail of distal portion of fifth pereiopod. Scale bar = 1 mm for a, b, c, and 0.31 mm for
ce, ft.

548 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

VOLUME 112, NUMBER 3

550

the appendix interna beyond the midpoint
in the remaining pleopods (Fig. 2c).

On the tip of third maxilliped, eight or
seven spines is the most frequent formula
(as in the holotype), but the allotype has
nine spines on one side and ten on the other.
The number of immovable teeth on the lat-
eral margin of the uropodal exopod varies
from three to six, with a mode of four; this
variation apparently relates to size and per-
haps sex, considering that the smaller spec-
imens have fewer teeth, and that the only
two specimens with six teeth were males.

Eggs and larva.—Eggs measure 0.9-—1.0
mm long, and about 0.6 mm wide. One
ovigerous female released larvae in the lab-
oratory. Seven larvae were recovered
swimming freely, but no attempts were
made to raise them beyond the first stage.
They were both preserved and stored in eth-
anol along with two eggs and another larva
with half its body still in the egg case.

Description of zoea I.—Carapace sub-
cylindrical (Fig. 6a), pterygostomian corner
projected into acute spine, low bump on
middorsal line, and angular projection prox-
imal to depression at base of rostrum; latter,
broadly triangular, pointed, directed down-
wards, shorter than, and somewhat hidden
by ocular peduncles; eyes not covered by
carapace.

Antennular peduncle with 3 segments.
Antennal scale with 2 outer plumose setae,
outer apical spine, and 9 plumose setae on
inner and distal margins; endopod reaching
to first inner seta on scale, entire, with one
strong and one small apical seta. Maxillae
and mandible buds present. All maxillipeds
with well developed endopods and strong
exopods, but setae present only on exopods.

First to fourth pairs of pereiopods bira-
mous, without apparent segmentation, exo-
pods without setae. Third and fourth pe-
reiopods rudimentary, folded anteriorly
against thorax. Fifth pair, long, turned for-
wards, uniramous, without setae.

Abdomen with sixth segment not clearly
separated from telson. Pleopods absent.
Uropods fused with telson. Telson (Fig. 6b),

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

broad, bilobate, with 7 + 7 setae, outer 2
pairs feathered on inner side only, remain-
ing 5 pairs plumose on both sides, inner-
most pair less than 0.25 as long as adjacent.

Ecology.—Most specimens of the new
species, S. williamsi, came from internal ca-
nals of the midnight-blue sponge Hymen-
iacidon caerulea. In a few cases (<25%)
we were not able to record the exact origin
of our specimens, since they were found
among debris in sampling containers. Only
once did we collect a specimen (male) of
S. williamsi, new species, from an uniden-
tified orange encrusting sponge of tubular
shape that was clearly different from H. ca-
erulea.

Traces of blue in the gut of some speci-
mens, and the anecdotal record in our field
notes of a voided fecal pellet containing
sponge spicules, are suggestive of a para-
sitic relation between the shrimp and the
sponge.

Etymology.—It is an honor and pleasure
to name this species after Dr. Austin B. Wil-
liams, National Oceanic and Atmospheric
Administration, National Marine Fisheries
Service Systematics Laboratory, National
Museum of Natural History, Smithsonian
Institution, Washington, D.C. This is in ap-
preciation for his many important contri-
butions to the knowledge of decapod crus-
taceans, and in recognition of his personal
generosity and integrity.

Remarks.—Synalpheus williamsi, new
species, is morphologically similar to S.
goodei Coutiére, but the following charac-
teristics of the latter serve to distinguish the
two species: the blade on the scaphocerite
is more reduced, the pollex on the large
chela is longer and without an accessory
lateral emargination (best seen in ventral
view), the small chela is more elongate, the
distal margin of the telson is narrower, the
lateral margin of the uropodal exopod has
more fixed spines and has a noticeably
stronger inner spine, adjacent to the mov-
able one (see Coutiére 1909, and Dardeau
1984). Also, unlike the new species, the
first larva of S. goodei hatches at a more

VOLUME 112, NUMBER 3

=e |

Fig. 6. Synalpheus williamsi, new species. Scanning electron micrographs of first zoea stage: a, lateral view;

b, ventral view. Magnification = 71x.

advanced stage with both pairs of chelae
and pleopods present (Gurney 1949).
Among the 12 species of Synalpheus
whose first larva has been described or fig-
ured, the zoea I from the new species, S.
williamsi, is most similar to that of S. neo-
meris (De Man, 1888), S. triunguiculatus
(Paulson, 1875), S. tumidomanus (Paulson,
1875), and S. scaphoceris Coutiére, 1910,
but it can be readily distinguished by the

acute projection of the pterygostomian cor-
ner; additionally, the first zoea of S. sca-
Phoceris has the sixth abdominal somite
clearly distinct (Dardeau 1986: fig. 3b).
Lack of pleopods and chelae on the first
pair of pereiopods is suggestive of a normal
development pattern, i.e., not abbreviated
nor direct as has been documented for other
species of Synalpheus (see Knowlton 1973,
and Bhuti et al. 1977).

352

As in the case of S. gambarelloides (Nar-
do, 1847), and probably S. longicarpus
(Herrick, 1891), recorded by Riietzler
(1976) and Erdman & Blake (1987) respec-
tively, the new species, S. williamsi, ap-
pears to be a parasitic endobiont of spong-
es; more careful observations on live spec-
imens might provide conclusive evidence
on its diet.

Acknowledgments

We thank Klaus Riietzler, Mike Carpen-
ter, and Brian Kensley for facilitating work
at Carrie Bow Cay, and Tripp MacDonald
for assisting in the field work. We are deep-
ly indebted to the people of Belize, and to
the Kuna Nation and the Republic of Pan-
ama for permission to work in San Blas.
This work was supported in part by the Na-
tional Science Foundation (Postdoctoral
Fellowship in Environmental Biology, and
DEB 9815785) to JED, and by the Smith-
sonian Institution’s Caribbean Coral Reef
Ecosystem Program, from which this is
Contribution #565. RR is a Fulbright-
Garcia Robles scholar at VIMS. Patrice
Mason at VIMS skillfully overcame the
poor preservation qualities of our larvae,
and produced the SEM photographs. This
is VIMS contribution #2210.

Literature Cited

Bhuti, G. S., S. Shenoy, & K. N. Sankolli. 1977. Lab-
oratory reared alpheid larvae of the genera Au-
tomate, Athanas and Synalpheus (Crustacea De-
capoda, Alpheidae).—Proceedings of the Sym-
posium on Warm Water Zooplankton. Special
Publication, National Institute of Oceanogra-
phy, Goa (India), Pp. 588—600.

Coutiére, H. 1909. The American species of snapping
shrimps of the genus Synalpheus.—Proceedings
of the United States National Museum 36:1-—93.

. 1910. The snapping shrimps (Alpheidae) of

the Dry Tortugas, Florida.—Proceedings of the

United States National Museum 37:485—487.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Dardeau, M. R. 1984. Synalpheus shrimps (Crustacea:
Decapoda: Alpheidae). I. The Gambarelloides
Group, with a description of a new species.—
Memoirs of the Hourglass Cruises 7(2):1—125.

. 1986. Redescription of Synalpheus scapho-
ceris Coutiere, 1910 (Decapoda: Alpheidae)
with new records from the Gulf of Mexico.—
Journal of Crustacean Biology 6:491—496.

De Man, J. G. 1888. On the podophtalmous Crustacea
of the Mergui Archipelago.—Journal of the
Linnean Society, Zoology 22:241-312.

Erdman, R. B., & N. J. Blake. 1987. Population dy-
namics of the sponge-dwelling alpheid Synal-
pheus longicarpus, with observations on S.
brooksi and S. pectiniger, in shallow-water as-
semblages of the eastern Gulf of Mexico.—
Journal of Crustacean Biology 7:328—337.

Gurney, R. 1949. The larval stages of the snapping-
shrimp, Synalpheus goodei Coutiére.—Proceed-
ings of the Zoological Society of London 119:
293-295.

Herrick, EK H. 1891. Alpheus: A study in the devel-
opment of Crustaceaw—Memoires of the Na-
tional Academy of Sciences, Washington 5:
370-463.

Knowlton, R. E. 1973. Larval development of the
snapping shrimp Alpheus heterochaelis Say,
reared in the laboratory.—Journal of Natural
History 7:273—306.

Nardo, G. D. 1847. Sinonimia moderna delle specie
registrate nell’opera intitolata: Descrizione
de’Crostacei, de’ Testacei e de’ Pesci che abitano
le lagune e golfo Veneto, rappresentati in figure,
a chiaro-scuro ed a colori dall’ Abate Stefano
Chiereghini Ven. Clodiense. i-1x + 64 pp. in
128 columns.

Paulson, O. 1875. Podophtalmata i Edriophtalmata
(Cumacea). Chast I in Izsledovaniya Rakoob-
raznykh krasnago morya s zametkami otnosi-
tel’no Rakoobraznykh drugikh morei. Kiev, 144
pp. [English translation: Por, E D. 1961 Po-
dophtalmata and Edriophtalmata (Cumacea).
Part I in Studies on Crustacea of the Red Sea
with notes regarding other seas. Jerusalem, Is-
rael. 164 pp.]

Pulitzer-Finali, G. 1986. A collection of West Indian
Demospongiae (Porifera). In appendix, a list of
Demospongiae hitherto recorded from the West
Indies.—Annali del Museo Civico di Storia Na-
turale ““Giacomo Doria’’, Genova 86:65—216.

Rtietzler, K. 1976. Ecology of Tunisian commercial
sponges.—Tethys 7:249—264.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(3):553-561. 1999.

Two new freshwater crabs of the genus Ptychophallus Smalley, 1964
(Crustacea: Decapoda: Brachyura: Pseudothelphusidae) from
Panama, with notes on the distribution of the genus

Martha R. Campos and Rafael Lemaitre

(MRC) Universidad Nacional de Colombia, Instituto de Ciencias Naturales,
Apartado Aéreo 53416, 114 Santa Fé de Bogota 2, Colombia, S. A.;
(RL) Department of Invertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560-0163, U.S.A.

Abstract.—Two new species of freshwater crabs of the genus Ptychophallus
Smalley, from Panama, P. uncinatus and P. kuna, are described and illustrated.
The addition of these two species means the genus now includes 13 species,
all of which are distributed in Panama and Costa Rica. The species are distin-
guished primarily by differences in characters of the male first gonopod. A
summary of the geographic distribution of the species in the genus is presented.

The systematics of the genus Ptycho-
phallus Smalley, 1964a, were reviewed by
Rodriguez (1982, 1994) who found that a
number of characters of the male first gon-
opod in species of this genus were homol-
ogous with those of Hypolobocera Ort-
mann, 1897 and Neostrengeria Pretzmann,
1965. These characters are the presence of
a caudal crest, a lateral lobe, and a central
papilla on the spermatic channel. Rodriguez
(1982) considered Ptychophallus to be tran-
sitional between Andean and Central Amer-
ican Pseudothelphusidae because species of
this genus present a fusion of the cephalic
and caudal borders forming a mesial pro-
cess, a morphological characteristic present
in all pseudothelphusids species from Cen-
tral America (including Mexico), but absent
in species of Hypolobocera and Neostren-
geria from South America.

Species of Ptychophallus are distin-
guished primarily by characteristics of the
third maxilliped and male first gonopod.
The exognath of the third maxilliped is 0.6
to 0.7 as long as the ischium. The male first
gonopod has a prominent apex that is bent
cephalically and is joined to the gonopod
by a narrow peduncle; the lateral projection
is large, and usually divided into two lobes;

and the caudal ridge extends longitudinally
on the distal half of the caudal surface.

A study of freshwater crabs in the Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington D.C.
(USNM), revealed two new species of
Ptychophallus from Panama, bringing to 13
the total number of species currently rec-
ognized in this genus. This number ex-
cludes P. campylos Pretzmann, 1965, a spe-
cies that was originally briefly described,
without illustrations (Pretzmann 1965).
This species was listed by Pretzmann
(1971), and he (Pretzmann 1972) subse-
quently added details with illustrations and
photographs. Rodriguez (1982:80) indicat-
ed in a footnote that based on the infor-
mation provided by Pretzmann (1965,
1971, 1972), P. campylos could not be sep-
arated from P. tristani (Rathbun, 1896);
however, Rodriguez did not formally syn-
onymize Pretzmann’s taxon with Rathbun’s
and excluded P. campylos from his treat-
ment of Ptychophallus species. The type of
P. campylos, deposited in the Naturhisto-
risches Museum, Vienna, needs to be ex-
amined in order to properly evaluate the va-
lidity of Pretzmann’s taxon.

The terminology used for the morpholo-

554

gy of the male first gonopod follows that of
Smalley (1964b) and Rodriguez (1982).
The abbreviations cb and cl stand for car-
apace breadth (measured at the widest
point) and carapace length (measured along
the midline), respectively. Color nomencla-
ture used follows Smithe (1975).

Family Pseudothelphusidae Rathbun, 1893
Tribe Hypolobocerini Pretzmann, 1971
Genus Ptychophallus Smalley, 1964a
Ptychophallus uncinatus, new species
Pigs. fy 2rd

Material.—Holotype: ¢, cl 16.4 mm, cb
26.5 mm, Rio San Pedro, Bocas del Toro,
Panama,, Jan,.1978,, lee... B. <l...Gordon
(USNM 276164). Paratype: 3 (juvenile) cl
9.5 mm, cb 15.2 mm, same locality as ho-
lotype (USNM 276165).

Type locality.—Rio San Pedro, Bocas del
Toro, Panama.

Diagnosis.—First male gonopod with
very broad lateral projection divided into 2
subequal lobes by deep depression on cau-
dal surface and lateral notch; distal lobe
with crest-like flange; distocaudal ridge
short and narrow, with lateral crest; sub-
apical mesial process with triangular base
and hook-like projection; central papilla of
spermatic channel robust, swollen.

Description of holotype.—Carapace (Fig.
1) with narrow, deep cervical groove curv-
ing posteriorly and not reaching lateral mar-
gin. Anterolateral margin with shallow si-
nus just posterior to anteroexternal orbital
angle; rest of margin with papillae. Post-
frontal lobes wide, delimited anteriorly by
transverse depression; median groove nar-
row, deep, with incision on upper margin
of front. Surface of carapace in front of
postfrontal lobes flat and inclined anterior-
ly. Upper border of front thin, marked with
row of tubercles; lower margin sinuous in
frontal view. Surface of front between up-
per and lower borders high and slightly ex-
cavated. Upper and lower orbital margins
each with row of tubercles. Surface of car-
apace covered with small papillae; limits

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

between regions demarcated. Third maxil-
liped (Fig. 2E) with merus of endognath
regularly curved, with shallow depression
on distal part of external margin; exognath
approximately 0.6 times length of ischium
of third maxilliped.

First pereiopods absent in holotype. In
paratype, right cheliped slightly larger than
left. Merus with 3 crests as follows: upper
crest with rows of tubercles, internal lower
crest with rows of teeth, and external lower
crest with few tubercles. Carpus with 4 tu-
bercles on internal crest and prominent
blunt spine distally. Palm of both chelipeds
smooth, moderately swollen. Fingers of
chelae not gaping when closed, tips cross-
ing, with rows of tubercles on dorsal side.

Surface of walking legs (pereiopods 2-5;
Fig. 1) with rows of minute setae. Dactyls
each with 5 rows of large spines slightly
diminishing in size proximally, rows ar-
ranged as follows: anterolateral and antero-
ventral rows each with 5 spines, external
row with 5 spines plus 2 proximal papillae,
posteroventral row with 4 spines, and pos-
terolateral row with 5 spines.

First male gonopod (Fig. 2A—C) wide in
caudal view. Lateral projection very wide,
divided in 2 subequal lobes by deep de-
pression on caudal surface and lateral
notch; distal lobe with crest-like flange di-
rected distally; proximal lobe almost round-
ed; margins with minute setae; distocaudal
ridge short and narrow, with lateral crest,
reaching only to middle of deep depression
dividing lateral projection. Subapical mesial
process with triangular base and hook-like
projection; apical mesial process triangular
(Fig. 2C, D). Apex bent cephalolaterally,
with field of spines directed cephalolater-
ally; caudal border of apex with shallow
notch near middle, and deep notch near lat-
eral side; central papillae of spermatic chan-
nel robust, swollen (Fig. 2B, C, D).

Color.—In alcohol, the dorsal side of the
carapace is light brown (near 239, Ground
Cinnamon). The walking legs are Buff
(124). The chelae are True Cinnamon (139)

VOLUME 112, NUMBER 3

Pig. .
ZIGLOS).

dorsally, and Buff (124) ventrally. The ven-
tral surface of the carapace is Buff (124).
Etymology.—The specific name is from
the Latin uwncinatus, meaning hooked, and
is given in reference to the form of the sub-
apical mesial process of the first gonopod.
Remarks.—This new species is most
similar to Ptychophallus tumimanus (Rath-
bun, 1898). The two can be distinguished
by differences in the male first gonopod. In
the new species the distal lobe of the lateral
projection is a crest-like flange directed dis-
tally, and the lateral border of the proximal
lobe has minute setae (not shown in Fig.
2A-C); in P. tumimanus the distal lobe is
rounded, and the lateral border of the prox-
imal lobe has rows of long setae. In the new
species the distocaudal ridge has a lateral
crest, and the subapical mesial process has
a triangular base and hook-like projection;
in P. tumimanus the distocaudal ridge has
a rounded surface, the subapical mesial pro-
cess is subtriangular, almost rounded.
Ptychophallus uncinatus, new species, is

JICPinzey a /

Ptychophallus uncinatus, new species, male paratype, cl 9.5 mm, cb 15.2 mm, dorsal view (USNM

also similar to P. montanus (Rathbun,
1898). The male first gonopod of the new
species differs markedly from that of P.
montanus in the shape of the apex. In the
new species the apex is wide, the caudal
border has a notch near the middle, and the
central papilla of the spermatic channel is
robust and swollen (Fig. 2D); in P. mon-
tanus the apex is narrow, the caudal border
is entire and lacks a median notch, and the
central papilla of the spermatic channel is
thin and curved. In addition, the male first
gonopod of the new species has lateral pro-
jections that extend laterally beyond the
distal end of the laterocaudal expansion.
(Fig. 2A), whereas the lateral projections do
not exceed the laterocaudal expansion in P.
montanus.

Ptychophallus kuna, new species
Figs. 3—5
Material.—Holotype: 6, cl 15.6 mm, cb
24.9 mm, Portobelo, Panama, 25 Feb 1973:
(USNM 184340).

556 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ABC

mi

E Kner

Fig. 2. Ptychophallus uncinatus, new species, male holotype, cl 16.4 mm, cb 26.5 mm (USNM 276164). A,
left first gonopod, caudal view; B, same, lateral view; C, same, cephalic view; D, apex of same, distal view; E,
left third maxilliped, external view. 1, subapical mesial process; 2, apical mesial process; 3, distocaudal ridge;
4, distal lobe of lateral projection; 5, lateral projection; 6, proximal lobe of lateral projection; 7, lateral border
of proximal lobe; 8, central papilla of spermatic channel; 9, caudal border of apex; 10, notch of caudal border.

VOLUME 112, NUMBER 3

Diagnosis.—First male gonopod with
lateral projection narrow, divided into 2
subequal lobes by shallow notch; distal lobe
small, subtriangular and with distal surface
close to gonopod axis; proximal lobe sub-
triangular; distocaudal ridge short, weakly
defined, almost continuous with distal lobe
of lateral projection; subapical mesial pro-
cess robust, flange-like; central papilla of
spermatic channel thin, delicate.

Description of holotype.—Carapace (Fig.
3A) with narrow, shallow cervical groove
curved posteriorly and not reaching lateral
margin. Anterolateral margin with shallow
sinus just posterior to anteroexternal orbital
angle; remaining margin with papillae.
Postfrontal lobes wide, delimited anteriorly
by transverse depression; median groove
narrow. Surface of carapace in front of
postfrontal lobes flat, inclined anteriorly.
Upper border of front thin, marked with
row of tubercles; lower margin visible in
dorsal view, sinuous in frontal view. Sur-
face of front between upper and lower bor-
ders high, slightly excavated. Upper and
lower orbital margins each with row of con-
spicuous tubercles. Surface of carapace
covered with small papillae; limits between
regions well demarcated. Third maxilliped
(Fig. 3D) with merus of endognath strongly
curved and with deep depression on distal
part of external margin; exognath approxi-
mately 0.6 times length of ischium of third
maxilliped.

First pereiopods heterochelous; right che-
liped larger than left. Merus with 3 crests
as follows: upper crest with rows of tuber-
cles, internal lower crest with rows of teeth,
and external lower crest with rows of tu-
bercles. Carpus with 4 tubercles on internal
crest, and prominent blunt spine distally.
Palm smooth, swollen. Fingers of chelae
(Fig. 3B) not gaping when closed, tips
crossing and with rows of tubercles on dor-
sal surface.

Walking legs (pereiopods 2—5; Fig. 3C)
strong. Dactyls each with 5 rows of large
spines diminishing in size proximally, rows
arranged as follows: anterolateral and an-

557

teroventral rows each with 5 spines, exter-
nal row with 5 spines and 2 proximal pa-
pillae, and posteroventral and posterolateral
rows each with 3 spines.

First male gonopod (Fig. 4) narrow in
caudal and cephalic views. Lateral projec-
tion narrow, divided in 2 subequal lobes by
shallow notch; distal lobe small, subtrian-
gular, distal surface close to gonopod axis;
proximal lobe subtriangular, lateral border
sinuous and with proximal papilla; disto-
caudal ridge very short, weakly defined and
almost continuous with distal lobe of lateral
projection. Subapical mesial process robust,
flange-like (Fig. 4C, D); apical mesial pro-
cess subtriangular, almost rounded. Apex
bent cephalolaterally, with field of spines
directed cephalolaterally; caudal border of
apex with deep incision near lateral side;
central papilla of spermatic channel thin,
delicate (Fig. 4B, C, D).

Color.—In alcohol, dorsal surface of the
carapace brown (near Kingfisher Rufous,
240). Walking legs Raw Sienna (136) dor-
sally, and True Cinnamon (139) ventrally.
Chelae Kingfisher Rufous (240) dorsally,
and True Cinnamon (139) with Olive-
Brown specks (28) ventrally. Ventral sur-
face of carapace Robin Rufous (340).

Etymology.—The species is named in
honor of the Kuna Indians, in whose terri-
tory this new species was found.

Remarks.—This new species can be dif-
ferentiated from other congenerics by the
first male gonopod. The gonopod is distinct
in that it has a short, weakly defined caudal
ridge which is almost continuous with the
distal lobe of the lateral projection.

Distribution of Ptychophallus Species

Distribution records of species of Ptycho-
phallus can be found in the original descrip-
tions as well as in various freshwater crab
reports from central America (Pretzmann
1965, 1968, 1971, 1972, 1980; Rodriguez
1982, 1994; Smalley 1964a; Villalobos
1974). The species of this genus are known
exclusively from Central America between

558 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ICFAnren

Fig. 3. Ptychophallus kuna, new species, male holotype, cl 15.6 mm, cb 24.9 mm (USNM 184340). A,
carapace, dorsal view; B, chela of large cheliped, external view; C, left third pereiopod, lateral view; D, third
maxilliped, external view.

VOLUME 112, NUMBER 3

ICPiazon

559

Fig. 4. Ptychophallus kuna, new species, male holotype, cl 15.6 mm, cb 24.9 mm (USNM 184340), left
first gonopod: A, caudal view; B, lateral view; C, cephalic view; D, apex, distal view.

Cana Mount, Darién Province, Panama, and
Monteverde, Puntarenas Province, Costa
Rica (Fig. 5). Six species occur in localities
of the Atlantic drainage: P. cocleensis

Pretzmann, 1965, P. exilipes (Rathbun,
1898), P. lavallensis Pretzmann, 1978, P.
tumimanus, P. kuna, new species, and P.
uncinatus, new species. Six species occur

560 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

hare

COSTA RICA OCEAN

Monteverde ;
* Volccn Pods
* Heredia
SanJosée|. sTurrialba
11 |g Cartago
4
9
|Son Isidro
3 Portobelo
Corg
Cerro Azul :
8 .Pdcora

Boruca
e

PACIFIC
OCEAN

OF
PANAMA

Fig. 5. Distribution of species of Ptychophallus Smalley, 1964: 1, P. cocleensis Pretzmann, 1965; 2, P.
colombianus (Rathbun, 1893); 3, P. costaricensis Villalobos, 1974; 4, P. exilipes (Rathbun, 1898); 5, P. gold-
manni Pretzmann, 1965; 6, P. kuna, new species; 7, P. lavallensis Pretzmann, 1978; 8, P. micracanthus Rod-
riguez, 1994; 9, P. montanus (Rathbun, 1898); 10, P. paraxanthusi Bott, 1968; 11, P. tristani (Rathbun, 1896);

12, P. tumimanus (Rathbun, 1898); 13, P. uncinatus, new species.

in localities of the Pacific drainage: P. col-
ombianus (Rathbun, 1893); P. costaricensis
Villalobos, 1974, P. goldmanni Pretzmann,
1965, P. micracanthus Rodriguez, 1994, P.
paraxanthusi Bott, 1968, and P. tristani.
One species, P. montanus, has been found
in both Atlantic and Pacific drainages.

Species of Pseudothelphusidae are usu-
ally found at altitudes ranging from 100 to
3000 m. However, when they occur in
coastal mountain ranges such as those
found along the Pacific or Atlantic coasts
of Panama and Colombia, some species can
reach sea level. Species of Ptychophallus,
for example, range from 0 to 2000 m above
sea level.

Acknowledgments

This study was made possible, in part, by
a visitor grant awarded to one of us (MRC)
by the Office of Fellowships & Grants,
Smithsonian Institution, Washington, D.C.
The illustrations were prepared by Juan C.
Pinzon.

Literature Cited

Bott, R. 1968. Fluss-Krabben aus dem 6stlichen Mit-
tel-Amerika und von den Grossen Antillen
(Crustacea, | Decapoda).—Senckenbergiana
Biologica 49(1):39—49.

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

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

. 1968. Neue Siidamerikanisch Siisswasserkrab-

ben der Gattung Pseudothelphusa.—Entomolo-

gisches Nachrichtenblatt, Wien 15(1):1—15.

. 1971. Fortschritte in der Klassifizierung der

Pseudothelphusidae.—Anzeiger der Mathema-

tisch Naturwissenschaftliche der Osterreichisch-

en Akademie der Wissenschaften (1)179(1—4):

14-24.

1972. Die Pseudothelphusidae (Crustacea

Brachyura).—Zoologica 42(120), pt. 1:1—182.

. 1978. Neue Potamocarcinini, Poglayen-Neu-

wall leg. 1975 (Vorlaufiger Mitteilung).—Sit-

zungsberichte der Osterreichischen Akademie
der Wissenschaften, Mathematisch Naturwis-
senschaftliche Klasse (1)1978(2):51—5S4.

. 1980. Von Dr. Ivo Poglayen-Neuall 1975 in

VOLUME 112, NUMBER 3

Mittelamerika gesammelte Krabben.—Annalen
des Naturhistorische Museum, Wien 83:651—
666.

Rathbun, M. 1893. Descriptions of new species of
American freshwater crabs.—Proceedings of
the United States National Museum 16(959):
649-661, pls. 73-77.

. 1896. Descriptions of two new species of

fresh-water crabs from Costa Rica.—Proceed-

ings of the United States National Museum 18:

377-379.

. 1898. A contribution to a knowledge of the
fresh-water crabs of America. The Pseudothel-
phusidae.—Proceedings of the United States
National Museum 21(1158):507—537.

Rodriguez, G. 1982. Les crabes d’eau douce
d’Amérique. Famille des Pseudothelphusi-
dae.—Faune Tropicale 22:1—223.

561

. 1994. A revision of the type material of some
species of Hypolobocera and Ptychophallus
(Crustacea: Decapoda: Pseudothelphusidae) in
the National Museum of Natural History, Wash-
ington D.C., with descriptions of a new species
and a new subspecies.—Proceedings of the Bi-
ological Society of Washington 107:296—307.

Smalley, A. 1964a. The river crabs of Costa Rica, and
the subfamilies of the Pseudothelphusidae.—
Tulane Studies in Zoology 12:5—13.

. 1964b. 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.

Villalobos, C. 1974. Ptychophallus costaricensis a new
freshwater crab from Costa Rica.—Revista de
Biologia Tropical 21(2):197—203.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(3):562—571. 1999.

A new species of Chevalia (Crustacea: Amphipoda: Corophiidae)
from the Indian Ocean with remarks on Chevalia carpenteri and the
C. aviculae superspecies complex

Eric A. Lazo-Wasem

Division of Invertebrate Zoology, Peabody Museum of Natural History, Yale University,
170 Whitney Avenue, P.O. Box 208118, New Haven, Connecticut 06520-8118

Abstract.—A new amphipod, Chevalia hirsuta, is described and figured from
material collected in the Indian Ocean by the Yale Seychelles Expedition. The
relationship of this species to all known Chevalia is discussed, along with a
review of several morphological characters of potential diagnostic value.

The genus Chevalia was reviewed by
Barnard & Thomas (1987), who described
a new species, Chevalia carpenteri from
Belize, adding to a genus previously re-
garded as monotypic. Their review also re-
established the validity of C. mexicana
Pearse; .1913, and Cs. megualis. (Stout,
1913), which were previously synony-
mized with Chevalia aviculae Walker,
1904 (Shoemaker 1942; Barnard 1962,
1970). Furthermore, Barnard & Thomas
(1987) showed that a Chevalia superspe-
cies complex exists, with variants known
from the Galapagos (Barnard 1979), Fiji
(Myers 1985) and Hawaii (Barnard 1970),
in addition to C. inaequalis and C. mexi-
cana. Clearly, Chevalia aviculae has been
a convenient taxonomic assignment for
what is actually a complex of sibling spe-
cies.

Another new species is herein described
from the Seychelles Archipelago in the In-
dian Ocean, collected during the Yale Sey-
chelles Expedition (YSE), 1957-1958. The
relationship of this new species to other
Chevalia (in particular to C. aviculae and
C. carpenteri), a discussion of diagnostic
characters and an emendment of the key
provided by Barnard & Thomas (1987) are
presented.

Chevalia hirsuta new species
Figs. 1—6

Diagnosis.—Article 2 of pereopods 5—7
ovate, broadly expanded, posterodistal cor-
ner not produced. Pereopods densely se-
tose; dense bundles of long setae on pos-
terior margin of article 2 pereopods 5-7,
anterodistal and posterodistal margin of ar-
ticle 4, pereopods 3—7, uropod 1 peduncle
and uropod 3 rami. Pleonal epimera round-
ed.

Description of holotype female.—Head
nearly twice as long as first pereon seg-
ment; coxae small, separated from each oth-
er. Mandible, palp article 1 short, article 2
slightly longer than article 3, articles 2 and
3 setose. Lower lip, inner lobes and outer
lobes separated, distal margins broadly con-
vex, setose. Maxilla 1, inner plate small,
lightly setose, outer plate with 11 distal
spines, palp 2-articulate, first article short.
Maxilla 2, inner plate with oblique row of
facial setae and setose inner margin, outer
plate distally setose. Maxilliped, inner plate
setose along inner margin, outer plate with
7 spines along inner margin, palp 4-articu-
late, second article long, article 3 with
oblique row of long facial setae, article 4
tapered, distally setose. Antenna 1 longer
than antenna 2, densely setose, accessory

VOLUME 112, NUMBER 3

flagellum 3-articulate, third article minute,
length of article 1 of flagellum nearly 300%
that of article 2, and 2 densely setose. An-
tenna 2, length of article 1 of flagellum
nearly 300% that of article 2. Gnathopod 1
much smaller than gnathopod 2, coxa pro-
duced, anterior margin weakly concave,
posterior margin broadly rounded, article 2
with dense posterodistal bundle of very
long setae, article 5 linear, longer than ar-
ticle 6, densely setose, article 6 subchelate,
ovate, palm oblique, convex. Gnathopod 2,
ventral margin of coxa excavate, anterodis-
tal corner rounded, article 2 shorter than ar-
ticle 6, article 5 short with small posterior
lobe, article 6 broad, palm oblique, sinu-
soidal, proximoposterior tooth strong,
pointed, posterior margin lined with fasci-
cles of setae and smaller curly facial setae.
Pereopod 3, length of gill and oostegite
subequal to that of article 2, article 2, an-
terior margin setose, article 4 expanded,
distal margin armed with many long setae,
article 5 short, dactyl simple. Pereopod 4,
gill 80% length of article 2, oostegite 125%
length of article 2, article 2, anterior margin
setose, article 4 expanded, article 5 short,
dactyl simple. Pereopod 5, gill 30% longer
than article 2, oostegite longer than entire
appendage, article 2 posterior margin ex-
panded, with dense medial bundle of long
setae, article 5, posterodistal corner with 4
stout submarginal spines. Pereopods 6 and
7 similar, basis expanded, posterior margin
setose, article 4, anterodistal and posterod-
istal corners each with bundles of very long
setae, article 5, posterodistal corner with 4
stout submarginal spines; pereopod 6, gill
nearly as long as basis. Uropod 1, peduncle
with dense distal bundle of long setae, outer
ramus shorter than inner, opposing inner
margins (viewed dorsally) with complexly
serrate teeth. Uropod 2, outer ramus shorter
than inner ramus, posterior margin setose,
apically spinose, inner ramus lanceolate,
margins naked. Uropod 3, distal margin of
inner and outer ramus with dense bundles
of extremely long setae. Telson subquad-
rate, distal corners weakly produced, with

563

scale-like plates, posterior margin with two
sparse bundles of long and short setae, lat-
eral margins lightly setose.

Male.—The males are smaller, but very
similar to the females in the major diag-
nostic features. However, in the largest
male (YPM 9741) the setal bundles are gen-
erally denser and the setae slightly longer
than in the largest females.

Variation.—In the smallest individuals of
both sexes, article 2 of pereopods 5-7 is
only slightly expanded, and the posterior
margin is nearly straight and parallel to the
anterior margin. In these smaller individu-
als the diagnostic long setae are present but
relatively shorter and fewer in number than
in larger specimens; smaller specimens,
however, typically bear longer setae on the
posterior margin of the antennae than those
exhibited by the holotype female.

Etymology.—The specific epithet is de-
rived from the Latin hirsutus, meaning
hairy, and alludes to the extremely long and
dense setae of the pereopods, urosome, and
telson.

Material examined.—Female holotype,
YPM 9206, 10.1 mm. Seychelles: Maliki
Pate, off Cerf Island, 4°33’N, 55°30’E, un-
der algae on dead coral. Yale Seychelles
Expedition Station 29. Collector, A. J.
Kohn, 2 November 1957. Male allotype,
YPM 9741, 9.1 mm. 7 female paratypes,
YPM 9742. 6 male paratypes, YPM 9743.
Female paratype, 7.0 mm, YPM 9744. 20
male and female paratypes, YPM 9828. Fe-
male paratype, 6.1 mm, YPM 9829. Allo-
type and paratypes with same data as ho-
lotype. All specimens deposited in the Yale
Peabody Museum.

Remarks.—Barnard & Thomas (1987)
subdivided the genus Chevalia into two
groups, based primarily on diagnostic fea-
tures of the oostegites and pereopods 5-7.
One group, represented by the C. aviculae
superspecies complex, was defined by hav-
ing three pairs of oostegites (pereopods 3—
5) and a produced posterodistal corner on
the basis of pereopods 5—7. Chevalia car-
penteri was placed as the sole member of a

564

S N/a
a 6 1 1 al (NR

=

SS
———

Pic. ©:

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

A: YPM 9206, Chevalia hirsuta new species, female holotype, 10.1 mm. Abbreviations are: A,

antenna; Md, mandible; LI, lower lip; Mx, maxilla; Mxpd, maxilliped; Gn, gnathopod:; P, pereopod; Pl, pleopod;
U, uropod; T, telson; ep, epimeron; in, incisor; mlr, molar; plp, palp. Gnathopods are numbered Gn1 and Gn2;

pereopods are numbered P3—P7.

second group because it possesses only 2
pairs of oostegites (on pereopods 4 and 5),
and the basis of pereopods 5-7 is ovate, the
posterodistal corner not produced. The new
species described herein, C. hirsuta, has
broadly rounded posterodistal corners of the
basis of pereopods 5-7 and three pairs of
oostegites; as such, it represents an inter-
mediate form with characters overlapping
both groups.

Based upon the ovoid condition of article
2 pereopods 5-7, C. hirsuta is morpholog-
ically allied to C. carpenteri of the Carib-
bean. However, based upon its possession
of three pairs of oostegites, and rounded
(i.e., not produced) epimeral plates, C. hir-
suta is similar to all forms of the C. avi-

culae complex as defined by Barnard &
Thomas (1987). Aside from the apparently
unique combination of oostegite and basis
characteristics of pereopods 5—7, C. hirsuta
clearly differs from all known species of
Chevalia in its possession of dense bundles
of exceptionally long setae on various parts
of the pereopods and uropods.

The type species of Chevalia, C. avicu-
lae, was described from Sri Lanka [former-
ly Ceylon] by Walker (1904) and has also
been reported from Madagascar by Ledoyer
(1982), although the latter record is now
provisionally attributed to C. mexicana
Pearse 1913 (Barnard & Thomas 1987).
Walker’s figures are not very detailed, and
the form he described is not easily distin-

VOLUME 112, NUMBER 3

565

NAN A) \

\

Velabitaee . ny
mete eM

4 Mine Ne Are ag ee
Q Ay NY nee AIAN
\ ENO EN
ofan a
\ ans
\\
\

ANH WW

Fig. 2. YPM 9206, Chevalia hirsuta, new species, female holotype, 10.1 mm. For abbreviations, see Fig. 1.

guishable from other members of the C. av-
iculae superspecies complex (Barnard &
Thomas 1987). The diagnostic posterodis-
tally produced article 2 of pereopod 7 (=
Walker’s pereopod 5, plate VII, figure 50),
is clearly figured, however, and serves to
distinguish that form from C. hirsuta. These

two variant forms of Chevalia (i.e., from
Madagascar and Sri Lanka), although col-
lected from areas relatively near the type
locality of C. hirsuta, are both easily distin-
guished from the latter by the diagnostic
condition of pereopods 5—7. Additionally,
although the examination of additional

566

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Pies:

YPM 9206, Chevalia hirsuta, new species, female holotype, 10.1 mm. For abbreviations, see Fig. 1.

VOLUME 112, NUMBER 3 567

Fig. 4. YPM 9206, Chevalia hirsuta, new species, female holotype, 10.1 mm. For abbreviations, see Fig. 1.

568 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

mA

P6

i

|

il

wy.
fi

Fig. 5. YPM 9206, Chevalia hirsuta, new species, female holotype, 10.1 mm. For abbreviations, see Fig. 1.

VOLUME 112, NUMBER 3 569

Fig. 6. Chevalia hirsuta, new species. Telson, YPM 9744, female paratype, 7.0 mm. A: YPM 9829, female
paratype, 6.1 mm; B: YPM 9741 male allotype, 9.0 mm.

570

specimens may prove otherwise, C. hirsuta
is very large compared to all other species
of Chevalia (adults); the largest specimen
of C. hirsuta examined (YPM 9206) is
nearly 2.5 times larger than any other
known species of the genus.

There is some similarity between small
specimens of C. hirsuta and C. pacifica, re-
cently described from Madang Lagoon,
Papua, New Guinea (Myers 1995); in both
species, article 2 of pereopod 7 is ovate,
and not produced posteroventrally. Cheva-
lia hirsuta differs from C. pacifica in hav-
ing a prominent palmar tooth on gnathopod
2, which is distinct even in the smallest
specimens (4—5 mm); a palmar tooth is
lacking in C. pacifica as figured by Myers
(1995). Furthermore, in small specimens of
C. hirsuta the rami of uropod 1 are sube-
qual in length, whereas in the figured spec-
imen of C. pacifica the inner ramus is near-
ly 30% longer than the outer ramus.

Barnard & Thomas (1987) defined a var-
iant of the C. aviculae complex, based upon
a description of C. aviculae from Fiji by
Myers (1985). Subsequently, Myers (1995)
erected C. pacifica to attribute full species
status to the Fijian variant; however, he
based this new taxon upon specimens col-
lected from New Guinea, rather than Fiji.
Myers’ (1995) figure of pereopod 7 of C.
pacifica from New Guinea differs from
what he had figured previously for the Fiji
specimens (Myers 1985). The Fiji speci-
mens, therefore, probably represent a dif-
ferent, and as yet undescribed, species.

Two characters are of potential future
value in evaluating the species assignments
of Chevalia: the number of submarginal
spines at the anterodistal corner, article 5,
pereopods 5-7; and the basic morphology
of the subserrations of the teeth on the inner
margin of the outer ramus, uropod 1. In C.
hirsuta there are 4 anterodistal submarginal
spines on article 5, pereopods 5—7. On the
holotype of C. carpenteri (USNM 195157)
there are only two spines. Furthermore,
these spines are bifid on the type specimen,
compared to simple on C. hirsuta. An ex-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

amination of several of the specimens of
Chevalia spp. described by Barnard &
Thomas (1987) has confirmed different pat-
terns of this character, depending on spec-
imen provenance. Strong differences were
also noted between the micromorphology
of the subserrations of the teeth on the inner
margin of the outer ramus of uropod 1. In
C. hirsuta, the marginal subserrations are
large and very irregular, whereas in C. car-
penteri the subserrations are small and very
regular. These two characters should prove
useful in helping to distinguish, perhaps as
new species, several of the morphs referred
to by Barnard & Thomas (1987) from var-
ious localities around the world.

Key to the taxa of Chevalia (modified
from Barnard & Thomas 1987)

1. Article 2 of pereopods 5—7 ovate, not
produced posteroventrally
Article 2 of pereopod 7 subquadrate,
produced or squared posteroventally ...

aviculae complex and pacifica

2. Oostegites 2 pairs, article 2 of pereopods
5—7 in fully developed adults moderately
expanded, length more than twice width,
pereopods 3—7, urosome and telson with
normal setae’ 2.4 3. oh = ae carpenteri
Oostegites 3 pairs, article 2 of pereopods
5-7 in fully developed adults greatly ex-
panded, width 90% length; pereopods 3—

7, urosome and telson with dense bun-
dles of exceptionally long setae ... hirsuta

Site ia, wt we: (ai te a) ce

Acknowledgments

The author thanks M. E Gable (Eastern
Connecticut State University) and J. D.
Thomas (Nova Southeastern University) for
reviewing a draft version of this manu-
script. I am grateful to E. Harrison-Nelson
(United States National Museum of Natural
History) for allowing me to examine spec-
imens of Chevalia, including the holotype
of Chevalia carpenteri.

Literature Cited

Barnard, J. L. 1962. Benthic marine Amphipoda of
Southern California.—Pacific Naturalist 3:1—72.

VOLUME 112, NUMBER 3

. 1970. Sublittoral Gammaridea (Amphipoda)
of the Hawaiian Islands.—Smithsonian Contri-
butions to Zoology 34:1—286.

. 1979. Littoral gammaridean Amphipoda from

the Gulf of California and Galapagos Islands.—

Smithsonian Contributions to Zoology 271:1l-

149.

, & J. D. Thomas. 1987. A new species of
Chevalia from the Caribbean Sea (Crustacea:
Amphipoda).—Proceedings of the Biological
Society of Washington 100:532—542.

Ledoyer, M. 1982. Crustacés amphipodes gammariens
families des Acanthonotozomatidae 4 Gammar-
idae.—Faune de Madagascar 59(1):1—598.

Myers, A. A. 1985. Shallow-water coral reef and man-
grove Amphipoda (Gammaridea) of Fiji.mRe-
cords of the Australian Museum, Supplement 5:
1-144.

. 1995. The Amphipoda (Crustacea) of Madang

Lagoon: Aoridae, Isaeidae, Ischyroceridae and

571

Neomegamphopidae.—Records of the Austra-
lian Museum, Supplement 22:25-—95.

Pearse, A. S. 1913. Notes on certain amphipods from
the Gulf of Mexico, with descriptions of new
genera and new species.—Proceedings of the
United States National Museum 43:369-379.

Shoemaker, C. R. 1942. Amphipod crustaceans col-
lected on the Presidential Cruise of 1938.—
Smithsonian Miscellaneous Collections
101(11):1-52.

Stout, V. R. 1913. Studies in Laguna Amphipoda.—
Zoologisches Jahrbucher fiir Systematik 34:
633-659.

Walker, A. O. 1904. Report on the Amphipoda col-
lected by Professor Herdman, at Ceylon, in
1902.—Ceylon Pearl Oyster Fisheries, Supple-
mentary Report 17:229-—300.

. 1909. Amphipoda Gammaridea from the In-

dian Ocean, British East Africa, and the Red

Sea.—Transactions of the Linnean Society of

London (2) Zoology 12:323-344.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(3):572—580. 1999.

Megagidiella azul, a new genus and species of cavernicolous
amphipod crustacean (Bogidiellidae) from Brazil, with remarks on its
biogeographic and phylogenetic relationships

Stefan Koenemann and John R. Holsinger

Department of Biological Sciences, Old Dominion University,
Norfolk, Virginia 23529-0266, U.S.A.

Abstract.—Megagidiella azul, a new genus and species, is described from
Gruta do Lago Azul, a cave in central-western Brazil. With a body length of
more than 16 mm, this species is the largest bogidiellid recorded to date. In
addition to its large size, the absence of a mandibular palp is a unique diag-
nostic character for the family Bogidiellidae and alone merits recognition of a
new genus. The occurrence of Megagidiella azul in an isolated, inland cave
habitat marks another exceptional biogeographic record of a bogidiellid am-

phipod from South America.

Recent biological exploration of caves by
speleologists in several Karst areas in Brazil
has revealed many new localities for gam-
maridean amphipod crustaceans and other
subterranean organisms (Pinto-da-Rocha
1995). One such investigation in the Serra
da Bodoquena Karst of central-western Bra-
zil resulted in the discovery of a new sty-
gobiont amphipod genus of the family Bog-
idiellidae, described below. The specimens
were collected from a deep, subterranean
lake in Gruta do Lago Azul (Blue Lake
Cave).

Megagidiella, new genus

Diagnosis.—Eyes absent. Body smooth,
unpigmented. Uronites not fused. Coxal
plates longer than wide, not overlapping.
Coxal gills occurring on pereopods 4—6;
sternal gills absent. Oostegites on pereo-
pods 2—5, sublinear. No sexual dimorphism.
Interantennal (lateral) lobe of head narrow-
ly rounded anteriorly. Mandibular palp ab-
sent. Maxilla 1: palp 2-segmented; outer
plate with 7 serrate spines; inner plate with
3 apical plumose setae. Gnathopod 1 pro-
podus much larger than gnathopod 2 pro-
podus. Pereopods 5—7 with narrow bases.

Pleopods and uropods unmodified. Pleo-
pods biramous; outer ramus 3-segmented;
inner ramus reduced, 1-segmented. Uro-
pods biramous; peduncle of uropod 1 with
several ventrolateral (basofacial) spines;
uropod 3 relatively long. Telson about as
long as broad, with shallow excavation.

Type species.—Megagidiella azul, new
species by monotypy; gender feminine.

Etymology.—The generic name, referring
to the relatively large size of the type spe-
cies, is a combination of the Greek prefix
‘““mega’’ (= large) and part of the family
name.

Remarks and _ relationships.—Bogidiel-
lids are relatively small amphipods, their
body lengths generally range between 1-3
mm, occasionally exceeding 5 mm. With
adult specimens reaching a body length of
16.2 mm, Megagidiella is an extraordinary
exception. The more significant diagnostic
character, however, is the absence of a man-
dibular palp, a morphological reduction to
date not reported in the family Bogidielli-
dae (sensu Stock 1981). Apart from its size
and absence of a mandibular palp, Mega-
gidiella closely resembles the typical mor-
phology of Bogidiella, s. str., e.g., gnatho-

VOLUME 112, NUMBER 3

aS

Se ‘

—_
—r

Fig. 1.
Grosso do Sul, Brazil. Note: buccal mass is shaded.

pod 1 larger than gnathopod 2; pereopods
3—7 with narrow bases; coxal plates not
overlapping, wider than long; 3-segmented
pleopodal outer ramus; reduced, 1-seg-
mented pleopodal inner rami. Minor excep-
tions from the general bogidiellid model are
a 1-segmented accessory flagellum and the
armature of the telson. Of all described
bogidiellid species, a 1-segmented acces-
sory flagellum is known only in 4 genera:
Artesia Holsinger (in Holsinger & Longley
1980), Kergueleniola Ruffo, 1970, Marigi-
diella Stock, 1981, and Parabogidiella Hol-
singer (in Holsinger & Longley 1980).
The armature of the telson shows a re-
markable resemblance to that of Spelaeo-
gammarus da Silva Brum, 1975, from caves
in eastern Brazil: Megagidiella has 2—3 api-
cal and 3-5 subapical (lateral) spines per
telsonic lobe in comparison with 2 apical
and 3—4 subapical spines in Spelaeogam-
marus. The combination of 2 apical spines
with more than 2 subapical spines is excep-
tional for bogidiellids. Moreover, the ar-
mature and shape of uropods 1-3 show
noteworthy similarities in both genera, for
example, a row of long setae on the medial

373

Megagidiella azul, n. sp., holotype female (16.2 mm) from Lago Azul Cave, Bonito, Estado Mato

margin of the outer ramus of uropod 3.
Along with Artesia, from an Artesian Well
in Texas, these are the only bogidiellids
known with setae on the rami of uropod 3.

Megagidiella azul, new species
Figs. 1-4

Material examined.—Holotype female
(16.2 mm), allotype male (15 mm), and 3
paratypes (1 male, 1 female, 1 juvenile),
collected by Adrian Boller, 1 July 1991.

Type locality.—Gruta do Lago Azul,
northwest of Bonito, Estado Mato Grosso
do Sul, Brazil.

The holotype is dissected and mounted
on microscope slides in Faure’s medium. It
is deposited in the Museu Nacional (UFRJ)
in Rio de Janeiro, Brazil (MNRJ 13339).
The allotype and paratypes are preserved in
alcohol and will be retained in the research
collection of JRH under the catalog no. H-
3487.

Diagnosis.—With the characters of the
genus. Largest male 15 mm, largest female
16.2 mm (Fig. 1).

Description.—Antenna 1 (Fig. 2a) about

574

50% length of body. Peduncular segments
1—3 gradually decreasing in length distally.
Primary flagellum longer than peduncle,
with up to 19 articles in adult specimens,
without aesthetascs. Accessory flagellum 1-
segmented.

Antenna 2 (Fig. 2b) about half as long as
antenna 1. Peduncular segment 4 longer
than segment 5. Flagellum as long as pe-
duncular segment 5, with 5 articles.

Upper lip (Fig. 2c) rounded apically, with
setules along distal margin.

Mandible (Fig. 2h, 1): palp absent; molar
prominent, rounded, weakly triturative,
bearing 1 long, finely serrate seta; left la-
cinia mobilis 5-dentate, right lacinia 2-den-
tate, with serrate upper margin; left and
right mandible with 4—6, variably plumose
accessory spines.

Lower lip (Fig. 2d) bearing setules on
outer lobes and on distal margins of inner
lobes; inner lobes small but distinct; lateral
processes short with bluntly rounded cor-
ners.

Maxilla 1 (Fig. 2e): Palp 2-segmented,
with 3 apical setae. Outer plate with 7
comblike spines (Fig. 2f), bearing loosely
inserted setules on surface and in row along
medial margin. Inner plate with marginal
setules and 3 apical plumose setae.

Maxilla 2 (Fig. 2g): Outer plate with ap-
proximately 24 naked apical setae; apical
margin of inner plate bearing about 17 na-
ked setae and 3 plumose setae; both plates
with fine setules.

Maxilliped (Fig. 2j): Palp 4-segmented;
3 blunt spines along apical margin of outer
plate; apical margin of inner plate with 2
bifid (y-shaped) spines, 4 plumose setae,
and 1 naked seta.

Gnathopod 1 (Fig. 3a): Basis naked,
bearing only 1 short seta at distoposterior
corner. Carpus short, triangular shaped,
with 2 setae on pointed posterior lobe. Pro-
podus almost twice as long as broad, ap-
proximately twice the size of gnathopod 2
propodus. Palmar margin oblique and even,
finely serrate along whole margin, with 5
medial and 5—6 lateral spines; medial mar-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

gin with about 27 short setae and 4 angular
spines of unequal length. Dactyl about 80%
length of propodus.

Gnathopod 2 (Fig. 3b): Basis naked,
bearing only 1 short seta at distoposterior
corner. Propodus bearing 18—20 short setae
(12-13 laterally and 6—7 medially), 5 spines
near corner, and a single lateral spine at
mid-palmer margin. Palm with distinctly
oblique, finely serrate margin. Dactyl about
60% length of propodus.

Pereopods 3 and 4 subequal (Fig. 4a, b).
Bases narrow, anterior margins little ex-
panded. Dactyls 24—27% length of propods.

Pereopods 5—7 (Fig. 4c—e) increasing in
length posteriorly. Bases narrow, posterior
margins very weakly expanded. Dactyls
about 22, 26, and 28% length of propods,
respectively.

All pereopod bases apparently without
lenticular organs.

Coxal plates small, wider than long;
plates 1—4 rectangular, plates 5—7 at least 2
times wider than long.

Coxal gills (Fig. 4a, d) present in 3 pairs,
ovate on pereopods 4 and 5 and sack-
shaped on pereopod 6.

Oostegites (Fig. 3b, 4a, c) small, subli-
near, occurring on pereopods 2—5 (not se-
tose in material examined).

Epimeral plates (Fig. 3c) with small, but
distinct distoposterior corners, bearing 1 se-
tule each in groove immediately above cor-
ner.

Pleopods 1-3 (Fig. 4f) similar. Inner ra-
mus reduced, 1-segmented, with terminal
plumose seta. Outer ramus 3-segmented,
with 2 terminal plumose setae per segment.

Uropod 1 (Fig. 4g) biramous, outer ra-
mus slightly shorter than inner ramus; rami
about 64% length of peduncle. Peduncle
bearing 14—15 spines, 3 of which inserted
along ventrolateral (basofacial) margin.
Outer ramus with 12 lateral spines and 4
apical spines. Inner ramus with 4—5 apical
and 5 dorsomedial spines.

Uropod 2 (Fig. 4h): Inner and outer rami
subequal, slightly longer than peduncle. Pe-
duncle with 6 spines. Outer ramus bearing

VOLUME 112, NUMBER 3

373

Fig. 2. Megagidiella azul, n. sp., holotype female (16.2 mm): a) antenna | (accessory flagellum enlarged),
b) antenna 2, c) upper lip, d) lower lip, e) maxilla 1, f) enlarged spine and seta types of maxilla 1, g) maxilla
2, h) left mandible, i) incisor, lacinia mobilis, and spine row of right mandible, j) maxilliped.

8 lateral spines and 4 apical spines (2 long
ones and 2 short ones). Inner ramus bearing
5 spines along medial and lateral margins
and 5 apical spines (3 long ones and 2 short
ones).

Uropod 3 (Fig. 41) long, with subequal,
l-segmented rami. Peduncle about 48%
length of rami, with 2—4 spines. Outer ra-
mus with 6 apical spines and 6 sets of
spines along lateral margin (with 1—5 spines
per set); medial margin bearing 4—5 long

plumose setae. Inner ramus with 6—7 apical
spines and about 19 medial and _ lateral
spines (some doubly inserted).

Telson (Fig. 3d, e) about as broad as
long; apex with shallow excavation (8%
length of telson); each half bearing 2 plu-
mose setae, 2 (sometimes 3) apical and 3—
5 subapical spines.

Variation.—Morphological variation,
apart from usual differences between juve-
niles and adults (e.g., number of spines on

576

Fie. 3
plates, d) telson, e) telson, allotype male (15 mm).

appendages, flagellum articles, etc.), was
observed most obviously in the armature of
the telson. The number of subapical spines
in the adult females (16 mm in length) var-
ied from 2 to 5 per side, whereas both adult
males (15 and 11 mm in length) had a con-
stant number of 3 subapical spines. In the
holotype female, a short third apical spine
was inserted on the left telsonic apex (Fig.
3d).

Etymology.—The epithet azul is based on
the name of the type locality and is used as
a noun in apposition.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

a

Megagidiella azul, n. sp., holotype female (16.2 mm): a) gnathopod 1, b) gnathopod 2, c) epimeral

Discussion

The type material was collected at a
depth between 6 and 12 m from a deep,
turquoise-blue lake inside Blue Lake Cave.
The cave is located at the southern edge of
the world’s largest wetland area along the
Serra da Bodoquena in central-western Bra-
zil (Pinto-da-Rocha 1995). Because of the
large cave entrance, the lake, about 50 m
inside the cave, receives light during some
hours of the day (Pires 1987). The water in
the lake presumably marks the upper por-
tion of a subterranean aquifer.

VOLUME 112, NUMBER 3 577

Fig. 4. Megagidiella azul, n. sp., holotype female (16.2 mm): a) pereopod 3, b) pereopod 4, c) pereopod 5,
d) pereopod 6, e) pereopod 7, f) pleopod 2, g) left uropod 1, h) left uropod 2, i) left uropod 3.

578

Blue Lake Cave was already biogeo-
graphically significant prior to the discov-
ery of Megagidiella azul, inasmuch as it is
the only known locality in the western
hemisphere for the extremely rare crusta-
cean order Spelaeogriphacea. Prior to the
discovery of Potiicoara brasiliensis Pires,
1987 in Blue Lake Cave, the only other
spelaeogriphacean known to science was
Spelaeogriphus lepidopus Gordon 1957
from caves on Table Mountain in South Af-
rica. One explanation for the occurrence of
freshwater stygobiont spelaeogriphaceans
in caves on opposite sides of the Atlantic
Ocean is that these species are derived from
a common ancestor which inhabited Gond-
wana prior to the separation of Africa and
South America in the Early Cretaceous. Al-
though it is tempting to speculate that bog-
idiellids and spelaeogriphaceans share a
similar evolutionary history affected by
continental drift, there is to date no evi-
dence that the ranges of these groups form
a generalized distribution track. Bogidiel-
lids are recorded only from a few localities
near coastal regions in northeastern and
northern Africa, whereas the freshwater
amphipod fauna in central and southern Af-
rica is composed primarily of epigean par-
amelitids, and stygobiont ingolfiellids and
sternophysingids.

From an ecological perspective, it is im-
portant to note that M. azul dwells in a large
lake of phreatic water. The extraordinary
size of this species might imply a correla-
tion of body size and available habitat
space. An interesting parallel example of
this phenomenon can be observed in the
amphipod family Ingolfiellidae. Most in-
golfiellids, like many bogidiellid taxa, are
less than 3 mm long and live in interstitial
habitats. In contrast to the norm, however,
species of the ingolfiellid genus Troglole-
leupia live in large “‘open” cave lakes in
central and southern Africa and may reach
23 mm in length (Griffiths 1989).

Bogidiellid amphipods have a near
world-wide distribution pattern, occuring
exclusively in subterranean habitats. Their

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

distribution pattern is characterized by sev-
eral regions with relatively dense concen-
trations of species. For example, the South
American continent shows the highest ge-
neric diversity as opposed to the Mediter-
ranean region where species richness is
higher but generic diversity is lower. To
date, 18 species, distributed among 10 gen-
era and subgenera, are known from South
America (Fig. 5).

The discovery of Megagidiella azul in
the interior of South America, approximate-
ly 1000 km from the nearest coast, is bio-
geographically significant because the vast
majority of bogidiellids occupy ranges be-
tween 100—200 km from marine coastal re-
gions. South America shows a remarkable
pattern of isolated aquatic habitats, and has
promise for the future study of stygobiont
organisms and their environments.

Subsequent to the completion of the
manuscript, we received additional mega-
gidiellids from several new localities in the
state Mato Grosso do Sul, Brazil. These
specimens match the description given in
this paper and show the same morphologi-
cal variation as observed in the type spe-
cies.

New localities.—2 juveniles (6 and 7
mm) from Gruta do Mimoso, Bonito, col-
lected by E. P. Costa, Jr., Feb and Jun, 1998;
1 female (12.5 mm) from Abismo do Pogo,
Jardin, collected by N. Moracchioli, Jun,
1998; 2 females (13 mm) from Buraco dos
Abelhas, Jardin, collected by E. P. Costa,
Jr., Apr and Jun, 1998.

Comments.—According to Dr. Eleonora
Trajano (pers. comm.), all specimens were
found in large cave lakes. They occurred in
the water column at depths of 20-52 me-
ters. Spelaeogriphaceans were also present
in Gruta do Mimoso, but they inhabited
only the benthic sediments of the lake.

Acknowledgments

We are grateful to Dr. Eleonora Trajano,
University of Sao Paulo, Brazil, for provid-
ing us with the specimens of the new bog-

VOLUME 112, NUMBER 3 a79

10N |

P
yi ‘ re VENEZUELA
Kis =

SEE me

0 ECUADOR
reel Spas

aN

10S

20S

Pacific Ocean

30S

Atlantic Ocean

40S

50S

80W 70W 60W 5O0W 40W

Fig. 5. Geographic distribution of bogidiellid amphipods in continental South America: (1) Bogidiella cooki
Grosso & Ringuelet (1979): (2) B. gammariformis Sket (1985); (3) B. neotropica Ruffo (1952); (4) B. (Dycti-
cogidiella) ringueleti Grosso & Fernandez (1988); (5) B. (Dyct.) talampayensis Grosso & Claps (1985); (6) B.
(Mesochthongidiella) tucumanensis Grosso & Fernandez (1985); (7) B. (Stygogidiella) hormocollensis Grosso
& Fernandez (1988); (8) B. (Styg.) lavillai Grosso & Claps (1984): (9) Eobogidiella purmamarcensis Karaman
(1982); (10) Marigidiella brasiliensis Stock (1981; see also Siewing, 1953); (11) Megagidiella azul, n. gen., n.
sp. (background darkened for emphasis); (12) Patagongidiella danieli Grosso & Fernandez (1993) and P. mauryi
Grosso & Fernandez (1993) (both in same locality); (13) Pseudingolfiella chilensis Noodt (1965); (14) Spelaeo-
gammarus bahiensis da Silva Brum (1975) and S. spp.

580

idiellid amphipod for identification and
study, and to Adrian Boller for collecting
the bogidiellids from Blue Lake Cave. We
also thank the Graphics Office at Old Do-
minion University for generating of the dis-
tribution map. This study was supported by
a PEET grant from the National Science
Foundation to JRH (DEB-9521752).

Literature Cited

da Silva Brum, I. N. 1975. Spelaeogammarus bahien-
sis g.n. sp.n. de Anfipodo Cavernicola do Brasil.
(Amphipoda-Bogidiellidae).—Atas da Socieda-
de de Biologia di Rio de Janeiro, 17:125—128.

Gordon, I. 1957. On Spelaeogriphus, a new cavernic-
olous crustacean from South Africa.—Bulletin
of the British Museum of Natural History (Zo-
ology) 5:31—47.

Griffiths, C. L. 1989. The Ingolfiellidae (Crustacea:
Amphipoda) of southern Africa, with descrip-
tions of two new species.—Cimbebasia 11:59—
70.

Grosso, L. E., & G. L. Claps. 1984. Tercer Bogidiel-
lidae (Crustacea Amphipoda) de la cuenca del
Rio Grande (Jujuy, Argentina).—Neotropica
30:223—231.

5 Ge . 1985. Distribuci6n geografica de

la familia Bogidiellidae (Crustacea, Amphipo-

da) en la Republica Argentina, con la descrip-

cidn de un nuevo subgénero y una nueva es-

pecie.—Physis (Buenos Aires), Secc. B, 43:49—

55.

, & H. R. Fernandez. 1985. Una nueva Bogi-

diella (Amphipoda Bogidiellidae) hiporreica de

la provincia de Tucumdan (Argentina).—Neotro-

pica 31:201—209.

,& . 1988. Un caso de simpatria de tres

especies del género Bogidiella (Crustacea, Am-

phipoda) en el noroeste Argentino, con la de-

scription de dos nuevoas especies.—Stygologia

4:64-78.

Mere . 1993. Nuevo género cavernicola

austral de Bogidiellidae; Patagongidiella n.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

gen. del noroeste Patagénico (Neuquén, Argen-

tina).—Bolletino del Museo Civico di Storia

Naturale, Verona 17:357—372.

, & R. A. Ringuelet. 1979. Fauna subterranea
de las aguas dulces de la Reptiblica Argentina.
I. Dos nuevas especies de Amphipodos del gé-
nero Bogidiella.—Limnobios 1:381—394.

Holsinger, J. R., & G. Longley. 1980. The subterranean
amphipod crustacean fauna of an artesian well
in Texas.—Smithonian Institution Press 308:i—
ili, 1-62.

Karaman, G. S. 1982. Critical remarks to the recent
revisions of bogidiella-group of genera with
study of some taxa (fam. Gammaridae). Contri-
bution to the knowledge of Amphipoda 126.—
Poljoprivreda I Sumarstvo Titograd, 28:31—57.

Noodt, W. 1965. Interstitielle Amphipoden der kon-
vergenten Gattungen J/ngolfiella Hansen und
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Crustaceana 9:17—30.

Pinto-da-Rocha, R. 1995. Sinopse da fauna cavernicola
do Brasil (1907—1994).—Papéis Avulsos de
Zoologia (S. Paulo) 39:61—173.

Pires, A. M. S. 1987. Potiicoara brasiliensis: a new
genus and species of Spelaeogriphacea (Crus-
tacea: Peracarida) from Brazil with a phyloge-
netic analysis of the Peracarida—Journal of
Natural History 21:225—238.

Ruffo, S. 1952. Bogidiella neotropica n. sp., nuovo
Anfipodo dell’ Amazonia.—Rivista Svizzera di
Idrologia 14/1:129-134.

. 1970. Descrizione di Kerguelenella macra n.
g. n. sp. (Amphipoda Gammaridae). Studi sui
Crostacei Anfipodi LX VIII.—Bolletino del Mu-
seo Civico di Storia Naturale, Verona 18:43—54.

Siewing, R. 1953. Bogidiella brasiliensis, ein neuer
Amphipode aus dem Kuestengrundwasser Bras-
iliens.—Kieler Meeresforschungen, Band 9,
Heft 2:243—247.

Sket, B. 1985. Bogidiella (s. 1.) gammariformis sp. n.
(Amphipoda) from Equador.—BioloSki Vestnik
33:81-88.

Stock, J. H. 1981. The taxonomy and zoogeography
of the family of Bogidiellidae (Crustacea, Am-
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dam) 51:345-—374.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(3):581-591. 1999.

The western and southern distribution of Mesocyclops edax
(S. A. Forbes) (Crustacea: Copepoda: Cyclopoida)

Janet W. Reid and Luis Moreno D.

(JWR) Research Associate, Department of Invertebrate Zoology, National Museum of Natural
History, Smithsonian Institution, Washington, D.C. 20560-0163, U.S.A.; (LMD) Departamento de
Hidrobiologia, CIRA/UNAN, Centro para la Investigaci6n en Recursos Acuaticos de Nicaragua,
Universidad Nacional Autonoma de Nicaragua, Apartado Postal 4598, Managua, Nicaragua

Abstract.—A review of published records and material of the cyclopoid co-
pepod Mesocyclops edax (S. A. Forbes) established its presence in the drainage
basins of the Colorado and Columbia Rivers in the western U.S.A., and verified
published records from Cuba, Guatemala, and Mexico. Mesocyclops nicara-
guensis Herbst is a junior synonym of M. edax. We provide supplementary
morphological observations and describe certain morphological variations ob-

served in some populations of M. edax.

The cyclopoid copepod Mesocyclops
edax (S. A. Forbes, 1891) is a common and
often numerous plankter in surface water-
bodies of North America, and its biology
and genetics have been the subject of many
reports (e.g., Williamson 1986; Wyngaard
1991; Wyngaard et al. 1985, 1995). Meso-
cyclops edax is widespread in south-central
and eastern Canada (Patalas 1986) and in
the conterminous U.S.A. east of the Rocky
Mountains; the animals undergo diapause in
winter in colder latitudes. The western and
southern limits of M. edax are less well
documented. Patalas (1986) failed to find it
in his own collections or in published re-
cords west of the Canadian Pacific conti-
nental divide. Published records of M. edax
from the western U.S.A. are few, and con-
firmed records from Mexico and Central
America even fewer. We review western
and southern records of M. edax, confirm-
ing those records where possible through
examination of available specimens.

The question of the synonymy of Meso-
cyclops nicaraguensis Herbst, 1960 with M.
edax, first raised by Cole (1976), has con-
tinued unresolved. We compared specimens
from the type-locality of M. nicaraguensis
in Lake Nicaragua, and from other water-

bodies in Nicaragua to populations of M.
edax from across North America.

Dussart (1985) and Dahms & Fernando
(1995a) redescribed adult M. edax. We add
observations on certain morphological fea-
tures of interest for the systematics of the
genus, and note some morphological vari-
ations observed during the course of this
study.

Specimens were deposited in the collec-
tions of the National Museum of Natural
History (USNM), and the Centro para la In-
vestigaci6n en Recursos Acuaticos de Nic-
aragua, Universidad Nacional Aut6noma de
Nicaragua (CIRA/UNAN).

Order Cyclopoida G. O. Sars, 1886
Family Cyclopidae Burmeister, 1834
Subfamily Cyclopinae Dana, 1853
Genus Mesocyclops G. O. Sars, 1914
Mesocyclops edax (S. A. Forbes, 1891)
Figs. 1-3

Cyclops edax S. A. Forbes, 1891:709-—710,
TS 7A Fe Fisys pl fie 1S Ppl TV) figs:
16-19.

Cyclops leuckarti Claus, 1857 (partim).—
Marsh 1893:191, 193, 209-211, 224, pl.
IV fig. 17, pl. V figs. 2—6.

582

Mesocyclops edax.—G. O. Sars 1914:58.

Cyclops leuckharti.—Coker 1926:230—234,
247-248, 257, pl. 40 figs. 5-7.

Mesocyclops Leuckarti edax.—Kiefer
1929a:306—307, figs. 4, 5.

Mesocyclops obsoletus (Koch, 1838).—C.
B. Wilson 1929:129.

Mesocyclops nicaraguensis Herbst, 1960:
27, 42—45, figs. 33-41.

Non Cyclops simplex Poggenpol, 1874.—
Herrick, 1887:14, 17-18, pl. VII fig. 1.
?Non Cyclops tenuissimus Herrick, 1883:
499-500, pl. V figs. 24, 25, pl. VI, figs.

20,218

Material examined.—Canada: USNM
79773, 10 2 2°, Lake-of-the-Woods, Ontar-
io, 17 Jul 1909. USNM 251908, 6 2°, 8
36, Red Bridge Pond, Halifax, Nova Sco-
tia, 8 Jul 1992, leg. G. A. Wyngaard.
U.S.A.: USNM Acc. No. 120079 (Marsh
Collection), 1 2, slide, Hutchins, Texas, C.
D. Marsh prep. no. 3101. USNM 62590, 10
2 2, Lake Erie, 12 Jul 1928, don. Buffalo
Society of Natural Science. USNM 78730,
10 2&, Welaka, Florida, 1 Apr 1938, leg.
L. Cable. USNM 216873, 10 2 2, Crane’s
Pond, Newport, North Carolina, 24 Aug
1984, leg. M. C. Swift. USNM 235221, 7
22, Lake Powell, Utah, 17 Jan 1985, leg.
L. Haury. USNM 251156, 60+ 22 36
and copepodids, Lake Union, Seattle, State
of Washington, 47°39'12”N, 122°19'24’W,
10 Oct 1991, leg. Parametrix Inc. for Mu-
nicipality of Metropolitan Seattle (MET-
RO), gift of J. R. Cordell. USNM 264309,
13 22, Hueco Spring I, near New Braun-
fels, Cornal Co., Texas,.8—10 Aug 1992,
leg. €. B. Bares USNMi2781335 152 26'S
2 copepodids, San Joaquin River, Fresno,
California, 10 Sep 1995, leg. S. Callison.
Mikropraparate 2993 and 2994, 1 °, partly
dissected on 2 slides, Lake Otay, San Die-
go, California, 29 Feb 1936, leg. Light,
prep. E Kiefer, Kiefer Collection, Staatlich-
es Museum fiir Naturkunde Karlsruhe. 20
22 $6, culture, New Orleans, Louisiana,
1989, collection of G. G. Marten. Mexico:
USNM 251674, 1 CV 6, Lake Patzcuaro,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Michoacan, 19°30'N, 101°38’W, 17 Oct
1991, leg. A. Orbe-Mendoza and P. Aguilar.
USNM 259699, 1 2, Cenote Viejo, Quin-
tana Roo, Mexico, 21 Jul 1987, leg. E. Sua-
rez-Morales. Guatemala: USNM 250931, 6
22 436 1 copepodid stage V (CV) @, all
mounted on 2 slides, 16 Mar 1972, and 3
22 1 6, all mounted on 3 slides, 19 Aug
1972, Lake Izabal, leg. L. G. Brinson, prep.
det. H. C. Yeatman. Nicaragua: USNM
259614, 1 2 (dissected on slide), 1 36 5
copepodids, Lake Nicaragua (Lake Coci-
bolca), Managua, 1991. USNM 259616, 6
22 1 3 57 copepodids, Lake Nicaragua,
1993. USNM 243696, 10 22, Lake Aso-
sosca, Managua, 22 Oct 1991. USNM
243700, 10 2 10 36, Lake Las Canoas,
21 Jan 1993. USNM 243701, 8 22 3464,
Lake Nicaragua, Sample 97-11, 13 Jun
1997. USNM 243707, 200+, Lake Nicara-
gua, Samples N97-09 and N97-11, 13 Jun
1997. USNM 243710, 100+, Lake Asosos-
ca, Managua, 22 Oct 1991. USNM 243713,
500+, Lake Managua, Point 7, 23 Mar
1995. USNM 243715, 100+, Lake Las
Canoas, 21 Jan 1993. Legs. L. Moreno. Ad-
ditional material deposited in the Plankton
Collection, CIRA/UNAN. Cuba: 1 2°, part-
ly dissected on slide, Sabanilla Reservoir,
15 May 1965, collection of C. H. Fernando.

Supplementary description of female.—
We describe certain morphological features
of the adult female, to supplement previous
redescriptions (Dussart 1985, Dahms &
Fernando 1995a).

Medians (ranges, n) of lengths, excluding
caudal setae, in mm: Canada: Ontario
(USNM 79773), 1.41 (1.24—-1.48, n = 10);
Nova Scotia (USNM 251908), 1.43 (1.38—
1.49, n = 6). U.S.A.: Lake Erie (USNM
62590), 1.23 (1.06—-1.33, n = 10); Florida
(USNM 78730), 1.27 (1.09—-1.35; n = 10);
North Carolina (USNM 216873), 1.31
(1.22—1.38, n = 10); Utah (USNM 235221),
1.17 (1.11—1.18, n = 7); Louisiana (Marten
collection), 1.40 (1.26—1.53, n = 10); Texas
(USNM 264309), 1.30 (1.28—-1.36, n = 10).
Mexico: Quintana Roo (USNM 259699),
1.08 (n = 1). Guatemala: Lake Izabaél

VOLUME 112, NUMBER 3

(USNM 250931), 1.26 (1.23—1.40, n = 9).
Nicaragua: Lake Nicaragua (Lake Cocibol-
ca) (USNM 259614), 1.18 (n = 1); Lake
Nicaragua (USNM 259616), 1.38 (1.26—
1.45, n = 6); Lake Asososca (USNM
243696), 1.02 (0.96—1.15, n = 10); Lake
Las Canoas (USNM 243700), 1.17 (1.09-—
1.31, n = 10).

Habitus (Fig. 1A) typically cyclopiform;
occasional individuals with circular pits on
prosomites. Pediger 5 (Fig. 1B—E) usually
without ornament except normal hair-sen-
silla, except group of hairs on lateral sur-
faces in one female from Nicaragua. Geni-
tal double somite (Fig. 1B—D) with group
of 6 tiny pores posterior to leg 6, pore pat-
tern otherwise as in figures. Pore-canal usu-
ally single as in Fig. 1D, but sometimes ap-
pearing divided (Fig. 1K G). Posterior pore-
canal usually represented as stout and pos-
teriorly directed, but strongly recurved in
oblique view (in direction of arrows, Fig.
1G). Anal somite (Fig. 1H) with continuous
row of spines along posterior margin; cau-
dal ramus usually divergent in preserved
specimens, with 4—5 irregular groups of
hairs on medial surface.

Hyaline membrane of antennule terminal
segment beginning at lateral seta, usually
with several large notches (Fig. 2A). Spec-
imen from Cuba with this condition exag-
gerated on one side, giving impression of
single deep notch (Fig. 2B).

Antenna basis (Fig. 2C—F) in most pop-
ulations with secondary curved row of
small spines (Fig. 2E, arrow; Dahms & Fer-
nando 1995a), row absent in some popula-
tions (Fig. 2C).

Maxilla (Fig. 2G) with group of tiny
spines on frontal side.

Maxilliped (Fig. 2H, I) usually with more
complex surface spine pattern than indicat-
ed by Dahms & Fernando (1995a).

Legs 1—4, caudal surfaces of coxa-bases
ornamented as in Fig. 3A—D. Medial spine
of leg 1 basipodite with slender spinules in
North American populations (Fig. 3A), with
more and stouter spinules in some southern
populations, e.g., Cuba (Fig. 3E). Couplers

583

(intercoxal sclerites) of legs 1—3 always
with small submarginal spinules (Fig. 3F—
H, J—L), number of spinules on leg 1 cou-
pler varying from 1 to 5 in North American
populations and from 7 to 10 on specimen
from Cuba, with similar, but fewer spines
on couplers of legs 2 and 3. These spines
usually stout, but slightly longer and slim-
mer in specimens from Guatemala and Nic-
aragua. Two projections on margin of leg 4
coupler usually rounded (Fig. 3M), but
sometimes acute and inward- or outward-
directed (Fig. 3D, I). Both terminal spines
of leg 4 endopodite 3 with spinulate mar-
gins, spinules of lateral terminal spine few-
er and coarser than spinules of medial ter-
minal spine in all specimens examined (Fig.
3N).

Discussion

Balcer et al. (1984), Coker (1943), Cole
(1960), and Patalas (1986) traced the
changing descriptions and taxonomy of Me-
socyclops edax. These changes are a story
of advances in understanding the taxonomy
of the genus Mesocyclops and the morpho-
logically similar genus Thermocyclops (see
also comments by Dahms & Fernando
1995a, 1995b). In spite of much previous
discussion, three points remain to be re-
solved.

The synonymy of Mesocyclops_ nicara-
guensis Herbst, 1960.—The question of the
possible synonymy of M. nicaraguensis
Herbst, 1960, which was described from
Lake Nicaragua, was first raised by Cole
(1976). Yeatman, who determined Cole’s
Nicaraguan copepods, affirmed Cole’s opin-
ion (H. C. Yeatman, in litt. to JWR, 1991).
Reid (1990) also proposed the synonymy.
Absent formal examination, however, ™.
nicaraguensis has continued to be treated as
a distinct taxon (e.g., Cisneros & Mangas
1991, Collado, 1984, Dussart & Fernando
1986). The type specimens of M. nicara-
guensis cannot be located (H.-V. Herbst, in
litt. to JWR, 1991), and there is no non-
type material from the original collection

584 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

— a ~=
——

BEG

Fig. 1. Mesocyclops edax, female. A-C, F—H, specimens from North Carolina, U.S.A. (USNM 216873); D,
Louisiana, U.S.A. (Marten Collection); E, Lake Nicaragua (USNM 259614). A, Habitus, lateral; B, Pediger 5
and genital double somite, left lateral; C, Pediger 5 and genital double somite, dorsal; D, Pediger 5 and genital
double somite, ventral (leg 5 setae foreshortened); E, Pediger 5, left lateral; EK Copulatory pore and pore-canal,
ventral, enlarged; G, Copulatory pore and pore-canal, ventral-oblique, enlarged; H, Anal somite and caudal rami,
dorsal. Scales equal 100 pm.

VOLUME 112, NUMBER 3

A B

585

a MUU radii

Fig. 2. Mesocyclops edax, female. A, C, D, G—I, specimens from North Carolina (USNM 216873); B, E, F
Specimen from Cuba (Fernando Collection). A, B, Antennule terminal segment; C, E, Antenna basis, caudal
side; D, EK Antenna basis, frontal side; G, Maxilla, frontal side; H, I, Maxilliped. Scales equal 100 um.

(G. Hartmann, in litt. to JWR, 1991). No
material remains from Cole’s (1976) study
of Lake Nicaragua (H. C. Yeatman, in litt.
to JWR, 1991).

There are a few differences between pub-
lished descriptions of M. edax and M. ni-
caraguensis. The most obvious is the

lengths of adult females, 0.83—1.1 mm for
M. edax (Dahms & Fernando 1995a, Dus-
sart 1985) and about 1.45 mm for M. ni-
caraguensis (Herbst 1960). This difference
in lengths was the only discriminator used
by Dussart (1987) and Petkovski (1986) in
their keys to American Mesocyclops. How-

586

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Pre_ 3:
(Fernando Collection); J-M, Guatemala (USNM 250931). Scale equals 100 um.

ever, we report populations of M. edax from
Ontario, Nova Scotia, and Louisiana with
individuals longer than 1.45 mm. Lengths
of females from lakes in Nicaragua varied
from 0.96 to 1.45 mm. There is then no
meaningful difference in the lengths of
northern and southern populations, nor is
there any correlation between length and
latitude.

Mesocyclops edax, female. A—D, N, specimens from North Carolina (USNM 216873); E-I, Cuba

The remaining apparent difference con-
sists of the presence of submarginal spines
on the leg 1 coupler in M. edax, while
Herbst (1960: fig. 37) indicated no orna-
ment on the leg 1 coupler in M. nicara-
guensis. However, all specimens from Lake
Nicaragua examined by us had such spines.
A single female from Nicaragua bore small
spines on the lateral surface of pediger 5

VOLUME 112, NUMBER 3

(Fig. 1E). The presence or absence of
spines at this location is a much used char-
acter in the genus systematics. However, the
facts that this condition is apparently ex-
tremely rare and found only in one Nicar-
aguan population, and that all females from
Nicaragua were otherwise congruent in all
respects with the diagnosis of M. edax, per-
suaded us to consider M. nicaraguensis
synonymous with the former taxon.

Habitat as well as genetic factors may ac-
count for some variation. Barwick & Hud-
son (1985) reported that individuals of M.
edax in the tailwater of a reservoir were
small and the posterior edges of the uro-
somes were sculptured, in contrast to a
‘reservoir form’’ that averaged larger and
had smooth urosomal edges. Specimens
were, however, unavailable (P. L. Hudson,
in litt. to JWR, 1998). Wyngaard et al.
(1985) noted differences in body size,
clutch size, and egg size among wild pop-
ulations from Michigan and Florida.

The taxon Cyclops tenuissimus (Herrick,
1883).—Cole (1960) suggested that Cy-
clops tenuissimus, described by Herrick
(1883) from Kentucky, U.S.A., is a senior
synonym of M. edax. Cole pointed out that
the combination of several characters, the
relationships of the lengths of the leg 5 ter-
minal seta and subterminal spine, the anten-
nular hyaline membrane with several small
notches, and the caudal rami hairs pos-
sessed by C. tenuissimus matches only M.
edax among North American cyclopoids.
However, Herrick showed the genital seg-
ment of C. tenuissimus as very slender and
the seminal receptacle with narrow lateral
and posterior extensions, and in his text de-
scribed (and named) this species as ‘“‘the
most slender cyclops known to me’’. Cole’s
argument might easily be accepted if not for
the contradictions between Herrick’s de-
scription and the relative robustness of M.
edax and the broad lateral and posterior ex-
pansions of its seminal receptacle. Since
Herrick’s description matches no presently
known American cyclopoid species in all

587

respects, in our opinion the identity of Her-
rick’s specimens remains in question.

The western and southern distribution of
M. edax.—The published records from Pa-
cific drainage basins in the U.S.A. and Can-
ada are few and relatively recent. Several
were confirmed by material made available
for this report, and we consider all of them
entirely reliable. Simenstad & Cordell
(1985) listed M. edax from the Columbia
River basin in Washington State. The pre-
viously unpublished record herein from
Lake Union, Washington, also comes from
the Columbia River basin. The few records
from the Colorado River basin include
those of Soto & Hurlbert (1991la, 1991b)
from California, and of Maddux et al.
(1987) and Sollberger & Paulson (1992)
from the Colorado River in Arizona and
Nevada. The records reported herein from
San Diego and Fresno, California are new.
In western Canada, M. edax was recently
reported from British Columbia (Chenga-
lath & Shih 1994, Shih & Chengalath
1994).

Only recent reports of M. edax from
Mexico, Central America, and the Antilles
are reliable. As noted by Cole (1960), prior
to and even following the revision of Coker
(1943) which established the distinction be-
tween M. edax and populations in the
Americas referred to as M. leuckarti, M.
edax was likely to be reported as M. leuck-
arti (e.g., Comita 1950). In the Americas,
records of M. leuckarti may refer to other
members of the /euckarti-group, such as M.
americanus Dussart, 1985, M. aspericornis
(Daday, 1906), or M. ruttneri Kiefer, 1981,
or even to any of several species of Ther-
mocyclops now known to be present in the
region (Dussart 1985, Dussart & Fernando
1990, Reid 1993, Reid & Reed 1994). Cole
(1960) pointed out inconsistencies in
Marsh’s (1910) description of M. leuckarti
which apparently led Marsh to synonymize
M. edax with the former species. Examples
of Marsh’s confusion can be found among
the slide-mounted specimens labeled “‘M.
leuckarti’’ in the Marsh Collection at the

588

National Museum of Natural History.
Among these specimens one of us (JWR)
has identified M. longisetus (Thiébaud,
1912), M. reidae Petkovski, 1986, M. ve-
nezolanus Dussart, 1987, and Thermocy-
clops decipiens Kiefer, 1929b, but neither
M. leuckarti nor M. edax. The extensive
Marsh Collection contains no unequivocal
specimen of M. edax from any locality
south of Texas. Pearse’s (1921) record of M.
leuckarti var. edax from Lake Valencia,
Venezuela, may refer to M. brasilianus Kie-
fer, 1933, as suggested by Infante et al.
(1979), or to some other congener with
haired caudal rami, such as M. aspericornis
which also occurs in Lake Valencia (Reid
& Saunders 1986). Pearse’s (1915) record
of M. leuckarti var. edax from Colombia is
similarly suspect.

Additional problems are the sparsity of
general collections of plankton from Cen-
tral America and the Antilles, and the pres-
ence of M. edax and M. leuckarti, but not
other congeners, in the widely used key of
Yeatman (1959). Before publication of the
keys to American Mesocyclops by Petkov-
ski (1986) and Dussart (1987) and a local
checklist by Reid (1990), any of the many
congeners in the region was likely to be
identified as one of those two species. Sim-
ilar doubts as to the validity of Central
American records were implied in the sum-
mary by Collado, Fernando, & Sephton
(1984) of the distribution of zooplankters
from this area, since those authors included
no records of M. edax other than those from
Cuba by Smith & Fernando (1978, 1980).

Smith & Fernando’s record from Cuba is
the only report of M. edax from the Antil-
les, and is supported by preserved material.

Most of the few Mexican and Central
American records of M. edax can be like-
wise verified. Coker (1943) and Osorio-Taf-
all (1944) reported M. edax from Lake
Patzcuaro, Mexico. Copepodids of Meso-
cyclops, including one stage V copepodid
referable to M. edax from Lake Patzcuaro
examined for this report substantiate their
records. Mesocyclops edax was reported

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

from the Yucatan Peninsula (and specimens
archived) by Suarez-Morales et al. (1996).
The reports by Brinson & Nordlie (1975)
and Deevey et al. (1980) from eight or more
lakes including Lake Izabal in the Guate-
malan lake district were substantiated by
the specimens furnished by H. C. Yeatman.
The scarcity of reports of M. edax south
of the U.S.A. may correctly reflect its dis-
tribution. General collections, mainly from
shallow waterbodies in Costa Rica (Collado
& Fernando 1984) and Honduras (Marten
et al. 1994) failed to record this species.
The presence of M. edax in several large
Central American lakes and its apparent ab-
sence from smaller waterbodies might im-
ply some direct temperature effect. On the
other hand, it was collected from Laguna
Sabanilla near Havana, Cuba, which is shal-
low (maximum depth 1.3 m) and near sea
level (Straskraba et al. 1969). Some of the
Guatemalan lakes are also shallow, such as
Petenxil (4 m maximum; Deevey et al.
1980). Better adapted neotropical cyclo-
poids may outcompete M. edax in most
conditions. One candidate might be Ther-
mocyclops decipiens, which is a pantropical
facultatively omnivorous plankter that is
frequently successful in the mesotrophic to
eutrophic conditions in which M. edax also
thrives (Reid 1989). It is usual for M. edax
and the much smaller Thermocyclops inver-
sus Kiefer, 1936 to co-occur (Coker 1943,
Cole 1976, Deevey et al. 1980, Osorio-Taf-
all 1944, and our samples from Nicaragua).
In Guatemala, T. decipiens (reported by
Deevey et al. [1980] as Mesocyclops hyali-
nus [Rehberg, 1880], a synonym) occurred
with M. edax in only one lake. Thermocy-
clops decipiens was also numerous in the
sample from Laguna Sabanilla examined
for this report. Thermocyclops decipiens
ranges from 0.75—0.99 mm in length, thus
approaching the length of M. edax. It is
common in the Antilles including Cuba and
in Central America (Reid 1985, 1989), but
does not extend northwards past southern
Mexico (Suaérez-Morales & Reid 1998).

VOLUME 112, NUMBER 3

Acknowledgments

C. B. Barr, S. Callison, G. A. Cole, J. R.
Cordell, C. C. Davis, C. H. Fernando, C. C.
Hakenkamp, G. Hartmann, L. R. Haury, P.
L. Hudson, A. de Infante, T. Ishida, V. Ko-
rinek, G. G. Marten, S. Montenegro-Guil-
len, A. Orbe-Mendoza, G. L. Pesce, J. E
Saunders III, E. Sudarez-Morales, M. C.
Swift, W. Wurtsbaugh, G. A. Wyngaard, H.
C. Yeatman, and the late H.-V. Herbst lent
their cordial assistance in answering inqui-
ries, searching literature, and lending or do-
nating specimens. H.-W. Mittmann lent
specimens from the E Kiefer Copepod Col-
lection in the Staatliches Museum fiir Na-
turkunde Karlsruhe. The Administrative
Counselor, American Embassy Managua
expedited shipment of specimens from Nic-
aragua. We are grateful for a Short-Term
Visit Grant awarded to L. Moreno D. by the
Smithsonian Institution, and to the Depart-
ment of Invertebrate Zoology, NMNH for
providing research facilities.

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(3):592-597. 1999.

A new species of Orbiniella (Orbiniidae: Polychaeta) from Marion
Island, Indian Ocean

Patrick Gillet

Laboratoire d’Ecologie Animale, Institut de Recherche Fondamentale et Appliquée,
U.C.O. BP 808, 49008 Angers Cedex 01, France

Abstract.—After the Marion Dufresnes MD/03 Expedition in 1974, a new
expedition Marion Dufresnes MD/08 was undertaken to study benthic inver-
tebrates from the South West Indian Ocean. Orbiniella marionensis, a new
species of Orbiniidae, Polychaeta from Marion Island, Indian Ocean is de-
scribed. This species was collected during the Marion Dufresnes BENTHOS
Expedition around subantarctic Islands. Orbiniella marionensis is compared
with other species of the genus Orbiniella Day, 1954 and differs in having only
one preanal segment and by the absence of modified setae in the posterior

region.

Materials and Methods

Polychaetes collected during the Marion
Dufresnes BENTHOS Expedition from
Marion and Prince Edward Islands were de-
scribed by Gillet (1991). A description of
the sampling site of the MD/08 BENTHOS
Expedition is provided by Arnaud & Hu-
reau (1979). Different sampling methods
were used: Charcot dredge, Okean grab,
Lithods nets and Trawl. Specimens were
conserved in 70% ethanol and observed by
light microscopy with an Olympus CHT2
and by scanning electron microscopy with
an Olympus JSM 5200.

Material has been deposited at the Insti-
tut de Recherche Fondamentale et Appli-
quée, Angers-France and a duplicate collec-
tion in the Museum National d’ Histoire Na-
turelle de Paris, France and in the South
African Museum, Cape Town, South Afri-
ca.

Family Orbiniidae Hartman, 1942

The name Orbiniidae was used by Hart-
man (1942) in place of Ariciidae with the
type genus Orbinia, Quatrefages, 1865.

The orbiniids have been studied by Hart-
man (1957), Pettibone (1957) and more re-

cently reviewed by Day (1973). The orbi-
niids have lateral parapodia in the thoracic
region and dorsal parapodia in the abdom-
inal region. The prostomium is without ap-
pendages and is followed usually by one or
two asetigerous segments. The setae are
simple capillaries and simple hooks, some-
times with furcate or crenulate setae. Hart-
man (1957) divided the family in two sub-
families: Orbiniinae, including the larger
orbiniids with one achaetous segment be-
hind the prostomium and, Protoariciniinae
with two achaetous segments behind the
prostomium. The Protoariciniinae was re-
vised by Solis-Weiss & Fauchald (1989).

Genus Orbiniella Day, 1954

Type species.—Orbiniella minuta Day,
1954: 22-23, fig. 3 g—k.

Prostomium round. First two segments
without setae. Setigerous segments without
gills or appendages. Setae include crenulate
capillaries and simple acicular. In Fauchald
(1977) the genus Orbiniella lacks branchi-
ae. Buzhinskaja (1992) suggested that Falk-
landiella annulata Hartman (1967) be
transferred to Orbiniella. Buzhinskaja
(1992) suggested that Orbiniella branchiata

VOLUME 112, NUMBER 3

be referred elsewhere. Blake (1996) sug-
gests that some species of Orbiniella might
be juveniles of other orbiniids such as Na-
ineris. In the natural world, 10 species of
Orbiniella are known.

Orbiniella marionensis new species
Figs. 1A—D, 2A—D; Table I

Material examined.—Holotype: Marion
Island 26.03.1976; station 18 (BB 108)
46°49.8'S, 37°56.2’E, complete, 29 setigers,
7 mm long, 1.3 mm wide with parapodia
and 1.0 mm wide without parapodia. Para-
types: Marion Island station 12 (BB 79),
46°55.7'S, 37°54.1'E, 6 exemplars; station
f5e(BB 88), 46°57.7'S, 37°59.9’E, 1 ex-
emplar; station 18 (BB 108) 46°49.8’S,
37°56.2'E, 12 exemplars; station 34 (BB
168), 46°50.2’S, 37°51.2’E, 4 exemplars
Material deposited IRFA Angers IRFA-
ORB.O15.

Description.—Holotype colorless in eth-
anol, with one preanal achaetous segment.
The prostomium is rounded, 0.2 mm long
and 0.4 mm wide, large and eyeless. The
second and third segments are achaetous
and without parapodium (Figs. 1A, 2A, B).
The following segments are setigerous,
without gills and ventral cirri and with dor-
sal cirri (Figs. 1D, 2D). Setae include cren-
ulate capillaries (Fig. 1B) and from the first
setiger 2 to 4 simple acicular spines (Fig.
1C). The pygidium is round with one pre-
anal segment (Fig. 2C).

The paratype material generally ressem-
bles the holotype closely. The number of
setigers varies between 14 (4 mm long) and
29 (7 mm long) and width varies from 1.3
mm with parapodia to 1.0 mm without
parapodia.

Discussion.—Orbiniella marionensis dif-
fers from Orbiniella minuta Day, 1954, de-
scribed from Tristan da Cunha (Day 1954)
and from Marion and Prince Edward Is-
lands (Day 1971) by the presence of dorsal
cirri (Table I). Orbiniella marionensis has
two to four acicular setae from the first se-
tiger. It differs by the absence of modified

395

crenulate setae in the posterior region and
by having only one preanal achaetous seg-
ment. Orbiniella uniformis Hartman, 1967
lacks dorsal cirri and has only one falcate
acicular spine from setiger 15. Orbiniella
drakei Hartman, 1967 has ventral and dor-
sal cirri and a modified posterior region
with very long segments. Orbiniella mari-
onensis also differs from O. dayi Branch,
1998 which has no preanal segments and
cirri are absent. For the same reasons, it dif-
fers from O. hobsonae Blake & Hilbig,
1990. These two species differ by the kind
of setae, simple acicular and crenulate cap-
illaries in O. dayi but barbed acicular and
bristled capillaries in O. hobsonae. Orbi-
niella aciculata Blake, 1985 has an unseg-
mented posterior region. O. annulata Hart-
man (1967) O. nuda Hobson, 1974 and O.
plumisetosa Buzhinskaja, 1992 have eyes in
the anterior region and have different setae:
O. plumisetosa has plumose cirrate setae,
O. nuda has spinous capillaries and O. an-
nulata has crenulate capillaries.

Habitat.—Infralittoral from 95 to 210
meters depth on sand, clay-sand or mud.

Subantarctic distribution.—Marion Is-
lands.

Etymology.—The specific name mari-
onensis refers to Marion Islands and is a
Latin adjective.

Key to the genera Orbiniella Day, 1954

Mem VeS OhESeMt yy terrae oe cc's thw he - 2
Eyes absent

2. Body colored, plumate cirrate setae pre-
SEM CS reg oer ae ads lis Sa '8 ia Riese O. plumisetosa
Body without pigment, plumate cirrate
setae absent

O. nuda
Acicular setae appearing from setiger 12—15
O. annulata

apiewisy ys) ‘ef e, innate) fey @1he .0.. 4/0) Ss (0:1 e816 oe Ly Sole ey ee

alle 676 ©8658 @Ce bere aie. ee 8). ee) ayie) se e's

4. Dorsal or ventral cirri present
Dorsal or ventral cirri absent
5. Both dorsal and ventral cirri present.
Two preanal segments one acicular seta
aac ee Th eee anal Ce Se O. drakei
Only dorsal cirri present, one preanal

594 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

0.2 mm ie ee
re

= |
B eS | 0.05mm
0.1mm

Fig. 1. Orbiniella marionensis, n. sp. (Holotype, IL.R.EA. ORB.O15) A: Anterior region; B: Crenulate seta;
C: Acicular seta; D: Parapodium.

395

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VOLUME 112, NUMBER 3

segment, two-four acicular setae ....
ESE REV ESE ys ote «2h a O. marionensis
G))Preanal ‘segments present) 0.2... 7
Preanal segments absent
7. Two preanal segments
Preanal segment unsegmented ......
I ee ogc gs Se vay ns O. aciculata
8. Crenulate setae present, two acicular se-
BAe nis cpa) wp IEE 2 O. minuta
Crenulate setae absent, only one acicular
S52) les os RAOee ee Bee eee O. uniformis
9. Acicular simple, crenulate capillaries
Acicular barbed, bristled capillaries
O. hobsonae

O. dayi

Acknowledgments

I am grateful to James A. Blake, Battelle
New England Marine Research Laboratory,
Duxbury, U.S.A. for his help in bibliogra-
phy and his careful review of the manu-
Script.

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dition to Tristan da Cunha 1937-1938. 29:1-35.

. 1971. Polychaeta. Marion and Prince Edward

Islands. E. M. Van Zinderen Bakker, J. M. Win-

terbottom, R. A. Dryer, eds., Cape Town: 383—

390.

. 1973. New Polychaeta from Beaufort with a
key to all species recorded from North Caroli-
na.—NOAA Technical Report 375:1—140.

Fauchald, K. 1977. The Polychaete worms definitions
and keys to the orders, families and genera.—
Natural History Museum of Los Angeles Coun-
ty, Science Series 28:160 pp.

Gillet, P. 1991. Biogeography and polychaete assem-
blages from Subantarctic Islands (Indian
Ocean). Marion Dufresne Expedition to Marion,
Prince Edward and Crozet Islands.—Bulletin of
Marine Science 48(2):358—368.

Hartman, O. 1942. A review of the type of polychae-
tous annelids at the Peabody Museum of Nat-
ural History, Yale University.—Bulletin of the
Bingham Oceanographic Collection 8:1—98.

. 1957. Orbiniidae, Apistobranchidae, Paraoni-

dae and Longosomidae.—Allan Hancock Pacif-

ic Expedition. 15:205-—393.

. 1967. Polychaetous annelids collected by the
USNS Eltanin and Staten Island Cruises, chiefly
from Antarctic Seas—Allan Hancock Mono-
graph in Marine Biology Los Angeles. 2:1—387.

Hobson, K. D. 1974. Orbiniella nuda new species (Or-
biniidae) and nine new records of other seden-
tariate polychaetous annelids from Washington
and British Columbia.—Canadian Journal of
Zoology 52:69-—75.

Pettibone, M. H. 1957. North American genera of the
family Orbiniidae (Annelida: Polychaeta) with
descriptions of new species.—Journal of the
Washington Academy of Science 47:159—167.

Solis-Weiss, V., & K. Fauchald. 1989. Orbiniidae (An-
nelida: Polychaeta) from mangrove root-mats in
Belize, with a revision of Protoariciin genera.—
Proceedings of the Biological Society of Wash-
ington 102:772-—792.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(3):598—623. 1999.

Redescriptions of Grania americana, G. bermudensis and descriptions
of two new species of Grania (Annelida: Clitellata: Enchytraeidae)

from Bermuda

J. M. Locke and K. A. Coates

Department of Zoology, University of Toronto, Toronto, Ontario, M5S 3G5 Canada; (JML, KAC)

Bermuda Biological Station for Research, Inc., GEO1, Bermuda

Abstract.—The diversity of the enchytraeid genus Grania is reexamined in
Bermuda. Herein, four species, G. bermudensis Erséus & Lasserre, 1976, G.
americana Kennedy, 1966, G. laxartus, new species, and G. hylae, new species,
are described from intertidal and subtidal sand habitats of Bermuda. Grania
americana and G. laxartus are also reported from the Caribbean area and G.
hylae from Florida. Grania bermudensis is recognized as a separate species
rather than as a subspecies of G. macrochaeta (Pierantoni, 1901). Grania amer-
icana and G. laxartus have distinctive spermathecal structures and the latter
has a penial stylet. Grania bermudensis and G. hylae can be easily separated
from their Bermudian congeners by the dorsal position of spermathecal pores.
Both G. americana and G. hylae have statocyst-like sensory structures asso-
ciated with their brains. These studies provide new information diversity, dis-
tribution and taxonomy of Grania species of the North Atlantic and on the
structural diversity of male duct systems and of statocyst-like sensory struc-

tures.

The taxonomic diversity of the abundant
and widespread marine oligochaete genus
Grania Southern, 1913 (Annelida: Clitel-
lata: Enchytraeidae) in Bermuda has been
poorly investigated, as have most genera of
the marine Enchytraeidae (Healy & Coates
1999). Only one species, Grania bermuden-
sis Erséus & Lasserre, 1976, has been de-
scribed from Bermuda and it was recog-
nized as a subspecies of G. macrochaeta
(Pierantoni, 1901) (Erséus & Lasserre 1976,
Locke & Coates 1998). However, a great
degree of anatomical variability among
Grania specimens collected from Bermuda
has been recognized for some time (Erséus,
pers. comm.). Furthermore, there are only
four publications referring primarily to the
Enchytraeidae of Bermuda (Moore 1902,
Lasserre & Erséus 1976, Giere 1979, Healy
& Coates 1999) and the most recent of
these shows the species diversity of this

family is much higher than previous studies
indicated.

The single species of Grania described
from Bermuda was reported prior to the
recognition of a variety of important taxo-
nomic characters for this genus. Coates
(1984) provided new criteria for categoriz-
ing forms of the male duct system found in
the genus. These characteristics have played
a significant role in the distinction of sev-
eral new species (e.g., Coates & Erséus
1985, Erséus 1990a, Coates & Stacey
1993). Recently, another interesting struc-
tural character was revealed by Rota & Er-
séus (1996). In a few new species from the
Antarctic and Subantarctic, a structure was
observed which they called the head organ
(Rota & Erséus 1996, 1997). It was de-
scribed as a vesicular body with refractile
inclusions, located within the peristomium
just anterior to the brain. Such advances in
recognition of morphological characters

VOLUME 112, NUMBER 3

have resulted in increased knowledge of
specific diversity and provide additional
clues that can be applied to discovering
evolutionary relationships among species of
Grania (Locke 1999a).

Observations of type specimens of G.
bermudensis revealed undescribed, diag-
nostic characteristics of the spermathecae of
this species. These observations, combined
with additional information about the form
of the male ducts (Coates 1984), suggested
that G. bermudensis was in need of rede-
scription.

A major marine faunal element in Ber-
muda is species with Caribbean and coastal
southern U.S. distributions (Fig. 1) (Sterrer
1986, 1998). Grania americana is the only
described species of Grania with such a
distribution. This species was first reported
from the shores of Florida and Bimini
(Kennedy 1966) and there have been no
subsequent records. In the most recent tax-
onomic discussion of this species, Erséus &
Lasserre (1976) stated that G. americana
must be considered a nomen dubium due to
the poor condition of the type material and
lack of detail in the original description.
They also suggested it probably was very
similar to G. bermudensis, implying that G.
americana might also be a subspecies of G.
macrochaeta. In fact, Lasserre (1967) had
designated G. americana as a subspecies of
G. macrochaeta along with many other spe-
cies of Grania. The basis for this designa-
tion, as discussed by Locke & Coates
(1998, also see Locke 1999a), was probably
largely historical and possibly eurocentric.
Oligochaetes have long been considered a
lineage that originated and diversified in
non-marine habitats and, thus, a low species
diversity in marine systems was expected.
In contrast to the early opinion of Lasserre
(1967), Erséus (1974) had noted G. ameri-
cana and G. macrochaeta were probably
separate species based on definite and con-
sistent morphological differences, but that
additional information about intraspecific
variation of each taxon was needed.

New taxonomic characteristics and com-

599

plete redescriptions are presented to sub-
stantiate the morphological differences
among G. americana, G. bermudensis and
G. macrochaeta sensu stricto. Redescrip-
tions of the first two taxa and a survey of
Grania species in Bermuda were undertak-
en in order to advance understanding of the
diversity, zoogeography and evolution of
marine oligochaetes.

Materials and Methods

Sediments were collected from three in-
tertidal sites and nine subtidal sites within
Bermuda. Intertidal sites were Ferry Point
Bridge, Whalebone Bay and Rocky Hill
Park, where samples were taken from an
accumulation of sand inside dissolution
tubes in limestone rock and from two rocky
limestone beach areas, respectively. Subti-
dal sites were from Castle Harbour: Eastern
Castle Roads, Rushy Island, and a patch
reef 100 m from Rushy Island; St. George’s
Harbour: ENE and SE Paget Island; Ferry
Reach; NW Pearl Island in Great Sound;
NE entrance of Smith’s Sound; and Bailey’s
Bay. Other specimens were provided by C.
Erséus, from Hutchinson Island, Florida;
Fowey Rocks, Miami, Florida; North of
Angelfish Key, Florida Keys; and Carrie
Bow Cay, Belize (Table 1).

Specimens collected by the authors in
Bermuda were obtained by washing sam-
ples on a 300 wm sieve and then sorting the
residue in seawater using a dissecting mi-
croscope. Specimens were fixed in Kahle’s
fluid, stained lightly with borax carmine,
dehydrated and cleared, and mounted in
Canada balsam for morphological obser-
vations. Only mature specimens were iden-
tified. Specimens of all Grania species
found in Bermuda were also observed
while alive.

Type specimens of G. americana
(USNM 33005) and G. bermudensis
(USNM 53202), Castle Island, were bor-
rowed from the United States National Mu-
seum of Natural History (USNM) (Smith-
sonian Institution), Washington, D.C., and

600

Fig. 1:

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

BERMUDA ¥#

Distribution of Grania species found in Bermuda in coastal North American, Bermudian and Carib-

bean waters. I G. americana, A G. laxartus, @ G. hylae, @ G. bermudensis.

examined for identification of and compar-
ison with the new specimens. Whole
mounted specimens of Grania monosper-
matheca Erséus & Lassetre, 1976, G. amer-
icana, G. bermudensis, and Grania atlan-
tica Coates & Erséus, 1985, and the new
species described herein were also exam-
ined from the collections of the authors
(KAC) and C. Erséus.

Observations of specimens were made
using differential interference contrast light
microscopy and videomicroscopy. Thirty-
five mm photography was done with a Wild
MPS46 photoautomat on Kodak Technical
Pan film ASA 100. Drawings were made

with the aid of a drawing tube. An Olympus
AX70 Provis microscope and Toshiba
3CCD camera with Image-Pro Plus® soft-
ware were utilized for video microscope
image capture analysis and morphological
measurements. Measurements were also
taken with the use of Jandel Video Analysis
Software (JAVA) and with the aid of an oc-
ular micrometer. The mean (X) and standard
deviation (SD) were calculated for morpho-
metric data.

Measurements for G. americana and one
new species were made from Bermuda and
Carrie Bow Cay whole mounted specimens;
for the other new species and G. bermu-

601

VOLUME 112, NUMBER 3

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602

densis, Measurements were taken from
whole mounts of Bermuda specimens. Ter-
minology for the head organ was adapted
from Rota and Erséus (1996, 1997) and for
penial bulb types from Coates (1984). Ob-
servations of the “‘head organ”’ indicate that
a more appropriate name for this structure
would be the anterior brain vesicle.

Grania Southern, 1913

For a recent synonymy of the genus refer
to Locke & Coates 1998.

Grania laxartus new species
Figs. 2—6; Table 1

Holotype.—USNM_ 185951, stained,
whole mounted specimen, collected Feb.
1996 by KAC.

Type locality.—Ferry Point Bridge, Ber-
muda. 32°21'70"N, 64°42'80"W. Intertidal
coarse biogenic sands within dissolution
tubes.

Paratypes.—Three whole mounted spec-
imens, USNM 185952 Ferry Reach, Ber-
muda, BBS-77-38, C. Erséus 1977; USNM
185953 Carrie Bow Cay, Belize, CBC 85-
41, C. Erséus 1985; and Bermuda Aquari-
um, Museum and Zoo, BAMZ 1999 180
006 Ferry Reach, Bermuda, BBS-77-41, C.
Erséus 1977.

Other material examined.—About 45
live specimens from the type locality col-
lected by the authors over the period 1996—
1998. Twenty-six whole mounts from type
locality, collected (KAC) February and
March, 1996. Fourteen whole mounts col-
lected by C. Erséus from Bermuda. Five
whole mounts collected from Carrie Bow
Cay, Belize, by C. Erséus (Table 1).

Description.—External characteristics:
Live specimens translucent white with a
pink tinge due to colored blood within their
vessels. Preserved length 4.7—8.3 mm (n =
18, X = 6.7, SD = 1.1). Number of seg-
ments 33-64 (n = 20, X = 50.0, SD = 9.3).
Diameter at 4/5 90.0—160.0 wm (n = 29, X
= 129.4, SD = 15.9), diameter at clitellum
100.0—198.2 um (n = 26, X = 138.0, SD

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

= 18.3) (Fig. 2). Head pore located in an-
terior region of peristomium. Setae (Figs. 2
& 3) present from IV ventrally and XVI-—
XVIII laterally; setae absent in XII of ma-
ture worms. Ventral setae anterior to the cli-
tellum appear more robust and thicker than
postclitellar setae. Setae are pointed distally
and somewhat broad at base and with a
curve and slight toe at the proximal end but
no distinct heel (Fig. 3). Length of anterior
ventral setae 38.0—56.0 um (n = 16, X =
48.4, SD = 5.1); midbody ventral setae
34.0—50.0 um long (n = 28, X = 42.0, SD
= 4.35); postclitellar lateral setae 38.0—48.0
um long (n = 18, X = 42.4, SD = 2.8). In
ten posterior-most segments ventral setae
ranged from 37.3-75.5 um (n = 9, X =
45.7, SD = 11.7) and lateral setae from
36.0—45.2 um in length (n = 11, X = 42.5,
SD = 3.2). Spermathecal pores paired, lat-
eral, immediately posterior to 4/5. Clitellum
extends from mid XI to mid XIII. Male
pores ventral, in line with ventral setae of
preceding segments, located at about mid-
point of XII. Female pores more lateral than
male pores, just posterior to 12/13.

Internal somatic characteristics: Brain
narrow anteriorly and indented posteriorly.
No anterior brain vesicle. Ventrolateral pha-
ryngeal gland lobes very small in IV. Small
dorsolateral lobes at 4/5; two pair of small
ventrolateral lobes in V and one dorsolat-
eral pair at 5/6; in VI, two or three pairs of
ventrolateral lobes and one lateral pair at 6/7
(Fig. 2). Coelomocytes numerous anterior-
ly, not found posterior to clitellum. Pat-
terned chloragogen cells covering posterior
regions of the intestine, which often con-
tains pennate diatoms. Nephridia not seen.
Dorsal blood vessel transition to blood si-
nus in XXVI. No glands on ventral nerve
cord.

Internal reproductive characteristics:
Spermathecae (Fig. 4) paired in anterior of
V, ampulla with narrow, ventrolateral, ental
connection to esophagus; connection more
ventral than line of ectal spermathecal
pores. Spermathecal ectal duct has large
subspherical dilation midway between pore

VOLUME 112, NUMBER 3

Biss:
from Bermuda. Semischematic ventrolateral view of
whole mounted specimen showing only one of the pe-
nial apparatuses; prostomium to segment XIV, illus-
trating position and relative sizes of diagnostic struc-
tures. Scale equals 100 pum. b, brain; bv, blood vessel;
c, clitellar gland cells; mk, muscle knot on vas defer-

Grania laxartus, new species, specimen

603

Vill

NS

Ip

Fig. 3. Grania laxartus, new species, from Ber-
muda. Setae from various positions, showing broad
base and slight toe. Scale equals 20 um. Ip, lateral
posterior seta, t, setal toe.

and ampulla (Fig. 4). Ectal duct runs at
about 90° from body wall to dilation, where
it turns 90° to run parallel to body wall. The
ectal duct then penetrates the ampulla as a
cone-like protrusion (Fig. 4). Diameter of
the ectal duct dilation 18.0—32.6 um (7 =
54, X = 25.0, SD = 3.0) (Fig. 4). Width of
spermathecal ectal duct 10.0—21.9 wm (n =
46, X = 15.5, SD = 2.7) between dilation
and ampulla and 10.0—14.0 um (n = 31, X
= 11.2, SD = 1.5) between dilation and ec-
tal pore. Ampulla roughly spherical, great-
est diameter 26.0-50.0 um (n = 54, X =
42.5, SD = 4.7). Rings of sperm in wall of
ampulla of reproductive specimens. No

—

ens; pa, penial apparatus; ph, pharyngeal glands; pr,
prostomium; px, pharynx; sf, sperm funnel; sp, sper-
matheca; spp, spermathecal ectal pore; v, vas deferens;
vs, ventral seta.

604

glands at ectal pore (Fig. 4). Sperm funnels
80.0—160.0 wm long (n = 25, X = 112, SD
= 23.5), 25-55 wm maximum width (n =
25, X = 41.4, SD = 9.2) or, length about
three times width; attached sperm with
straight heads (acrosome & nucleus) (Fig.
5A). Laterally paired vasa deferentia pos-
sess a muscular “‘knot”’ at one quarter their
total length, near sperm funnel (Fig. 5A);
muscle bands run around vas and constrict
it, but there are no obvious longitudinal ex-
tensions of the muscle bands. Vas confined
to XI-XIII. Width of vas 6.3-8.8 um (n =
26, X = 7.6, SD = 0.8) both before and
after “‘knot’’. Penial apparatus with small
gland and lateral aglandular sac formed by
extended invagination of male pore, mus-
cles extend from body wall to outer wall of
sac (Fig. 5A, B). Vas enters gland from a
dorsolateral position, extends through glan-
dular material to medial side. Penial stylet
present, walls of vas deferens thin, with
pronounced nuclei where stylet begins. Sty-
let extends from ectal end of vas deferens,
tapering toward blind end of sac-like ex-
tended invagination of male pore. Length of
stylet 30.0—47.7 wm (n = 12, X = 33.9, SD
= 6.5); maximum width of stylet 1.7 um
(measured for only two specimens) (Fig.
5B). Posterior extension of sperm sac X VIII
through XXI. Egg sac extends to XXIII,
with primary oocytes located only at pos-
terior end of sac.

Etymology.—The species is named in al-
lusion to the unique shape of the sperma-
thecal ectal duct; laxartus from the Latin
laxus meaning wide or broad, combined
with artus meaning joints; a noun in ap-
position.

Taxonomic remarks.—Lateral setae are
reported as commencing in XVI-XVIII,
however, one specimen was an exception
with lateral setae beginning in XXV and
with ventral setae lacking from XVIII-—
XXIII. The ventral setal distribution for G.
laxartus is similar to G. bermudensis, G.
americana and G. hylae (Fig. 6) from the
Northwest Atlantic. The last three species
are easily distinguished from G. laxartus by

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Grania laxartus, new species, from Ber-
muda. Dorsal view of spermatheca in V. Scale equals
20 wm. a, ampulla; cp, conical protrusion of sperma-
thecal duct; e, spermathecal ectal duct; ed, ectal dila-
tion; en, spermathecal ental duct; see Fig. 2 for other
symbols.

dorsal spermathecal pores and a muscular
sac-like penial apparatus in both G. ber-
mudensis and G. hylae, and by the absence
of both a penial stylet and muscular vas de-
ferens (Coates 1984) in G. americana. Fur-
thermore, G. bermudensis, G. hylae and G.
americana each have very different sper-
mathecal morphologies, lacking the distinct
dilation of the spermathecal ectal duct that
is present in G. laxartus (Fig. 6).

This form of dilation of the spermathecal
ectal duct is reported in one other species
of Grania from the Northwest Atlantic, G.
monospermatheca, and is not known in any
other species of Grania (Locke & Coates
1998; Rota & Erséus 1996, 1997; Coates &
Stacey 1993, 1997; Coates 1990; Erséus
1990a; Coates & Erséus 1985; Coates & Er-
séus 1980; Erséus & Lasserre 1976). The
ectal duct dilation of G. monospermatheca

VOLUME 112, NUMBER 3

—

—

605

Biss:

Grania laxartus, new species, from Bermuda. A, ventral view of male duct system showing sperm

funnel, vas deferens and penial apparatus. B, lateral view of penial apparatus. Scales equal 20 pm. bw, body
wall; co, sperm funnel collar; i, invagination at male pore; m, muscle; p, male pore; pg, penial gland; st, stylet;

see Fig. 2 for other symbols.

occurs immediately before the ampulla and
is not as distinctly bulb-like as the medial
dilation of the spermathecal ectal duct in G.
laxartus. Grania monospermatheca is also
readily distinguished from G. laxartus by
the absence of lateral setae and the presence
of a single spermatheca with a dorsal pore
instead of a pair of spermathecae which
open laterally.

Habitat and distribution.—Intertidal
pools with accumulations of sand, in dis-
solution tubes formed by the erosion of
limestone rock. Also occurs subtidally.
Found in sand samples with debris along
with G. americana and various Tubificidae,
Nematoda and Sipunculida. Sand particle
size for samples ranged from fine (occur-
ring with Thalassia testudinum Koenig) to
medium coarse calcareous coral sand.
Known from Ferry Point Bridge, Whale-
bone Bay, Pearl Island, Ferry Reach and

Smith’s Sound, Bermuda; and Carrie Bow
Cay, Belize (Table 1).

Grania hylae new species
Figs. 6, 7-11; Table 1

Holotype.—USNM_ 185954, stained,
whole mounted specimen, BBS-77-42, col-
lected: by Gy Erséus 1977:

Type locality—ENE of Paget Island,
Bermuda, 32°22'38”"N, 64°38'57”"W. Subti-
dal in 17 m, medium to coarse coral sand.

Paratypes.—Three_ stained, whole
mounted specimens, USNM 185955 M77-
6, North of lighthouse, Fowey Rocks USA,
C. Erséus 1977; USNM 185956 BBS-77-15
Castle Island, Bermuda, C. Erséus 1977;
and BAMZ 1999 180 007 BBS-77-15 Cas-
tle Island, Bermuda, C. Erséus 1977.

Other material examined.—Live material
from Rocky Hill Park, Bermuda. Fourteen

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606

Fig. 6. Comparison of Grania species. Measurements are shown as averages. Spermathecae of G. bermu-

enification used for G. hylae and G. laxartus.

half the mag

densis and G. americana are shown at one

VOLUME 112, NUMBER 3

sb

VS

Fig. 7. Grania hylae, new species, specimen from
Bermuda. Semischematic lateral view of whole mount-
ed specimen; prostomium to segment XIV. Scale
equals 100 pm. sb, subesophageal ganglion; see Fig.
2 for other symbols.

whole mounted specimens from the collec-
tion of C. Erséus, from three different lo-
calities in Bermuda, and Fowey Rocks,
Florida, U.S.A. (Table 1).
Description.—External characteristics:
Fixed specimen length 5.3—-11.5 mm (n =
7, X = 8.0, SD = 2.1). Number of segments
41-61 (n = 6, X = 52, SD = 8.1). Diameter
at 4/5 139.5-192.0 um (n = 7, X = 155.9,
SD = 17.3), diameter at clitellum 154.0—
196.0 im (7 = 7, X = 171.4, SD = 17.3)

607
ew wan Pe AX kV
Fig. 8. Grania hylae, new species, from Bermuda.

Setae from different segments. Scale equals 20 pm.
vpc, ventral postclitellar seta; see Fig. 3 for other sym-
bols.

(Fig. 7). Head pore in anterior of peristom-
ium. Ventral setae (Figs. 7 & 8) present
from IV and laterally from XV—XVII. No
setae in XII of mature specimens. Length
of anterior ventral setae 37.0—66.4 wm (n
= 10, X = 51.3, SD = 8.9); midbody ven-
tral setae 49.8—91.0 um (n = 35, X = 71.8,
SD = 8.5); postclitellar lateral setae 59.8—
80.0 um: lone (2 — 31,.X, = 69.6, SD =
5.8). In ten posterior-most segments ventral
Seiac 53.770 @ = 172 X = 62:8, SD
= 5.2) and lateral setae 49.0—74.0 um (n =
14, X = 62.2, SD = 8.4). Setal shaft wid-
ening proximally, with sloping toe, distinct
heel absent (Fig. 8). Spermathecal pores
paired, dorsal, immediately posterior to 4/5
(Fig. 7), distance between pores 46.0—37.5
pm (@ = 2; XK = 41.8, SD = 6.0), no epi-
dermal papilla at pore. Clitellum extending
from post XI to mid XIII. Male pores ven-
tral, in line of setae, at midpoint of XII.
Female pores posterior to 12/13, opening
just lateral to position of male pores.
Internal somatic characteristics: Brain in-
dented posteriorly with narrow transition to
small anterior brain vesicle. Membrane-
bound, anterior, brain vesicle lies below
head pore, with one refractile inclusion
(Fig. 9). Pharyngeal glands present in seg-
ments IV through VI; one ventrolateral pair
of lobes in IV and one dorsolateral pair at

608

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Big; 9:
Scale equals 20 pm. ab, anterior brain vesicle; hp, head pore; ri, refractile inclusion; see Fig. 2 for other symbols.

4/5, two paired ventrolateral lobes in V and
one dorsolateral pair at 5/6; three pairs of
ventrolateral lobes in VI and one dorsal pair
at 6/7. No nephridia observed. Dorsal blood
vessel transition to blood sinus at XX or
X XI. Gland on ventral nerve cord in XIV.

Internal reproductive characteristics:
Spermathecae lie dorsal to pharyngeal
glands in segment V (Figs. 7 & 10). Sper-
mathecal ectal duct of uniform, narrow
width with slight curvature toward pore,
length of duct about 1.2 times maximum
diameter of ampulla. Glands lacking at ectal
pore. No protrusion of duct tissue into am-
pulla. Spermathecal ampulla small and
round to ovoid, maximum diameter 32.0—
52.0 pm (vn = 11, X= 43.858D = 63):
ampullae lie close together, dorsal to gut,
sometimes even touching; sperm rings pre-
sent (Fig. 10). Narrow ental connection of
spermathecal ampulla to esophagus. Sperm

Grania hylae, new species, from Bermuda. Lateral view of anterior brain vesicle with one inclusion.

funnel length two and a half times the
width, with non-granular section posterior
to collar. Vasa deferentia confined to XII
and XIII. Wall of vas deferens constricted
at three almost equally spaced points im-
mediately posterior to union with sperm
funnel; with slight muscularization ectal to
constrictions (Fig. 11A); vas possesses
large, obvious nuclei along length, diameter
of vas uniform along length 6.7—10.0 wm
(n = 13, X = 8.4, SD = 0.9)#Penaiyap
paratus bipartite, with horizontal and erect
sections. Penial stylet present, originating
within the vas deferens, extending just into
the penial apparatus (Fig. 11A, B). Stylet
long, 55.4-70.9 um (n = 6, X = 61.8, SD
= 7.1). Vas terminates at ental end of the
saccate part of the bipartite penial appara-
tus, which lies parallel to body wall and
gut. This sac may appear longitudinally
ridged or smooth, depending on whether it

VOLUME 112, NUMBER 3

Fig. 10. Grania hylae, new species, from Bermu-
da. Lateral view of spermatheca in V, with dorsal pore.
Scale equals 20 pm. sr, sperm ring; see Fig. 4 for other
symbols.

is contracted or relaxed. At anterior end of
sac, there is an acute to right angle bend
that unites the ridged, contracted sac to an
erect extension of the male pore (standing
more or less perpendicular to the body
wall). Muscles run from the body wall to
insert on both the longitudinal sac and onto
the lateral wall of the perpendicular exten-
sion (Fig. 11A, B). Sperm sac extends to
XVIII. Egg sac extends to XXII.
Etymology.—Named hylae in appreciation
of Hyla Tibbitts’ dedication and keen interest
in the subject matter of her illustrations.
Taxonomic remarks.—Grania hylae has
similarities to G. bermudensis in subtidal
habitat, although it has also been found in
the lower intertidal of rocky beaches, and
in the structure of both spermathecae and
the penial apparatus. Both species have dor-
sal spermathecal pores and round to ovoid
ampullae. The average maximum diameter
of the spermathecal ampulla for G. hylae,
however, is only half that for G. bermudensis.
An epidermal bump at the ectal pore is also
seen only in G. bermudensis as well as a wid-
ening of the ectal duct near the ampulla.

609

Grania hylae possesses an anterior brain
vesicle which is lacking in G. bermudensis.
Differences between the two species are
also seen in the details of the male duct
systems. The walls of the vas deferens in
G. bermudensis are highly muscularized
whereas those of G. hylae have only a thin
muscle layer. The maximum width of the
muscular part of the vas deferens in G. ber-
mudensis averages 22.9 m and the non-
muscular sections 12.3 wm; G. hylae has an
average vas width of 8.4 wm. Both species
do possess bipartite penial apparatuses with
longitudinal sac-like parts but the junctions
of vas deferens and penial sac are quite dif-
ferent.

There are a few other species of Grania
which also have dorsally positioned sper-
mathecal pores, including Grania maricola
Southern, 1913 (see also Locke & Coates
1998) in which the pores are distinctly dor-
sal to the midlateral line although not fully
dorsal. Both Grania inermis Erséus, 1990
and Grania stilifera Erséus, 1990 described
from around Hong Kong (Erséus 1990a)
have dorsal spermathecal pores but the first
of these does not have a penial stylet and
the second is reported to have a large gland
associated with the penial apparatus. There
are no separate glands on the penial appa-
ratus of either of the species with dorsal
spermathecal pores which are found in Ber-
muda, G. hylae and G. bermudensis (see
below).

Habitat and distribution.—Subtidal,
found in fine, medium and coarse coral
sand, many Tubificidae also present. Inter-
tidal, rocky limestone beach area, medium
to course sand, with G. americana, Tubifici-
dae and Nematoda. Known from Rocky Hill
Park, Castle Island, Bailey’s Bay, and ENE
and SE Paget Island, Bermuda; and Fowey
Rocks, Miami, Florida, U.S.A. (Table 1).

Grania bermudensis Erséus & Lasserre,
1976
Figs. 6, 12-15; Table 1
Grania macrochaeta bermudensis Erséus &

Lasserre, 1976:122—124, fig. 3; Lasserre
& Erséus 1976:453.

610

CO

Fig. 11.
funnel, vas deferens and penial apparatus. B, photo of live specimen showing stylet orientation within vasa
deferentia and bipartite penial apparatus. Scale equals 20 wm. bu, union of two sections of bipartite penial
apparatus; vc, vas deferens constrictions; see Figs. 2 & 5 for other symbols.

Type material.—Holotype USNM
53202; paratypes USNM 53203; whole
mounts. Collected by P. Lasserre, 12 Nov
LOS:

Type locality.—Castle Island, Bermuda,
32°20'40"N, 64°40'25”W. Coarse coral sand
with ripple marks, 8 m depth.

Other material examined.—Live speci-
mens from the southeastern tip of Paget Is-
land, Bermuda, collected March 1998 by
JML. Eleven whole mounted specimens
from the collections of C. Erséus collected

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Grania hylae, new species, from Bermuda. A, lateral view of male duct system showing sperm

from two sites in Bermuda, including the
type locality (Table 1).
Description.—External characteristics:
Length of preserved specimens 5.8—10.1
mm (n = 7, X = 7.8, SD = 1.4). Number
of segments 31-59 (n = 7, X = 51.4, SD
= 9.4). Diameter at 4/5 138.0—216.0 um (n
= 6, X = 180.7, SD ="27;5); diameter at
clitellum 159.0-258.9 um (n = 7, X =
217.9, SD = 36.4) (Fig. 12). Ventral setae
from IV, lateral setae from XVI-XX (n =
7). No setae in XII of mature specimens.

VOLUME 112, NUMBER 3

Ventral setae at IV—V 48.0-—72.0 pm (n =
14, X = 59.2, SD = 7.2); length of all other
ventral setae 69.0-92.2 um (n = 32, X =
80.6, SD = 5.9); postclitellar lateral setae
75.0-96.0 um (n = 19, X = 83.4, SD =
6.3) (Fig. 13). In ten posterior-most seg-
ments ventral setae 68.9-74.4 um (n = 6,
X = 71.5, SD = 1.8) and lateral setae 64.7—
75.6 wm (n = 6, X = 70.3, SD = 3.5). Setae
thin with blunt proximal part possessing an
upturned toe, no distinct heel present (Fig.
13). Head pore in anterior part of peristom-
ium. Spermathecal pores paired, dorsal just
posterior to 4/5, with thickened, glandular,
epidermal papilla posterior to each pore
(Fig. 12). Dorsal spermathecal pores about
60.0—64.4 wm apart (n = 2, = 621. SD
= 3.1). Clitellum extends from posterior XI
to mid XIII. Male pores ventral, in line of
setae, at mid XII. Female pores slightly
more lateral than male pores, just posterior
to 12/13.

Internal somatic characteristics: Brain in-
dented posteriorly. Anterior brain vesicle
lacking. Pharyngeal glands present in IV—
VI: with one pair of dorsolateral lobes on
anterior faces of each of 4/5 through 6/7;
ventrolateral lobes, one small pair in IV,
two pairs in V and three pairs in VI (Fig.
12). Nephridia not observed in live speci-
mens, however, in fixed material five to
seven pairs of complete nephridia were ob-
served in segments posterior to egg sac,
about four pairs in posterior-most segments
and about two to three pairs near egg sac;
small overall, with small funnels. Dorsal
blood vessel transition to blood sinus as far
back as XX. One specimen with small
gland on ventral nerve cord in XIV.

Internal reproductive characteristics:
Spermathecae paired in V with narrow ental
connection of ampulla to esophagus (Figs.
12 & 14). Ampulla large and round, walls
very grainy, maximum diameter 42.9—
106.5 um @ = 13, X = 80.7, SD = 23.5);
sperm rings present (Fig. 14). Glands lack-
ing at spermathecal ectal pore. Spermathe-
cal ectal duct diameter wider closest to am-
pulla, diameter ranges from 10.5-19.5 wm

611

G1 2X =449,.SD = 3.0); Ental duct
runs ventrolateral to reach wall of gut,
width about one half ampullar width.
Sperm funnel length about four times
width, non-staining, aglandular region pos-
terior to collar (Fig. 15A). Vas deferens ex-
tending into XIII, thick and muscular along
about one-third of length. Vas deferens rel-
atively wide at sperm funnel, but lacking
muscle on walls; then progressing to a short
section with a spiral of muscle bands, first
wound tightly around circumference of vas
then unwinding so that the thick muscle
bands run almost along length of the vas
(Fig. 15B); muscular section followed by a
thin-walled and narrow section, lacking
muscle, which extends to a pad-like glan-
dular junction with the penial apparatus
(Fig. 15A, C). Maximum diameter of lon-
gitudinally muscled part is 18.0—25.5 wm
Gy 82x = 02 OSD =' 2:6) (is. 15A, B),
non-muscular ectal and ental vas deferens
diameter 7.5—18.0 pm (n = 9, X = 12.3,
SD = 3.2). Penial stylet present, extends
within ectal end of vas deferens causing
that portion to be curved (Fig. 15A, C). Sty-
let length 91.8-130.1 pm (n = 6, x=
107.9, SD = 16.8); maximum width at or-
igin (Fig. 15C) within vas deferens 7.3 wm
(one specimen); stylet width along length
30-42 mm (= 3, XK = 3245S) = 0.7).
Bipartite penial apparatus consists of an
erect sac-like extension from the male pore
running to a lateral, anterior-posterior ori-
ented, longitudinally ridged sac lying par-
allel to the body wall; stylet terminates at
pore on pad in posterior, ental end of ridged
sac, and can be protruded through pore and
into penial apparatus (Fig. 15C). Extensions
of muscles from the body wall run to the
erect extension and the lateral sac, and the
whole of the penial apparatus can be evert-
ed through the male pore at the body wall.
Sperm sac extends to XIX. Egg sac extends
to XXI.

Taxonomic remarks.—The only species
of Grania recorded in Bermuda prior to this
study, G. bermudensis, was described as a
subspecies of G. macrochaeta by Erséus &

612 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

sf:

Fig. 12. Grania bermudensis, specimen from Bermuda. Semischematic dorsolateral view of whole mounted
specimen, from prostomium to segment XIV. Scale equals 100 wm. See Figs. 2 & 7 for symbols.

VOLUME 112, NUMBER 3

WAM,

Fig. 13. Grania bermudensis, specimen from Ber-
muda. Various setae. Scale equals 20 pm. Ipc, lateral
postclitellar seta; see Figs. 3 & 8 for other symbols.

Lasserre (1976). The original description of
the new subspecies was brief and indefinite
about some characteristics, which in part
explains the classification of G. bermuden-
sis within G. macrochaeta s. |. The group-

613

ing of four subspecies, G. m. macrochaeta,
G. m. bermudensis, G. m. pusilla and G. m.
trichaeta was recently reevaluated as taxo-
nomically invalid based on morphological
characteristics (Locke & Coates 1998) and
cladistically invalid as it is polyphyletic
(Locke 1999a). Grania pusilla has been ful-
ly redescribed (Locke & Coates 1998). This
redescription of G. bermudensis provides
details of the dorsal spermathecal pore po-
sition and male duct structures which were
omitted from the original description. The
distribution of lateral setae is amended from
previous records of their originating in
X VITI-XIX to XVI-XX.

The dorsal spermathecal pore position,
bipartite penial apparatus and highly mus-
cularized vas deferens separate this species
from G. macrochaeta s. s. (Locke & Coates
1998) and other Bermudian congeners (Fig.
6). Grania hylae does share comparable
modifications of the first two characters
with G. bermudensis, however, G. hylae

Fig. 14. Grania bermudensis, specimen from Bermuda. Lateral view of spermatheca with epidermal papillae.
Scale equals 20 um. ep, epidermal papillae; see Figs. 2, 4 & 10 for other symbols.

614

3
cence |}
MS OR iA BEEBE Ze

A

Pies. 15,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

C

Grania bermudensis, specimen from Bermuda. A, male duct system and penial apparatus. B, thick

muscular area of vas deferens, in slightly compressed live specimen. C, lateral view of bipartite penial apparatus.
Scale equals 20 ym. bu, union of two sections of bipartite penial apparatus; cc, ciliated canal; cv, cellular vas
wall; g, gut; pj, pad-like glandular junction; rs, ridged sac; sm spiral of muscle; see Figs. 2 & 5 for other

symbols.

lacks the extreme muscular modifications of
the vas deferens. Grania bermudensis is
named here as a distinct species.

Other species with dorsal spermathecal
pores are considered in the taxonomic re-
marks following the description of G. hylae
(above) and in the Discussion.

Habitat and distribution.—Subtidal, me-
dium to coarse coral sand and gravel. Not
found intertidally. Found in samples with
Tubificidae. Known only from Castle Is-
land, East Castle Roads, Rushy Island, a
patch reef 100 m south of Rushy Island and
SE Paget Island, Bermuda (Table 1).

VOLUME 112, NUMBER 3

Grania americana Kennedy, 1966
Figs. 6, 16—20; Table 1

Grania americana Kennedy, 1966:404—
405, fig 3; Erséus 1974:90—93, table 1.
Grania macrochaeta americana: Lasserre

1967:278—280.
Grania americana nomen dubium: Erséus
& Lasserre 1976:123.

Type material.—Holotype, USNM
33005, paratype USNM 33039. Whole
mounts. Collected by M. L. Jones 27 Aug
1962.

Type locality.—0.5 km from North En-
trance point, West side of North Bimini,
Pa N; 79°19.0'W.

Other material examined.—Five
stained, whole mounted specimens, BAMZ
1999 180 O09, Pearl Island, Bermuda;
USNM 185957, Ferry Reach, Bermuda;
USNM 185958 Hutchinson Island, U.S.A.;
and USNM 185959 and USNM 185960,
Carrie Bow Cay, Belize. About 55 live
specimens from Ferry Point Bridge, Ber-
muda, collected by the authors over the pe-
riod 1996-1998. Twenty-four whole
mounted specimens from the collections of
C. Erséus, collected from Bermuda; Carrie
Bow Cay, Belize; Angelfish Key, Florida
Keys, and Hutchinson Island, Florida (Ta-
ble 1).

Description.—External characteristics:
Live specimens notably long, with pink
blood in vessels, posterior segments yel-
lowish to light brown due to colored gran-
ules within chloragogen cells. Prostomium
pointed with anterior thin region; with
many multi-ciliate epidermal papillae.
Fixed length 8.7-12.6 mm (n = 10, X =
10.8, SD = 1.4). Number of segments 60-—
74 (n = 9, X = 64, SD = 4.8). Diameter at
4/5 140.0-196.5 pm (n = 15, X = 168.7,
SD = 18.7); clitellum diameter 150.0—
208.9 wm (n = 15, X = 188.1, SD = 19.1)
(Fig. 16). Setae from IV ventrally and X VI—
XX laterally (n = 13); no setae in XII of
mature specimens. Setae broad at base,
sometimes with slight toe and slight heel in
posterior setae (Fig. 17). Anterior ventral

615

setae 32.6—53.3 um long (n = 10, X = 44.2,
SD = 8.2); midbody ventrals from 53.2—
60.8 wm in length (n = 8, X = 56.3, SD =
2.4). In ten posterior-most segments ventral
setae 56.6—71.3 um long (n = 21, X = 63.4,
SD = 3.7) and lateral setae 49.2—64.1 um
long (n = 16, X = 55.6, SD = 4.2). Head
pore at anterior of peristomium. Spermathe-
cal pores paired, lateral, posterior to 4/5
(Fig. 16). Clitellum extending over XII-—
XIII. Male pores in line with ventral setae,
at mid XII. Female pores at 12/13, slightly
more lateral than male pores.

Internal somatic characteristics: Brain in-
dented posteriorly. Anterio-medial brain
vesicle present (Figs. 16 & 18). Divided
into distinct lateral halves, each containing
two inclusions (for a total of four) (Fig. 18).
Inclusions in each compartment located at
different levels, maximum diameter of in-
elusiens;2-5-2. 8 pm) (n= 16, X= 2:7, SD
= 0.1) (Fig. 18). Total length across both
compartments of head organ 20.5 wm (one
specimen). Coelomocytes flattened and
ovoid. Chloragogen cells, especially in pos-
terior segments, very distinctive with pe-
ripherally clumped granules giving the cells
the doughnut appearance of mammalian red
blood cells, granules reddish-brown by
transmitted light in both live and fixed spec-
imens; outer surface of intestine, as a result,
with leopard-spot pattern. Pharyngeal
glands (Figs. 16 & 19) with one pair of
small ventrolateral lobes in IV and a pair of
dorsolateral lobes at 4/5; two pairs of ven-
trolateral lobes in V and a dorsolateral pair
at 5/6; and with three pairs of ventrolateral
lobes in VI and a dorsolateral pair at 6/7.
Nephridia observed only in posterior seg-
ments, often unpaired, with small elongate
funnel. Solitary, internalized, setae present
in very few segments. Dorsal blood vessel
transition around XXVI-XXVIII. Glands
on nerve cord at XIV and/or XV.

Internal reproductive characteristics:
Spermathecae (Figs. 16 & 19) paired in V
with ental duct connecting ampulla to
esophagus. Ampulla oviform with a maxi-
mum diameter of 42.0—90.0 wm (n = 51, X

616 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

pr
ab

pa

Fig. 16. Grania americana, specimen from Bermuda. Semischematic ventral twisted to lateral view of whole
mounted specimen from prostomium to segment XIV. Scale equals 100 wm. eg; ectal gland of spermatheca; see
Figs. 2 & 9 for other symbols.

VOLUME 112, NUMBER 3

Fig. 17. Grania americana, specimen from Ber-
muda. Setae from various segments. Scale equals 20
ym. h, setal heel; vp, ventral posterior seta; see Figs.
3 & 8 for other symbols.

617

= 68.5, SD = 11.8). Spermathecal ectal
duct with two glands at pore, one small and
one large (Fig. 19). Ectal duct of sperma-
theca consists of two distinct parts: shorter
part immediately adjacent to body wall with
external glands and with wide canal; ental
or proximal part with canal occluded by
glandular cells (Fig. 19) that run as longi-
tudinal segments around the narrow canal.
Spermathecal ectal duct distal width 8.7—
145 im Gp =6.X = 10:8; SD = 2.1); ectal
duct medial width (max) 28.5—83.7 wm (n
— Se — 42 SD — 1 7-4) proximal por-
tion of ectal duct 21.2—46.0 wm in width (n
= 21, X = 34.0, SD = 6.7). Spermathecal
ectal duct length 54.0-94.7 um (n = 16, X
= 74.3, SD = 15.7). Sperm rings present in
walls of ampulla (Fig. 19). Sperm funnel
length about eight times width, often dis-
placed from XI to XII or XIII (Figs. 16 &
20A). Sperm funnel with a non-staining
aglandular region immediately posterior to
the collar; more posteriorly glandular tissue
loose, giving funnel a very irregularly lobed
surface (Fig. 20A). Sperm heads straight.
Vas deferens long, extending through XIII,
non-muscular, 6.8—11.4 wm wide (n = 8, X

Fig. 18.

Grania americana, specimen from Bermuda. Dorsal view of anterior brain vesicle showing inclu-
sions. Scale equals 20 wm. See Figs. 2 & 9 for symbols.

618

Fie. Lo:
muda. Dorsolateral view of spermatheca with glands
at ectal pore. Scale equals 20 ym. See Figs. 4 & 10
for symbols.

Grania americana, specimen from Ber-

= 9.6, SD = 1.7). No stylet. Penial appa-
ratus relatively simple (Figs. 16, 20A, B),
with a distinct bulbous gland and an elon-
gate, saccate dorsal extension from the male
pore which extends along the lateral side of
the gland (Fig. 20A, B); vas enters gland
from dorsomedial side and runs to the lat-
eral side, extending to the elongate exten-
sion. Ventrally, near epidermal pore, the pe-
nial apparatus is narrower with little glan-
dular mass and is surrounded by very
strong elaborations of the muscle bands of
the body wall (Fig. 20B). Sperm sac ex-
tends to XXI. Egg sac extends to XXVII.
Taxonomic remarks.—Kennedy’s (1966)
original description of G. americana differs
in a number of respects from our descrip-
tion. This is probably due to his shortage of
good specimens, which limited collection
of both morphometric and structural data.
Kennedy did provide approximate measure-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ments for the overall length of the species
(10 mm), the number of segments (50), and
the ratio of length to width of the sperm
funnels (several times longer than wide).
However, the description was based on two
specimens only. These were missing a few
segments from their posterior ends and
were eventually mounted in a macerating
medium (pers. obs.).

The original description of G. americana
states that the dorsal setae begin at XVI or
XVIII (Kennedy 1966). In our redescrip-
tion, the ‘“‘dorsal’’ setae are described as lat-
eral and the range of segments at which
they begin is extended to XVI-XX. The
presence of pharyngeal glands in VII (Ken-
nedy 1966, Erséus 1974) was not seen in
the type material when it was re-examined
by us. The peptonephridia originally noted
for G. americana have since been deter-
mined to be absent from the genus Grania
(Erséus & Lasserre 1976) and what Ken-
nedy observed, most likely, were the thick
bundles of ducts running from the pharyn-
geal glands to the dorsal pharyngeal pad.

In more recent literature the taxonomic
importance of the structure of the penial ap-
paratus (Coates 1984) has become apparent,
and therefore, G. americana which had a
brief subspecies status, is not a subspecies
of G. macrochaeta (sensu Lasserre, 1967).
The penial apparatus of G. americana has
a distinct bulbous gland, an elongate, sac-
cate, epidermal invagination and is lacking
a penial stylet. Grania americana also has
ectal glands on the spermathecal ectal duct
at the pore and these glands are visible in
the type material of G. americana. Erséus
& Lasserre (1976) had stated that these
were absent in Grania even though they in-
cluded G. americana. Similar glands have
been noted in other recently described spe-

cies (Coates & Erséus 1980, Erséus 1980,

Rota & Erséus 1997) and the generic de-
scription amended accordingly.

Grania americana differs from other
Bermudian congeners by the possession of
glands at the spermathecal ectal pore, lack
of penial stylet and muscular vas deferens

VOLUME 112, NUMBER 3

619

Pg

60S

ae Stra N wy aan a :

ye

OULU

ovare 1 00)9 sw NO aA

p

Fig. 20. Grania americana, specimen from Bermuda. A, dorsal view of male duct system and penial ap-
paratus. B, lateral view of penial apparatus. Scale equals 20 pm. See Figs. 2 & 5 for symbols.

(Fig. 6). In addition, the length of G. amer-
icana is much greater than that of the other
species described from Bermuda. Grania
americana possesses an anterior brain ves-
icle with four inclusions, described as the
‘head organ” in other species of Grania by
Rota & Erséus (1996, 1997). Grania hylae
which is also found in Bermuda has a dif-
ferent form of anterior brain vesicle with
only one visible inclusion. In G. americana
this organ is located in the anterior of the

peristomium, just below the boundary with
the prostomium, as an anterior, medial bul-
bous extension of the brain. Previous ac-
counts of this structure in recent species de-
scriptions (Rota & Erséus 1996, 1997) have
also noted the presence of four refractile
bodies, but these authors (1996) also noted
that some damaged specimens showed few-
er inclusions or none. Quite possibly this
organ has a geotactic function (Locke
1999b). A head organ has only been re-

620

ported for eight subtidal species of Grania:
five from the Ross Sea, Antarctica, two
from South Georgia in the subantarctic, and
one from the western Atlantic. It is possible
that G. americana shares a unique apomor-
phy with this subgroup of Grania, but is an
intertidal representative of the lineage.

Grania americana differs from all of the
other species of Grania which are known
to have head organs by its spermathecal
form, in particular large glands at the ectal
pore, and distinctive ectal duct structure,
penial apparatus form, and setal distribu-
tion. Two species of Grania are known to
have single, small glands at the spermathe-
cal ectal pore, G. monochaeta (Michaelsen,
1888; amended Rota & Erséus 1997) an At-
lantic species and G. paucispina (Eisen,
1904). The first does not have a head organ
with refractile inclusions, although a vesicle
is present, and for the second no head organ
has been described.

Habitat and distribution.—Coarse inter-
tidal sand within dissolution tubes at Ferry
Point Bridge and rocky limestone beach ar-
eas at Whalebone Bay, Ferry Reach, and
Rocky Hill Park, Bermuda. In Bermuda
also recorded from Pearl Island and Bai-
ley’s Bay in fine to medium subtidal sands.
Found in samples with G. laxartus, various
Tubificidae and Nematoda. First reported
from Bimini, Bahamas; Panama City and
Miami, Florida, U.S.A. Habitat at type lo-
cation in Bimini was probably subtidal as
Kennedy (1966) recorded that the speci-
mens were collected “‘off the coast’’. In
Florida, collected from coarse sand in Thal-
assia beds, and in shell rubble (Table 1). At
Carrie Bow Cay, Belize, also found in me-
dium to coarse sands and Thalassia beds
(Table 1).

Discussion

This is the first addition to our knowl-
edge of the species diversity within Grania
for the Northwestern Atlantic since the
study by Coates & Erséus (1985). Many
other species of the North Atlantic remain

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

to be described (pers. obs.) and further stud-
ies are required to develop an understand-
ing of the speciation patterns and evolution
of this genus. An illustrated dichotomous
key to the species of Grania of the North-
west Atlantic can be found on the world
wide web at http://www. bbsr.edu/users/
kcoates/Grania/Grania.html (Locke 1999a).

The distributions of Grania species
found in Bermuda pose some interesting
questions about speciation and dispersal
patterns, particularly because of the rela-
tively young geological age of Bermuda,
52-34 MY. Grania americana, G. laxartus
and G. hylae are all known from the Ca-
ribbean and Bermuda. This distribution
pattern is considered typical for Bermudi-
an fauna (Sterrer 1986, 1998). The absence
of a larval stage in the development of
these worms makes it difficult to explain
how regular genetic exchange between
Bermudian and coastal populations might
be maintained. Genetic studies of Bermu-
dian and Caribbean coastal populations
could reveal information about effective
‘“‘population’’ sizes, frequency of coloni-
zation events, and whether any existing ge-
netic flow is unidirectional, as would seem
likely due to prevailing currents. Morpho-
logical measurements for conspecific spec-
imens from different localities did not in-
dicate any notable differences between
populations.

The intertidal pools formed within dis-
solution tubes at Ferry Point Bridge create
an intertidal refuge for G. laxartus and G.
americana which is rich in detrital matter.
Healy (1996) noted the importance of crev-
ices within the rocky intertidal as moist mi-
cro-climates during low tide. The dissolu-
tion tubes fulfill the same role as Healy’s
rock crevices in that they protect from wave
disturbance, displacement by water currents
and retain organic matter, but have high ox-
ygen levels (Healy 1996), conditions in
which enchytraeids thrive (Healy & Coates
1999). These small pots or tubes may also
effectively isolate adjacent populations as
the rock surface between is very exposed

VOLUME 112, NUMBER 3

and barren. This is in contrast to the sub-
tidal habitat occupied by G. bermudensis
and G. hylae. These species are protected
from wave action by the greater depth of
the water in which they are found and the
habitat is relatively continuous. The subti-
dal sites in Bermuda were located to the
side of major channels into bays and har-
bours. Tidal currents through these open-
ings bring nutrients and detrital matter to
the areas, and these get deposited in the
slower water and back eddies that border
the main channel.

The presence of G. laxartus and G. amer-
icana within the same samples throughout
the year could conflict with commonly held
ideas about ecological divergence between
congeneric species. There is some evidence,
other than morphological discontinuities, to
suggest that co-occurring populations of the
two species are biologically isolated by
temporal separation of reproductive peri-
ods. High numbers of mature specimens of
each species were found at different times
of the year.

The muscular, bipartite penial apparatus-
es found in G. bermudensis and G. hylae
are different from the many types of glan-
dular, bulb-like apparatuses previously de-
scribed for Grania species (Coates 1984).
This form of penial apparatus appears to be
associated with species which possess dor-
sal spermathecal pores. It also is found in
G. monospermatheca which possesses a
single dorsally displaced spermathecal pore.
Coates (1984) had previously designated a
penial bulb ““Type 6” as glandular penial
bulbs reduced or absent, aglandular sac of-
ten absent; with a long penial stylet plus
considerable modifications to other parts of
the vasa deferentia. Representative taxa
were G. bermudensis and G. monosperma-
theca. This grouping included more struc-
tural diversity than the other “‘types’’ rec-
ognized. Additional information from the
species described herein is considered and
Coates (1984) “‘Type 6” is revised. Penial
bulb ““Type 6” is now described as: bipar-
tite, saccate penial apparatus lacking a dis-

621

tinct glandular bulb; saccate part with hor-
izontal and erect parts. Stylet long and con-
tained within vasa deferentia. Muscular
modifications of the vasa deferentia. Sper-
mathecal pores dorsal. Representative taxa:
G. bermudensis, G. monospermatheca, G.
hylae, and two undescribed species from
Australia (Coates 1984). Documentation of
dorsal spermathecal pores and complex pe-
nial apparatuses in G. stilifera Erséus, 1990
and G. inermis Erséus, 1990 present the
possibility of their being included within
the Type 6 grouping. Whereas, Erséus
(1990a) designated these as ““Type 5”’ and
“Type 3” (sensu Coates 1984), respective-
ly, penial apparatuses. Other forms origi-
nally “‘lumped”’ in Type 6 await further
clarification and typification.

The presence of an anterior brain vesicle
in G. americana and G. hylae are the first
records in readily collected species. This
structure, first reported as the “‘head organ”’
by Rota & Erséus (1996), has only been
found in one other species from the North
Atlantic, G. atlantica (Rota & Erséus 1996;
pers. obs). The function of the anterior
brain vesicle is still unknown, but it appears
to be statocyst-like in structure (Locke
1999a, 1999b). Phylogenetic studies of
Grania species which possess this structure
may provide ecological and functional hy-
potheses underlying the origin of such a
structure in clitellates.

Although Erséus & Lasserre (1976) stat-
ed their understanding of the morphology
of Grania had improved, they still believed
that morphological differences between
widely separated populations were small,
sufficient only to justify their recognition as
subspecies. Nonetheless, they (Erséus &
Lassetrre 1976:131) stated that with further
studies the reinstatement of their subspecies
to species could easily be made and Erséus
(1977) noted that the structural variation
within G. macrochaeta s. |. was very broad.
Erséus & Lasserre (1976) also noted the im-
portance of spermathecae in the taxonomy
of enchytraeids. Locke & Coates (1998)
resurrected G. pusilla from subspecies sta-

622

tus. Herein, G. bermudensis is recognized
as even more distinct and G. americana is
redeemed from nomen dubium (Lasserre &
Erséus 1976) to full species status. The dif-
ferences in spermathecal structures and in
the male duct systems (Coates 1984) among
these species provides sufficient support for
the recognition of all the subspecies of G.
macrochaeta as morphologically distinct
species. Even more damning for G. macro-
chaeta s. |. is that this grouping of subspe-
cies is clearly polyphyletic. Only one taxon
in this group, Grania macrochaeta trichae-
ta Jamieson, 1977, awaits a full redescrip-
tion.

Acknowledgments

BBSR NSF-Research Experience for Un-
dergraduate student, Mr. Brandon Falk, for
his initial studies of Bermuda species; and
Christer Erséus for the loan of specimens.
The Canadian Associates of the Bermuda
Biological Station and National Science
Foundation grants #DEB-9615211 and
#DBI 9724335 for support and funding for
research facilities. Invertebrate Zoology
Section, CBCB, Royal Ontario Museum for
the use of the videomicroscope and speci-
mens from the accessioned collections of
KAC. Ms. Hyla Tibbitts for her detailed il-
lustrations and The Rhode Island School of
Design Internship program. Hank Trapido-
Rosenthal (BBSR) for logistic and financial
support. Appreciation to Pat Pocklington
for our joint subtidal collection trip and Ja-
mey Penney-Ritter for her graphical skills.
This is contribution number 1500 of the
Bermuda Biological Station for Research,
Inc.; contribution number 25 of the Ber-
muda Biodiversity Project (BBP), Bermuda
Aquarium, Natural History Museum and
Zoo; and contribution number 567 of the
Caribbean Coral Reef Ecosystems Program,
National Museum of Natural History,
Smithsonian Institution, deriving from the
collections made by C. Erséus.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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. 1990b. The marine Tubificidae (Oligochaeta)

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840.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(3):624—631. 1999.

Maractis rimicarivora, a new genus and species of sea anemone
(Cnidaria: Anthozoa: Actiniaria: Actinostolidae) from an Atlantic
hydrothermal vent

Daphne G. Fautin and Brian R. Barber

Division of Biological Sciences and KU Natural History Museum, University of Kansas,
Lawrence, Kansas 66045, U.S.A.

Abstract.—Maractis rimicarivora is a new genus and new species of me-
dium-sized sea anemone (Actiniaria) from the TAG (Trans-Atlantic Geotra-
verse) hydrothermal vent fields (26°08.3'N, 44°49.6’W; 3650 m). The genus,
which belongs to family Actinostolidae, is distinguished by the following com-
bination of features: six pairs of complete mesenteries, mesenteries not arrayed
according to the Actinostola rule, all mesenteries gametogenic, tentacles not
thickened at the base, and no tentacular mastigophores. The species is distin-
guished by long, longitudinally-furrowed tentacles, and a roughened column.
Anemones of this species encircle black smokers at distances from 15 to 60
m. This is the second species to be reported from hydrothermal vents of the
Atlantic. Four species have been reported from Pacific hydrothermal vents.

Five species of sea anemones have been
documented from hydrothermal vents (Des-
bruyeres & Segonzac 1997). Cyananthea
hydrothermala Doumenc & Van-Praét,
1988, and Marianactis bythios Fautin &
Hessler, 1989, were described from Pacific
vents. Doumenc & Van-Praét (1988) also
identified from Pacific vents Actinostola
callosa (Verrill, 1882) and Chondrophellia
coronata (Verrill, 1883), species described
from trawled specimens before the exis-
tence of hydrothermal vents was known.
The only sea anemone thus far reported
from Atlantic vents is Parasicyonis ingolfi
Carlgren, 1942 (Segonzac 1992), another
species described from trawled specimens.
Certainly three and probably four of these
five species belong to family Actinostoli-
dae, one of the two richest families of deep-
sea anemones (the other is Hormathiidae, to
which C. coronata belongs).

We describe another member of Actino-
stolidae as a new genus and new species
from the TAG (Trans-Atlantic Geotraverse)
vent fields, located at 26°08.3’'N,
44°49.6'W, and 3650 m. Part of the Mid-

Atlantic Ridge (MAR), TAG is one of the
largest submarine hydrothermal fields
known (German et al. 1995, Van Dover
1998). We infer this to be the species Van
Dover et al. (1997) reported preying on a
shrimp of the species Rimicaris exoculata
at the TAG site, and that is depicted in Fig.
3b of Van Dover (1995). Shrimp (along
with fish and crabs) dominate high-temper-
ature regions of the TAG vent field near
active black smokers, and anemones occu-
py peripheral regions where the water is
cooler (Van Dover 1995), attached to the
crumbly substratum of oxidized sulfide.
Materials and methods.—The five spec-
imens we studied were collected on 21 Sep-
tember 1994 using the claw of the sub-
mersible Mir 1. Within an hour of being
brought to the surface, they were fixed in
10% seawater formalin; later they were
transferred to isopropanol. Morphological
aspects of this description are based on
these preserved specimens; biological in-
formation is from the published literature
and from P. A. Tyler, who provided the
specimens to us. One well-expanded spec-

VOLUME 112, NUMBER 3

imen, which we designated the holotype,
was missing a few tentacles, but was oth-
erwise in excellent condition. The four par-
atypic specimens were missing many ten-
tacles and patches of ectoderm. Three of
them were in good condition otherwise and
were moderately expanded; the other was
moderately contracted and its partly-everted
actinopharynx was damaged.

Preparation of histological sections and
the gathering of cnidae data followed meth-
ods of Fautin & Hessler (1989). Histologi-
cal sections 8 pm thick were stained with
hematoxylin and eosin. Cnidae were mea-
sured in smash preparations at 1000.

Maractis, new genus

Definition.—Maractis has the following
combination of diagnostic characters: mem-
ber of family Actinostolidae having six
pairs of complete mesenteries; mesenteries
not arrayed according to the Actinostola
rule; all mesenteries gametogenic; tentacles
not thickened at the base; and no tentacular
mastigophores. Further defining features in-
clude a broad base, at least 48 tapered ten-
tacles, and column ectoderm that lacks pa-
pillae or other such structures. Animals of
this genus are supple, unlike many deep-sea
anemones, because the mesoglea is rela-
tively thin.

Differential diagnosis.—In the key to
some genera of Actinostolidae of Fautin &
Hessler (1989), Maractis falls under op-
tions B), ff), hh), jjj), and kk) by virtue of
having “All or all stronger mesenteries fer-
tile ...,’ “Longitudinal tentacle muscles
ectodermal,’ “‘No microbasic b-mastigo-
phores in tentacles,” “‘No tentacular mas-
tigophores,”’ and “‘Six pairs of mesenteries
perfect,’ respectively.

Three genera fall under that option in the
key: Bathydactylus, Cnidanthea, and Epi-
paractis. Bathydactylus, like Maractis, has
a broad base. However, Bathydactylus has
a very strong marginal sphincter muscle
that forms a projecting wall, stout tentacles
that are few in number, mesenteries that are

625

not hexamerously arrayed, and papillae in
the distal part of the column (at least in
some species). Cnidanthea, like Maractis,
has hexamerously arrayed mesenteries, but,
unlike Maractis, has papillose nematocyst
batteries on its column. Epiparactis, like
Maractis, has more than 48 tentacles that
are closely packed at the rim, and has broad
pedal and oral discs. However, it has a
smooth column and “thick, cartilaginous
mesogloea”’ (Carlgren 1921, page 198).

The other genera of actinostolid sea
anemones that have been reported from hy-
drothermal vents are Marianactis, Parasi-
cyonis, and Actinostola; Cyananthea prob-
ably also belongs to this family. Maractis
is similar to Marianactis Fautin & Hessler,
1989, in lacking microbasic b-mastigo-
phores in the tentacles, but differs in lack-
ing microbasic amastigophores in the ten-
tacles. Parasicyonis Carlgren, 1921, and
Actinostola Verrill, 1883, differ from Mar-
actis both externally (both have short ten-
tacles) and internally (in the former, mes-
enteries of only the last cycle are fertile,
whereas in the latter those of the first two
cycles are sterile). Cyananthea is incom-
pletely known, but has twice as many mes-
enteries at the margin as at mid-column, in
contrast to Maractis.

Etymology and gender.—Maractis is a
composite of MAR (Mid-Atlantic Ridge)
and “‘actis,’’ Greek for ray or beam, a term
often applied to anemones. Its gender is
feminine.

Type species.—Maractis rimicarivora,
new species.

Maractis rimicarivora, new species
Figs. 1—4

Description.—Pedal disc flat, margin
scalloped because pulled inward at mesen-
terial insertions (Fig. 1), which are visible
through disc. Diameter in animals exam-
ined 25—55 mm. Color in preservation same
as that of column, oral disc, and tentacles—
uniformly dull pink.

Column diameter 20—30 mm, length 10—

626

} |

| j

TET TT
i | bi di
/ pti

1 | 2
CENTIMETERS

Riga:

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Maractis rimicarivora: portion of holotype (KUNHM 01149). Note scalloped edge of pedal disc and

long, longitudinally-furrowed tentacles. Raised circular spots are inferred to be artifacts—impressions of the

perforations in a collecting device or storage container.

20 mm. P. A. Tyler (pers. comm.) remarked
in life “‘the column looks very short’’ but
was difficult to see because animals “al-
ways appear to have the [tentacle] crown
[directed] to the camera.”’

Mesenterial insertions visible through
body wall where ectoderm is missing. Ec-
toderm thin, eroded from many specimens.
Surface texture rough: mesoglea and ecto-
derm thrown into short, relatively deep,
complex folds (Figs. 2A, 3).

Oral disc torn or obscure in specimens
examined. No fossa: outermost tentacles
arise from margin (Fig. 2A). Tentacles 5—
25 mm long (longest roughly equal to or
slightly longer than column height). Flac-
cid, flattened in preserved specimens, lon-
gitudinally furrowed, tapering toward distal
end (Fig. 1); number to nearly 100 in spec-
imens examined. Cannot be withdrawn and
covered when animal retracted. Longitudi-
nal muscles ectodermal, continuous with ra-
dial muscles of oral disc (Fig. 2A).

Mesenteries regularly arrayed in 4 cycles,
not according to the Actinostola rule; those
of fourth cycle may not reach margin. Two
pairs of directives, each attached to a si-
phonoglyph, lie diametrically opposite one
another. Only mesenteries of first cycle com-
plete. All fertile Gncluding directives and
those of highest order) (Fig. 3); sexes sepa-
rate. No stomata. Retractor muscles well-de-
veloped, diffuse, arrayed in bolster-like
strands (Fig. 3). Parietobasilar muscles de-
veloped only along stronger mesenteries;
broad at base; each extends about half length
of column, has very narrow free edge.

Marginal sphincter muscle mesogleal,
moderately well developed. May fill entire
width of mesoglea at margin but for most
of length does not extend to ectodermal
edge of mesoglea (Fig. 2B); at proximal
end, occupies about half width of mesoglea,
lying along endoderm (Fig. 2A, B). Small
alveoli of equal size scattered along ecto-
dermal side of sphincter; may be reticulate

VOLUME 112, NUMBER 3

on endodermal side (Fig. 2A, B). Muscle
cells relatively denser in larger individuals.
Musculature weakly transversely stratified
in parts of muscle of some individuals.

Actinopharynx about half length of col-
umn.

Animals, including tentacles, may remain
fully extended upon preservation.

Cnidom.—spirocysts, basitrichs, micro-
basic p-mastigophores, holotrichs.

Size and distribution of cnidae.—Letters
refer to illustrations in Fig. 4. Measure-
ments in parentheses are of single capsules
that fell outside the usual range. Number of
capsules measured is indicated by “‘n’’; ra-
tio of the number of animals in which a
particular type of cnida was found to the
total number examined for that tissue is in-
dicated by #.

Tentacles—distal end:

Spirocysts (A, B) 30.0-78.5 X n= 91
2.8—7.3 (7.8) wm # = 5/5
Basitrichs (C) (15.6, 15.9) n= 54
17.2—25.5 X 1.6—2.9 um # = 5/5
Basitrichs (D) 31.0-48.4 n= 50
(54.6) X 2.5—3.8 wm # = 5/5
Holotrichs (E) 30.4-53.9 X n= 39
4.5—6.4 wm # = 5/5
Tentacles—proximal end:
Spirocysts (A, B) 30.3-64.1 XK n= 85
2.9-6.9 (7.7, 8.1) wm #= 5/5
Basitrichs (F) (16.8) 19.2—- n= 74
43.4 X 1.9-3.8 um # = 5/5
Column:
Basitrichs (F) 19.5-31.5 X n=7I1
1.9-3.9 (4.9) pm # = 5/5
Microbasic p-mastigophores n = 54
(G) 21.3-31.9 X 3.4-6.2 um # = 4/5
Actinopharynx:
Basitrichs (F) 23.4-34.9 Xx n=9
2.3-3.2 pm # = 3/4
Microbasic p-mastigophores n = 31
(G) 28.6—-42.1 xX 4.1-5.2 #= 3/4
(5.7) wm
Mesenterial filaments:
Basitrichs (C) 16.1-34.1 n= 6l

(39.3, 45.1) X 1.8-4.5 um #= 5/5

627

Microbasic p-mastigophores n= 61
(G) 26.9-38.9 (43.1) Xk #= 5/5
3.3—-5.7 (6.0) wm

Most spirocysts could be assigned to one of
two types (both shown in Fig. 4A), al-
though rare ones shared characteristics of
both. This distinction was not noticed until
many measurements had been gathered, so
our data are not divided by type. Spirocysts
that were, on average, longer and wider had
tubules wound in an untidy fashion and
lacked space between the tubule and the
capsule’s posterior end (Fig. 4A, at left).
Spirocysts that were, on average, shorter
and narrower had neatly wound tubules and
space between the tubule and the posterior
end of the capsule (Fig. 4A, at right, and
Fig. 4B). For example, in tentacles of par-
atype KUNHM 01150, 9 of the “‘neat’’ spi-
rocysts measured 36.0—49.9 X 3.3—5.3 um,
and 10 of the “untidy’® ones measured
37.3-59.2 X 5.1-7.7 wm.

Etymology.—The name rimicarivora re-
fers to the ingestion (Latin vora: to eat) by
this anemone of the shrimp Rimicaris exo-
culata Williams & Rona, 1986. [The tax-
onomy of vent shrimps is considered by
Shank et al. (1998).]

Type locality.—TAG (Trans-Atlantic
Geotraverse) hydrothermal vent fields:
26°08.3'N, 44°49.6'W; 3650 m.

Type specimens.—Holotype (Fig. 1) Di-
vision of Invertebrate Zoology, University of
Kansas Natural History Museum (KUNHM)
01149: a dissected male specimen and 16
microscope slides made from it.

Paratypes: KUNHM 01150: a dissected
specimen of indeterminate sex and 20 mi-
croscope slides made from it. The Natural
History Museum, London 1999.80: a dis-
sected male specimen and 9 microscope
slides made from it. The Santa Barbara Mu-
seum of Natural History, Santa Barbara,
California 145140: a dissected female spec-
imen and 10 microscope slides made from
it. The U.S. National Museum of Natural
History 100132: a dissected specimen of in-

628

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fion2:
KUNHM 01150. Tentacles have ectodermal longitudinal musculature; outermost tentacle arises at margin. The

surface of the animal is thrown into short, complex folds. B: Proximal end of sphincter muscle of holotype
(KUNHM 01149).

determinate sex and 5 microscope slides
made from it.

Discussion

The holotype (KUNHM 01149), a quar-
ter of which is illustrated in Fig. 1, has con-
spicuous raised circular patches on part of
its column, especially the lower portion. We
initially thought these might be adhesive
structures to which sulfide particles could
adhere (see below). We concluded, how-
ever, that they are artifacts from a collecting
device or storage container. Their position
does not correspond to any anatomical fea-
ture of the animal, they are on only one side
of the holotype, and they are not visible in
the other specimens (although the ectoderm
is eroded from several of them, patches per-
Sist).

Mesogleal sphincter muscle of Maractis rimicarivora. A: Longitudinal section at margin of paratype

We, like Van Dover et al. (1997), found
no evidence of bacterial symbionts in Mar-
actis rimicarivora: there were neither bod-
ies that could be interpreted as bacteria nor
structures of the animal that were likely to
harbor bacteria. In addition, Van Dover et
al. (1997) found no biochemical or physi-
ological evidence of bacterial symbiosis.
We infer, therefore, that M. rimicarivora is
a typical cnidarian in depending on prey for
energy and nutrients. Shrimp and anemones
dominate the biota of TAG and most other
Atlantic vent fields that have been studied,
in contrast to Pacific vent fields, which are
dominated by tubeworms, mussels, and
clams (Van Dover 1995). At TAG, there ap-
pears to be little aside from shrimp on
which the anemones could feed (P. A. Tyler,
pers. comm.). The observation of predation

VOLUME 112, NUMBER 3

Biz.,32

Cross-section at mid-column level of the holotype (KUNHM 01149). Spermaries associated with
mesenteries of the third and fourth (highest) orders are visible. Well-developed retractor muscles are diffuse.
Scale bar = 1 mm.

C D E F G

Fig. 4. Cnidae of Maractis rimicarivora. See text for explanation. Scale bar = 9 wm.

aeeetttt |.
»

ne th be
J 2
4 sae"

~

A B

630

on a shrimp (Van Dover et al. 1997) is con-
sistent with these data and inferences.

The size and color of the predatory
anemone, and the length of its tentacles,
were consistent with our specimens. How-
ever, our specimens lacked the black spots
on the column that are conspicuous in the
videotape on which the report of Van Dover
et al. (1997) is based. Van Dover (pers.
comm.) suggested they were sulfide parti-
cles. Although the specimens we studied
lacked discrete adhesive structures, the
large surface area of their wrinkled ecto-
derm (Figs. 2, 3) would seem likely to have
adhesive properties. Indeed, some fine de-
bris adhered to one specimen we studied.
We therefore concluded the animal that is
the subject of the report by Van Dover et
al. (1997) is M. rimicarivora. Van Dover et
al. (1997) found the anemone most abun-
dantly 30—40 m from the center of a black
smoker, where its density reached nearly
200 m~’; its average density in the ring it

formed around the black smoker at a dis-
tance of about 15—60 m was 3 m~*, so the
anemones probably do not experience water
of elevated temperature.

Van Dover (1995) inferred there might be
two species of anemones at TAG, one small
and one large. One small anemone speci-
men we examined had a short, strongly re-
ticulate mesogleal sphincter muscle, which
was like that of neither M. rimicarivora nor
Paractinostola ingolfi, the anemone col-
lected at the Snake Pit hydrothermal vent
field 3° south of TAG (Segonzac 1992).
Therefore, there may be at least three spe-
cies of anemones at TAG. Anemones also
occur at the Broken Spur vent field 3° north
of TAG (Van Dover 1995), but not at the
Lucky Strike vent field 11° north of TAG
(Van Dover et al. 1996).

Acknowledgments

We are grateful to Paul A. Tyler for spec-
imens of anemones from TAG and for a
copy of the videotape showing an individ-
ual of Maractis rimicarivora preying on a

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

shrimp. We thank Tracy White for help with
histology and Adorian Ardelean for tech-
nical assistance. Cindy L. Van Dover and
Paul A. Tyler provided important informa-
tion and helpful comments. Cadet Hand
commented on an earlier version of this
manuscript. Stanley E Lombardo helped
with the Latin. This research was supported
by NSF grant DEB95-21819 (PEET) to
Daphne Fautin. Collections were made un-
der the auspices of NERC Grant BRIDGE
21 to Paul A. Tyler.

Literature Cited

Carlgren, O. 1921. Actiniaria. I—The Danish Ingolf-
Expedition 5(9):1—241.

. 1942. Actiniaria. Il—The Danish Ingolf-Ex-
pedition 5(12):1—92.

Desbruyéres, D., & M. Segonzac, eds. 1991. Hand-
book of deep-sea hydrothermal vent fauna. Edi-
tions IFREMER, Brest, France, 279 pp.

Doumenc, D., & M. Van-Praét. 1988. Actinies abys-
sales d’un site hydrothermal du Pacifique ori-
ental.—Oceanologica Acta special volume 8:
61-68.

Fautin, D. G., & R. R. Hessler. 1989. Marianactis by-
thios, a new genus and species of actinostolid
sea anemone (Coelenterata: Actiniaria) from the
Mariana vents.—Proceedings of the Biological
Society of Washington 102:815—825.

German, C. R., E. T. Baker, & G. Klinkhammer. 1995.
Regional setting of hydrothermal activity. Pp.3—
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on, eds., Hydrothermal vents and processes.
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Segonzac, M. 1992. Les peuplements associés a
Vhydrothermalisme océanique du Snake Pit
(dorsale médio-atlantique; 23°N, 3480 m): com-
position et microdistribution de la mégafau- -
ne.—Comptes Rendus Hebdomadaires des Sé-
ances de l’Académie des Sciences, Paris, series
III 314:593—600.

Shank, T. M, R. A. Lutz, & R. C. Vrijenhoek. 1998.
Molecular systematics of shrimp (Decapoda:
Bresiliidae) from deep-sea hydrothermal vents,
I: Enigmatic “‘small orange” shrimp from the
Mid-Atlantic Ridge are juvenile Rimicaris ex-
oculata.—Molecular Marine Biology and Bio-
technology 7(2):88—96.

Van Dover, C. L. 1995. Ecology of Mid-Atlantic Ridge
hydrothermal vents. Pp. 257—294 in L. M. Par-
son, C. L. Walker, & D. R. Dixon, eds., Hydro-
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Saldanha, A. Fiala-Médioni, & C. Langmuir.
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field.—Deep-Sea Research I 43(9):1509-1529.
, M. Polz, J. Robinson, C. Cavanaugh, D. Kad-
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at TAG.—BRIDGE Newsletter 12:33-—34.
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631

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. 1883. Report on the Anthozoa, and on some
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
112(3):632—633. 1999.

INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE
Applications published in the Bulletin of Zoological Nomenclature

The following Applications were published on 18 December 1998 in Vol. 55, Part 4
and on 30 June 1999 in Vol. 56, Part 2 of the Bulletin of Zoological Nomenclature.
Comment or advice on any of these applications is invited for publication in the Bulletin
and should be sent to the Executive Secretary (I.C.Z.N.), % The Natural History Museum,
Cromwell Road, London SW7 S5BD, U.K. (e-mail: iczn@nhm.ac.uk).

Case No.

3036 Haliotis clathrata Reeve, 1846 (non Lichtenstein, 1794) and H. elegans Philip-
pi, 1844 (Mollusca, Gastropoda): proposed conservation of the spe-
cific names.

3080 Polydora websteri Hartman in Loosanoff & Engle, 1943 (Annelida, Poly-
chaeta): proposed conservation of the specific name by a ruling that
it is not to be treated as a replacement for P. caeca Webster, 1879,
and designation of a lectotype for P. websteri.

Spherillo Dana, 1852 (Crustacea, Isopoda): proposed designation of S. vitiensis
Dana, 1853 as the type species, with designation of a neotype.

Terebratula Miiller, 1776 (Brachiopoda): proposed designation of Anomia ter-
ebratula Linnaeus, 1758 as the type species.

Coluber infernalis Blainville, 1835 and Eutaenia sirtalis tetrataenia Cope in
Yarrow, 1875 (currently Thamnophis sirtalis infernalis and T. s. te-
trataenia; Reptilia, Squamata): proposed conservation of the subspe-
cific names by the designation of a neotype for T. s. infernalis.

Crotalus ruber Cope, 1892 (Reptilia, Serpentes): proposed precedence of the
specific name over that of Crotalus exsul Garman, 1884.

Arctocephalus EF Cuvier, 1826 and Callorhinus Gray, 1859 (Mammalia, Pinnipe-
dia): proposed conservaton by the designation of Phoca pusilla Schreber,
[1775] as the type species of Arctocephalus; and Otaria Péron, 1816 and
Eumetopias Gill, 1866: proposed conservation by the designation of Pho-
ca leonina Molina, 1782 as the type species of Otaria.

Bulinus wrighti Mandahl-Barth, 1965 (Mollusca, Gastropoda): Proposed con-
servation of the specific name.

Sphaerius Waltl, 1838 and SPHAERIUSIDAE Erichson, 1845 (Insecta, Coleoptera):
proposed conservation by the partial revocation of Opinion 1331.

Blennocampa Hartig, 1837, Cryptocampus Hartig, 1837, Taxonus Hartig, 1837,
Ametastegia A. Costa, 1882, Endelomyia Ashmead, 1898, Monsoma
MacGillivray, 1908, Gemmura E.L. Smith, 1968, BLENNOCAMPINI Ko-
now, 1890 and CALIROINI Benson, 1938 (Insecta, Hymenoptera): pro-
posed conservation by setting aside the type species designations by
Gimmerthal (1847) and recognition of those by Rohwer (1911).

Macrophya Dahlbom, 1835 (Insecta, Hymenoptera): proposed designation of
Tenthredo montana Scopoli, 1763 as the type species; and Tenthredo
rustica Linnaeus, 1758: proposed conservation of the usage of the
specific name by the replacement of the syntypes with a neotype.

Apis proava Menge, 1856 (currently Electrapis proava; Insecta, Hymenoptera):
proposed conservation by designation of a neotype.

VOLUME 112, NUMBER 3 633

Opinions published in the Bulletin of Zoological Nomenclature

The following Opinions were published on 18 December 1998 in Vol. 55, Part 4
and on 30 June 1999 in Vol. 56, Part 2 of the Bulletin of Zoological Nomenclature.
Copies of these Opinions can be obtained free of charge from the Executive Sec-
retary, I.C.Z.N., % The Natural History Museum, Cromwell Road, London SW7
SBD, U.K. (e-mail: iczn@nhm.ac.uk).

Opinion No.

1910. Roeslerstammia Zeller, 1839 and Acrolepiopsis Gaedike, 1970 (Insecta, Lep-
idoptera): conserved by the designation of Alucita erxlebella Fabri-
cius, 1787 as the type species of Roeslerstammia; and A. erxlebella
and Tinea imella Hubner, [1813] (currently Roeslerstammia erxle-
bella and Monopis imella): specific names conserved by the desig-
nation of a neotype for A. erxlebella.

Dasineura Rondani, 1840 (Insecta, Diptera): Tipula sisymbrii Schrank, 1803
designated as the type species.

Pseudofoenus Kieffer, 1902 (Insecta, Hymenoptera): Foenus unguiculatus
Westwood, 1841 designated as the type species.

Trachelocerca Ehrenberg (Ciliophora): authorship conserved as Ehrenberg
(1840), and Vibrio sagitta Miiller, 1786 fixed as the type species.

Helix draparnaudi Beck, 1837 (currently Oxychilus draparnaudi; Mollusca,
Gastropoda): specific name conserved.

Turrilites gravesianus d’Orbigny, 1842 (currently Hypoturrilites gravesi-
anus; Mollusca, Ammonoidea): specific name conserved and a re-
placement lectotype designated; Turrilites tuberculatus Bosc, [1802]
(currently Hypoturrilites tuberculatus): placed on the Official List.

DASYPODIDAE Borner, 1919 (Insecta, Hymenoptera): spelling emended to DA-
SYPODAIDAE, So removing the homonymy with DASYPODIDAE Gray,
1821 (Mammalia, Xenarthra).

Lactura Walker, 1854 (Insecta, Lepidoptera): conserved, and the specific
name of Eustixis pupula Hubner, [1831] (currently Lactura pupula):
conserved.

Waagenoconcha Chao, 1927 and Gruntoconcha Angiolini, 1995 (Brachio-
poda): conserved.

Cnemidophorus neomexicanus Lowe & Zweifel, 1952 (Reptilia, Squamata):
specific name conserved.

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REVIEWERS

The following people reviewed manuscripts for the Proceedings in 1998. S. Ahyong, C. C. Baldwin, R. C.
Banks, W. C. Banta, R. Bauer, R. W. Baumann, E M. Bayer, J. A. Blake, J. P. Blanco-Rambla, G. A. Boxshall,
A. Brandt, R. Brinkhurst, B. Brown, N. L. Bruce, G. Brusca, D. R. Calder, D. K. Camp, M. D. Carleton, M.
Carvalho, FE A. Chase, Jr., A. H. Cheetham, C. M. Cleveland, K. A. Coates, C. O. Coleman, P. Cornelius, F B.
Crandall, N. Cumberlidge, E D’Incao, M. Dillon, M. Elias-Gutierrez, D. Erwin, D. G. Fautin, D. L. Felder, E
D. Ferrari, J. FE Fitzpatrick, Jr., K. Fitzhugh, O. S. Flint, Jr., C. H. J. M. Fransen, S. L. Gardiner, A. L. Gardner,
O. Gon, N. Gotelli, G. R. Graves, M. J. Grygier, D. Guinot, C. Hand, E. Hatfield, B. Healy, R. W. Heard, Jr.,
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Kathman, B. Kensley, W. Kim, N. Knowlton, W. Kobusch, R. K. Kropp, J. H. Leal, D. B. Lellinger, R. Lemaitre,
B. J. MacFadden, R. B. Manning, W. N. Mathis, J. Mauchline, J. McCosker, R. W. McDiarmid, J. McEachran,
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R. Nelson, J. Nelson, P K. L. Ng, W. Ngoc-Ho, J. Norenburg, S. Ohtsuka, G. Pohle, W. W. Price, J. Pruski, C.
E. Ray, N. A. Rayner, Y. R. Reddy, J. W. Reid, J. V. Remsen, M. E. Retzer, R. P- Reynolds, K. Riemann-
Ziirneck, R. Rios, C. B. Robbins, C. R. Robins, G. Rodriguez, R. Roman-Contreras, D. A. Rossman, M. C.
Sanz, W. B. Saunders, J. M. Savage, S. Schaefer, R. B. Searles, E. K. Shea, D. Smarzly, J. P. S. Smith, I, H.
Squires, K. C. Stuck, H. Suzuki, M. Tavares, C. A. Taylor, E G. Thompson, C. Tudge, J. C. Tyler, S. Tyler, R.
Vargas, V. S. Vassilenko, J.-W. Waegele, X. Wang, S. H. Weitzman, M. J. Wetzel, J. T. Williams, K. Wittman,
M. Wolsan, K. Wouters, E. L. Yochelson, G. R. Zug.

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112(3):735-736. 1999.

BIOLOGICAL SOCIETY OF WASHINGTON

126th Annual Meeting, 20 May 1999

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The Society continues to make modest fi-
nancial gains from the sale of back issues
of the Proceedings and from interest on So-
ciety accounts.

Editor Brian Robbins then reported that
Volume 111 of the Proceedings comprised
87 papers totaling 1008 pages, up from 70
and 692, respectively, in Volume 110. As
of 1 May, there were 37 submissions for
Volume 112. There is no backlog for ac-
cepted papers. The Guidelines for submis-
sions that was printed in 1990 as a Supple-
ment to issue no. 1 will be revised and re-
printed; Robbins solicited input from the
Society regarding modifications or addi-
tions that should be incorporated.

Custodian of Publications, Storrs Olson,

then provided an update on his progress of
sorting the Society’s stock of unbound Pro-
ceedings separates from Volumes 6-89.
Separates are being arranged by taxonomic
category for future sale as sets. Olson asked
that a committee be established to deal with
issues relating to pricing, advertising, and
marketing of the sets when complete. Pres-
ident Vari so agreed, and it was decided af-
ter much discussion that this same commit-
tee should look into the fate of the Society’s
stock of bound Proceedings issues and con-
sider a proposal that the number of extra
copies of each volume printed be reduced
from about 100 to 50.

Before calling for the Report from the
Finance Committee, President Vari an-
nounced that Richard Banks had recently
resigned from the Finance Committee and
has been replaced by Bruce Collette. Be-
cause of a work-related absence, however,
Collette was not able to participate in the
preparation of the 1998 Finance Committee
report, and Don E. Wilson joined Finance
Committee member Austin Williams in this
endeavor. Williams reported that accounting
of the Society’s funds by the Treasurer is in
good order and that continued attention will
be given to establishing a meaningful an-
nual budget for the Society, a task made
difficult by the unpredictability of number
of Proceedings pages published, amount of
page charges recovered from authors, etc.
Williams also noted that the Society’s con-
tract with Allen Press will expire soon and
recommended that other publishers be con-
sidered along with Allen Press in efforts to
achieve an expedient new contract for the
Society.

During the President’s call for New Busi-
ness, Storrs Olson suggested that the Soci-
ety consider revamping the cover of the

636 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Summary financial statement for 1998.

General Endowment Total

fund fund assets
ASSETS: JANUARY 1, 1997 14,722.89 O1-557 53 106,280.22
TOTAL RECEIPTS FOR 1997 98,904.60 12 554.27 111,458.87
TOTAL DISBURSEMENTS FOR 1997 88,121.22 —.— $8,121.22
ASSETS: DECEMBER 31, 1997 25,506.27 104,111.60* 129,617.87
NET CHANGES IN FUNDS 10,783.38 1255457 23,331.65

* The income from publication inventory ($3,797.50) and Merrill Lynch income ($8,756.77) are the income
for the Endowment Fund.

Proceedings. Much discussion ensued re- and Storrs Olson and Austin Williams were
garding possible cover designs, and Vari recognized for recent contributions.

asked Editor Robbins to investigate with President Vari adjourned the meeting at
Allen Press the cost of various options pro- 12:15 p.m.
posed.

Chad Walter and Brian Robbins were Respectfully submitted,
commended for their diligent efforts and Carole C. Baldwin

longstanding dedication to Society affairs, Secretary

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CONTENTS

Cytogeography of Philippine bats (Mammalia: Chiroptera)
Eric A. Rickart, Jennifer A. Mercier, and Lawrence R. Heaney
On the valid name of the lesser New Zealand short-tailed bat (Mammalia: Chiroptera)
Gregory C. Mayer, John A. W. Kirsch, James M. Hutcheon,
Francois-Joseph Lapointe, and Jacinthe Gingras
Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 8. A provisional hypothesis for the

hybrid origin of Zodalia glyceria (Gould, 1858) Gary R. Graves
A new species of pelican (Aves: Pelecanidae) from the Lower Pliocene of North Carolina and
Florida Storrs L. Olson
A new species of Platymantis (Amphibia: Ranidae) from the Sierra Madre Mountains, Luzon
Island, Philippines W. C. Brown, A. C. Alcala, P. S. Ong, and A. C. Diesmos
Two new species of the Eleutherodactylus rugulosus group (Amphibia: Anura: Leptodactyli-
dae) from Honduras James R. McCranie and Larry David Wilson

Taxonomic status and geographic distribution of Bryconamericus eigenmanni Evermann &
Kendall, 1906 (Characiformes: Characidae)
Amalia M. Miquelarena and Adriana E. Aquino
Caridina clinata, a new species of freshwater shrimp (Crustacea: Decapoda: Atyidae) from
northern Vietnam Yixiong Cai, Nguyen Xan Quynh, and Peter K. L. Ng
Inclusion of the austral species Pinnotheres politus (Smith, 1869) and P. garthi Fenucci, 1975
within the genus Calyptraeotheres Campos, 1990 (Crustacea: Brachyura: Pinnotheridae)
Ernesto Campos
Description of Synalpheus williamsi, a new species of sponge-dwelling shrimp (Crustacea:
Decapoda: Alpheidae), with remarks on its first larval stage
Rubén Rios and J. Emmett Duffy
Two new freshwater crabs of the genus Ptychophallus Smalley, 1964 (Crustacea: Decapoda:
Brachyura: Pseudothelphusidae) from Panama, with notes on the distribution of the genus
Martha R. Campos and Rafael Lemaitre
A new species of Chevalia (Crustacea: Amphipoda: Corophiidae) from the Indian Ocean with
remarks on Chevalia carpenteri and the C. aviculae superspecies complex
Eric A. Lazo-Wasem
Megagidiella azul, a new genus and species of cavernicolous amphipod crustacean (Bogidiel-
lidae) from Brazil, with remarks on its biogeographic and phylogenetic relationships
Stefan Koenemann and John R. Holsinger
The western and southern distribution of Mesocyclops edax (S. A. Forbes) (Crustacea: Cope-
poda: Cyclopoida) Janet W. Reid and Luis Moreno D.
A new species of Orbiniella (Orbiniidae: Polychaeta) from Marion Island, Indian Ocean
Patrick Gillet
Redescriptions of Grania americana, G. bermudensis and descriptions of two new species of
Grania (Annelida: Clitellata: Enchytraeidae) from Bermuda
J. M. Locke and K. A. Coates
Maractis rimicarivora, a new genus and species of sea anemone (Cnidaria: Anthozoa: Acti-
niaria: Actinostolidae) from an Atlantic hydrothermal vent
Daphne G. Fautin and Brian R. Barber
International Commission on Zoological Nomenclature
Reviewers, 1998
Biological Society of Washington: 126th Annual Meeting

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BIOLOGICAL SOCIETY

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By é

VOLUME 112 NUMBER 4
29 DECEMBER 1999

ISSN 0006-324X

THE BIOLOGICAL SOCIETY OF WASHINGTON
1998-1999
Officers

President: Richard P. Vari Secretary: Carole C. Baldwin
President-elect: Brian F. Kensley Treasurer: T. Chad Walter

Elected Council

Michael D. Carleton Rafael Lemaitre
W. Duane Hope Roy W. McDiarmid
Susan L. Jewett James N. Norris

Custodian of Publications: Storrs L. Olson

PROCEEDINGS

Editor: C. Brian Robbins

Associate Editors

Classical Languages: Frederick M. Bayer Invertebrates: Stephen L. Gardiner
Plants: David B. Lellinger Frank D. Ferrari
Insects: Wayne N. Mathis Rafael Lemaitre

Vertebrates: Gary R. Graves

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(4):637—651. 1999.

Abyssal sea anemones (Cnidaria: Actiniaria) of the northeast Pacific
symbiotic with molluscs: Anthosactis nomados, a new species, and
Monactis vestita (Gravier, 1918)

Tracy R. White, April K. Wakefield Pagels, and Daphne G. Fautin

(TRW) Department of Ecology and Evolutionary Biology, and Division of Invertebrate Zoology,
Natural History Museum and Biodiversity Research Center, University of Kansas,
Lawrence, Kansas 66045, U.S.A.; (AKWP) Department of Systematics and Ecology, and
Division of Invertebrate Zoology, Natural History Museum and Biodiversity Research Center,
University of Kansas, Lawrence, Kansas 66045, U.S.A.;

(DGF) Department of Ecology and Evolutionary Biology, and Division of Invertebrate Zoology,

Natural History Museum and Biodiversity Research Center, University of Kansas,
Lawrence, Kansas 66045, U.S.A., and
Kansas Geological Survey, 1930 Constant Avenue, Lawrence, Kansas 66047, U.S.A.

Abstract.—We describe Anthosactis nomados, new species, which belongs
to family Actinostolidae, and redescribe Monactis, vestita (Gravier, 1918), a
species belonging to family Hormathiidae. Anemones of both species live at-
tached to molluscs on the soft-sediment abyssal plain of the northeastern Pacific
Ocean. Anthosactis nomados is known from 260 specimens, most of which
were still attached to scaphopod shells when studied, and the others of which
show evidence of having been attached to one. By altering the definition of
genus Anthosactis to include all tentacles being of equal length, this species
can be accommodated in it. The species also differs from the other six species
previously ascribed to the genus in the size of its nematocysts. We examined
about 600 specimens of Monactis vestita, more than 350 of which show evi-
dence of having been attached to a gastropod shell but none of which was still
associated with the host when studied. This is the first report of Monactis
vestita in such a symbiosis and from the Pacific Ocean. We hypothesize that
both anemone-mollusc symbioses are mutualistic. In that between Anthosactis
nomados and the scaphopod, which is the third anemone-scaphopod symbiosis
to be documented, the protection provided by the sea anemone may allow the
scaphopod to avoid the corrosive environment of sub-surface sediments and to
forage at the surface where it can obtain energy and calcium carbonate. For
both symbioses, the mollusc may resuspend sediments and carry the actinian
to food-rich areas, and the anemone may protect its host from predators.

Specimens of the two species of sea
anemones we studied were collected as part
of long-term research on the biota of the
abyssal northeastern Pacific. Specimens of
both Monactis vestita (Gravier, 1918) and a
new species we describe below were col-
lected by A. G. Carey, Jr. from stations cen-
tered around 45°N, 135°W, approximately
1200 km off the coast of Oregon, at 3700—

3900 m (Table 1). Many more specimens of
the new species were collected by K. L.
Smith, Jr. at a site referred to as Station M
by us and Smith (Smith et al. 1994, Lauer-
man et al. 1996) and as Station N by Rei-
mers et al. (1992), 220 km off the coast of
California, at 34°50’N, 123°00’W, and 4100
m (fig. 1 in Lauerman et al. 1996). We ex-
amined 161 specimens of the new species

638

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Data for specimens of Monactis vestita (M. v.) and Anthosactis nomados, new species (A. n.)
collected off the Oregon coast. Shape of Monactis vestita specimens are coded as follows: d = dome-shaped, f
= flat, co = wrapped around cylindrical object. Shapes are illustrated in Fig. 2. Museum abbreviations are

provided in the text.

Catalog number Station number Coordinates Depth Number of specimens
SBMNH 144459 BMT 233 44°40'48’N, 133°26'18”"W 3717 m 8d, 10f, lco M. v.
SBMNH 144422 BMT 303 45°05'06"N, 133°10'54”"W 3700 m Od, 1f, ico May:
SBMNH 345404 . ‘i : 43 A. n.

SBMNH 144424 BMT 305 45°05'12’"N, 134°43’24"W 3900 m 20d, 4f, 15co M. v.
SBMNH 144423 BMT 306 45°02'00’N, 134°42'12”"W 3900 m 67d, 83f, 3co M. v.
RBCM 999-262-1 q ‘, ‘ 2d, 1f M. v.
SBMNH 345405 as ¥ 45 A. n.

SBMNH 144461 BMT 307 45°03'30"N, 134°45'00”W 3900 m 52d, 18f, lco M. v.
USNM 100314 g \ 2d, 2f, lee Nity:
SBMNH 345406 4 a 3 ALS a:

SBMNH 144419 BMT 308 45°01'06"N, 135°13'36"W 3932 m 173d, 65f, 10co M. v.
CAS 119154 : ‘ * 3d, 2f, lco M. v.
KUNHM 01210 % i : 8d, 1f, 2co M. v.
SBMNH 345407 x ‘ % YASH

SBMNH 144460 BMT 309 45°02'00"N, 135°23'12”W 3990 m 63d, 5f, 6co M. v.

LACM 1972-386.001 4 f
SBMNH 345408

from 36 collections made at Station M from
1989 to 1995. No specimens of M. vestita
were collected at Station M. At least four
specimens from Station M_ superficially
looked like M. vestita, but differed signifi-
cantly in characteristics of their nemato-
cysts.

Each specimen of the new species ex-
amined from Station M was attached to the
concave surface of a shell of the scaphopod
Fissidentalium actiniophorum Shimek,
1997. Sixty-two percent of scaphopods we
examined carried a sea anemone and anoth-
er 30% of the shells showed evidence of
recent anemone attachment—a brown ma-
terial secreted by the anemone’s pedal disk.
This is the third documented sea anemone-
scaphopod symbiosis. As part of the rede-
scription of Hormathia pectinata, Riemann-
Ziirneck (1973) mentioned that some spec-
imens collected in 1970-1971 from Sta-
tions 251 (53°51’S, 59°54'’W, 535 m) and
ZIV (52°40'S20.60°39 W, 405 “m)- of “the
Walther Herwig Expedition were attached
to a Dentalium shell. Shimek & Moreno
(1996) reported the symbiosis between Fis-
sidentalium megathyris and the species we

‘ 2d. 2& Se0KVax:
fs PA ce

describe below. In addition, Zibrowius
(1998) reported a symbiosis in the north-
western Pacific between the solitary scler-
actinian coral Heterocyathus japonicus
(Verrill, 1866) and the scaphopod Fissiden-
talium vernedei (Sowerby, 1860), and Carl-
gren (1928b) found the anemone Paracal-
liactis stephensoni Carlgren, 1928b, on a
large Dentalium shell inhabited by the her-
mit crab Parapagurus pilosimanus Smith,
1879, off the Irish coast.

Specimens of both species we studied
from the Oregon sites had been removed
from their substrata. However, we were giv-
en a photograph (Fig. 1) of a specimen of
M. vestita attached to the shell of Buccinum
sp. (probably B. strigillatum Dall, 1891;
Henry Chaney, Santa Barbara Natural His-
tory Museum, pers. comm.). We do not
know whether M. vestita associates with
more than one species of snail, but we infer
from variation in concavity of the pedal
disk that M. vestita attaches to substrata
other than snail shells (Fig. 2). For instance,
the pedal disk of some specimens had been
wrapped around a cylindrical object and a

VOLUME 112, NUMBER 4

METRIC 1 2
TO

Fig. 1. Rounded specimen of Monactis vestita on
Buccinum, probably B. strigillatum.

flat specimen (Fig. 3) was likely attached to
a rock.

Paractis vestita Gravier, 1918 was de-
scribed from 14 specimens collected during
the voyages of the Hirondelle and the Prin-
cess-Alice from 1888 to 1913. Twelve spec-
imens were collected in 1895 at Station 749
(38°54'00"N, 21°06'45”W, 5005 m) and two
specimens were collected in 1905 at Station
2044 (32°28'00"N, 16°37'30"W, 2286 m).
Eleven of the 12 syntypes from Station 749
bear catalog number 13 0022 and the two
syntypes from Station 2044 bear catalog
number 13 0080 in the Institut Océanogra-
phique of Monaco. Gravier’s (1918) de-
scription was accurate except that he found
no acontia. He therefore placed the species
in family Paractidae, which is no longer
valid. Recognizing acontia in 208 speci-
mens she studied from six stations in the
Venezuela deep-sea basin (3476-5060 m)
and three stations off the Atlantic coast of
Spain (4706-5320 m), Riemann-Ziirneck
(1986) created the genus Monactis for this
species, which she placed in family Hor-
mathiidae. The specimens Riemann-Ziir-
neck (1986) studied were poorly preserved,
so she could not provide information on in-
ternal anatomy or sources of cnidae. In
1992, Zamponi & Acufia described the re-
productive structures of three specimens of
M. vestita collected from 200-250 m near

639

Mar del Plata, Argentina. Thus, M. vestita
has been collected from only the Atlantic
Ocean; we extend the range of M. vestita to
the Pacific Ocean.

We infer these symbioses are mutualistic.
Both species of sea anemones have in-
creased access to food as they are carried
across the sea floor by the mollusc. The sca-
phopod or snail, when covered by an ex-
panded sea anemone, receives protection
from predators. Scaphopods typically live
buried within the sediment, but the symbi-
osis allows the scaphopod to live at the sur-
face, where the sediment contains more cal-
cium carbonate and energy (Smith et al.
1994). We initially hypothesized that the
anemone also protects the aragonitic sca-
phopod shell from dissolution in this high-
pressure, low-temperature environment
where the water is undersaturated with re-
spect to calcium carbonate (Edmond &
Gieskes 1970, Berger 1976, Grotsch et al.
1991), but this hypothesis was not borne
out.

Materials and Methods

Immediately after being trawled, some
specimens of the new species from Califor-
nia and their host scaphopods were fixed in
10% buffered formalin and were later trans-
ferred to 70% EtOH; others were preserved
immediately in 70% EtOH. Specimens of
M. vestita were fixed at sea in buffered 10%
formalin and later transferred to 70% EtOH.

Histological study was done of 41 spec-
imens of the new species from California
and 13 from Oregon. Five were embedded
whole; the others were cut in half longitu-
dinally through the center of the body, and
the half on the larger-diameter (anterior)
end of the scaphopod shell was removed for
histology. Histological study was done of
16 specimens of M. vestita. Tissue was em-
bedded in Paraplast® (melting point 56°C);
8 wm longitudinal sections or 10—20 wm
cross sections were stained with hematox-
ylin and eosin (Humason 1979). Egg di-
mensions are reported as the average of the

640

a

en?
Pid

Saal Fahne tc

Fig. 2.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Specimens of Monactis vestita (voucher: KUNHM 01210) demonstrating variety of shapes. (d) Dome-

shaped specimens. (f) Flat specimen. (co) Specimens wrapped around cylindrical objects. Note cuticle on smaller

specimen labeled d.

maximum diameter and the largest perpen-
dicular diameter in one section of each egg
with a visible nucleolus.

Tissue for cnidae squashes was taken
from mesenterial filaments, acontia (in M.
vestita), actinopharynx, tentacle tips, and
oral and aboral sides of tentacle bases. Cni-
dae were studied with differential interfer-
ence contrast optics; measurements are of

undischarged capsules in squash prepara-
tions.

Anthosactis nomados, new species
Figs. 4—8

Column.—Freshly collected and pre-
served specimens white. All specimens
contracted. Column smooth; ectoderm

VOLUME 112, NUMBER 4

crac Wy 2

641

i : * :

tf

i] Caaenanne
HENBaR

Fig. 3.

Fig. 4. Longitudinal section of Anthosactis noma-
dos, new species. s = sphincter muscle, t = tentacle,
g = gametes. Note that column ectoderm and pedal
disk are absent (holotype: KUNHM 01019).

Contracted specimens of Monactis vestita attached to rocks.

sloughed off in collected specimens (Fig.
4), a common condition in sea anemones
collected from the deep sea (Carlgren
1928a, 1956; Fautin & Hessler 1989; Rie-
mann-Ziirneck 1993, 1994).

Sphincter.—Mesogleal. Thick at margin,
without external evidence of projecting
wall. Typically long: tapered at proximal
end (Fig. 5A) but truncated in some (Fig.
5B). Commonly centered in mesoglea but
typically closer to the endodermal side at
proximal end; closer to ectodermal side in
rare specimens. Muscle bundles dense and
arranged transversely in most, scattered in
others; reticulate in nature along endoder-
mal side of some. Variability in appearance
may be related to state of contraction or an-
gle of section.

Pedal disk.—Broad; well-developed; so
tightly adherent to shell of scaphopod Fis-
sidentalium actiniophorum (Fig. 6) that it
typically remains attached to shell when sea
anemone is removed (Fig. 4). Maximum di-
ameter never greater than length of scapho-
pod shell, ranging from 5 to 65 mm; me-
dian 44 mm. Midway along scaphopod
Shell, pedal disk typically covers 75% cir-
cumference of shell; coverage increases at

642

Fig. 5. Cross sections through mesogleal sphincter
muscle of Anthosactis nomados, new species. Both fig-
ures are oriented so external side of sphincter is up-
ward. (A) voucher: KUNHM 01020; (B) holotype:
KUNHM 01019.

posterior end of shell, decreases at anterior
end. Secretes brown cuticle onto scaphopod
shell.

Oral disk and tentacles.—Oral disk not
visible in specimens examined due to con-
tracted state. Radial musculature of oral
disk and longitudinal musculature of ten-
tacles ectodermal (Fig. 7). Tentacles taper
to a point; not thickened at base. Tentacle
length uniform within an individual; range
from 2 to 5 mm. Ratio of number of ten-
tacles to number of mesenteries variable,
ranging from 0.5:1 to 1:1, mean 0.7:1. Ten-
tacles of preserved animals brittle and eas-
ily damaged or detached, which may ex-
plain variation in tentacle: mesentery ratio.
Due to contracted state and morphology,
number of siphonoglyphs and arrangement
of tentacles could not be determined.

Mesenteries and internal anatomy.—
Mesenteries hexamerously arrayed, in 3 cy-
cles; first 2 cycles, including directives,
complete, fertile, and filament-bearing;
mesenteries in third cycle weak, sterile. Di-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

rective mesenteries aligned with long axis
of scaphopod. Gametogenic tissue easily
damaged or detached so pattern of fertile
mesenteries difficult to determine. More
mesenteries proximally than distally. Tertia-
ry mesenteries develop asynchronously; ei-
ther not all pairs of tertiary mesenteries
present or not all reach distal portion of an-
imal. Retractor muscles diffuse; due to mor-
phology, structure of parietobasilar muscles
could not be determined.

Sexes separate; 6 males, 1 female ob-
served among sectioned specimens. Range
of minimum and maximum diameter of
sperm packets (n = 45) 34 X 36 pm—132
x 175 pm. In large packets, cells in various
stages of spermatogenesis layered, with
spermatozoa central. Diameter of three eggs
51-73 wm.

Cnidom.—Spirocysts, basitrichs, micro-
basic p-mastigophores, and microbasic b-
mastigophores. Sizes and distribution of
cnidae given in Table 2; cnidae illustrated
in Fig. 8. No difference in cnidae on oral
and aboral sides of tentacle base. All indi-
viduals have basitrichs of two size classes
in tentacle tips; most have two size classes
in tentacle bases. Robust spirocysts differ-
entiated from typical ones by their thick and
spiny tubules.

Etymology.—The specific epithet noma-
dos is a Greek word for roving (Brown
1956), and was inspired by the nickname
“crawling white anemone” bestowed on
this sea anemone by Lynn M. L. Lauerman,
then a graduate student at Scripps Institu-
tion of Oceanography. Its gender is femi-
nine.

Specimens

Holotype.—University of Kansas Natural
History Museum (KUNHM), Division of
Invertebrate Zoology 01019, half a male in-
tact on scaphopod shell and longitudinal
histological sections of the other half (5
slides); collected 7 Nov 1993. Type locality
34°42'N, 123°08’W, 4100 m.

Paratypes.—United States National Mu-

VOLUME 112, NUMBER 4

643

f PLE UTR ARR LAL UAL

‘CENTIMETERS

Fig. 6. Contracted specimen of Anthosactis nomados, new species, attached to scaphopod shell. The oral
disk of the sea anemone is approximately centered on the concave surface of the scaphopod shell. In a minority
of the collected specimens, the pedal disk of the anemone is not wrapped tightly around the scaphopod shell

but is bunched as shown in Shimek (1997: fig. 1).

seum of Natural History (USNM), Depart-
ment of Invertebrate Zoology 96574, half a
male intact on scaphopod shell and longi-
tudinal histological sections of the other
half (5 slides); collected 10 Feb 1994 from
34°41'N, 123°11’W at 4100 m. California

Big: 7;
Anthosactis nomados, new species (voucher: KUNHM
01020). White arrow indicates ectodermal longitudinal
muscle. A spirocyst is indicated by the black arrow. c
= ectoderm, m = mesoglea, n = endoderm.

Longitudinal section through tentacle of

Academy of Sciences (CAS), Department
of Invertebrate Zoology and Geology
106264, half a male on scaphopod shell and
longitudinal histological sections of the oth-
er half (5 slides); collected 17 Oct 1992
from 34°46'N, 123°08'’W at 4100 m. Royal
British Columbia Museum (RBCM), De-
partment of Invertebrate Zoology 996-24-1,
half a female removed from its scaphopod
host and longitudinal histological sections
of the other half (5 slides); collected 19 Jul
1993 from 34°43’N, 123°06’W at 4100 m.
RBCM 996-25-1, one specimen intact on
scaphopod shell; collected 1 May 1995
from 34°40'N, 123°03’W at 4100 m. Santa
Barbara Museum of Natural History
(SBMNH), Department of Invertebrate Zo-
ology 143214, half a specimen intact on
scaphopod shell and longitudinal histologi-
cal sections of the other half (5 slides); col-
lected 24 Feb 1993 from 34°45’'N,
123°02'’W at 4100 m. Los Angeles County

644 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Size and distribution of cnidae of Anthosactis nomados, new species (letters refer to Fig. 8). *‘n’’
is the number of capsules measured and ‘‘#’’ is the proportion of animals examined in which that type of cnida
was present. A single capsule that fell considerably outside the range of the others is distinguished by parentheses,
following the convention of Dunn (1982).

Tissue/Cnida type Capsule length (1m) Capsule width (tm)

TENTACLE TIPS

Spirocysts (A) 102 10/10 14.6-44.7 (51.5) 1.8—5.0

Robust spirocysts (B) ‘ke 9/10 (16.7) 20.3-62.9 (66.6) 3.4-9.1 (9.6)

Basitrichs (C) 68 10/10 9.7-19.4 1.5—4.]

Basitrichs (D) 91 10/10 17.8-31.0 (32.8) 2.2—5.8

Microbasic b-mastigophores (F) 43 7/12 18.1—31.2 4.6-6.8
TENTACLE BASES

Spirocysts (A) 101 10/10 14.8-37.8 (42.9) 235.7

Robust spirocysts (B) 114 10/10 (15.1) 18.2-45.6 3.0-10.9 (11.9)

Basitrichs (C) 60 7/10 10.6—17.9 (20.6) 2.3—-4.1

Basitrichs (D) 1S 9/10 18.5—30.9 2.6—5.1
ACTINOPHARYNX

Microbasic p-mastigophores (E) 102 10/10 (30.2) 35.8—50.3 3.4-6.7
MESENTERIAL FILAMENTS

Microbasic p-mastigophores (E) 100 10/10 28.9-44.7 (50.8) (2.8) 3.3-6.6 (7.1)

Basitrichs (C) 102 10/10 = (10.5) 11.3-15.9 (18.8) 2.2—5.4 (6.1)

Museum of Natural History (LACM), De-
partment of Malacology and Invertebrate
Paleontology 92-113.1, half a specimen re-
moved from its scaphopod host and longi-
tudinal histological sections of the other
half (5 slides); collected 17 Oct 1992 from
34°46'N, 123°08’W at 4100 m.

Other material examined.—Sea anemo-
nes and scaphopods collected from August
1989 to October 1991 were borrowed from
LACM. Scaphopods collected during 1992,
from February 1993 to February 1994, dur-
ing August 1994, and during February 1995
are housed at LACM; sea anemones from
these collections are deposited in KUNHM.
Scaphopods and sea anemones collected
during July, September, and October of
1994, and May and June of 1995, are de-
posited in KUNHM.

Taxonomic Issues

Family Actinostolidae currently includes
22 genera (Fautin & Hessler 1989). The
new species conformed in most respects to
the definition of genus Anthosactis. To ac-
commodate this species, we amend Carl-

gren’s (1949) definition of Anthosactis
slightly (change underlined):

Actinostolidae with well developed pedal disc. Col-
umn smooth, rather low, often longitudinally sul-
cated in the contracted state. Sphincter strong, me-
sogloeal. Tentacles short, rather few, conical, robust,
hexamerously or octamerously arranged, the outer
only a little shorter than the inner ones or all ten-
tacles of equal length. Outer tentacles on their ab-
oral sides at the base provided with a stinging bat-
tery of microbasic b-mastigophors, which are some-
times distributed along the whole aboral side of the
tentacle, though more numerous at the base. Lon-
gitudinal muscles of tentacles ectodermal, strongest
on the oral side. Radial muscles of oral disc ecto-
dermal to ecto-mesogloeal. Actinopharynx short
with two well developed siphonoglyphs. Pairs of
perfect mesenteries 6, 6+2 single, 8, or 12. 2 pairs
of directives. Retractors rather weak, parietobasilar
muscles fairly well developed. At least the younger
mesenteries growing from the basal disc upwards.
All stronger mesenteries fertile. Cnidom: spirocysts,
basitrichs, microbasic b- and p-mastigophors.

See Table 3 for the differences between
A. nomados and five of the six species listed
by Carlgren (1949); a description of A.
georgiana has never been published so is a
nomen nudum.

645

VOLUME 112, NUMBER 4

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Height of

Cnidae of Anthosactis nomados, new spe-
Figs. 1-3, 9-11

Monactis vestita (Gravier, 1918)
Column.—Preserved specimens brown;
most contracted. Scapus and scapulus dis-

tinct in extended specimens.

Fie. 8.
cies. A and B are spirocysts; C and D are basitrichs;

E is a microbasic p-mastigophore; F is a microbasic b-
mastigophore. Size and distribution of cnidae are
ered in grainy cuticular material that flakes
off (as reported also by Riemann-Ziirneck
1986). Column of some specimens not en-
tirely covered with cuticular material; may
have been sloughed off during collection
[as is common in sea anemones collected
from the deep sea (Carlgren 1928a, 1956;

shown in Table 2.
specimens 1.5 mm to 10 mm. Column cov-

646

Fig. 9. Cross section through mesogleal sphincter
of Monactis vestita oriented so external side of sphinc-
ter is upward (voucher: SBMNH 144423).

Fautin & Hessler 1989; Riemann-Ziirneck
1993, 1994)]. Foraminiferans attached to
pedal disk of few specimens; Gravier
(1918) reported foraminiferans covering the
column.

Sphincter.—Strong, mesogleal (Fig. 9).
Centered in mesoglea; proximal end tapered
and closer to endodermal side. Alveoli uni-
form in size.

Pedal disk.—Well-developed; so tightly
adherent it usually remains attached to sub-
stratum when anemone is removed. Broad
or narrow depending on substratum. Width
(measured across surface of specimen) of
flatter anemones 14.5 mm to 33 mm; 15
mm to 42 mm in anemones with concave
pedal disks.

Oral disk and tentacles.—Oral disk not
visible due to contracted state. Tentacles
conical; base broad, tapering to pointed tip.
One expanded specimen with 32 tentacles:
16 large inner tentacles and 16 small outer
tentacles.

Mesenteries and internal anatomy.—
Mesenteries hexamerously arranged in 4

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 10. Cross section of Monactis vestita (vouch-
er: SBMNH 144460).

cycles; only those of primary cycle com-
plete. Two distinct siphonoglyphs moder-
ately deep. More mesenteries proximally
than distally. Retractor muscles diffuse,
strong (Fig. 10). Acontia on primary mes-
enteries (as reported by Riemann-Ziirneck
1986), inconspicuous; lighter in color than
mesenterial filaments.

Sexes separate; 4 males, 8 females
among those on which histology was done.
Diameter of 25 eggs with visible nucleolus
53-107 pm.

Cnidom.—Spirocysts, basitrichs, and mi-
crobasic p-mastigophores. Sizes and distri-
bution of cnidae given in Table 4; cnidae
illustrated in Fig. 11.

Specimens

For repositories of voucher specimens,
see Table 1.

Taxonomic Issues

Because Gravier (1918) did not find
acontia in the specimens he studied, he
identified them as Paractis; the new species
differed from known species in number of
complete mesenteries. In 1986, Riemann-
Ziirneck found acontia and established a
new genus for the species now termed Mo-
nactis vestita. Carlgren (1949) listed 15
genera in family Hormathiidae; Monactis
brings the number to sixteen.

VOLUME 112, NUMBER 4

647

Table 4.—Size and distribution of cnidae of Monactis vestita (letters refer to Fig. 11). ‘‘n’’ is the number of
capsules measured and ‘‘#’’ is the proportion of animals examined in which that type of cnida was present.

Tissue/Cnida type n # Capsule length (jm) Capsule width (jm)

TENTACLE TIPS

Spirocysts (A) 105 9/9 21.4—75.7 3.2-8.8

Robust spirocysts (B) 109 I) 21.8-69.3 3.2-11.9

Basitrichs (C) 80 8/9 7.6—21.1 1.4-3.5

Basitrichs (D) 92 9/9 18.6-41.2 2.3-4.4
TENTACLE BASES

Spirocysts (A) 71 TUT 23.8-62.3 3.4-8.8

Robust spirocysts (B) 81 TH 24.1—53.1 4.1-12.6

Basitrichs (C) 62 7/7 9.8—-17.7 1.4-3.3

Basitrichs (D) HA 7/7 19.3—32.7 2.5-4.4

Microbasic p-mastigophores (E) 67 the 16.7—23.4 3.0—5.4
ACTINOPHARYNX

Basitrichs (D) 55 6/6 23.4—-34.9 2.6—3.9

Microbasic p-mastigophores (E) 44 4/4 15.4—25.4 3.2—-5.0

Microbasic p-mastigophores (F) 36 4/4 27.2—32.8 4.3-7.7
MESENTERIAL FILAMENTS

Microbasic p-mastigophores (E) 66 7/7 19.8—25.6 3.4-5.1

Basitrichs (C) 68 7/7 10.6—19.3 2.0-3.1
ACONTIA

Basitrichs (C) 40 4/4 10.9-20.9 1.8—3.2

Basitrichs (D) 40 4/4 27.6—37.7 3.14.6

The pedal disk was missing from most Discussion

specimens we examined; presumably it was
torn off when they were removed from their
substrata. Acontia were found only in spec-
imens with pedal disks that were intact or
present at least at the margins. Gravier’s
(1918) not having found acontia may be ex-
plained if the specimens he studied had torn
pedal disks.

Riemann-Ziirneck (1986) could not de-
termine the source tissue of cnidae except
for acontia in the 208 poorly-preserved
specimens of M. vestita she studied. Of
more than 600 specimens we examined,
many were in fine condition, so we could
ascertain the cnidom of each tissue. Our
data differ in two particulars from those of
Riemann-Ziirneck (1986): we found gracile
spirocysts (in the tentacles) as well as the
robust type found by Riemann-Ziirneck,
and the basitrichs we found fell into two
size classes rather than three.

Sea anemones of both species and their
molluscan hosts from Oregon were separat-
ed by the time we obtained the specimens.
We were therefore unable to ascertain
which species of scaphopod hosted the
specimens of Anthosactis nomados collect-
ed there. Likewise, we could not determine
if Buccinum strigillatum is the only species
of snail that hosts Monactis vestita. Gravier
(1918) and Riemann-Ziirneck (1986) stud-
ied specimens of M. vestita that were at-
tached to objects such as stones. We are the
first to document specimens of M. vestita
attached to a snail as well as the first to
record specimens from the Pacific Ocean.
Perhaps only individuals from the northeast
Pacific Ocean are symbiotic with molluscs.
Despite the existence of other hard surfaces
at Station M and off the Oregon coast, such
as brachiopods, Mn nodules, snails, and
sponge spicules, individuals of the sea

648

Jars JE
spirocysts; C and D are basitrichs; E and F are micro-
basic p-mastigophores. Size and distribution of cnidae
are shown in Table 4.

Cnidae of Monactis vestita. A and B are

anemone A. nomados were found only on
the shells of Fissidentalium actiniophorum.

We infer from the specificity of A. no-
mados that the scaphopod offers the anem-
one a benefit in addition to the hard sub-
stratum for attachment. By contrast, we in-
fer that individuals of M. vestita attach to a
variety of substrata. We hypothesize that A.
nomados and M. vestita gain access to food
by attaching to a mobile animal: food par-
ticles may be resuspended as the mollusc
host moves through the sediment; if the sea
anemone can pick prey off of the sea floor,
it has access to a larger area than it would
if it were stationary; and the mollusc host
presumably can detect and move to food.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Such benefits have been proposed for other
sea anemones that are carried by hermit
crabs or snails (Ross 1971, Dunn et al.
1980, Chintiroglou & Koukouras 1991,
Riemann-Ziirneck 1994).

We infer that the scaphopod and the snail
are protected from predators by their actin-
ian symbionts (e.g., Ross 1971, Brooks
1988) and thus that the symbioses are mu-
tualistic. We further hypothesize that this
protection provided by A. nomados enables
its scaphopod host to forage in the upper
layers of sediment that are rich in energy
relative to sediments at depths in which
such scaphopods typically live. The sedi-
ment at the surface has three times more
ATP (an indicator of microbial activity)
than that at a depth of 35 mm (Smith et al.
1994).

Living at the surface may also be favor-
able to formation and maintenance of the
scaphopod’s aragonitic shell. Scaphopods
prey on foraminiferans (e.g., Shimek 1990,
Langer et al. 1995). There are 1-18 X 10°
m~? calcareous foraminiferans in the sur-
face sediments of Station M (Smith et al.
1994). Foraminiferans may be a source of
calcium carbonate, as well as energy and
nutrients, for the scaphopods (Shimek, pers.
comm.). Very little carbonate carbon occurs
in surface sediments of Station M (Reimers
et al. 1992); absolute values of CaCO, in
the upper 10 mm of sediment range from
5.14 to 16.36 mg (g dry weight)! (Smith
et al. 1994). The difficulty of producing
and/or maintaining a calcareous shell at this
site is reflected by a dearth of shelled or-
ganisms such as gastropods, bivalves, and
brachiopods (Lauerman et al. 1996). The
anoxic sediments below 2.5 to 3 cm (Rei-
mers et al. 1992), the depth at which a sca-
phopod of this size might typically live,
may be expected to be more corrosive to
calcareous organisms than the surface sed-
iments.

We had hypothesized the sea anemone
might insulate the aragonitic shell of a sur-
face-dwelling scaphopod from the sur-
rounding sea water that is undersaturated

VOLUME 112, NUMBER 4

with respect to CaCO,. Geological thin-sec-
tions were made to compare shell thickness
of host scaphopods in areas that had been
covered by a specimen of A. nomados with
areas that had not been covered. Fifteen
scaphopod shells, 12 that had been partially
covered by a sea anemone and three that
showed no evidence of sea anemone attach-
ment, were vacuum-embedded in clear ep-
oxy resin. A 30 pm thin-section was made
at one-third of the total shell length from
the anterior end of each shell (one shell was
sectioned at one-fourth of the total shell
length) by Spectrum Petrographics, Inc.
(Winston, Oregon). Two shells were thin-
sectioned at two-thirds as well as one-third
of the way along the shell. We compared
cross-sectional area of the shell thin-section
that had underlain the anemone with the
portion that had been exposed. As a proxy
for area, we used weight of the enlarged
image of the thin-section traced onto matte
board. Tracings of shells that had been bare
or completely covered by a sea anemone
were also divided into two parts, a portion
that extended 270° around the shell, and a
90° portion. We normalized the weights to
the length of the inner perimeter of the shell
at that position. The ratios of normalized
weights of the covered and exposed por-
tions of each shell tracing were analyzed
with the Sign Test (Samuels 1989).

Shell thickness in the areas covered and
not covered by the anemone did not differ
significantly (Sign Test, p > 0.20). The
Shells of F. actiniophorum are, however,
unusually thin for a scaphopod, ranging in
thickness from 189 pm to 479 wm (mean
309 wm; n = 15), which is a third to half
as thick as the shells of F. erosum collected
at 3000—3300 m (Shimek & Moreno 1996).
[Shimek (1997:185) stated the thickness of
the F. actiniophorum shell to be ‘‘about 150
wm’’; he did not specify in which part of
the shell his measurements were made. In
one shell from which we had two sections,
the anterior end was slightly thicker, and in
the other shell there was no difference. ]

649

Acknowledgments

We thank K. L. Smith, Jr, R. Baldwin,
L. Lauerman, and S. Beaulieu, then of
Scripps Institution of Oceanography, for
collecting sea anemones and scaphopods,
and for generously sharing information
about the collection site and the deep sea.
Their research was supported by National
Science Foundation grants OCE89-22620
and OCE92-17334 to K. L. Smith. We
thank A. G. Carey, Jr. for making the spec-
imens from Oregon available to us. His re-
search was supported by Atomic Energy
Commission contracts AT (45-1)-1750 and
AT (45-1)-2227; National Science Founda-
tion grants GA-42655, GA-43202, GB-531,
and GB-4629; and National Oceanic and
Atmospheric Administration Sea Grant 2-
5187. We thank C. Hand, K. Riemann-Ziir-
neck, D. Alexander, C. Annett, and S.
Hauswaldt for comments on a previous ver-
sion of the manuscript, D. A. Doumenc for
helpful suggestions, R. L. Shimek for shar-
ing his knowledge of scaphopods, H. Feld-
man for instruction in the art of thin-sec-
tioning, and H. Chaney for the gastropod
identification. We also thank M. Schonew-
eis, of the Kansas Geological Survey, and
S. Pagels for help with the figures. A. G.
Carey, Jr. acknowledges M. A. Ashcraft, R.
E. Ruff, R. R. Paul, M. A. Kyte, D. R. Han-
cock, and other technicians and graduate
students for their invaluable assistance in
the field and laboratory. Some of this re-
search was done by AKWP in partial ful-
fillment of the Master of Arts degree in the
Department of Systematics and Ecology at
the University of Kansas. Financial support
of this research by the University of Kansas
to AKWP included a Panorama Society
Small Grant from the KUNHM and an Ida
H. Hyde Scholarship from the Division of
Biological Sciences. Financial support from
National Science Foundation grant DEB95-
21819 (PEET) to DGF is also gratefully ac-
knowledged.

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(4):652—660. 1999.

Urticina mcpeaki, a new species of sea anemone
(Anthozoa: Actiniaria: Actiniidae)
from the North American Pacific coast

J. Susanne Hauswaldt and Katherine E. Pearson

The University of Kansas Natural History Museum and Biodiversity Research Center and
Department of Ecology and Evolutionary Biology, The University of Kansas,
Lawrence, Kansas 66045, U.S.A.

Abstract.—Urticina mcpeaki, a new species of sea anemone, is described
based on specimens collected from the Coronados Islands, Baja California del
Norte, Mexico; Santa Cruz and Anacapa Islands, Channel Islands, California;
and Point Loma, California. Mesenteries and tentacles of the new species are
arranged hexamerously, rather than decamerously as is typical for other species
of Urticina. The new species is distinguished externally from other species in
the genus by a radial banding pattern on the oral disc, longitudinal striping on
the tentacles, and typically debris-covered adhesive verrucae on a bright-red
column. We include the first detailed description of the two morphologically
different types of microbasic p-mastigophores in the mesenterial filaments. Ad-
ditionally, size ranges of holotrichs in the tentacle tips are given for the first
time for a species in the genus Urticina.

Urticina Ehrenberg, 1834 is the most
speciose genus of sea anemones on the Pa-
cific coast of North America. Gotshall
(1994) recognized five species: Urticina
columbiana Verrill, 1922; U. coriacea (Cu-
vier, 1798); U. crassicornis (Miiller, 1776);
U. lofotensis (Danielssen, 1890); and U.
piscivora (Sebens & Laakso, 1977). Sebens
& Laakso (1977) recognized the same five
species under the generic name Tealia
Gosse, 1858, a junior synonym of Urticina
(Williams in Manuel, 1981). In addition to
these species, Zamponi & Acufia (1996) re-
ported Urticina asiatica (Averincev, 1967)
and U. tuberculata (Cocks, 1851) from
Barkley Sound, British Columbia, Canada.
On the west coast of North America, the
genus Urticina ranges from Alaska to Baja
California. Within this range, animals occur
either in the subtidal or in both the intertidal
and subtidal zones (Cutress 1949, Hand
1955, Sebens & Laakso 1977, Gotshall
1994, Zamponi & Acufia 1996). Sea anem-
ones in the genus Urticina also occur in the

Northwest Pacific and on both sides of the
North Atlantic. Though originally applied
to North Atlantic species, the names U. cor-
iacea, U. crassicornis, and U. lofotensis are
applied to Northeast Pacific species sensu
Hand (1955).

External characters that have been used
for distinguishing species in the genus Ur-
ticina are color pattern and the presence and
nature of verrucae on the column (Cutress
1949, Hand 1955, Sebens & Laakso 1977,
Gotshall 1994, Zamponi & Acufia 1996).
Internal characters include size and distri-
bution of cnidae, number and arrangement
of mesenteries, and distribution of fertile
mesenteries (Hand 1955, Sebens & Laakso
1977, Zamponi & Acuna 1996).

Materials and Methods

Divers collected nine animals in Septem-
ber 1995 at 25 m from 1.85 km southeast
of the tip of South Coronado Island, the Co-
ronados Islands, Baja California del Norte,

VOLUME 112, NUMBER 4

Mexico. Ten animals were collected in May
1997 from Point Loma, San Diego, Cali-
fornia at 15 m. Animals were held in tanks
for observation before relaxation with mag-
nesium sulfate and fixation with 10% for-
malin. Five museum specimens from the
California Academy of Sciences Depart-
ment of Invertebrate Zoology and Geology
(CASIZ) were also examined: CASIZ
61487 (two specimens) and 61488 (one
specimen) from Santa Cruz Island, Channel
Islands, California, and CASIZ 61491 (two
specimens) from Anacapa Island, Channel
Islands, California.

Undischarged cnidae capsules from the
column and structures at the margin, the ac-
tinopharynx, mesenterial filaments, and the
tips and bases of the outer and inner ten-
tacles were measured from squash prepa-
rations at 1000 magnification. Cnidae no-
menclature follows Mariscal (1974). Dis-
play of cnidae data follows Dunn, Chia, &
Levine (1980). Histological sections 8 ~m
thick were stained with hematoxylin and
eosin (Humason 1979). Oocytes were mea-
sured following the method of Wedi &
Dunn (1983).

Urticina mcpeaki, new species
Figs. 1A, B, 2—5

Pedal disc.—Circular, strongly adhesive.
Color bright red; mesenterial insertions vis-
ible as darker red radiating lines. Well de-
veloped basilar muscles along each mes-
entery.

Column.—Bright red with many small,
white, adhesive verrucae. Verrucae usually
heavily encrusted with debris (Fig. 1A) and
arranged in longitudinal rows correspond-
ing to endocoels and exocoels. No verrucae
on limbus. Well developed margin and
fosse. Distinct round marginal structures; in
life yellow, with dark-colored depression in
center (Figs. 1A and 2A). Histology (Fig.
2B) and cnidae complement of marginal
structures is not distinct from rest of col-
umn. When expanded, column generally
wider than tall. Maximum size of animals

653

smaller than other species in genus found
in Northeast Pacific; not larger than 5—6 cm
pedal disc diameter when expanded. Height
generally not greater than 3—4 cm; oral disc
diameter when expanded similar to pedal
disc diameter. No cinclides present.

Tentacles.—Long and slender when ex-
panded, short and blunt with longitudinal
furrows when contracted. Arrangement
hexamerous with up to 5 cycles; in most
specimens fifth cycle incomplete. Number
of tentacles 68—93. Base color of tentacles
in most individuals red; in some pink, tan,
or brown. Each tentacle encircled with
light-colored band at base. Longitudinal
stripe extends upward from band at base 1/
2—2/3 length of tentacle on aboral side (Fig.
1A), then broadens, creating chevron shape
on oral side (Fig. 1B). In most individuals
stripe white; other colors such as orange
may occur. Bases of tentacles outlined by
dark red bands corresponding to mesenter-
ial insertions (Fig. 1B). Longitudinal mus-
cles strong, mesoectodermal; circular mus-
cles weak, endodermal (Fig. 3).

Oral disc.—Color in most individuals
red; in some orange or pink. Mesenterial
insertions clearly visible. Broad, tentacle-
free region around mouth, usually white or
light olive green, gives way to distinctive
radial banding pattern, such that 12 thick
bands extend outward to spaces between
first two cycles of tentacles (Fig. 1B). Ac-
tinopharynx red, ribbed, with 2 red siphon-
oglyphs. Radial muscles strong, mesoecto-
dermal.

Mesenteries and internal anatomy.—
Mesenteries arranged hexamerously in 4
cycles. Fourth cycle usually incomplete;
fourth cycle mesenteries added in exocoels
between first and third cycles first, then be-
tween second and third cycles. Typical ar-
rangement from half of an animal, directive
to directive: D4 3234143243414
3 2 3 4 D. Number of mesentery pairs 29-—
44; one irregular individual with 52 pairs.
Number of mesenteries same proximally
and distally. Two pairs of directive mesen-
teries attached to siphonoglyphs. First and

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VOLUME 112, NUMBER 4

Fig. 2. Urticina mcpeaki, new species. A, View of
column showing marginal structures (slanted arrow)
and verrucae (horizontal arrow). Scale bar = 1.5 cm.
B, Histological section showing marginal structure and
sphincter (s). Scale bar = 0.2 mm. C, Histological sec-
tion showing verrucae with modified glandular ecto-
derm (arrow). Scale bar = 0.2 mm.

second cycles perfect. First cycle sterile; all
subsequent cycles may bear gametogenic
tissue. Sexes separate. Maximum diameter
of ripe oocytes in section approximately
450 wm. Retractor muscles strong, diffuse
to restricted (Fig. 4). Parietobasilar muscles
well developed and extending from half to
entire distance from column to retractor
muscle, terminating in free flap (Fig. 4).
Marginal stomata present; oral stomata not
seen.

655

Endodermal marginal sphincter muscle
circumscribed, large, strong; circular to
oval in shape (Fig. 3) and attached to col-
umn wall roughly halfway between fosse
and parapet. Laminae arranged pinnately
and of equal length on both sides of lamel-
la.

Cnidom.—Spirocysts, basitrichs, holo-
trichs, and microbasic p-mastigophores
(two types).

Distribution and size of nematocysts and
spirocysts.—In the following table, n sig-
nifies the number of individual capsules
measured, N signifies the ratio of the num-
ber of specimens in which a type of cnida
was found to the number of specimens ex-
amined, and measurements in parentheses
indicate individual capsules that fell outside
the range.

Tentacles (tip and base of inner and out-
er):
Spirocysts (Fig. 5A) 12.5—39.9 X 1.4—-
4.4 wm; n = 528, N = 15/15
Basitrichs (Fig. 5B) 6.5—32.3 x 1.1-
4.8 pm; n = 709, N = 15/15
Holotrichs (Fig. 5C) (found only in
tips, particularly of outer tentacles)
13.8-23.3 X 2.9-4.9 pm; n = 111,
N = 11/15

Column, including margin:

Basitrichs (Fig. 5B) 4.5—9.2 X 1.0—2.5
pm; n = 46, N = 10/11

Basitrichs (Fig. 5B) 13.4—23.1 (35.4)
xX 1.5—4.6 pm; n = 152, N = 11/11

Actinopharynx:

Basitrichs (Fig. 5D) 19.2—38.9 xX 2.5—
6.0 wm; n = 245, N = 14/14

Microbasic p-mastigophores type I
(Fig. 5E) 16.5—29.2 X 4.0—7.6 um;
n = 64,N = 8/14

Mesenterial filaments:

Basitrichs (Fig. 5B) 11.5—29.8 (34.4,
36.2) X 1.9—4.1 wm; n = 207, N =
ivi

Microbasic p-mastigophores type I
(Pig. SE) 11.7-30.3.G3A) X 3.5-
6.8 wm; n = 122, N = 11/11

656 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

a Yt ;
I MAN t.
rN %, al

Xx

Fig. 3. Urticina mcpeaki, new species. Histological section showing sphincter muscle and cross section of
tentacle. Scale bar = 0.25 mm.

Microbasic p-mastigophores type II ifornia, south at least to Asuncion, Baja
(Fig. SF) 19.8—42.9 X 3.6—-8.6 pm; California, Mexico.
n= 113, N = 11/11 Habitat.—Subtidal, in shallow debris-

Distribution.—From Santa Barbara, Cal- _ filled depressions of rocks.

Fig. 4. Urticina mcpeaki, new species. Histological section showing oocytes and three pairs of mesenteries
with retractor muscles and parietobasilar muscles. Scale bar = 1 mm.

VOLUME 112, NUMBER 4

Fig. 5. Urticina mcpeaki, new species. Cnidae sig-
nature. See text for explanation. Scale bar = 5 wm.

Type material and locality.—Holotype
collected May 1997 from Point Loma, San
Diego, California at 15 m; deposited at the
CASIZ 114595. Histological slides of spec-
imen also deposited.

Paratypes from type locality CASIZ
114596 and 114597, Santa Barbara Muse-
um of Natural History (SBMNH) 145138
and 145139, University of Kansas Natural
History Museum (KUNHM) 001041, Na-
tional Museum of Natural History (NMNH)
99971 and 99972, and Royal British Co-
lumbia Museum (RBCM) 998-221-1. Par-
atype from the Coronados Islands also de-
posited at RBCM 998-222-1. Additional

657

specimens examined deposited at KUNHM
001038, 001162 through 001168.
Etymology.—The _ specific epithet
mcpeaki honors Mr. Ron McPeak, who first
noticed this species in 1958 and suspected
it was a new species of Urticina. He pro-
vided many of the specimens examined.

Discussion and Differential Diagnosis

Urticina mcpeaki, new species, fits the
generic description of Tealia (= Urticina)
sensu Carlgren (1949). It is an actiniid with
a well-developed pedal disc, adhesive ver-
rucae on the column, and a well-developed
fosse. It has a strong, circumscribed, en-
dodermal sphincter, and the longitudinal
musculature of the tentacles and radial mus-
culature of the oral disc are mesoectoder-
mal. The basitrichs of the actinopharynx are
larger than those of the tentacles.

The major morphological difference be-
tween U. mcpeaki and other species in this
genus is that mesenteries and tentacles of
the first three cycles are always arranged
hexamerously. Mesenteries in the fourth cy-
cle are not added simultaneously, resulting
in a break of the hexamerous pattern. In
other species of Urticina, mesenteries and
tentacles are usually arranged decamerously
(Carlgren 1893, 1921, 1949; Hand 1955,
Sebens & Laakso 1977, Manuel 1981).
Hexametry in U. mcpeaki is unusual, but it
is not unique within Urticina. Several au-
thors (e.g., Carlgren 1902, 1949; Sebens &
Laakso 1977, Zamponi & Acuna 1996) re-
fer to specimens of Urticina in which the
arrangement of either the mesenteries or the
tentacles is not strictly decamerous, but
rather hexamerous.

The new species can also be distin-
guished from other species in this genus by
its geographic range. Urticina mcpeaki oc-
curs as far north as Santa Barbara, Califor-
nia (Jack Engle, Ron McPeak, pers.
comm.). The geographic ranges of all other
species of Urticina on the west coast of
North America extend at least as far north
as Vancouver Island, British Columbia,

658

Canada. Most species are found as far north
as Alaska (Gotshall 1994). The range of U.
mcpeaki overlaps with the ranges of U. co-
lumbiana, U. coriacea, U. lofotensis, and
U. piscivora (Ron McPeak, pers. comm.).

Of all species of Urticina on the North
American Pacific coast, U. mcpeaki most
closely resembles U. lofotensis externally.
Both species have a bright red column with
distinct white verrucae arranged in longi-
tudinal rows. However, U. mcpeaki can be
distinguished externally from U. lofotensis
by the color pattern of the tentacles and oral
disc. Urticina mcpeaki has a distinct lon-
gitudinal stripe on the otherwise uniformly
colored tentacles (Fig. 1A). Urticina lofo-
tensis does not have stripes on the tentacles
and the color changes from yellow at the
base to red at the tip (Fig. 1D). In U.
mcpeaki, the oral disc has a distinct radial
banding pattern (Fig. 1B), while the oral
disc of U. lofotensis lacks this pattern (Fig.
1D).

The structures at the margin of U.
mcpeaki differ from those of U. lofotensis.
In U. mcpeaki they are yellow and have a
dark-colored depression in the center (Figs.
1A and 2A). The structures in U. lofotensis
are white and do not differ in appearance
from the verrucae on the column (Fig. 1C).
Histologically the marginal structures in
both species differ from verrucae. Verrucae,
sensu Carlgren (1949), have modified ec-
toderm (e.g., Fig. 2C), which is lacking in
the marginal structures. The distinct mar-
ginal structures of U. mcpeaki may be con-
fused externally with other marginal spe-
cializations such as spherules or pseudo-
spherules. However, they do not fit the
definition of marginal spherules or pseu-
dospherules, sensu Carlgren (1949), be-
cause they have no distinctive histology
(Fig. 2B) or cnidae.

The verrucae of U. mcpeaki usually have
debris attached to them (Fig. 1A). Although
the verrucae of U. lofotensis are capable of
attaching debris (pers. observation), speci-
mens are usually found with no debris at-
tached (Fig. 1C). In U. mcpeaki, verrucae

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

are absent from the limbus. In U. lofotensis,
verrucae extend to the limbus where they
are smaller and closer together than the ver-
rucae of the column.

Lastly, the two species differ in size. The
pedal disc diameter in the largest specimens
of U. mcpeaki is 6 cm, while in U. lofoten-
sis it can reach to 18 cm (Sebens & Laakso
LOT):

This is the first detailed description of the
two types of microbasic p-mastigophores
found in the mesenterial filaments. Micro-
basic p-mastigophore type I (Fig. 5E) is
more oval than type II (Fig. 5F) and has
pronounced serrations all along its robust
shaft. Cnidae of type II are usually larger
than those of type I; however, there is some
overlap in size within individuals. Type II
are more slender than type I, in proportion
to their lengths, and the serrations of the
shaft are not as distinct. In a few individ-
uals, small numbers of microbasic p-mas-
tigophores type I were also found in other
tissues (column, actinopharynx, and tenta-
cle base). These capsules were considered
contaminations of those tissues. Microbasic
p-mastigophore type II were found only in
the mesenterial filaments. We have subse-
quently found both types of microbasic p-
mastigophores in U. columbiana, U. cori-
acea, U. crassicornis, U. lofotensis, and U.
piscivora from the Northeast Pacific. In his
monograph on sea anemones of central Cal-
ifornia, Hand (1955) included line drawings
of the two types of microbasic p-mastigo-
phores he found in the mesenterial fila-
ments of Tealia crassicornis (p. 74) and T.
lofotensis (p. 85) and the one type he found
in T. coriacea (p. 81). He did not describe
the differences between the two types. Our
type I corresponds to Hand’s line drawing
labeled “‘g’”’ for 7. crassicornis and T. lo-
fotensis, and *‘k”’ for T. coriacea. Our type
II corresponds to Hand’s line drawing la-
beled “th”? for T. crassicornis and T. lofo-
tensis.

This is also the first description of a spe-
cies of Urticina that includes a size range
for tentacle holotrichs. In their report of

VOLUME 112, NUMBER 4

several species of Urticina from British Co-
lumbia, Zamponi & Acuna (1996) did not
give sizes and probably identified the type
of cnidae we term holotrichs as atrichs.
Holotrichs have armature along the entire
tubule, as opposed to atrichs, which have
no armature (Weill 1930). If the spines are
very fine, holotrichs can be confused with
atrichs (Cutress 1955). As Bigger (1976,
1982) pointed out, atrichs and holotrichs
have been confused repeatedly throughout
the literature. Sebens & Laakso (1977)
mentioned a personal communication with
C. Hand regarding the size of holotrichs
found in the tentacle tips of several species
of Tealia. The holotrichs were reportedly
larger than other nematocysts in the tentacle
tips. We found in U. mcpeaki that holotrichs
overlap in size with the basitrichs and are
never larger. Preliminary data suggest this
is also true for U. coriacea, U. crassicornis,
and U. lofotensis from the Northeast Pacif-
ic:

Acknowledgments

This work was funded by National Sci-
ence Foundation grant #DEB95-21819
(PEET) awarded to Dr. Daphne G. Fautin,
University of Kansas. We are grateful to Dr.
Fautin for valuable suggestions and discus-
sions of this work and for reviewing this
manuscript. We thank the following people:
Ron McPeak, research associate of the San-
ta Barbara Museum of Natural History, and
currently of Global Biological Consultants,
Battle Ground, Washington, for specimens,
distribution and natural history data, and for
reviewing the manuscript; Constance
Gramlich of San Diego State University for
specimens and for hospitality; Dr. Jack En-
gle of the Marine Science Institute at the
University of California, Santa Barbara, and
the Tatman Foundation Channel Islands Re-
search Program, for supplying distribution
data; M. en C. Gustavo Hernandez Car-
mona of the Instituto Politecnico Nacional,
Mexico for help with the collections in
Mexico; and Dr. C. Hand of Bodega Marine

659

Laboratory for helpful comments on the
manuscript. Order of authorship was deter-
mined by coin toss.

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(4):661—666. 1999.

Morphological observations on a hatchling and a paralarva of the
vampire squid, Vampyroteuthis infernalis Chun
(Mollusca: Cephalopoda)

Richard E. Young and Michael Vecchione

(REY) Department of Oceanography, University of Hawaii, Honolulu, Hawaii 96734, U.S.A.;
(MV) National Marine Fisheries Service, Systematics Laboratory,
National Museum of Natural History, Washington D.C. 20560, U.S.A.

Abstract.—We present morphological observations on two very small vam-
pire squids. One, a trawl-caught hatchling of 8 mm mantle length (ML), had
embryonic features including internal yolk, broad arm-like filaments, and lack
of mantle-head fusion. The other was a 9 mm ML paralarva collected by
submersible and photographed alive aboard ship. Although only slightly larger
than the hatchling, its morphology was much more like that of larger specimens
than that of the hatchling, except for the oblique orientation of its fins.

The vampire squid, Vampyroteuthis in-
fernalis Chun, 1903, is the only living
member of the Vampyromorpha, the sister
group to the Octopoda (Young & Vecchione
1996, Young et al. 1998). Through out-
group and ontogenetic comparisons, this
species holds important clues to our under-
standing of cephalopod evolution and to the
reconstruction of ancestral character states.
Very little has been published, however,
concerning its anatomy, behavior or embry-
ology. As a result, any new information is
valuable. We report here on two very small
vampire squid. One, a hatchling captured
by trawl, was dead but in excellent condi-
tion when retrieved. The other was a para-
larva (= “‘stage 1 larva’, Pickford 1949a)
captured alive from a submersible and pho-
tographed in a shipboard aquarium. These
are the first observations of a live vampire
squid paralarva. Together, these observa-
tions support many morphological charac-
ters described by Pickford (1949a) from
damaged specimens.

Materials and Methods

The hatchling Vampyroteuthis (8 mm
ML, USNM 885891) was captured near the

Hawaiian Islands on a cruise of the R/V
New Horizon off leeward Oahu on 1 July
1996. The collecting gear was an opening—
closing net that fished between 0130 and
0743 h at a depth of about 1050-1300 m
(Sta. 62). The specimen was sorted from the
catch soon after collection, and photographs
were taken aboard ship shortly thereafter.
The paralarva (9 mm ML, USNM 816886)
was captured in the eastern Gulf of Mexico
using a pelagic suction sampler from the
manned submersible Johnson Sea Link II
during dive 1453, 30 August 1987,
24°30'5"N, 83°45'12”W at 830 m depth. It
was photographed while still alive imme-
diately after the submersible returned to its
support ship.

Observations

Hatchling.—The trawl-caught animal
was clearly a hatchling (i.e., there was no
chorion surrounding it), but morphological-
ly it was an advanced embryo presumably
incapable of feeding except from its inter-
nal yolk supply. A large mass of yolk glob-
ules, presumably within a large internal
yolk sac, was seen through the transparent
gladius ventral and lateral to the esophagus.

B

%
Filament
ae i

y , *
JP
m@ XS

Pig, l:

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Esophagus

Photographs of freshly collected Vampyroteuthis infernalis hatchling (8 mm ML). A, Lateral view

with dissecting pin laying across mantle to prevent motion from the ship’s rolling; B, Dorsolateral closeup of

head and anterior mantle.

During attempts to photograph the embryo
aboard ship, a pin was placed on top of the
mantle to keep the animal from rolling (Fig.
1A). This resulted in a sausage-shaped ex-
trusion of yolk from the buccal region.
The body of the hatchling was very large
relative to the head and arms (compare Fig.

Fig. 2.

1A with Fig. 3), the eyes were very small
and the mantle was not fused to the head in
the nuchal region (Fig. 2). The eye fold
formed a circular fold around the base of
each eye, not yet enclosing the eye. Al-
though squid eyes often are forced out
through circular eyelids during capture, this

Break in tissue

Schematic drawings of Vampyroteuthis infernalis hatchling, lateral, dorsal, and oral views. From top

to bottom on the oral view the arm pairs are dorsal (Arms I), filaments (Arms II), dorsolateral (Arms III),

ventrolateral (Arms IV), and ventral (Arms V).

VOLUME 112, NUMBER 4

was not a case in this instance. The epithe-
lial body could clearly be seen surrounding
the lens of the eye even though the lens was
partially covered by the iris.

Gelatinous tissue surrounded the arms
and separated muscle layers in the mantle
but did not cover the mantle muscle exter-
nally. At the posterior dorsal apex of the
mantle a small knob (ca. 15 ym) was pres-
ent in the epithelium and partially surr-
ounded by a groove. This knob is in the
position of the hatching organ of other
cephalopods but we find it reminiscent of
the peculiar circular feature in the photos of
the paralarva (Fig. 3B—D) described below.

The filaments (= ‘‘velar filaments’’,
Hochberg & Nixon 1992) were intact. Each
was broad, short, of uniform thickness to
the rounded tips (Figs. 1B, 2), and lacked
both suckers and cirri. Their attachment to
the brachial crown was as in the adult ex-
cept that the filaments did not reside in a
pocket, as a web was virtually absent. A
slight ridge between arms III, [V and V
(Fig. 2) appeared to be the web precursor.
The arm tips were slightly damaged but all
seemed to be subequal and armed with cirri
and suckers. Four developing suckers were
counted on one arm and we assume this to
apply to all arms. Suckers had constricted
bases but lacked obvious stalks. Four pairs
of cirri preceded the first sucker and the
suckers and cirri alternated as in the adult.

On the oral surface of the brachial crown,
two circular lips and a more distal fold sur-
rounded the beak. A break in the tissue ven-
tral to the lips and medial to this fold was
presumably the point where yolk came out.
Unlike in adults, the large funnel was not
embedded in the head, but no free adductor
muscles were present. The exact orientation
of the very small, but damaged, fins could
not be determined. The pigmented esopha-
gus, where it emerged from the brain, was
clearly visible through the transparent glad-
ius. The esophagus gradually increased in
diameter posteriorly. The gladius was the
same general shape as in adults but had a
somewhat narrower median field.

663

The animal’s coloration was faint pink-
brown and fairly translucent. Black chro-
matophores were scattered at several points
on the animal, especially on the oral surface
of the arm bases and anterodorsally on the
eye. A light reddish-brown epidermal pig-
ment was present. The pigment was most
obvious on the covering of the visceral
mass, the ventral surface of the head and
the oral surfaces of the arms. The animal
seemed to be neutrally buoyant although
small bubbles attached to the epidermis pre-
vented actual determination.

Paralarva.—While nearly the same size
(9 vs. 8 mm ML), the morphology of the
paralarva was very different from that of
the hatchling. Except as noted below, the
overall appearance was similar to that of
larger juveniles and adults (e.g., head/man-
tle fusion, pigmentation, etc.). The oblique
angle of the fin noted by Pickford (1949a)
is confirmed here in photographs (Fig. 3B,
C) of the living paralarva. The fins were the
first of two pairs to develop (= “‘juvenile
fins” or “larval fins’, Young & Vecchione
1996); the second pair (“‘adult fins’”’ Ibid)
were present only as minute precursors an-
terior to the fin-base photophore. Also ap-
parent in these live photos (Fig. 3B—D) is a
peculiar circular feature at the apex of the
mantle. Examination of the preserved spec-
imen, however, revealed no obvious struc-
ture at this location. Figure 3A shows the
arms in a forward orientation, and Fig. 3B
shows them in the lateral orientation. The
filaments were very thin and emerged from
pockets in the well-developed web, as in
older specimens. The incompletely retract-
ed but coiled filaments are seen most clear-
ly in Fig. 3C. The eyes of this young vam-
pire squid were relatively small (Fig. 3A,
C), compared with those of an adult.

Discussion

Pickford (1949a) described a young vam-
pire squid in the same stage of development
as the hatchling described here. However,
her specimen was badly damaged (mantle

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Filament,

(Detritus)

Mantle-apex structure

Buccal area

D

Fig. 3. Photographs of live paralarval Vampyroteuthis infernalis (9 mm ML). All views dorsal. A, with arms
in anterior orientation; B, with arms and web spread laterally; C, showing coiled, partially retracted filament;
D, with arms and web folded posteriorly revealing buccal area.

VOLUME 112, NUMBER 4

length was thought to be 5.7 mm) and, as
a result, some of her conclusions have been
in doubt. The description here confirms
some of her more important findings (e.g.,
mantle free from head; thick filaments).

The vampire hatchling also confirms the
fusion of the head and mantle in octopods
and vampyromorphs to be independent, as
predicted from morphology (Young & Vec-
chione 1996). The broad filaments are more
arm-like than in older vampire squids and
add support to the interpretation that the fil-
aments are the second pair of arms. If they
were ocular filaments, as suggested by
Young (1967), one might expect that they
would be especially small at this stage. The
embryonic eye fold in cephalopods is con-
tinuous with the arm bases and the partic-
ular arms associated with the fold have
been used to determine the homology of
arm pairs in decapods and octopods (Bol-
etzky 1993). Unfortunately, in the vampire
hatchling the eye fold does not continue
onto the arm crown, presumably because of
its advanced stage of development. A fold
surrounding the lips could be the remnant
of a buccal crown that is otherwise absent
in Vampyroteuthis (see Fig. 3D). More like-
ly, however, this is the demarcation be-
tween the gelatinous tissue and the more
muscular region where the arm muscles
fuse with one another and has no phyloge-
netic significance.

Vampire hatchlings may drift freely in
deep water until they change into the form
of a typical paralarva. However, the speci-
men examined here could be an advanced
embryo, hatched prematurely within the
trawl. Pickford (1949b) stated that the egg
size of spawned Vampyroteuthis eggs is up
to 4.0 mm in diameter. Clearly the present
hatchling with its embryonic features (e.g.,
yolk) was not derived from such a small
egg. An egg size of ca. 8 mm (i.e., com-
parable to the mantle length) would be ex-
pected. The discrepancy cannot be ex-
plained at present.

Young et al. (1998) proposed that relaxed
orientation of the arms in Vampyroteuthis

665

might provide evidence to choose between
alternative hypotheses about octopodiform
evolution. However, the photographs of the
paralarva with its arms both spread laterally
and projecting anteriorly are equivocal on
this question.

Vampyroteuthis infernalis currently is
considered to comprise a single panmixic
species. Therefore, although our specimens
were collected in different oceans, we do
not believe that the morphological differ-
ences between them reflect interspecific
variability. The strong differences in mor-
phology between our two specimens, which
are nearly the same size, demonstrate the
changes that occur in the early post-hatch-
ing ontogeny of Vampyroteuthis. Pickford
(1949a) described the paralarval develop-
ment of V. infernalis. One peculiarity of
this species is that early paralarvae (Pick-
ford’s “‘stage 1 larvae’? have one pair of
fins; then a second pair develops (“‘stage
2”’) followed by absorption of the original
pair (“‘stages 3 and 4’’). She observed that
the fins of the paralarva are set obliquely
on the body, unlike the horizontal attach-
ment of the adult fin, and suggested that the
young vampire hangs head downward in
the water. The present observations, which
are the first live of a paralarval vampire
squid, confirm Pickford’s descriptions, such
as fin orientation, which were based on pre-
served specimens.

The pucker-like circular structure seen
near the posterior apex of the paralarva’s
mantle is similar to that recorded in a sub-
adult Vampyroteuthis videotaped as part of
a televison program about cephalopods
(“Incredible Suckers’’). No discrete struc-
ture is apparent on the preserved paralarva
and we do not know the function of the
structure.

Acknowledgments

We thank J. Childress, University of Cal-
ifornia, Santa Barbara, for the opportunity
to participate in one of his cruises and we
acknowledge the keen eye of Eric Thuesen

666

that detected the small hatchling vampire
amidst a large bucket of midwater animals.
Richard Harbison, Woods Hole Oceano-
graphic Institution, provided the paralarval
vampire and Ronald Gilmer took the pho-
tos. Bruce Collette, Michael Sweeney, Eliz-
abeth Shea, and a helpful anonymous re-
viewer provided suggestions for improve-
ment of drafts of this paper. The research
reported here was funded in part by a grant/
cooperative agreement from the National
Oceanic and Atmospheric Administration.
project #R/MR-51, which is sponsored by
the University of Hawaii Sea Grant College
Program, SOEST, under institutional grant
No. NA86RG0041 from the NOAA Sea
Grant Office, Department of Commerce.
UNIHI-SEAGRANT-JC-99-03.

Literature Cited

Boletzky, S. von. 1993. The arm crown in cephalopod
development and evolution: a discussion of

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

morphological and behavioral homologies.—
American Malacological Bulletin 10:61—69.

Hochberg, EF G., & M. Nixon. 1992. Order Vampyro-
morpha Pickford, 1939. Pp. 211-212 in M. J.
Sweeney, C. FE E. Roper, K. M. Mangold, M.
R. Clarke, & S. von Boletzky, eds., ‘Larval’ and
juvenile cephalopods: a manual for their iden-
tification.—Smithsonian Contributions to Zool-
ogy 513:282 pp.

Pickford, G. E. 1949a. Vampyroteuthis infernalis
Chun. An archaic dibranchiate cephalopod. II.
External anatomy.—Dana Report 32:1—132.

. 1949b. The distribution of the eggs of Vam-
pyroteuthis infernalis Chun.—Journal of Ma-
rine Research 8:73-83.

Young, R. E. 1967. Homology of retractile filaments
of vampire squid.—Science 156:1633—1634.

, & M. Vecchione. 1996. Analysis of morphol-

ogy to determine primary sister-taxon relation-

ships within coleoid cephalopods.—American

Malacological Bulletin 12:91—112.

: , & D. T. Donovan. 1998. The evolu-

tion of coleoid cephalopods and their present

biodiversity and ecology.—South African Jour-

nal of Marine Science 20:393—420

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(4):667—681. 1999.

Nereidids (Polychaeta) from the Caribbean Sea and adjacent Coral
Islands of the southern Gulf of Mexico

Jesus Angel de Leén-Gonzalez, Vivianne Solis-Weiss, and Victor Ochoa Rivera

(JAL-G) Centro de Investigaciones Biolégicas del Noroeste, S.C.;
Laboratorio de Zoologia de Invertebrados, Facultad de Ciencias Biolégicas, UANL, Ap. Postal 5 “‘F’’,
San Nicolas de los Garza, N.L. 66451, México; (VS-W, VO-R) Laboratorio de Ecologia Costera,
Instituto de Ciencias del Mar y Limnologia, (ICML) UNAM, Apdo. Postal 70-305,
México, D.F, 04510, México

Abstract.—In this study 23 species of nereidids from the Greater Caribbean
region collected in different expeditions from 1930 to 1970 are recorded: six
species of Ceratonereis, five species each of Nereis and Perinereis, four species
of Neanthes and one species each of Platynereis, Pseudonereis and Stenoni-
nereis. Neanthes egregicirrata is removed from the genus Nereis, because no
homogomph falcigers are present in the atokous form, which is herein de-

scribed for the first time.

In a recent study, Salazar-Vallejo &
Jiménez-Cueto (1997) reported that for the
Greater Caribbean region 48 species of ner-
eidids are known. In the same study, 17
species of nereidids were recorded for the
state of Quintana Roo’s (Mexico) littoral
waters, and a key was given for species of
the Greater Caribbean Region.

In this study, 23 species of nereidids are
recorded. The material comes from several
oceanographic programs: collections made
at Caribbean islands between 1930 and
1973 and deposited in the Instituut voor
Taxonomische Zodlogie (Zodlogisch Mu-
seum) University of Amsterdam (ZMUA);
collections made by the Smithsonian-Bre-
din Caribbean Expedition IV in 1960 in
Quintana Roo, Mexico, and deposited in the
National Museum of Natural History,
Smithsonian Institution, Washington, D.C.
(USNM); and collections made by two of
us (VS-W, VO-R) as part of the Imca IV
(March 1989) and Dinamo I and III (March
90 and 91) oceanographic expeditions (see
Ochoa-Rivera 1996) and deposited in the
Laboratorio de Ecologia Costera, Instituto
de Ciencias del Mar y Limnologia, Univ-
ersidad Nacional Aut6énoma de Mexico

(ICML-UNAM), in the coral islands from
the southwestern region of the Gulf of Mex-
ico.

In addition, some type specimens from
the Naturhistoriska Rijksmuseet Stockholm
(NRS) and other specimens deposited at the
Smithsonian Institution are included here.

Under Material Examined, the records
are reported in the following order and as
the case may be: the type material is listed
first, then the material coming from the In-
stituut voor Taxonomische Zodlogie, fol-
lowed by the Smithsonian-Bredin Caribbe-
an Expedition IV material, and ending with
the ICML-UNAM material. Dates are ab-
breviated in order of day, month and year.
For each sampling station the number of
specimens studied is in parentheses.

Ceratonereis (Ceratonereis) excisa
(Grube, 1874)

Nereis excisa Grube, 1874:72.

Ceratonereis excisa.—Perkins, 1980:13,
fig. 6.—Hartmann-Schréder, 1985:47.
Salazar-Vallejo & Jiménez-Cueto, 1997:
363.

Material examined.—ZMUA: Bonaire,

668

Lac entrance, 200 m West of Cai, 6 m
depth, 11 Aug 1967 (3).
Distribution.—Western Atlantic, Cuba,
Bonaire and Santa Catarina Island, Brazil.
Habitat.—Shallow waters, in sand
trapped among Thalassia roots.

Ceratonereis (Composetia) irritabilis
(Webster, 1879)

Ceratonereis irritabilis.—Taylor, 1984:
31.30, figs. 31.25, 31.26.—Salazar-Val-
lejo & Jiménez-Cueto, 1997:363.

Material examined.—ZMUA: Barbados,
0.5 mi off Holetown, 100 m depth, 19 Nov
1964 (14).

Distribution.—From Virginia, U.S.A.,
south to Panama.

Habitat.—Soft bottoms, intertidal to con-
tinental shelf.

Ceratonereis (Ceratonereis) longicirrata
Perkins, 1980

Ceratonereis longicirrata Perkins, 1980:26,
figs. 11-12.

Material examined.—ZMUA: Puerto
Rico, Paraguera Mata de la Gata, 12 Sep
1963 (4); St. Martin, Etang aux Poissons,
lV’ Embouchure, 3 Oct 1963 (1); Little Cay-
man, Owen Island, ca. 0.5 m depth, 7 Jun
1973 (5); Grand Cayman, south of Jack-
son’s Point, 0O-O0.5 m, 9 Jun 1973 (1);
Smithsonian-Bredin Caribbean Expedition
IV, Est. 85-60, Ascencién Bay, 200-300
yds. southwest of Suliman Point, 17 Apr
1960 (8); ICML-UNAM: Dinamo I and IIT
expeditions, Alacran Reef, West bay 18
Mar 1990 (1); 18 Mar 1991 (1); Triangulos
Oeste reef, West bay, 19 Mar 1991 (1).

Distribution.—Tropical Western Atlantic.
Known from the Gulf of Mexico, and Ca-
ribbean Sea.

Habitat.—Soft bottoms in the continental
Shelf, in sediments accumulated among
roots of algae attached to coral rocks.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Ceratonereis (Ceratonereis) mirabilis
Kinberg, 1866

Ceratonereis mirabilis Kinberg, 1866:
170.—Perkins 1980:4, figs. 1—4.—Tay-
lor, 1984:31.30, figs. 31.27—31.28.—
Hartmann-Schroéder, 1985:43, fig. 23.—
Salazar-Vallejo & Jiménez-Cueto, 1997:
363, figs. 5, 20, 21, 42, 44.

Material examined.—Brazil, 09°S, 33 m,
Werngren Coll., (Syntype, NRS 539).
ZMUA: Bonaire, Kralendijk, near Pasang-
grahan, Sep 1930 (1); St. Barts, S. Public
near Gustaria, 4 Jun 1949 (1); North Bim-
ini, Bahamas, 1 km offshore, 17 Aug 1949
(29); 18 Aug 1949 (13); Florida, east of
Soldier Key, 2 m, 5 Sep 1963 (1); Barba-
dos, 0.5 mi of Holetown, 100 m, 19 Nov
1964 (28) Little Cayman, South Hole
Sound, The Bight; 0.5-1 m, 5 Jun 1973
(71); Jamaica, Drunkemans, Key, O—0.5 m,
15 Jun 1973 (7). USNM: Smithsonian-Bre-
din Caribbean Expedition IV, Est. 11-60,
Mujeres Island, 29 Mar 1960 (1); Est. 41-
60, Espiritu Santo Bay, north shore, near
Lawrence Point, 6 Apr 1960 (3); Est. 44-
60, North end of Ascencién Bay, small inlet
behind Allen Point, 7 Apr 1960 (4); Est. 52-
60, Ascencion Bay, just behind center of
Nicchehabin Reef, east of Allen Point, 10
Apr 1960 (6); Est. 53-60, Ascencién Bay,
10 Apr 1960 (6); Est. 61-60, near Allen
Point light, 12 Apr 1960 (129); Est. 67-60,
Ascencion Bay, 13 Apr 1960 (12); Est. 68-
60, Ascencién Bay, Allen Point, 13 Apr
1960 (2); Est. 72-60, Ascencié6n Bay, cen-
tral part of Nicchehabin Reef, 14 Apr 1960
(7); Est. 77-60, North end of Ascencién
Bay, beach just east of Halfway Point, 15
Apr 1960 (26); Est. 79-60, Ascencién Bay,
the electric light over ship’s side at anchor-
age, 15 Apr 1960 (2); Est. 83-60, Ascen-
cion Bay, mangrove inlet behind Allen
Point light, 16 Apr 1960 (2); Est. 85-60,
Ascencion Bay, along shore near Suliman
Point, 17 Apr 1960 (2); Est. 91-60, Inner
side, Nicchehabin Reef, 1/3 distance from
north end Ascencion Bay, 18 Apr 1960
(20); Est. 93-60, north end Ascencién Bay,

VOLUME 112, NUMBER 4

shore near Halfway Point, 18 Apr 1960 (2);
Est. 95-60, Ascencion Bay, Suliman Point,
19 Apr 1960 (5); Est. 100-60, north of San-
ta Maria Point, south end of Cozumel Is-
land, 21 Apr 1960 (3); Est. 109-60, south
end of Cozumel Island, 114 miles north of
Santa Maria Point, 22 Apr 1960 (2).

Distribution.—Western Atlantic. Greater
Caribbean region, Bermuda, Florida, Ba-
hama Islands, south to Barbados and the
Netherlands Antilles. Santa Catarina, Bra-
zil.

Habitat.—Mixed bottoms, coral reefs,
macroalgae, seagrass beds, shallow waters
down to 60 m.

Ceratonereis (Ceratonereis) singularis
(Treadwell, 1929)

Ceratonereis singularis Treadwell, 1929:1,
figs. 1—8.—Perkins, 1980:17, figs. 7-
10.—Hartmann-Schréder, 1985:45, figs.
36—39.—Salazar-Vallejo et al., 1990:
213.—Bastida-Zavala, 1993:29.

Material examined.—ZMUA: Antigua,
Dickinson Bay, 26 Jul 1967 (5); 28 Jul
1967 (19).

Distribution.—Amphi-American. Gulf of
California south to Panama, North Carolina
south to Colombia.

Habitat.—Rocky substrates of the inter-
tidal zone, sandy bottoms in the continental
shelf.

Ceratonereis (Composetia) versipedata
(Ehlers, 1887)

Nereis (Ceratonereis) versipedata Ehlers,
1887:116, pl. 36, figs. 5-10.

Ceratonereis versipedata.—Day, 1973:
39.—Gardiner, 1976:148, fig. 140.—Tay-
lor, 1984:31.30, figs. 31.23; 31.24a-f.

Material examined.—ZMUA: Curacao,
Piscadera Baai, north part, 2nd buoy, 31
Oct 1963 (2); central part, 3rd buoy, 31 Oct
1963 (2); Piscadera Baai, northeast point,
11 Dec 1963 (3). USNM: Smithsonian-Bre-
din Caribbean Expedition IV, Est. 41-60,
Espiritu Santo Bay, westward side of reef

669

to east of anchorage, 6 Apr 1960 (3); Est.
52-60, Ascencion Bay, just behind center of
Nicchehabin Reef, east of Allen Point, 10
Apr 1960 (1); Est. 53-60, Ascencién Bay,
10 Apr 1960 (1); Est. 77-60, north end of
Ascencion Bay, shore just east of halfway
point, 15 Apr 1960 (1).). ICML-UNAM:
Dinamo III expedition, Cayo Nuevo, 24
Mar 1991 (2); Cayo Arcas, 17 Mar 1991
(6).

Distribution.—Western Atlantic. Geor-
gia, U.S.A., Gulf of Mexico and Caribbean
Sea.

Habitat.—Soft bottoms and among inter-
stices of coralline rocks, intertidal to 40 m.

Neanthes acuminata (Ehlers 1868)

Nereis acuminata Ehlers 1868:552.

Nereis (Neanthes) arenaceodonta: Petti-
bone, 1963:162, figs. 441, 45e.

Nereis (Neanthes) acuminata: Day, 1973:
41.—Gardiner, 1976:149, fig. 15e-f.

Neanthes acuminata: Taylor, 1984:31.15,
figs. 31.11; 31.12a-—e.

Material examined.—ZMUA: Florida,
Key Biscayne, northwest swamp (mangrove
ditch), 31 Aug 1963 (2); Puerto Rico, Par-
guera, Mata de the Gata, 12 Sep 1963 (2);
Curacao, Piscadera Baai, northeast point,
25 Nov 1963 (1); 11 Dec 1963 (3); 18 Dec
1963 (6); central part, southeast point, 13
Dec 1963 (7); Jamaica, Kingston Harbour,
Inlet W. of airport, O—1 m, 7 May 1973 (1).
USNM: Smithsonian-Bredin Caribbean Ex-
pedition IV, Stn. 17-60, Mujeres Harbor, 30
Mar 1960 (1); Stn. 83-60, Ascencion Bay,
mangrove inlet behind Allen Point light, 16
Apr 1960 (1); Stn. 100-60, north of Santa
Maria Point, south end of Cozumel Island,
21 Apr 1960 (5).

Distribution.—Western Atlantic. From
North Carolina down to the Gulf of Mexi-
co.

Habitat.—Soft bottoms of the continen-
tal shelf.

670

Neanthes egregicirrata (Treadwell, 1924)
(Fig. la—h, 2a—f)

Nereis (Leptonereis) egregicirrata Tread-
well, 1924:13, fig. 24.

Material examined.—Antigua Island,
English Harbor, Coll. University of Iowa,
07/1918 (1, Lectotype USNM 071733), (4
Paratypes USNM 020224); ZMUA: Bar-
bados Island, Conset Bay (St. John), 7 Jul
1967 (1); Litthe Cayman, South Hole
Sound, The Bight, 0.5—1 m, 5 Jun 1973 (2).
USNM: Smithsonian-Bredin Caribbean Ex-
pedition IV, Stn. 67-60, Ascencion Bay, be-
hind central part of Nicchehabin Reef, 13
Apr 1960 (2); Stn. 72-60, Ascencion Bay,
central part of Nicchehabin Reef, 14 Apr
1960 (2); Stn. 79-60, Ascencién Bay, the
elelectric light over ship’s side at anchor-
age, 15 Apr 1960 (2); Stn. 85-60, Ascen-
cidn Bay, along shore near Suliman Point,
17 Apr 1960 (1); ICML-UNAM: Ascencio6n
Bay, Pajaros Point, 6 Oct 1983 (1); Puerto
Morelos, La Bocana, Francisco Solis-Marin
coll. 17,Feb 199574).

Diagnosis.—Atokous specimen collected
in Littke Cayman complete, light yellow, 14
mm long, 0.5 mm wide, with 83 setigers,
prostomium and palps pigmented. Anterior
median region of each segment with trans-
verse diffusely pigmented stripe. Prosto-
mium slightly longer than wide, with one
pair of frontal small cirriform antennae.
Two pairs of large eyes with a lens, in tra-
pezial arrangement. Palps biarticulate, glo-
bose, with conical palpostyle. Peristomium
with 4 pairs of tentacular cirri, largest pair
extending to setiger 5 (Fig. 1a).

Paragnaths arranged on pharyngeal areas
as follows: I: 0; II: 15 cones in 2 rows; III:
3 in one row; IV: 14 in two rows; V: 0; VI:
4 cones in a row; VII—VIII: 6 in one row.
Jaws yellowish with 7 teeth.

First two parapodia uniramous, third bi-
ramous, notopodia with two ligules, dorsal
one digitiform, short; median ligule larger,
basally wide. Neuropodia with presetal lobe
distally conical, postsetal lobe rounded,
ventral ligule subulate. Dorsal cirrus wide,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

inserted proximally on ligule, larger than
ventral cirrus (Fig. 1b). Sixth parapodium
with dorsal cirrus elongate, with constric-
tion in median part of cirrus, which results
in distal cirristyle. Notopodial median lig-
ule and neuropodial ventral ligule rounded
distally; neuropodial postsetal lobe conical.
Ventral cirri inserted proximally on body
(Fig. 1c). Notopodial ligules of anterior
parapodia posterior to setiger 6 rounded,
shorter than in parapodia anterior to setiger
6. Neuropodial ventral ligule subulate (Fig.
1d). Notopodial ligules of posterior para-
podia triangular, median ligule larger, slen-
derer than dorsal ones. Neuropodial postse-
tal lobes conical distally, presetal one
rounded, ventral ligule slender (Fig. le).

All notopodial setae homogomph spini-
gers with long and slender appendix, finely
spinulate on margin. Supraacicular neuro-
setae from first biramous parapodium, and
from modified setiger 6, homogomph spi-
nigers similar to notopodial ones, and het-
erogomph spinigers with short appendix,
with long spines on margin (Fig. 1f); su-
praacicular neurosetae posterior to sixth
parapodia, homogomph spinigers and het-
erogomph falcigers, with long and slender
appendix, with spinulations along margin,
distally ending in blunt tooth (Fig. 1g).
Neuropodial infraacicular setae from first
biramous parapodium to setiger 5, hetero-
gomph spinigers with short appendix; neu-
ropodial infraacicular seta from modified
setiger 6 throughout body including heter-
ogomph spinigers and falcigers in anterior
parapodia, with latter similar to supraaci-
cular ones, and in median and posterior
parapodia falcigers with short, triangular
appendix, distally ending in curved tooth,
with margin covered by one row of long
spines (Fig. 1h).

Pygidium with pair of slender anal cirri.
Anus terminal.

Epitokous stage.—Epitokous male, 8 mm
long and 1.7 mm wide including parapodia,
with 70 setigers, no coloration pattern.
Body divided into 3 regions: pre-epitocal
formed by two regions: anterior one, from

VOLUME 112, NUMBER 4

Fig. 1.

671

£

Neanthes egregicirrata (atoke): a, Anterior end, dorsal view; b, 3rd seriger, anterior view; c, 6th

setiger, anterior view; d, 10th setiger, anterior view; e, 32th setiger, anterior view; f, Neuropodial heterogomph
spiniger of supraacicular position from setiger 3; g, Neuropodial heterogomph falciger of infraacicular position
from setiger 10; h, Neuropodial heterogomph falciger of infraacicular position from setiger 32. Scale bars: a =

0.5 mm; b—-e = 150 pm; f—h = 15 pm.

setigers 1 to 5, with notopodial and neuro-
podial ventral ligules digitate, neuropodial
postsetal lobes conical. Dorsal cirrus en-
larged, with a small cirrostyle attached near
distal end (Fig. 2b). In setiger 6, as in ato-
kous material, dorsal cirrus modified, wide
proximally slender distally (Fig. 2a). From
setigers 7 to 16, parapodia with notopodium
formed by two triangular ligules, and a no-
topodial digitiform superior lobe (Fig. 2c).
From setiger 17, parapodia modified for

swimming, notopodia with one small la-
mella associated with dorsal cirrus, with
crenulations. Neuropodia with elongate la-
mella associated with the postsetal lobe.
Ventral cirri with two small lamellae (Fig.
2d).

In pre-epitokous setigers notosetae hom-
ogomph spinigers. First 5 setigers with su-
praacicular neurosetae homogomph spini-
gers and heterogomph falcigers with slen-
der appendix (Fig. 2e); infraacicular spini-

672 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fy
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me
a

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Ge |
Pye

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.
oh

-

Fig. 2. Neanthes egregicirrata (epitokous male): a, Anterior end, dorsal view; b, 5th setiger, anterior view;
c, 15th setiger, anterior view; d, 44th setiger, anterior view; e, Neuropodial heterogomph falciger of supraacicular
position from setiger 5; f, Neuropodial heterogomph falciger of supraacicular position from setiger 15. Scale
bars: a = 1 mm; b-d = 150 pm; e-f = 15 pm.

gers heterogomph. Parapodia posterior to cigers in infraacicular position. Setae from
modified setiger with supraacicular neuro- epitocal region paleae with homogomph ar-
setae homogomph spinigers and hetero-_ ticulation, appendix oar-shaped.

gomph falcigers with short and triangular Pygidium with pair of short anal cirri.
appendix (Fig. 2f); only heterogomph fal- Anus terminal.

VOLUME 112, NUMBER 4

Discussion.—The holotype of this spe-
cies was described from an epitokous spec-
imen. Before this study, the atokous form
was unknown. When examining the ato-
kous specimens, it was noted that no hom-
ogomph notopodial falcigers are present,
which means that it should be included in
the genus Neanthes. The dorsal elongate
cirrus in the sixth setiger persists in the ato-
kous phase and as previously thought, is not
exclusive of the epitokes. FE Solis-Marin
(pers. comm.) observed the epitokes in the
water column and reported that they ap-
peared to gyrate one around the other, until
they stopped completely. Apparently, the
modified dorsal cirri are used to hold the
animals to each other during copulation.

In the type material examined from Pla-
tynereis abnormis Horst (1924) (ZMA Pol
730) it was noted that the dorsal cirri are
modified as in N. egregicirrata, except that
this happens on setiger 7, rather than on
setiger 6.

Distribution.—Western Atlantic. Known
from the Caribbean Sea, Quintana Roo,
Mexico, Antigua Island, Puerto Rico, Bar-
bados and Little Cayman Island. This is the
first record for Mexico.

Habitat.—Atokous specimens collected
among corals, epitokes collected close to
the surface with zooplankton net attracting
the organisms with a bright light.

Neanthes micromma (Harper, 1979)

Nereis (Neanthes) micromma Harper 1979:
91, figs. 1-11.

Neanthes micromma: Taylor, 1984:31.17,
figs. 31.14a—h.—Hermdandez-Alcantara &
Solis-Weiss, 1991:255.

Material examined.—ICML-UNAM:
Imca IV expedition, Cayo Arcas, 3 Oct
1989 (4); 4 Oct 1989 (1); Dinamo III ex-
pedition, Cayo Nuevo, 24 Mar 1991 (1).

Distribution.—Amphi-American. West-
ern coasts of Mexico, Gulf of Mexico.

Habitat.—Sandy muds, shallow waters
down to 50 m.

673

Neanthes succinea (Frey & Leuckart,
1847)

Nereis (Neanthes) succinea: Pettibone,
1967:165, fig. 44a-e.—Day 1967:321,
fig. 14a—e.—Hartmann-Schroder, 1971:
200, figs. 64-65.

Neanthes succinea: Hartman, 1945:17, pl.
3, figs. 1-2.—Rioja, 1946:205, pl. 1, figs.
1—2.—1962:165.—Imajima, 1972:108,
fig. 32a—k.—Taylor, 1984:31.17, figs.
31.16a—h.

Material examined.—ZMUA: Margarita,
Punta de Piedra, 21 Jan 1964 (2); Surinam,
Suriname River, wreck of Goslar, 7 Mar
1964 (2); Jamaica, Kingston Harbour, inlet
west of airport, 7 May 1973 (1).

Distribution.—Cosmopolitan in tropical
to temperate waters. Atlantic from the
North sea south to South Africa, and from
Massachusetts south to Uruguay; Pacific
Ocean from California south to Panama; In-
dian Ocean.

Habitat.—Among rhizoids of algae at-
tached to rocks.

Nereis allene Pettibone, 1956

Nereis allene Pettibone, 1956:287, figs. 4b—
f, 6a—h.

Material examined.—Puerto Rico, 15
Aug 1955 (Holotype, USNM 27778); An-
tigua, Jul 1918 (4 Paratypes USNM 27779).
ZMUA: North Bimini, Bahamas, 17 Aug
1949 (56); 18 Aug 1949 (20).

Discussion.—The examined specimens
are epitokes collected with a zooplancton
net in the water column. Those are mor-
phologically similar to the type specimens.
However, in parapodia of setigers 1—6, the
notopodium is slightly modified, with cylin-
drical dorsal cirri. In the original descrip-
tion, Pettibone mentioned that, in those
specimens, paragnaths are present in both
pharyngeal rings, but she does not detail
their distribution. In the specimens collect-
ed in North Bimini the following pharyn-
geal arrangement is found: Area I: 1 cone,
If: 135=15conesi in two: rows; 11: -18=22

674

small cones in an oval; IV: 25 cones in a
crescent, with 3—4 rows; V: 0; VI: 3 cones
in an irregular row; VII—VIII: 5 cones in a
row. Jaws amber colored with 6 stout teeth.

Distribution.—Known only from the
greater Caribbean region, Puerto Rico, An-
tigua, Barbados and Bahamas.

Habitat.—Undescribed for atokous spec-
imens, epitokes collected with zooplankton
net in the water column.

Nereis falsa Quatrefages, 1865

Nereis falsa: Fauvel, 1923:337, fig. 129e—
m.—Day, 1967:317, fig. 14.7k—o (with
synonymy).—Day 1973:41.

Nereis pelagica occidentalis Hartman,
1945:20, pl. 4, figs. 1-6.—1951:46.

Nereis (Nereis) occidentalis: Pettibone,
1956:291, fig. 7a—d, 8a-f.

Nereis occidentalis.—Duefias, 1981:88, pl.
8, fig. a-f.

Material examined.—ZMUA: Bonaire,
Lagoen Southeast, corner, 28 Oct 1930 (3);
Bonaire, Lagoen south, 19 Sep 1967 (10);
Curacao, Piscadera Baai, N., 2nd buoy, 31
Oct 1963 (16); Curacao, Ironbeam of
mouth of sewer-pipe, 31 Oct 1963 (47); Cu-
racao, Piscadera Baai, northeast, 11 Oct
1963 (86); Curacao, Piscadera Baai, north-
east, 25 Nov 1963 (66); Curacao, Piscadera
Baai, northeast, 11 Dec 1963 (2); Curagao,
Piscadera Baai, central part, southeast, 31
Oct 1963 (10); Curacao, Piscadera Baai,
central part, southeast, 13 Dec 1963 (5);
Curacao, Piscadera Baai, central part,
southeast, 18 Dec 1963 (1); Curacao, St.
Jorisbaai, northwest bay, 25 Feb 1970 (46);
Curacao, Piscadera Baai, inner bay north,
islet, 26 Sep 1967 (20); Margarita, Puente
de La Restinga, 11 Jan 1964 (1); Guade-
loupe, Riviere Salée la Manche a Eau, 16
Jul 1967 (8); Jamaica, Kingston Harbour,
inlet west of airport, 7 May 1973 (7).

Distribution.—Warm and tropical Atlan-
tic from France to West Africa and North
Carolina to the Gulf of Mexico; Mediter-
ranean; South Africa to Madagascar.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Habitat.—Rocky bottoms, associated
with algal mats and Rhizophora roots.

Nereis panamensis Fauchald, 1977
(Fig. 3a—g)

Nereis panamensis Fauchald, 1977:29, fig.
6d—i1.—San Martin, 1994:6, fig. 2.—Sa-
lazar-Vallejo & Jiménez-Cueto, 1997:
374.

Material examined.—ZMUA: Klein Cu-
ragao, west shore, 1 Oct 1948 (19); An-
guilla, north sandy ground, 19 Apr 1949
(2); Anguilla, Cocus bay, north, 3 Aug
1973 (5); St. Barts, S. Public, near Gustaria,
4 Jun 1949 (21); Saba, East Fort Bay, 21
Jul 1949 (8) Saba, Covebay at flat point
(tidal area), 5 Oct 1963 (22); Aruba, Mat-
mok, Arasji, 14 Aug 1955 (5); St. Vincent,
Calliwater Bay near Johnson Pt., 10 Jul
1967 (2); Jamaica, Drunkemans Key, 15
Jun 1973 (1);). ICML-UNAM: Imca IV ex-
pedition, Cayo Arcas, 3 Oct 1989 (27); 4
Oct 1989 (23); Dinamo I, III expeditions,
Triangulos Oeste reef, East bay, 19 Mar
1990 (4); 19 Mar 1991 (5); Triangulos Oes-
te reef, West bay, 19 Mar 1991 (2); Trian-
gulos Oeste reef, East bay, 19 Mar 1990
(1); Cayo Arcas, North bay, 10 Mar 1990
(12); 17 Mar 1991 (6); Cayo Arcas, South
bay, 10 Mar 1990 (50); 17 Mar 1991 (12);
Cayo Arcas, Southeast bay, 23 Mar 1991
(117), 10 Mar 1990 (2); Cayo Arcas, South
bay, 17 Mar 1991 (166); Cayo Arcas,
Northeast bay, 17 Mar 1991 (134); Cayo
Arenas, Southwest bay, 21 Mar 1991 (1).

Diagnosis.—Best preserved specimen
complete, 16.5 mm long, 1.5 mm wide, 71
setigers, no pigmentation pattern. Prosto-
mium longer than wide, with pair of digi-
tiform antennae extending from distal re-
gion of prostomium slightly past the palps.
Two pairs of rounded eyes in rectangular
arrangement, anterior pair clearly visible,
posterior pair covered by peristomium.
Palps long with conical palpostyle. Peris-
tomium as long as first two setigers, with 4
pairs of tentacular cirri, longest pair extend-
ing to setiger 3 (Fig. 3a).

VOLUME 112, NUMBER 4

675

Fig- 3:
view; d, 58th setiger, anterior view; e, Notopodial homogomph falciger from setiger 30; f, Neuropodial heter-
ogomph falciger of supraacicular position from setiger 10; g, Neuropodial heterogomph falciger of infraacicular
position from setiger 58. Scale bars: a = 1 mm; b—d = 150 pm; e-g = 15 pm.

Paragnaths arranged on pharyngeal areas
as follows: I: 0; II: 3 cones in a row; III: 3
in a row; IV: 9 3 in 2 rows; V: 0; VI: 3 in
one row; VII-VIII: 6 in one row. Jaws dark
brown with 8 teeth.

In anterior parapodia, notopodia with 2
ligules, dorsal one smaller, small flange rep-
resenting the superior lobe present. Postse-
tal lobe of neuropodia conical distally, ven-
tral ligule wide basally. Dorsal and ventral
cirri subequal (Fig. 3b). Notopodial dorsal
ligule in median and posterior parapodia
represented only by small flange associated

Nereis panamensis: a, Anterior end, dorsal view; b, 10th setiger, anterior view; c, 30th setiger, anterior

with basal region of dorsal cirrus, notopo-
dial median ligule subulate. Neuropodia
with postsetal lobe distally conical, ventral
ligule slender (Fig. 3c—d).

Homogomph notopodial spinigers in an-
terior parapodia substituted in median and
posterior parapodia by homogomph falci-
gers, appendage of latter with two blunt dis-
tal teeth, and small tooth on margin (Fig.
3e). Neuropodia with homogomph spini-
gers and heterogomph falcigers in supra-
acicular position in all parapodia, latter with
slender appendix, strongly spinulate on

676

Margin in anterior parapodia (Fig. 3f); In-
fraacicular ones, including heterogomph
spinigers and falcigers, appendix similar in
anterior parapodia to these of supracicular
falcigers, in posterior ones appendix some-
what smaller, and spinulation on the margin
shorter (Fig. 3g).

Pygidium with pair of long anal cirri.
Anus terminal.

Discussion.—Fauchald (1977) men-
tioned that in this species only one pair of
eyes is present, but in the specimens ex-
amined for this study, a second pair was
observed, covered by the anterior part of
the peristomium.

Salazar- Vallejo & Jimenez-Cueto (1997)
noted that San Martin’s record (1994) for
Cuba, could be Nereis kauderni Fauvel, but
the absence of the notopodial superior lobes
in posterior parapodia suggests that the Cu-
ban record does belong to N. panamensis,
since in N. kauderni two notopodial lobes
in posterior parapodia are present.

Distribution.—Western Atlantic. Known
from the Caribbean region, reported from
Panama and Cuba as well as from Klein
Curacao, Anguilla, St. Barts, St. Vincent,
Aruba, Jamaica and Saba. In this study its
range is extended to the coral islands of the
southern Gulf of Mexico, and it is the first
record for Mexico.

Habitat.—In interstices of coralline rock
fragments.

Nereis pelagica Linnaeus, 1758

Nereis pelagica: Fauvel, 1923:336, fig.
130a—f.—Imajima 1961:85.—1967:422.—
1972:142, fig. 48a—m.—Imayjima & Hart-
man, 1964:147.—Pettibone, 1963:179,
fig. 42d—h.—Day, 1967:315, fig. 14.7f-
j.—tTaylor, 1984:31.42, fig. 31.40a—g.

Material examined.—ICML-UNAM:
IMCA IV expedition, Cayo Arcas, 3 Oct
1989 (7); 4 Mar 1989 (1).

Distribution.—Cosmopolitan. Known
from the Arctic, from Norway to the Med-
iterranean Sea, Azores, western coast of Af-
rica, Atlantic coast of North America from

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Hudson Bay to the Gulf of Mexico, Behring
Sea to Panama, Japan, Kerguelen Islands,
Strait of Magellan, Mexican littoral areas,
recorded from Baja California Norte, Baja
California Sur, Sonora, Sinaloa, and Vera-
cruz.

Habitat.—Rocky substrates, associated
with algal mats, soft and hard bottoms from
intertidal zone down to continental shelf.

Nereis riisei Grube, 1857

Nereis riisei Grube 1857:162.—Monro,
1933:43.—Hartman, 1940:221, pl. 33,
fig. 37.—Fauchald, 1977:31, fig. 8c—e.
Taylor, 1984:31.38, fig. 31.36a—g.

Nereis (Nereis) riisei: Day, 1973:39, fig.
5g—j.—Gardiner, 1976:152; fig. 150-1.

Material examined.—ZMUA: Bonaire,
Kralendijk, near Pasanggrahan, Sep 1930
(1); Anguilla, north sandy ground, 19 Jun
1949 (1); St. Eustatius, south Tumble
Down, Dick Bay, 10 Jul 1949 (1); St. Eu-
statius, Billy Gut, 13 Jul 1949 (8); Aruba,
Seroe Colorado, Oostpunt, 2 May 1955 (8);
Curacao, Piscadera Baai, entrance east, 28
Sep 1963 (10); Curagao, Piscadera Baai,
north, 2nd buoy, 31 Oct 1963 (6); Curagao,
Piscadera Baai, central part, 3rd buoy, 31
Oct 1963 (26); Curacao, Piscadera Baai,
northeast, small bay, 25 Nov 1963 (2); Cu-
ragao, Piscadera Baai, northeast, 11 Dec
1963 (13); Curacao, Piscadera Baai, south-
east part, 18 Dec 1963 (6); the Désirade,
Grande Anse, near breadge, 23 Jan 1964
(1); Barbados, 0.5 mi. off Holetown, 19
Nov 1964 (3); St. Vincent, Calliwater bay,
near Johnson Pt., 10 Jul 1967 (1); Grenada,
Hog Island near Pt. Salines, 8 Jul 1967 (7);
Barbuda, Great Lagoon entrance at Billy
Pt., west, 22 Jul 1967 (2); Antigua, Dick-
inson Bay pier, 28 Aug 1967 (3); Curag¢ao,
St. Joris Baai, south, shore, 23 Oct 1968
(40); Curacao, St. Joris Baai, northwest bay,
25 Feb 1970 (6); Jamaica, Drunkemans
Key, 15 Jun 1973 (2); E Bonet coll., Ala-
cran Reef, 18 Apr 1955 (2); USNM: Smith-
sonian-Bredin Caribbean Expedition IV,
Stn. 52-60, Ascencién Bay, just behind cen-

VOLUME 112, NUMBER 4

ter of Nicchehabin Reef, east of Allen
Point, 10 Apr 1960 (23); Stn. 67-60, As-
cencion Bay, behind central part of Nicche-
habin Reef, 13 Apr 1960 (19); Stn. 72-60,
Ascencion Bay, central part of Nicchehabin
Reef, 14 Apr 1960 (46); Stn. 77-60, north
end of Ascencién Bay, shore, just east of
Halfway Point, 15 Apr 1960 (5); Stn. 83-
60, Ascencién Bay, mangrove inlet behind
Allen Point light, 16 Apr 1960 (1); Stn. 82-
60, behind central part of Nicchehabin
Reef, 16 Apr 1960 (9); Stn. 87-60, north of
Ascencion Bay, southwest of Suliman
Point, 17 Apr 1960 (2); Stn. 91-60, inner
side Nicchehabin Reef, 18 Apr 1960 (1);
Stn. 95-60, Ascencién Bay, Suliman Point,
19 Apr 1960 (10); Stn. 100-60, north of
Santa Maria Point, south end of Cozumel
Island, 21 Apr 1960 (3); Stn. 118-60, San
Miguel, Cozumel Island, 29 Apr 1960 (1).
ICML-UNAM: Imca IV expedition, Cayo
Arcas, 4 Oct 1989 (1); Dinamo [, III ex-
peditions, 10 Mar 1990 (1); Alacran Reef,
West bay, 18 Mar 1990 (10); 23 Mar 1991
(23); Alacran Reef, East bay, 23 Mar 1991
(4); Triangulos Oeste reef, 19 Mar 1991 (7);
Cayo Nuevo, 24 Mar 1991 (1).

Distribution.—Amphi-American. Gulf of
California to Panama, Gulf of Mexico down
to the Caribbean Sea.

Habitat.—Soft bottoms, intertidal zone,
under rocks, among sponges on mangrove
roots, in the interstices of algae-sponges as-
sociation, in algal rhizoids.

Perinereis anderssoni Kinberg, 1866

Perinereis anderssoni Kinberg, 1866:175;
Fauchald, 1977:31, fig. 8a—b; Hartman,
1951 47, pl. 13, fig. 6; de Leén-Gonzélez
& Solis-Weiss, 1998:675, fig. la—g.

Material examined.—MZUA: Bonaire,
Kralendijk, near Pasanggrahan, 5 Sep 1930
(1).

Distribution.—Amphi-American. From
the Gulf of Mexico to Uruguay, and for
Juan Fernandez Island, Chile.

Habitat.—Rocky substrate, associated
with algal mats.

677

Perinereis cariboea de Le6n-Gonzalez &
Solis-Weiss, 1998

Perinereis cariboea de Leoén-Gonzalez &
Solis-Weiss, 1998:677, fig. 3a—e.

Material examined.—ZMUA: Klein Bo-
naire, East coast at landing place, 13 Sep
1948 (1); St. Eustatius, Tumble Down, Dick
Bay, 10 Jul 1949 (1).

Distribution.—Antilles and Mexican Ca-
ribbean area.

Habitat.—Among rocks and in sands, in-
tertidal.

Perinereis cariacoensis Linero-Arana,
1983

Perinereis cariacoensis Linero-Arana 1983:
5, fig. a—e.

Material examined.—ZMUA: Bonaire,
Kralendijk, near Pasanggrahan, 09/1930
(3).

Distribution.—Known only from Vene-
zuela and Bonaire.

Habitat.—Rocky beaches, intertidal.

Perinereis elenacasoae Rioja, 1947

Perinereis elenacasoi: Rioja 1947:531.

Perinereis elenacasoae: Salazar- Vallejo,
1989:50.—de Leoén-Gonzaélez & Solis-
Weiss, 1998:680, figs. 4A—-E, S5A—D.

Perinereis obfuscata: Berkeley & Berkeley,
1960:359 (in part).

Perinereis anderssoni: Rioja, 1961:295,
figs. 12—15 (in part).

Material examined.—ZMUA: Bonaire,
South of Kralendijk, near Hoop, May 1930
(3); Bonaire, Kralendijk, near Pasanggra-
han, 5 Sep 1930 (57); Klein Bonaire, east
coast at landing place, 13 Sep 1948 (2); Bo-
naire, Paloe Lechis of Salinja, 24 Feb 1949
(10); St. Eustatius, Tumble Down, Dick
Bay, 10 Jul 1949 (1); Curacao, Piscadera
Baai, entrance east, 28 Sep 1963 (1); Cu-
racao, Piscadera Baai, north., 2nd buoy, 31
Oct 1963 (2); Curacao, Piscadera Baai, cen-
tral part, 3rd buoy, 31 Oct 1963 (3); Bar-
bados, Conset Bay (St. John), 7 Jul 1967

678

(4); Curagao, St. Joris baai, northwest bay,
25 Feb 1970 (14).

Distribution.—Amphi-American. Mexi-
can Pacific from Puerto Pefiasco, Sonora to
Salina Cruz, Oaxaca; Gulf of Mexico, Ca-
ribbean Sea, Brazil.

Habitat.—Rocky substrates, among rhi-
zoids of algae attached to rocks in break-
waters and in coral rocks.

Perinereis floridana (Ehlers 1868)

Nereis floridana: Ehlers, 1868:503.
Perinereis floridana: de Leén-Gonzalez &
Solis-Weiss, 1998:683, figs. 6a—e, 7a—e.

Material examined.—ZMUA: St. Barts,
South Public, near Gustaria, 4 Jun 1949 (1);
Klein Bonaire, east coast of landing place,
13 Sep 1948 (1); Klein Curacao, west
shore, 1 Oct 1948 (2); Barbados, Conset
Bay (St. John), 7 Jul 1967 (2).

Distribution.—Westerm Atlantic. Known
from the greater Caribbean region.

Habitat.—Associated with coral rocks,
intertidal.

Platynereis dumerilii (Audouin & Milne-
Edwards, 1834)

Platynereis dumerilii: McIntosh, 1885:224,
pl. 35, figs. 7-9, pl. 16A, figs. 14-16.
Day, 1967:306, figs. 14.4d—k.—Fau-
chald, 1977:31, fig. 4d-f.

Material examined.—ZMUA: Bonaire,
Kralendijk, near Pasanggrahan, 5 Sep 1930
(2); Margarita, Punta Mozquito, near Por-
lomar, 4 Jun 1936 (10); Klein Bonaire, east
coast at landing place, 13 Sep 1948 (3);
Klein Bonaire, west shore, 1 Oct 1948 (5);
Bonaire, Paloe Lechis of Salinja, 24 Feb
1949 (1); St. Barts, South Public, near Gus-
taria, 4 Jun 1949 (5); St. Eustatius, South
Tumble Down, Dick Bay, 10 Jul 1949 (25);
St. Eustatius, Billy Gut, 13 Jul 1949 (84);
Florida, east of Soldier Key, 5 Sep 1963
(7); Puerto Rico, Parguera, Mata de the
Gata, 12 Sep 1963 (5); Curagao, Piscadera
Baai, central Part, 3rd buoy, 31 Oct 1963
(2); Curacao, Piscadera Baai, north, 2nd

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

buoy, 31 Oct 1963 (1); Curacao, Piscadera
Baai, northeast, 11 Oct 1963 (1); Margarita,
Puente de La Restinga, 11 Jan 1964 (2);
Barbados, Conset Bay (St. John), 7 Jul
1967 (34); Guadeloupe, Riviere Salée, La
Manche a Eau, 16 Jul 1967 (1); Antigua,
Dickinson Bay pier, 28 Jul 1967 (2); Bo-
naire, Lac entrance, 200m west of Cai, 11
Aug 1967 (3); Curagao, St. Jorisbaai, north-
west bay, 25 Feb 1970 (3); Jamaica, Drun-
kemans Key, 15 Jun 1973 (15); Anguilla,
Cocus Bay, north, 3 Aug 1973 (14);
USNM: Smithsonian-Bredin Caribbean Ex-
pedition IV, Stn. 34-60, San Miguel, Coz-
umel Island, near anchorage northwest of
main dock, 03 Apr 1960 (3); Stn. 35-60,
Espiritu Santo Bay, 5 Apr 1960 (9); Stn.
44-60, north end of Ascencién Bay, small
inlet behind Allen Point, 7 Apr 1960 (2);
Stn. 52-60, Ascencion Bay, just behind cen-
ter of Nicchehabin Reef, east of Allen
Point, 10 Apr 1960 (2); Stn. 60-60, Allen
Point, Ascenciédn Bay, where last sand
beach adjoins mangroves, 12 Apr 1960 (1);
Stn. 61-60, near Allen Point, 12 Apr 1960
(1); Stn. 72-60, Ascencion Bay, central part
of Nicchehabin Reef, 14 Apr 1960 (2); Stn.
82-60, behind central part of Nicchehabin
Reef, 16 Apr 1960 (2); Stn. 87-60, north of
Ascencion Bay, 200—300 yds. southwest of
Suliman Point, 17 Apr 1960 (2); Stn. 91-
60, inner side, Nicchehabin Reef, 18 Apr
1960 (10); Stn. 95-60, Ascencion Bay, Su-
liman Point, 19 Apr 1960 (11); Stn. 100-
60, north of Santa Maria Point, Cozumel
Island, 21 Apr 1960 (1). ICML-UNAM:
Dinamo I expedition, Cayo Arcas, North
bay, 10 Mar 1990 (1).

Distribution.—Cosmopolitan in tropical
and subtropical waters.

Habitat.—Associated with hard sub-
strates between algal mats. Specimens
forming mucous tubes were collected in
brown algal fronds.

Pseudonereis gallapagensis Kinberg, 1866

Pseudonereis gallapagensis: Hartman,
1940:231.—Fauvel, 1953:215, fig. 110a—

VOLUME 112, NUMBER 4

c—Rioja 1961:297.—Wesenberg-Lund,
1962:84, fig. 32.—Imajima, 1972:97, fig.
28a—j.—Fauchald, 1977:32, fig. 4g—h.—
Rozbaczylo & Bolados, 1980.—219, fig.
a—d.—Wut et al., 1985:220, fig. 124a—f.—
Bastida-Zavala, 1993:30.

Material examined.—ZMUA: Bonaire,
Lagoen, southeast corner, 28 Oct 1930 (1);
Margarita, Punta Mozquito, near Porlamar,
4 Jun 1936 (1); Klein Curacao, west shore,
1 Oct 1948 (2); St. Eustatius, South Tumble
Down, Dick Bay, 10 Jul 1949 (34); Saba,
East Fort Bay, 21 Jul 1949 (36); Aruba,
Seroe Colorado, Oostpunt, 2 May 1955
(14); Saba, Covebay at flat point, 5 Oct
1963 (9); Curacao, Piscadera Baai, central
part, 3rd buoy, 31 Oct 1963 (1); Grand
Cayman, south of Jackson’s Point, 9 Jun
1973.G3).

Distribution.—Cosmopolitan in tropical
to temperate waters Galapagos Islands,
Peru, Chile, Panama, Hawaii Islands, Mar-
shall Islands, Samoa, New Caledonia, Chi-
na, Japan, India, Sri Lanka, Madagascar,
Cape of Good Hope, Cameroon, Brazil and
Gulf of Mexico.

Habitat.—Associated with rocky bot-
toms, and coral substrates.

Stenoninereis martini Wesenber-Lund,
1958

Stenoninereis martini Wesenberg-Lund,
1958:9, fig. 2—4.—Pettibone, 1971:39,
figs. 23—24.—Williams et al., 1976:83.—
Hartmann-Schroder, 1977.—Gardiner &
Wilson, 1979:165.—Hernandez-Alcan-
tara & Solis-Weiss, 1991:251.—1995:
117.—de Leén-Gonzdalez & Solis Weiss,
1998:199, fig. la—d.

Material examined.—ZMUA: St. Martin,
Devils Hole Swamp, 14 Oct 1963 (32).

Distribution.—Western Atlantic greater
Caribbean region (type locality St. Martin
Island and Sarasota, Florida), western Gulf
of Mexico (Texas), Cuba, North Carolina
and Laguna de Términos, Campeche.

679

Habitat.—Soft bottoms, associated with
roots of grassbeds.

Acknowledgments

We would like to thank Harry ten Hove
and J. Bleeker (ZMA), Kristian Fauchald
and Linda Ward (USNM), Karin Sindemark
(RNS) for their help in making available for
examination the material under their care.
Thanks are also due to the personnel of the
Laboratorio de Ecologia Costera and others
from the Imca-Dinamo expeditions who
helped in the collection of the organisms of
the Coral islands if the Gulf of Mexico. We
also thank the anonymous reviewers whose
comments greatly improved the manuscript.
Part of this study was funded by the ““Com-
isi6n Nacional para el Conocimiento y uso
de la Biodiversidad’”” (CONABIO, Project
H-011).

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(4):682—686. 1999.

A new stygobitic Calanoida (Crustacea: Copepoda) of the genus
Stygodiaptomus Petkovski, 1981 from the Balkan Peninsula

Tomislav Karanovic

Institute of Marine Biology, PO. Box 69, 85335 Kotor, Montenegro, Yugoslavia

Abstract.—A new species of the genus Stygodiaptomus Petkovski, 1981 is
described. The genus now includes three species, all endemic to the Balkan
Peninsula: S. kieferi Petkovski, 1981, S. petkovskii Brancelj, 1991, and S. ferus
n. sp. The male of S. ferus can be distinguished from males of the other species
by the setation of swimming legs 1—4, and morphology of the exopodite and
endopodite of leg 5. The diagnosis for the genus Stygodiaptomus is revised.

There are only a few true stygobitic spe-
cies of calanoid copepods. Most reports of
calanoid copepods from freshwater subter-
ranean habitats are of one or a few speci-
mens of epygean species, which are col-
lected in subterranean habitats accidentally.
Bowman (1986) cited only eight stygobitic
freshwater calanoids of the world. In Eu-
rope, until now, five species have been de-
scribed. Borutzky (1962) described Speo-
diaptomus birsteini from Crimea as a new
species and new genus in a new subfamily,
Speodiaptominae. Dussart (1970) described
Spelaeodiaptomus rouchi from southern
France as a new species and new genus in
the recognized subfamily Diaptominae, and
Kiefer (1978) redescribed the species.
Three other stygobitic species of freshwater
calanoids are known from the Dinaric Alps.
Petkovski (1978) described Troglodiapto-
mus Sketi as a new species and genus in the
subfamily Speodiaptominae. Subsequently
it has been recorded by Stoch (1984), Pet-
kovski (1984), and Brancelj (1987, 1991)
from other localities in the northern and
central Dinaric Alps. Petkovski (1981) de-
scribed Stygodiaptomus kieferi as a new
species and new genus in the subfamily
Diaptominae. Brancelj (1991) collected it
again from the type locality in central the
Dinaric Alps, and described Stygodiapto-

mus petkovskii as a new species from two
caves in northern Dinaric Alps. During an
investigation of the copepod fauna in Mon-
tenegro, one male specimen of a new spe-
cies of the genus Stygodiaptomus was col-
lected in Skadar Lake Valley (southern Di-
naric Alps).

In the description, standard abbreviations
of the characters are used as follows: Al,
antennule; A2, antenna; Md, mandible;
Mxp, maxilliped; P1—4, first to fourth
swimming legs; P5, fifth leg.

Family Diaptomidae Baird, 1850
Subfamily Diaptominae Kiefer, 1932
Genus Stygodiaptomus Petkovski, 1981
Stygodiaptomus ferus, new species

Material examined.—One male (Holo-
type) from a small lake in the cave Sutim-
ska Jama (42°25'50’N, 19°10’40’E), near
the town Podgorica, Montenegro, coll. T.
Karanovic, 18 Sep 1997. The specimen is
completely dissected, mounted on a slide in
Faure’s medium, and deposited in the au-
thor’s collection (No. 8/60/0584/c) at the
Institute of Marine Biology, Kotor, Monte-
negro.

Description. —Male (Holotype): Length,
including furcal rami (excluding furcal se-
tae), 1.18 mm. Body colorless; without in-
tegumental windows, or paired sensila on

VOLUME 112, NUMBER 4

the lateral extensions of thoracic somite 6
or on the right side of the genital somite).
Naupliar eye absent. Cephalothorax subcy-
lindrical; about 1.4 times as long as wide,
equaling 36.5% of body length (Fig. 9).
Five thoracic somites of unequal width and
length. Urosome cylindrical, 5-segmented,
and dorsoventrally symmetrical (Fig. 10).
Genital somite small. Caudal rami smooth,
subcylindrical, and about 2.6 times longer
than wide.

Left Al 25-segmented, reaching to mid-
dle of the furca, with cylindrical aesthe-
tascs on articulating segments 1, 2, 3, 5, 9,
12, 14, and 19 (Fig. 12). Right Al 23-seg-
mented (Figs. 6, 7, and 8), prehensile, with
aesthetascs on articulating segments 1, 2,
4,5, 7, 9, 12, 14, 18, and 23. Strong spines
on segments 10, 11, and 13, as well as
small ones on segments 7 and 12 (Fig. 6).
Segments 15 and 16 with an attenuation on
inner margin (Fig. 7). A2 with 2-segment-
ed endopodite, and 7-segmented exopodite
(Fig. 13). Md with 2-segmented endopo-
dite, and 4-segmented exopodite (Fig. 14).
Basis bears 4 setae; coxa with a sharp tooth
separated from a number of small teeth.
Maxillule and maxilla typical for the sub-
family. Maxilliped slender, with 3 plumose
setae on the basis, and 8 naked setae (2
long, and 6 small) on the coxa (Fig. 11).

All swimming legs without setae on the
inner margin of the coxa. Pl with 3-seg-
mented exopodite, and 2-segmented endo-
podite (Fig. 5). Exopodite of P1 with outer
spine only on distal segment; first exopodite
segment without inner seta. P2—4 almost
identical (Figs. 2, 3, and 4); exopodite with
outer spines on middle and distal segments;
proximal segment unarmed. Middle endop-
odite segment of P2 without Schmeil’s or-
gan. Spine and setal formula on exopodite
and endopodite of P1—P4 as follows (num-
ber to left of first slash refers to the inner
spines or setae; number to right of second
slash refers to the outer spines or setae;
number between two slashes refers to the
apical spines or setae):

683

exopodite endopodite

segments 1 Zz 3 1 2 3

Pl 0/0 1/0 2/2/1 1/0 2/2/)0 —
P2 OO T/T 2/2/72 Alo “Th: -2/2/0
PS O70 I 27272 TWO LO 2/20
P4 DOSAM 22D WO. 1/0 22/0

Right P5 basis without hyaline mem-
brane and almost as long as wide (Fig. 1).
Proximal segment of exopodite as long as
wide and with rounded distal-lateral corner.
Distal segment of exopodite about 2 times
longer than wide, with long lateral spine at
3/4th of its length, and a very long apical
claw. Endopodite of right P5 undifferenti-
ated. Left P5 with quadriform basis and 2-
segmented rami. Distal exopodite segment
bears 2 digitiform processes. The inner pro-
cess is naked and longer than outer, while
the outer one has 5 teeth on inner margin
(Piper).

Remarks.—Stygodiaptomus ferus is
clearly distinguished from the other two
species in the genus Stygodiaptomus by the
following: absence of armature elements on
the first exopodite segment of P1—P4; pres-
ence of an outer spine on the second exo-
podite segment of P2; undifferentiated en-
dopodite on male right P5; and basal seg-
ment of the exopodite on male right P5 with
rounded distal-lateral corner. Other differ-
ences between S. kieferi Petkovski, 1981,
and S. ferus n. sp. are: 2-segmented exo-
podite of Pl in S. kieferi (in S. ferus 3-seg-
mented); inner margin of basis on male
right P5 in S. kieferi with a strong spine
(not developed in S. ferus); and completely
fused 22nd and 23rd segments of the male
right Al in S. kieferi (not fused in S. ferus).
Stygodiaptomus ferus n. sp. is more similar
to S. petkovskii Brancelj, 1991; in addition
to the differences noted above they differ
only in details in armature of the male right
Al, and male P5. A close relationship
among these three species is suggested by
the shape of male P5, Al, and identical ar-
mature of the P1—P4 endopodites.

684 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

2

Figs. 1-5. Stygodiaptomus ferus, n. sp., Holotype: 1, P5; 2, P4; 3, P3; 4, P2; 5, Pl. Scale bars = 0.1 mm.

6.7810

Figs. 6-10. Stygodiaptomus ferus, n. sp., Holotype: 6, right Al (segments 1—13); 7, right Al (segments 14—
18); 8, right Al (segments 19-23); 9, habitus, dorsal view; 10, abdomen, dorsal view. Scale bars = 0.1 mm.

VOLUME 112, NUMBER 4

11

SS

—
SS
12

11,13

685

Figs. 11-14. Stygodiaptomus ferus, n. sp., Holotype: 11, Mxp; 12, left Al; 13, A2; 14, Md. Scale bars =

0.1 mm.

Etymology.—The specific name is from
the Latin adjective ferus, which means
wild, agreeing in gender with the masculine
generic name.

Geographic distribution.—The new spe-
cies is known only from the type locality.
We presume that it inhabits a wide area of
Skadar Lake Valley in southeastern Mon-
tenegro and northwestern Albania.

Habitat notes.—Samples collected on
two other occasions (3 Feb 1997, and 10
Aug 1998) from the type locality and earlier
from more than 300 other localities in Mon-
tenegro did not yield additional specimens
of the new species, or any other stygobitic
calanoid. This suggests that either S. ferus
is a very rare species, or that a cave envi-
ronment is not the optimal habitat for it. We
believe that its optimal habitat may be the
large artesian reservoir which is located un-
der Skadar Lake Valley. Unfortunately ar-
tesian waters have not yet been investigated
in Montenegro.

Revised diagnosis for the genus Stygo-
diaptomus.—Diaptomids with 2- or 3-seg-
mented endopodite of Pl. Second segment
of P2—P4 endopodites each with only 1 seta
on the inner margin. Terminal segments on
exopodites P2—P4 with 5 setae and 1 spine.
Terminal segments of endopodites P2—P4
with 4 setae. All swimming legs without
setae on the inner margin of the coxa. Male
right P5 with 1-segmented, very small, or
completely reduced endopodite. Male right
Al with short and strong spines on seg-
ments 10, 11, and 13, as well as a smaller
one on 12th segment. This diagnosis sepa-
rates the genus Stygodiaptomus from the
most similar genus Spelaeodiaptomus, as
well as from all other genera in the subfam-
ily Diaptominae.

Key to the species of Stygodiaptomus

1. Right male P5 with smooth inner margin
on basis

686

Right male P5 with a strong spine on

inher. marci Or Basis .... oSars ws le

1: deca: Sac eee eer S. kieferi Petkovski, 1981
2. First exopodite segment of P2—P4 with

Spihe Of OUTEr Mareing 2.3.5 <5. 96

io 6 Uae S. petkovskii Brancelj, 1991

First exopodite segment of P2—P4 with-

out spine on outer margin .. S. ferus n. sp.

Literature Cited

Borutzky, E. V. 1962. Pervoe nahozhdenie troglobion-
ta iz Calanoida (Crustacea, Copepoda) v pod-
zemnyh vodah.—Doklady Akademii Nauk
SSSR 147:1499-1502.

Bowman, T. E. 1986. Copepoda: Calanoida. Pp. 295—
298, in L. Botosaneanu, ed., Stygofauna Mundi,
a faunistic, distributional, and ecological syn-
thesis of the world fauna inhabiting subterra-
nean waters (including the marine interstitial).
E. J. Brill/Dr. W. Backhuys, Leiden, 740 pp.

Brancelj, A. 1987. Cyclopoida and Calanoida (Crus-
tacea, Copepoda) from the Postojna-Planina
Cave System (Slovenia).—Bioloski vestnnik,
Ljubljana 35:1—16.

. 1991. Stygobitic Calanoida (Crustacea: Co-

pepoda) from Yugoslavia with the description

of a new species Stygodiaptomus petkovskii

from Bosnia and Hercegovina.—Stygologia 6:

165-176.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Dussart, B. H. 1970. Un nouveau Calanoide en eaux
souterraines (Crustace, Copepode).—Annales
de Speleologie 25:155—159.

Kiefer, E 1932. Versuch eines Systems der Diaptomi-
den (Copepoda Calanoida).—Zoologische Jahr-
bericht, Systematik, Jena 63:451—520.

. 1978. Das Zooplankton der Binnengewasser;
Freilebende Copepoda.—Die Binnengewasser
26: 1-343.

Petkovski, T. K. 1978. Troglodiaptomus sketi n. gen.,
n. sp., ein neuer Hohlen-Calanoide vom Karst-
gelande Istriens (Crustacea, Copepoda).—Acta
Musei Macedonici Scientiarum Naturalium,
Skopje 15:151—165.

. 1981. Stygodiaptomus kieferi n. gen. et n. sp.,

zweiter Hohlen-Calanoide vom Dinarischen

Karstgebiet (Crustacea, Copepoda).—Fragmen-

ta Balcanica Musei Macedonici Scientiarum

Naturalium, Skopje 11:63—74.

. 1984. Neue und seltene Copepoden (Crusta-
cea) aus Jugoslawien.—Acta Musei Macedonici
Scientiarum Naturalium, Skopje 17:135—164.

Sars, G. O. 1903. Copepoda Calanoida.—An Account
of the Crustacea of Norway with short descrip-
tions and figures of all the species 4:145-171,
pls. 97-102, suppl. pls. 1—6.

Stoch, FE 1984. Sulla presenza di Troglodiaptomus ske-
ti Petkovski, 1978 (Copepoda, Calanoida) in
una grotta del Carso Triestino (Italia Nordorien-
tale).—Atti e Memorie della Commissione
Grote ““E. Boegan”’ 23:65—67.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(4):687—694. 1999.

The freshwater centropagid Osphranticum labronectum Forbes, 1882
(Crustacea: Copepoda: Calanoida) in Mexico with description of a

new subspecies

M. A. Gutiérrez-Aguirre & E. Suarez-Morales

El Colegio de la Frontera Sur (ECOSUR), A.P. 424, Chetumal, Quintana Roo 77000, Mexico

Abstract.—From a zooplankton survey carried out during January 1998 in
several ponds of the Mexican state of Tabasco, southern Gulf of Mexico, sev-
eral specimens of the freshwater centropagid copepod Osphranticum labronec-
tum Forbes, 1882 were collected. This is a widespread species in North Amer-
ica. There is a single record south of 25°N, in Guatemala (ca. 15°N). The
species has not been reported from the large area lying between these records,
mostly represented by Mexico. This finding in southeast Mexico is helpful to
understand the distributional range of the species. It is also the first record of
a freshwater species of the Centropagidae for Mexico. The Mexican specimens
show morphological differences when compared with previously described ma-
terial, mainly on the armature of male and female fifth legs, and in their size.
These differences, and the assumed isolation of the Mexican populations from
the species main distribution area in the Nearctic region, suggest that the Mex-
ican population is a new subspecies, O. labronectum mexicanum. Taxonomic
illustrations, including structures not depicted or described before for the spe-

cies, are also provided herein.

The calanoid copepod family Centropa-
gidae is represented in the Americas by six
freshwater genera (Parabroteas Mrazek,
1901; Limnocalanus Sars, 1863; Osphran-
ticum Forbes, 1882; Boeckella De Guerne
& Richard, 1889; Neoboeckella Bayly,
1992a; Sinodiaptomus Kiefer, 1932). The
genus Pseudoboeckella Mrazek, 1901 is
now considered a synonym of Boeckella
(Bayly, 1992b). However, only Limnoca-
lanus, Sinodiaptomus, and Osphranticum
are known to be distributed in North Amer-
ica (Williamson 1991, Bayly 1992a, Reid
& Pinto-Coelho 1994).

Osphranticum labronectum, the only
species of this genus, is one of the most
widely distributed freshwater centropagids
in the United States and Canada (Wilson &
Yeatman 1959, Williamson 1991). This spe-
cies was described originally by Forbes
(1882) from material collected in Lake
Michigan. It is mainly a Nearctic copepod,

reported from several localities of the Unit-
ed States and Canada (Marsh 1933; Wilson
& Yeatman 1959; Dussart & Defaye 1983,
1985; Bayly 1992a). Its southernmost rec-
ord in North America is in the Everglades,
Florida (Reid 1992). South of this, there is
one record from Guatemala (Juday 1915,
Reid 1990) which was overlooked by Fer-
nando & Smith (1982) in their revision of
the Central America freshwater copepod
fauna.

From a zooplankton survey carried out
to investigate the copepod diversity of
freshwater environments of the state of Ta-
basco, southeastern Mexico, several speci-
mens of this copepod were recorded. A
geographic record of this species is pre-
sented along with illustrations of the ma-
terial examined, and the description of a
new subspecies, O. labronectum mexica-
num.

Zooplankton was collected 12 January

688

1998, during a survey of several aquatic en-
vironments in the state of Tabasco, south-
eastern Mexico. Samples were collected us-
ing a standard plankton net with a 0.072
mm mesh, deployed in near-shore plankton
trawls. The zooplankton specimens were
fixed in 4% formalin, and then processed
for identification. The taxonomically rele-
vant structures for the identification of these
copepods, and those not previously depict-
ed were illustrated with the aid of a camera
lucida. Descriptions are provided of the ce-
phalic appendages. Specimens were depos-
ited in the collection of the National Mu-
seum of Natural History (USNM), Wash-
ington, D.C., in the the Muséum National
d’ Histoire Naturelle (MNHN), and in the
zooplankton collection of El Colegio de la
Frontera Sur, Chetumal (ECO-CHZOO).

Family Centropagidae
Osphranticum labronectum Forbes, 1882
O. labronectum mexicanum, new
subspecies
Figs. 1-13

Material examined.—Holotype: One
adult female, ethanol-preserved. Laguna
Matillas, Tabasco, Mexico, 12 Jan. 1998,
USNM 288458. Paratypes: Six adult fe-
males from same site, deposited in MNHN-
Cp1777. One adult male in the same insti-
tution (MNHN-Cp1776). One adult female
and three adult males from same locality,
12 Jan 1998, ethanol-preserved, access
number ECOCH-ZOO-00463. Two adult
males from same locality, USNM 288459.

Type locality—Laguna Matillas, Tabas-
co, southeastern Mexico: 17°53’'45.8’N,
92°31'19.6"W, 12 Jan 1998. Environmental
conditions of the sampling site associated
with this record were: temperature: 27.7°C;
pH::6.32, oxygen: 7-2, ppt.

General description.—Body shape and
proportions as described and depicted by
Marsh (1933) and Dussart & Defaye
(1995). Average length (n = 5) of females:
1.44 mm, cephalothorax 0.97 mm long,
0.15 mm wide. Antennule length 1.06 mm.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Genital somite 0.14 mm long, 0.15 mm
wide. Average length (n = 5) of males:
1.26 mm, cephalothorax 0.85 mm long,
0.34 mm wide. Antennules 0.79 mm. Gen-
ital somite 0.07 mm long, 0.09 mm wide.

Description of cephalic appendages.—
Female: Antennules (Fig. 4) relatively
short, 24-segmented, reaching posterior
margin of third pediger (Fig. 3), longer in
some specimens. Spiniform seta on first
segment long, stout. Long setae on seg-
ments 17, 19, and 22. Appendages per seg-
ment as follows (numerals = segment, nu-
merals between brackets = number of se-
tae, ae = aesthetasc, sp = spine): 1(sp),
2(2+2ae),. 33); 40) « SC ae oer
7(2+ae), 8(1), 9(2+ae), 10(1), 11-16(2+ae),
17(2),._ 18(1 Fae), 19@-+ae), 20D
22(2), 23(2+ae), 24(5+ae) (Fig. 4).

Male: Antennules 22-segmented, left an-
tennule as in female, right antennule (Fig.
2) geniculated between segments 18-19,
segments 13-17 relatively wider. Strong
muscular striation on segments 14 and 15.
Appendages per segment as follows: 1(sp),
2(1+3ae), 3(1+2ae), 4(1), 5(1+2ae), 6(1),
7(2+ae), 81), 9@+ae), IOGear
13(@2+ae), 14 and 150. +2ae); 16@ Fae
and 18(1+ae), 19(1+ae), 20(2), 21(2+ae),
22(5+ae).

Antennae with exopod longer than en-
dopod (Fig. 5). Coxa with one long seta.
Basis wide, with two subequal setae on out-
er distal margin. Endopod with two seg-
ments, proximal with two setae on distal 1/
3. Distal portion of terminal endopodal seg-
ment with two lobes, internal with row of
hairlike spinules, with five setae, one small-
er than others. External lobe with one short
and seven long setae. Exopod 8-segmented,
with one seta on first segment, two on sec-
ond, and one seta each on segments 3-7.
Distal segment with one short seta on prox-
imal 1/3, and three long terminal subequal
setae.

Mandible (Fig. 6) with seven bluntly-
pointed teeth on gnathobase and a movable
tooth at the tip. Basis with three setae; en-
dopod of two segments, proximal segment

VOLUME 112, NUMBER 4 689

Le
4 We

Y
LM Ne
a ZA,
Zit

¥Y
wa Z,

Osphranticum labronectum mexicanum. Adult male, Laguna Matillas, Tabasco, Mexico: 1, Hab-

Figs. 1-5.
itus, dorsal; 2, right antennule; Adult female. 3, habitus, dorsal view; 4, right antennule; 5, antenna.

690

with four setae on distal portion of outer
margin; distal segment slightly longer, with
7 subequal terminal setae plus one posterior
seta. Exopod 5-segmented, with a 1,1,1,1,2
setation pattern.

Maxillule (Fig. 7) with praecoxal arthrite
with 15 spiniform setae, two anterior, nine
terminal, four posterior. Coxal epipodite
with nine setae, inner two smaller and thin-
ner than the others, coxal endite with four
setae. Basis with one internal lobe bearing
four setae, basal exite with one seta. Basal
endite with five setae. Endopod 3-segment-
ed, articulating with basis, setation pattern
as: 4,4,7. Exopod with 12 setae, three of
five proximalmost larger, plumose.

Maxilla (Fig. 8) indistinctly segmented,
with two praecoxal and two coxal lobes,
and a well developed basal lobe. Setation
pattern of five lobes as: 4, 3 (first and sec-
ond praecoxal endites), 3, 3 (first and sec-
ond coxal endites), 3 (basal endite); endo-
pod 4-segmented, with setation pattern:
ee Byes

Maxilliped (Fig. 9) well developed. Coxa
fused with praecoxa, with anterior protu-
berance projecting over next segment, with
row of short spinules surrounding process.
Coxa with 3 distinct lobes, proximalmost
low, with two setae, second with three, third
with four. Basis with group of three setae
on middle of inner margin. Endopod 6-seg-
mented, with first segment partially fused
to basis, bearing two subequal setae. Sec-

ond and third endopodal segments with four -

subequal setae, fourth with three, fifth with
four; terminal segment with four subequal
setae.

Caudal rami (Fig. 10) structure and pro-
portions as described and depicted by
Marsh (1933), furcal seta relatively long,
length of ramus/setal length ratio: 1.75.
Other caudal seta as depicted by Marsh
(1933).

Male fifth legs (Figs. 11, 12).—As de-
scribed by Marsh (1933), Bayly (1992a),
and Dussart & Defaye (1995). Relative
length ratios of lateral spine on terminal
exopodal segment, and of the outermost ter-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

minal spine of same segment as presented
in Table 1. Inner rounded protuberance on
terminal exopodal segment relatively low
and not pronounced.

Female fifth legs (Fig. 13).—As de-
scribed by Marsh (1933), Bayly (1992a),
and Dussart & Defaye (1995) but differs in
the relative length of the spines on first and
second exopodal segments, the innermost
apical spine, and the spine on third endo-
podal segment (see Table 1).

Etymology.—The subspecific epithet re-
fers to the country in which the specimens
were collected and for which they represent
the first record of a freshwater centropagid.
The gender of the genus and species names
is neuter, and so is the corresponding Latin
form used on the subspecies name.

Remarks.—The species can be readily
distinguished from most genera of fresh-
water Centropagidae by having, in the fifth
legs, 3-segmented endopodites on both
sides in males and females. Additionally,
the male left distal exopodal segment has
an inner rounded protuberance at its base
(Fig. 12). The caudal rami in both sexes are
less than 2.5 times as long as maximum
width (Wilson & Yeatman 1959, Bayly
1992a). The right caudal ramus has the sec-
ond inner terminal seta clearly wider and
longer than the others (Williamson 1991).
Several structures, including the antennules,

-antennae, and mouthparts (Figs. 2, 4—9) are

illustrated here; some of these appendages
have not been depicted before.

We detected some differences in the
Mexican specimens with respect to the il-
lustrations of the species by Marsh (1933),
Wilson & Yeatman (1959), Dussart & De-
faye (1995), and Bayly (1992a), all from
material of the Nearctic region. The relative
length of the spines of both the female and
male fifth legs are different in both groups
of specimens (see arrowed spines on Figs.
12, 13), as shown in Table 1. The antennule
relative length is 70.8% of the total body
length and 89.5% of the cephalothorax
length in Wilson & Yeatman (1959, fig.
20.29). It is 63.5%, and 85.7%, respective-

VOLUME 112, NUMBER 4 691

Figs. 6-13. Osphranticum labronectum mexicanum. Adult female, Laguna Matillas, Tabasco, Mexico: 6,
mandible; 7, maxilla; 8, maxillule; 9, maxilliped; 10, urosome, dorsal view; 11, male right fifth leg; 12, male
left fifth leg; 13, female fifth leg.

692

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1—Comparison of length ratios of some morphological features described for Osphranticum labro-
nectum in the Nearctic region, and O. labronectum mexicanum from Tabasco, Mexico. P5 = fifth leg.

Wilson & Yeatman Dussart & Defaye Marsh Specimens from
Features (1959) (1995) (1993) Tabasco

Female P;:Re 1 spine/Re2 0.87 0.77 0.85 LSz
Female P;:Re 2 outer spine/Re 2 0.78 0.74 0.69 1k
Female P; : apical inner spine/Re 3 L.87 1.26 1.78 eh
Female P; : spine on Ri 3/Ri 3 1.0 0.83 0.6 1,23
Female furca : dorsal seta/furcal ra-

mus 0.40 0.75 0.41 TS
Male P; Re 2 lateral spine/Re 2 @:53 0.52 0.56 0.68
Male P; Re: apical outer spine/api-

cal medial and inner spines f 1 1 Le
Male P; Re : process on inner mar- high, high, high, low, not

gin pronounced pronounced pronounced pronounced

ly, in Dussart & Defaye (1995, fig. L7). Al-
though relatively shorter in the figure by
Dussart & Defaye (1995), in both, anten-
nules do not reach beyond the posterior
margin of the fourth pediger. In our speci-
mens the corresponding relative lengths are
ca. 67% (60-73.9%), and ca. 95% (78.9—
more than 100%). Antennules are relatively
longer in our specimens, reaching at least
the posterior margin of the third pediger,
but in some they reach the second uroso-
mite. However, the variability of this char-
acter seems to make it taxonomically un-
important.

The dorsal furcal setae are much longer
in the Tabasco specimens (1.6—1.7 the
length of the furcal rami versus 0.4—0.7)
than in the North American material. This
character is not as variable as the antennule
length and could be used to recognize the
subspecies.

Both, our female and male specimens
body size (1.4 mm, 1.26 mm, respectively)
seem to be well below the species range
when compared with the measurements re-
ported by Wilson & Yeatman (1959) for
specimens from the United States (total
length: females 1.7—2.5 mm; males: 1.4—2.3
mm). As found in the Tabasco material,
males tend to be smaller than females.

The Tabasco specimens show consistent
but slight differences when compared with
illustrations of North American material.

These differences, which include secondary
sexual structures such as the male and fe-
male fifth legs, the smaller size, the relative
length of the dorsal furcal seta, and the geo-
graphic isolation of the Mexican population
(and probably that of Guatemala), seem to
be enough evidence to justify the erection
of a new subspecies of O. labronectum. Of
course, further studies on comparative mor-
phology and even interbreeding experi-
ments should be carried out in order to de-
termine the status of the neotropical popu-
lations of O. labronectum.

The species record permits a southward
extension of the known latitudinal distri-
bution of this species into the tropical zone
of Middle America (17°N), from the sub-
tropical Florida area (30°N). It also repre-
sents a modest northwards range extension
from the two localities in which it was re-
corded in two localities of Guatemala, Los
Amates (15°16'N, 89°05’) and Puerto Ba-
trios (15°43’N, 88°36’W), both close to the
Gulf of Honduras, on the Atlantic coast (Ju-
day 1915). The present record consolidates
the regional distributional range of this spe-
cies in the region (Fig. 14).

The present record of O. labronectum
represents the first report of a freshwater
representative of the family Centropagidae
in Mexico (see Sudrez-Morales & Reid
1998). Osphranticum labronectum can be
collected in littoral areas, contrasting with

VOLUME 112, NUMBER 4 693

Osphranticum labronectum

@4

9
4
®
9@
4@
4@ 046
PACIFIC OCEAN
e\8
GULF OF a
MEXICO men
2S nip
10 >
© 457
f = CARIBBEAN SEA

Fig. 14. Records of O. labronectum and authors reporting the species from different areas of North America.
1. Forbes (1882); 2. Herrick (1895); 3. Juday (1915); 4. Marsh (1933); 5. Wilson & Yeatman (1959); 6. Dussart
& Defaye (1983); 7. Reid (1990); 8. Reid (1992); 9. other records from data of the Wilson Copepod Library,
Washington, D.C.; 10. record in this paper (arrowed).

694

the diaptomid calanoid copepods which
tend to prefer open waters.

Acknowledgments

We thank Manuel Elias-Gutiérrez, I. Cas-
tellanos, and C. Curiel ECOSUR-Chetumal,
for collecting these samples at Tabasco.
This work is part of the postgraduate pro-
ject supporteed by CONACYT by the first
author to survey the freshwater copepod di-
versity of the Mexican state of Tabasco.
Sampling was supported by ECOSUR. Ja-
net Reid kindly provided relevant literature
from the Wilson Copepod Library. Frank
Ferrari made useful comments on the mor-
phology of the species.

Literature Cited

Bayly, I. A. E. 1992a. The non-marine Centropagidae.
Guides to the Identification of the Microinver-
tebrates of the Continental Waters of the World.
SPB Academic Publishers, The Hague, The
Netherlands, 30 pp.

. 1992b. Fusion of the genera Boeckella and
Pseudoboeckella (Copepoda) and revision of
their species from South America and sub Ant-
arctic islands.—Revista Chilena de Historia
Natural 65:17—63.

De Guerne, J., & J. Richard. 1889. Revision des cal-
anides d’eau douce.—Mémoires de la Société
Zoologique de France 2:53—231.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Mrazek, A. 1901. Siisswassercopepoden.—Ergebnisse
der Hamburger Magelhaensischen Sammelreise
1892-93, 2:1-29.

Reid, J. W. 1990. Continental and coastal free-living
Copepoda (Crustacea) of Mexico, Central
America and the Caribbean region. Pp. 175-213
in D. Navarro & J. G. Robinson, eds., Diversi-
dad Bioldgica en la Reserva de la Biosfera de
Sian Ka’an, Quintana Roo, Mexico. CIQRO/
University of Florida, Mexico, 471 pp.

. 1992. Copepoda (Crustacea) from fresh wa-
ters of the Florida Everglades, U.S.A., with a
description of Eucyclops conrowae n. sp.—
Transactions of the American Microscopical So-
ciety 111:229—254.

Reid, J. W., & R. M. Pinto-Coelho. 1994. An Afro-
Asian continental copepod, Mesocyclops ogun-
nus, found in Brazil; with a new key to the spe-
cies of Mesocyclops in South America and a
review of intercontinental introductions of co-
pepods.—Limnologica 24:359—-368.

Sars, G. O. 1863. Oversigt af de indenlandske fer-
skvandscopepoder.—Forhandlinger i Viden-
skabsselskabet i Kristiania. 1863:212—262.

Suarez-Morales, E., & J. W. Reid. 1998. An updated
list of the freshwater copepods of Mexico.—
Southwestern Naturalist 43:256—265.

Williamson, C. E. 1991. Copepoda. Pp. 787-822 in J.
H. Thorp & A. P. Covich, eds., Ecology and
classification of North American freshwater in-
vertebrates, Academic Press Inc., San Diego,
911 pp.

Wilson, M. S., & H. C. Yeatman. 1959. Free-living
Copepoda. Pp. 735-861 in W. T. Edmondson,
ed., Ward’s & Whipple’s Freshwater biology,
John Wiley & Sons, Inc., New York, 1248 pp.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(4):695-713. 1999.

New records of isopods from the Indian River Lagoon, Florida
(Crustacea: Peracarida)

Brian Kensley and Marilyn Schotte

Department of Invertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560, U.S.A.

Abstract.—Fifteen species of isopod are recorded for the first time as oc-
curring in the Indian River Lagoon. Two species are described as new: the
janirid asellote Jais floridana, n. sp., which occurs commensally with Sphae-
roma terebrans in low salinity water, and the sphaeromatid flabelliferan Sphae-
romopsis sanctaluciae, n. sp., which is also recorded from the Orange River,
Lee County, Florida, and from Islas de Juventud, Cuba. A brief discussion of
protogyny in the sphaeromatid Paradella dianae is included.

The Indian River Lagoon, Florida, is the
most biologically diverse estuarine system
on the east coast of North America. As part
of the Intra-coastal Waterway, it is subject
to heavy usage by commercial and sport/
recreational water traffic, and has seen
heavy residential development along its
shores. Given its important mixed-use re-
sources, intensive study of the lagoon has
been carried out for some time (see Rich-
ards 1995). Ongoing investigations by the
authors of the crustacean fauna of the la-
goon have revealed a number of isopod
species not recorded in earlier studies (e.g.,
Kensley, Nelson, & Schotte 1995), al-
though some of these may be known from
the wider Florida region (see Camp, Lyons,
& Perkins 1998). In part, these new records
are the result of sampling in a wide variety
of habitats, both in the main lagoon as well
as in its tributary rivers and in the inlets that
open to the sea. Twenty-five marine isopod
species had previously been recorded from
the IRL. The present paper documents 15
additional species and adds to the knowl-
edge of the biodiversity of the Indian River
Lagoon. Restricted synonymies, which in-
clude the original description plus any Flor-
ida records, and references that contain ful-
ler synonymies are provided for most spe-
cies. Collecting stations designated ‘FTP’

are those of the authors’. Unless otherwise
stated, all material is deposited in the col-
lections of the National Museum of Natural
History, Smithsonian Institution.

Suborder Anthuridea
Family Anthuridae
Cyathura polita (Stimpson, 1855)

Anthura polita Stimpson, 1855:393.—Har-
ger, 1880:398—402, pl. XI, figs. 68-69.
Cyathura polita: Burbanck, 1959:507.—
Kruczynski & Subrahmanyam, 1978:

93.—Camp et al., 1998:132.

Material examined.—1 2°, FTP-1, St. Lu-
cie River, rotten wood in mangroves, 0.5 m,
salinity 15-20 ppt., 29 May 1995.—1 2°,
FEP-22, Fort Pierce,, Taylor's Creek near
Rt. 1, rotten wood on muddy bank with cat-
tails and Spartina, intertidal, 10 ppt., 25
Apr 1996.

Previous records.—East coast of Amer-
ica from the Gulf of Mexico to Canada.

Mesanthura pulchra Barnard, 1925

Mesanthura pulchra Barnard, 1925:145,
fig. 9e—Kensley & Schotte, 1989:49,
fig. 19b; 52—53.—Camp et al., 1998:132.

Mesanthura decorata Menzies & Glynn,
1968:26, fig. 8a—-i.

696

Mesanthura floridensis Menzies & Kruc-
zynski, 1983:28—30, fig. 9a-.

Material examined.—1 °, 1 3, FTP-4,
Fort Pierce Inlet, south of South Jetty,
mixed algal turf on sand bag barrier on
beach, intertidal, 30 May 1995.—1 oviger-
ous 2, FTP-89, Sebastian Inlet State Park,
100 ft inside south jetty, encrusting orange
sponge with red branching alga on jetty
rocks, 0.5 m, 19 Sep 1996.—1 2, 1 juv.,
FTP-51, Sebastian Inlet State Park, south
side, algal clumps on granite boulders on
shore inside bridge, 0.5 m, 25 Jun 1997.

Previous records.—Belize; Puerto Rico;
Florida; intertidal to 36 m.

Ptilanthura tenuis Harger, 1878
Fig. 1

Ptilanthura tenuis Harger, 1878:377.—
Kensley, 1996a:763, figs. 1, 2.—Kensley,
1996b:278, fig. SB—D.—Camp et al.,
1998:132.

Material examined.—5 &°, Indian River
Lagoon, south of Sebastian Inlet,
27°49.64'N, 80°27.04'W, 1.6 m, salinity
30.15 ppt., 1995, coll. R. Heard.

Previous records.—Florida, Alabama, to
Maine; intertidal to 253 m.

Remarks.—The material from the region
of Sebastian Inlet had a distinctive and per-
sistant color pattern, which is recorded
here.

Suborder Asellota
Family Janiridae
Carpias algicola (Miller, 1941)

Janira algicola Miller, 1941:318—320, fig.
4a-n.

Bagatus algicola.—Pires, 1982:247—250,
fig. 47-55.

Carpias algicola.—Kensley & Schotte,
1989:82-83, fig. 38a—g.—Camp et al.,
19 9G1 55:

Material examined.—1 6,4 2, FTP-38,
Sebastian Inlet State Park, gravel and peb-
bles in pockets around boulders, infratidal,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Ptilanthura tenuis: Female in dorsal view,
showing persistent color pattern. Scale = 1 mm.

O-—0.8 m, 19 Sep 1996.—2 6, 1 ovigerous
?, 11 immature, FTP-40, Sebastian Inlet
State Park, clumps of Caulerpa racemosa
and branching red alga on granite boulders
inside inlet, 0.5—1.0 m, 19 Sep 1996.—2 ¢,

VOLUME 112, NUMBER 4

4 2, 2 immature, FTP-42, Sebastian Inlet
State Park, off south jetty about 50 yds.
west of bridge, on Caulerpa 0.25 m, 19 Sep
1996.—18 specimens, FTP-51, Sebastian
Inlet State Park, south side, granite boulder
shore inside bridge, algal clumps on boul-
ders, 0.5 m, 25 Jun 1997.—3 6, 7 oviger-
ous 2, 60+ immature, FTP-53, same lo-
cality, algal clumps and sponge on boulders
outside of bridge, 0.5-1.0 m, 26 Jun
1997.—3 specimens, FTP-65, Sebastian In-
let State Park, orange sponge on rocks in-
side of inlet, low tide level, O—50O cm, 18
Aug. 1998.—1 ovigerous 2, 5 immature,
FTP-68, Sebastian Inlet State Park, gravel
rubble, empty shells in pockets between
rocks inside inlet, 0-20 cm, 18 Aug
1998.—12 ovigerous 2, 8+ immature,
FTP-69, Sebastian Inlet State Park, algal
turf and hydroids on granite boulders inside
inlet, O-50 cm, 18 Aug. 98.

Previous records.—Venezuela; Jamaica;
Yucatan, Mexico; Belize; Looe Key, Flori-
da; intertidal to 2 m.

Carpias triton (Pires, 1982)

Bagatus triton Pires, 1982:251—254, figs.
72-84. .

Carpias triton.—Kensley & Schotte, 1989:
83, 87, fig. 39e-f.

Material examined.—20+ specimens,
FTP-5, Ft. Pierce Inlet, large barnacle
clumps with orange sponge and algal turf
on boulders inside inlet, shallow infratidal,
30 May 1995.

Previous records.—Belize, intertidal.

Tais floridana, new species
Figs. 2, 3

Material examined.—Holotype, USNM
243843, 1 ¢ tl 1.35 mm, Paratypes, USNM
243844, 10 ¢ tl 1.26-1.40 mm, 12 ovig-
erous @ tl 1.71—1.82 mm, 10 non-ovigerous
2, 50+ juveniles, sta FTP-22, on Sphae-
roma terebrans in rotten wood, Taylor
Creek near Rt. 1, intertidal, 10 ppt., 25 Apr
1996.

697

Additional material.—10 specimens, sta
FTP-2, on Sphaeroma terebrans in rotten
wood, Port St. Lucie, Florida, 29 May
1995.—100+ specimens, sta FTP-12, on
Sphaeroma terebrans in rotten wood, Riv-
erside Park at Port St. Lucie Boulevard, in-
tertidal—1 m, 10 ppt., 1 Jun 1995.—1 spec-
imen, sta FTP-13, on Sphaeroma terebrans
in dead wood, north fork of St. Lucie River,
at Prima Vera Boulevard, 2.5 ppt, 0.1 m, 1
Jun 1995.—2 specimens, sta FTP-17, on
Sphaeroma terebrans in rotten wood, Riv-
erside Park on St. Lucie River, O ppt, 0.1—
0.5 m, 23 Apr 1996.—3 specimens, sta
FTP-27, on Sphaeroma terebrans in rotten
wood, near mouth of St. Sebastian River,
15 ppt, 0.5 m, 17 Sep 1996.—2 specimens,
sta FTP-29, on Sphaeroma terebrans in rot-
ten wood, island in mouth of St. Sebastian
River, 15 ppt., 0.5 m, 17 Sep 1996.—14
specimens, Estero River mouth, Lee Coun-
ty, Florida, 26°26'05’N, 81°50’52’W, coll.
A. S. Walton.—1 specimen, Hendry Creek,
off Estero Bay, Lee County, Florida, coll.
A. S. Walton, 14 Apr 1993.

Description.—Male: Body about 2.5
times longer than greatest width at pereo-
nite 4. Cephalon about twice wider than
long, anterolateral corners rounded. Antero-
lateral corners of pereonites, especially of
3—5, rounded, setose, coxae visible on all
pereonites in dorsal view. Pleon consisting
of short anterior pleonite lacking free lateral
margins, plus subcircular pleotelson. Eye
consisting of 2 ommatidia. Antennular fla-
gellum of 3 articles, terminal and subter-
minal articles each bearing single aesthe-
tasc. Antennal flagellum of 13 articles.
Mandibular palp of 3 articles, article 2 with
2 stout setae, article 3 with distal row of 7
setae; incisor of about 6 sclerotized cusps;
spine row of 4 or 5 setae; lacinia of 2 cusps;
molar distally truncate with 2 distomesial
setae. Maxilla 1, inner ramus with 4 distal
setae, outer ramus with about 10 pectinate
distal setae. Maxilla 2, 2 outer lobes each
with 4 distal fringed setae. Maxilliped; en-
dite broad, distolaterally convex, with about

698

7 fringed setae and 8 simple setae; palp ar-
ticles setose on mesial margins.

Pereopod 1, dactylus bearing 2 claws; pe-
reopods 2-3, 5-7, dactyli each with 3
claws. Pereopod 4 considerably shorter than
3 or 5, dactylus with 2 claws, propodus
with single stout distal claw. Pleopod 1,
rami fused for about 4/5 of total length, dis-
tal lobes rounded, bearing 9 setae distally
per side. Pleopod 2, protopod semicircular,
canula not reaching beyond distal angle of
protopod. Uropodal rami both longer than
protopod, exopod about 1/3 longer than en-
dopod, each with 4 elongate distal simple
setae.

Female: Brood pouch containing up to 8
eggs. Pleonal operculum ovate, midlength
about 2/3 greatest width, with 4 or 5 fine
marginal setae.

Remarks.—Of the eight described spe-
cies of Jais (see Wilson & Wagele 1994),
at least three occur commensally with
sphaeromatid isopods, as does the present
species, which is found in association with
Sphaeroma terebrans. Several species (e.g.,
I. aquilei Coineau, 1977; I. elongata Si-
vertsen & MHolthuis, 1980; see Kensley
1994) also perform mate-guarding as is
seen in the present material, with the male
clasping a manca female with the shortened
specialized pereopod 4.

Given that some species of Sphaeroma,
especially those that bore into mangroves,
have wide distributions, and have been im-
plicated in introductions along with their
commensals (Rotramel 1972, 1975), it is
necessary to compare the present material
closely with I. californica (found on Sphae-
roma quoyanum), in case the present spe-
cies was somehow introduced to the east
coast of the United States, where Sphaero-
ma terebrans is the available host. How-
ever, Jais floridana more closely resembles
I. singaporensis Menzies & Barnard, 1951
(see Miiller & Brusca 1992) especially in
the general habitus and in possessing
rounded anterolateral lobes on the pereoni-
tes, than J. californica (Richardson, 1904).
Comparison with recently collected mate-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

rial of both L. californica and I. singapo-
rensis reveals several differences that rein-
force the view that the Florida material rep-
resents an undescribed species. The two
distal articles of the antennule differ in pro-
portions, the penultimate articles especially
being more slender and elongate on the two
previously described species. The antennal
flagellum has fewer articles in the Florida
material (13) than in I. californica (20) and
I. singaporensis (24). The distal propodal
spine of pereopod 4 in the male of I. cali-
fornica is noticeably more elongate than in
the Florida and Asian material. Jais califor-
nica is a larger species (¢ 2.49 mm mean
length, n = 15; ovigerous 2 2.55 mm mean
length, m = 11) than either the Florida spe-
cies (6 1.34 mm mean length, n = 10;
ovigerous 2 1.76 mm mean length, n = 10)
or I. singaporensis (36 1.3-1.7 mm, ovig-
erous 2 1.4—1.7 mm). The stylet of pleopod
2 of the male is more slender and elongate
in the Florida species than in J. singaporen-
SiS:

Etymology.—The specific name derives
from Florida, from whence the species is
recorded.

Family Joeropsidae
Joeropsis coralicola Schultz &
McCloskey, 1967

Joeropsis coralicola Schultz & McCloskey,
1967:103-—107, figs. 1-39.—Kensley &
Schotte, 1989:88, fig. 40g Camp et al.,
1998:133.

Material examined.—11 specimens, sta
FTP-5, Fort Pierce Inlet, on large barnacle
clumps with orange sponge and algal turf
on boulders inside inlet, shallow infratidal,
30 May 1995.

Previous records.—North Carolina to
Florida Middle Grounds, Gulf of Mexico,
25-33 m.

Joeropsis tobagoensis Kensley & Schotte,
1994

Joeropsis tobagoensis Kensley & Schotte,
1994:482, 486, fig. la—o.

VOLUME 112, NUMBER 4 699

Fig. 2. Jais floridana, new species: A, habitus, dorsal view, scale = 0.5 mm; B, antenna; C, antennule; D,
maxilliped; E, maxilla 1; FE left mandible; G, right mandible (palp omitted); H, maxilla 2.

700 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Jais floridana, new species: A, pereopod 1 propodus and dactylus; B, pereopod 2, propodus and
dactylus; C, pereopod 3; D, male pereopod 4; E, uropod; EK male pleopod 1; G, male pleopod 2; H, female
operculum.

VOLUME 112, NUMBER 4

Material examined.—18 specimens, sta
FTP-5, Fort Pierce Inlet, on large barnacle
clumps with orange sponge and algal turf
on boulders inside inlet, shallow infratidal,
30 May 1995.

Previous records.—Tobago, intertidal to
Sm:

Suborder Flabellifera
Family Cirolanidae
Anopsilana jonesi Kensley, 1987

Anopsilana jonesi Kensley, 1987:565—568,
fig. 5a—j, 6a—h.—Camp et al., 1998:135.

Material examined.—1 °, sta FTP-12,
North Fork St. Lucie River at Riverside
Park on Port St. Lucie Blvd., rotten wood
around dock, intertidal to 1 m, salinity 10
ppt., 1 Jun 1995.—1 4, 1 juv., sta FTP-27,
Indian River Lagoon near mouth of Sebas-
tian River, rotten submerged wood on small
island, in low turf of Enteromorpha and
Ceramium, 0.5 m, salinity 15 ppt., 17 Sep
1996.—1 6, 1 &, sta FTP-29, Sebastian
River, first island inside mouth, on rotten
wood at shore, salinity 15 ppt., 17 Sep
1996.

Previous records.—Belize; Florida; in
estuarine mangroves.

Cirolana parva Hansen, 1890

Cirolana parva Hansen, 1890:340—341, pl.
II, fig. 6—6b, pl. III, fig. 1-1d.—Bruce &
Bowman, 1982:325-333, figs. 1, 2.—
Kensley & Schotte, 1989:135, fig. 59d—
e, 60.—Camp et al., 1998:135.

Material examined.—2 2°, sta FTP-38,
Sebastian Inlet State Park, gravel and peb-
bles in pockets around granite boulders, in-
fratidal, 19 Sep 1996.—1 @, 1 juv., sta
FTP-51, Sebastian Inlet State Park, south
side of inlet, algal clumps on granite boul-
ders, 0.5 m, 25 Jun 1997.—1 juv., sta FTP-
52, Sebastian Inlet State Park, south side,
shallow embayment at campsite in State
Park, 1/2 mile from mouth in lagoon, 0.5
m, 25 Jun 1997.—1 juv., sta FTP-57, Se-
bastian Inlet State Park, lagoon near Co-

701

conut Point, sweep through Syringodium on
Inlet side, 0.5—1 m, 26 Jun 1997.—1 juv.,
sta FTP-60, Wabasso Causeway Park, sub-
merged rotten wood, 20-40 cm, 26 Jun
1997.

Previous records.—Panama; Belize;
Cozumel, Mexico; Antilles to Florida Keys;
Gulf of Mexico; N. & S. Carolina; intertidal
to 55 m.

Family Corallanidae
Excorallana sexticornis (Richardson,
1901)

Corallana_ sexticornis Richardson,
518, fig 9.

Excorallana sexticornis: Delaney, 1989:
38.—Kensley & Schotte, 1989:165, figs.
75e—-f, 76d—f.—Camp et al., 1998:135.

Material examined.—1 @, sta FTP-14,
Ft. Pierce Inlet, barnacles, sponges, algal
turf on blocks in inlet, intertidal, 23 Apr
1996.—1 ovigerous @2, sta FTP-15, Ft.
Pierce State Recreational Area, rotten wood
piles with encrusting algae, intertidal, 23
Apr 1996.—1 @, 3 juvs., sta FTP-17, North
Fork St. Lucie River, Riverside Park, rotten
submerged wood on mud with numerous
barnacles and shells, 10-50 cm, 23 Apr
1996.—2 °, sta FTP-33, Jim Island near Ft.
Pierce Inlet, dead submerged wood at edge
of mangrove island, with algal mat, 0.—0.5
m, 18 Sep 1996.—1 &, sta FTP-38, Sebas-
tian Inlet State Park, gravel inlet, south side,
east of bridge, algal clumps, sponge on
boulders, strong wave and wash action,
0.5—-1.0 m, 26 Jun 1997.—1 &, sta FTP-61,
North Hutchinson Island, near causeway,
rocks with algal turf, 0.1 m, 27 Jun 1997.
kes lo 9. 4.g0v.. Sta. JZRP-71. Ft Piesce
Inlet, north bank, algal turf on boulders,
low tide level, 19 Aug 1998.

Previous records.—Belize; Puerto Rico;
Cuba; Florida; shallow infratidal.

1901:

Family Sphaeromatidae
Cassidinidea ovalis (Say, 1818)

Naesa ovalis Say, 1818:484—485.—Rich-
ardson, 1900:224, 1901:537.

702

Cassidena lunifrons: Richardson, 1900:
222.

Cassidina lunifrons: Richardson, 1901:533,
fig. 14.

Cassidisca lunifrons: Richardson, 1905:
273, figs. 283—284.—Schultz, 1969:115,
fig. 158.

Cassidinidea lunifrons: Hansen, 1905:
130.—Menzies & Frankenberg, 1966:44,
fig. 20.—Kussakin, 1979:336, figs. 199—
200.—Bruce, 1994:1151.

Cassidinidea ovalis: Schultz, 1969:115, fig.
159.—Kensley & Schotte, 1989:208, fig.
92b—e.—Bruce, 1994:1151, fig. 45.—
Camp et al., 1998:136.

Dies arndti Ortiz & Lalana, 1980:161—164,
figs. 1-8.

Dies barnardi Carvacho, 1977:14—17, figs.
4a-f, 5a—i.

Material examined.—1 2°, FTP-1, North
Fork St. Lucie River, rotten wood in man-
groves, 0.5 m, salinity 1—20 ppt., 29 May
1995.—3 specimens, FTP-12, North Fork
St. Lucie River at Riverside Park, Port St.
Lucie Boulevard, rotten wood around dock,
intertidal, salinity 10 ppt., 1 Jun 1995.
specimen, FTP-13, North Fork St. Lucie
River at marina on Prima Vera Boulevard,
dead submerged wood in shore grass at riv-
er’s edge, 0.1 m, salinity 2.5 ppt., 1 Jun
1995.—4 specimens, FTP-17, Riverside
Park on North Fork St. Lucie River, rotten
submerged wood on mud with numerous
barnacles and shells, 10—50 cm, salinity 0
ppt., 23 Apr 1996.—30+ specimens, FTP-
23, mouth of North Fork St. Lucie River at
U.S. Rt. 1 and Fern Rd., oysters shells and
rocks on muddy bank, intertidal, salinity 0
ppt., 25 Apr 1996.—1 ovigerous 2, FTP-
29, Sebastian River, first island west of
mouth, rotten submerged wood at shore, sa-
linity 15 ppt., 17 Sep 1996.—1 specimen,
FTP-30, same locality as above, in organic
detritus, intertidal, 17 Sep 1996.—1 speci-
men, FTP-31, Sebastian River, island op-
posite MacDonald State Campground, sub-
merged leaf litter, Typha and Crinum in
shallow water, salinity 0 ppt., 17 Sep 1996.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Previous records.—Panama; Belize;
Trinidad; Dominica; Cuba; Gulf of Mexico;
Florida to New Jersey; intertidal—1 m.

Paradella dianae (Menzies, 1962)
Figs. 4, 5

Dynamenopsis dianae Menzies, 1962:341,
fe. 3:

Paradella dianae: Harrison & Holdich,
1982:103, fig. 6—Kensley & Schotte,
1989:224, fig. 98a—c.

Material examined.—3 ¢, FTP-14, Ft.
Pierce Inlet, barnacles, sponges, algal turf
on blocks in inlet, intertidal, 23 Apr
1996.—6 6, 14 ovigerous ¢, 30+ imma-
ture, FTP-15, Ft. Pierce Recreational Area,
rotten wood piles with encrusting algae, in-
tertidal, 23 Apr 1996.—6 6, 4 ovigerous
2, 25+ immature, FTP-17, Riverside Park
on North Fork St. Lucie River, rotten sub-
merged wood on mud with barnacle shells,
salinity 0 ppt., 10-50 cm, 23 Apr 1996.
1 6, 2 ovigerous 2, 15+ immature, FTP-
19, Jack Island near Ft. Pierce Inlet, Cau-
lerpa and empty shells near oyster bank, 0.5
m, 24 Apr 1996.—100+ specimens, FTP-
38, Sebastian Inlet State Park, gravel and
pebbles in pockets around granite boulders
lining inlet, infratidal, 0-32", 19 Sep
1996.—92 specimens, FTP-39, same local-
ity, on encrusting orange sponge with red
branching algae on jetty rocks ca. 100 ft
from end of south jetty, 0.5 m, 19 Sep
1996.—2 <6, 1 immature, FTP-42, same lo-
cality, 50 m west of bridge off south jetty,
in Caulerpa, depth 6”, 19 Sep 1996.—38
specimens, FTP-44, same locality, 50 ft in-
land from bridge, mixed algae on sandy/
Shelly bottom with rocks and boulders, 0.5
m, 19 Sep 1996.—1 6, 7 2,2 juvs EEE
45, same locality, red filamentous alga on
rocks and south jetty wall, 30 cm, 19 Sep
1996.—1 6, 4 ovigerous 2, 3 immature,
FTP-46, Sebastian Inlet State Park, north
side, gravel and pebbles among granite
boulders ca. 100 m inside inlet, 10—50 cm,
20 Sep 1996.—12 6, 11 ovigerous 2, 73
2, 40 juvs., FTP-48, Sebastian Inlet State

VOLUME 112, NUMBER 4

Park, south side, rubble and stones in 3”
pools at top of shore, with blue-green alga,
25 Jun 1997.—1 6,5 &, 1 juv., FTP-50,
same locality, granite boulder shore inside
of bridge, stones and rubble with low algal
turf, at bottom of shore with strong wave
and wash action, 25 Jun 1997.—6 subadult
6, 3 ovigerous 2, 4 2, 6 juvs., FTP-51,
same locality, algal clumps on boulders in-
side of bridge, 0.5 m, 25 Jun 1997.—2 9°,
FTP-53, same locality, outside of bridge, al-
gal clumps and sponge on boulders in
strong wave and wash action, 26 Jun
1997.—1 6, 1 subadult ¢, FTP-54, same
locality, boulders outside bridge, chunks of
reef worm rock, 26 Jun 1997.—7 ovigerous
2, 14 juvs., FTP-56, Sebastian Inlet State
Park, lagoon near Coconut Point, Entero-
morpha/Ulva mats exposed at low tide on
boulders at top of shore, surface, 26 Jun
1997.—1 6, 4 ovigerous 2, FTP-63, large
boat canal at Smithsonian Marine Station,
in floating Sargassum, at surface, 25 Jun
1997.1 6, 4 ovigerous ?, FTP-66, Se-
bastian Inlet State Park, chunks of reef
worm tubes on rocks at low tide level, in-
side inlet, 0-50 cm, 18 Aug 1998.—4 sub-
adults, 1 ovigerous °, FTP-68, same local-
ity, gravel rubble and empty shells in pock-
ets between rocks, inside inlet, 0-20 cm, 18
Aug 1998.—3 ovig 2, FTP-69, same lo-
cality, algal turf with hydroids on granite
boulders inside inlet, 0-50 cm, 18 Aug
1998.—3 ovigerous 2, FTP-70, Ft. Pierce
Inlet, north bank, reef worm tubes on boul-
ders in inlet, low tide, surface, 19 Aug
1998.—2 ovigerous ¢, FTP-71, same lo-
cality, algal turf at low tide level on boul-
ders, surface, 19 Aug 1998.

Previous records.—Baja California,
Mexico; Queensland, Australia; Western
Australia; Marshall Islands; Hong Kong;
Puerto Rico; Florida; intertidal.

Remarks.—While Paradella dianae has
previously been recorded from the IRL, an
aspect of its biology has come to light that
demands mention.

Fifty-one ovigerous females out of 182
examined (about 28%) were observed to

703

possess penes, suggesting that a protogyn-
ous sex change occurs in P. dianae. In Fig.
4C, a scanning electron micrograph, the
ovigerous female shows both the opening
of the marsupium between the fourth pe-
reopod bases, and penes that are character-
istic of a subadult male. The penes of the
adult male are long, very slender in the dis-
tal half, tapering to acute apices and ex-
tending beyond the endopod of pleopod 1
by nearly 50%. The ovigerous hermaphro-
dites show no retention of either appendix
masculina or adult penes, which suggests
that protandry is not the condition here.
This would seem to be the first record of
protogyny in the sphaeromatid subfamily
Dynameninae. Among the Isopoda, protan-
drous sex change is well known in the fam-
ilies Anthuridae (Wagele 1979), Cymothoi-
dae (Brusca 1981),.several families of the
suborder Epicaridea (Kozloff 1987), and in
at least one oniscidean (Brook et al. 1994).
Members of the Sphaeromatidae known to
exhibit protogyny are members of other
subfamilies: Gnorimosphaeroma oregon-
ense (Dana, 1853), G. luteum Menzies,
1954 (both Sphaeromatinae), and Paralep-
tosphaeroma glynni Buss & Iverson, 1981
(Cassidininae). Bruce (1994:1132) further
mentions observing hermaphroditism in
Paracassidina munna, having ‘developed
male characters in ... pleopod 2” as well
as oostegites in the same specimen. Pleopod
2 in the ovigerous females of P. dianae did
not display any male characters. The pro-
portion of ovigerous females with penes in
G. oregonense (31% of females collected in
the field) is comparable to that of P. dianae
recorded here. Brook et al. (1994) provide
a discussion of the adaptive value of pro-
togyny as compared to protandry, the com-
moner reproductive strategy in Crustacea.

Paradella quadripunctata (Menzies &
Glynn, 1968)
Fig. 6

Dynamenella quadripunctata Menzies &
Glynn, 1968:60—61, fig. 28a—n.

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Paradella dianae: A, ventral cephalon; B, ovigerous female, ventral view, arrow indicating opening
of brood pouch between fourth pereopods; C, ovigerous female with brood pouch opening and penes; D, ovig-
erous female, close-up of ventral pleotelson and penes.

Paradella quadripunctata: Harrison &
Holdich, 1982:101.—Kensley & Schotte,
1989:224—225, fig. 98f—-g.—Camp et al.,
1993: 136:

Material examined.—1 ovigerous 9°,
FTP-38, Sebastian Inlet State Park, gravel
and pebbles in pockets around granite boul-
ders lining inlet, infratidal, 0-32”, 19 Sep
1996.—1 immature, FTP-50, Sebastian In-
let State Park, south side, granite boulder
shore inside of bridge, in stones and rubble
with algal turf at bottom of shore with
strong wave action, 25 Jun 1997.—24 im-

mature, FTP-51, same locality, algal clumps
on boulders, 0.5 m, 25 Jun 1997.—25+ im-
mature, same locality, FTP-53, south side,
outside bridge, algal clumps and sponge on
boulders in strong wave and wash action,
0.5-1.0 m, 26 Jun 1997.—1 subadult <6,
80+ immature, FTP-54, same locality,
south side, boulders in inlet, outside of
bridge, in chunks of reef worm rock, 26 Jun
1997.—5 immature, FTP-59, Sebastian In-
let State Park, lagoon near Coconut Point,
26 Jun 1997.—25+ immature, FTP-62,
North Hutchinson Island, Recreation Park,

VOLUME 112, NUMBER 4

Figs5:
indicating immature penes and appendix masculina; C, ovigerous female, dorsal pleotelson; D, mature male,
dorsal pleotelson.

rotten wood in shallow water, < 1 m, 27
Jun 1997.—8 subadult 6, FTP-66, Sebas-
tian Inlet State Park, chunks of reef worm
tubes on rocks at low tide, inside inlet, O—
50 cm, 18 Aug 1998.—1 ovigerous 9°,
FTP-67, same locality, algal turf at low tide
inside inlet, 0.5-1 m, 18 Aug 1998.—1
ovigerous 2, FTP-68, same locality, gravel
rubble, empty shells between rocks inside
inlet, O—20 cm, 18 Aug 1998.—1 ovigerous
?, 3 immature, FTP-69, same locality, algal
turf mixed with hydroids on granite boul-
ders inside inlet, 0-50 cm, 18 Aug 1998.—
2 subadult ¢, 1 ovigerous 2, FTP-70, Ft.

Paradella dianae: A, sub-adult male, dorsal pleotelson; B, sub-adult male, ventral pleotelson, arrows

Pierce Inlet, north bank, reef worm tubes
on boulders in inlet, low tide, 19 Aug
1998.—2 subadult ¢, 2 immature, FTP-71,
same locality, algal turf on boulders at low
tide level, 19 Aug 1998.—2 subadult d, 3
ovigerous 2, FTP-72, Warton Beach rocks
off Rt. AlA, algal turf growing on beach
rock at bottom of shore, 0-50 cm, 20 Aug
1998.

Previous records.—Dominican Republic;
Puerto Rico; U.S. Virgin Is.; Florida; Ber-
muda; intertidal—1 m.

Remarks.—Although no adult males
were collected, identification was based on

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Paradella quadripunctata: A, allotype, ovigerous female, ex USNM 119307, dorsal pleotelson; B,
ventral pleotelson; C, ovigerous female, Indian River specimen; D, ovigerous female, ventral cephalon lamina;
E, sub-adult male, dorsal pleotelson; K sub-adult male, ventral pleotelson.

VOLUME 112, NUMBER 4

comparison of ovigerous females and sub-
adult males to the ovigerous allotype (Fig.
6A-F).

Sphaeromopsis sanctaluciae, new species
Figs. 7-9

Material examined.—Holotype, USNM
285356, 1 6 tl 3.1 mm, Allotype USNM
285357, 1 2 tl 2.0 mm, Paratypes, USNM
285358, 45 3d, 29 2, 39 juvs., sta FTP-13,
North Fork of St. Lucie River at Prima Vera
Boulevard, Port St. Lucie, Florida, in dead,
submerged wood, 0.1 m, 1 Jun 1995.

Additional material.—USNM 285359, 2
ovigerous 2, 2 @, 1 juv., sta K-CUBA-64,
Islas de Juventud, Ensenada de la Siguanea,
Cuba, in algal carpet on Rhizophora roots,
0.5 m, 9 Jun 1995.—USNM 285360, 5 2°,
1 juv., sta: FTP-8, Merritt Island at boat
ramp, Indian River Lagoon, on dead wood,
0.5 m, 31 May 1995.—USNM 285361, 1
2, 3 juvs., sta FTP-11, on ring of metal
plates in Banana River off Merritt Island,
Indian River Lagoon, amongst encrusting
oysters, barnacles and algal turf, 0.5 m, 31
May 1995.—USNM 285362, 1 2, 2 juvs.,
sta FTP-24, mouth of North Fork of St. Lu-
cie River at US 1 and Fern Rd., in algal
turf with Enteromorpha on boulders, inter-
tidal, 25 Apr 1996.—USNM 285363, 1 °,
Indian River Lagoon at Jensen Beach, in
algae in mangrove roots, 29 May 1995.—
USMN 285364, 3 6, 1 ovigerous 2, 1 &,
Orange River, Lee County, Florida, coll. A.
Walton, 15 Aug 1994.—USNM 285365, 1
6, Orange River, Lee County, Florida, coll.
A. Walton, 18 Jan 1995.

Diagnosis.—Sexes similar, cephalon and
pereon smooth, pigmented; pleotelson
domed, smooth, apex broadly truncate.
Margins of uropodal rami entire. Appendix
masculina of male broad proximally, taper-
ing to narrowly rounded apex. Rami of pe-
nes elongate, widening in proximal half be-
fore tapering to narrowly rounded apices.

Description.—Adult male: Body length
about 1.9 times greatest width. Cephalon
broader than long, frontal margin undulat-

707

ing, rostral point small and acute. Frontal
lamina narrowly truncate distally. Brown
pigment pattern somewhat variable, strong
in fresh specimens, densest on pleotelson.
Pleon with two short suture lines reaching
posterior margin. Pleotelson broadly trian-
gular, domed, with apex broadly truncate in
posterior view.

Antennule with basal article equal in
length to articles 2 and 3 combined; flagel-
lum of 9 articles; articles 5—8 each bearing
single aesthetasc. Antenna with articles 1
and 2 subequal in length; article 3 shorter
than 2; article 4 somewhat shorter than 2
and 3 together; article 5 longest; flagellum
of 13 articles. Mandible having incisor of 3
cusps, spine row of 5 spines, 3 of which
fringed, molar process with numerous small
teeth; palp, article 2 having 4 fringed setae,
terminal article with 7 fringed setae. Max-
illa 1, inner ramus with 4 fringed setae, out-
er ramus with 4 stout spines and 4 slender,
fringed spines. Maxilla 2 bearing 3 un-
armed and 4 fringed spines on inner ramus;
outer ramus having 4 fringed spines on each
lobe. Maxillipedal endite with 1 coupling
hook on mesial margin; dense, fine setae
distally; distal margin with 5 blunt spines
and several fringed setae; palp of 5 articles,
articles 2 and 3 with distomesial lobe bear-
ing several setae, article 4 longer and more
slender than 3, article 5 short with terminal
setae.

Pereopods with fringe of short setae on
posterior margins of propodus, carpus and
merus, very sparse on merus of pereopod
5. Pereopod 1, propodus and carpus each
with single plumose seta at anterodistal
margins; merus with anterodistal lobe bear-
ing 3 long setae. Pereopod 2 slender, longer
than 1; propodus with single plumose seta
anterodistally; carpus with single plumose
setae on posterodistal margin; merus having
anterodistal lobe bearing 4 long setae; is-
chium with several long setae on anterior
margin. Pereopods 3—7 equally robust, in-
creasing in length posteriorly. Pereopod 3
shorter than 2; propodus with 2 fringed pos-
terodistal spines; carpus triangular with

708

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Pisege
C, antennule; D, antenna; E, mandible; EK maxilliped; G, uropodal rami; H, maxilla 2; I, maxilla 1; J, ventral
cephalon; K, penes of sub-adult male; L, penes of adult male; M, pleopod 1; N, pleopod 2; O, pleopod 3; P,
pleopod 4; Q, pleopod 5.

fringed spine at posterodistal corner; merus
with small anterodistal lobe and single
fringed, posterodistal spine. Pereopod 4,
propodus having single plumose seta at an-
terodistal angle; carpus with single plumose
seta at each distal margin; merus with an-

Sphaeromopsis sanctaluciae, new species: A, adult male, habitus. Scale = 1 mm; B, lateral view;

terodistal lobe bearing several long setae.
Pereopod 5, carpus with single stout fringed
Spine at posterodistal margin; anterodistal
lobe of merus having several long setae. Pe-
reopod 6, carpus with plumose seta and
stout fringed spine at antero- and poster-

VOLUME 112, NUMBER 4

709

Fig. 8.
4; E, pereopod 5; FE pereopod 6; G, pereopod 7.

odistal margins respectively; anterodistal
lobe of merus with several long setae; is-
chium bearing 3 long setae on anterodistal
margin. Pereopod 7, propodus with single
plumose seta at anterodistal corner; distal
margin of carpus with 3 fringed spines, 3

Sphaeromopsis sanctaluciae, new species: A, pereopod 1; B, pereopod 2; C, pereopod 3; D, pereopod

fringed, 2 unarmed, and 1 plumose setae;
anterodistal margin of merus with 2 long
and 1 short setae; ischium and basis with
several setae on anterior margins.

Penial rami basally fused, elongate, wid-
ening at 1/3 length and tapering to narrowly

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 9. Sphaeromopsis sanctaluciae, new species: A, adult male, dorsal pleotelson; B, ventral cephalon; C,
adult female, dorsal pleotelson; D, adult female, ventral pleotelson.

rounded apices. In subadult male, penial
rami fused basally with subparallel margins
and rounded apices. Pleopod 1, basis with
3 coupling hooks; exopod subrectangular in
shape, slightly broader than endopod. Ple-
opod 2, basis and exopod as in pleopod 1;
endopod broad, subrectangular; appendix
masculina articulating basally, broad prox-
imally, tapering to narrow apex extending
somewhat beyond apex of endopod. Pleo-
pod 3, basis broadly rectangular with 3 cou-
pling hooks; endopod triangular without ar-
ticulation; exopod somewhat ovate. Pleo-
pod 4, both rami broad with transverse
pleats. Pleopod 5, both rami with transverse

pleats and setae on mesial margins; exopod
with transverse suture and 3 spinulose boss-
es. Rami of uropods subequal in length,
margins entire, bearing short setae.
Female: As in male except in secondary
sexual characters; generally smaller.
Remarks.—The new species is the sev-
enth member of the genus to be described
and the third from the western hemisphere,
following S. mourei Loyola e Silva, 1960
and S. heardi Kensley & Schotte, 1994.
Like S. minutus Javed & Yousef, 1995, it
lacks the “dense pads of setae’’ on posterior
margins of the pereopods, previously
thought to be a generic character (Holdich

VOLUME 112, NUMBER 4

& Harrison 1981). The comparatively
sparse nature of the setal fringe plus mor-
phological details of the appendix mascu-
lina and penes serve to separate S. sanctal-
uciae from S$. mourei and S. amathitis
Holdich & Jones, 1973, both of which it
superficially resembles. In contrast to the
other two species, the appendix masculina
in S. sanctaluciae is markedly inflated near
the base and extends beyond the apex of
the pleopod endopod.

Whereas almost all cogeners have been
collected from sandy beaches and intertidal
habitats (Holdich & Harrison 1981), S.
sanctaluciae, like S. serriguberna Holdich
& Harrison 1981, can also tolerate low sa-
linity, e.g., 2.5 ppt. in the St. Lucie River.

Etymology.—The species is named for its
type locality, St. Lucie River.

Suborder Valvifera
Family Idoteidae
Erichsonella filiformis (Say, 1818)

Stenosoma filiformis Say, 1818:424.

Erichsonella filiformis: Kensley & Schotte,
1989:258, fig. 108c.—Camp et al., 1998:
TSP.

Material examined.—7 specimens, 1
ovigerous ¢, FTP-40, Sebastian Inlet State
Park, clumps of Caulerpa racemosa and
branching red alga on granite boulders in-
side of inlet, 0.5—-1.0 m, 19 Sep 1996.—1
°, FTP-41, Sebastian Inlet State Park,
south side, sheltered cove ca. 1/2 mile from
mouth, small boulders with clumps of al-
gae, 0.5—1 m, 19 Sep 1996.—3 @&, 2 juvs.,
FTP-42, same locality, 50 yds. west of
bridge, in Caulerpa sp., depth 6”, 19 Sep
1996.—1 @, 1 juv., FTP-44, same locality,
50 ft west of bridge, mixed algae on sandy/
shelly/rocky bottom, 0.5 m, 19 Sep 1996.—
1 2, 4 juvs., FTP-51, Sebastian Inlet State
Park, south side, granite boulder shore in-
side of bridge, algal clumps on boulders,
sm, 25 June 199/—2 6, 4.9. 7. juvs.,
FTP-52, same locality, south side, shallow
embayment at camp site 1/2 mile from
mouth in lagoon, algal clumps on stones

eit

and rocks, 25 Jun 1997.—1 9°, 1 juv., FTP-
53, same locality, south side, boulders out-
side of bridge, algal clumps and sponge on
boulders in strong wave and wash action,
0.5—1.0 m, 26 Jun 1997.—3 ovigerous &,
1 2°, FTP-55, Sebastian Inlet State Park, la-
goon near Coconut Point, algal clumps in
shallow embayment, 1 m, 26 Jun 1997.—1
juv., FTP-65, Sebastian Inlet State Park, or-
ange sponge on rocks at low tide level, in-
side of inlet, 0-50 cm, 18 Aug 1998.—1 9°,
FTP-68, same locality, gravel rubble, empty
shells between rocks inside inlet, O—20 cm,
18 Aug 1998.—1 6, FTP-69, same locality,
algal turf mixed with hydroids on granite
boulders inside inlet, 0-50 cm, 18 Aug
1998.

Previous records.—Brazil; Yucatan,
Mexico; Puerto Rico; Turks & Caicos Is.;
Bahamas; Gulf of Mexico; shallow infrati-
dal—109 m.

Acknowledgments

We are grateful to the staff of the Smith-
sonian Marine Station, and especially
Woody Lee and Sherry Reed for assistance
with this project in the field, and for pro-
viding ideal working conditions at the lab.
We thank Dr. David Camp, late of the Flor-
ida Marine Research Institute, and Dr. A. S.
Walton of the Florida Department of Envi-
ronmental Regulation for making material
available for study. Susann Braden assisted
with the SEM’s, for which we are grateful.

This paper is Contribution Number 470
from the Smithsonian Marine Station at
Link Port, Florida.

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(4):714-721. 1999.

A new species of Pseudostegias Shiino, 1933
(Crustacea: Isopoda: Bopyridae: Athelginae)
parasitic on hermit crabs from Bali

Jason D. Williams and Christopher B. Boyko

(JDW, CBB) Department of Biological Sciences, University of Rhode Island,
Kingston, Rhode Island 02881-0816, U.S.A.; (CBB) Department of Invertebrates,
American Museum of Natural History, Central Park West @ 79th St.,

New York, New York 10024, U.S.A.

Abstract.—A single male and female bopyrid pair was collected in Sanur,
Bali, Indonesia in August 1997, from the abdomen of a hermit crab, Calcinus
gaimardii (H. Milne Edwards). Examination of these specimens showed that
they belong to the athelgine genus Pseudostegias Shiino, but cannot be placed
into any described species. We describe this new species as P. macdermotti
and compare it to P. dulcilacuum Markham, which it most closely resembles.
This is the first record of a Pseudostegias from the genus Calcinus. A list and
key of all described Pseudostegias species are provided.

Bopyrid isopods of the subfamily Athel-
ginae are obligate parasites found on the ab-
domen of hermit crabs and lithodids (De-
capoda: Paguroidea). The seven genera in
this subfamily are defined primarily on the
combination of lateral plates and pleopods
on the pleomeres of the female. The genus
Pseudostegias is distinguished by the pres-
ence of long uniramous lateral plates on
pleomeres 1—4, and biramous pleopods.
The lateral plates on pleomere 5 are re-
duced to a dorsally produced globular or
bifurcated lobe.

One of us (JDW) collected a series of
hermit crabs in Sanur, Bali, Indonesia dur-
ing August 1997. One of these crabs was
found to bear an immature female bopyrid
with a male inside her brood plates. The
specimens belong to the genus Pseudoste-
gias and represent a new species most sim-
ilar to P. dulcilacuum Markham, 1982. A
list of the species of Pseudostegias, with
their known localities and hosts, and a key
to their identification are provided.

Methods

Hermit crabs inhabiting gastropod shells
were collected intertidally in Sanur, Bali,
Indonesia on 5—6 August 1997. Specimens
were relaxed in 3% magnesium chloride,
fixed in 10% formalin-seawater solution
and stored in 70% ethanol. The shells were
cracked using a hammer and pliers and the
crabs removed and examined for parasites.

Camera lucida sketches made of speci-
mens were scanned into a Macintosh®
computer. Images were then prepared using
the programs Adobe Photoshop® and Ado-
be Illustrator®.

Shield length (SL) is provided as an in-
dicator of size for the host crabs. Isopod
size is given as total body length (anterior
margin of head to posterior margin of pleo-
telson). Measurements were made to 0.01
mm using an ocular micrometer.

Specimens are deposited in the Department
of Invertebrates, American Museum of Nat-
ural History, New York, U.S.A. (AMNH).

VOLUME 112, NUMBER 4

715

Fig. 1.

Pseudostegias macdermotti, new species. Female, 3.6 mm, AMNH 17877, holotype. A, dorsal view;

B, ventral view. L = lateral plate; P = pleopod; T = telson; U = uropod; numbers indicate pleon segment.

Scale = 0.5 mm.

Family Bopyridae Rafinesque, 1815
Subfamily Athelginae Codreanu and
Codreanu, 1956
Genus Pseudostegias Shiino, 1933
Pseudostegias macdermotti, new species
Figs. 1-4

Material examined.—Holotype: female
(3.6 mm), infesting male Calcinus gaimar-
dii (H. Milne Edwards) (2.66 mm SL;
AMNH 17879), inhabiting shell of Drupel-
la cornus (RGding), 08°41’S, 115°15’E,
Sanur, Bali, Indonesia, intertidal, coll. J. D.
Williams, 6 Aug 1997 (AMNH 17877). Al-
lotype: male (1.06 mm), same data as ho-
lotype (AMNH 17878).

Type locality.—Sanur, Bali, Indonesia,
Pacific Ocean.

Description.—Female (Figs. 1—2). Body
length 3.60 mm, maximal width 1.83 mm,
head length 0.75 mm, head width 0.44 mm,
pleon length 1.64 mm. Pereon not distorted
due to immaturity; pleon deflected to the
right. Body outline narrow and elongated.
All body regions and most pereomeres dis-
tinctly segmented (Fig. 1A, B).

Head not produced due to immaturity.
Eyes present approximately 1/4 distal from
anterior margin. Antenna (Fig. 2C) of 7 ar-
ticles; antennule (Fig. 2C) of 3 articles, dis-
tal margins of segments with setae. Maxil-
liped (Fig. 2E) with low rounded spur; palp
absent; posterior margin setose. Barbula un-
developed.

Pereon composed of 7 pereomeres,

716 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Pseudostegias macdermotti, new species. Female, 3.6 mm, AMNH 17877, holotype. A, right pereo-
pod 1; B, left pereopod 7; C, right antenna and antennule; D, dorsal view of fifth lateral plates, pleotelson, and
uropods (L = lateral plate; P = pleopod; T = telson; U = uropod; numbers indicate pleon segment); E, left
maxilliped. Scale = 0.1 mm (A, B, D), 0.05 mm (C), and 0.02 mm (E).

VOLUME 112, NUMBER 4

broadest across pereomeres 4 and 5, taper-
ing anteriorly and posteriorly. Pereomeres
2-7 incompletely fused. Coxal plates on
sides of pereomeres all similar. Oostegites
completely enclosing brood pouch; poste-
riormost oostegite with fringe of setae on
posterior margin. Pereopods 1—4 (Fig. 2A)
of about same size, pereopods 5-7 (Fig. 2B)
slightly smaller and shorter. Propodus of all
pereopods with distally-directed medio-
ventral projection (Fig. 2B); distal region of
projection with small, rounded tubercles;
short setae at distal tip. First 2 pereopods
surrounding head region; no large gaps be-
tween any pereopods.

Pleon with 6 distinct pleomeres. Pleo-
meres 1—4 with extended lanceolate, distal-
ly rounded, biramous pleopods and unira-
mous lateral plates (Fig. 1B); pleomere 5
(Figs. 1A, 2D) with biramous pleopods and
dorsally produced globular plate, only
slightly bifurcated posteriorly and with pa-
pillate surface; pleotelson (Fig. 2D) with
pair of large lanceolate, distally rounded,
uniramous uropods and rounded, dorsally
produced, papillate pleotelson.

Male (Figs. 3—4). Length 1.06 mm, head
length 0.09 mm, head width 0.21 mm,
pleon length 0.33 mm. Occurring under
oostegites of female (Fig. 1B); directed an-
teroposteriorly.

Head suboval, widest posteriorly, incom-
pletely fused with lst segment of pereon.
Large eyes near posterolateral margin. An-
tenna (Fig. 4C) of 7 articles, distally setose;
extending posterolaterally from head; an-
tennule of 3 articles (Fig. 4C).

Pereomeres 2—6 broadest, tapering ante-
riorly and posteriorly. Pereomeres 1—4 di-
rected laterally; 5—7 directed posterolater-
ally. All segments of body except for head
with irregular dark pigmentation pattern.
All pereopods (Fig. 4A, B) of equal size,
all articles distinctly separated.

Pleon tapering posteriorly, pleomeres di-
rected laterally and fringed with setae. First
segment incompletely fused to 2nd seg-
ment; all other segments fused. No midven-
tral tubercles; 5 pairs of tuberculiform ple-

V9,

opods (Fig. 3A). Pleotelson (Fig. 4D)
notched medially, produced distolaterally,
distal ends of lobes with setae; uropods ab-
sent.

Distribution.—Found on hermit crab,
Calcinus gaimardii from Sanur, Bali, In-
donesia; intertidal.

Etymology.—This species is named in
honor of Dr. John J. McDermott (Franklin
and Marshall College, Pennsylvania,
U.S.A.) for his considerable contributions
to the biology and systematics of numerous
marine invertebrates, including bopyrids.
The specific name is spelled to conform to
the ICZN Recommendations on the For-
mation of Names, Appendix D 21(a) (ICZN
1985-7197):

Remarks.—Pseudostegias macdermotti
most closely resembles females of P. dul-
cilacuum Markham, 1982 from Hong
Kong, Thailand and South Korea, in the
number of segments in the antennae, shape
of pleopods, and presence of a papillate
pleotelson. Pseudostegias macdermotti dif-
fers from females of P. dulcilacuum in the
degree of subdivision of the 5th pleomere
lateral plate. In P. macdermotti, the undi-
vided globular 5th pleomere lateral plate
has only a hint of a medial seam, while P.
dulcilacuum has a strongly bifurcated 5th
pleomere lateral plate. The only other spe-
cies in the genus which has an undivided
5th pleomere lateral plate is P. otagoensis
Page, 1985, from New Zealand. Pseudo-
stegias otagoensis differs from P. macder-
motti in both the shape of its 5th pleomere
lateral plate, which is extended anteropos-
teriorly rather than mesiolaterally as in P.
macdermotti, and in the distally pointed
shape of the pleopods which are rounded in
P. macdermotti. Some authors (Shiino
1933, Lemos de Castro 1965) have incor-
rectly described species of Pseudostegias as
lacking Sth pleomere lateral plates, al-
though all species in this genus possess
such plates (Markham 1982). We consider
the shape of the 5th pleomere lateral plate
to be diagnostic in species of Pseudoste-
gias, since it has been shown to be constant

718

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3:
B, ventral view. Scale = 0.1 mm.

within species, including P. dulcilacuum,
between juveniles and adults (Lemos de
Castro 1965: figs. 2—4, Markham 1985:
figs. 26-27). The key given below provides
additional characters for separating the spe-
cies of Pseudostegias based on females.
Male P. macdermotti differ from those of
P. dulcilacuum in having a fused head with
the first pereon segment and in pleotelson
shape. The pleotelson of the male P. mac-
dermotti is notched medially and produced

Pseudostegias macdermotti, new species. Male, 1.06 mm, AMNH 17878, allotype. A, dorsal view;

distolaterally while P. dulcilacuum possess-
es a pleotelson which tapers to a blunt
point. Male P. macdermotti differ from
those of P. otagoensis in posessing eyes
and in pleotelson shape. Pseudostegias ota-
goensis lacks eyes and has a tapered, point-
ed pleotelson. The location of the male in-
side the brood plates of the female is con-
sistent with other reports for this genus
(Shiino 1933, Markham 1982), although the
male of P. hapalogasteri was found outside

VOLUME 112, NUMBER 4

Porshe
A |

719

Fig. 4. Pseudostegias macdermotti, new species. Male, 1.06 mm, AMNH 17878, allotype. A, left pereopod
1; B, left pereopod 7; C, right antenna and antennule; D; Dorsal view of pleon. Scale = 0.05 mm.

the brood plates on the posterior end of the
female (Shiino 1950).

Ecology.—A total of 43 hermit crabs
were collected from Sanur, Bali on Aug. 5—
6, 1997. The majority were Calcinus gai-
mardii (34 specimens) and only a single
specimen of this crab was found with a bo-
pyrid parasite (3% prevalence). This is the

first report of a species of Pseudostegias on
a species of Calcinus. The other crabs col-
lected were Calcinus latens (Randall) (4
specimens), Clibanarius sp. (1 specimen),
Dardanus sp. (2 specimens), and 2 uniden-
tified hermits. No specimens of any of these
other species were parasitized. The overall
rate of bopyrid occurrence was 2.3%, which

720

is comparable to rates found in large
(1000+) sample sizes (Thompson 1901,
Pike 1961).

The species of Pseudostegias.—The ge-
nus Pseudostegias now contains the follow-
ing six species:

P. atlantica Lemos de Castro, 1965, Bra-
zil, on Clibanarius sp. (Lemos de Castro
1965).

P. dulcilacuum Markham, 1982, Hong
Kong, on Diogenes aff. edwardsii (de
Haan) (Markham 1982); Thailand, on Cli-
banarius merguiensis de Man (Markham
1985); South Korea, on Diogenes sp. (Kim
& Kwon 1988).

P. hapalogasteri Shiino, 1950, Japan, on
Hapalogaster dentata (de Haan) (Shiino
1950).

P. macdermotti, n. sp., Bali, Indonesia,
on Calcinus gaimardii (H. Milne Edwards).

P. otagoensis Page, 1985, New Zealand,
on Paguristes barbatus Heller (Page 1985).

P. setoensis Shiino, 1933, Japan, on Cli-
banarius bimaculatus (de Haan) (Shiino
1933); Taiwan, on C. striolatus Dana (Shi-
ino 1958); Hong Kong, on C. bimaculatus
(de Haan) and C. ransoni Forest (Markham
1982); Thailand, on C. padavensis de Man
(Markham 1985); New Caledonia, on “‘Tri-
zopagurus” sp. (Markham 1994) =Strio-
pagurus boreonotus Forest, 1995.

Discussion.—There is a strong possibil-
ity that the material reported as Pseudoste-
gias setoensis by Shiino (1933), Markham
(1985), and Markham (1994) actually rep-
resents three distinct species. Based on the
published illustrations and descriptions, the
5th pleomere lateral plates on each speci-
men are quite different, the number of pe-
reopods found overlapping the head region
is not identical, and the shape of the pleo-
pods is more variable than has been dem-
onstrated for other athelgine species. Spec-
imens would need to be examined to make
a final determination on their status. How-
ever, neither P. setoensis sensu Markham
(1985) or Markham (1994) closely resem-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

bles our specimens of P. macdermotti n. sp.
The key below contains only P. setoensis
sensu Shiino (1933).

All available evidence indicates that spe-
cies of Pseudostegias, like all athelgine bo-
pyrids, are obligate abdominal parasites and
records from hermit crab branchial cavities
(e.g., Kim & Kwon 1988: p. 215) should
be regarded as erroneous.

Key to the species of Pseudostegias
(based on females)

1. Pleomere 5 lateral plate undivided
— Pleomere 5 lateral plate divided
2. Pleomere 5 lateral plate extended antero-
posteriorly, pleopods pointed distally ..
whe SO ee P. otagoensis
— Pleomere 5 lateral plate extended mesi-
olaterally, pleopods rounded distally ...
ohicic Gee Sh Cee ee ee P. macdermotti
3. Pleomere 5 lateral plate posteriorly sep-
arated P. atlantica
— Pleomere 5 lateral plate wholly separated

4. Pleomere 5 lateral plate lobes large, pa-
pillate P. dulcilacuum
— Pleomere 5 lateral plate lobes small,
widely separated
5. Pleotelson large, uropods shorter than
pleopods = 2245s" .. Aamee P. hapalogasteri
— Pleotelson small, uropods longer than
pleopods .. P. setoensis sensu Shiino, 1933

oe © © © © © ee ee ee ele

Acknowledgments

Drs. Paul Cassidy (Western Washington
University) and Alan Harvey (Georgia
Southern University) kindly assisted in the
identification of hermit crabs. Two anony-
mous reviewers contributed greatly to the
final product. This work was supported by
a grant from the Lerner-Gray Fund for Ma-
rine Research (American Museum of Nat-
ural History) and a Libbie Hyman Memo-
rial Scholarship (Society for Integrative and
Comparative Biology) to JDW.

Literature Cited

Codreanu, M., & R. Codreanu. 1956. Sur l’ Anisarthrus
pelseneeri, épicaride parasite abdominal de la

VOLUME 112, NUMBER 4

crevette Athanas nitescens; sa présence dans la
Mer Noire et la dispersion du genre Anisar-
thrus.—Bulletin Biologique de la France et de
Belgique 90: 111-121.

Forest, J. 1995. Crustacea Decapoda Anomura: Révi-
sion du genre Trizopagurus Forest, 1952 (Di-
ogenidae), avec |’établissement de deux genres
nouveaux. /n A. Crosnier, ed., Résultats des
Campagnes MUSORSTOM, vol. 13.—Mémo-
ires du Muséum National d’Histoire Naturelle
163:9-149.

[ICZN] International Commission of Zoological No-
menclature. 1985. International Code for Zoo-
logical Nomenclature, 3rd edition. International
Trust for Zoological Nomenclature: London,
338 pp.

Kim, H. S., & D. H. Kwon. 1988. Bopyrid isopods
parasitic on decapod crustaceans in Korea.—
Korean Journal of Systematic Zoology special
issue 2:199—221.

Lemos de Castro, A. 1965. Crustaceos is6podos epi-
carideos do Brasil. I: descrigao de uma espécie
nova do género ““Pseudostegias’’ Shiino (Iso-
poda, Bopyridae).—Revista Brasileira de Biol-
ogia 25(1):105—108.

Markham, J. C. 1982. Bopyrid isopods parasitic on
decapod crustaceans in Hong Kong and south-
ern China. Pp. 325-391 In B. S. Morton and C.
K. Tseng, eds., The marine flora and fauna of
Hong Kong and Southern China, vol. 1. Hong
Kong University Press, Hong Kong.

#21

. 1985. Additions to the bopyrid fauna of Thai-

land.—Zoologische Verhandelingen 224:63 pp.

. 1994. Crustacea Isopoda: Bopyridae in the
MUSORSTOM collections from the tropical
Indo-Pacific I. Subfamilies Pseudioninae (in
part), Argeiinae, Orbioninae, Athelginae and
Entophilinae. Jn A. Crosnier, ed., Résultats des
Campagnes MUSORSTOM, vol. 10.—Mémo-
ires du Muséum National d’Histoire Naturelle
161:225—253.

Page, R. D. 1985. Review of the New Zealand Bo-
pyridae (Crustacea: Isopoda: Epicaridea).—
New Zealand Journal of Zoology 12(2):185—
Ze.

Pike, R. B. 1961. Observations on Epicaridea obtained
from hermit-crabs in British waters, with notes
on the longevity of the host-species.—Annals
and Magazine of Natural History, ser. 13, 4(40):
225-240.

Rafinesque-Schmaltz, C. S. 1815. Analyse de la nature
ou tableau de l’univers et des corps organisés.
Palermo, 224 pp.

Shiino, S. M. 1933. Bopyrids from Tanabe Bay.—
Memoirs of the College of Science, Kyoto Im-
perial University, ser. B, 8(3):249—300.

. 1950. Notes on some new bopyrids from Ja-

pan.—Mie Medical Journal 1:151—167.

. 1958. Note on the bopyrid fauna of Japan.—
Report of Faculty of Fisheries, Prefectural Uni-
versity of Mie 3(1):67—73, pl. 3.

Thompson, M. T. 1901. A new isopod parasitic on the
hermit crab.—Bulletin of the U.S. Fish Com-
mission 21:53—56.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(4):722—738. 1999.

Caprella kuroshio, a new species (Crustacea: Amphipoda:
Caprellidae), with a redescription of Caprella cicur Mayer, 1903, and
an evaluation of the genus Metacaprella Mayer, 1903

Atsushi Mori

Seto Marine Biological Laboratory, Kyoto University, Shirahama, Wakayama 649-2211, Japan

Abstract.—A new species of Caprella, C. kuroshio, is described based on
the specimens collected from the Pacific coast of central Japan. Caprella cicur
Mayer, 1903, closely related to the new species, is redescribed based on the
type material. The new species differs from C. cicur in the shape of the per-
eonites and the second gnathopod. The female of the new species has a pair
of abdominal appendages, which is a diagnostic character of the genus Meta-
caprella Mayer, 1903. However, the abdominal appendage is considered to be
a polymorphic and symplesiomorphic character in Caprella and Metacaprella.
Therefore, the genus Metacaprella is not recognised, and the new species is

assigned to the genus Caprella.

In 1991, during a study of the caprellid
fauna of the west coast of Shikoku, western
Japan, numerous specimens of a caprellid
species were collected by the author from
the red algae, Gelidium elegans Kiitzing, at
Kashiwajima Island, Kochi Prefecture. Af-
terward, many individuals of the same spe-
cies also were found on G. elegans in Ku-
shimoto and Shirahama, Wakayama Prefec-
ture, in 1992 and 1996 respectively. Fur-
thermore, a single specimen of the species
was collected from the drifting brown al-
gae, Sargassum sp., in Suruga Bay in 1993.
Detailed examination revealed that all of
these specimens belong to a new species.
The description of the species is presented
here.

The type series is deposited in the Seto
Marine Biological Laboratory, Kyoto Uni-
versity (SMBL).

Caprella kuroshio, new species
Figs. 1-5

Type material.—Holotype (SMBL Type
No. 391): male, 26 Apr 1991, on Gelidium
elegans, 2 m in depth, Sankaku-bae, Kash-
iwajima Island (32°46'N, 132°38’E), Kochi

Prefecture, Japan. Paratypes (SMBL Type
No. 392): 10 males, 10 females, collected
together with holotype.

Additional material examined.—10
males, 10 females, 4 Jun 1992, on Gelidium
elegans, 1 m in depth, Sabiura, Kushimoto
(33°28'’N, 135°46’E), Wakayama Prefec-
ture, Japan; 1 male, 10 Apr 1993, on drift-
ing Sargassum sp., off the mouth of Fuji
River, Suruga Bay (35°04’N, 138°39’B),
Shizuoka Prefecture, Japan; 5 males, 5 fe-
males, 16 Jun 1996, on Gelidium elegans,
depth unrecorded, Rinkai, Shirahama
(33°42'N, 135°21'E), Wakayama Prefec-
ture, Japan.

Description of holotype.—Body (Fig.
1A) 9.65 mm long, smooth. Head with
blunt triangular projection directed straight
forward; pereonite 1 shorter than head; per-
eonite 2 longer than pereonite 3, with single
acute projection on ventral surface between
insertions of second gnathopods (Fig. 1B).

Antenna 1 (Fig. 1A) longer than a half
of the body length; flagellum consisting of
14 segments, shorter than peduncle. Anten-
na 2 (Fig. 1A) shorter than peduncle of an-
tenna 1; peduncular segments 3—5 with 2

VOLUME 112, NUMBER 4 723

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Z

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rw

Fig. 1. Caprella kuroshio, new species, holotypic male: A, body, lateral view; B, pereonite 2, lateral view;
C, abdomen. Scales: A, B = 1.0 mm; C = 0.1 mm.

724

rows of dense long setae on ventral surfac-
es; flagellar segment 1 with 2 rows of dense
long setae on ventral surface.

Upper lip (Fig. 2A) semicircular, slightly
concave on apical margin, densely pubes-
cent. Inner and outer lobes of lower lip
(Fig. 2B) well-developed, round, densely
pubescent on apicomedial margins; man-
dibular process developed.

Mandible (Fig. 2C—D) without palp. Left
and right sides consisting of 5- and 4-
toothed incisor, 5- and 6-toothed lacinia
mobilis, and setal row of 3 and 2 plumose
setae respectively. Molar process well-de-
veloped, with 1 plumose seta; left molar
with triangular stout projection; right molar
with pubescent molar extension.

Maxilla 1 (Fig. 2E) without inner lobe.
Its outer lobe rectangular, with 3 simple and
4 serrate spines on apical margin. Palp 2-
segmented. Its distal segment bearing a row
of teeth and 6 spines on apical margin, 7
setae on inner margin, 2 rows of setae on
ventral surface.

Inner lobe of maxilla 2 (Fig. 2F) shorter
than outer lobe, oval, with 2 rows of nu-
merous setae on apicomedial margin. The
outer lobe rectangular, with 2 rows of setae
on apical margin.

Inner lobe of maxilliped (Fig. 3A) small,
with 3 short spines on apical margin, and
numerous marginal setae. Outer lobe small,
subequal to palpal segment 1 in length,
bearing 1 long stout seta on apex, 7 short
spines on medial margin, and numerous
marginal setae. Palp 4-segmented; segment
1 short; segments 1—2 bearing numerous
long setae on medial margins, and segment
3 on apicomedial surface; segment 4 fal-
cate, longer than apical setae on segment 3,
with rows of setulae on grasping margin.

Gnathopod 1 (Fig. 3B) robust. Its ischi-
um and merus short, with setae on ventral
surfaces. Carpus expanded ventrally, bear-
ing numerous setae. Propodus bearing rows
of setae on medial surface. Palm bearing a
pair of grasping spines at the proximal cor-
ner, its grasping margin straight and mi-
nutely serrate, bearing rows of setae. Dac-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

tylus falcate, bearing rows of numerous
teeth on grasping margin.

Gnathopod 2 (Figs. 1A, 3C) attached to
midpoint of pereonite 2. Basis shorter than
pereonite 2 in length, bearing laterodistal
and mediodistal triangular projections. Is-
chium bearing lateral triangular projection.
Propodus oblong, two times longer than ba-
sis. Palmar corner produced, bearing single
large grasping spine on apex. Palmar mar-
gin slightly concave, bearing a pair of small
grasping spines on proximal end, subdistal
poison tooth, and distal triangular projec-
tion. Dactylus curved and narrowed distal-
ly.
Gills (Fig. 1A) present on pereonites 3—
4, large, oval.

Pereopods 5—7 (Figs. 4A—C) morpholog-
ically similar, but increasing in size poste-
riorly. Basis bearing triangular dorsal pro-
jection at posterior end. Merus inflated dor-
sally. Propodus longest; palm bearing a pair
of grasping spines on proximal corner, and
rows of short setae on slightly concave
grasping margin. Dactylus bearing minute
serration on curved grasping margin.

Abdomen (Fig. 1C) consisting of a pair
of appendages, a pair of lateral lobes, and
single dorsal lobe. Abdominal appendage 2-
segmented; three setae present on both
proximal and distal segments; tip of distal
segment round. Lateral lobes bearing 6—7
proximal and 4—5 distal setae. Penes large,
situated medially.

Description of paratypic female.—Body
(Fig. 5A—-B) 6.90 mm long. Pereonite 2
subequal to pereonite 3 in length.

Antenna 1 (Fig. 5A) subequal to a half
of the body length. Flagellum consisting of
13 segments, subequal to peduncle in
length. Antenna 2 (Fig. 5A) longer than pe-
duncle of antenna 1.

Gnathopod 2 (Figs. 5A, C) attached to
anterior end of pereonite 2. Propodus oval.
Palmar margin slightly convex, bearing
subdistal small poison tooth and distal tri-
angular projection. Dactylus slightly
curved.

Abdomen (Fig. 5D) consisting of a pair

VOLUME 112, NUMBER 4 72S

Fig. 2. Caprella kuroshio, new species, holotypic male: A, upper lip; B; lower lip; C, left mandible; D, right
mandible; E, maxilla 1; EK maxilla 2. Scales: 0.1 mm.

726 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Caprella kuroshio, new species, holotypic male: A, maxilliped; B, gnathopod 1; C, gnathopod 2 with
details of palmar projections indicated by arrows. Scales: A, B = 0.1 mm; C = 1.0 mm.

VOLUME 112, NUMBER 4 Tat

Fig. 4. Caprella kuroshio, new species, holotypic male: A, pereopod 5; B, pereopod 6; C, pereopod 7.
Scales: 0.1 mm.

728 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

=e ea 7
A a aN
WN oe

Fig. 5. Caprella kuroshio, new species, female: A, body, lateral view; B, pereonite 2, lateral view; C,
gnathopod 2; D, abdomen. Scales: A, B = 1.0 mm; C, D = 0.1 mm.

VOLUME 112, NUMBER 4

of appendages, a pair of lateral lobes, and
single dorsal lobe. Abdominal appendage
unsegmented, small, bearing 1 subterminal
seta. Lateral lobe with 2 distal setae.

Color in life-—Specimens occurring on
the red algae Gelidium elegans are scarlet
to wine red. A single male collected from
the drifting Sargassum sp. in Suruga Bay is
yellowish.

Etymology.—The specific name is from
the Kuroshio current.

Remarks.—Caprella kuroshio resembles
Caprella penantis Leach, 1814 in general
appearance. However, the former is distin-
guishable from the latter by: pereonite 2
with ventral projection; male antenna 1 lon-
ger than half the body length; basis of male
gnathopod 2 with mediodistal triangular
projection; palmar margin of male gnatho-
pod 2 with subdistal poison tooth; female
abdomen bearing a pair of appendages.

Caprella kuroshio is also similar to Ca-
prella cicur Mayer, 1903, an endemic spe-
cies to South African waters (Griffiths
1974a, 1974b, 1975, 1976). The species has
never been redescribed in detail since May-
er’s original description. Here, C. cicur is
redescribed below using the type material
and compared to C. kuroshio.

The type series has been deposited in the
South African Museum, Cape Town
(SAM).

Caprella cicur Mayer, 1903
Figs. 6—10

Caprella cicur Mayer, 1903:75, 97, pl. 4
(figs. 5—7), pl. 8 (figs. 3-5); Griffiths,
1974a:255; Gmriffiths, 1974b:331; Grif-
fiths, 1975:174; Griffiths, 1976:86, 104.

Material examined.—Syntypes (labeled
as “‘Cotypes’’): 1 male, 1 female (SAM
1312), Table Bay, Cape Town, on Palinurus
lalandi, collected by W. E Purcell, June
1897.

Description of male.—Body (Fig. 6A)
10.40 mm long. Head with small, acute tri-
angular projection directed straight for-
ward; pereonite 1 longer than head; pereo-

729

nite 2 longer than pereonite 3, with acute
ventral projections between insertions of
second gnathopods, and acute lateral pro-
jection on insertion of second gnathopod;
pereonites 3—4 with ventrolateral small pro-
jections on anterior end; pereonite 5 with
dorsolateral small projections and lateral
blunt knobs on anterior end; pereonite 6
with a pair of dorsal minute projections.

Antenna 1 (Fig. 6A) longer than a half
of the body length; flagellum consisting of
14 segments, shorter than peduncle. Anten-
na 2 (Fig. 6A) shorter than peduncle of an-
tenna 1; peduncular segments 3—5 with 2
rows of dense long setae on ventral surfac-
es; flagellar segment 1 with 2 rows of dense
long setae on ventral surface.

Upper lip (Fig. 7A) semicircular, slightly
concave on apical margin, densely pubes-
cent. Inner and outer lobes of lower lip
(Fig. 7B) well-developed, round, densely
pubescent on apicomedial margins; outer
lobe with an apicolateral incomplete fold;
mandibular process developed.

Mandible (Figs. 7C—D) without palp. Its
left and right sides consisting of 5-toothed
incisor, 5-toothed and irregularly-toothed
lacinia mobilis, and setal row of 3 and 2
plumose setae respectively. Molar process
well-developed, with 1 plumose seta, and
triangular stout projection.

Maxilla 1 (Fig. 8A) without inner lobe.
Its outer lobe rectangular, with 7 bifid
spines on apical margin. Palp 2-segmented.
Its distal segment bearing a row of teeth
and 8 spines on apical margin, 4 setae on
inner margin, rows of setae on ventral sur-
face.

Inner lobe of maxilla 2 (Fig. 8B) shorter
than outer lobe, oval, with 2 rows of nu-
merous setae on apicomedial margin. The
outer lobe rectangular, with 2 rows of nu-
merous setae on apical margin.

Maxilliped removed.

Gnathopod 1 (Fig. 8C) robust. Its ischi-
um short. Ventral margin of merus pro-
duced anteriorly, bearing numerous setae on
anteroventral surface. Carpus expanded
ventrally, bearing numerous setae. Propo-

730 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Caprella cicur Mayer, 1903, syntypes: A, male, lateral view; B, male abdomen, ventral view; C,
female, lateral view; D, female abdomen, ventral view. Scales: A, C = 1.0 mm; B, D = 0.1 mm.

VOLUME 112, NUMBER 4

Fig. 7.
mandible. Scales: 0.1 mm.

dus bearing rows of setae on medial sur-
face. Palm bearing a pair of grasping spines
at the proximal corner, its grasping margin
straight, bearing rows of setae. Dactylus fal-
cate.

Gnathopod 2 (Fig. 6A) attached to per-
eonite 2 posteriorly. Basis shorter than per-
eonite 2 in length, bearing laterodistal and
mediodistal triangular projections. Ischium
bearing lateral triangular projection. Merus
acutely produced anteroventrally. Propodus
oblong, 2.5 times longer than basis. Palmar
corner produced, bearing single large grasp-
ing spine on apex. Palmar margin without

731

l

cana)

es
pes
=
SS
ser
=
=
ES oo
~-

Caprella cicur Mayer, 1903, syntypic male: A, upper lip; B, lower lip; C, left mandible; D, right

poison tooth, with distal triangular projec-
tion. Dactylus curved and narrowed distal-
ly.

Gills (Fig. 6A) present on pereonites 3—
4, large, elliptical.

Pereopods 5-7 fell off.

Abdomen (Fig. 6B) consisting of a pair
of appendages, a pair of lateral lobes, and
single dorsal lobe. Abdominal appendage 2-
segmented, bearing 10 setae; tip of distal
segment round. Lateral lobe bearing 4 prox-
imal and 1 distal setae. Penes large, situated
medially.

Description of female.—Body (Fig. 6C)

‘32 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 8. Caprella cicur Mayer, 1903, syntypic male: A, maxilla 1; B, maxilla 2; C, gnathopod 1. Scales: 0.1

nm.

8.40 mm long. Pereonite 1 slightly shorter
than head; pereonite 2 slightly longer than
pereonite 3; pereonite 4 without ventrolat-
eral small projections on anterior end; per-
eonite 6 without dorsal projections.

Antenna 1 (Fig. 6C) longer than a half of
the body length. Flagellum consisting of 15
segments, slightly shorter than peduncle.
Antenna 2 (Fig. 6C) slightly longer than pe-
duncle of antenna 1.

VOLUME 112, NUMBER 4

\
x

i \
‘ \ Mi i ’ \\

4)
Py

NN)

\

Fig9!

Inner lobe of maxilliped (Fig. 9A) small,
with 2 short spines on apical margin, and
numerous marginal setae. Outer lobe small,
subequal to palpal segment 1 in length,
bearing 1 long stout seta on apex, 8 short
spines on medial margin, and numerous
marginal setae. Palp 4-segmented; segment
1 short; segment 2 bearing numerous long
setae on medial margin, and segment 3 on
apicomedial surface; segment 4 falcate, lon-
ger than apical setae on segment 3.

Gnathopod 2 (Figs. 6C, 9B) attached to
pereonite 2 anteriorly. Propodus oval. Pal-
mar margin slightly convex. Dactylus
slightly curved.

Abdomen (Fig. 6D) consisting of a pair
of lateral lobes and single dorsal lobe, with-
out appendages.

Pereopods 5-7 (Fig. 10) morphologically
similar. Basis bearing triangular dorsal pro-
jection at posterior end. Merus inflated dor-
sally. Palm of propodus bearing a pair of
grasping spines on proximal corner, and
rows of short setae on slightly concave
grasping margin. (Pereopods 5—7 were de-

733

Caprella cicur Mayer, 1903, syntypic female: A, maxilliped; B, gnathopod 2. Scales: 0.1 mm.

tached from the pereonites, and scattered in
the vial. Therefore, it is indeterminable to
which specimen, and to which pereonite,
each pereopod was attached.)
Remarks.—Caprella kuroshio is distin-
guishable from C. cicur by: pereonites 1—2
not elongate; pereonites 5—6 without dorsal
projections; pereonites 3—5 without lateral
projections; insertion of gnathopod 2 with-
out lateral projection; palmar margin of
gnathopod 2 with poison tooth; female ab-
domen bearing a pair of appendages.

Discussion

The most remarkable feature of the new
species is the presence of paired abdominal
appendages in the female. If the generic di-
agnosis of Mayer (1903) had been adopted,
this species would be assigned to the genus
Metacaprella Mayer, 1903. However, Me-
tacaprella has been considered a question-
able genus among the Caprellidea, because
it was established only on the basis of the
presence of the abdominal appendages in

734

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 10. Caprella cicur Mayer, 1903, pereopods 5—7 of syntypes. Scales: 0.1 mm.

the female (Laubitz 1970). In this section,
I assess the taxonomic value of the abdom-
inal appendages based on a survey of the
literature to determine whether the genus
Metacaprella is a valid taxon or not.
History of Metacaprella.—Mayer (1903)
suggested the possibility of splitting the ge-
nus Caprella, and tentatively proposed the
genus Metacaprella for Caprella anomala
Mayer, 1903 and Caprella kennerlyi Stimp-
son, 1864, based on the presence of paired
abdominal appendages in the female. At
that time, however, he hesitated separating
the two species from Caprella. Subsequent-
ly, Dougherty & Steinberg (1953) adopted
Mayer’s generic distinction. They included
the two species to Metacaprella, and des-
ignated M. kennerlyi as the type species of
the genus. They also referred Caprella fer-
rea Mayer, 1903 to Metacaprella provision-
ally because of its general resemblance to
M. kennerlyi and M. anomala, even though

no females of C. ferrea had been found at
that time.

In the catalog of the world caprellids,
McCain & Steinberg (1970) listed these
three species in the genus Metacaprella.
McCain (1968) and Laubitz (1970) criti-
cized Mayer’s generic distinction on the ba-
sis of unreliable characters such as the or-
ganization of the abdomen. Laubitz (1970)
stated, ““‘Thus the separation of Metaca-
prella from Caprella, solely on the basis of
the female abdomen, is highly unsatisfac-
tory.’ However, she inconsistently retained
the genus Metacaprella consisting of M.
anomala and M. kennerlyi. She restored M.
ferrea to Caprella, because there was no
sign of the presence of appendages on the
female abdomen of the species. Laubitz
(1972) transferred Caprella horrida G. O.
Sars, 1877 to Metacaprella based on the
presence of a pair of small appendages on
the female abdomen. Subsequent authors

VOLUME 112, NUMBER 4

also have allowed the genus Metacaprella
to stand in their caprellid faunal studies
(Vassilenko 1974, Arimoto 1976, Martin
197 1).

Ontogeny of the abdominal appendag-
es.—Sakaguchi (1989) described the post-
marsupial development of Caprella scaura
diceros Mayer, 1890, reared in the labora-
tory. All the hatchlings (body length 1.25
mm, flagellum of antenna 1 2-segmented,
sexes still indistinct) had paired abdominal
appendages bearing a single apical seta
(Fig. 11A). In males, the appendage was
still 1-segmented and small in the specimen
with 5-segmented flagellum of antenna 1
(body length 2.62 mm), but became larger
and 2-segmented in the older specimens
(Figs. 11B—E). In contrast, the appendages
of the female had been reduced in the spec-
imen in the early stage (body length 2.66
mm, flagellum of antenna 1 5-segmented),
and only their apical seta was left as a ves-
tigial structure (Figs. 11F—H).

Takeuchi (1989) also thoroughly de-
scribed the postmarsupial development of
Caprella danilevskii Czerniavski, 1868, C.
okadai Arimoto, 1930, and C. generosa Ar-
imoto, 1977, reared in the laboratory. In
these three species, all the early juveniles
had paired abdominal appendages bearing
single apical setae. The appendages grew
larger in males, whereas they were reduced
in females, as in C. scaura diceros.

The postmarsupial development of Me-
tacaprella species have not been investi-
gated, except for Mayer’s (1903) illustration
of the abdomen of a juvenile of M. kenner-
lyi. The abdomen of the juvenile bears a
pair of unsegmented small appendages with
single apical setae. It should be noticed that
the morphological features of the append-
age are almost identical to those of the
above-mentioned four Caprella species.

Intra-generic morphological variations
of abdominal appendages.—Consultation
of the literature (above) shows that the in-
tra-generic variations for the morphology of
abdominal appendages exist in both Ca-
prella and Metacaprella.

735

In Caprella, the degree of reduction of
the female abdominal appendages varies
specifically. In C. okadai and C. cristi-
brachium Mayer, 1903, the appendages are
completely reduced (Takeuchi 1989, Lee &
Lee 1993), whereas in C. generosa, a pair
of setae is present as a vestigial structure
(Takeuchi 1989). Furthermore, the intraspe-
cific morphological variation of the abdo-
men exists in females of C. danilevskii. The
female from the Japanese waters has only a
pair of vestigial setae on the abdomen (Tak-
euchi 1989), whereas those from the west-
ern North Atlantic and the Mediterranean
have a pair of small abdominal appendages
(McCain 1968, Krapp-Schickel 1993). In
the male, on the other hand, the morphol-
ogy of the abdominal appendages varies
specifically. In C. simia Mayer, 1903, they
are distinctly 2-segmented (Mori & Yamato
1993), whereas in C. monoceros Mayer,
1890 and C. santosrosai Sanchez-Moyano,
Jiménez-Martin, & Garcia-Gomez, 1995,
their segmentation is not distinct (Aoki &
Kikuchi 1990, Sdanchez-Moyano et al.
1985):

In Metacaprella, the degree of develop-
ment of the female abdominal appendages
also varies specifically. The female of M.
anomala has relatively large, incompletely
2-segmented appendages on the abdomen
(Mayer 1903, Laubitz 1970). In females of
M. horrida and M. kennerlyi, the append-
ages are small and unsegmented (Mayer
1903; Laubitz 1970, 1972). The simular
morphology also can be seen in the juvenile
of M. kennerlyi (Mayer 1903).

Concluding remarks.—lt is apparent that
the morphology of the abdominal append-
age changes ontogenetically in the above-
mentioned four Caprella species (Sakagu-
chi 1989, Takeuchi 1989). Furthermore, the
intrageneric, and even intraspecific, mor-
phological variations of the abdominal ap-
pendage exist in Caprella and Metacaprella
species. These facts emphasize that the ex-
ternal morphology of the abdominal ap-
pendage is not an ontogenetically and mor-
phologically stable character enough to di-

736 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 11. Caprella scaura diceros Mayer, 1890, juveniles, body and abdomen: A, hatchling (body length 1.25
mm); B, male (body length 2.62 mm); C, male (body length 3.75 mm); D, male (body length 5.40 mm); E,
male (body length 8.86 mm); FE female (body length 2.66 mm); G, female (body length 5.28 mm); H, female
(body length 9.11 mm) (modified from Sakaguchi, 1989).

VOLUME 112, NUMBER 4

agnose a genus, and its taxonomic value
seems to be relatively low.

In the Caprellidea, the loss and reduction
of characters are considered to be apo-
morphic (Larsen 1997). The ontogenetic in-
formation confirms that the reduction of ab-
dominal appendages is an apomorphic char-
acter state in the genus Caprella. Morpho-
logical features of the juvenile abdomens of
the four Caprella species are almost iden-
tical to that of M. kennerlyi. Female abdom-
inal appendages are present in early devel-
opmental stages of both genera. They are
reduced to various degree in later juvenile
stages in Caprella, while they remain in
adults in Metacaprella. Because characters
appear earlier in the ontogeny are consid-
ered to be ancestral and those appear later
derived (Eldredge & Cracraft 1980), the
presence of the abdominal appendage in the
adult female of Metacaprella is not an au-
tapomorphy of the genus, but a symple-
siomorphy of the species assigned to the ge-
nus.

Metacaprella and Caprella shares almost
all diagnostic characters, and the morphol-
ogy of female abdominal appendages varies
gradually within these two genera, namely,
from incompletely 2-segmented (M. anom-
ala) through unsegmented (M. kennerlyi,
M. horrida and Atlantic C. danilevskii) or
vestigial (C. generosa and Japanese C. dan-
ilevskii) to completely reduced (C. okadai
and C. cristibrachium). No gaps enough to
separate Metacaprella from Caprella could
be found. Therefore, the establishment of
the genus Metacaprella only on the basis of
a plesiomorphic character, i.e., abdominal
appendage, is considered to be inadequate.
Consequently, I decline to adopt the genus
Metacaprella as a separate taxon, and as-
sign the present new species to the genus
Caprella.

Acknowledgements

I would like to thank Mr. Kikuo Okita,
Seto Marine Biological Laboratory, Kyoto
University, and the officers and crews of the

TS)

R/V Tansei-maru, Ocean Research Institute,
University of Tokyo, for collecting seaweed
samples, Dr. Ichiro Takeuchi, Otsuchi Ma-
rine Research Center, University of Tokyo,
for providing me with a copy of his unpub-
lished Ph.D. thesis, and Mr. Masaki Saka-
guchi, Nishinomiya-higashi High School,
for kindly permitting me to use the drawing
of Caprella scaura diceros. Mses Michelle
van der Merwe and Liz Hoenson, South Af-
rican Museum, are gratefully acknowledged
for permitting me to examine the type ma-
terial of Caprella cicur. Thanks are also
due to Prof. Yoshihisa Shirayama, Associ-
ate Prof. Shin Kubota, Drs. Shigeyuki Ya-
mato and Katsumi Miyazaki, Seto Marine
Biological Laboratory, Kyoto University,
and Dr. Masakazu Aoki, Shimoda Marine
Research Center, University of Tsukuba, for
their invaluable advice and critical reading
of the manuscript.

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Arimoto, I. 1976. Taxonomic studies of caprellids
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738

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. 1972. The Caprellidae (Crustacea, Amphipo-
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ed States National Museum Bulletin 278:i—iv +
1-147.

, & J. E. Steinberg. 1970. Amphipoda I. Ca-
prellidea I. Fam. Caprellidae. Jn H.-E. Gruner
& L. B. Holthuis, eds., Crustaceorum Catalogus
2:1-78.

Mori, A., & S. Yamato. 1993. Caprella simia Mayer,
1903 (Crustacea: Amphipoda: Caprellidae) col-
lected from the body surface of a frogfish An-
tennarius striatus (Shaw & Nodder, 1794).—
Nanki Seibutu 35:41—46 (in Japanese with En-
glish summary).

Sakaguchi, M. 1989. [New knowledge of Caprella
scaura diceros (Crustacea, Amphipoda, Caprel-
lidae) (I).J—Hyogo Biology 9:280—285 (in
Japanese).

Sanchez-Moyano, J. E., J. A. Jiménez-Martin, & J. C.
Garcia-Gémez. 1995. Caprella santosrosai n.
sp. (Amphipoda: Caprellidea) from the Strait of
Gibraltar (southern Spain).—Ophelia 43:197—
204.

Takeuchi, I. 1989. Taxonomic and ecological studies
of the Caprellidea (Crustacea, Amphipoda) in-
habiting the Sargassum zone. Unpublished
Ph.D. thesis, Graduate School of Agriculture,
The University of Tokyo, Tokyo, 244 pp (in
Japanese).

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

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112(4):739-745. 1999.

Palaemonella hachijo, a new species of shrimp
(Crustacea: Decapoda: Palaemonidae)
from a submarine cave in southern Japan
Junji Okuno

Coastal Branch of Natural History Museum and Institute, Chiba, 123 Yoshio,
Katsuura, Chiba 299-5242, Japan

Abstract.—A new species of the pontoniine shrimp genus Palaemonella
Dana, P. hachijo, is described and illustrated on the basis of single male spec-
imen from a submarine cave at Hachijo-jima Island, Izu Islands, southern Ja-
pan. It closely resembles P. dolichodactylus Bruce, but is readily distinguished
by the antennal spine overreaching the distal margin of the antennal basicerite,
the posteroventral margin of the fifth abdominal somite armed with an acute
tooth, and the ambulatory propodi not segmented.

The genus Palaemonella Dana, 1852, is
distinguished from other pontoniine genera
in having a hepatic spine, a mandibular
palp, a strong ventromesial spine on the
fourth thoracic sternite, and slender, simple
fingers of the first pereiopod (Chace &
Bruce 1993, Holthuis 1993, Bruce 1994).
Until now, it contained 13 species world-
wide (Chace & Bruce 1993).

During a survey of the caridean shrimp
fauna of Hachijo-jima Island, the Izu Is-
lands, southern Japan, in the summer of
1998, a specimen of Palaemonella was cap-
tured in a submarine cave at a depth of 20
m. Although the specimen closely resem-
bled P. dolichodactylus Bruce, 1991,
known only from the New Caledonian wa-
ters at the depths of 44-250 m (Bruce
1991a, 1991b), the specimen differed mor-
phologically from P. dolichodactylus and
was found to represent a new species. Here-
in this new species is described and illus-
trated. The abbreviation CL indicates the
postorbital carapace length measured in
mm. The specimen is deposited in the
Coastal Branch of Natural History Museum
and Institute, Chiba (CMNH).

Palaemonella hachijo, new species
(Figs. 1-3)
Material.—Holotype, male, 3.3 mm CL,
33°03.5'N, 139°47.9’E, southern Japan, Izu

Islands, Hachijo-jima Island, Occho-ga-
hama, submarine cave, 20 m, 2 Sep 1998,
with SCUBA gear, coll. J. Okuno, CMNH-
ZC 00017.

Diagnosis.—Small sized pontoniine
shrimp with subcylindrical body form. Car-
apace without supraorbital spine. Rostrum
well developed, horizontal, dentate. Anten-
nal spine acute, slightly overreaching dor-
sodistal margin of antennal basicerite. Pos-
teroventral angle of fifth abdominal somite
pointed, armed with acute tooth. Second
pereiopod with merus armed with distov-
entral tooth and carpus lacking strong sub-
marginal tooth. Ambulatory pereiopods
long, slender, propodi not segmented.

Description.—Carapace (Fig. 1A)
smooth, glabrous; orbit feebly developed,
with distinct posterior marginal ridge, with-
out supraorbital spine; antennal spine well
developed, acute, slightly overreaching dor-
sodistal margin of antennal basicerite; he-
patic spine acute, situated at lower and pos-
terior levels of antennal spine; epigasteric
Spine acute; pterygostomian margin round-
ed.

Rostrum (Fig. 1C) well developed, hori-
zontal and straight, 0.85 times as long as
CL, falling short of distal margin of sca-
phocerite. Dorsal carina with 8 acute teeth,
distal tooth subapical, smaller than other

740 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

i
Fig. 1. Palaemonella hachijo, new species. Holotype male (CMNH-ZC 00017). A, carapace and left cephalic
appendages, lateral view; B, anterior part of carapace with cephalic appendages, dorsal view; C, anterior part of

carapace with rostrum, lateral view; D, fourth to sixth abdominal somites, right view; E, telson with uropods,
dorsal view; F posterior margin of telson, dorsal view. Scales: A, 3 mm; B—E, 2 mm; E 0.5 mm.

VOLUME 112, NUMBER 4

posterior teeth, second to eighth teeth equi-
distant, seventh tooth just above orbit, prox-
imal tooth posterior to postorbital margin,
interspaces setose. Ventral carina with 2
acute teeth, situated at distal half of length;
lateral carina feebly developed.

Abdominal somites smooth, glabrous;
pleura of first to third somites broad, round-
ed. Posteroventral margin of fourth somite
with pleuron produced posteriorly. Fifth so-
mite (Fig. 1D) armed with acute tooth at
posteroventral margin of posteriorly pro-
duced pleuron. Sixth somite (Fig. 1D) 0.51
times as long as CL, posteroventral margin
armed with acute tooth. Telson (Fig. 1E)
0.67 times as long as CL, 2 pairs of acute
submarginal dorsal spine; posterior margin
(Fig. 1F) rounded, with 3 pairs of spines,
intermediate spines longest, lateral and in-
termediate spines simple, submedian spines
with setules.

Eye with large globular cornea, stalk
slightly longer than corneal diameter.

Antennular peduncle (Fig. 2A) falling
slightly short of rostral apex. Proximal seg-
ment with anterolateral margin produced,
distal lobe slightly convex, armed sparsely
with short setae, with acute distolateral
spine overreaching distal lobe, ventrome-
dial margin armed with acute tooth, stylo-
cerite well developed, acute, reaching level
of proximal third of the segment, statocyst
well developed, rounded; intermediate seg-
ment with lateral margin feebly convex,
armed densely with short setae, medial mar-
gin fringed sparsely with moderately long
setae; distal segment non-setose, obliquely
articulated with flagella. Upper flagellum
(Fig. 2B) biramous, divided at distal third
of flagellum length, longer ramus about 4
times as long as shorter ramus, with seg-
ments neighboring shorter ramus fringed
with simple setae (Fig. 2C).

Antenna (Fig. 2D) with well developed
scaphocerite 3.82 times as long as maxi-
mum width, with distolateral tooth acute,
overreaching distal margin of lamella; car-
pocerite reaching proximal fifth of scapho-

741

cerite; basicerite armed anterolaterally with
acute tooth.

Mandible (Fig. 2E) robust, with one-seg-
mented palp with sparse setae; molar pro-
cess obliquely truncated distally, with blunt
teeth; incisor process well developed, with
3 acute distal teeth.

Maxillula (Fig. 2F) with bilobed palp,
upper lobe tapering distally, lower lobe
slightly curved; upper lacinia slender, with
distal margin truncated, armed with 2 rows
of acute spines; lower lacinia broken, but
probably typical of Palaemonella.

Maxilla (2G) with simple palp tapering
distally; basal endite developed, deeply bi-
lobed, with numerous short setae distally;
coxal endite obsolete; scaphognathite
fringed densely with setae.

First maxilliped (Fig. 2H) with simple
seta on palp; caridean lobe well developed,
with sparse setae, exopod with well devel-
oped flagellum; basal endite with medial
margin with 3 rows of dense setae, coxal
endite with medial margin with numerous
short setae; epipod large, triangular, feebly
bilobed.

Second maxilliped (Fig. 21) with oval
epipod without podobranch. Dactylar seg-
ment laterally with 2 rows of dense setae,
distal margin truncated, with dense setae.
Propodal segment with broadly rounded
distal margin, with dense setae; dorsodistal
and ventrodistal margins of carpal segment
acute. Distal margin of ischiomeral segment
about twice as long as carpal segment. Bas-
al segment with exopodal flagellum well
developed.

Third maxilliped (Fig. 2J) with oval epi-
pod. Endopod slender, slightly overreaching
midlength of scaphocerite; ultimate seg-
ment tapering, with group of short distolat-
eral setae medially, fringed with long setae
distally; penultimate segment slightly lon-
ger than ultimate segment, lateral and me-
dial margins fringed with setae; ischio-
merus widening, with 6 distolateral spi-
nules. Exopod well developed, fringed dis-
tally with long setae. Small arthrobranch
present.

742 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

H

Fig. 2. Palaemonella hachijo, new species. Holotype male (CMNH-ZC 00017). A, right antennular peduncle,
dorsal aspect; B, same, upper flagellum; C, same, shorter ramus; D, right antenna, dorsal aspect; E, right man-
dible; F right maxillula; G, right maxilla; H, right first maxilliped; I, right second maxilliped; J, right third
maxilliped. Scales, A, D, J, 1 mm; B, 1.5 mm; C, E-I, 0.5 mm.

VOLUME 112, NUMBER 4

Table 1.—Branchial formula of Palaemonella
hachijo, new species.

Maxillipeds Pereiopods
en a FE now
Pleurobranchs —_—- — — 1 1 1 1 1
Arthrobranchs — —Y1| | — — ~ ~~ —
Podobranchs —- — — — ~— ~~ — —
Epipods 1%tskbe—— —~ — —
Exopods beet OL SS SSS

Branchial formula as in Table 1.

First pereiopod (Fig. 3A) slender, over-
reaching distal margin of scaphocerite by
proximal fourth of carpus. Chela (Fig. 3B)
0.52 times as long as CL, palm with trans-
verse rows of setae proximally; fingers ta-
pering with small hooked tip, with dense
setae distally, cutting edges entire. Carpus
0.73 times as long as CL, 1.63 times as long
as chela, slightly widening distally, distal
margin not denticulate, distoventrally with
dense setae.

Second pereiopods (Fig. 3C) long, slen-
der, symmetrical, overreaching distal mar-
gin of scaphocerite by distal fourth of meri.
Chela 1.70 times as long as CL, 1.26 times
as long as carpus, with palm subcylindrical,
compressed, entire, flexor margin expanded
proximally, slightly shorter than fingers;
movable finger (Fig. 3D) with acute hooked
tip, distal 2/3 of cutting edge entire, proxi-
mal third armed with 2 large blunt teeth;
fixed finger (Fig. 3D) with acute hooked
tip, distal 3/4 of cutting edge entire, proxi-
mal fourth armed with large blunt tooth and
quadrate conical projection, marginally ser-
rate, Opposite to the teeth on movable fin-
ger. Carpus slender, entire, 1.30 times as
long as CL, widening distally, without sub-
terminal spine. Merus slender, armed with
acute distal tooth on flexor margin, with
long setae dorsodistally. Ischium unarmed.

Third pereiopod (Fig. 3E) long, slender,
overreaching distal margin of scaphocerite
by level of midlength of carpus. Dactylus
(Fig. 3F) slightly sinuous, uniunguiculate,
midlength to distal third of external margin
armed sparsely with long setae. Propodus

743

4.75 times as long as dactylus, 1.39 times
as long as carpus, not segmented, disto-
ventral and dorsodistal angles with long se-
tae, dorsal margin armed with sparse setae.
Carpus 0.70 times as long as CL, fringed
with sparse setae dorsodistally. Merus 1.05
times as long as CL, 1.48 times as long as
carpus, dorsodistal angle fringed with 2
long setae (Fig. 3G). Ischium with sparse
setae ventrally.

Fourth thoracic sternite (Fig. 3H) armed
with long slender median spine between
coxae of first pereiopods. Fifth thoracic
sternite (Fig. 3H) armed with pair of acute
submedian spines with sinuous lateral bor-
ders proximal coxae of second pereiopods.
Eighth thoracic sternite (Fig. 3H) armed
with stout median process directed anteri-
orly.

Endopod of first pleopod (Fig. 31) ob-
long, fringed with sparse setae distally.

Endopod of second pleopod armed with
appendices interna and masculina (Fig. 3J);
appendix interna slightly overreaching tip
of appendix masculina, with cincinnuli dis-
tally; appendix masculina with numerous
long setae.

Uropod (Fig. 1E) with protopodite pos-
teroventrally acute; exopod broad, reaching
subequal to posterior telson margin, poste-
rior margin feebly rounded, lateral margin
Straight, with acute distolateral tooth, with
strong mobile spine medially; endopod
reaching subequal to distal margin of exo-
pod, slightly tapering distally.

Coloration.—Body and appendages
transparent. Posterodorsal region of cara-
pace with short, oblique rusty red line. An-
terolateral angle of carapace with oblique,
feebly undulate rusty red line. Ventral mid-
line and external margin of telson colored
with rusty red. Abdominal somites each
with transverse rusty red line along poster-
odorsal margin. Second pereiopod with red-
dish brown patches at distal margins of
merus and carpus, and junction of palm and
movable finger.

Etymology.—From locality of capture,
Hachijo-jima Island. This island is com-

744

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Rig: 3:
same, chela; C, left second pereiopod; D, same, fingers; E, left third pereiopod; EK same, dactylus; G, same,
carpomeral articulation; H, thoracic sternites, ventral aspect; I, left first pleopod; J, left second pleopod, appen-
dices interna and masculine. Scales: A, E, 1.5 mm; B, 0.5 mm; C, 3 mm; D, H, 1 mm; E G, I, J, 0.25 mm.

monly abbreviated to “‘Hachio’’. In this
case, the word hachijo is used as a noun in
apposition.

Distribution.—Known only from the
type locality.

Ecological notes.—The collection site of

Palaemonella hachijo, new species. Holotype male (CMNH-ZC 00017). A, left first pereiopod; B,

the specimen was in a small crevice in a
submarine cave at a depth of 20 m. The
crevice was naked, without any sponges or
coelenterates, therefore, P. hachijo may be
considered as a free-living species. This
point was about 15 m distance from the

VOLUME 112, NUMBER 4

cave entrance, and not influenced by inci-
dence of daylight. Several specimens of
Urocaridella sp. (Palaemonidae) and some
Cinetorhynchus concolor (Okuno) and C.
erythrostictus Okuno (Rhynchocinetidae)
were also collected.

Remarks.—All ambulatory pereiopods
but the left third pereiopod were detached
before the specimen was scrutinized. Three
unnumbered (probably right fourth and fifth
and left fourth) ambulatory pereiopods re-
mained in the bottle. These appendages
were similar in form to that of the third pe-
reiopod.

Palaemonella hachijo closely resembles
P. dolichodactylus sharing the following
characters: supraorbital spine absent; merus
of second pereiopod armed with distal tooth
on flexor margin; carpus of second pereio-
pod lacking strong submarginal tooth; and
ambulatory pereiopods long and slender.
The new species differs from P. dolicho-
dactylus in the antennal spine overreaching
the distal margin of the antennal basicerite,
posteroventral angle of the fifth abdominal
somite armed with acute tooth posteroven-
trally, and the ambulatory propodi not seg-
mented. In P. dolicodactylus, the antennal
spine falls slightly short of the level of the
distal margin of the antennal basicerite, and
the fifth abdominal somite has the pleuron
rounded posteroventrally, and the propodi
of ambulatory pereiopods are obscurely
segmented (Bruce 1991a, 1991b).

Acknowledgements

I thank Messrs. S. Kato and K. Tanaka
of Hachijo-jima Island for their help to cap-

745

turing the specimen. My thanks go to Dr.
Mary K. Wicksten of the Texas A & M
University for reading the manuscript, and
Mr. John P. Hoover of the Hawaii Medical
Library for his help in obtaining literature.
This study was supported by a Grant-in-Aid
for Encouragement of Young Scientists
(No. 10740402).

Literature Cited

Bruce, A. J. 1991a. Shallow-water palaemonid shrimps
from New Caledonia (Crustacea: Decapoda).
Pp. 221—279 in B. Richer de Forges, ed., Le
Benthos des Fonds Meubles des Lagons de
Nouvelle-Calédonie, 1. Etudes et Théses, Paris,
ORSTOM, 311 pp.

. 1991b. Crustacea Decapoda: Further deep-sea

palaemonid shrimps from New Caledonian wa-

ters. Pp. 299-411 in A. Crosnier, ed., Résultats
des Campagnes MUSORSTOM, 9.—Mémoires

Muséum National d’ Histoire Naturelle, (A) 152,

520 pp.

. 1994. A synopsis of the Indo-Pacific genera
of the Pontoniinae (Crustacea: Decapoda: Pa-
laemonidae). Koeltz Scientific Books, Konig-
stein, Germany, 172 pp.

Chace, E A. Jr, & A. J. Bruce. 1993. The caridean
shrimps (Crustacea: Decapoda) of The Alba-
tross Philippine Expedition 1907-1910 Part 6:
Superfamily Palaemonoidea.—Smithsonian
Contributions to Zoology 543:i—tiv+ 1-152.

Dana, J. D. 1852. Conspectus Crustaceorum quae in
Orbis Terrarum circumnavigatione, Carolo
Wilkes e Classe Republicae Foederatae Duce,
lexit et descripsit.—Proceedings of the Acade-
my of Natural Sciences of Philadelphia 1852:
10-28.

Holthuis, L. B. 1993. The recent genera of the caridean
and stenopodidean shrimps (Crustacea, Deca-
poda): with an appendix of the order Amphion-
idacea. National Natuurhistorisch Museum, Lei-
den, The Netherlands. 328 pp.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(4):746—753. 1999.

A new species of Macrobrachium
(Crustacea: Decapoda: Palaemonidae), with
abbreviated development, from Veracruz, Mexico

José Luis Villalobos and Fernando Alvarez

Coleccion Nacional de Crustaceos, Instituto de Biologia,
Universidad Nacional Autonoma de México, Apartado Postal 70-153,
México 04510 D.E, México

Abstract.—Macrobrachium tuxtlaense, a new epigean species of freshwater
prawn with abbreviated development from Los Tuxtlas region in the state of
Veracruz, Mexico, is described. The new species is most similar to the cave-
dwelling M. acherontium Holthuis and M. villalobosi Hobbs, the two previ-
ously known species from Mexico with abbreviated development.

Although abbreviated development is a
widespread characteristic among species of
Palaemoninae (Pereira & Garcia 1995),
which comprises mostly freshwater and es-
tuarine shrimps, only two cave-dwelling
species of Macrobrachium Bate, 1868 from
North America (Mexico and the United
States) present this kind of development: M.
villalobosi Hobbs, 1973 and M. acheron-
tium Holthuis, 1977. Until now, within the
Palaemoninae from North America, the
only epigean species with abbreviated de-
velopment belonged to Palaemonetes, Hell-
er, 1869; and Cryphiops (Bithynops) Villa-
lobos et al., 1989. In this study we describe
a new epigean species of Macrobrachium
from Los Tuxtlas region, Veracruz, in
which females carry few (10—20), large
eggs (mean size 2.8 mm by 2.5 mm), and
an abbreviated development is present as
evidenced by the morphology of the first
larval stage (Fig. 1). The presence in the
first larval stage of all five pairs of pereio-
pods, well developed pleopods, and a fan
shaped telson, indicate a partially abbrevi-
ated development with two larval stages ac-
cording to Jalihal et al. (1993).

All specimens used are deposited in the
Coleccion Nacional de Crustaceos, Instituto
de Biologia, Universidad Nacional Auton-

oma de México (CNCR). Abbreviations
used are: cl, carapace length; tl, total length.

Macrobrachium tuxtlaense, new species
Figs. 1-3

Material examined.—Holotype: 1 ¢ cl
10.4 mm, tl 40.5 mm, small stream 1 km S
of Coyame (18°26'N, 95°7'W), tributary of
lake Catemaco, Municipio de Catemaco,
Veracruz, México, Aug 1994, colls. E Al-
varez, M. E. Camacho, J. L. Villalobos,
CNCR 13174. Allotype: 1 ¢ cl 8.6 mm, tl
34.3 mm, same locality, date, and collectors
as holotype, CNCR 13174. Paratypes: 6 3
cl 7.7—10.0 mm, tl 31.5—37.6 mm; 3 @ cl
8.5—9.7 mm, tl 33.4—35.3 mm; same local-
ity, date, and collectors as holotype, CNCR
13175. Non-paratypes: 8 ¢ cl 6.4—10.8
mm, tl 26:8—42.2 mm; 12 ¢ chi6e4=ies
mm, tl 25.5—40.8 mm, stream 1 km S of
Las Margaritas, tributary of lake Catemaco,
Municipio de Catemaco, Veracruz, México,
Mar 1997, colls. J. C. Molinero, R. Robles,
CNCR 17428.

Description.—Small sized shrimp, max-
imum known size 42.2 mm tl (Fig. 2a).
Rostrum reaching beyond distal margin of
third segment of antennular peduncle, me-
dian rib present along inferior half. Dorsal
margin arched, distal third elevated, proxi-

VOLUME 112, NUMBER 4 747

Fig. 1. Macrobrachium tuxtlaense new species, first larval stage CNCR 13174b: a, lateral view; b, telson;
c, first pereiopod; d, second pereiopod; e, third pereiopod; f, fourth pereiopod; g, fifth pereiopod. Scale bars
represent 1 mm.

748

mal *% oriented downwards; with 9 to 11
short, acute teeth, pointing forward, with
setae in between; first 2 teeth located pos-
terior to orbit. Distal *% of ventral margin
arched, with 2 teeth, distal one under sec-
ond and third teeth of dorsal margin and
proximal one under third and fourth teeth
of dorsal margin.

Carapace smooth, with antennal and he-
patic spines. Antennal spine slightly below
inferior orbital angle, submarginal; hepatic
spine slightly behind and under antennal
spine. Inferior orbital angle rounded. He-
patic groove deep, anterior half oriented to-
wards antennal spine.

Abdomen smooth, inferior margin of
pleurae of first 3 somites rounded. Postero-
lateral angle of fourth and fifth somites sub-
acute. Sixth somite 1.7 times as long as
fifth; posterior margin with rounded notch
flanked by acute angles, and adjacent to ar-
ticulation of external ramus of uropods.

Telson (Fig. 2e) reaching distal fourth of
internal ramus of uropods, narrowing dis-
tally; armed with 2 pairs of small, acute
dorsal spines on posterior half, first pair at
¥% the length of telson, second pair at %4 the
length of telson. Posterior margin subacute,
bearing 10—12 thick, plumose setae; disto-
lateral angles armed with 2 articulated
spines, internal pair longest reaching be-
yond tip of telson, external pair % length of
internal pair.

Eyes pigmented, cornea normally devel-
oped, globose, wider than ocular peduncle.

Antennules (Fig. 2b) with slender, acute
stylocerite, tip reaching beyond half of first
peduncular segment. First peduncular seg-
ment depressed, concave dorsally to fit eye;
lateral margin ending in slender, acute
spine; second and third segments subcy-
lindrical.

Antennae with basicerite armed on ante-
rior margin with acute, slender spine. Sca-
phocerite (Fig. 2c) tapering anteriorly, 2.8
times as long as wide; lateral margin ending
distally in short spine, not overreaching dis-
tal margin.

Mandibles (Fig. 3a) with 3-segmented

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

palp bearing long setae on tip and articu-
lations; incisor process with 3 conical teeth,
molar process with 2 wide, rounded teeth
on mesial border.

Maxillules (Fig. 3b) with bilobed palp;
distal lobe slender with long, single, sub-
apical seta on anterior margin; proximal
lobe rounded, with 1—3 apical setae. Mesial
margin of anterior lacinia armed with spi-
niform teeth, anterior and posterior margins
bearing thin setae; posterior lacinia curved
with abundant thin setae on distal half.

Maxillae (Fig. 3c) with scaphognathite
bordered with plumose setae, anterior lobe
narrower and longer than posterior; palp de-
void of setae, thick proximally, tapering
abruptly on distal half, ending in sharp tip;
endite bilobed, divided by deep incision
along % of its length, both lobes slender and
with terminal tuft of setae.

First maxilliped (Fig. 3d) with 2 endites
bearing marginal and submarginal setae,
anterior endite twice as long as posterior
endite. Exopod slender, 4 times as long as
palp, distal third bearing long setae; cari-
dean lobe well formed, fused to base of ex-
opod, bordered with long, plumose setae;
palp strong basally, tapering distally; epi-
podite subtriangular, distal tip rounded.

Second maxilliped (Fig. 3e) subpediform
with well developed podobranch; endopo-
dite 4-segmented, distal 2 segments orient-
ed mesially, gnathal border armed with se-
tae and spines; exopodite slender, approxi-
mately twice as long as endopodite, tip
bearing long, plumose setae.

Third maxilliped (Fig. 3f) pediform,
slender, reaching distal margin of second
segment of antennular peduncle; with well
developed arthrobranch; coxa with round-
ed, chitinized plate; basis short, bearing ar-
ticulations with endopodite and exopodite
at same level. Endopodite 3-segmented,
bearing tufts of setae, becoming denser dis-
tally; first segment slender, 1.15 times as
long as second segment; third segment
shorter than second, ending in rounded tip.
Exopodite slender, longer than first segment

VOLUME 112, NUMBER 4 749

Mj

Me

Fig. 2. Macrobrachium tuxtlaense new species, male holotype CNCR 13174: a, lateral view; b, distal portion
of antennular peduncle; c, distal portion of antennal peduncle; d, appendix masculina on second pleopod; e,
telson and uropods, dorsal view. Scales equal 5 mm (a), | mm (b—d), and 2 mm (e).

“Ye

Udy
Vb

| gir eae

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Macrobrachium tuxtlaense new species, male holotype CNCR 13174: a, right mandible; b, right

maxillule; c, right maxilla; d, first maxilliped; e, second maxilliped; f, third maxilliped. Scale bars represent 1

mm.

of endopodite, distal third bearing long se-
tae.

First pair of chelipeds slender, shorter
than second, distal margin of carpus reach-
ing beyond scaphocerite. Ischium short,
0.65 times as long as merus, 0.57 as long
as carpus, subequal in length to chela; ven-

tral margin with tufts of setae on proximal
half. Merus shorter than carpus, 1.4 times
length of chela, dorsal and ventral margins
parallel. Carpus smooth, slender, widening
distally, 1.54 times length of chela. Chela
wider than distal portion of carpus; palm
slightly longer than fingers, with line of se-

VOLUME 112, NUMBER 4

tae on proximoventral margin; fingers
straight, slightly gaping, with tufts of setae
on distal half, tips subacute and corneous,
bearing short setae; cutting edge hard, chi-
tinized, devoid of teeth.

Second pair of chelipeds subequal, long,
covered with granules and small spines;
proximal third of carpus reaching beyond
scaphocerite. Ischium moderately com-
pressed, dorsal and ventral margins parallel,
of same length as merus and carpus, 0.53
times length of chela. Merus cylindrical, 5
times as long as wide, of same length as
carpus, half as long as chela. Carpus be-
coming slightly wider distally, 3.7 times as
long as distal width, 0.52 times length of
chela. Chela moderately compressed, dorsal
and ventral margins parallel, 6 times as
long as high; palm as long as ischium, mer-
us, and carpus, 0.52 times the length of che-
la, 1.18 times length of movable finger. Fin-
gers slender, covered with scattered setae,
tips acute and corneous; cutting margin
with conical teeth on proximal half, becom-
ing continuous smooth ridge distally; dactyl
with 4 teeth, fixed finger with 5 teeth, distal
2 teeth the largest.

Third to fifth pair of pereiopods slender,
increasing in length posteriorly; with scat-
tered short setae, 2 rows of spinules along
posterior margin of propodus, tuft of setae
on propodus-dactyl articulation, and row of
setae along superior border of dactyl; dactyl
the shortest segment, ending in sharp tip.

Third pair of pereiopods with distal third
of propodus reaching beyond scaphocerite;
ischium half length of merus and propodus,
subequal in length to carpus, 1.5 times
length of dactyl; merus 1.9 times length of
carpus, 3.2 times length of dactyl, of same
length as propodus; carpus 0.52 times
length of propodus, 1.6 times length of dac-
tyl; propodus 3.2 times length of dactyl.

Fourth pair of pereiopods with distal
fourth of propodus reaching beyond sca-
phocerite; ischium half length of merus and
propodus, of same length as carpus, 1.6
times length of dactyl; merus 1.63 times
length of carpus, 3.4 times length of dactyl,

731

of same length as propodus; carpus 0.6
times length of propodus, twice as long as
dactyl; propodus 3.4 times length of dactyl.

Fifth pair of pereiopods with distal mar-
gin of propodus reaching beyond scapho-
cerite; ischium 0.52 times length of merus,
0.75 times length of carpus, 0.41 times
length of propodus, 1.57 times length of
dactyl; merus 1.47 times length of carpus,
3.1 times length of dactyl, 0.8 times length
of propodus; carpus half as long as propo-
dus, twice as long as dactyl; propodus 4
times as long as dactyl.

Pleopods with both rami well developed,
except for first pair in both sexes; first pair
with reduced endopod, lacking appendix
interna. Second pleopod (Fig. 2d) of male
with slender appendix masculina, reaching
distal third of endopod, mesial border bear-
ing 2 rows of 9 acute setae reaching apex;
appendix interna moderately robust, reach-
ing proximal third of appendix masculina.

Uropods (Fig. 2e) with protopodite with
external lobe ending in sharp tip. Endopod
shorter than exopod, with setae along pos-
terior margin and distal half of lateral mar-
gins, posterior margin rounded. Exopod
with bare, straight, lateral margin; ending in
fixed spine, flanked internally by long,
sharp, movable spine; diaeresis incomplete,
marked by weak furrow extending through
more than half the surface of ramus; inter-
nal and posterior margins with setae.

Remarks.—Macrobrachium tuxtlaense is
probably more closely related to the cave-
dwelling M. acherontium and M. villalo-
bosi, all of which exhibit abbreviated de-
velopment and are similar morphologically.
These three species can be distinguished by
the shape of rostrum, length of telson, ex-
tent of development of the eyes, and the
different proportions of the segments of the
first and second pair of pereiopods (Table
L):

Similarly to other species of Macro-
brachium from the Americas with abbrevi-
ated development (e.g., M. aracamuni Rod-
riguez, 1982; M. cortezi Rodriguez, 1982;
M. reyesi Pereira, 1986; M. pectinatum Pe-

752

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Comparison of selected characteristics of Macrobrachium tuxtlaense, new species, from Los Tuxtlas
region, Veracruz, and M. acherontium and M. villalobosi.

M. tuxtlaense

M. acherontium

M. villalobosi

Rostrum

Ist pair of pe-
reiopods

2nd pair of pe-
reiopods

9-11 dorsal teeth/2 ventral
teeth, reaching distal
margin of scaphocerite

1.2 times length of 6th ab-
dominal somite
Normally developed

Carpus 1.54 times length of
chela

Covered with granules and
small spines; ischium as
long as merus and car-
pus, 0.53 length of chela

8—11 dorsal teeth/3 ventral
teeth, reaching distal
margin of scaphocerite

1.2 times length of 6th ab-
dominal somite

Reduced, globose cornea,
narrower than peduncle

Carpus less than twice
length of chela

Smooth with scattered se-
tae; ischium 0.93 as long
as merus, 0.75 as long as
carpus, and 0.63 as long
as chela

9-11 dorsal teeth/2 ventral
teeth, reaching beyond
distal margin of scapho-
cerite

1.4 times length of 6th ab-
dominal somite

Cornea absent

Carpus twice length of che-
la

Smooth with scattered se-
tae; ischium 0.76 as long
as merus, 0.5 as long as
carpus and chela

reira, 1986; M. atabapense Pereira, 1986;
M. dierythrum Pereira, 1986; M. rodriguezi
Pereira, 1986; M. pumilum Pereira, 1986),
M. tuxtlaense is a small-sized species with
a maximum total length of 42.2 mm. In M.
tuxtlaense, the egg size (2.8 X 2.5 mm) is
the second largest among all Macrobrach-
ium species from the Americas, the largest
eggs (3.66 X 2.44 mm) being those of M.
ferreirai Kensley & Walker, 1982 (see Ma-
galhaes & Walker 1988, Pereira & Garcia
1995).

The evolution of abbreviated develop-
ment in freshwater palaemonid shrimps has
been linked to the invasion of nutrient-poor
waters where planktotrophic larvae would
die in the absence of phyto and zooplankton
upon which to feed (Magalhaes & Walker
1988). The distribution pattern of species
with abbreviated development coincides
with the presence of oligotrophic waters
such as small mountain streams or bodies
of water inside caves. The three Mexican
species of Macrobrachium with abbreviated
development fit well this pattern, occurring
in small streams where species of Macro-
brachium with planktotrophic larvae do not
occur.

Macrobrachium tuxtlaense possibly can
be considered an endangered species as it

posseses very limited dispersal capabilities
and occurs only in two small tributaries of
Lake Catemaco, an area that increasingly is
being devoted to cattle ranching.

Acknowledgments

We thank Rolando Mendoza for prepar-
ing the drawings, and Rafael Robles and

Juan Carlos Molinero for their help in the
field.

Literature Cited

Bate, C. S. 1868. On a new genus, with four new spe-
cies, of freshwater prawns.—Proceedings of the
Zoological Society of London 1868:363—368.

Heller, C. 1869. Zur Naheren Kenntniss Der in Den
Sussen Gewassern de Sudlichen Europa Vor-
kommenden Meerescrustaceen.—Zeitschrift
fuer Wissenschaftliche Zoologie 19:156—162.

Hobbs, H. H., Jr. 1973. Two new troglobitic shrimps
(Decapoda: Alpheidae and Palaemonidae) from
Oaxaca, Mexico.—Association for Mexican
Cave Studies, Bulletin 5:73-—80.

Holthuis, L. B. 1977. Cave shrimps (Crustacea: Dec-
apoda: Natantia) from Mexico.—Accademia
Nazionale dei Lincei 171:135—142.

Jalihal, D. R., K. N. Sankolli, & S. Shenoy. 1993. Evo-
lution of larval development patterns and the
process of freshwaterization in the prawn genus
Macrobrachium Bate, 1868 (Decapoda, Palae-
monidae).—Crustaceana 65:365-—376.

Kensley, B., & I. Walker. 1982. Palaemonid shrimps

VOLUME 112, NUMBER 4

from the Amazon basin, Brazil (Crustacea: Dec-
apoda: Natantia).—Smithsonian Contributions
to Zoology 362:1—28.

Magalhaes, C., & I. Walker. 1988. Larval development
and ecological distribution of central Amazo-
nian palaemonid shrimps (Decapoda, Cari-
dea).—Crustaceana 55:279-—292.

Pereira, G. A. 1986. Freshwater shrimps from Vene-
zuela I: seven new species of Palaemoninae
(Crustacea: Decapoda: Palaemonidae).—Pro-
ceedings of the Biological Society of Washing-
ton 99:198—213.

, & J. V. Garcia. 1995. Larval development of

Macrobrachium reyesi Pereira (Decapoda: Pa-

fe

laemonidae), with a discussion on the origin of
abbreviated development in Palaemonids.—
Journal of Crustacean Biology 15:117—133.

Rodriguez, G. 1982. Fresh-water shrimps (Crustacea,
Decapoda, Natantia) of the Orinoco basin and
the Venezuelan Guayana.—Journal of Crusta-
cean Biology 2:378-391.

Villalobos, J. L., J. C. Nates, & A. Canti. 1989. Re-
vision de los géneros Cryphiops Dana, 1852 y
Bithynops Holthuis, 1973 de la familia Palae-
monidae (Crustacea: Decapoda), y descripcidn
de una especie nueva para el estado de Chiapas,
México.—Anales del Instituto de Biologia,
Universidad Nacional Aut6énoma de México,
Serie Zoologia 60(2):159—184.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(4):754-—758. 1999.

Description of a new spider crab, Maja gracilipes, from the South
China Sea, with notes on the taxonomic validity of M. brevispinosis
Dai, 1981 (Crustacea: Decapoda: Brachyura: Majidae)

H.-L. Chen and Peter K. L. Ng

(H-LC) Institute of Oceanology, Chinese Academy of Sciences, 7 Nan-Hai Road,
Qingdao 266071, People’s Republic of China; (PKLN) Department of Biological Sciences,
National University of Singapore, Kent Ridge, Singapore 119260, Republic of Singapore

Abstract.—A new species of majid crab, Maja gracilipes, is described from
the South China Sea. It is similar to M. confragosa Griffin & Tranter, 1986,
described from the Kei Islands, Indonesia, but differs in proportions of the
ambulatory legs, structure of the intercalated spine, third maxilliped, male ab-
domen, and male first pleopod. Maja brevispinosis Dai, 1981, is synonymised

with M. compressipes (Miers, 1879).

Seventeen species of spider crabs of the
genus Maja Lamarck, 1801, are known, of
which 14 occur in the Indo-West Pacific
(Griffin & Tranter 1986, Dai 1981). Dai et
al. (1986) and Dai & Yang (1991:150) re-
corded five species from China, M. japon-
ica Rathbun, 1932, M. spinigera de Haan,
1839, M. gibba Alcock, 1895, M. sakaii
Takeda & Miyake, 1969, and M. brevispi-
nosis Dai, 1981. A sixth species also known
from China and previously placed in the ge-
nus Leptomithrax, L. compressipes Miers,
1879, originally described from China, was
transferred to Maja by Griffin & Tranter
(1986).

A new species of Maja is here described
from the South China Sea. The species is
similar to M. confragosa Griffin & Tranter,
1986, but can easily be distinguished by the
structure of the intercalated spine (between
orbital eave and postorbital spine), third
maxilliped, ambulatory legs, male abdomen
and male first pleopod. The identity of M.
brevispinosis Dai, 1981, described from
southern China, is re-appraised, and the
species is here synonymised with M. com-
pressipes (Miers, 1879).

The following abbreviations are used:
male first pleopod (G1), male second ple-

opod (G2). The terminology used follows
Griffin & Tranter (1986). The leg length is
the combined maximum lengths of merus,
carpus, propodus and dactylus (measured in
a straight line point to point). The type
specimen is in the Institute of Oceanology,
Chinese Academy of Sciences (OCAS) in
Qingdao, China.

Maja gracilipes, new species
Figs. 1, 2

Material examined.—Holotype, male,
carapace width 43.5 mm, carapace length
45.4 mm, rostral length 14.4 mm, IOCAS
K33B-34, South China Sea, station 6080,
180 m, on gravel, 21 Apr 1959.

Description of male holotype.—Carapace
longitudinally subovate, relatively broad,
width subequal to postrostral length; dorsal
surface densely covered with numerous
granules of various sizes and strength; mid-
line weakly but distinctly elevated, with 6
low spines anteriorly, 2—3 pairs of low,
rounded submedian granules posteriorly.
Rostrum with 2 long, slender spines, ca. 0.3
times postrostral length; spines gently curv-
ing outwards; outer margin with 1—2 small,
submedian rounded granules; cleft between
base of rostral spines deep.

VOLUME 112, NUMBER 4

Orbital eave strongly expanded, anteri-
orly rounded, posteriorly developed into
long, sharp antorbital spine; intercalated
spine subtruncate with proximal part much
narrower than median part, much shorter
than antorbital spine; preorbital spine ab-
sent. Eyestalks relatively narrow; cornea
large, mostly ventral, ca. half length of en-
tire eye.

Basal antennal segment longitudinally
subrectangular; outer lateral spine large, di-
rected obliquely outwards, anterior margin
with 3-4 small granules; distal median
spine short; anterolateral margin granular
but without distinct spine or larger granule;
median surface with 2-3 large rounded
granules. Suborbital lobe separated from
basal antennal segment by distinct fissure.
Pterygostomial region relatively smooth;
margin with 5 large rounded granules. Ep-
istome smooth, unarmed.

Postorbital tooth reaching slightly but
distinctly further anteriorly than antorbital
spine, basally broad, partially excavated,
narrow and acutely triangular distally, tooth
directed anteriorly and obliquely outwards;
basal part with several small granules. He-
patic spine much shorter, ca. 0.3 times
length of postorbital tooth; base and area
posterior to it covered with numerous large
rounded granules.

Branchial region with 4 low but distinct
spines (or large sharp granules), 3 clearly
marginal and 1 submarginal in position;
dorsal surface with 1 large, rounded granule
which is smaller than marginal ones, adja-
cent to median cardiac spine. Mesogastric
region with 4 low, large, rounded median
granules, not spiniform; urogastric region
with 1 low, large median granule. Cardiac
region with 1 median and 1 anterior low,
large, rounded median granules; posterior
surface with 2 sharper granules. Anterior
surface of intestinal region with 2 large sub-
median granules; posterior surface with 2
small sharp spines.

Merus of third maxilliped with postero-
median margin raised, not distinctly gran-
ulated, anteroexternal angle rounded, su-

TAS

bauriculiform; inner posterolateral angle tri-
angular, strongly produced. Ischium with
broad, shallow oblique median depression;
margins of depression with low rounded
granules. Exopod relatively broad, medially
with low longitudinal ridge, not distinctly
granular.

Chelipeds elongate, subequal, ca. sube-
qual postrostral carapace length. Chelae not
swollen, surfaces smooth; fingers 0.7—0.8
times length of palm; cutting edges un-
armed, not pigmented. Carpus relatively
short, dorsal surface with distinct sulcus;
dorsal margin with very low crest, more
prominent on proximal part, crest with 2
low sharp teeth. Merus with one very low
submedian granule on dorsal margin, oth-
erwise smooth. Basis-ischium smooth, un-
armed.

Surfaces of ambulatory legs smooth, first
leg ca. subequal in length to postrostral car-
apace length; second to fourth legs 1.6, 1.5—
1.6, 1.4 and 1.1—1.2 times postrostral cara-
pace length respectively. Merus of first leg
slender, ca. 7.5 times longer than high; dor-
sal margin with low, rounded distal tooth,
otherwise unarmed, inner surfaces with
scattered long stiff setae. Carpus on all legs
with median sulcus, inner surface (espe-
cially of first leg) with numerous long, stiff
setae; propodus laterally flattened, slender.
Second to fourth legs with merus ca. 7.5,
7.0, 5.8 times longer than high respectively;
dorsal margin of merus smooth, without
distinct distal tooth; inner surfaces of merus
and carpus with scattered long, stiff setae.
Dactylus of all legs gently curved, un-
armed, ventral margin with numerous short,
stiff setae.

Thoracic sternites 1 and 2 fused, demar-
cated by granular ridge; sternites 2-4 com-
pletely fused, without trace of sutures, sur-
face smooth, lateral margins cristate; sur-
faces of sternites 5—8 covered with numer-
ous small granules. Telson semicircular;
segments 4—6 longitudinally rectangular,
lateral margins of each segment gently con-
cave; segment 3 trapezoidal, anterolateral
margin gently concave, posterolateral mar-

756 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Maja gracilipes, new species. Holotype male, carapace width 43.5 mm, IOCAS K33B-34. a, left
frontal part of carapace (dorsal view); b, left frontal part of carapace (ventral view); c, right third maxilliped; d,
left first ambulatory leg; e, left second ambulatory leg; f, right third ambulatory leg; g, right fourth ambulatory
leg.

VOLUME 112, NUMBER 4

gin rounded; surfaces with scattered gran-
ules.

G1 relatively long; basal part dilated;
median part almost straight from ventral
view but gently curving when viewed in
situ; distal part with one perpendicular
tooth-like projection, dorsal margin with
numerous stiff setae. Male second pleopod
very short.

Etymology.—The name is derived from
the Latin gracilis (for slender) and pes (for
leg), alluding to the relatively long ambu-
latory legs of the species. Used as a noun
in apposition.

qv

med

ae
=

rad

\
)
<<
le e

Fig. 2. Maja gracilipes, new species. Holotype male, carapace width 43.5 mm, IOCAS K33B-34. a, b, left
Gl; c, d, distal part of G1; e, left G2; f, abdomen (schematic outline only).

Remarks.—Maja gracilipes, new species,
is closest to M. confragosa, especially with
regards to the general carapace morpholo-
gy, presence of a large spine on the lateral
margin of the basal antennal segment, and
general structure of the G1 (Griffin & Tran-
ter 1996:210). Maja gracilipes can easily be
separated from M. confragosa by the pres-
ence of a subtruncate intercalated tooth
with the median part broader than the prox-
imal part (Fig. la) (against acutely trian-
gular with the median part distinctly nar-
rower than the proximal part); anteroexter-
nal angle of the third maxilliped ischium is

758

angular (Fig. 1c) (against rounded); propor-
tionately longer ambulatory meri and pro-
podi (Figs. 1d—g); relatively broader male
abdomen, notably segment 3 (Fig. 2f); and
the distal part of the G1 (beyond the lateral
subdistal projection) is proportionately lon-
ger (Figs. 2a—d) (cf. Griffin & Tranter 1986:
214, 215, Figs. 72a—c, 73a, pl. 16).

Maja confragosa was originally de-
scribed by Griffin & Tranter (1986) from
only one male (carapace length 39.0 mm)
from the Kei Islands in Indonesia, and con-
sidering its slender chelipeds, appears to be
a subadult specimen. Its Gl, however, is
fully developed. The male specimen of M.
gracilipes is somewhat larger (carapace
length 45.4 mm) and the chelipeds are al-
ready relatively enlarged. The differences
observed between the male specimens, es-
pecially in the proportions of the legs, can-
not be accounted for by size-associated var-
jation.

On another matter, M. brevispinosis Dai,
1981, bears a striking resemblance to M.
compressipes (Miers, 1879) in its carapace
features, short ambulatory legs and posses-
sion of an ambulatory carpus which is very
broad and triangular in shape. Interestingly,
M. compressipes is known only from the
holotype female from Canton (= Guang-
dong), China, while M. brevispinosis was
described from two males and a female
from the same area. From the descriptions
and figures of the holotype female of M.
compressipes (cf. Griffin & Tranter 1986:
211, pl. 16) and types of M. brevispinosis
(see Dai 1981:37, figs. 1:6—-10; Dai & Yang
1991:151, pl. 18(2), figs. 77(1—4)), we can
find no reason to separate the two species.
As such, Maja brevispinosis Dai, 1981, is
here synonymised with M. compressipes
(Miers, 1879).

Acknowledgements

The senior author’s work in Singapore
was partially supported by a travel grant
from the Raffles Museum, while the junior

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

author was supported by a research grant
from the National University of Singapore.
This is contribution number 3598 from the
IOCAS.

Literature Cited

Alcock, A. 1895. Materials for a carcinological fauna
of India, 1. The Brachyura Oxyrhyncha.—Jour-
nal of the Asiatic Society of Bengal 64:157—
291, pls. 3-5.

Dai, A.-Y. 1981. Two new species of Chinese Majidae
(Crustacea: Brachyura).—Acta Zootaxonomica
Sinica 6(1):36—38, pl. 1.

, & S. Yang. 1991. Crabs of the China Seas.

China Ocean Press (Beijing). 682 pp, figs. 1—

295, pls. 1-74.

: , Y. Song, & G. Chen, 1986. Crabs of
the China Seas. China Ocean Press, Beijing.
17+642 pp., pls. 1-74. [In Chinese]

Griffin, D. J. G., & H. A. Tranter. 1986. The Decapoda
Brachyura of the Siboga Expedition. Part VIII.
Majidae.—Siboga-Expeditie 39C4:1—335, 22
pls.

Haan, W. De. 1833—1849. Crustacea. Jn: P. E Siebold,
Fauna Japonica sive Descriptio Animalium,
quae in Itinere per Japoniam, Jussu et Auspiciis
Superiorum, Qui Summum in India Batava Im-
perium Tenent, Suscepto, Annis 1823-1830
Collegit, Notis, Observationibus et Adumbra-
tionibus Illustravit, (Crustacea): i-xvii + 1-xxx1
+ i-ix-xvi + 1-243, pls. A-J, L—Q, 1-55, circ.
tabl. 2. Lugduni-Batavorum.

Lamarck, J. B. P. A. 1801. Syst€me des animaux sans
vertébres, ou tableau général des classes, des
ordres et des genres de ces animaux’ présentant
leurs caractéres essentiels et leur distribution,
d’apres la considération de leurs rapports natu-
rels et de leur organisation, et suivant
l’arrangement établi dans les galeries du Mu-
séum d’Hist. Naturelle, parmi leurs dépouilles
conservées; précédé du discours d’ ouverture du
cours de zoologie, donné dans le Muséum na-
tional d’Histoire naturelle l’an 8 de la Répub-
lique. Paris, viiit 432 pp.

Miers, E. J. 1879. On a collection of Crustacea made
by Capt. H. C. St. John, R. N. in the Korean
and Japanese seas, 1. Podophthalmia.—Pro-
ceedings of the Zoological Society of London
1879:18-61, pls. 1-3.

Rathbun, M. J. 1932. Preliminary descriptions of new
species of Japanese crabs.—Proceedings of the
Biological Society of Washington 45:29-38.

Takeda, M., & S. Miyake. 1969. Crabs from the East
China Sea. 3. Brachygnatha Oxyrhyncha.—
Journal of the Faculty of Agriculture, Kyushu
University 15:469—522.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(4):759-767. 1999.

On the identities of two Pacific species of deep-water porter crabs,
Hypsophrys longirostris Chen, 1986, and Homologenus donghaiensis
Chen, 1986 (Crustacea: Decapoda: Brachyura: Homolidae)

Peter K. L. Ng and H.-L. Chen

(PKLN) Department of Biological Sciences, National University of Singapore, Kent Ridge,
Singapore 119260, Republic of Singapore; (H-LC) Institute of Oceanology,
Chinese Academy of Sciences, 7 Nan-Hai Road,

Qingdao 266071, People’s Republic of China

Abstract.—The identities of two species of poorly known deep-water hom-
olid crabs, Hypsophrys longirostris Chen (presently in the genus Lamoha) and
Homologenus donghaiensis Chen from the East and South China Seas are
clarified. The publication validating these taxa is not well circulated, and both
names have been missed by most carcinologists. Hypsophrys futuna Guinot &
Richer de Forges, described from the Wallis and Futuna Islands in the central
Pacific, is shown to be a junior synonym of Lamoha longirostris.

Chen (1986) published a short abstract in
Chinese on the deep-water homolids from
the East China Sea in the proceedings of a
meeting of the Chinese Crustacean Society.
The text of this short abstract listed five
species: Homola orientalis Henderson,
1888, Paromola macrochira Sakai, 1961,
Paromolopsis boasi Wood-Mason & A\l-
cock, 1891, and two new species, Hypso-
phrys longirostris and Homologenus don-
ghaiensis. The two new species were treat-
ed separately, with their key characters
highlighted, albeit very briefly by Chen
(1986). Although no specimens were men-
tioned or sizes indicated in Chen (1986),
her two names are nevertheless valid under
the International Code of Zoological No-
menclature (1985). Chen had treated the ab-
stract only as a provisional document in the
proceedings of a local meeting and did not
regard it as a scientific publication or the
names as having any validity. She had orig-
inally intended to formally describe both
species in a full paper later on, but for var-
ious reasons, this was never done. The pro-
ceedings of the meeting was not easily
available outside China and as such, has
been missed by most carcinologists.

Since Chen’s (1986) publication, many
species of homolids have been described
from the Pacific in the important revision
by Guinot & Richer de Forges (1995). Un-
fortunately, Guinot & Richer de Forges
(1995) were not aware of Chen’s (1986) ab-
stract. In any case, full descriptions and fig-
ures of the two species in question were not
provided by Chen (1986), and it would not
have been possible to ascertain the actual
identities of Hypsophrys longirostris and
Homologenus donghaiensis just from her
brief text.

The present paper clarifies the outstand-
ing taxonomic problems with Chen’s (1986)
two taxa, Hypsophrys longirostris and
Homologenus donghaiensis. Both species
are redescribed based on Chen’s (1986)
original material, figures provided and
types designated. Measurements indicated
are of the carapace length (tip of rostrum of
posterior carapace margin) and width (max-
imum, between tips of lateral-most spines)
respectively. The terminology used follows
Guinot & Richer de Forges (1995). The ab-
breviations Pl—5 refer to the pereiopods
(P1, cheliped; P2—5, first to fourth ambu-
latory legs respectively). Specimens exam-

760

ined are deposited in the Institute of Ocean-
ology, Chinese Academy of Sciences,
Qingdao (IOCAS); Taiwan Museum, Taipei
(TMCD); Muséum national d’ Histoire na-
turelle, Paris (MNHN); and Zoological Ref-
erence Collection of the Raffles Museum,
National University of Singapore (ZRC).

Systematic Account

Family Homolidae
Genus Lamoha Ng, 1988a

Type species.—Hypsophrys superciliosa
Wood-Mason, in Wood-Mason & Alcock,
1891.

Remarks.—The better known generic
name Hypsophrys Wood-Mason & Alcock,
1891, was found by Ng (1998a) to be pre-
occupied by Hypsophrys Agassiz, 1859, a
genus of American freshwater fish, and
Wood-Mason & Alcock’s name was re-
placed by Lamoha Ng, 1998a. Nine species
of Lamoha are now recognized from the
Indo-Pacific and Atlantic (Guinot & Richer
de Forges 1995; Ng 1998a, 1998b).

Lamoha longirostris (Chen, 1986)
Figsa ly 2

Hypsophrus longirostris Chen, 1986:227
(misspelling of Hypsophrys).

Hypsophrys futuna Guinot & Richer de
Forges, 1995:456, figs. 611, 66a, g.

Material examined.—Lectotype male
(22.7 by 18.2 mm) COCAS KY8B-73),
East China Sea, 28°45’N, 127°30’E, 900 m,
on soft mud, coll. 3 Jan 1981.

Paralectotypes.—1 male (23.5 by 19.0
mm) (ZRC 1999.007), same data as lecto-
type. 1 male (21.2 by 18.0 mm), 1 oviger-
ous female (22.0 by 18.5 mm) (IOCAS
K69B-30), South China Sea, 19°00'N,
113°30’E, 1100 m, on soft mud, 13 Jul
15

Others.—2 males (18.3 by 14.8 mm,
14.6 by 11.7 mm), 1 female (23.7 by 17.6
mm), paratypes of Hypsophrys futuna
Guinot & Richer de Forges, 1995: MU-
SORSTOM 7, station CP 623, Wallis and

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Futuna Islands, 12°5’S, 178°11.5'W, 1300
m, 28 May 1992 (MNHN B-24696). 1 male
(24.7 by 19.0 mm) (TMCD), 1 male (26.3
by 20.5 mm) (ZRC 1999.410), Tungsha Is-
lands, South China Sea, 1265 m, 25 Apr
1996, coll. P-H. Ho.
Description.—Carapace longitudinally
subrectangular, regions clearly defined (Fig.
la). Entire carapace and pereiopod surfaces
with scattered stiff, simple setae which do
not obscure margins or surface, those on
lateral surfaces longer, those on dorsal sur-
face very short. Rostrum well developed,
simple, slightly bent downwards, dorso-me-
dian part furrowed; lateral margins gently
converging to rounded tip (Fig. 1a, b, f, g).
Pseudorostral spines absent (Fig. 1a, f). Su-
praorbital margin sinuous, without spine
but with low subdentiform median lobe
(Fig. la, f). Basal part of eyestalk dilated,
outer surface finely granular (Fig. la, f).
Anterolateral margin with 1 short but dis-
tinct, slightly curved lateral spine. Postero-
lateral margin weakly convex, finely gran-
ulated (Fig. 1a). Posterior margin of cara-
pace weakly concave (Fig. 1a). Protogastric
and epigastric regions with scattered low
granules. Mesogastric and metagastric re-
gions unarmed. Branchial regions covered
with scattered coarse granules (Fig. 1a).
Antennal spine strong, anteriorly directed.
Subhepatic region with 1 large, anteriorly
directed inner spine and 1 small, often
small outer spine or granule; subventral sur-
face with small, sharp spiniform or very
low granule (Fig. 1b, g). Posterior part of
pterygostomial region covered with small
granules (Fig. la, b, g). Gastro-cervical
groove deep, contiguous medially, meeting
below distinct gastric pits. Branchio-cardiac
groove deep. Linea homolica distinct, sin-
uous (Fig. la). Basal antennal article un-
armed. Antennular peduncle subglobular,
unarmed. Proepistome with low, subtrun-
cate lamelliform tooth, otherwise unarmed
(Fig. 1b, g). Merus of third maxilliped un-
armed, almost smooth; ischium subrectan-
gular; palp when appressed against ischium
reaching to proximal margin of ischium;

VOLUME 112, NUMBER 4

exopod slender, reaching to 1/3 length of
outer margin of merus, with long flagellum
(Fig. 1b).

P1 (chelipeds) subequal, elongate, rela-
tively slender (Fig. 1c). Basis-ischium 3-
faceted; inner surface distinctly granular,
other surfaces almost smooth; inner ventral
margin with 4 distinct spines and 1 small
distal spinule; outer ventral margin with 2
spines; dorsal margin with 2 spines (Fig.
1c). Merus with all margins distinctly spi-
niform; inner surface granular; outer and
ventral surfaces almost smooth. Surfaces of
carpus spiniform, with 2 large spines on in-
ner distal angle (Fig. 1c). Outer and inner
surfaces of palm with numerous spines and
spinules; inner surface with 1 distinct lon-
gitudinal row of strong spines (Fig. Ic).
Fingers elongate, distal part gently curving
inwards, anterodorsal margin of dactylus
flattened; cutting edge of dactylus with 1
distinct subproximal, obliquely directed
tooth which fits anterior to distinct proximal
tooth on cutting edge of pollex, rest of cut-
ting edges of both fingers blade-like (Fig.
lc, h). Inner and outer surfaces of base of
pollex of both chelae with large ovate pig-
mented spot each (Fig. Ic, h).

P2 and P3 longest. Coxa of P2—P5 un-
armed. Dorso-distal margin of basis-ischi-
um of P2—P5 with 2 small spines bracketing
merus. Dorsal margin of M2—4 with 14 or
15 distinct spines, distal 0.14—0.17 of seg-
ment usually unarmed; proximal part of
ventral margin with 2 rows of spines which
converge into 1 uneven row of spines along
distal part; anterior surface granular; pos-
terior surface almost smooth. Distal edge of
propodus with 2 stiff, movable setae brack-
eting proximal edge of dactylus. Ventral
margin of dactylus with row of stiff but
movable setae. P5 very slender; when me-
rus articulated forwards, distal end reaching
to base of anterolateral spine; all segments
unarmed; hooked dactylus and distal part of
propodus forming distinct subchelate struc-
ture (Fig. 1d).

Male and female abdomens covering en-
tire thoracic sternum; female abdomen

761

slightly broader than male abdomen (Fig.
le); surfaces of all segments and telson al-
most smooth; telson subtriangular, lateral
margins sinuous, with distal part weakly
concave, and proximal part convex (Fig. le,
i).

Male first pleopod stout, relatively
straight; distal part dilated into distinct flap
(Figs. 1j, 2a, b). Male second pleopod stout,
short, distally cupped (Fig. 1k).

Remarks.—Lamoha longirostris was de-
scribed by Chen (1986) as Hypsophrys lon-
girostris without designating types or indi-
cating the number of specimens available.
The original syntype series consisted of
three males and a female. One of the male
specimens from the East China Sea is here
designated as the lectotype. Lamoha futuna,
as Hypsophrys futuna, was described by
Guinot & Richer de Forges (1995) from
three males and two females collected from
the Wallis and Futuna Islands in the central
Pacific.

Although all the known male specimens
of L. futuna are relatively small, the largest
male specimen being the holotype measur-
ing 18.0 by 15.0 mm; and the present male
specimens of L. longirostris from the East
and South China Seas are all much larger
than the holotype male of L. futuna, we
could discern no major differences between
the two taxa. As such, we here regard La-
moha futuna (Guinot & Richer de Forges,
1995) as a junior subjective synonym of
Lamoha longirostris (Chen, 1986).

In describing their species, Guinot &
Richer de Forges (1995:444) recognized
two main groups of Hypsophrys, but felt
that H. superciliosa Wood-Mason & AIl-
cock, 1891, and H. futuna (= L. longiros-
tris) had intermediate characters which ex-
cluded them from either group. Lamoha su-
perciliosa and L. longirostris are distin-
guished from all other Lamoha species in
having the merus of the last ambulatory leg
completely unarmed and the rostrum being
simple (i.e., not bifurcated at the tip) (Fig.
la, f). Guinot & Richer de Forges (1995:
458) used three main characters to separate

762 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Lamoha longirostris (Chen, 1986). a—e, paralectotype female (22.0 by 18.5 mm) (IOCAS K69B-30);
f—k, paralectotype male (21.2 by 18.0 mm) (IOCAS K69B-30). a, carapace (dorsal view); b, buccal cavity
showing third maxillipeds; c, right cheliped (outer view); d, subchelate dactylus and propodus of P5; e, abdomen;
f, frontal region of carapace; g, epistome, orbit, antennae and antennules; h, distal end of left chela (inner view);
i, abdomen; j, left male first pleopod (ventral view); k, left male second pleopod (dorsal view). Scales equal 5.0
mm (a, b, e, g-i), 1.0 mm (c, d, f), 0.5 mm G, k).

VOLUME 112, NUMBER 4

Fig’: 2:

763

Lamoha longirostris (Chen, 1986). Male (18.3 by 14.8 mm), paratype of Hypsophrys futuna Guinot

& Richer de Forges, 1995 (MNHN B-24696). Left male first pleopod (setae not drawn). a, ventral view; b,

dorsal view. Scales equal 0.5 mm.

L. longirostris from L. superciliosa: sub-
hepatic region with one spine (Fig. la, b, f,
g) (two spines in L. superciliosa), proto-
gastric region smooth (Fig. la) (with some
short spinules in L. superciliosa), and ab-
sence of a spine on the supraorbital margin
(Fig. la, f) (present in L. superciliosa).
These characters are evident on all the spec-
imens examined.

The two pigmented spots at the base of
the pollex on the inner and outer surfaces
of each chela are large and distinctive (Fig.
1c, h) in Lamoha longirostris. In the freshly
preserved specimens (in 70% ethanol) from
the Tungsha Islands, they are dark blue in
the center and bluish along the edges. The
spots in the types of L. longirostris and
Tungsha specimens are much larger than

those figured by Guinot & Richer de Forges
(1995:fig. 66g) and on the types of L. fu-
tuna examined, but this is probably asso-
ciated with size. As noted above, the South
and East Chinese Seas specimens speci-
mens are larger than the holotype male of
L. futuna available to Guinot & Richer de
Forges (1995).

The type specimens of L. longirostris
from the East and South China Seas vary
in several features, none of which are re-
garded as significant. The lateral margins of
the rostrum are subparallel in the paralec-
totype male (ZRC 1999.007) from the East
China Sea but gently converging distally in
the lectotype male (IOCAS KY8B-73). The
pteryogostomial region has a small sharp
granule on each side in the lectotype but in

764

a paralectotype male (ZRC 1999.007), the
right granule is spiniform while the left one
is only granular. The right inner subhepatic
spine of a paralectotype male (ZRC
1999.007) is bifurcated distally but all the
other subhepatic spines in the other speci-
mens are simple.

The known distribution of L. longirostris
is disjunct, being known only from the East
and South China Seas, as well as over 7500
km to the east in the Wallis and Futuna Is-
lands. There are no other records. This,
however, may be due to inadequate sam-
pling, especially considering the depths in
which this species occurs. The type speci-
mens of L. longirostris were collected by
trawls from a depth of about 900-1100 m.
Guinot & Richer de Forges (1995:200) ob-
tained their specimens of H. futuna (= La-
moha futuna) from 1300 m. None of the
specimens were recorded carrying any ob-
ject although members of this genus are
known to carry sea anemones (Guinot et al.
1995):

Genus Homologenus A. Milne-Edwards,
in Henderson, 1888

Type species.—Homolopsis rostratus A.
Milne-Edwards, 1880.

Remarks.—Nine species of Homologen-
us are currently known, of which one oc-
curs in the Indian Ocean and six are present
in the Pacific (Guinot & Richer de Forges
1995). Homologenus donghaiensis Chen,
1986, is the seventh species known from
the Pacific.

Homologenus donghaiensis Chen, 1986
Fig. 3

Homologenus donghaiensis Chen, 1986:

Pd

Material examined.—Holotype male
(11.0 by 10.3 mm), East China Sea, 900 m,
on soft mud, 3 Aug 1981 GOCAS KY8B-
DAY:

Description of holotype male.—Carapace
longitudinally subrectangular, regions well

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

defined (Fig. 3a). Entire dorsal carapace
surface with numerous small granules and
scattered short, stiff setae which do not ob-
scure surface (Fig. 3a). Rostrum well de-
veloped, 3-pronged; accessory spines long,
directed obliquely upwards; median projec-
tion very long, reaching well beyond ac-
cessory spines, directed gently downwards.
Pseudorostral spines very strong, directed
laterally, reaching well beyond eyes (Fig.
3a). Supraorbital margin sinuous, granulat-
ed, inner part with very small slender spine.
Basal part of eyestalk not distinctly dilated
(Fig. 3a). Anterolateral margin with well
developed, obliquely directed long spine;
surfaces of spine finely granular. Postero-
lateral margin gently convex, finely granu-
lated, some granules sharp. Posterior mar-
gin of carapace distinctly concave (Fig. 3a).
Protogastric and epigastric regions with
scattered low granules, those on epigastric
region especially sharp. Mesogastric region
with distinct, long sharp spine. Branchial
regions covered with numerous small gran-
ules. Subhepatic region with 1 large,
obliquely directed spine; ventral surface
with oblique ridge armed with 4 or 5 small
spines (Fig. 3a). Antennal spine long, di-
rected obliquely, longer than subhepatic
spine. Pterygostomial and sub-branchial re-
gions covered with small granules. Gastro-
cervical groove shallow but discernible
(Fig. 3a). Branchio-cardiac groove very
broad, shallow, hardly discernible. Linea
homolica sinuous (Fig. 3a). Basal antennal
article unarmed; peduncle with opening for
gland directed inwards. Antennular pedun-
cle subglobular, with 3 spinules. Proepi-
stome with low, subtruncate lamelliform
tooth. Anterior edge of buccal cavity with
2 small spines. Outer surfaces of third max-
illiped covered with granules; merus elon-
gate, unarmed; ischium subrectangular,
granules on outer margin larger; inner mar-
gin of palp with several spinules; exopod
slender, reaching to mid-point of outer mar-
gin of merus, with long flagellum (Fig. 3f).

P1 (chelipeds) subequal, elongate, slen-
der; surfaces covered with numerous short,

VOLUME 112, NUMBER 4 765

.
.

-t
.
.
>
c
c
c

Fig. 3. Homologenus donghaiensis Chen, 1986. Holotype male (11.0 by 10.3 mm) (IOCAS KY8B-71). a,
carapace (dorsal view); b, right cheliped (outer view); c, right chela (outer view); d, right P2; e, right P5; f, left
third maxilliped; g, abdomen; h, left male first pleopod (ventral view); i, left male second pleopod (dorsal view).

Scales equal 1.0 mm.

766

hooked setae which do not obscure margins
(Fig. 3b). Coxa with inner distal margin dis-
tinctly expanded, armed with 2 small
spines. Basis-ischium with 3 spines on ven-
tral surface; 1 spine on dorsal surface. Inner
margin of merus with 5 or 6 sharp spines;
outer margin with 5 spines; inner surface
granular or with scattered spines. Outer sur-
face of carpus with 2 distinct median spines
and 1 long subdistal spine; inner surface
with 3 evenly spaced spines; ventral surface
with 1 distal spine (Fig. 3b). Outer and in-
ner surfaces of palm with numerous gran-
ules; with 4 short spines on subventral sur-
face; dorsal margin with 1 small distal spine
and 1 longer submedian spine. Fingers
elongate, cutting edges of both fingers
blade-like (Fig. 3b, c).

Surfaces of P2—P5 covered with numer-
ous short, hooked setae which do not ob-
scure margins (Fig. 3d, e). P2 and P3 lon-
gest. Ventral surface of basis-ischium of P2
and P3 with 2 small spines (Fig. 3d). Dorsal
margin of merus of P2—4 with 4 strong,
posteriorly directed spines along proximal
half, distal half mostly unarmed except for
small subdistal spine and longer distal
Spine; proximalmost part of ventral margin
with 2 spines with small subdistal spine
(Fig. 3d, e). Basis-ischium with small spine.
Carpus unarmed (Fig. d, e). P5 slender,
when articulated forwards, distal edge of
merus reaching to gastric spine; merus and
carpus unarmed; propodus with a strong
knob on proximoventral margin which is
tipped with large, slightly movable seta;
curved dactylus and propodus forming sub-
chelate structure (Fig. 3e).

Abdomen covering entire thoracic ster-
num; surface of segment 1 weakly granular;
segment 2 with large, sharp median gran-
ule; surfaces of segments 3—4 with large,
sharp granule on side of each segment, rest
of surface with scattered low granules; seg-
ment 5 with 2 groups of submedian gran-
ules; surfaces of segment 6 and telson al-
most smooth; telson subtriangular, lateral
margins sinuous, distal part concave and
proximal part convex (Fig. 3g).

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Male first pleopod stout, distal part con-
ical, gently curved (fig. 3h). Male second
pleopod stout, short, distally cupped (Fig.
Sil).

Remarks.—Homologenus donghaiensis
belongs to a group of species which have a
long rostrum, a strong gastric spine and the
dorsal margin of the ambulatory merus with
a pronounced distal spine (Guinot & Richer
de Forges 1995:470). Two species are cur-
rently known from this group, H. malay-
ensis Thle, 1912, and H. levii Guinot &
Richer de Forges, 1995.

With regards to its unarmed ambulatory
meri and long P5, H. donghaiensis is mor-
phologically most similar to H. malayensis
from Sulawesi, Ceram, Kei Islands, Phil-
ippines and possibly Japan (Chle 1912,
1913; Guinot & Richer de Forges 1995;
Nagai 1994). Homologenus donghaiensis,
however, differs markedly from H. malay-
ensis in having proportionately much lon-
ger first to third ambulatory meri (Fig. 3d)
(maximum length of P4 merus is ca. 33
times maximum width in H. donghaiensis;
22 times in H. malayensis, cf. Ihle 1913:pl.
2, figs. 13-15). In addition, the P5 of H.
donghaiensis (Fig. 3e) is proportionately
longer, with the merus, when articulated
above the carapace, reaching to the base of
the gastric spine (vs. reaching to below the
anterolateral spine in H. malayensis, cf. Ihle
1913:pl. 2, figs. 13-15); and the distal part
of the male first pleopod is relatively stouter
(Fig. 3h) (vs. more slender in H. malayen-
sis, cf. Ihle 1913:fig. 31). The specimen fig-
ured by Nagai (1994:50, pl. 1, fig. 3) from
Kii Peninsula in Japan appears to be closer
to H. malayensis sensu stricto with regards
to its relatively shorter legs, although it
should be re-examined.

The holotype specimen of H. donghaien-
sis was trawled from some 900 m. The re-
lated H. malayensis has been recorded from
a depth range of 769-1190 m (Guinot &
Richer de Forges 1995).

Acknowledgments

The present study was supported by a re-
search grant (RP 910410) to the first author

VOLUME 112, NUMBER 4

from the university. The second author was
partially supported by a grant from the Raf-
fles Museum during her tenure in Singa-
pore. We are grateful to Daniéle Guinot for
the loan of MNHN specimens. Thanks are
due to Ho Ping-Ho for generously passing
us his Tungsha specimens for study as well
as the associated data and photographs.

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(4):768-777. 1999.

Neocallichirus raymanningi, a new species of ghost shrimp
from the northeastern coast of Venezuela
(Crustacea: Decapoda: Callianassidae)

Juan Pablo Blanco Rambla and Rafael Lemaitre

(JPBR) Instituto Oceanografico de Venezuela, Universidad de Oriente, Apartado Postal 245,
Cumana, Estado Sucre, Venezuela; (RL) Rafael Lemaitre, Department of Invertebrate Zoology,
National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0163

Abstract.—A new species of ghost shrimp, Neocallichirus raymanningi, is
described based on specimens collected in subtidal habitats in the Gulf of
Cariaco, northeastern Venezuela. This new species is most similar to N. rath-
bunae (Schmitt), but differs from it primarily in having a broadly rounded
rostrum, and in the case of males, unique armature of merus and dactylus on
the major cheliped. A checklist of all known species in the world currently
assigned to the genus Neocallichirus Sakai is included.

Recent studies of thalassinideans from
the northeastern coast of Venezuela have
shown the existence of at least 18 species
of these ecologically important infaunal
crustaceans commonly called ghost
shrimps. Several species had not been re-
corded previously from this coast or were
found to be undescribed (Blanco Rambla &
Linero Arana 1994, Blanco Rambla et al.
1995, Blanco Rambla (unpublished data)).
During a recent faunistic survey of the Gulf
of Cariaco, specimens were obtained of yet
another undescribed species as here de-
scribed.

The genus Neocallichirus Sakai, 1988
was originally proposed for nine Indo-West
Pacific and two Atlantic species. Manning
& Felder (1991) later assigned to this genus
four more western Atlantic species. Subse-
quently, Manning & Lemaitre (1994) re-as-
signed four western Atlantic species previ-
ously in Neocallichirus to the genus Sergio
Manning & Lemaitre, 1994. With the de-
scription of the new species, there are now
18 species recognized worldwide in Neo-
callichirus. In addition to the new species,
five other Neocallichirus species occur in
the western Atlantic: N. rathbunae
(Schmitt, 1935), N. grandimana (Gibbes,

1850), N. nickellae Manning, 1993, N. le-
maitrei Manning 1993, and N. cacahuate
Felder & Manning 1995. Species of this ge-
nus are characterized primarily by: carapace
with well defined oval, lacking rostral ca-
rina or cardiac protuberance; corneae dor-
sal, subterminal, disk-shaped; antennular
peduncles no longer or stouter than anten-
nal peduncles; third maxillipeds lacking ex-
opod, with ischium-merus subpediform,
and merus not projecting beyond articula-
tion with carpus; major cheliped lacking
meral hook; in both sexes, first pleopod uni-
ramous, and second pleopod biramous; sec-
ond pleopod with appendix interna well de-
veloped in females, with appendix interna
reduced or short and slender in males; and
endopod of third to fifth pleopods with
stubby, embedded appendix internae (Sakai
1988, Manning & Felder 1991).

Specimens of the new species were cap-
tured with yabby pumps at three localities
in northeastern Venezuela on the southern
and northern coasts of the Gulf of Cariaco
(approximately at 10°30’N, 64°00’W). The
Gulf has about 170 km of shoreline, and
includes diverse habitats.

Measurements (mm) were made with an
ocular micrometer. Carapace length (cl) was

VOLUME 112, NUMBER 4

measured along the middorsal line of the
carapace from the anterior margin of ros-
trum to the posterior margin of the cara-
pace. Total length (tl) was measured from
the tip of the rostrum to posterior margin
of the telson. Specimens are deposited in
the National Museum of Natural History,
Smithsonian Institution, Washington, D.C.
(USNM), and Laboratorio de Carcinologia,
Instituto Oceanografico de Venezuela,
Universidad de Oriente (IOV).

Family Callianassidae Dana, 1852
Subfamily Callichirinae Manning &
Felder, 1991
Neocallichirus Sakai, 1988
Neocallichirus raymanningi, new species
Figs. 1—4

Holotype.—Playa Cristal, south coast of
Gulf of Cariaco, Sucre State, Venezuela,
10°27'07,0"N, 63°57'01,7"W, sandy sub-
strate, 0.5 m, 5 Aug 1997, coll. J. P. Blanco
Rambla, M. Gomez, & A. De La Rosa: 1
3 cl 23.1 mm, tl 76.1 mm, USNM 276147.

Paratypes.—Same locality data as holo-
type: 1 2 (dry, damaged) cl 20.6 mm, tl 69
mm, USNM 276150.—Las Maritas, north
coast of Gulf of Cariaco, Sucre State, Ven-
ezuela, 10°33'05.2”N, 63°50'45.5’W, subti-
dal sand-mud substrate, 4 Feb 1997, colls.
J. PR Blanco Rambla, M. Gomez, & A. De
Mapkesa 1 2 cl.23.1 mm, tl 80.1 mm,
USNM 276148.—La Pefia, south coast of
Gulf of Cariaco, Sucre State, Venezuela,
10°28'08.5”N, 63°42'26.4"W, mud-sand
substrate, 0.5 m, 26 Jun 1997, coll. J. P.
Blanco Rambla, M. Gémez & A. De La
Rosa: 1 @ cl 21.9 mm, tl 79.0 mm, USNM
276149; 1 2 cl 19.1 mm, tl 70.6 mm, IOV
1IS76:

Diagnosis.—Front of carapace with ros-
trum broadly rounded, each lateral projec-
tion armed with terminal spine slightly ex-
ceeding rostrum. Inferior portion of lateral
wall of carapace with 3 small sharp or blunt
spines: 1 on hepatic boss, and 2 just pos-
terior to hepatic boss. Major cheliped merus
armed with row of strong bifid teeth on in-

769

ferior margin proximally; prehensile margin
of dactylus with large proximal subrectan-
gular tooth separated from distal margin by
deep quadrate cleft. Male second pleopod
with distinct, slender appendix interna bear-
ing short stiff setae terminally.

Description.—Mature male and females
of relatively large size, ranging in known
specimens from cl 19.1 to 23.1 mm, and tl
from 69.0 to 80.1 mm.

Frontal margin of carapace (Fig. la, b)
consisting of broadly rounded rostrum, and
subtriangular lateral projections to each
side; rostrum with short marginal setae; lat-
eral projections each with terminal spine
slightly exceeding rostrum and reaching to
about midline of basal segment of antennal
peduncle. Carapace lacking rostral carina,
with distinct linea thalassinica; dorsal oval
with distinct tubercle on each side of ante-
rior half, oval distinctly marked posteriorly
by deep transverse cardiac furrow, latter ex-
tending anteroventrally to either side above
linea thalassinica as shallow groove mark-
ing posterior half of dorsal oval. Frontal
margin of carapace continued ventrolater-
ally beyond intersection with linea thalas-
sinica as thickened oblique ridge ending an-
teriorly to prominent hepatic boss, latter
surmounted with small sharp or blunt tu-
bercle followed posteriorly by 2 small sharp
or blunt spines. Subantennular region of ep-
istome bearing dense tuft of long setae.

Eyestalks (Fig. la, b) subtriangular, with
flattened and moderately pigmented cor-
neae occupying less than half width of
stalk. Length of exposed eyestalk in dorsal
view about 1.5 times basal width; tips lo-
bate to obtusely angular, reaching distal end
of basal segment of antennular peduncle;
mesial margins of eyestalks closely ap-
pressed proximally, diverging terminally.

Antennular peduncle shorter and heavier
than antennal peduncle; terminal article
about 1.8 times length of penultimate,
reaching to about proximal third of terminal
antennal article; penultimate and terminal
articles with ventromesial and ventrolateral
rows of long setae; rami of flagellum sub-

FD

Pig.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Neocallichirus raymanningi, new species, Gulf of Cariaco, Venezuela. a, c, e, Male holotype (cl 23.1

mm), USNM 276147; b, d, Female paratype (23.1 mm), USNM 276148. a, Anterior carapace, eyestalks and
peduncles of cephalic appendages, dorsal view; b, Anterior carapace, eyestalks and portion of peduncles of
cephalic appendages, dorsal view; c, Male major cheliped, internal surface; d, Female major cheliped, internal
surface; e, Minor cheliped, internal surface. Scales equal 2 mm.

equal in length, dorsal ramus with sparse
setae, ventral ramus with dense long setae.

Antennal peduncle with terminal article
slightly shorter than penultimate; basal ar-
ticle with slightly produced excretory pore;
second article with distinct oblique ventral
suture, distolaterally with tuft of setae; third
article narrower than second; fourth article
longest, slightly narrower than third, and
with tuft of long setae distolaterally.

Mandible (Fig. 2a) with large, 3-seg-
mented palp; third article of palp elongated,
rounded distally, and with short, weakly
hooked setae on extensor surface; second
article of palp with long setae distally; first
article as long as wide, without setae. In-
cisor process with cutting margin consisting
of well defined blunt corneous teeth on
proximal half, and long, low tooth on distal
half; internal surface with lip giving rise to

VOLUME 112, NUMBER 4

molar process with about 5 blunt teeth; par-
agnath (Fig. 2b) scaliform, with small scler-
otized projection on upper distal half, set
against and below molar process. First max-
illa (Fig. 2c) with endopodal palp long, nar-
row, and with terminal article deflected
proximally at articulation; proximal endite
with dense fine setation on most of lower
mesial margin, terminal lobe with long, dis-
tally bifurcate setae; distal endite elongate,
narrow proximally, mesial margin with
short bristles often bifurcated distally; ex-
opod obsolete, marked by low truncate lobe
with setae. Second maxilla (Fig. 2d) with
endopod terminally flexed, rounded distal-
ly; proximal and distal endites each longi-
tudinally subdivided and densely setose ter-
minally, setae on second endite consisting
of dense bristles; proximal endite with ar-
cuate setose crest across internal surface;
exopod forming large, broad, scaphogna-
thite. First maxilliped (Fig. 2e) with proxi-
mal endite narrowly produced, marginally
setose; distal endite robust, subrectanglar,
external surface with median longitudinal
row of long setae directed mesially; exopod
ovoid, marginally setose, with marked
notch on mesial margin, and short arcuate
row of setae on external surface; epipod
large, posterior lobe broad, anterior end
narrowing terminally. Second maxilliped
(Fig. 2f) with long, narrow endopod; en-
dopodal merus more than 4 times as long
as broad, flexor margin with dense fringe of
long setae; carpus short; propodus about
half as long as merus, slightly arcuate, wid-
ening distally; dactylus elongate, about half
as long as propodus, with terminal brush of
short stiff setae; exopod narrow, slightly ar-
cuate, overreaching end of endopodal car-
pus, and fringed with long setae; epipod
small. Third maxilliped (Fig. 2g) without
exopod; endopod with long setation on me-
sial margin; terminal 3 articles also with
long setation on extensor margins; length of
merus-ischium more than 2.5 times its
width; ischium subrectangular, distinctly
longer than broad, proximomesial margin
rounded, internal surface with well defined

771

crista dentata consisting of curved row of
sharp spines, distalmost spines closely set,
median spines distinctly larger and wide
apart; merus subtriangular, broader than
long; carpus subtriangular, with setose lobe
on flexor margin, internal surface with
dense semicircular field of fine setae; pro-
podus subrectangular, height more that 1.3
its length, internal surface with median sub-
circular field of fine dense setae, anterior
margin setose and ending distally as lobe;
dactylus narrow, slightly arcuate, longer
than propodus, with small brush of stiff
bristles; lacking epipod.

Branchial formula includes exopods and
epipods as described for first to third max-
illipeds. Other branchiae consisting of 1 ru-
dimentary arthrobranch on second maxilli-
ped, pair of arthrobranchs on third maxil-
liped, and pair of arthrobranchs on each of
first to fourth pereopods.

First pereopods forming markedly dis-
similar chelipeds. Major cheliped of male
(Fig. 1c) heavy, strongly calcified; ischium
with row of well spaced sharp or blunt
spines on inferior margin; merus about 2.3
times as long as high, superior margin
slightly arcuate distally, inferior margin
keel-like and armed with row of strong
teeth decreasing in size distally, teeth bifid
proximally and simple distally; carpus
short, about 1.9 times higher than long, su-
perior margin keel-like and with tufts of
short setae on inner side, posterior and
proximal part of inferior margins crenulate
with tufts of setae on inner side, evenly
curved and forming blunt angle with distal
part of inferior margin; chela about 1.8
times as long as high (greatest height on
palm proximally); palm about 1.1 times as
high as long (length measured on superior
margin), outer surface smooth, glabrous ex-
cept for scattered tufts of short setae, inner
surface with scattered tufts of setae and
cluster of low tubercles medially near base
of fixed finger, superior margin broadly ar-
cuate and with tufts of setae on inner side,
inferior margin keel-like and turned in-
wards, crenulate and with tufts of long setae

TG2.

\ \ WY oy
77}
SA. Wi Yj
= \ a y
=> /
SSW
= SSS
Cay

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Neocallichirus raymanningi, new species, male holotype (cl 23.1 mm), Gulf of Cariaco, Venezuela,
USNM 276147, left mouthparts: a, Mandible, external surface; b, Paragnath, external surface; c, First maxilla,
external surface; d, Second maxilla, external surface; e, First maxilliped, external surface; f, Second maxilliped,
external surface; g, Third maxilliped, internal surface. Scales equal 1 mm (a—e), and 2 mm (f, g).

on inner side; fixed finger with tip curved
upward, inferior margin with tufts of setae
on inner side, prehensile margin with small
rounded teeth proximally, and small or in-
conspicuous rounded teeth distally; dacty-
lus slightly longer than palm, outer and in-
ner surfaces with tufts of long setae in prox-
imity to opposable margin, superior margin
arcuate and with tufts of setae on inner and
outer sides, tip curved downward, prehen-
sile margin with large subrectangular tooth
proximally and separated by deep subquad-
rate cleft from low rounded tooth, followed
distally with row of small rounded teeth.
Major cheliped of female (Fig. 1d) not as
heavy and massive as in male; differing
from that of male as follows: merus slightly
more slender, about 2.4 times as long as
high; teeth on inferior margin of merus sim-
ple; carpus about as long as high; chela
more slender, about 2.3 times as long as

high, palm about 1.2 times as long as high
(length measured on superior margin); fixed
finger with prehensile margin forming a
broadly triangular tooth at midline; oppos-
able margin of dactylus with proximal tooth
broader and lower.

Minor cheliped (Fig. le) similar between
sexes, well calcified much less massive and
more slender than in male; outer and inner
surfaces of segments glabrous except for
scattered tufts of short setae on chela. Is-
chium with inferior margin unarmed. Merus
about 2.3 times as long as high, inferior
margin slightly sinuous. Carpus about twice
as long as high, slightly longer than merus,
Superior and inferior margins unarmed,
with row of tufts of setae on interior side.
Chela about 2.7 times as long as high. Palm
about 1.3 times as long as high (length mea-
sured on superior margin), superior margin
with row of tufts of setae; palm and fixed

VOLUME 112, NUMBER 4

finger with inferior margin slightly concave
at base of fixed finger, with row of tufts of
setae. Fixed finger shorter than dactylus, tip
curved upward; prehensile margin minutely
denticulate. Dactylus slightly longer than
palm, tip curved downward; superior mar-
gin with tufts of setae; opposable margin
minutely denticulate on proximal half.

Second pereopod (Fig. 3a) chelate. Is-
chium with sparse long setae on inferior
margin. Merus with longitudinal row of
well-spaced tufts of short setae on outer
surface; flexor margin with dense row of
long setae over most of margin, setae di-
minishing in length distally. Carpus with
long setae on superior and inferior margins;
outer surface with scattered setae in addi-
tion to row of. tufts of short setae parallel
to distal margin; inner surface with well
spaced tufts of setae. Chela with fingers
straight prehensile and opposable margins
of fingers corneous, micropectinate proxi-
mally and smooth distally; outer and inner
surfaces of palm and fingers with well
spaced tufts of setae.

Third pereopod (Fig. 3b) with merus
about 2.8 times as long as high. Carpus
widening distally, twice as long as high,
with patches of long setae terminally. Pro-
podus with inferior margin produced into
large lobe directed proximally; outer sur-
face with numerous tufts of short setae ex-
cept on longitudinal median portion; inner
surface with scattered tufts of short setae;
superior margin with tufts of long setae; in-
ferior margin with dense tufts of long setae.
Dactylus tear-shaped, terminating in cor-
neous tip hooked toward external side; out-
er surface densely covered with long setae;
inner surface with scattered setae.

Fourth pereopod (Fig. 3c) subchelate. Is-
chium about 2.6 times as long as high. Me-
rus longer and heavier than carpus, about 4
times as long as high. Carpus slender, near-
ly 6 times as long as high, slightly widening
distally. Propodus about 3.5 times as long
as high; inferodistal corner produced into
short, blunt fixed finger; outer surface
densely covered with long microserrate se-

higis.

tae except on longitudinal median portion;
inner surface with scattered setae. Dactylus
about twice as long as high, terminating in
short corneous tip hooked toward external
side; outer surface densely setose.

Fifth pereopod (Fig. 3d) minutely che-
late. Merus and carpus with scattered setae.
Propodus with dense patch of dense setae
on distal half, upper limit of patch oblique.
Fixed finger and dactylus with opposable
surfaces spooned; prehensile lip of fixed
finger corneous, minutely pectinate. Dac-
tylus curving inward, densely setose on out-
er surface.

Abdominal somites smooth, glabrous
dorsally except for short setae near poste-
rior margin. Third to fifth somites laterally
with semicircular tuft of dense short setae
laterally. Sixth somite with 3 dorsal pairs of
setae or tufts of setae (1 pair anteriorly, 1
pair medially, and 1 pair of longer setae on
posterior margin), and short fine marginal
setation; with short posterior mid-dorsal
sulcus reaching posterior margin.

First pleopod of male and female unira-
mous, 2-segmented; in male (Fig. 4a), distal
segment spatulate, shorter than proximal,
subdivided into 2 lobes by weak longitu-
dinal furrow, anterior lobe terminally
rounded and setose, posterior lobe termi-
nally acute and setose with tip directed an-
teriorly; in female (Fig. 4d), both articles
narrow and elongate, proximal article
curved outward and setose distally, terminal
article marginally setose on distal half and
with distinct setose shoulder at midlength.
Second pleopod of male and female bira-
mous; in male (Fig. 4b), exopod setose dis-
tally, slender and curved inward, endopod
with distal lobe demarcated by weak trans-
verse suture and longer than poorly demar-
cated appendix masculina, appendix interna
(Fig. 4c) short and slender, with 3 short stiff
setae distally; in female (Fig. 4e), exopod
with marginal setae distally, endopod with
well developed appendix interna (Fig. 4f)
about 0.5 as long as distal lobe of endopod
and with short hooked setae distally. Third
to fifth pairs of pleopods forming large,

774

Fig. 3.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

y\\ty!
Nah

Z *
Ly :
Ly,

Ly
tH
ey

ANS
AN
\
~

\
Ne

Neocallichirus raymanningi, new species, male holotype (cl 23.1 mm), Gulf of Cariaco, Venezuela,

USNM 276147: a, Right second pereopod, external surface; b, Right third pereopod, external surface; c, Right
fourth pereopod, external surface; d, Right fifth pereopod, external surface; e, Telson and uropods, dorsal view.

Scales equal 3 mm.

posteriorly cupped fans when coupled at
mesial margins of endopods; endopod of
each (Fig. 4g, h) subtriangular, short, with
stubby appendix interna embedded into me-
sial margin of endopod.

Telson (Fig. 3e) about 1.3 times as broad
as long, broadest at lateral lobes on anterior
half; posterior margin broadly rounded or
weakly sinuous, setose; posterolateral cor-
ners rounded, each bearing tuft of long se-
tae; dorsal surface anteromedially elevated
and with tuft of setae medially. Uropod
with posterolaterally directed rounded lobe
of protopod overreaching anterior margin of
endopod; endopod broader than long, sub-
rectangular, posterior margin truncate, near-
ly straight dorsal surface with longitudinal

carina and broad tuft of long setae on pos-
terolateral corner; exopod with anterodorsal
plate with dense spiniform setae distally,
posterior plate marginally with dense, long
spiniform setae grading posteriorly into
thinner setae.

Known range and habitat.—Known only
from the northern and southern shores of
the Gulf of Cariaco, Venezuela. The speci-
mens were found in shallow subtidal habi-
tats at about 0.5 m in depth, on sand or
sand-mud substrates. Salinity and tempera-
ture ranged from 34 to 35 ppt, and 23.6° to
25°C, respectively.

Etymology.—The specific name is given
in honor of our colleague and friend, Ray-
mond B. Manning, in recognition of his

VOLUME 112, NUMBER 4

T7135

Fig. 4. Neocallichirus raymanningi, new species, Gulf of Cariaco, Venezuela. a—c, g, h, Male holotype (cl
23.1 mm), USNM 276147; d—f, Female paratype (23.1 mm), USNM 276148. a, Male left first pleopod, posterior
surface; b, Male left second pleopod, posterior surface (arrow indicates appendix masculina); c, Appendix interna
of same; d, Female first right pleopod, posterior surface; e, Female second right pleopod, posterior surface; f,
Appendix interna of same; g, Third pleopod, posterior surface; h, Appendix interna of same, posterior surface.
Scales equal 1 mm (a, b, d, c, h), 2 mm (e, g) and 0.5 mm (f).

contributions to the systematics of the Cal-
lianassidae. Over the years he has helped
and encouraged us to collect and study
these poorly known crustaceans.
Remarks.—Among the western Atlantic
species of Neocallichirus, N. raymanningi,
new species, is most similar to N. rathbu-
nae (Schmitt, 1935). The two can be sepa-
rated easily by differences in the frontal
margin of the carapace and, in the case of
males, the major cheliped. In the new spe-
cies the rostrum is broadly rounded and un-
armed, whereas in N. rathbunae the rostrum
is subtriangular and has a terminal spine.

The major cheliped of the only known male
of the new species has a row of irregular,
bifid teeth on the inferior margin of the
merus; the prehensile margin of the dacty-
lus has a large proximal subrectangular
tooth, which is separated from the distal
part of the margin by a deep subquadrate
cleft. In contrast, the major cheliped in
males of N. rathbunae have simple spines
on the inferior margin of the merus; the pre-
hensile margin of the dactylus has a large,
low subrectangular tooth proximally, which
is separated from the rest of the margin by
a Shallow rounded (U-shaped) cleft.

776

The second pleopod of the male in Neo-
callichirus raymanningi, new species, and
N. rathbunae, is distinct from that of other
western Atlantic congeners in having a well
developed, slender appendix interna with
short stiff setae terminally (Fig. 4b). In oth-
er western Atlantic congeners the appendix
interna is obsolete or at most a vestige is
present in the form of a small patch of short
microsetae.

The following is a checklist of all Neo-
callichirus species currently known from
the world, and their general distribution:

N. cacahuate Felder & Manning, 1995:
western Atlantic (Florida, U.S.A.).

N. caechabitator Sakai, 1988: western Pa-
cific (Australia).

N. darwinensis Sakai, 1988: western Pacific
(Australia).

N. denticulatus Ngoc-Ho, 1994: western
Pacific (Australia).

N. grandimana (Gibbes, 1850): western At-
lantic (Bermuda; Caribbean Sea; Brazil),
and eastern Pacific (Panama to Ecuador).

N. horneri Sakai, 1988: western Pacific
(Australia).

N. indica De Man, 1905: Indo-West Pacific
(Mauritius; Java Sea; Japan).

N. lemaitrei Manning, 1993: western Atlan-
tic (Colombia).

N. limosa (Poore, 1975): western Pacific
(Australia).

N. manningi Kazmi & Kazmi, 1992: Indian
Ocean (Arabian Sea, Pakistan).

N. moluccensis (De Man, 1905): western
Pacific (Indonesia).

N. natalensis (Barnard, 1947): western In-
dian Ocean (Natal, South Africa).

N. nickellae Manning, 1993: western Atlan-
tic (Tobago).

N. pachydactyla (A. Milne-Edwards, 1870):
eastern Atlantic (Cape Verde Islands;
Senegal; Ghana).

N. rathbunae (Schmitt, 1935): western At-
lantic (Florida, U.S.A.; Bahamas; U.S.
Virgin Islands; Jamaica).

N. raymanningi, new species: western At-
lantic (Venezuela).

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

N. sassandrensis (Le Loeuff & Intés, 1974):
eastern Atlantic (Ivory Coast).

N. taiaro Ngoc-Ho, 1995: South Pacific
(French Polynesia).

Acknowledgments

We thank R. B. Manning (USNM), for
his helpful advice and valuable suggestions;
and A. De La Rosa and M. Gémez (IOV),
for their assistance during field work. This
study was possible thanks to support pro-
vided to one of us (JPBR) through grant CI-
5-1803-0767/96-97 from Consejo de Inves-
tigaci6n, Universidad de Oriente, Venezuela
(CIUDO). Travel to the National Museum
of Natural History, Smithsonian Institution,
Washington, D.C., was possible thanks to
funds provided by CIUDO, and Consejo
Nacional de Investigaciones Cientificas y
Tecnolo6gicas (CONICIT), Caracas, Vene-
zuela.

Literature Cited

Barnard, K. H. 1947. Descriptions of new species of
South African decapod Crustacea, with notes on
synonymy and new records.—Annals and Mag-
azine of Natural History 13(11):361-392.

Blanco Rambla, J. PB, & I. Lifero Arana. 1994. New
records and new species of ghost shrimps (Crus-
tacea: Thalassinidea) from Venezuela.—Bulle-
tin of Marine Science 55:16—29.

; , & L. B. Lares M. 1995. A new cal-
lianassid (Decapoda: Thalassinidea) from the
southern Caribbean Sea.—Proceedings of the
Biological Society of Washington 108:102—106.

Dana, J. D. 1852. Macroura. Conspectus Crustaceorum
& Conspectus of the Crustacea of the Exploring
Expedition under Capt. C. Wilkes, U.S.N.—
Proceedings of the Academy of Natural Scienc-
es of Philadelphia 6:10—28.

Felder, D. L., & R. B. Manning. 1995. Neocallichirus
cacahuate, a new species of ghost shrimp from
the Atlantic coast of Florida, with reexamina-
tion of N. grandimana and N. lemaitrei (Crus-
tacea: Decapoda: Callianassidae).—Proceedings
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Gibbes, L. R. 1850. On the carcinological collections
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tained in them, with notes on the most remark-
able, and descriptions of new species.—Pro-

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ceedings of the American Association for the
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Manning, R. B. 1993. Two new species of Neocalli-
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, & D. L. Felder. 1991. Revision of the Amer-

ican Callianassidae (Crustacea: Decapoda:

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TH

anassa (Leach) et description des plusieurs es-
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. 1995. Une espéce nouvelle de Neocallichirus
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(4):778—786. 1999.

Cucumaria flamma, a new species of sea cucumber from the central

eastern Pacific (Echinodermata: Holothuroidea)

Francisco A. Solis-Marin and Alfredo Laguarda-Figueras

Laboratorio de Sistematica y Ecologia de Equinodermos, Instituto de Ciencias del Mar y
Limnologia, Universidad Nacional Aut6noma de México, Apdo. Post. 70-305
México, D.F 04510

Abstract.—Cucumaria flamma, n. sp. from the central eastern Pacific is of
medium size with mouth and anus terminal, directed upward; a conspicuous
deep antero-posterior groove in the dorsal skin extending from near base of
tentacles to anus. Ventral skin ossicles are knobbed buttons and smooth per-
forated plates, dorsal skin ossicles are smooth perforated plates and smooth
buttons. Distributed from Mazatlan, Sinaloa, México to Isla de la Plata, Ec-
uador from 4 to 12 m on rocky substrata.

Resumen.—Cucumaria fiamma, n. sp. del Pacifico este central, es una es-
pecie de talla mediana, con boca y ano en posicién terminal, dirigidos hacia
arriba, con un surco antero-posterior en la superficie dorsal, que se extiende
desde cerca de la base de los tentaculos hasta el ano. Los osiculos de la pared
ventral del cuerpo son botones abollonados y placas lisas perforadas, los de la
pared dorsal son placas lisas perforadas y botones lisos. Distribuido desde
Mazatlan, Sinaloa, México hasta Isla de la Plata, Ecuador, de los 4 alos 12 m

de profundidad, sobre substratos rocosos.

Of the two hundred and forty seven spe-
cies currently named in the family Cucu-
mariidae, the genus Cucumaria (Subfamily
Cucumariinae), embraces one hundred and
sixty four. As far as we know, three of these
species occur in the central eastern Pacific:
Cucumaria crax Deichmann, 1941, C.
pseudocurata Deichmann, 1938 and C. sal-
ma Yingst, 1972. Lambert (1998a) removed
C. curata Cowles, 1907 from the genus
based on morphology and DNA evidence.
The systematic status of certain groups
within the family remains somewhat con-
fused, but the works of Panning (1949—
1971), Pawson & Fell (1965), Pawson
(1970, 1982) and Lambert (1998a, 1998b)
are steadily eliminating the confusion. The
purpose of this paper is to describe a new
species of Cucumaria from the eastern Pa-
cific coast.

Material and Methods

Live specimens were relaxed with chloral
hydrate and fixed with 70% ethyl alcohol.
Ossicles were extracted from pieces of skin
(4 mm square) from mid-dorsal and mid-
ventral body wall, tentacles and introvert.
The skin was dissolved with fresh house-
hold bleach in centrifuge tubes. After cen-
trifugation at 2750 rpm for 10 minutes,
bleach was pipetted off and the ossicles
were flooded and centrifuged three times
with ethyl alcohol. After drying, the ossi-
cles were observed by scanning electron
microscopy.

Results

Order Dendrochirotida Grube, 1840
Family Cucumariidae Ludwig, 1894

Diagnosis.—Body wall essentially soft,
ossicles small and inconspicuous. Calcare-

VOLUME 112, NUMBER 4

ous ring simple, lacking posterior process-
es. Usually 10-20 tentacles are present;
tube feet most commonly restricted to the
radii, where they may be scattered or ar-
ranged in regular rows. In several species
some feet may also be scattered in dorsal
interradii (after Pawson 1982).

Remarks.—The family was emended by
Pawson & Fell (1965).

Subfamily Cucumariinae Panning, 1949

Diagnosis.—Ten tentacles. Calcareous
ring simple, low, without forked tails, or at
most medium high, with short undivided
forked-tails, the radials and inter radials un-
divided. In the skin, only plates, no cups,
no tables, no table-like constructions (after
Panning 1949).

Genus Cucumaria de Blainville, 1834

Diagnosis.—Ten tentacles, of equal size,
or the ventral ones smaller. Feet large, soft,
in 5 bands, in some forms also scattered in
the interradii, particularly dorsally. Calcar-
eous ring low, simple. Ossicles plates or
buttons, knobbed or smooth, often re-
duced. Feet with simple end plates or none
at all; walls of feet supported by rods, of-
ten 3 armed, or plates. Introvert and ten-
tacles with perforated plates and rods. Os-
sicles show tendency to become reduced in
many forms (after Deichmann 1941). Type
species: Cucumaria frondosa Gunnerus,
1770:

Remarks.—The diagnosis embraces only
the members of the genus Cucumaria sensu
stricto. The genus contains about 165 de-
scribed and attributed species, of which
about 43 are currently recognized. The ge-
nus was emended by Panning (1949).

Cucumaria flamma, new species
Figs. 1—5

Pattalus mollis Nepote, 1998: 50-52, fig.
16. Non: Pattalus mollis Selenka, 1868.

Diagnosis.—Body curved, mouth and
anus terminal, directed upward. Medium

a9

size form, reaching a length of approxi-
mately 150 mm; skin soft, smooth; feet soft,
large, retractile, particularly abundant in
ventral region, not arranged in bands, also
present in interambulacra. A conspicuous
antero-posterior deep groove in dorsal skin
extends from near base of tentacles to anus.
Ten large dendritic tentacles with broad
base, soft, richly branched, bushy; often
deep orange, speckled with black. Calcare-
ous ring simple, low and stout with short
anterior projections, lacking posterior pro-
cesses. Radial and interradial pieces almost
same size. Ventral skin ossicles four to six-
holed knobbed buttons (110-150 wm) and
less abundant smooth perforated plates (200
zm). Dorsal skin ossicles smooth perforat-
ed plates and smooth buttons with two cen-
tral holes bigger than rest. Introvert with
star-shaped (~340 wm) or elongated per-
forated plates (200 wm). Tentacle ossicles
in two forms: abundant large, perforated,
robust rods, tree-branched, curved or elon-
gated, and scarce oval perforated plates,
with scalloped margins.

Material examined.—Thirty specimens
from 9 localities on the Mexican Pacific
Coast: four in Sinaloa, one in Nayarit, one
in Colima and three in Guerrero, and two
localities in Central America: one in Pana-
ma and one in Ecuador (Fig. 5). Number of
specimens in parentheses after the cata-
logue number; depth in meters (m). The ab-
breviations used in this paper are: United
States National Museum (USNM), Califor-
nia Academy of Sciences (CASIZ), Royal
British Columbia Museum (RBCM), Natu-
ral History Museum of Los Angeles County
(LACM), Instituto de Ciencias del Mar y
Limnologia, Colecci6n Nacional de Equi-
nodermos, Universidad Nacional Autonoma
de México (CML-UNAM).

Holotype.-—USNM E48072 (1), collect-
ed by FE A. Solis-Marin & R. Ramirez-Mu-
rillo, 18 Jun 1993, length 142 mm (mea-
sured along the outside of the curved body),
female.

Type locality.—Mexican Pacific coast,
La Pedregosa Beach, Zihuatanejo, Guerre-

780

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Big. te

Paratype of Cucumaria flamma collected by scuba from 6 m, Playa La Pedregosa, Zihuatanejo,

Guerrero, Mexico, (17°37'06"N, 101°31'52”W), length 140 mm, ICML-UNAM 5.8.0. Female. 1A, left lateral

view. 1B, dorsal view. Note conspicuous dorsal groove.

ro, Mexico, (17°37'06’N, 101°31'52”W), 6
m, rocky substrate.

Paratypes.—USNM E48073 (1), collect-
ed by E A. Solis-Marin & R. Ramirez-Mu-
rillo, 18 Jun 1993, La Pedregosa Beach,
Zihuatanejo, Guerrero, Mexico, (17°37'06'N,
101°31'52”W), 6 m, rocky substrate, spec-
imen length 141 mm. ICML-UNAM 5.8.0
(2), collected by FE A. Solis-Marin & R. Ra-
mirez-Murillo, 18 Jun 1993, Playa La Ped-
regosa, Zihuatanejo, Guerrero, Mexico,
(17°37 '06'N :10173.1'52" W) 6 amis stocky.
substrate, specimens length from 130 to 151
mm. ICML-UNAM 5.8.1 (1), collected by
Elaine M., Playa La Peruana, Morro Pul-
pito, Colima, Mexico, 6 m, 10 Jan 1995,
length 62 mm. ICML-UNAM 5.8.2 (1),
Collected by J. M. Salcedo, 22 Nov 1981,
Morro del Tigre, Bahia de Zihuatanejo,

Guerrero, Mexico, (17°38'04’N, 101°33'07’W),
5 m, rocky substrate, specimen length 90
mm. ICML-UNAM 5.8.3 (6), Collected by
Facultad de Ciencias, UNAM, 5 Sep 1965,
Playa Las Gatas, Zihuatanejo, Guerrero,
Mexico, (17°38'10"N, 101°33’0"W), 4 m,
rocky substrate, specimens length from 55
to 132 mm. ICML-UNAM 5.8.4 (1), Col-
lected by Caso, M. E., 17 Jun 1980, Isla
Cardones, Bahia de Mazatlan, Sinaloa,
Mexico, (23°11'09"N, 106°24'24”W), 8 m,
rocky substrate, specimen length 85 mm.
ICML-UNAM 5.8.5 (2), Collected by O.
L6épez, 19 Mar 1992, Isla Venados, Bahia
de Mazatlan, Sinaloa, Mexico, (23°15'39’N,
106°28'38”W), 12 m, rocky substrate, spec-
imens length from 99 to 130 mm. ICML-
UNAM 5.8.6 (3), Collected by O. Lopez,
22 Mar 1992, Off Isla Cardones, Bahia de

VOLUME 112, NUMBER 4

Fig. 2. Calcareous ring of Cucumaria flamma;
scale bar 6 mm. From paratype (USNM E48073), one
radial and two interradials pieces.

Mazatlan, Sinaloa, Mexico, (23°11'04’N,
106°24'23"W), 10 m, rocky substrate, spec-
imens length from 80 to 125 mm. ICML-
UNAM 5.8.7 (1), Collected by O. L6pez,
25 Jan 1992, Isla Chivos, Bahia de Maza-
tlan, Sinaloa, Mexico, (23°13'46’'N,
106°27'53"W), 8 m, rocky substrate, spec-
imen length 85 mm. ICML-UNAM 5.8.8
(1), Collected by M. E. Caso, 25 Jan 1983,
Isla Cardones, Bahia de Mazatlan, Sinaloa,
Mexico, (23°11'03"N, 106°24'10’W), 8 m,
rocky substrate, specimen length 102 mm.
ICML-UNAM 5.8.9 (1), Collected by M. E.
Caso, 24 Jan 1983, Isla Pajaros, Bahia de
Mazatlan, Sinaloa, Mexico, (23°15’00’N,
106°28'52”W), 10 m, rocky substrate, spec-
imen length 90 mm. ICML-UNAM 5.8.10
(2), Collected by M. E. Caso, 20 Jan 1983,
Isla Venados, Bahia de Mazatlan, Sinaloa,
Mexico, (23°15'39"N, 106°28'38"W), 4 m,
rocky substrate, one specimen length 72
mm, the other specimen was partially de-
stroyed. ICML-UNAM 5.8.11 (1), Collect-
ed by A. C., Nepote, 20 May 1996, Islas
Marietas (Isla Larga), Bahia de Banderas,
Nayarit, Mexico (20°41'98"N,
105°34'72"W), 4 m, rocky substrate, spec-
imen length 120 mm. ICML-UNAM 5.8.12
(1), Collected by E A. Solis-Marin, 21 Mar
1996, Islas Marietas (Isla Redonda), Bahia

781

de Banderas, Nayarit, Mexico (20°42'04’N,
105°33'53”"W), 10 m, rocky substrate, spec-
imen length 132 mm. USNM E47607 (1),
Collected by Emlet, R., 14 Nov 1983,
North Pacific Ocean, Panama, Taboguilla
Island, depth unknown, subtidal rock holes,
specimen length 139 mm. USNM E47750
(1), Collected by E Rivera, May 1998, Isla
de La Plata, Ecuador, (1°16’N, 81°06’W), 5
m, rocky substrate and coral debris, speci-
men length 104 mm (female). CASIZ
102949 (1), Collected by O. L6pez, 22 Mar
1982, Off Isla Cardones, Bahia de Maza-
tlan, Sinaloa, Mexico, (23°11'04’"N,
106°24'23”"W), 10 m, subtidal rock holes,
specimen length 100 mm. RBCM 999-
00015-001 (1), Collected by Facultad de
Ciencias, UNAM, 5 Sep 1965, Playa Las
Gatas, Zihuatanejo, Guerrero, Mexico,
(17°38'10"N, 101°33'0"W), 4 m, rocky sub-
Strate, specimens length 80 mm. LACM 92-
193.1 (1), Collected by O. Lépez, 19 Mar
1992, Isla Venados, Bahia de Mazatlan, Sin-
aloa, Mexico, (23°15'39"N, 106°28'38”"W),
12 m, rocky substrate, specimens length 99
mm.

Description.—Total length 55-151 mm
(measured along the greater curvature of
the body); mean length 101 mm. Holotype
141 mm long. Body curved, mouth and
anus terminal, directed upward; anterior re-
gion wider than posterior. Deep groove in
dorsal region, runs from near base of ten-
tacles to anus (Fig. la, b). Skin soft,
smooth. Podia slender, soft, large, retractile,
widely dispersed and particularly larger and
abundant in ventral region, not arranged in
bands; also present in interambulacra. Spec-
imens typically deep red, tentacles deep or-
ange with black tips in life and reddish
brown in alcohol. Ten tentacles almost
equal. Madreporite in dorsal mesentery
about one-half of distance from anterior end
of calcareous ring. Circular, bilobate mad-
reporite with a long narrow stone canal.
Two polian vesicles on ventral side of ring.
From left and right sides of cloaca arise two
respiratory trees; each splits into a dorsal
and ventral branch; in left side tree, ventral

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

200 um

200 pum

Bis. 3.

200 um

Ossicles of Cucumaria flamma. (a—d) dorsal skin ossicles, smooth perforated plates and smooth

buttons with two central holes; (e—f) Introvert with star-shaped and elongated perforated plates. All from holotype

(USNM E48072).

branch runs full length of body, the dorsal
branch three-quarters of length, in right
tree, ventral branch runs the full length of
body but the dorsal branch runs to middle
of body because of presence of gonad mass
(in mature specimens). Respiratory trees
usually light brown or yellowish, each with
a main trunk. Cloaca occupies approxi-
mately one-tenth of length of the body. Go-
nads arranged in two tufts of unbranched

tubules, attached by a mesentery to right-
mid dorsal side of the body.

Thin retractor muscles join to body wall
at a point level with anterior end of calcar-
eous ring. Plates of calcareous ring almost
of same size, with anterior projections, ra-
dials longer than interradials; anterior pro-
jection of interradials narrower than radials
(Fie 2).

Dorsal skin ossicles smooth perforated

VOLUME 112, NUMBER 4

200 um

100 um

100 um

100 um

Fig. 4. Ossicles of Cucumaria flamma. (a—d) tentacle ossicles; (e—g) ventral skin ossicles; (h) end plate of
podia. All from holotype (USNM E48072).

784

110°

30°

GULF OF
MEXICO

Mazatlan Bay
20°
Banderas Bay
(Manetas Islands)
ima +
(Pulpito Beach)

Zihuatanejo Bay

10°

°

110° 100

Fig. 5. Collection sites for Cucumaria flamma.

plates and smooth buttons with two central
holes (Fig. 3a—d). Introvert with star-shaped
(~340 pm) and elongated perforated plates
(200 zm) (Fig. 3e, f). Tentacle ossicles per-
forated, robust rods, three-branched, curved
or elongated, and scarce oval perforated
plates, with scalloped margins (Fig. 4a—d).
Ventral skin ossicles: abundant knobbed
buttons (110-150 wm) and smooth perfo-
rated plates (200 wm). Knobbed buttons
usually with four or sometimes six holes
(Fig. 4e—g). Curved perforated rods in po-
dia, end plates of 270—290 pm in diameter
(Fig. 4h). Skin ossicles seem to disappear
as the animal grows.

Etymology.—The specific epithet flamma
in Latin means ‘‘flame, blaze or fire’’. It is
here used as a noun in apposition and refers
to the color of the skin and the tentacle
crown of live specimens.

Distribution and habitat.—Cucumaria
flamma is known from the Mexican Central
Pacific, from Mazatlan, Sinaloa (23°15'39'N,
106°28'38’W) to Isla de la Plata, Ecuador
(1°16’N, 81°06’W), (Fig. 5). It ranges in
depth from 4 to 12 m. The majority of col-

90°

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

80° 70° 30°

Takoguilla Island

90° 80" 70°

lections are from less than 10 m. Usually
occurs on rocky substrates, in cavities and
holes.

Discussion

Cucumaria flamma is distributed from
Sinaloa, Mexico (in the mouth of the Gulf
of California) to Isla de la Plata, Ecuador.
As far as we know, eight species of the ge-
nus that could share this distribution area
are: C. frondosa japonica (Gunnerus,
1767); C. miniata (Brandt, 1835); C. piper-
ata (Stimpson, 1864); C. pseudocurata
Deichmann, 1938; C. salma Yingst, 1972;
C. crax Deichmann, 1941; C. pallida Kir-
kendale & Lambert, 1995 and C. vegae
Théel, 1886 (Deichmann 1938, 1941; Ber-
gen 1996; Lambert 1997). C. flamma is the
largest of the Cucumaria species on Mexi-
can coasts. It has the typical Cucumaria
shape (Fig. la, b). It can attain a length of
150 mm and longer, depending on its state
of relaxation.

Externally, C. flamma is similar to C.
miniata, but the arrangement of the tube

VOLUME 112, NUMBER 4

feet distinguishes them. C. miniata has five
bands of tube feet separated by broad ex-
panses of smooth skin, and occasional scat-
tered tube feet. C. flamma has very widely
dispersed tube feet not arranged in bands,
particularly abundant in the ventral region.
No other cucumariid exhibits the antero-
posterior dorsal groove characteristic of C.
flamma.

Cucumaria flamma is allied to C. pseu-
docurata in the shape of the smooth buttons
ossicles of the body wall with two central
holes, but those of C. flamma have broad
and knobbed edges. The buttons of C. pseu-
docurata are smaller (about one-third of di-
ameter) than those of C. flamma. C. pseu-
docurata also resembles C. flamma in the
shape of the calcareous ring, but the radial
plates in C. pseudocurata are slender, lon-
ger and with long anterior prolongations in
comparison to C. flamma, that have broad
radial plates with short anterior prolonga-
tions.

The knobbed plate ossicles present in the
body wall may be reminiscent of Cucumar-
ia lubrica H. L. Clark (formally Pseudoc-
nus lubricus); Lambert (1997, 1998a)
placed this species in the genus Pseudocnus
based on the distribution of tube feet, the
complex button ossicles and DNA evi-
dence.

Acknowledgments

Our thanks to Dr. Philip Lambert from
the Royal British Columbia Museum, to Dr.
David L. Pawson and Cynthia Ahearn,
Smithsonian Institution, Washington, D.C.
for the comments and suggestions provided;
to M. en C. Alicia de La Luz Duran-Gon-
zalez for helped in the techniques to get os-
sicles from the specimens; to Biol. Sara
Fuentes Soriano, academic technician of the
Laboratorio de Microscopia Electrénica,
Inst. de Biologia, UNAM for the electron
microscope photographs; to Dr. Antonio
Garcia Cubas and Dr. Sergio Licea Duran
for taking the photographs of the complete
specimen. Magdalena Alvarez Alpizar and

785

Jorge A. Ornelas Ibafiez arranged and pre-
pared the figures.

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lie Cucumariidae (Ordnung Dendrochirota).—

Mitteilungen aus dem Hamburgischen Zoolo-

gischen Museum und Institut 55:25-38.

. 1962. Bemerkungen Uber die Holothurien-

Familie Cucumariidae (Ordnung Dendrochiro-

ta).—Mitteilungen aus dem Hamburgischen

Zoologischen Museum und Institut 60:57—80.

. 1964. Bemerkungen Uber die Holothurien-

Familie Cucumariidae (Ordnung Dendrochiro-

ta).—Mitteilungen aus dem Hamburgischen

Zoologischen Museum und Institut 61:159—174.

. 1966. Bemerkungen Uber die Holothurien-

Familie Cucumariidae (Ordnung Dendrochiro-

ta).—Mitteilungen aus dem Hamburgischen

Zoologischen Museum und Institut 63:51—69.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

. 1971. Bemerkungen Uber die Holothurien-
Familie Cucumariidae (Ordnung Dendrochiro-
ta).—Mitteilungen aus dem Hamburgischen
Zoologischen Museum und Institut 67:29—51.

Pawson, D. 1970. The marine fauna of New Zealand:
sea cucumbers (Echinodermata: Holothuroi-
dea).—Bulletin of the New Zealand Department
of Scientific and Industrial Research 201:1—69.

. 1982. Holothuroidea. Pp. 813-818 in S. P

Parker, ed., Synopsis and classification of living

organisms, vol. 2. McGraw-Hill, New York,

1232 pp.

. & H. B. Fell. 1965. A revised classification
of the Dendrochirote Holothurians.—Brevioria
214:1-7.

Selenka, E. 1868. Nachtrag dazu Zeitschrift fuer
Wissenschaftliche Zoologie 18:109—-118.
Stimpson, W. 1864. Descriptions of new species of
marine invertebrata from Puget Sound, collect-
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ary Commission, A. H. Campbell, Esq., Com-
missioner.—Proceedings of the Academy of

Natural Sciences of Philadelphia 16:153—161.

Théel, H. 1886. Report on the Holothurioidea dredged
by the HMS Challenger during the years
18873-1876 Part Il—Report of the Scientific
Results of the Voyage of H.M.S. Challenger
1873-1876, Zoology 14:1—290.

Yingst, Josephine Y. 1972. A new species of the rock
dwelling dendrochirote holothurian from Cata-
lina Island—Bulletin of Southern California
Academy of Sciences 71(3):145—150.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(4):787-792. 1999.

A new species of rainfrog of the Eleutherodactylus cruentus group
from eastern Honduras (Amphibia: Anura: Leptodactylidae)

Jay M. Savage, James R. McCranie, and Larry David Wilson

(JMS) Department of Biology, PO. Box 249118, University of Miami,
Coral Gables, Florida 33124, U.S.A.; (JRM) 10770 SW 164th Street, Miami, Florida 33157-2933,
U.S.A.; (LDW) Department of Biology, Miami-Dade Community College, Kendall Campus,
Miami, Florida 33176, U.S.A.

Abstract.—A new species of Eleutherodactylus (E. operosus) of the E.
cruentus group is described from eastern Honduras. The new species differs
from the remaining species in this group by the following combination of
characters: heels with one small pustular tubercle; outer tarsus smooth; tuber-
cles absent on upper eyelids; discs of fingers II—-IV narrower than tympanum;
disc covers and pads on fingers HI—IV and all toes somewhat truncated; tym-
panum distinct in females; loreal region long; vomerine teeth present; dorsal
surfaces brown in life; groin mottled, without distinct spots; groin and anterior
and ventral surfaces of thighs brown in life; and heels pale copper in life. The
forests around the known locality for the new species are under heavy human
assault, even though the region is part of the Rio Platano Biosphere Reserve.

Frogs of the genus Eleutherodactylus
(s.l.) constitute the largest genus of verte-
brates with about 600 valid species cur-
rently recognized. The genus has an exten-
Sive geographic range from Arizona,
U.S.A., to southern Ecuador on the Pacific
slope and from Texas, USA, to Bolivia and
southern Brazil on the Atlantic slope, in-
cluding the West Indies.

Most mainland species belong to one of
two lineages (Lynch 1986, Savage 1987),
the Middle American clade (I; Craugastor)
or the South American clade (II; Eleuthero-
dactylus s.s.). These two groups are recog-
nized on the basis of distinctive synapo-
morphies in the jaw muscles (Lynch 1986,
Savage 1987) and karyotypes (DeWeese
1976, Savage 1987).

The majority of species found in North
and Central America belong to clade I, but
14 forms, mostly confined to lower Central
America, are representatives of clade II.
Within that clade, twelve Central American
species, characterized externally by having
strongly areolate ventral integument, no toe

webs, and toe III much shorter than toe V,
with the tip of the latter reaching the level
of the distal subarticular tubercle on toe IV,
may be referred to the Eleutherodactylus
martinicensis series (Lynch & Duellman
1997). Prior to the present report only one
member of this series, FE. ridens, was
known to range as far north as Honduras.
In July 1997, JRM and LDW collected a
single subadult female Eleutherodactylus
sharing the ventral skin texture and toe fea-
tures of the E. martinicensis series during
three nights of searching the environs of a
small stream in northeastern Olancho, Hon-
duras. This specimen differed significantly
from E. ridens in several characters and ap-
peared to represent an undescribed species.
Thus, in July—August 1998, JRM and LDW
returned to the same stream hoping to col-
lect additional material of this Eleuthero-
dactylus. Despite six nights of searching
along a broader area of the stream and sur-
rounding forest, JRM and LDW were able
to secure only one additional subadult fe-
male of this form. The second specimen

788

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Bie.

agrees in all details with the first specimen
and both differ significantly from all known
Central American species of the E. martin-
icensis series. Given the difficulty in col-
lecting these frogs, plus the fact that JRM
and LDW are extremely unlikely to ever
return to this rapidly deforested locality, we
describe the new taxon herein.

Materials and Methods

Measurements were made to the nearest
0.1 mm with dial calipers under a dissecting
microscope. Comparative material exam-
ined is listed in Appendix I. Comparative
data for E. museous was taken from Ibdafiez
et al. (1994). Abbreviations used are EL
(eye length), EN (anterior border of eye to
posterior edge of nostril; equals loreal
length), HL (head length; tip of snout to
angle of jaw), HW (greatest width of head),
SHL (shank length), SL (snout length; an-
terior border of eye to tip of snout), SVL
(snout-vent-length), and TPL (tympanum
length). Disc terminology follows that of
Savage (1987, 1997) and color codes are

Subadult female holotype of Eleutherodactylus operosus (USNM 530555), SVL 18.8 mm.

those of Smithe (1975-1981). Museum ab-
breviations follow those of Leviton et al.
(1985), except for CRE (Costa Rican Ex-
peditions, presently being catalogued into
the LACM collection).

Systematics

Eleutherodactylus operosus, new species
Pig.

Holotype.—National Museum of Natural
History (USNM) 530555, a subadult fe-
male, from near a small dam along a small
tributary of the Quebrada de Las Marias
(15°18’N, 85°21’W), about 12 airline km
NNE La Colonia, Departamento de Olan-
cho, Honduras, 680 m elev., collected 31
Jul 1998 by J. R. McCranie, K. L. Williams,
and L. D. Wilson. Original number LDW
11354.

Paratype.-—USNM 530556, a subadult
female with the same data as the holotype,
except. collected’ 31. Jul -1997 sby Menke
McCranie and L. D. Wilson.

Diagnosis.—Eleutherodactylus opero-

VOLUME 112, NUMBER 4

sus is referred to the E. cruentus group
(sensu Savage 1980) within the E. martin-
icensis series by having large digital discs,
non-triangular shaped disc pads, finger I
shorter than finger II, and lacking toe
webs. Eight valid Central American spe-
cies (E. altae, E. caryophyllaceus, E.
cruentus, E. moro, E. museous, E. pardal-
is, E. ridens, and E. taeniatus) belong to
this group, which is a subset of the large
(about 150 species) E. unistrigatus species
group (sensu Lynch & Duellman 1997).
Eleutherodactylus operosus differs from
both E. cruentus and E. museous (character
states for latter two species in parentheses)
by having the heels with one small pustular
tubercle (a distinct well-developed pointed
tubercle), the outer tarsus smooth (two to
four well-developed tubercles), no tuber-
cles on the upper eyelids (distinct well-de-
veloped pointed tubercles), the discs of fin-
gers IIJ-IV narrower than the tympanum
length (broader than tympanum length),
and the groin region lacking distinct spots
(usually one to several distinct yellow
spots in E. cruentus; a black spot in E. mu-
seous). Eleutherodactylus operosus differs
from E. altae and E. pardalis in having an
essentially brown ground color with darker
markings (nearly uniformly black), the
groin area mottled dark brown and pale
brown (groin marked with a large pale
spot; spot red in life in E. altae, silvery-
white in life in E. pardalis), and the ante-
rior thigh surface dark brown with some
pale brown mottling (marked with dark
vertical bars separating large pale areas of
the same color as spots in groin).
Eleutherodactylus operosus differs from E.
caryophyllaceus by lacking a superciliary
tubercle (enlarged pointed superciliary tu-
bercle located on free margin of upper eye-
lid) and by having the heel with one small
pustular tubercle (distinct well-developed
pointed tubercle). Eleutherodactylus op-
erosus is distinguished from E. moro by
having brown dorsal surfaces in life
(green), the heels pale copper in life (no
red on hind limbs), and the disc covers and

789

pads on fingers III-IV and all toes some-
what truncated (round). Eleutherodactylus
operosus is distinguished from E. ridens
by having the disc covers somewhat trun-
cate (round), the heel with one small pus-
tular tubercle (heels smooth to rugose), the
upper eyelids smooth or rugose (one to
several low tubercles), the groin and an-
terior and ventral surfaces of the thighs
brown in life (these surfaces some shade
of red in life), ventral surfaces of head and
body with sparse brown flecking from chin
to anterior portion of belly (ventral surfac-
es moderately to heavily flecked with
brown), and a long loreal region (EN/EL
>1.00 versus <0.90 in females). The new
form differs from E. taeniatus in having
the posterior surface of the thighs dark
brown with small distinct pale spots (es-
sentially uniformly pale brown), a smaller
tympanum (TPL/EL 0.30—0.31 versus
0.45—0.48 in females), and a longer loreal
region (EN/EL 1.04—1.09 versus 0.88—1.00
in females).

Description of holotype (stored in alco-
hol after formalin fixation).—A subadult
female with the following measurements
(percentages of SVL in parentheses): SVL
18.8 mm; HL 7.7 mm (41.0); HW 6.7 mm
Gao), Blet222) mmin(1h:7); Sibo3 4 mm
ChSel); EN°24 mm *@l2.8);eRPE: 0-7 ‘mm
(357);, SHE’ 11:30mm) (60:1)3;)and FL. 8:1
mm (43.1). Snout long, nearly rounded in
dorsal aspect, vertical with rounded upper
end in profile; top of head flat; canthus
rounded, distinct; loreal region concave;
upper lip not flared; nostrils directed lat-
erally, situated at point about two-thirds
distance between anterior border of eye
and tip of snout; cranial crests absent; su-
pratympanic fold weak, narrowly obscur-
ing upper edge of tympanum; tympanum
indistinct, located posterior to lower half
of eye, separated from eye by distance
about equal to tympanum length; upper
arm more slender than moderately slender
forearm; transverse dermal fold present on
upper surface of wrist; transverse dermal
fold absent on elbow; tubercles or dermal

790

ridge absent along posterior ventrolateral
edge of forearm; finger discs strongly ex-
panded (disc on finger III about 3.0 times
width of digit just proximal to disc); disc
covers on fingers somewhat truncated; disc
pads on fingers truncate; subarticular tu-
bercles on fingers round, conical to pun-
gent; supernumerary tubercles absent on
fingers; palmar tubercle low, bifid, larger
than thenar tubercle; 3—4 accessory palmar
tubercles present; thenar tubercle ovoid to
elongate, elevated, barely visible from
above; pollex not enlarged; relative length
of fingers I < II < IV < III; fingers un-
webbed, bearing weak lateral keels on fin-
gers II-IV; heels broadly overlapping
when hind limbs held at right angles to
body; weak vertical dermal fold present on
outer lateral edge of heel; heels bearing a
single, small pustular tubercle; tubercles or
dermal ridge absent along posterior ventro-
lateral edge of tarsus; inner tarsal fold ab-
sent, although one low elongated tubercle
present on one leg at about one-third
length of tarsus; subarticular tubercles on
toes round, globular; supernumerary tuber-
cles absent on toes; 8—10 very small plan-
tar tubercles present; inner metatarsal tu-
bercle elongate, elevated, visible from
above; outer metatarsal tubercle absent;
relative length of ttesI< I< HWI<V<
IV, disc on toe V extending to level of dis-
tal subarticular tubercle on toe IV; toe
discs moderately expanded (disc on toe IV
about 2.5 times width of digit just proxi-
mal to disc); disc covers on toes somewhat
truncated; disc pads on toes somewhat
truncated; toes unwebbed, bearing very
weak lateral keels; inguinal gland not ev-
ident; vent opening directed posteroven-
trally near upper level of thighs, skin be-
low vent granular; skin of dorsal surfaces
smooth to weakly granular, including that
of upper eyelids; distinct ridge extending
posteriorly from upper eyelids to level
above axilla, ridge broken posteriorly with
small laterally offset portion just posterior
to level of axilla; suprascapular fold ab-
sent; dorsolateral ridges absent; skin of

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

throat, chest, and ventral surface of thighs
smooth, that of belly coarsely areolate;
ventral disc absent; pupil horizontally el-
liptical; palpebral membrane translucent,
unpatterned; tongue ovoid, barely notched
posteriorly, free posteriorly for about one-
half of its length; vomerine teeth on very
low, ovoid ridges located posteromedially
to ovoid choanae, tooth patches separated
by distance greater than width of either
patch; maxillary teeth present.

Color in life: dorsal surfaces of head and
body Hair Brown (119A), with raised Ma-
hogany Red (132B) postocular ridge; dorsal
surface of forelimbs Hair Brown (119A);
dorsal surface of hind limbs Army Brown
(219B), with Sepia (219) crossbars; heels
pale copper; groin mottled Hair Brown
(119A) and Army Brown (219B); chin
transparent with small brown punctations;
belly colorless with small white puncta-
tions; iris yellowish brown on upper half,
bronze with brown reticulations on lower
half.

Color in alcohol: dorsal surfaces of head
and body brown with darker brown outlin-
ing postocular ridge; dorsal surfaces of
limbs pale brown with distinct, dark brown
crossbars; lateral surface of head barred
with pale brown and dark brown; posterior
surface of thighs dark brown, with distinct
pale brown spots; anterior surface of thighs
dark brown, with some pale brown mot-
tling; heel region very pale brown; ventral
surfaces of head, body, and thighs pale
cream-colored, with sparse brown flecking
from chin to anterior portion of belly, fleck-
ing more pronounced on thighs.

Variation in paratype.—The paratype
agrees well with the holotype in all mor-
phological characters and in color pattern,
except that the heel tubercles are not as well
developed and the vomerine tooth patches
are better developed than in the holotype.
The following measurements (percentages
of SVL in parentheses) were recorded: SVL
20.2 mm; HL 8.1 mm (40.1); HW 6.8 mm
(33.7); EL 2.6 mm) (12:9); SE) 3:33am
(18.8); EN 2.7 mm (13.4); TPL 0.8 mm

VOLUME 112, NUMBER 4

(4.0); sSHE-11.9 (58,9); and FL 8.6, mm
(42.6).

Natural history notes.—Both specimens
were taken at night from vegetation about
1.5 m above the ground along a trail near
where it crosses a small stream at 680 m
elevation. Although the specimens were
taken in different years, both were found at
nearly the identical spot along the trail. The
locality is in the Premontane Wet Forest
formation of Holdridge (1967). The vege-
tation in the immediate vicinity was in a
nearly pristine condition in 1997 and 1998.
However, the area alongside this stream was
completely deforested at 660 m, as were
most of the hillsides surrounding the forest
on both sides of the portion of the stream
where E. operosus was collected. Also,
considerably more deforestation was evi-
dent along this stream on JRM’s and
LDW’s second trip in 1998 than was evi-
dent in 1997. This extensive deforestation
is occurring even though the region is part
of the “‘protected’’ Rio Platano Biosphere
Reserve. Other species of Eleutherodacty-
lus collected in the environs of this trail and
stream were E. epochthidius, E. fitzingeri,
E. mimus, E. ridens, and Eleutherodactylus
sp. (E. rugulosus group). Males of E. fitzin-
geri and E. ridens could be located by their
advertisement call. No other species of
Eleutherodactylus were heard to call at this
locality.

Etymology.—The name operosus is a
Latin word meaning laborious or difficult
(Brown 1991). The name is used in refer-
ence to the difficulty experienced in efforts
to collect a series of these frogs.

Remarks.—Lynch & Duellman (1997)
referred all species in the E. martinicensis
series to two subdivisions, the E. diastema
group (now composed of seven species,
Savage 1997) and the E. unistrigatus group
(about 150 species). It is clear that the latter
is a composite that will be further divided
into additional species groups as knowledge
increases. The species placed here in the E.
cruentus group are the same as those in-
cluded by Savage (1980), with the exclu-

791

sion of E. cerasinus, transferred by Lynch
& Duellman (1997) to the E. conspicillatus
series, and the additions of E. museous, E.
operosus, and E. taeniatus. It remains to be
seen if the E. cruentus group, as utilized
here, is monophyletic, but for the present it
conveniently clusters all Central American
species of the E. martinicensis series, aside
from E. diastema and its allies, belonging
to the South American clade into a single

group.

Acknowledgments

Collecting and exportation permits were
provided by M. Acosta, A. Barahona, A. P.
Martinez, C. Romero, and T. L. Garcia of
COHDEFOR, Tegucigalpa. Valuable assis-
tance with these permits was provided by
M. R. Espinal. Field assistance was provid-
ed by D. Almendarez and K. L. Williams.
Comparative material was provided by B.
T. Clarke (BMNH), K. Beaman and D. Ki-
zirian (LACM), E T. Burbrink (LSUMZ),
G. Kohler (SMF), and A. G. Kluge and G.
Schneider (UMMZ).

Literature Cited

Brown, R. W. 1991. Composition of scientific words.
A manual of methods and a lexicon of materials
for the practice of logotechnics. Smithsonian In-
stitution Press, Washington, 882 pp.

DeWeese, J. E. 1976. The karyotype of middle Amer-
ican frogs of the genus Eleutherodactylus (An-
ura: Leptodactylidae); a case study of the sig-
nificance of the karyologic method. Unpub-
lished Ph.D. dissertation, The University of
Southern California, Los Angeles, 210 pp.

Holdridge, L. R. 1967. Life zone ecology. Revised edi-
tion. Tropical Science Center, San José, Costa
Rica, 206 pp.

Ibanez, D. R., C. A. Jaramillo, & EF A. Arosemena.
1994. A new species of Eleutherodactylus (An-
ura: Leptodactylidae) from Panama.—Amphib-
ia-Reptilia 15:337-—341.

Leviton, 2 Bio R. (oY Gibbs) Jn, E Heal, é&"C.E.
Dawson. 1985. Standards in herpetology and
ichthyology: part I. Standard symbolic codes for
institutional resource collections in herpetology
and ichthyology.—Copeia 1985:802—832.

Lynch, J. D. 1986. The definition of the middle Amer-
ican clade of Eleutherodactylus based on jaw

492

musculature (Amphibia: Leptodactylidae).—

Herpetologica 42:248-—258.

, & W. E. Duellman. 1997. Frogs of the genus
Eleutherodactylus (Leptodactylidae) in western
Ecuador: systematics, ecology, and biogeogra-
phy.—University of Kansas Natural History
Museum Special Publication 23:1—236.

Savage, J. M. 1980. A handlist with preliminary keys
to the herpetofauna of Costa Rica. Allan Han-
cock Foundation, The University of Southern
California, Los Angeles, 111 pp.

1987. Systematics and distribution of the

Mexican and Central American rainfrogs of the

Eleutherodactylus gollmeri group (Amphibia:

Leptodactylidae).—Fieldiana Zoology, New Se-

ries 33:1—57.

. 1997. A new species of rainfrog of the Eleu-
therodactylus diastema group from the Alta Tal-
amanca region of Costa Rica.—Amphibia-Rep-
tilia 18:241—247.

Smithe, E B. 1975-1981. Naturalist’s color guide. Part
I. Color guide. The American Museum of Nat-
ural History, New York, 182 color swatches.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Appendix I
Comparative Material Examined

Eleutherodactylus altae. Costa Rica—San José:
CRE 3181, 3212, 3380-81, 7048.

Eleutherodactylus caryophyllaceus. Costa Rica—
Cartago: CRE 7053, 7058 (2), 7060, 7075, 7095. San
José: CRE 7035, 7048.

Eleutherodactylus cruentus. Costa Rica—Alajuela:
CRE 693, 696, 767, 2929, 2932, 7014. Cartago: CRE
191, 2284. San José: CRE 490, 3181, 6467-68.

Eleutherodactylus moro. Costa Rica—San José:
CRE 765, UCR 6000.

Eleutherodactylus pardalis. Costa Rica—Puntare-
nas: CRE 916 (2), 919, 3182 (2), 3485, 3488, 7224,
LACM 119586. San José: CRE 7097.

Eleutherodactylus ridens. Honduras—Atlantida:
FMNH 236380, SMF 77632—33, USNM 514529-S0.
Colon: BMNH 1985.1232, 1985.1453, LSUMZ 33647,
UMMZ 89461, USNM 514551-54. Olancho: SMF
78781-82, USNM 343720—-22, 344791, 514555-—59,
530557-—60.

Eleutherodactylus taeniatus. Colombia—Choc6:
LACM:-50551;5 73208, 73212; 73222273

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(4):793-803. 1999.

A new species of arboreal viper (Serpentes: Viperidae: Atheris) from
Cameroon, Africa

Dwight P. Lawson

Department of Biology, The University of Texas at Arlington,
Arlington, Texas 76019-0498, U.S.A.

Abstract.—A distinctive new species of the arboreal viper genus Atheris (A.
broadleyi) is described from Cameroon, Africa. The new taxon inhabits moist
evergreen and semideciduous transition forest in southern Cameroon, Congo,
Central African Republic and Gabon. The new species previously has been
confused with the widespread A. squamigera, but is distinguished from this
species and other congeners by a combination of scale characteristics and a

unique color pattern.

In a recent review, Broadley (1998) com-
mented on an unusual color morph of Ath-
eris Squamigera from southern Cameroon.
Perret & Mertens (1957) had previously
mentioned the same population of Atheris
from Moloundou in the extreme southeast
of the country, and suggested that addition-
al material was needed to ascertain this
population’s taxonomic status. Both of
these descriptions agree with a series of
Atheris 1 obtained during a herpetological
survey of the proposed Lac Lobeké pro-
tected area of extreme southeastern Came-
roon. Comparison of this material with mu-
seum specimens from Cameroon, Demo-
cratic Republic of the Congo, Central Af-
rican Republic (CAR), topotypic A.
anisolepis Mocquard (1887) and the holo-
type of A. squamigera (Hallowell, 1856) re-
veals that the southern, inland Cameroon
population is distinct from all other Atheris
Species.

Methods

Specimens of the new Atheris were fixed
in 3.7% formalin within 24 hr of collection.
Specimens were transferred into 70% eth-
anol within two months after preservation.

Snout-vent length (SVL) and tail length
(TL) were measured to the nearest mm us-

ing a meter stick. Other standard measure-
ments were taken to the nearest 0.1 mm us-
ing dial calipers held under a dissecting mi-
croscope. Features of scalation were ex-
amined by holding preserved specimens
under a dissecting microscope. Scale ter-
minology generally follows Klauber
(1956). Suprarostrals and interorbitals are
used in keeping with Broadley (1998) for
other members of the genus. Interrictal
counts refer to scales across the back of the
head between the posterior supralabials.
Anterior body dorsal scale row counts were
made approximately one head-length pos-
terior to the head. Posterior body dorsal
scale row counts were made approximately
one head-length anterior to the vent. Color
descriptions in preservative and life are
based on a published standard (Smithe
1975). Institutional abbreviations refer to
Academy of Natural Sciences of Philadel-
phia (ANSP), American Museum of Natu-
ral History (AMNH), University of Texas
at Arlington Live Collection (DPL), Uni-
versity of Texas at Arlington Collection of
Vertebrates (UTA). A list of specimens ex-
amined is given in the Appendix.

Systematic Account

During the summer of 1994 I made her-
petological collections at various proposed

794

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1.
mm.

and existing protected areas throughout
Cameroon. Specimens of Atheris taken at
Lac Lobeké confirm long-standing impres-
sions that this population is distinct. I pro-
pose that this new snake be known as

Atheris broadleyi, new species
Figs. 1—2 and 5 (A)

Atheris squamigera squamigera.—Pertret &
Mertens, 1957:597

(A) Dorsal and, (B) left lateral aspect of Atheris broadleyi holotype, UTA R-37801, head length 29.0

Atheris squamigera ssp. Perret, 1961:137
Atheris squamigera.—Broadley, 1998: fig. 8

Holotype.—University of Texas at Ar-
lington Collection of Vertebrates (UTA) R-
37801, mature female collected vicinity Li-
pondji village, East Province, Cameroon
(2°23'05"N, 15°25’41”E), 1 Jun 1994 by D.
P. Lawson (original field number DPL
3608).

VOLUME 112, NUMBER 4

795

Fig. 2. Atheris broadleyi, adult male from the type-locality, ca. 525 mm SVL, DPL 3646. Photo by D. P.

Lawson.

Paratypes.—UTA R-37798-37800 and
44911-21, 10 adult and 4 juvenile females
from the type-locality, collected between
May and December 1994, UTA R-44957,
an adult female collected in the vicinity
Mbanjani village, East Province, Cameroon
(ca. 5 road km south of the type-locality),
7 Sep 1997, and ANSP 20334, an adult
male from Nola, Central African Republic
(3°32'22’"N, 16°02'51”E), collected 27 Oct
1934 by J. A. G. Rehn of the G. Vanderbilt
Africa Expedition.

Definition and diagnosis.—A species of
Atheris distinguished from all other mem-
bers of the genus by the following combi-

nation of characters: lateral scales without
serrated Keel; supralabials in contact with
suboculars (not separated by one or more
scale rows); 14—18 interrictals; interorbitals
keeled; rostral 3.5 to 4 times broader than
high; scales in loreal region smooth or with
only slight keel or knob; a dark postocular
stripe persisting in adults; maximum total
length of 765 mm or more; and a dark tail
tip in adult females.

Specifically, Atheris broadleyi appears to
be most similar to A. squamigera, but dif-
fers consistently by the combination of: a
distinctive and consistent dorsal color pat-
tern of citrine to greenish olive; checked

796

black, white and sky blue venter, and a dark
postocular stripe; more oculars (mode of 15
versus modes of 12—14, see Table 1); and,
greater length of 765 mm or more (versus
650 mm maximum for Cameroon A. squa-
migera). Additionally, A. broadleyi usually
has more ventrals (mean of 162 + 3.3 ver-
sus 157 + 3.6) and fewer subcaudals (mean
of 51 2 4.9 versus 52> =" 3-2) and more
infralabials (mode of 11 versus 10) than A.
squamigera.

Atheris broadleyi \acks serrated keels on
its lateral scales differentiating it from an
East African group comprised of A. cera-
tophora, A. desaixi, A. katagensis, A. nit-
schei, and A. rungweensis with pronounced
serrations on the lateral scales. Elongate su-
praocular scales forming a horn-like projec-
tion over the eye further distinguish Atheris
ceratophora. Similarly, Atheris broadleyi
lacks lanceolate or acuminate dorsal scales
and fused lateral scale rows, differentiating
it from the distinctive East African A. his-
pida and A. acuminata. Atheris broadleyi is
distinguished from A. chlorechis by having
fewer interrictals (14-18 versus 23 or
more) and fewer mid-body dorsal scale
rows (17-23 versus 25—36). In addition to
differences in color pattern, Atheris broad-
leyi is distinguished from A. anisolepis by
having completely keeled interorbitals
(smooth in A. anisolepis), supralabials in
contact with oculars (separated by a scale
row in A. anisolepis), and usually fewer su-
prarostrals.

Description of holotype.—Rostral flat-
tened, dorsal margin slightly concave me-
dially, 3.5 times broader than high, con-
tacting anteriormost supralabials, three un-
keeled suprarostrals and divided left nasal;
medial suprarostral the smallest, 1.3 times
wider than high, right and left suprarostrals
as wide as high; right and left nasals par-
tially divided above the nares, left nasal fur-
ther divided at anteriormost edge forming a
separate small scale; nares large, covering
approximately 30% of nasal, directed lat-
erally and slightly posteriorly; internasals 5,
strongly keeled, medial scale lying directly

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

above medial suprarostral but not separat-
ing two innermost internasals; interrictals
17; interorbitals 6, strongly keeled, those
not contacting oculars enlarged and irregu-
larly shaped; oculars 15/13, supra- and
postoculars keeled, terminating in blunt
knobs, suboculars and ventralmost preocu-
lar dorsoventrally compressed, knobbed but
not keeled, two dorsalmost preoculars fee-
bly knobbed, enlarged, extending anteriorly
into loreal region; oculars separated from
nasals by 6/5 smooth or feebly knobbed
scales loosely arranged in three rows of 1/
1, 1/2, 4/2; supralabials 11/10, 3—5 contact-
ing suboculars; infralabials 11/11, 5—10 on
each side feebly keeled, anteriormost in
contact at the midline, separating mental
from 5 pairs of chin shields; mental 2.3
times wider than deep; gulars strongly
keeled; ventrals 159; subcaudals 50, entire;
anal entire; dorsal scales about twice as
long as wide, becoming shorter posteriorly,
in 19-21-16 rows; keel on dorsals increas-
ing in height from base, declining rapidly
at apex, ending in slight knob; paraventrals
larger than other dorsals, keeled, angled
slightly ventrally.

Measurements (mm): SVL 550; tail
length 110; eye diameter 5.1; head length
29; head width 21.2; distance from anterior
margin of eye to nares 4.7 mm, to tip of
snout 6.8 mm.

Color in preservative (ethanol after for-
malin) has not changed appreciably from
life: dorsum of head uniformly citrine with
olive-yellow keels; sides of head spectrum
yellow extending onto lateral aspect of
neck; face straw yellow; a diffuse dark
stripe extending from the ventral posterior
margin of the eye caudad to the rictus, in-
corporating the last three supralabials and
posterior margin of the eighth, barely ex-
tending onto the last three infralabials; dor-
sum of body citrine anteriorly becoming
greenish olive posteriorly, broken by 29
black-bordered sulphur yellow crossbands,
crossbands on anterior half of body do not
meet on midline, becoming indistinct on
neck, posterior 15 crossbands conspicuous,

VOLUME 112, NUMBER 4

complete across the dorsum; tail with 9
moderately distinct, complete crossbands;
tail tip black; interstitial skin color varied,
corresponds to dorsal color pattern; throat
and neck white; venter sky blue, irregularly
suffused with white anteriorly, becoming
increasingly darker posteriorly; white
blotch on lateral portion of every third and/
or fourth (occasionally second) ventral
scale extending onto paraventral, blotches
become more pronounced as venter darkens
posteriorly, forming a checkered pattern;
ventral color extends onto paraventrals and
occasionally onto adjacent dorsal scale.

Variation.—The 15 paratypes closely re-
semble the holotype in color, pattern, and
scalation. Posterior portions of the venter in
some individuals Gncluding the only male
in the type series) are more uniformly dark
than that of the holotype, but always have
distinct white lateral blotches. Specimens
from the western extent of the range (see
below) may be less boldly patterned, and
are almost uniformly citrine/greenish olive
with barely discernable crossbands and
postocular stripe (DPL 5508). Juveniles less
than 300 mm SVL (UTA R-37798, 44913,
44918, 44920) have white tail tips, and
remnants of a pale tail tip are present in two
subadult specimens (UTA R-44915 and
44919). UTA R-37799 is the largest of the
series (768 mm total length), UTA R-37798
the smallest (247 mm total length). One to
three isolated scales are present in the in-
teroculabial region in five of the paratypes,
but do not completely separate oculars and
supralabials. Scales in the loreal region are
smooth in eight specimens and feebly
knobbed in seven.

The paratypes exhibit the following var-
iation (means and standard deviations in pa-
rentheses, modes in brackets, see Table 1):
ventrals 157-169 (162 + 3.3); subcaudals
45-59 [50]; midbody dorsal scale rows 17—
23 (20 + 1.6); anterior body dorsal scale
rows 16—21 [19]; posterior body dorsal
scale rows 13-18 [16]; ratio rostral width
to height 3.2—-5 (3.9 + 0.5); ratio medial
suprarostral width to height 1—2.2 (1.2 =

17

0.3); ratio right suprarostral width to height
0.65—1.3 (0.91 + 0.2); suprarostrals 3-7
[3]; internasals 3—5 [5]; interorbitals 3—8
[6]; oculars 12—16 [15]; total scales in lo-
real region 3—7 (5.3 + 1.2) [6]; supralabials
9-12 [10]; infralabials 9-12 [11]; ratio of
mental width to depth 1.3-2.5 (2 + 0.3);
chin shields 4—7 [5]; interrictals 14-18
[17]; ratio eye diameter to head length
0.15-—0.24 (0.18 + 0.02); ratio head length
to SVL 0.05—0.07 (0.06 + 0.01); ratio head
width to head length 0.64—0.84 (0.78 +
0.05); ratio eye-nares to head length 0.13-—
0.16 (0.15 + 0.01); ratio eye-snout to head
length 0.21—0.27 (0.24 + 0.01).

Distribution.—Atheris broadleyi is
known from southern Cameroon from the
Nyong River just southeast of Yaounde
Git hi 12, 3°27 38 N, DPE. 5508), south
through the Dja Forest Reserve (Broadley
1998) and east at least as far as the Sangha
region of the Central African Republic (Fig.
3). This distribution corresponds to a region
previously mapped for Cameroon as moist
evergreen and semideciduous forest (tran-
sition forest) comprised of Baillonella tox-
isperma and Gilbertiodendron dewevrei
(CENADEFOR 1985). This region also fea-
tures palm swamps and large scrub grass-
land clearings associated with seasonal high
water tables (D. Thomas, pers. comm.). The
distribution of Atheris broadleyi presum-
ably follows this forest type into northern
Gabon and Congo.

Atheris broadleyi was the most common
snake in the Lac Lobeké, Cameroon survey,
but does not appear in a slightly larger col-
lection from the vicinity of Oesso, Congo,
just across the Sangha River (Kate Jackson,
pers. comm.). Atheris broadleyi is abundant
near the type-locality. Local collectors ob-
tained 24 specimens in 48 hours, indicating
remarkable densities for a snake by Central
African forest standards.

Etymology.—The specific epithet is a
patronym for Donald G. Broadley in rec-
ognition of his considerable contributions to
the herpetology of Africa in general, and

798 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Nj 10° ss ° ° : ™° 3

igeria ~~.

bel Central

we

way :
YW African

Cameroon .

Republic

Yaounde Fi
Douala 2

Nola oN,
A . "aaa

Fernando v
Po

0°

Neem.

Equatorial i

Libreville
a

7 Lukolela
/
i aa
/
i
Brazzaville .
6 Atlantic is Kinshasa
-~./ Democratic
Republic of
the Congo
aT SRS 8°

Ocean

14 Angola 16° \ 18°

Fig. 3. Localities of Atheris broadleyi (triangles), A. squamigera (inverted triangles), and A. anisolepis
(squares) in west Central Africa. Outlined symbols indicate type-localities.

VOLUME 112, NUMBER 4

his recent clarifications of the genus Atheris
in particular.

Comparisons.—Broadley (1998) provid-
ed a key to the species of Atheris by which
A. broadleyi is readily diagnosed from all
members of the genus except A. anisolepis
and the widely distributed and highly vari-
able A. squamigera. Atheris broadleyi is
also distinctly allopatric from all but A.
squamigera. Although recognized in recent
treatments (e.g., McDairmid et al. 1999,
Broadley 1998, Trape & Roux-Estéve
1995), A. anisolepis is similar to A. squa-
migera and its validity remains uncertain
(Broadley 1998). Descriptive statistics and
differences among A. broadleyi, A. aniso-
lepis and A. squamigera are summarized in
Table 1. For the purpose of comparisons
with the new taxon, information for four
populations of A. squamigera is presented
separately in Table 1. Atheris squamigera is
widely distributed in forests from Senegal
to western Kenya (Spawls and Branch
1995). As evidenced by this species de-
scription and the intraspecific variability of
A. squamigera (Table 1), previous descrip-
tions of this taxon may have inadvertently
increased the reported variation in many
characters by combining distinct species
with A. squamigera.

With the exception of occasional speci-
mens which uniformly darken in preserva-
tive over time, color and pattern features of
Atheris in preservative closely resemble
those in life (pers. obs.). Atheris squami-
gera are most often apple green to turquoise
blue with yellow crossbands above and yel-
low, green or blue ventrally (Fig. 4,
Schmidt 1923, pers. obs.). Occasional spec-
imens of A. squamigera from the Came-
roon-Nigeria frontier (UTA R-44963,
44926, Stuck-Stirn 1979, Lawson 1993)
and Congo (AMNH 45940, 51840), and
specimens of A. anisolepis (AMNH 11898-
99) are uniform spectrum yellow sparsely
flecked with yellow-green spots. Atheris
squamigera from the Congo River above
Kinshasa (AMNH 45940, 45943, 45945,
51840) may be uniform yellow, turquoise

799

green, violet or a mottled combination of
yellow and violet. Unlike A. sguamigera, A.
broadleyi appears to be highly consistent in
color pattern.

South-central Cameroon is an area of
possible sympatry between Atheris broad-
leyi and A. squamigera for which material
is available. Specimens from Metet and Bi-
tye (AMNH 5254, 51841-—43) have a pro-
nounced, dark postorbital stripe and several
features of scutellation overlapping A.
broadleyi. I have referred the south-central
Cameroon material to Atheris squamigera
on the basis of dorsal color pattern and scu-
tellation. However, this area lies at the tran-
sition between the coastal moist evergreen
forest of A. squamigera and the inland tran-
sition forest of A. broadleyi, and may rep-
resent an integration zone between the taxa
(Fig. 3). Collections from other areas of
possible sympatry at the southern limit of
A. broadleyi are lacking.

In addition to differences in color and
scalation, Atheris broadleyi differs from
Cameroon A. squamigera in behavior and
reproduction. Both in the field and in cap-
tivity, A. broadleyi are more aggressive, ac-
tive and alert than A. squamigera, striking
readily and repeatedly with little provoca-
tion (pers. obs.). Neonate Atheris broadleyi
(Fig. 5 A) resemble the adults in coloration,
but have a pale colored tail tip that disap-
pears as they mature. This is in contrast to
A. squamigera from western Cameroon that
produce almost uniform charcoal-black off-
spring with pale green eyes and a pale tail
tip (Fig. 5 B). These neonates attain the ap-
ple green or turquoise blue adult coloration
with successive post-natal sheds. The pale
tail tip persists in female A. squamigera
from western Cameroon, but is lost in
males from this population as they mature.

A pair of Atheris broadleyi collected as
adults at the type-locality in 1994 and main-
tained in captivity in the United States have
reproduced twice. Courtship and copula-
tions were observed between 28 June and
11 July 1995, and 13 young (one deformed)
were born on 10 February 1996. Courtship

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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VOLUME 112, NUMBER 4

801

Fig. 4. Atheris squamigera, adult male from vicinity Nguti, Southwest Province, Cameroon, ca. 375 mm

SVL, DPL 5348. Photo by D. P. Lawson.

was observed on 13 July 1996, but no off-
spring were produced that year. Courtship
was observed again on 10 June 1997 and
continued for approximately one week.
Five live and three stillborn young were
produced on 17 February 1998. The male
was observed courting and trying to copu-
late approximately one week prior to par-
turition. Specimens of A. squamigera from
western Cameroon maintained under simi-
lar captive conditions to the A. broadleyi
above have been observed in courtship in
September—October and have given birth in
April (pers. obs.). Pitman (1974) reported
similar timing of reproduction for A. squa-
migera in Uganda.

Like its congeners, Atheris broadleyi is
primarily nocturnal (Spawls and Branch
1995, pers. obs.). Specimens have been
found as they were either active on the
ground during or following evening rains,

or coiled among vine tangles, sitting in ap-
parent ambush along arboreal rodent run-
ways (pers. obs.). The few direct observa-
tions of this species in the wild occurred at
the forest edge.

There is a preponderance of females in
samples of both Atheris broadleyi and A.
squamigera from Cameroon indicating a
strong collecting bias or skewed sex ratio
in the populations. Males of both species
are considerably smaller than females.
Males and females of A. broadleyi are
much longer and more robust than their A.
squamigera counterparts (Table 1, Figs. 2
and 4).

Acknowledgments

The manuscript benefitted from the com-
ments and assistance of J. A. Campbell and
R. L. Gutberlet. P. Ustach provided the

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. (A) Neonate Atheris broadleyi born to parents from the type-locality; (B) Neonate A. squamigera
from wild-caught gravid female DPL 5318, Banyang-Mbo Wildlife Sanctuary, Southwest Province, Cameroon.
Photos by J. A. Campbell.

VOLUME 112, NUMBER 4

drawing of the holotype, and M. Fost doc-
umented captive behavior and reproduction.
I thank J. A. Powell and M. D. Powell for
their friendship and logistic support in
Cameroon, and P. Elkan and J. Ako for their
help in the field. Field work was funded by
the Wildlife Conservation Society through
a grant from the U.S. Agency for Interna-
tional Development. Cameroon permits
were issued by the Ministry of Environ-
ment and Forest. I also thank L. Ford
(AMNH) and T. Daeschler (ANSP) for the
loan of specimens examined in this study.

Literature Cited

Broadley, D. G. 1998. A review of the genus Atheris
Cope (Serpentes: Viperidae), with the descrip-
tion of a new species from Uganda.—Herpeto-
logical Journal 8:117—135.

Hallowell, E. 1856. Descriptions of new reptiles from
Guinea.—Proceedings of the Academy of Nat-
ural Sciences of Philadelphia 1854:193—194.

Klauber, L. M. 1956. Rattlesnakes their habits, life his-
tories, and influence on mankind, vol. 1. Uni-
versity of California Press, Berkely, 740 pp.

Lawson, D. P. 1993. The reptiles and amphibians of
the Korup National Park Project, Cameroon.—
Herpetological Natural History 1(2):27—90.

McDiarmid, R. W., J. A. Campbell, & T. A. Touré.
1999. Snake species of the world. A taxonomic
and geographic reference, vol. 1. Herpetolo-
gists’ League, Washington, D.C. (in press).

Mocquard, M. F 1887. Sur les ophidiens rapportés du
Congo par la mission de Brazza.—Bulletin de
la Société Philomatique de Paris 11:62—92.

National Centre for Forestry Development (CENA-
DEFOR). 1985. Ecological map of the vegeta-
tion cover of Cameroon (Based on landsat im-
ages).

Perret, J.-L. 1961. Etudes herpétologiques Africains

803

I1I.—Bulletin de la Société Neuchatel des Sci-
ences Naturelles 84:133—138.

Perret, J.-L., & R. Mertens. 1957. Etude d’une collec-
tion herpétologique faite au Cameroun de 1952
a 1955.—Bulletin de l'Institut Francais
d’Afrique Noire, Serie A, Sciences Naturelles
19:548-601.

Pitman, C. R. S. 1974. A guide to the snakes of Ugan-
da. Revised edition. Wheldon and Wesley, Ltd.,
290 pp.

Schmidt, K. P. 1923. Contributions to the herpetology
of the Belgian Congo based on the collection of
the American Museum Congo expedition,
1909-1915 Part I1—Snakes.—Bulletin of the
American Museum of Natural History 49:1-—
146.

Smithe, FE B. 1975. Naturalist’s color guide. American
Museum of Natural History, 14 pp.

Stucki-Stirn, M. C. 1979. Snake report 721. Herpeto-
Verlag, Teuffenthal, Switzerland, 650 pp.

Spawls, S., & W. R. Branch. 1995. The dangerous
snakes of Africa. Ralph Curtis Publishing, Inc.,
Sanibel Island, Florida, 192 pp.

Trape, J. EF, & R. Roux-Estéve. 1995. Les serpents du
Congo: liste commentée et clé de détermina-
tion.—Journal of African Zoology 109:31—50.

Appendix
Specimens Examined

Atheris anisolepis (3): Democratic Republic Of The
Congo: Banana, Lower Congo; AMNH 11898-99. Ku-
kolela, Congo R.; AMNH 45941.

Atheris broadleyi (5): Cameroon: East Province;
DPL 3646, 3657, 5300, 5302. Center Province; DPL
5508.

Atheris squamigera (28): Cameroon: Bitye; AMNH
5254. Metet; AMNH 51841—43. Southwest Province;
UTA R-31346, 44922—-930, 44963, 45003—04. Demo-
cratic Republic Of The Congo: Lukolela, Congo R.;
AMNH 45940, 45943, 45945, 51840. Gabon: ANSP
27253. Near the River Gaboon, Guinea; ANSP 6949
Holotype. Kenya: Kakamega; UTA R-12782, 13037,
19510, 26612, 32074.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

112(4):804—812. 1999.

Taxonomic notes on hummingbirds (Aves: Trochilidae)
2. Popelairia letitiae (Bourcier & Mulsant, 1852) is a valid species

Gary R. Graves

Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560, U.S.A.

Abstract.—Popelairia letitiae (Bourcier & Mulsant, 1852), known from two
specimens supposedly from Bolivia, appears to be a valid species. Analysis of
plumage color and morphometrics indicates that P. letitiae does not represent
an immature plumage or geographic variant of Discosura longicauda, although
this species and P. letitiae may be sister taxa. No credible evidence for a hybrid
origin of P. letitiae was discovered. “‘Letitia’s Coquette”? is proposed as the

common English name for P. letitiae.

Known from two specimens of vague
provenance, Popelairia letitiae (Bourcier &
Mulsant, 1852), has been neither observed
nor collected during the 20th century. This
fact alone is reason enough to question the
taxonomic validity of Letitia’s Coquette, as
many nominal trochiline taxa of compara-
ble rarity have proven to be hybrids (Meyer
de Schauensee 1947; Graves 1996, 1997a,
1997b, 1998a). Nonetheless, the systematic
status of P. letitiae remains unchallenged
(Mulsant & Verreaux 1876, Elliot 1878,
Boucard 1893, Cory 1918, Simon 1921, Pe-
ters 1945, Morony et al. 1975, Sibley &
Monroe 1990), although Salvin (1892) may
have been the last taxonomic authority to
critically examine the type specimen. The
relevance of this issue was brought to the
forefront by the inclusion of P. letitiae in a
recent survey of threatened avian species
(Collar et al. 1992). Here I offer an apprais-
al of the systematic status of P. letitiae.

Despite its current placement in the ge-
nus Popelairia (Peters 1945, Morony et al.
1975, Sibley & Monroe 1990), letitiae more
closely resembles Discosura longicauda in
plumage color (Elliot 1878). Generic limits
in the Trochilidae are based primarily on
male plumage traits (Taylor 1909). This has
resulted in a proliferation of genera (n =
109), more than 1/4 of which are monospe-

cific (Sibley & Monroe 1990). Zimmer
(1950) advocated merging all band-rumped
coquettes and thorntails in a single genus,
Lophornis. Even under a narrow interpre-
tation of generic limits, Popelairia would
have been merged with Discosura if not for
the racket-tipped rectrices of the latter (El-
liot 1878). In that eventuality, Popelairia
Reichenbach 1854 would become a junior
synonym of Discosura Bonaparte 1850. In
order to avoid nomenclatural confusion,
however, I use the binomial, Popelairia le-
titiae, throughout this paper.

Methods

The type of Popelairia letitiae (BMNH
1888.7.25.83 in The Natural History Mu-
seum, formerly British Museum of Natural
History) was obtained from John Gould,
who procured it from Bourcier (Gould
1858, Warren 1966). A second specimen in
the American Museum of Natural History
(AMNH 38060) was part of the Daniel Gi-
raud Elliot collection cataloged in 1888
(fide Paul Sweet). Both specimens appear
to be males in definitive plumage as judged
by their brilliant gorgets, crowns, elongated
rectrices, and unstriated maxillary ram-
phothecae (Figs. 1-3). I compared them
with male specimens of all species of hum-

VOLUME 112, NUMBER 4

a, i iN SS

yy, BY,

Figs I.
Mulsant, 1852).

mingbirds deposited in The Natural History
Museum and the American Museum of
Natural History. It was not possible to com-
pare directly the two specimens. However,
I compared photographs of the AMNH
specimen of P. letitiae with the type spec-
imen (BMNH), and vice versa. Because
previous characterizations of P. letitiae
were brief and somewhat contradictory, I
provide a more detailed description in Ap-
pendix 1. Measurements of wing chord, bill

805

Multiple exposures of the type specimen (BMNH 1888.7.25.83) of Popelairia letitiae (Bourcier &

length (from anterior extension of feathers),
and rectrix length (from point of insertion
of central rectrices to the tip of each rectrix)
were made with digital calipers and round-
ed to the nearest 0.1 mm. Rectrices are
numbered from innermost (R1) to outer-
most (R5) (Table 1).

I evaluated the color of the plumage
(forecrown, back above white band, rump
below white band, upper throat, lower side
of throat, lower breast along midline) with

806 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

-i

PE’ Mus ae

fe ee ree og

At

Fig. 2. Dorsal and ventral views of Elliot’s specimen of Popelairia letitiae (AMNH 38060).

Fig. 3. Lateral view of Elliot’s specimen of Popelairia letitiae (AMNH 38060).

VOLUME 112, NUMBER 4

807

Table 1.—Measurements (mm) of the two known specimens of Popelairia letitiae and male specimens of
Discosura longicauda (range; mean + standard deviation) in definitive plumage.

BMNH AMNH Discosura longicauda
1888.7.25.83 38060 n = 158
Wing Chord 3728 38.0 42.1—47°5 (45.2 = 1:6)
Bill length 10.5 10.9 10.2-12.4 (11.4 + 0.6)
Rectrix | 14.8 Missing 15.2—19.2 (16.7 + 0.9)
Rectrix 2 Loy 16.1 24.3-28.2 (26.0 + 1.1)
Rectrix 3 26.7 DDL, 29) 2-35:.6 G20 = F.2)
Rectrix 4 30.7 293 923-3) (39.0) = 1-4)
Rectrix 5 36.2 38.9 49.9-56.0 (52.5 + 2.0)

4 Bahia, Brazil (n = 4), Brazil (n = 2), ‘‘Cayenne’’

locality unknown (n = 1).
> Tip (~0.2—0.4 mm) missing.

a calibrated colorimeter (CR-221 Chroma
Meter, Minolta Corporation) equipped with
a 3.0 mm aperture. The measuring head of
the CP-221 uses 45° circumferential illu-
mination. Light from the pulsed xenon arc
lamp is projected onto the specimen surface
by optical fibers arranged in a circle around
the measurement axis to provide diffuse,
even lighting over the measuring area. Only
light reflected perpendicular to the speci-
men surface is collected for color analysis.
Colorimetric data from iridescent gorget
feathers are acutely dependent on the angle
of measurement, the curvature of the gorget
surface in museum skins, and the degree of
pressure applied to the plumage surface by
the Chroma Meter aperture. In order to re-
duce measurement variation, I held the ap-
erture flush with the plumage surface with-
out depressing the plumage surface. The
default setting for the CR-221 Chroma Me-
ter displays mean values derived from three
sequential, in situ measurements. I repeated
this procedure three times for each area of
plumage, removing the aperture between
trials. Thus, each datum summarized in Ta-
ble 2 represents the mean of three indepen-
dent measurements, each of which repre-
sents the average of three sequential default
measurements.

Colorimetric characters were described in
terms of opponent-color coordinates (L, a,
b) (Hunter & Harold 1987). This system is

(n = 3), ‘‘British Guiana’’ (n = 1), Guyana (n = 4),

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
‘““greenness’’ 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 O 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 Z value will be. Vi-
sual systems in hummingbirds (e.g., Gold-
smith & Goldsmith 1979) differ signifi-
cantly from those of humans. The relevance
of opponent color coordinates to colors per-
ceived by hummingbirds is unknown.

I considered four hypotheses: Popelairia
letitiae represents (1) an immature plumage
of Discosura longicauda; (2) a geographic
variant of D. longicauda; (3) a hybrid; or
(4) a valid species. In investigating the pos-
sibility of hybridization, I considered the
band-rumped coquettes and thorntails that
occur in South America (1.e., Lophornis or-
natus, L. gouldii, L. magnificus, L. delattrei,
L. stictolophus, L. chalybeus, L. pavoninus,
Popelairia popelairii, P. langsdorfi, P.

808 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—ZL a b opponent color coordinates for plumage characters of the two known specimens of Popelairia

letitiae: L = lightness; a/—a = red/green; b/—b = yellow/blue.

BMNH AMNH
BMNH 1888.7.25.83 38060
a b EE a b
5 | 12:0 228). —4.0 1333
4.3 15.0 21.4 S20 10.5
B72 20.1 19.6 8.5 11.4
= Oe 4.5 16.8 —3.4 8.9
= 9.0 13.4 =05 6.7
Deal 15.8 PLES) 3.9 10.7

Plumage character Color L
Forecrown golden-green io
Lower back above band coppery-bronze 24.5
Rump below band coppery-bronze Pray |
Upper throat golden-green 11.6
Lower throat (side) golden-green 3h)
Lower breast (midline) bronze-green 25.6

4 General color observed when specimen is held in a position that yields the greatest apparent brilliance.

conversii, Discosura longicauda) as poten-
tial parental species (taxonomy of Sibley &
Monroe 1990). Fewer than half (23 of 55)
of the possible pairwise combinations of
the aforementioned species actually occur
in nature (i.e., species sympatric at the res-
olution of 1° X 1° latitude-longitude
blocks). Unless otherwise noted, assess-
ments of plumage characters refer to those
of males in definitive plumage. Assump-
tions and methods of hybrid diagnosis fol-
low Graves (1990) and Graves & Zusi
(1990).

Results

Immature plumage or geographic variant
of Discosura longicauda?—-A review of
plumage and mensural characters demon-
strates that Popelairia letitiae is not an im-
mature of Discosura longicauda. Immature
males of D. longicauda possess rounded
rectrices (USNM 328627, AMNH 46737)
that are replaced in subsequent molts by
sharply attenuated (R2—R4) and _ racket-
tipped (R5) rectrices. The outer rectrices
(R4—R5) of Popelairia letitiae are sharply
attenuated and lack rackets. P. letitiae also
differs from D. longicauda in lacking a
black chin spot, in possessing a yellowish-
brown mandibular ramphotheca (black in
D. longicauda), coppery-bronze back plum-
age (green in D. longicauda), white rump
band (buff in sub-definitive plumages of
Discosura longicauda), coppery-bronze
rump (green in definitive plumages, pur-

plish-black in sub-definitive plumages of D.
longicauda), and dull bronze-green lower
breast and abdomen (spangled with irides-
cent golden-bronze disks in both definitive
and sub-definitive plumages of D. longicau-
da). Finally, bill length is similar in P. le-
titiae and D. longicauda, but the wing and
rectrices are substantially shorter in P. le-
titiae (Table 1). The qualitative differences
between P. letitiae and D. longicauda far
exceed the magnitude of geographic varia-
tion exhibited within species of Popelairia
and Lophornis, and approximate the level
of morphological divergence observed
among the largely allopatric rufous-crested
species of Lophornis (Zimmer 1950).
Hybrid?—I failed to discover convincing
evidence for a hybrid origin of Popelairia
letitiae. Among the pool of potential paren-
tal species, the pale mandibular rampho-
theca (possibly orange in life) of P. letitiae
is shared with the rufous-crested species of
Lophornis (ornatus, gouldii, magnificus,
delattrei, stictolophus). These same species
also exhibit varying amounts of rufous in
the gorget. Because rufous pigmentation
appears to be inherited in a codominant
fashion in hummingbird hybrids (see Banks
& Johnson 1961, Graves & Newfield 1996),
I would expect Lophornis hybrids in defin-
itive plumage to exhibit traces of ochra-
ceous or rufous pigments in crown and gor-
get feathers. However, none were found in
P. letitiae (10X magnification). This force-
fully suggests that P. letitiae shares no im-

VOLUME 112, NUMBER 4

mediate genealogical relationship with the
rufous-crested species of Lophornis. Like-
wise, I could see no manifestation of a co-
ronal apterium (blue in life) present in L.
chalybeus (Ruschi 1962) or the elongated
and spectacularly spotted auriculars of L.
pavoninus in either specimen of P. /etitiae.
Further, none of the pairwise species com-
binations drawn from the subset consisting
of Discosura longicauda and Popelairia
(popelairii, langsdorfii, conversii) could
have produced the suite of characters ex-
hibited by P. letitiae (e.g., pale mandibular
ramphotheca, coppery-bronze rump).

Additional evidence arguing against the
hybrid hypothesis is provided by feather
length and shape, both of which appear to
be controlled polygenically in hummingbird
hybrids (Banks & Johnson 1961, Graves
1990). Lophornis ornatus, L. gouldii, L.
magnificus, L. delattrei, and L. stictolophus
possess elongated crests (>11 mm), and L.
chalybeus and L. gouldii have lateral gorget
feathers that exceed 15 mm in length.
Crown (4.3—5.5 mm) and lateral gorget
feathers (6.8-7.0 mm) of P. letitiae are
rounded, similar in size and shape to those
of Discosura longicauda. These data pro-
vide further grounds for excluding Lophor-
nis species from the pool of potential pa-
rental species. All combinations of species
drawn from the subset of thorntails (Pope-
lairia popelairii, P. langsdorfii, P. conver-
sii) and Discosura longicauda can again be
eliminated from consideration because their
outer rectrices are substantially longer than
those of P. letitiae.

The two specimens of Popelairia letitiae
are similar in size and shape (Table 1).
Wing length differs by 0.5%, whereas the
difference in rectrix lengths vary from 4.7%
(R4) to 22.4% (R2). These values fall with-
in the normal range of variation found
among museum samples of trochiline hum-
mingbirds (e.g., Graves 1996, 1997a,
1998c). Plumage pattern and color are near-
ly identical, agreeing in such minor char-
acters as tibial feathering and undertail co-
verts (Table 2, Appendix 1). These obser-

809

vations are consistent with the hypothesis
that P. letitiae is a valid species.

In conclusion, analysis of plumage and
size characters indicates that Popelairia le-
titiae does not represent an immature plum-
age or geographic variant of Discosura lon-
gicauda, although the close resemblance of
the two suggests a sister species relation-
ship. As noted in the introduction, the very
rarity of P. letitiae in museum collections
raises the specter of hybridization. How-
ever, based on known patterns of phenotyp-
ic inheritance in trochiline hybrids (Banks
& Johnson 1961; Graves 1990, 1998c,
1999; Graves & Zusi 1990) and the char-
acteristics of phenotypic variants (e.g.,
Graves 1998b), the possibility that P. leti-
tiae represents a hybrid seems remote. Bar-
ring discovery of contradictory data, P. le-
titiae should be regarded a valid species.

Geographic origin.—Both specimens of
Popelairia letitiae were thought to have
been collected in “Bolivia”? (Bourcier &
Mulsant 1852, Elliot 1878), and Remsen &
Traylor (1989) suggested “northeastern Bo-
livia” as a possible site. The purveyor of
the type specimen is unknown (Bourcier &
Mulsant 1852), whereas ‘‘Verreaux’’ was
listed as the collector of the AMNH speci-
men. Both are relaxed taxidermy mounts
prepared in a similar style, perhaps by the
Verreaux brothers, who operated a thriving
import/export business in natural history
specimens in Paris during the middle de-
cades of the 19th century.

Collecting localities inscribed on labels
of 19th century hummingbird specimens
are frequently unreliable (Berlioz & Jouan-
in 1944). For example, in the same paper
in which Popelairia letitiae was described,
Bourcier & Mulsant (1852) reported the
type locality of Ramphodon dohrnii as “‘la
République de l’Equateur,” although this
Species is apparently restricted to the Atlan-
tic coastal forest of Brazil. Consequently, it
would be unwise to confine a contemporary
search for P. letitiae to Bolivia.

Common English name.—Bourcier &
Mulsant (1852:144) dedicated Popelairia

810

letitiae to the “‘jeune enfant de M™ la mar-
quise Delgallo, fille de l’un des ornitholo-
gistes les plus célébres de |’Europe, M. Le
prince Charles Bonaparte.”’ The authors of-
fered no common name and Gould (1858)
referred to the taxon simply as “Letitia” in
his Monograph of the Trochilidae. Sixty
years later, Cory (1918) proposed “‘Letitia’s
Thorn-bill.”” The group name “‘thornbill”’ is
now restricted to short-billed species in two
related Andean genera, Chalcostigma and
Ramphomicron (Sibley & Monroe 1990).
No other common English name was used
in taxonomic literature until Meyer de
Schaunsee (1966) coined a new name,
‘““Coppery Thorntail,’’ which has been used
sporadically since then (e.g., Collar et al.
1992). Meyer de Schauensee’s name was
inappropriate because other species in the
complex possess “‘coppery”’ plumage, and
because Popelairia letitiae does not possess
a “‘thorntail’’ on par with those of the so-
called thorntails (Popelairia popelairii, P.
langsdorfii, P. conversii). I recommend
““coquette’’ as a group name for the small
band-rumped species currently placed in the
genera, Lophornis, Popelairia, and Disco-
sura (Sibley & Monroe 1990), and the com-
mon English name, “‘Letitia’s Coquette,”’
for Popelairia letitiae.

Acknowledgments

I thank Robert Pr¥s-Jones, Michael Wal-
ters, Mark Adams, Don Smith, and Frank
Steinheimer (The Natural History Museum,
Tring) and George Barrowclough, Mary
LeCroy, and Paul Sweet (American Muse-
um of Natural History, New York) for ac-
cess to collections in their care. Many
thanks are owed to Leslie Overstreet
(Smithsonian Institution Libraries) for bib-
liographic assistance, and to Don Hurlbert
(Smithsonian Photographic Services) for
help with the color plate. Travel was sup-
ported by the Department of Vertebrate Zo-
ology, Research Opportunities Fund, and
the Alexander Wetmore Fund, Smithsonian
Institution. Finally, I thank Richard C.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Banks and Richard L. Zusi for comments
on the manuscript.

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Appendix 1

Description of male Popelairia letitiae in definitive
plumage based on the two known specimens (type,
BMNH 1888.7.25.83; AMNH 38060). Characteriza-
tion of structural colors is unusually subjective as color
seen by the observer varies according to the angle of
inspection and direction of light. Color descriptions
were made under natural light.

Forecrown and crown (to a line drawn behind the
eyes) are brilliant golden-green. The crown color
blends smoothly into dark bronze-green on the hind-
neck and back. Crown feathers are of moderate length
(4.3—5.5 mm long) and rounded. The mantle emits a
coppery-bronze iridescence when viewed in direct
light. Wing coverts are the same color as back plum-
age. A narrow white band crosses the lower back.
Band feathers are gray, broadly tipped with silky white
barbs (especially apparent at the sides). The white
band is bordered posteriorly by a coppery-bronze (cop-
pery-purple or coppery-red at some angles) rump,
which in turn is bordered by bronze-green uppertail
coverts.

The chin to upper breast is brilliant golden-green
(about the same as crown; see Table 2), with coppery-
gold reflections at the sides of the throat (when viewed
head-on). Iridescent terminal disks are bordered prox-
imally by a narrow subterminal bronze-green zone, a
broader band of dull white (obscured by imbricated
feather tips), and finally by gray basal barbs. Obscured
portions of gorget feathers become progressively gray-

812

er toward the sides of the throat; the outer vane of
lateral gorget feathers is gray below the iridescent disk.
Lateral gorget feathers are of moderate length (6.8—7.0
mm), the iridescent disks are slightly wider (~2.8—3.1
mm) than long (~2.3—2.4 mm). Green terminal disks
are reduced or absent along the posterior border of the
gorget, producing a mottled green and white pectoral
band. The breast below the pectoral band is bronze-
green along the midline. This area is burnished with
coppery-gold (AMNH specimen) and a few spangles
of coppery-red immediately below the pectoral band.
Feathers of the lower belly and sides are broadly
tipped with buffy-white and grayish-white barbs. Vent
feathers are dark gray, tipped with white. The undertail
coverts are dark green with gray bases and rufous tips.
Tibial feathers are of moderate length (reaching about
half way to the base of the hallux), dark gray, tipped
with a mixture of white and cinnamomeous barbs.
The outermost rectrix (R5) is brownish-black with
a bluish sheen on the medial vane. A pale stripe bor-
dering the rachis becomes wider and buffier near the
base of the shaft. Dorsal rachis color is pale cinna-
momeous, becoming browner distally. The inner rec-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

trices (R2—R4) are similar in color and pattern. The
outer vanes of R4 & R3 in the AMNH specimen are
faintly glossed with bronzy-green. Both vanes of R2
in the AMNH specimen are glossed with bronze-green
(less pronounced in the type). There is a small V-
shaped buffy spot at the tip of R2 in both specimens.
The innermost pair of rectrices is absent in the AMNH
specimen. Those of the BMNH specimen are bronze-
green, broadly tipped with black, and faintly marked
with terminal V-shaped cinnamomeous spot. From be-
low, the medial vanes are bluish-black (R3—R5); the
rachises are white, becoming very pale buffy-white
proximally. The remiges, which lack emarginations or
markedly thickened rachises, are black with faint pur-
ple and bronze glossing under strong light.

The maxillary ramphotheca is black becoming dark
brown at the nares. The nares are completely obscured
by adpressed feathers. The mandibular ramphotheca is
brownish-yellow (slightly darker in the type) becoming
dark brown about half way to the bill tip. Scutes of
toes and tarsometatarsus are medium brown—less
heavily melanized than in Popelairia langsdorfii or
Discosura longicauda.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
112(4):813-856. 1999.

Identification and distribution of cotton rats, genus Sigmodon
(Muridae: Sigmodontinae), of Nayarit, México

Michael D. Carleton, Robert D. Fisher, and Alfred L. Gardner

(MDC) Division of Mammals, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560, U.S.A.; (RDE ALG) Patuxent Wildlife Research Center, Biological

Resources Division, U.S. Geological Survey, National Museum of Natural History,
Washington, D.C. 20560, U.S.A.

Abstract.—Morphological, chromosomal, distributional, and ecological data
are presented for three species of Sigmodon (S. alleni, S. arizonae, and S.
mascotensis) from the state of Nayarit, México. The species were collected in
all possible pairwise combinations of sympatry, including the first record of
such documented for S. arizonae and S. mascotensis. Emphasis is devoted to
the discrimination of S. arizonae and S. mascotensis, from each other and from
typical S. hispidus, using qualitative features of the skin and skull and mor-
phometric analyses of craniodental measurements. Based on these results and
examination of type specimens, additional synonyms of S. mascotensis are
identified, with reassignment of two forms, tonalensis Bailey (1902) and ob-
velatus Russell (1952), currently mistaken as subspecies of S. hispidus. Sig-
modon mascotensis emerges as a species distributed from southern Nayarit and
Zacatecas to extreme western Chiapas, where it inhabits deciduous or semi-
deciduous tropical vegetation having a pronounced dry season. These reallo-
cations and other reidentifications remove any documentation for S. hispidus
along the entire Pacific versant of México.

A useful form of research communica-
tion that sees less application today is the
brief expeditionary account or short faunal
note. Aside from the practical enhance-
ments in knowledge of a taxon’s distribu-
tion and habitat, such reports offer the ad-
vantage of bringing regionally focussed no-
menclatural clarity to complex taxonomic
problems that seem incomprehensible over
a broader geographic scale. One recalls that
the prolific literature appearing over the
past two decades on the Peromyscus boylii
group emanated from Hooper’s (1955) im-
memorable commentary in ‘‘Notes on
Mammals of Western México,”’ in which he
recorded the sympatric occurrence of vari-
ous “‘morphological types”’ of boylii at sev-
eral collecting localities in Jalisco, Nayarit,
and Sinaloa (see systematic reviews by
Carleton 1989, and Bradley et al. 1996).

The regional focus here is Nayarit, Mé-

xico, and the taxon of interest is the genus
Sigmodon, the ecologically abundant cotton
rats that inhabit open landscapes from the
southern United States, through México and
Middle America, to northern South Amer-
ica (Hall 1981, Voss 1992). Situated along
coastal westcentral México, the state of
Nayarit encloses a varied topography and
diverse natural environments, a biogeo-
graphical setting that has proven pivotal for
illuminating the systematics of other small
mammals (for example, Fisher & Bogan
1977, Gardner 1977, Bogan 1978, Diersing
& Wilson 1980, Carleton et al. 1982, Wil-
son 1991).

The excellent series of Nayarit cotton rats
collected by personnel of the U.S. Fish and
Wildlife Service in the middle 1970s war-
rant report in view of Zimmerman’s (1970)
seminal report on Sigmodon taxonomy. His
study, and the subsequent contributions of

814

Severinghaus & Hoffmeister (1978) and
Hoffmeister (1986), revealed the greater
species diversity and distributional com-
plexity of Sigmodon found in the south-
western United States and northwestern
México. In this report, we document the
kinds and distribution of Sigmodon species
in Nayarit (S. alleni, S. arizonae, and S.
mascotensis); review morphological and
chromosomal characteristics for identifying
the species, with emphasis on discrimina-
tion of S. arizonae and S. mascotensis; and
amplify the known geographic range of S.
mascotensis in western México, including
the reallocation of forms currently classified
as subspecies of S. hispidus (namely, Sig-
modon hispidus tonalensis Bailey, 1902,
and Sigmodon hispidus obvelatus Russell,
1952):

Materials and Methods

The 214 specimens of Nayarit Sigmodon
that form the nucleus of this report are con-
tained in the National Museum of Natural

Table 1.—Gazetteer of collecting localities (see Fig.

Locality

. Tacote, 1.4 mi N

Rio Canas, near La Concha
. Acaponeta

. Cuautla, 1 mi S

Santa Cruz, 6 km S

. Playa Colorado

. Playa Los Corchos, 4 mi E
San Blas

. Aticama, 4 km §

. Chacala

. Lo de Marcos, 1 mi S

. EL Venado, 3.5 mi E

. Arroyo de Jiguite

> EI Casco; 1:2) oS

pvica, Vallita, IukisS

a epic

. San Pedro Lagunillas, 2 mi E
. Estanzuela

. Coapan, 1.8 mi NW

. Ahuacatlan, 8 mi S

. Mesa del Nayar

. Ocota Airstrip

. Rancho Sapotito

SNe Ne ee ee
OMIDKHRWNKOCOOWMANADAWAWNE

NN WY VY
WNrFe ©

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

History, Smithsonian Institution, Washing-
ton, D.C. (USNM, the abbreviation for the
former United States National Museum). A
few originated from the pioneering Biolog-
ical Survey of México conducted by E. A.
Goldman and E. W. Nelson (1897 expedi-
tion to Tepic; see Goldman 1951), but most
were collected recently (1975-1977) by
personnel associated with the U.S. Fish and
Wildlife Service (now part of the USGS Pa-
tuxent Wildlife Research Center), pursuant
to a faunal analysis of Nayarit mammals.
Other specimens reported here, including
holotypes and type series, are housed in the
American Museum of Natural History, New
York (AMNH); the Field Museum of Nat-
ural History, Chicago (FMNH); Museum of
Natural History, University of Kansas,
Lawrence (KU); University of Michigan
Museum of Zoology, Ann Arbor (UMMZ);
The Museum, Michigan State University,
East Lansing (MSU); and Texas Coopera-
tive Wildlife Collections, College Station
(TCWC).

1).

Elevation Coordinates

(meters)

ils) 22°38 (1052
20 21°31 105265
46-75 22°29") aa
0 22°12 /1OS:38)

0 2.1°56'/105%35¢

0) 21°53 1052344

0 21°43'/105°25’

0 21°32'/1054

50 21°27 FOS
30 21°10 M053"

0 20°57 /1 05° 2s

100 22°57 (S04 Sie
100 21°49'/104°48’

60 21°45'/104°51’
760 21°35'/104°56'
1000 21°30'/104°53’
1300 21°12'/104°43’
1380 21°16'/104°28’
1560 21°09'/104°29'
1500 20°58'/104°28’
1300 22°12'/104°39’
1900 21°50'/104°13’
1100 2 120/103 S8-

VOLUME 112, NUMBER 4

815

Rio de
Acaponeta

Pacific
Ocean

Eig. I.

22@ \

Rio Grande
de Santiago

State of Nayarit, México, illustrating collecting localities of Sigmodon specimens housed in the

National Museum of Natural History (numbers refer to sites listed in Table 1).

Twenty-three Nayarit localities are rep-
resented in the preserved material (Table 1,
Fig. 1). Coordinates of collecting sites were
later determined from 1:50,000 topographic
maps used by the field teams and annotated
with the localities visited; most elevations
were recorded in the field with an altimeter

and later verified against the same topo-
graphic series.

Animals were generally preserved as
conventional study skins and skulls but also
as complete skeletons and formalin-fixed,
whole carcasses stored in alcohol. Prepa-
ration of standard chromosomal spreads,

816

definition of fundamental number (FN), and

descriptive terminology for chromosomal

morphology follow Patton (1967) and

Gardner & Patton (1976). The 32 voucher

specimens for the karyotypic variation re-

ported are contained in USNM< and repre-
sent the localities listed next.

Sigmodon alleni: Nayarit, Arroyo de Jigui-
te, Rio Santiago, 1 d (523935); 4 km S
Aticama, 1 3 (524480), 2 2 (524478,
524479); Chacala, 1 2 (523934); 1.2 mi
S El Casco, 1 2 (511699): 1 mi S Lo de
Marcos, 1 ¢ (523943), 3 2 (523940-
523942).

S. arizonae: Nayarit, Mesa del Nayar, 1 @
(511700); Ocota Airstrip, 2 3 (523948,
523950), 4 2 (523946, 523947, 523949,
523951). Sinaloa, Microondas La Mur-
alla, 1 ¢ (524487), 1 2 (524486).

S. mascotensis: Nayarit, 8 mi S Ahuacatlan
tid. ©239935)F 22 (6239929523954):
Arroyo de Jiguite, Rio Santiago, 4 4
(523962, 523963, 523965, 523966), 1 2
(523964); Rancho Sapotito, 2 ¢ (511703,
yt 704). *3, 2 GL 7O2. 511705, 511706).

We recorded 5 external, 18 cranial, and
3 dental dimensions (in millimeters, mm) to
summarize patterns of variation within and
between the populations sampled. Total
length, length of tail vertebrae, hindfoot
length, length of pinna from notch, and
mass (weight in grams, g) were transcribed
from skin labels. Crania were viewed under
a dissecting microscope when measuring
the 21 craniodental variables to 0.01 mm
with hand-held, digital calipers accurate to
0.03 mm. These measurements, their abbre-
viations as used in tables and figures, and
their landmark definitions where clarifica-
tion is necessary, include (see Carleton and
Musser, 1995, for illustration of most ana-
tomical endpoints): occipitonasal length
(ONL); zygomatic breadth (ZB); least in-
terorbital breadth (LOB); breadth of brain-
case (BBC)—taken behind the squamosal
root of the zygomatic arches, the caliper’s
jaws resting on the squamosal bones just
above the flange (inferior temporal ridge)

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

extending from the arches; breadth across
occipital condyles (BOC); depth of brain-
case (DBC); distance between temporal
ridges (DTR)—vertical distance between
the dorsal margin of the superior temporal
ridge and the ventral edge of the inferior
temporal ridge (see Fig. 8); length of ros-
trum (LR)—measured oblique to the mid-
longitudinal cranial axis, from the inner-
most bevel of the right zygomatic notch to
the end of the nasals at their midsagittal
junction; breadth of rostrum (BR)—dis-
tance across the lateralmost convexity of
the nasolacrimal capsules; postpalatal
length (PPL); length of bony palate (LBP);
breadth of bony palate (BBP); length of in-
cisive foramen (LIF); length of diastema
(LD); breadth of zygomatic plate (BZP);
length of zygomatic spine (LZS)—distance
between the anterior orbital rim and the tip
of the zygomatic spine; length of auditory
bulla (LAB); coronal length of maxillary
toothrow (LM1-3); coronal width of upper
first molar (WM1); depth of upper incisor
(DI); depth of mandible (DM)—distance,
taken on the lateral surface of the dentary,
from the rim of the m1 alveolus to the ven-
tralmost projection of the mental symphy-
SiS.

Relative age was coarsely indexed by de-
gree of molar wear according to the four
age-classes recognized by Carleton and
Musser (1989)—juvenile (J), young (Y),
full (A), and old-adult (O). The distinction
between juvenile and young adult cohorts
based on upper third molar eruption consti-
tuted a more objective criterion of age rec-
ognition than did the assignment of individ-
uals among the three adult classes based on
gradations of wear. Among specimens with
annotation of their reproductive state, many
we assigned as young or full adult based on
tooth wear exhibited signs of reproductive
maturity (testis scrotal, evidence of lacta-
tion, counts of embryos or embryo scars),
whereas those classified as juveniles did
not.

To augment sample sizes for the various
morphometric comparisons, Nayarit speci-

VOLUME 112, NUMBER 4

mens were grouped into eight analytical
samples (operational taxonomic units,
OTUs), as defined below (locality numbers
correspond to those identified in Table 1
and Fig. 1). To provide a comparative stan-
dard, we included a homogeneous series of
Sigmodon hispidus, sensu stricto, as a ninth
OTU (U.S.A., Florida, Wakulla Co., St.
Marks National Wildlife Refuge; n = 53;
USNM 526059-526106, 527358, 527359,
527362—527364). Since most collectors at
this locality also participated in the Nayarit
survey, one can expect procedural confor-
mity in the measurement of external vari-
ables.

Sigmodon alleni: OTU 1, n = 20, localities
8-15.

S. arizonae: OTU 2, n = 9, localities 2-7;
OTU 3, n = 25, locality 17; OTU 4, n =
6, locality 22.

S. mascotensis: OTU 5, n = 27, locality 13;
OTU 6, n = 46, locality 17; OTU 7, n =
12, localities 18-20; OTU 8, n = 6, lo-
cality 23.

S. hispidus: OTU 9, n = 53, Florida.

Standard descriptive statistics (mean,
range, standard deviation) were obtained
for the OTUs. Principal components and ca-
nonical variates were extracted from the
variance-covariance matrix and computed
using natural logarithmic transformations of
the 21 craniodental variables. Loadings are
expressed as Pearson product-moment cor-
relation coefficients of the principal com-
ponents or canonical variates with the orig-
inal skeletal and dental variables. All uni-
variate and multivariate computations were
generated using Systat (Version 7.0, 1997),
a series of statistical routines programmed
for microcomputers.

Discrimination of Nayarit Sigmodon and
Comparisons with Sigmodon hispidus

Intrasample age and sex variation.—
Field and lab workers who study Sigmodon
populations have regularly commented on
the considerable ontogenetic variation en-
countered (e.g., Chipman 1965, Baker

817

1969, Voss 1992, Zelditch et al. 1992). Cot-
ton rats are highly precocial and weaned an-
imals soon enter the trappable population;
seasonal differences in age composition of
a population, as well as differences in
growth rates between age cohorts, may be
remarkable (Layne 1974, Slade et al. 1984).
In view of such demographic factors, we
found age variation to be equally conspic-
uous in all species samples of Sigmodon re-
ported here, an impression quickly formed
from superficial observation of crania and
their wide range in size, shape, and rugos-
ity. Casual visual impressions are rein-
forced by statistical comparisons among
age groups in the large sample of S. hispi-
dus (Table 2). Nearly all measurements dis-
play regular, incremental increases in size
across the four age classes we defined, pro-
ducing age-correlated differences that con-
tribute substantially to nongeographic vari-
ation within our locality samples. Notable
exceptions include dimensions of the mo-
lars (LM1-3, WM1), which once erupted
decrease in crown height with occlusal use
but do not grow in size; in contrast, girth
of the incisor (DI) enlarges appreciably as
cotton rats age.

The contribution of sexual dimorphism
to intrasample variation, on the other hand,
is hardly apparent, at least given the unbal-
anced nature of analyzable material usually
consolidated from museum collections.
Only two (OB, DTR) of the 21 cranio-
dental measurements yielded significant
differences according to sex (Table 2), and
these are sufficiently infrequent and inci-
dental to suggest sampling error as an ex-
planation.

Other than procedural elimination of the
youngest age class, juveniles, we did not
adjust for size in the morphometric analy-
ses. Although variation attributable to post-
weaning growth may be substantial within
samples of Sigmodon, it is typically negli-
gible relative to the interspecific contrasts
that proved to be taxonomically important.
In this respect, patterns of morphometric
differentiation among cotton rats, their con-

818

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Arithmetic means of craniodental variables and results of one-way ANOVAs for sex and age cohorts
in a large sample of Sigmodon hispidus from Florida (OTU 9, n = 53).

M F
Variable (31) (22) f (sex)
ONL 34.8 34.3 1.0 3
ZB 19.5 19.3 1:5 1
IOB S22 Sl 4.2*
BBC 14.4 14.2 ao 1
BOC tS aS) 0.4
DBC 10.7 10.7 LC 1
DTR 3:1 2.9 oe
IER TER Ld 0.2
BR 67, 6.6 0.9
PPE 2 12.0 0.4 1
LBP 6.6 (ome) ty
BBP Ta is 0.2
LIB 7.8 Tak 0.1
LD 9.6 9.3 1.1
BZP 3.9 Shy 2.8
LZS 4.6 4.4 s(S)
LAB 6.3 6.3 0.0
LM1-3 6.4 6.4 0.2
WM1 Det Da | 0.5
DI O24) 2.0 0.4
DM 6.4 6:3 1.6

<P 01052 = iP = 00k = * = P = 0.000

Spicuous age variety notwithstanding, are
much like those derived for other closely
related, congeneric species of New World
muroids (e.g., Carleton & Musser 1989,
1995; Voss & Marcus 1992, Hoofer et al.
L909):

Morphometric variation.—Covariation
patterns derived from the 21 original cra-
niodental variables suggest two kinds of
large Sigmodon among Nayarit populations
(Fig. 2). These clusters correspond to the
species §. arizonae and S. mascotensis
(Figs. 3, 4), identifications independently
corroborated by agreement of karyotyped
specimens with currently understood differ-
ences in diploid number (2N = 22 or 28;
Zimmerman 1970) and by contrasts in cer-
tain qualitative features (see below). The
“correct”? phenetic association of relevant
holotypes—arizonae, major, and mascoten-
sis (Fig. 2)—bolsters the use of these names
for the Nayarit populations. Greater size in
all dimensions accounts for separation of S.

J
(3)
0.2
te
4.8
S)-5)
TS
0.1
2.4
9.8
6.1
0.5
6.1
7.0
6.7
8.1
S72
3.6
6.1
6.4
2.1
1.6
3.6

Ni A O
(13) (29) (8) Ff (age)
32.8 34.9 37.0 3321 ee
18.7 £95 20.3 DAWA
5.0 a2 553 OP
14.2 14.4 14.7 83755
7.4 7.4 Wie!) £9
10.6 10.7 Pi 8.084
3.0 Oe 333 L2ASSe
10.9 11.7 125 2AEES
6.3 6.7 WES 22 Sma
1p3 122 133 34. Lee
6.3 6.7 6:7 6232
es 7.8 8.1 24. Ves
E33 #4 8.5 EOS ae
Sah RL 10:5 31a
a7 a9 4.1 Ak oes
4.3 4.6 49 i Be ae
6.1 6.4 6.5 Gas
6.4 6.4 6.3 0.5
2.1 21 pe) 0.5
1.9 2.0 2A 49 .0***
6.1 6.5 6.6 14.0725

mascotensis and S. arizonae on the first
principal component (loadings uniformly
large and positive—Table 3); the generous
proportions of the latter’s molars provide
most discrimination on the second compo-
nent (LM1-3 and WM 1 correlations rela-
tively large and negative). The resulting
constellations of specimen scores conform
to the now familiar ordination pattern evi-
denced between morphologically similar,
closely related (congeneric) species of Mu-
roidea; furthermore, their elongate spread
and orientation, oblique to the bivariate plot
of PC I and PC II, suggest the interplay of
consistent interspecific shape differences
and age-related size increases (Voss et al.
1990, 1992). Included among our samples
is one locality where the two species were
collected in sympatry (2 mi E San Pedro
Lagunillas).

Unambiguous specific discrimination is
preserved in the discriminant function anal-
ysis of the eight OTUs representing S. ar-

VOLUME 112, NUMBER 4

4A S.mascotensis

PC II (7.7 %)

819

@o S. arizonae

A
mascotensis

AA

Ar 1

aX A

@
mayor
+

arizonae
A & +

O

0) 1 2 3

PC | (75.8 %)

Fig. 2. Plot of first and second principal components extracted from analysis of 21 log-transformed cran-
iodental variables measured on intact specimens of Nayarit Sigmodon arizonae (n = 29) and S. mascotensis (n
= 79). Open symbols indicate specimens collected in close proximity at 2 mi E San Pedro Lagunillas, 1300 m;
crosses refer to relevant type specimens discussed in the text. Regression lines of PC II on PC I differ significantly
between species in their y-intercepts (0.54 versus — 1.50; F = 144.9, P < 0.001) but not their slopes (0.42 versus

eee >. P = 0.553). See Table 3.

izonae, S. mascotensis, and S. hispidus (Fig.
5). Separation of Floridian S. hispidus on
the first canonical variate extracted primar-
ily results from differences in three vari-
ables (Table 4)—the larger size of the au-
ditory bullae (LAB), the narrower distance
between the temporal ridges (DTR), and,
perhaps in correlation with the latter, the
shallower braincase (DBC). LAB also gen-
erated the largest f-value in one-way anal-

yses of variance of the 21 craniodental mea-
surements among the three species. Less
conspicuously, the relatively greater length
of the facial region in S. hispidus is reflect-
ed in the moderate, positive loadings for
lengths of rostrum and incisive foramen,
whose univariate means match those of the
bigger S. arizonae. The generally larger
values and comparable range (mostly —0.5
to —0.7) of correlations on the second ca-

820

Table 3.—Results of principal component analysis
and one-way ANOVAs performed on all intact speci-
mens of Sigmodon arizonae (n = 29) and S. masco-
tensis (n = 79) from Nayarit, México (see Fig. 2).

Correlations
ANOVAs

Variable PC I PC II Ff (species)
ONL 0.97 0.15 DOIG
ZB 0.98 0.02 Soli
IOB (OTS) 0.07 4.1*
BBC 0.88 —0.20 TOO"
BOC 0.78 —=(0:22) Bales os
DBC 0.81 0.08 DOs
DTR 0.82 O01 SOLA
LR 0.94 0.16 Gea Es
BR 0.95 0.16 ETatserees
PB 0.96 0.13 Dray Ao REE
LBP 0.78 0.13 Srila
BBP 0.94 —=()) 113) AI OESS
LIF 0.94 0.01 44.3***
LD 0.96 0.18 DOO
BZP 0.86 0.28 Dea)
ZS 0.88 0.16 rg orsenes
LAB 0.85 = 0533 144.5***
LM 1-3 0.64 =—O:70 SOBs
WMI 0.53 —0.79 206.6***
DI 0.92 0.15 iis
DM 0.97 0.10 Does
Eigenvalue 15.8) 1.6
% Variance WDas Tedl

#105052 42° P= 00k 424 =P 30.001

nonical variate are more suggestive of gen-
eral size and again underscore the robust
proportions of S. arizonae apparent in vi-
sual inspection of skulls; the substantial di-
mensions of its molars (LM1-3, WM1) are
particularly noteworthy in this regard. Sam-
ples of S. mascotensis approximate that of
S. hispidus on the second axis.

A posteriori classifications of type spec-
imens historically associated with the S.
hispidus complex, as implemented in dis-
criminant function analysis of the same
eight OTUs, accord with some present tax-
onomic alignments and dispute others. The
nomenclatural significance of these statis-
tical assignments is considered in the Dis-
cussion and Taxonomic Summary (Fig. 17,
Table 6—see below).

Sorting examples of Nayarit S. alleni
from those of S. arizonae and S. mascoten-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

sis 1s Straightforward based on pelage color
(see below) or diploid number (2QN = 52;
Zimmerman 1970). Yet in cranial size and
discrete characteristics, S. alleni (Fig. 6) un-
expectedly proved somewhat difficult to
distinguish from S$. mascotensis; for exam-
ple, most univariate measurements of the
two overlap extensively (Appendix) and
disclose relatively few significant differenc-
es between means (Table 5). Principal com-
ponent analysis of log-transformed vari-
ables nonetheless supplied clear separation
of the two species based on crania alone
(Fig. 7). The variables most influencing dis-
crimination represent measurements taken
on the braincase (DBC, DTR), zygoma
(LZS), and molars (LM1-3, WM1), each of
which is notably greater in S. mascotensis
(Table 5; Appendix). Holotypes of interest
within the region appropriately associate
with their respective Nayarit populations,
those of mascotensis and colimae more so
and that of alleni less so. Although the dis-
position of the type specimen of alleni was
somewhat peripheral as divulged in PCA,
discriminant function analysis based on all
nine OTUs (not illustrated) indisputably
placed the holotype with the Nayarit sample
of the taxon.

Although external dimensions were ex-
cluded from multivariate examination, they
generally reflect the same pattern of inter-
specific size contrasts noted for cranioden-
tal variables and supply helpful guidance
for first-approximation field or museum
identification (Appendix). The absolutely
short and relatively narrow hind foot of S.
alleni, for example, readily separates that
species from young examples of S. masco-
tensis. As noted by Bailey (1902), hindfoot
size and tail length, absolute and relative,
help to distinguish S. mascotensis from ex-
amples of S. h. hispidus and h. berlandieri.
Relative length of tail in S. mascotensis (TL
ca. 45% of TOTL) also exceeds that in the
larger-bodied S. arizonae (TL ca. 40% of
TOTL), a species which otherwise stands
apart for its exceptional mass and size in all
other external variables quantified.

VOLUME 112, NUMBER 4

821

Rig. 3.

Dorsal and ventral cranial views (about 1.75) of adult Sigmodon: left pair, S. mascotensis (USNM

510026), a male from 2 mi E San Pedro Lagunillas, Nayarit; middle pair, S. hispidus berlandieri (USNM
157382), a male from 8 mi E Deming, New México; and right pair, S. arizonae (USNM 510040), a female from

2 mi E San Pedro Lagunillas, Nayarit.

Qualitative cranial traits.—Zimmerman
(1970) identified several consistent cranial
differences among Sigmodon populations
that corresponded to the chromosomal con-
trasts he documented and to the three spe-
cies he recognized among his samples—
namely, S. hispidus, S. arizonae, and S.
mascotensis. Other useful qualitative traits

were advanced by Severinghaus & Hoff-
meister (1978) and Hoffmeister (1986), par-
ticularly for separation of S. hispidus from
S. arizonae in the southwestern United
States. We here extend the utility of select
cranial features to the Nayarit populations
formerly included under S. hispidus.

The vertical distance between the supe-

Fig. 4. Lateral cranial view of adult Sigmodon:
top, S. mascotensis; middle, S. hispidus berlandieri;
and bottom, S. arizonae (same specimen numbers as
in Fig. 3).

rior and inferior temporal ridges (the latter
called the occipital crest by Zimmerman
1970) provides a reliable means for dis-
criminating examples of S. hispidus from
those of S. arizonae and S. mascotensis. As
noted by Zimmerman, this distance is visi-
bly narrower in S. hispidus relative to the
latter two (see Appendix), a difference un-
derscored by our multivariate results in
which distance between the temporal ridges
(DTR) heavily influenced the separation of
OTUs along the first canonical variate (Fig.
5, Table 4). The temporal ridges, together
with the lateral arc of the transversely ori-
ented lambdoidal ridge, define the size and
shape of the temporal fossa, an area on the
lateral wall of the braincase from which

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

originates the temporalis, an important mas-
ticatory muscle (Rinker 1954). In S. hispi-
dus (and S. alleni), the temporal fossa ta-
pers conspicuously toward the rear of the
skull, forming a trapezoidal shape over the
posterior half of the parietal and squamosal
bones; whereas, the fossa in S$. mascotensis
and S. arizonae occupies a larger area, ap-
proximately rectangular in outline on the
lateral braincase wall (Fig. 8). As expected
for osseus features that provide muscular
attachment, some change in fossa shape ac-
companies individual aging, such that ju-
venile animals of all three species exhibit a
stronger rearward convergence of the su-
perior and inferior temporal ridges; conse-
quently, the size and shape differences not-
ed are best appreciated when comparing
crania of similar age classes.

The anterodorsal edge of the zygomatic
plate forms a spinous process in all Sig-
modon examined here, but its projection,
and the degree of concavity imparted to the
plate’s anterior border, can aid specific iden-
tification, as demonstrated by Severinghaus
and Hoffmeister (1978) for Arizonan pop-
ulations of S. hispidus and S. arizonae.
Among Nayarit samples, the spine is lon-
gest and most acute in specimens of S. ar-
izonae, shortest in those of S. alleni. Con-
figuration of the anterior zygoma in S. mas-
cotensis more closely resembles S. arizonae
but is not so extreme. Such interspecific
contrasts are partially conveyed by certain
variable loadings (BZP, LZS) generated
from principal component and discriminant
function analyses (Tables 3, 4), as well as
by their mean differences (Appendix). Nev-
ertheless, the expression of the spinous pro-
cess has a strong age component (Table 2),
which must be considered when comparing
and identifying individuals. In S. hispidus,
compared with S. arizonae, the spinous pro-
cess is not only shorter, but its dorsal border
appears wider and its tip is usually blunt
(Severinghaus & Hoffmeister 1978:868, fig.
1; Hoffmeister 1986). In some individuals
of S. hispidus, the spinous process is even
expanded anteriorly to produce a knoblike

VOLUME 112, NUMBER 4

@® S. arizonae

CV 2 (26.4 %)

Rig. 5:

A S.mascotensis

823

@ S.hispidus

CV 1 (54.5 %)

Plot of the first and second canonical variates extracted from discriminant function analysis of

eight samples of Sigmodon arizonae (OTUs 2-4), S. mascotensis (OTUs 5-8), and S. hispidus (OTU 9). Each
of the eight OTU centroids is surrounded by a polygon that encloses maximal dispersion of sample scores.

See Table 4.

tip. The contrast is similar in kind, if some-
what less pronounced in degree, between
samples of S. hispidus and S. mascotensis.

The relative volume of the auditory bul-
lae, difficult to capture accurately in a linear
dimension (i.e., LAB) but easy to see in
side-by-side comparisons of skulls, is an-
other feature that separates S. hispidus from
both S. arizonae and S. mascotensis—no-
ticeably more inflated in the former and less
so in the latter two (Fig. 3). Still, LAB con-
tributed even more heavily than distance
between the temporal ridges (DTR) to the
segregation of the S. hispidus sample along

the first canonical variate (Fig. 5, Table 4).
Voss (1992) characterized the auditory bul-
lae of S. hispidus as “‘small,’’ but his tax-
onomic context involved contrast with the
manifestly rotund capsules possessed by
South American S$. peruanus, a species in-
digenous to dry habitats in western Ecuador
and northwestern Pert. Within the genus,
the auditory bullae in members of the S.
hispidus complex may be loosely graded as
medium-sized, those of S. alleni, S. arizo-
nae, and S. mascotensis as small, and those
of S. peruanus as large.

The anatomy of the posterior palatal re-

824 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 4.—Results of discriminant function analysis and one-way ANOVAs performed on eight OTUs repre-
senting intact specimens of Sigmodon arizonae (n = 29) and S. mascotensis (n = 79) from Nayarit, México,
and §. hispidus (n = 50) from Florida, U.S.A. (see Fig. 5).

Correlations

ANOVAs
Variable CV CV 2 f (OTU)

ONL 0.28 (NS 15.8***
ZB 0.08 (0), 74k 19°:4*55
IOB 0.30 =O0°55 3.95
BBC 0.24 0.67 13.9%2%
BOC —0.19 AOL OI, 10104454
DBC = (5) =(0) 59s) 16.82%
DTR —0.58 —0.61 39:85
LR 0355 0.93 140s
BR 0.13 —0.53 CASES
PPE 0.19 —0.61 Lees
LBP 0.38 =0395 8.258
BBP 033 —0.46 TROzes
LIF 0.40 —0.60 1OOe
(AD) 0.30 055 14 eas
BZP 0.10 =0.36 5:4
LZS = (0083 = (595) 104e
LAB OF =055 T3BtSs
LM1-3 0.22 —0.78 38.5845
WM1 O20 =0.79 32.3534
DI 0.31 —0.43 7.788%
DM O83 —0.58 13.48
Canonical correlations 0.94 0.89
Eigenvalue fies) 3.8
% Nariance 54.5 26.4

** = P= (0.01; *** = P = 0.001.

gion affords several points of contrast that
aid specific separation. In examples of S.
arizonae, the palatine bones are distinctly
keeled and terminate medially as a pro-
nounced spine that projects into the meso-
pterygoid fossa (Fig. 9). A well formed me-
sopterygoid spine is atypical of S. masco-
tensis skulls (Figs. 9, 10), although the pos-
terior border of their palatines may be
gently curved or occasionally bluntly point-
ed. Even in the latter condition, however,
the bony palate is relatively flat, unmarked
by the conspicuous palatal gutters and
raised keel observed in specimens of S. ar-
izonae. The ventral opening of the foramen
ovale, situated at the posterolateral corner
of the parapterygoid fossa, is notably large
in most S. arizonae and smaller in S. mas-
cotensis (Figs. 9, 10). Although we oppor-
tunistically used a 0.9 mm-diameter probe

to convey this difference, some objective
measure of foraminal area would better un-
derscore the size distinction between the
species. With regard to both palatal con-
struction and size of the foramen ovale, the
Floridian sample of S. hispidus resembles
S. arizonae, but its variability is greater, at
least according to the character states we
have defined.

Another useful characteristic, one not
mentioned by Zimmerman (1970) or Sev-
eringhaus & Hoffmeister (1978), involves
the occurrence of an oval-shaped vacuity or
fenestra on the parapterygoid fossa. Such
an opening, situated just laterad to the pter-
ygoid process and astride the palatine-pter-
ygoid suture (Fig. 9), occurs commonly in
samples of S. mascotensis (ca. 70%) but un-
commonly in specimens of S. arizonae and
S. hispidus (<25%). Nevertheless, this

VOLUME 112, NUMBER 4

Fig. 6. Dorsal, ventral, and lateral cranial views
(about 1.75X) of an adult Sigmodon alleni (USNM
524479), a female from 4 km S Aticama, Nayarit.

character varies in all three species, so that
the fenestra’s presence or absence is not
alone decisive for identification but infor-
mative when applied in combination with
other qualitative and quantitative variables.
In some individuals, especially those of S.
mascotensis, the position of a presumptive
fenestra is suggested by an oval area of
thin, translucent bone (“‘Present, covered”
per histogram, Fig. 11). By its location and
orientation, the parapterygoid fenestra ap-
pears to correspond to the path of the anas-
tomotic artery that crosses the dorsal sur-
face of the pterygoid plate to supply the
distal cephalic circulation in muroids hav-
ing a reduced stapedial branch, as is true in
most Sigmodon (Bugge 1970, Voss 1992,
and see Carleton and Musser 1989: fig. 21).
Why the impression of this artery’s passage
should usually ossify fully in some Sigmo-
don species but not in others is unknown;

825

the opening, however, does not appear to
transmit nerves or blood vessels.

With regard to the typical conditions we
reviewed above for S. arizonae, S. hispidus,
and S. mascotensis, our series of S. alleni
present an interesting mosaic of conditions.
Like examples of S. mascotensis, the fora-
men ovale is small and the parapterygoid
fenestra is typically present (on both sides
in 10 individuals; one side in 2; and absent
in 3); a mesopterygoid spine is usually
present but small with shallow palatal cor-
rugations. The temporal fossa, on the other
hand, is narrow, convergent posteriorly in
the manner of S. hispidus. The spinous pro-
cess is short, slightly overhanging the zy-
gomatic plate and tapering to a point (not
wide and blunt as in S. hispidus). The au-
ditory bullae of S. alleni are absolutely the
smallest of the four species we have ex-
amined (Appendix) and in proportion near-
ly match the capsules of S. mascotensis.

As for other cranial features noted by
Zimmerman (1978) or Severinghaus &
Hoffmeister (1978), we subjectively as-
sessed their variability apropos the Nayarit
samples, but did not attempt to quantify
their diagnostic utility because of their
shape complexity or definitional arbitrari-
ness (e.g., curvature of the lateral nasal
margins, width of the presphenoid, shape of
the occipital shield). Of these, the angular-
ity (S. hispidus) or not (S. arizonae and S.
mascotensis) of the dorsal rim of the occip-
ital shield, as described by Severinghaus
and Hoffmeister (1978), seems to provide
consistent contrast, at least for the regional
examples we examined.

Pelage color and texture.-—Among the
three species of Nayarit Sigmodon, S. alleni
visually stands apart based on the uniformly
rich brown color, occasionally with rufous-
or cinnamon-brown tones, of its dorsal pel-
age. The common name, brown cotton rat,
is aptly descriptive of the species. Rufes-
cent tints are most evident over the rump,
with medium brown on the middle dorsum
that fades on the flanks to create a paler
tawny hue. In texture, the dorsal fur of S.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

4 S. mascotensis

aap “se

Ansa

mascotensis

A

0 1 2 3

PC | (64.0 %)

eS. allen

3

2
Oe 1
<<
co
(je)
=e
O
OW

={

)

-3 2 =|

Biees ie

Plot of first and second principal components extracted from analysis of 21 log-transformed cran-

iodental variables measured on intact specimens of Nayarit Sigmodon alleni (n = 15) and S. mascotensis (n =
79). Crosses indicate relevant type specimens discussed in the text. Regression lines of PC II on PC I differ
significantly between species in their y-intercepts (1.84 versus —0.42; F = 156.0, P < 0.001) but not their slopes

(0.18 versus 0.28; F = 0.4, P = 0.544). See Table 5.

alleni is comparatively soft and fine, only
slightly stiff to the touch; guard hairs on the
rump are mostly unicolored and project lit-
tle above the cover hairs.

The upperparts of S. mascotensis and S.
arizonae suggest some shade of brown: typ-
ically a paler saturation, brighter tone, and
distinctly grayish hue in the former species;
and a darker saturation, more somber cast,
and yellowish hue in the latter. In speci-
mens of S. arizonae, there is greater inter-
mixture of darkly tipped cover hairs over
the mid-dorsum, which contrasts more no-
ticeably with the yellow browns of the
flanks; in examples of S. mascotensis, the
grayish brown pelage color is usually even-

ly expressed across the dorsum. The subtle
difference in shade of brown also results
from the band colors of their agouti-pat-
terned cover hairs: the basal band a pale
plumbeous gray and middle band medium
buff in S. mascotensis versus a dark gray
basal band and deep buff middle band in S.
arizonae. The chromatic accent of the buffy
middle bands against the darker bases and
tips of the cover hairs imparts a more griz-
zled or flecked appearance to the upperparts
of S. arizonae and S. mascotensis; in indi-
viduals of S. alleni, the middle band is
ochraceous and less dramatically set off
from the umber tips. In further contrast to
S. alleni, the dorsal fur in the two larger

VOLUME 112, NUMBER 4

Table 5.—Results of principal component analysis
and one-way ANOVAs performed on all intact speci-
mens of Sigmodon alleni (n = 15) and S. mascotensis
(n = 79) from Nayarit, México (see Fig. 7).

Correlations

ANOVAS

Variable PC I rea Ff (species)
ONL 0.93 0.24 0.5
ZB 0.95 0.01 iie*%
IOB 0.48 0.27 0.8
BBC 0.73 (05 SiGtt
BOC 0.44 =0:10 OrF*
DBC 0.61 —0.43 ail Le
DTR 0.78 —0.44 Age Te
LR 0.85 0.39 05
BR 0.82 0.37 1.0
PPLE 0.90 0.26 0.5
LBP 0.44 0.17 0.0
BBP 0.85 0.03 Ss
LIF 0.87 0.19 2.6
LD 0.89 0.36 0.0
BZP 0.85 0.08 3.0
EZS 0.77 —0.44 Arf es
LAB 0.67 =0413 ees
LM1-3 0.42 —0.49 46.4***
WM1 0.43 =().52 SOON
DI 0.87 0.17 D22)|
DM 0.90 0:25 0.4
Eigenvalue 0.073 0.012
% Variance 64.0 10.8

ee 09, — P= 0.01; *** = P= 0.001.

827

species is coarser and sparser, especially so
in S. arizonae, and can be tactually appre-
ciated as hispid. Guard hairs are conspicu-
ously longer than the cover hairs over the
rump and consist of both monocolored
black and agouti-banded types.

Ventral cover hairs of all three species
are bicolored, having a plumbeous gray
base and a pale tip. The general appearance
of the ventrum and the color differences ob-
served between species principally depends
upon the pigmentation of the tips. In S. al-
leni, the tips are buffy to pale ochraceous
and the bases dull plumbeous, imparting a
somber, dark gray color to the venter. In S.
mascotensis, the cover hairs terminate in
pure white, creating a light gray impression
and conveying brighter tones than the un-
derparts of the other species, especially S.
alleni. Ventral pelage color is more variable
in S. arizonae; some individuals have a pre-
dominance of pale buffy tips, while in oth-
ers they are dull white. The overall impres-
sion is one of dull to medium gray, in con-
trast to the brighter grays of S. mascotensis.

The upper surfaces of the hindfeet are
generally well haired in all three species but

Fig. 8.

Left lateral view of the temporal region in adult examples of Sigmodon hispidus (left; USNM 526071,

Florida, St. Marks National Wildlife Refuge) and S. mascotensis (right; USNM 523954, Nayarit, 8 mi S Ahu-
acatlan). Abbreviations: ab, auditory bullae; exo, exoccipital; ip, interparietal; itr, inferior temporal ridge; Ir,
lambdoidal ridge; pa, parietal; rza, squamosal root of the zygomatic arch; sq, squamosal; str, superior temporal
ridge. The superior temporal, inferior temporal, and lambdoidal ridges outline a trapezoidal shape of the temporal
region in S. hispidus in contrast to the rectangular shape observed in S. arizonae and S. mascotensis. The distance
between the temporal ridges (DTR) was measured between the points of the arrows denoting the superior (str)

and inferior (itr) ridges.

828 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 9. Ventral view of the bony palate and adjoining parapterygoid and mesopterygoid fossae in Nayarit
examples of Sigmodon arizonae (top; USNM 510012, 2 mi E San Pedro Lagunillas) and S. mascotensis (bottom;
USNM 510022, 2 mi E San Pedro Lagunillas). Abbreviations: ab, auditory bullae; al, alisphenoid bone; bo.
basioccipital bone; bs, basisphenoid bone; fen, fenestra of parapterygoid fossa; fo, foramen ovale; if, incisive
foramen; max, maxillary bone; mf, mesopterygoid fossa; ms, mesopterygoid spine; pal, palatine bone; ppf,
parapterygoid fossa; spv, sphenopalatine vacuity; sq, squamosal bone. In S. arizonae, as compared with S.

VOLUME 112, NUMBER 4

chromatic differences are apparent. Hairs
covering the dorsal metatarsus and phalan-
ges of S. alleni have dusky bases with buffy
to pale ochraceous tips, such that hindfoot
color generally blends with the reddish-
brown appearance of the dorsum. Like the
hairs of its ventral pelage, those on the
hindfeet of S. mascotensis have plumbeous
gray bases and bright white tips; thus, the
feet appear pale gray and distinctly contrast
with the grayish browns of the rump. Upper
surfaces of the hindfeet of S. arizonae ap-
pear medium gray to dull brown over the
metatarsum and grayer on the phalanges,
blending with the dorsum more so than in
S. mascotensis but less so compared with S.
alleni.

Although Bailey (1902) characterized all
of these taxa as having semi-naked and
coarsely annulated tails, we found clear dif-
ferences in caudal pilosity and scutellation.
Epidermal scales appear dark brown, above
and below, in all three; consequently, ex-
pression of bicoloration issues from the in-
terplay of hair color, density, and length.
None of the species, however, possesses a
truly bicolored tail (e.g., like that of Pero-
myscus maniculatus) with sharp linear de-
marcation between dark dorsal and pale
ventral surfaces. Instead, transition from
darker dorsal to the paler undersurface is
gradual. In examples of S. alleni, the tail is
sparsely covered, the caudal hairs extending
over 2—3 annuli; the color is dark brown
dorsally and slightly paler below, bicolora-
tion weakly expressed and effectively uni-
color in some individuals; caudal scales are
small but typically visible to the naked eye
given the relatively sparse investiture of
hairs. Individuals of S. mascotensis possess
a similarly fine scalar pattern, but caudal
hairs are longer, about 4 annuli in length,
and scutellation is mostly obscured. This

829

species most nearly approaches a bicolored
condition, brown above and pale gray be-
low. In samples of S. arizonae, the tail is
dark brown above and grades imperceptibly
to paler brown below. Caudal scales are no-
ticeably larger, and the annulations accord-
ingly coarser and readily evident macro-
scopically. Paradoxically, the annulation
pattern remains visibly appreciable in S. ar-
izonae even though its caudal hairs are ab-
solutely longer (about 4—5 annuli) in con-
trast to those of S. mascotensis. The im-
pression of semi-nakedness arises from the
sparser density of caudal hairs, arranged in
triplets as in the other species but more
widely spaced as a result of the larger tail
scales.

Chromosomal variation.—Three well
marked chromosomal formulae are appar-
ent among Nayarit Sigmodon. The diploid
number (2N) of S. arizonae is 22, consist-
ing of largely biarmed chromosomes (fun-
damental number, FN, = 38). The auto-
somes number four pairs of very large
metacentrics and submetacentrics, two pairs
of large subtelocentrics, two pairs of me-
dium-sized metacentrics, one pair of small
submetacentrics, and a single pair of small
acrocentrics (Fig. 12). The X chromosome
iS a medium-sized acrocentric; the Y, a
small acrocentric with discernible second-
ary arms, is the smallest element in the
complement. This karyotype is identical to
that described by Zimmerman (1970) for
the 22-chromosome cotton rats he studied
from Arizona, Sinaloa, and Nayarit.

The diploid number in examples of S.
mascotensis is invariantly 28, nearly all of
the autosomes uniarmed (FN = 28). Auto-
somes consist of 12 pairs of acrocentrics,
gradated in size from medium to large, and
one pair of small subtelocentrics (Fig. 12).
The X chromosome is a medium-sized ac-

<_

mascotensis, note the occurrence of a pronounced mesopterygoid spine, the conspicuously larger size of the
foramen ovale, and the absence of a parapterygoid fenestra.

830 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

CONDITION OF BONY PALATE

100%, .. aiZonae

mascotensis
Fae AS. Ne <3 hispidus

50% ee ee eee Poe te ee ce oe |e cee ede Ip iS kena Pe? chico Core Ole Ske Bo ce ee eS RP ee SO eee ce ee ee,
hispidus
(yl Ae PR ean (MRC oe rk: een ees A eee Bee brea ene Been a) 2
aoe mascotensis
25, arizonae
Pa AOE ela iB cies hia ea Cl a
Mesopterygoid Spine Acute Mesopterygoid Spine Blunt/Absent
Palatal Gutters Deep Palatal Gutters Shallow
SIZE OF FORAMEN OVALE
100% Oe ee et 8 Oo Cee es Oise feces ae is ee eM Ee Gh AR te Wun os 2c
arizonae
hispidus mascotensis
75% Paras a Pee ere re Om eEranaT | lo OMe) Clic Ceara Cer Aer ener are Thee 6 Oh DEED oO pais ceca ao fe 2 te
50% ee tee era eee eee nce ee | eee ema Pe ee) oo te ey RCA Ce ire cole cce oe PARE Oe
DE O/T: aes ae ene Gee eee ee Sos DS Sere Be Pepi u ai hee cote ie
arizonae
0% a eh tay ar ne Baie re

Foramen Ovale > 0.9 mm Foramen Ovale < 0.9 mm

Fig. 10. Percent occurrence of certain qualitative cranial characters in samples of Sigmodon arizonae (n =
36-40) and S. mascotensis (n = 88-91) from Nayarit, México, and in S. hispidus (n = 53) from Florida, U.S.A.

Top, condition of the bony palate; bottom, size of the foramen ovale (See Fig. 9 for illustration and text for
description of character states).

VOLUME 112, NUMBER 4

831

PATENCY OF PARAPTERYGOID FENESTRA

arizonae

hispidus ns eee es ne. Ae

mascotensis

25% et a Ba ie fatln| va Vi gw aes me) cakle! Waa ee ce, epiep we eee. see) we 3 6 8 ce a se we sre wwe mascotensis Acree wt ener ey of | f> ) © | Of OO [> ee) Se
arizonae
mascotensis
arizonae
hispidus
0% Ste a wie cbc eto e Soe etal one ie ag Pee | tee Be aS) fe eo ene ate eas) too ate en ao) Cyr Mei) (CaPEurcmicnin me ewe (nie Sl ae jo Ae ea
Absent Present, Covered Present

Fig. 11. Percent occurrence of character states of the right parapterygoid fenestra in samples of Sigmodon

arizonae (n = 38) and S. mascotensis (n = 86) from Nayarit, México, and in S. hispidus (n = 52) from Florida,
U.S.A. (See Fig. 9 for illustration and text for description of character states).

rocentric; the Y, a tiny acrocentric, is small-
er than any of the autosomal acrocentrics.
A female (USNM 511706) from Rancho
Sapotito is heteromorphic for a centric fu-
sion involving two similarly sized larger ac-
rocentrics; two other females and two males
karyotyped from this locality possess typi-
cal karyotypes. Lee & Zimmerman (1969)
noted similar examples of heteromorphic
karyotypes resulting from centric fusion in
S. fulviventer, and Zimmerman (1970) re-
ported the same phenomenon in S. hispidus.

Zimmerman (1970) characterized all au-
tosomes in S. mascotensis as acrocentric
(that is, FN = 26), but we follow Elder
(1980) in describing the smallest of these
as subtelocentric (FN = 28). In most
spreads, this pair has clearly discernible
second arms (see Fig. 12), which Elder
(1980) found to be heterochromatic in most
of his C-banded preparations. Excepting
this minor discrepancy in autosomal clas-
sification, the karyotype of Nayarit animals

conforms to those described by Zimmer-
man (1970) from scattered localities in
southern Jalisco, Michoacan, Guerrero, and
western Oaxaca.

In S. alleni, the 2N is consistently 52 and
the FN is typically 64. Autosomes in the
64-FN karyotype consist of one pair of
large, four pairs of medium-sized, and one
pair of small subtelocentrics; one pair of
very small metacentrics; and 18 pairs of ac-
rocentrics graded in size from small to me-
dium (Fig. 13). At Lo de Marcos, two FNs,
64 and 66, are apparent. The autosomal
complement in the 66-FN karyotype in-
cludes an additional pair of small submeta-
centrics and lacks a pair of acrocentrics.
The X chromosome is a moderately large
subtelocentric; the Y is a small subtelocen-
tric. The karyotype Zimmerman (1970) re-
ported for S. alleni from Michoacan is sim-
ilar to our 64-FN pattern, except that the Y
chromosome in our Nayarit samples ap-
pears to be smaller.

832 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

KUURKKRX FA

an

oh ro oe Ak

AA AN AGRO ARAN

DAD
xX XxX
Fig. 12. Representative karyotypes of Nayarit Sigmodon: a, S. arizonae (USNM 511700), a female from

Mesa del Nayar; b, S. mascotensis (USNM 523953), a male from 8 mi S Ahuacatlan; c, heteromorphic variant
of S. mascotensis (USNM 511706), a female from Rancho Sapotito.

b

C

y

VOLUME 112, NUMBER 4

Fig. 13.

Discussion

Bailey’s (1902) revision of Sigmodon es-
tablished the view that most North Ameri-
can populations with coarsely annulated,
partially naked tails are geographic races of
S. hispidus, including the distinctive com-
plex of large-bodied cotton rats found in the
southwestern U.S.A. and western México.
His classification prevailed over subsequent
decades and accommodated most new de-
scriptions of taxonomic variety as subspe-
cies (Miller 1924, Miller & Kellogg 1955,
Hall & Kelson 1959). Zimmerman’s (1970)
fine synthesis of chromosomal and morpho-
logical information abruptly overturned this
comfortable arrangement and resurrected
both S. arizonae and S. mascotensis as valid
species distinct from S. hispidus. The mor-
phological basis for specific stature of these
forms was amplified by Severinghaus &
Hoffmeister (1978) and Hoffmeister (1986),
who also refined our understanding of their
intermingling distributions in the south-
western U.S.A. and northwestern México.
Morphometric, morphological, and chro-
mosomal differences observed among Nay-
arit populations of Sigmodon further cor-
roborate the taxonomic insights and nomen-
clatural realignments advanced by Zimmer-
man (1970).

Examples of S. alleni, S. arizonae, and

833

Jy AE AK 2D tb ms
AA 44 AR AR AN MA

£a AR AA AR AK HA
an AR AA QA 20

),

Representative karyotype of Sigmodon alleni (USNM 523940), a male from 1 mi S Lo de Marcos.

S. mascotensis were collected in all possible
pairwise combinations of sympatry, but at
no place were all three species documented.
Of particular note is the co-occurrence now
recorded for S. arizonae and S. mascotensis
at a locality east of San Pedro Lagunillas in
southcentral Nayarit. Although collected
*““sympatrically”’ in terms of bearing a sin-
gle skin-tag provenience, the two species
are apparently segregated according to mi-
crohabitat at this place. The vegetation in
the vicinity includes succulent dicots, par-
ticularly water hyacinth, and marsh grasses
bordering the small lake; and dryer brush,
low sparse woodland, and bunchgrasses
covering the hillsides that overlook the lake
(Fig. 14). Most examples of S. arizonae
originated from the former habitat and
those of S. mascotensis from the latter (Fig.
15). Whether the two species at this place
are entirely nonoverlapping in their micro-
habitat occurrence is unknown, since field
identifications were tentative at the time of
collection. Furthermore, whether the specif-
ic habitat affinities recorded at this one lo-
cality reflect general ecological differences
of the species, particularly their humidity
tolerances, will require extended site inves-
tigations.

The distinctive cranial and pelage fea-
tures so apparent in Nayarit S. mascotensis

834

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 14. Panoramic view of collecting site at the eastern side of the small lake San Pedro Lagunillas and
two miles east of the village of the same name. Traplines were placed both along the emergent lakeside vegetation
and on the dryer hillside and tableland.

encouraged our review of its distribution in
western México. We concur with all spe-
cies-group epithets so far allocated to the
species by Zimmerman (1970) and Hall
(1981): these are atratus, colimae, inexo-
ratus, and ischyrus. Like Allen’s (1897b) S.
mascotensis, populations represented by
each of these names are large, grayish
brown cotton rats with strongly hispid fur
and relatively long tails; their crania possess
relatively small auditory bullae and expan-
sive temporal fossae; the occurrence of par-
apterygoid fenestrae is common within lo-
cality samples, but a large foramen ovale,
pronounced mesopterygoid spine, and deep
palatal grooves are uncommon. The forms
tonalensis Bailey (1902) and obvelatus
Russell (1952), currently maintained as
subspecies of S. hispidus (Hall 1981, Mus-
ser & Carleton 1993, Alvarez-Castafieda
1996), also fit well with this morphology

and properly belong as junior synonyms of
S. mascotensis (see remarks under Taxo-
nomic Summary).

The emergent picture of S. mascotensis
distribution reveals a species confined to
western México, documented from south-
ern Nayarit and Zacatecas to extreme west-
ern Chiapas (Fig. 16). Although its range
boundaries appear convoluted and constit-
uent populations are certainly discontinu-
ous, the occurrence of the species corre-
sponds closely to deciduous or semidecid-
uous tropical woodlands having a pro-
nounced dry season. Comments by
collectors typically mention the semiarid to
arid environment and-or xerophilous vege-
tational character of localities where S.
mascotensis has been captured (Hooper
1947, 1957; Goldman 1951; Russell 1952;
Goodwin 1969; Alvarez et al. 1987). Ac-
cording to Goldman’s (1951) life-zone di-

VOLUME 112, NUMBER 4 835

Fig. 15. Microhabitat preferences of Sigmodon captured at 2 mi E San Pedro Lagunillas (see Fig. 14). Top:
hillside covered with brush, low broken woodlands, and bunch grasses where examples of S. mascotensis were
captured. Bottom: dense growth of water hyacinth fringing the lake and inhabited by S. arizonae.

836

Ocean

Pacific

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Gulf of Mexico

Fig. 16. Generalized distribution of Sigmodon mascotensis in western México based on specimens examined
herein. Enclosed stars identify the type localities of the six species-group taxa that we view as regional synonyms
of S. mascotensis Allen (1897), including two newly associated herein (tonalensis Bailey, 1902, and obvelatus

Russell, 1952).

visions of México, nearly all of these dis-
tributional records represent either Arid
Lower Tropical Subzone or Arid Upper
Tropical Subzone, a few the Lower Austral
Zone; by Leopold’s (1959) categorization
of broad vegetational units, they fall within
Tropical Deciduous Forest and Arid Tropi-
cal Scrub. Not unexpectedly, in view of
such climatic and vegetational associations,
the geographic boundary of the species
conforms well to certain biotic patterns dis-
tilled from distributions of the Méxican her-
petofauna (Flores-Villela 1993).

The range of S$. mascotensis collectively
overlies three physiographic regions: Pacif-
ic coastal plain and the contiguous lowlands
of the Isthmus of Tehuantepec; the drainage
basin and surrounding tablelands of Lago
de Chapala (Meseta Central de Anahuac or
Altiplano Sur), including the Tequisquiapan

Valley; and the Balsas Basin, including the
Tehuacan Valley. Except for those records
in the Tehuacaén valley of western Puebla,
collecting sites of S. mascotensis predomi-
nantly occur within hydrologic systems that
ultimately drain into the Pacific Ocean; the
Rio Salado of the Tehuacan Valley joins the
Rio Santa Domingo, a major effluent to the
Gulf of México. The Tehuacan-Cuicatlan
Valley is presently considered part of the
same physiographic domain as the Balsas
Basin (Sierra Madre del Sur Morphotecton-
ic Province), but persuasive argument for
past biotic connections drawn from geolog-
ical evidence is elusive, given both the
daunting complexity of México’s physical
landscapes and the continuing need for geo-
logic-tectonic research (see review by Fer-
rusquia-Villafranca 1993).

Although our study is focussed on means

VOLUME 112, NUMBER 4

for interspecific discrimination, we are as
much impressed by the morphological sim-
ilarities that S. arizonae and S. mascotensis
share, notably the configuration of the tem-
poral fossa and degree of bullar inflation.
Goodwin (1969:199) had earlier appreciat-
ed the fundamental affinity among these
populations and summarized their external
resemblances: “‘Sigmodon h. mascotensis is
in a group of several subspecies of very
large, long-tailed, coarse-haired cotton rats
that range from Arizona [1.e., arizonae] in
western North America south along the
slopes and plains of the Pacific coast in
México to Tonala, Chiapas.’’ Such general
notions of kinship, along with their geo-
graphically complementary distributions,
suggest a sister-group relationship, a hy-
pothesis which has received more rigorous
endorsement from traditional and banding
chomosomal studies (Zimmerman 1970,
Zimmerman & Sihvonen 1973, Elder
1980). Zimmerman (1970), for example, di-
vided North and Middle American Sigmo-
don into two species groups based on a
combination of diploid and fundamental
numbers: those with a low range (2N = 22-—
30, FN = 26-38), consisting of S. arizonae,
S. fulviventer, and S. mascotensis; and those
with a high range (2N = 52-56, FN = 52-
66), consisting of S. alleni, S. hispidus, S.
leucotis, and S. ochrognathus. Assuming
derivation from a S. hispidus-like ancestor,
G-banding evidence supports the common
ancestry of S. arizonae and S. mascotensis
(Elder 1980). The phyletic affinity of S. ful-
viventer with S. arizonae and S. mascoten-
sis as postulated by Zimmerman (1970) re-
mains untested.

More importantly, our nomenclatural and
distributional amendments of S. mascoten-
sis bear on a taxonomically more entangled
issue: what is S. hispidus proper? As now
documented, populations of medium to
large Sigmodon known to occur throughout
the Pacific coastal lowlands and west-facing
slopes from Sonora to western Chiapas,
represent either the species S. alleni, S. ar-
izonae, or S. mascotensis. Supposed west-

837

ern records for S. hispidus proper have
proven incorrect upon reexamination. Spec-
imens from Jalisco and Querétaro that had
been assigned to S. h. berlandieri (Bailey
1902, Hall 1981) are in fact S. mascotensis,
as enumerated below in the Taxonomic
Summary; Goodwin’s (1969) two vouchers
(KU 63075, 63076) of S. h. saturatus in
southern Oaxaca (1 mi NNW Soledad) are
instead S. alleni, as corrected by Baker
(1969). To our knowledge, these reidentifi-
cations, together with reallocation of obve-
latus and tonalensis and their referred se-
ries, remove any documentation for S. his-
pidus along the entire Pacific versant of
México.

The inability of discriminant coefficients
to associate Middle American holotypes of
nominal “‘hispidus” with Floridian S. his-
pidus indirectly attests the still composite
nature of the species as arranged. While our
selection of this Floridian population sam-
ple as a standard for S. hispidus proper was
a reasonable choice, on geographic and
morphologic grounds, our expectation of its
agreement with Neotropical taxa proved
wonderfully naive. Most holotypes repre-
senting such populations clustered among
or nearer the Nayarit samples of S. masco-
tensis (Fig. 17) and were classified a pos-
teriorly with that species (Table 6). None of
these numerically probabilistic assign-
ments, except Bailey’s (1902) tonalensis,
merits serious consideration as biological
truth—the various taxa are not conspecific
with S. mascotensis of western México.
Those craniodental variables that heavily
influenced multivariate separation among
centroids—particularly LAB on the first ca-
nonical variate and LM1-3 and WM1 on the
second—are small in most of the Middle
American type specimens, as compared
with typical S. hispidus (OTU 9). By de-
fault these were associated with examples
of S. mascotensis, since among the eight
predefined OTUs, individuals of that spe-
cies possess the smallest bullae, shortest
toothrows, and narrowest molars compared
with typical S. hispidus or S. arizonae (Ap-

838 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

@ S. arizonae 4 S.mascotensis @ S. hispidus
6
4
a
A
inexoratus
0 masco nsis. 55 + M
jacksoni ealimae
+
25) ischyrus arizonae
+
plenus
-4
-6
6 borucae
+
microdon
4 +
zanjonensis,
confinus
2 Saturatus s +
A chiri }
ene. komarecki
eremicyé + .
0 lonialensis + + virginianus
; florid
griseus on aes
-2
-4
-6

6 4 ©) 0 2 4 6

Fig. 17. Phenetic dispersion of type specimens of Sigmodon based on discriminant coefficients extracted
from the same discriminant function analysis of eight samples of Sigmodon arizonae (OTUs 2-4), S. mascotensis
(OTUs 5-8), and S. hispidus (OTU 9). Polygons in this instance enclose the maximal dispersion of all specimen
scores around the grand centroid for each species (compare with Fig. 5). Top, type specimens of forms currently
considered synonyms of S. arizonae (jacksoni, plenus, and major) and S. mascotensis (colimae, inexoratus, and
ischyrus); bottom, type specimens of forms currently considered synonyms of S. hispidus (two groups plotted
separately to avoid visual congestion). See Table 6 and text for discussion.

VOLUME 112, NUMBER 4

839

Table 6.—A posteriori classification of type specimens of Sigmodon derived from discriminant function anal-

ysis of OTUs 2-9 (see Fig. 17).

Current
Holotype classification

arizonae Mearns S. arizonae
(AMNH 2370/1841)

major Bailey S. arizonae
(USNM 96275)

Jacksoni Goldman S. arizonae
(USNM 214121)

plenus Goldman S. arizonae
(USNM 181086)

mascotensis Allen S. mascotensis
(AMNH 8321/6655)

colimae Allen S. mascotensis
(AMNH 2648/2318)

inexoratus Elliot S. mascotensis
(FMNH 8651)

ischyrus Goodwin S. mascotensis
(AMNH 10045)

borucae Allen S. hispidus
(AMNH 171600)

eremicus Mearns S. hispidus
(USNM 60319)

tonalensis Bailey S. hispidus
(USNM 75144)

saturatus Bailey S. hispidus
(USNM 99998)

microdon Bailey S. hispidus
(USNM 108467)

chiriquensis Allen S. hispidus
(AMNH 18789)

griseus Allen S. hispidus
(AMNH 28497)

confinus Goldman S. hispidus
(USNM 204241)

zanjonensis Goodwin S. hispidus
(AMNH 69277)

floridanus Howell S. hispidus
(USNM 261624)

virginianus Gardner S. hispidus
(USNM 273535)

komareki Gardner S. hispidus

(USNM 207210)

pendix). By way of instructive contrast,
those holotypes originating from the south-
eastern U.S.A. (floridanus, komareki, vir-
ginianus) did group sensibly with the Flo-
ridian sample of S. hispidus (Table 6).
Members of the hispidus complex, so far
as known from the southern U.S.A. to
northwestern South America, possess a rel-
atively narrow temporal fossa, tail notably

oT’ assignment P
2 (arizonae) 0.71
2 (arizonae) 0.99
5 (mascotensis) 1.00
3 (arizonae) 1.00
8 (mascotensis) 1.00
7 (mascotensis) 0.67
7 (mascotensis) 1.00
5 (mascotensis) O73
8 (mascotensis) 0.63
7 (mascotensis) Chey)
8 (mascotensis) 1.00
5 (mascotensis) 0.70
9 (hispidus) 0.92
7 (mascotensis) 0.65
5 (mascotensis) 0.64
5 (mascotensis) 0.69
7 (mascotensis) 0.98
9 (hispidus) 1.00
9 (hispidus) 0.99
9 (hispidus) 1.00

Shorter than head-and-body length, and a
diploid number of 52 (Zimmerman & Lee
1968, Kiblisky 1969, Zimmerman 1970),
but in pelage color and texture and in cra-
nial form and size, they vary substantially.
Based on our review of most relevant types
and casual examination of museum series,
typical S. hispidus appears to comprise
those populations in the southeastern

840

U.S.A. (floridanus, komareki, virginianus),
the southern Great Plains (alfredi, texi-
anus), and the central plateau of northern
México (berlandieri). To the southwest and
south of this moiety, the taxonomic picture
becomes muddled. Populations in Arizo-
na—those bearing the names confinus, er-
emicus, and jacksoni—deserve additional
systematic clarification, with respect to their
synonymy as S. hispidus, as S. arizonae, or
as something else (Zimmerman 1970, Hoff-
meister 1986). Along the Gulf coast of east-
ern México, across the Isthmus of Tehuan-
tepec, and into the lowlands of the Yucatan
Peninsula and northern Guatemala, one
finds a group of populations characterized
by shorter fur, generally somber brown in
color, shorter toothrows, and smaller audi-
tory bullae. The oldest name for these is
Saussure’s (1860) toltecus, which may in-
clude the forms saturatus, microdon, and
perhaps furvus (Bangs, 1903); Bailey
(1902) had mentioned a foltecus group
within his broad polytypic definition of S.
hispidus. Hispid cotton rats described from
the highlands of Chiapas and western Gua-
temala, villae and zanjonensis, respectively,
resemble those of the toltecus complex in
size but possess a longer, more brightly col-
ored pelage; the elevation of the type lo-
cality of zanjonensis, ca. 2700 m, alone ad-
vises caution in accepting the homogeneity
of populations now grouped under the epi-
thet in conventional range maps (Hall
1981). The relationships of the Méxican
hispidus-like forms to populations farther
south in Central America—griseus, boru-
cae, and chiriquensis—similarly invite de-
tailed study. As noted by Voss (1992); Bur-
meister’s (1854) hirsutus, whose type lo-
cality is in northern Venezuela, may assume
importance in deciding nomenclatural is-
sues that involve these southernmost pop-
ulations of the hispidus complex.

The number and distribution of species
still mistakenly subsumed under “‘hispidus”’
are topics that require much additional in-
vestigation, incorporating other kinds of
data and fine-scale geographic sampling.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Like Hooper’s (1955) early perceptions of
taxonomic differences among Peromyscus
boylii populations, unraveling the intricate
alpha systematic problems that yet surround
Sigmodon will likely unfold from firm an-
swers attained first on select regional bases.

Taxonomic Summary

Partial synonymies that trace first author-
ities for new name combinations are given
below for S. arizonae and S. mascotensis.
Species-group taxa given under S. alleni
and S. hispidus are intended only as a list
of the type specimens we examined, and are
not a statement of junior synonyms dem-
onstrated to be conspecific. For the latter,
we include names that have been applied
only to populations in the southwestern
U.S.A., México, and Central America.

Sigmodon arizonae Mearns

Sigmodon hispidus arizonae Mearns, 1890:
287 (type locality—United States, Ari-
zona, Yavapai County, 3 mi SE Camp
Verde, Bell’s Ranch [as restricted by
Hoffmeister 1986]; holotype—AMNH
2370/1841).

Sigmodon arizonae, Zimmerman, 1970:435
(elevation to species).

Sigmodon hispidus major Bailey, 1902:109
(type locality—México, Sinaloa, Sierra
de Choix, 50 mi NE Choix; holotype—
USNM 96275).—Zimmerman, 1970:446
(synonymy with S. arizonae).—Hall,
1981:742 (retention as a subspecies of S.
arizonae).

Sigmodon hispidus jacksoni Goldman,
1918:22 (type locality—United States,
Arizona, Yavapai County, 3 mi N Fort
Whipple, near Prescott, 5000 ft; holo-
type—USNM 214121).—Zimmerman,
1970:446 (synonymy with S. arizo-
nae).—Hall, 1981:742 (retention as a
subspecies of S. arizonae).

Sigmodon hispidus cienegae Howell, 1919:
161 (type locality—United States, Ari-
zona, Pima County, 4 mi E Fort Lowell,
Bullock’s Ranch; holotype—A. B. How-

VOLUME 112, NUMBER 4

ell collection 1531).—Zimmerman, 1970:
446 (synonymy with S. arizonae).—Hall,
1981:741 (retention as a subspecies of S.
arizonae).

Sigmodon hispidus plenus Goldman, 1928:
205 (type locality—United States, Ari-
zona, Yuma County, Parker, 350 ft; ho-
lotype—USNM 181086).—Zimmerman,
1970:446 (synonymy with S. arizo-
nae).—Hall, 1981:742 (retention as a
subspecies of S. arizonae).

Distribution.—Extreme southeastern
California and southcentral Arizona,
U.S.A., southwards along coastal plain and
contiguous foothills of the Sierra Madre
Occidental in northwestern México, to
southcentral Nayarit. In Nayarit, most col-
lecting sites situated in the coastal plain that
dominates the northwestern sector of the
State, but also at intermediate elevations
(1000—1900 m) along the upper drainage of
the Rio Grande de Santiago (Fig. 18).

Apart from the unresolved status of jack-
soni (see below), collection records, as re-
ported here and by Hoffmeister (1986),
likely delimit the full geographic extent of
S. arizonae. In our museum searches to re-
define the distribution of S$. mascotensis in
western México, we have not encountered
examples of S. arizonae to the south of the
localities now documented from Nayarit. To
the northwest—in Sinaloa, Durango, and
Sonora—places where the species has been
collected lie on coastal plain and lower Pa-
cific-facing slopes of the Sierra Madre Oc-
cidental. Along eastern slopes of that moun-
tain axis and onto adjoining plateaus, one
finds populations of S. hispidus proper, cor-
responding to the race S. h. berlandieri
(Baker 1956, Baker & Greer 1962, Ander-
son 1972, Severinghaus & Hoffmeister
1978, Matson & Baker 1986).

The presence of S. arizonae at moderate
elevations in eastern Nayarit (Mesa del
Nayar, 1300 m; Ocota Airstrip, 1900 m) ap-
pears anomalous in light of the usual oc-
currence of the species in coastal plain of
the westcentral region (Fig. 18). Both kar-

841

yotypic and morphological data, however,
confirm the specific assignment of these
outlier populations. Carleton et al. (1982)
had noted similar disjunct distributional
records among certain Nayarit Peromyscus
and speculated that riparian vegetation
along major rivers, like the Rio Grande de
Santiago and Rio Huayanamota, may pro-
vide dispersal corridors to the state’s inte-
rior. Matson (1982) offered a similar expla-
nation for certain tropical rodent taxa that
reach the canyon region of southwestern
Zacatecas.

Remarks.—Nayarit samples of this large-
bodied Sigmodon most closely resemble
Bailey’s (1902) S. hispidus major, de-
scribed from nearby Sinaloa. Zimmerman’s
(1970) argument for the taxon’s synonymy
with Mearn’s (1890) S. arizonae is persua-
sive, and our observations and results sup-
port his conclusion. While the discriminant
scores of the holotypes of arizonae and
plenus fall outside the limits of craniodental
variation obtained for the Nayarit series
(Fig. 17), a posteriori classification casts the
morphological affinity of the Arizonan taxa
with them and major, not with representa-
tives of S. hispidus or S. mascotensis (Table
6). Further, some allowance must be duly
given for geographic variation, and Zim-
merman (1970) had previously remarked
that the Sinaloan and Nayarit samples av-
eraged slightly larger than those from Ari-
zona, a distinction reflected in his cluster
analysis. Verification of a diploid count of
22 for animals from the type locality of ar-
izonae, where specimens have not been col-
lected since 1932 according to Hoffmeister
(1986), would help to cement the usage of
this name; to date, reported karyotypes rep-
resent the taxa cienegae, major, and plenus
(Zimmerman 1970, this study).

The invocation of geographic variation
less easily explains the unexpected assign-
ment of the type of S. hispidus jacksoni to
Nayarit S. mascotensis (Table 6). Zimmer-
man (1970) arranged jacksoni, also named
from Arizona, as another junior synonym of
S. arizonae, but Hoffmeister (1986) later

842

Table 7—Measurements (mm) of type specimens
of taxa currently associated with Sigmodon arizonae.

arizonae major plenus jJacksoni

Sex M M M M
Age A O A J
TOTL 320.0 365.0 316.0 249.0
LT 116.0 156.0 147.0 108.0
LHF 36.0 40.5 36.5 33.0
ONL 40.2 40.4 37.1 ais
ZB pay | 235 2A 18.1
IOB 3-5 os. 5 4.9
BBC 14.6 16.0 14.4 14.2
BOC 8.1 8.0 8.2 74
DBC 11a 12.8 23 jm |
DTR 4.5 Sai 3.6 532
LR 13.6 14.0 12.0 10.0
BR 8.2 8.6 74 6.7
PPE 14.0 14.7 13.4 10.0
LBP TS TS Tz 6.2
BBP 8.2 8.7 8.5 6.7
LIF 8.6 8.8 8.7 6.3
LD 143 11.4 1 8.5
BZP 44 4.6 4.6 3.3
LZS 4.7 6.2 = 4.2
LAB 6.4 6.5 6.6 5.8
LM1-3 6.62 6.96 6.54 6.35
WM1 2.36 23 PIPL? 2.05
DI ZAP 255 2.03 1.59

DM 7.4 7 7.0 6.1

drew attention to the problematic character-
istics of the type, a juvenile and so far the
only known specimen. We can only echo
Hoffmeister’s reservations while assuring
that the specimen is not an individual of S.
mascotensis. It was statistically associated
with that species as the best multivariate
choice among the taxonomic alternatives
offered in our discriminant function analy-
sis and given the weight accorded certain
pivotal variables, especially LAB and LM1-
3. With regard to both, the molar row and
bulla of jacksoni are small compared with
other holotypes and with the Nayarit series
of S. arizonae (Table 7, Appendix). Nor
does the individual’s immature age suffi-
ciently explain the erroneous numerical
classification—length of molar row, at
least, is age invariant. And we reiterate that
inclusion of juveniles in preliminary mor-
phometric comparisons, whether principal

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

component or discriminant function analy-
ses, did not obscure taxonomic separation
among Nayarit species and S. hispidus. The
conformation of the zygomatic spine in
jJacksoni more closely resembles the con-
dition typical of S. hispidus than that of S.
arizonae. We doubtfully retain Goldman’s
(1928) jacksoni in the synonymy of S. ar-
izonae for want of more convincing evi-
dence of its specific relationships and sta-
tus.

Ecological notes.—Localities where
specimens of S. arizonae were captured in
Nayarit generally fall within the Arid Upper
Tropical Subzone (sensu Goldman 1951).
The dominant vegetation at these sites was
characterized by field personnel as savanna-
woodlands, deciduous tropical forest, palm
forest, or mangrove swamp. For localities
where microhabitat origin can be reason-
ably determined, however, individual Sig-
modon were consistently trapped in open
grassy and herbaceous settings, usually in
the presence of ample water—whether in
the form of rivers and their floodplains,
lakes and ponds, or drainage sloughs
through agricultural fields and pastures. As
noted above, S. arizonae taken near San Pe-
dro Lagunillas inhabited a dense stand of
water hyacinth bordering the small lake,
some of the traps placed on floating mats
of the plant. At Ocota Airstrip in south-
eastern Nayarit (1900 m), surrounded by
pine and oak forest, S. arizonae were
trapped only in grassy patches at the bottom
of a nearby arroyo with standing pools of
water. Near La Concha, the single specimen
of S$. arizonae was swimming in the shal-
low Rio Canas and caught by hand, after
diving straight to the bottom to escape the
first attempt at capture.

In view of its presence on coastal plain
and along river valleys into Nayarit’s inte-
rior, S. arizonae was collected with a wide
variety of rodent species: Sciurus aureo-
gaster, S. colliaei, Spermophilus annulatus,
S. variegatus, Pappogeomys bulleri,
Thomomys umbrinus, Chaetodipus pernix,
Liomys irroratus, L. pictus, Baiomys tay-

VOLUME 112, NUMBER 4

lori, Hodomys alleni, Neotoma mexicana,
Oryzomys palustris, Peromyscus boylii, P.
maniculatus, P. melanophrys, P. pectoralis,
P. simulus, P. spicilegus, and Reithrodon-
tomys fulvescens. It was documented with
S. alleni in southern coastal plain (Chacala,
San Blas) and with S. mascotensis in the
southern ridge and valley country (2 mi E
San Pedro Lagunillas).

Specimens examined.—Durango: Chaca-
la (USNM 96735-96742). Nayarit: Acapo-
neta (USNM 91398, 91399); near La Con-
cha, Rio Canas, Nayarit-Sinaloa state line
(USNM 509978); 1 mi S Cuautla (USNM
509980, 510765); Mesa del Nayar, 4500 ft
(USNM 511700); Ocota Airstrip (USNM
523944523951, 524385); Playa Colorado,
15 km S Santa Cruz (USNM 553988
553995, 554152, 554153); 4 mi E Playa
Los Corchos (USNM 553996, 554158); San
Blas (USNM 89214); 2 mi E San Pedro La-
gunillas, east side of lake (USNM 509999,
510003, 510012, 510013, 510017—510019,
510028-—5 10033, 510035-—510037, 510039,
510040, 510042-—510045, 510047, 510764,
510765, 510768, 510769, 510771, 510775,
510777, 510780—510783, 510785, 510788,
510791); 6 km S Santa Cruz (USNM
553987, 554151); 1.4 mi N (by rd) Tacote
(USNM 509979, 510764); Tepic (USNM
88237). Sinaloa: Ahomé (USNM 131541-—
131543); Culiacan (USNM 96741); Escui-
napa (AMNH 24072-24078, 24080-24084,
24086-24094, 24096-24103, 24105-
24117, 24484-24486, 24488, 24490-—
24492, 24494-24497, 24777, 24778,
24476-24478, 24480-24483, 24889-
24891, 25887; FMNH 20098); Microondas
La Muralla (USNM 524486, 524487); Pi-
nos Gordo (UMMZ 75265); Plomosas
(USNM 91401); Rosario (AMNH 13755;
USNM 91400); Sierra de Choix, 50 mi NE
Choix (USNM 96270-96275). Sonora: Al-
amos (USNM 96276-96278); Bacerac
(USNM 250901); Nogales (USNM 2213);
Magdalena (USNM_ 17806/24743-—17808/
24745); 23 km S Nogales (USNM 251032);
Oputo (USNM 250902—250903).

843

Sigmodon mascotensis Allen

Sigmodon mascotensis Allen, 1897b:54
(type locality—Meéxico, Jalisco, Mineral
San Sebastian, near Mascota, 3300 ft; ho-
lotype—AMNH 8321/6655).—Bailey,
1902:108 (reallocated as a subspecies of
S. hispidus).—Zimmerman, 1970:435
(reinstatement as species).—Hall, 1981:
740 (arrangement as nominate subspe-
cies).

Sigmodon colimae Allen, 1897b:55 (type
locality—Mexico, Colima, plains of Co-
lima; holotype—AMNH 2648/2318).—
Bailey, 1902:108 (synonymy with S. his-
pidus mascotensis).—Allen, 1906:209
(reinstatement as subspecies of S. hispi-
dus).—Zimmerman, 1970:446 (synony-
my with S. mascotensis).—Hall, 1981:
741 (full synonymy with S. m. mascoten-
SiS).

Sigmodon hispidus tonalensis Bailey, 1902:
109 (type locality—Meéxico, Chiapas,
Tonala; holotype—USNM 75144).

Sigmodon hispidus inexoratus Elliot, 1903:
144 (type locality—Meéxico, Jalisco,
Ocotlan; holotype FMNH 8651).—Hall,
1981:740 (reallocation as a subspecies of
S. mascotensis).

Sigmodon hispiduas [sic] atratus Hall,
1949:149 (type locality—México, Mi-
choacan, 6.5 mi W Zamora, 5950 ft; ho-
lotype—MVZ 100628).—Russell, 1952:
82 (synonymy with S. hispidus inexora-
tus Elliot).—Hall, 1981:740 (full synon-
ymy with S. mascotensis inexoratus).

Sigmodon hispidus obvelatus Russell, 1952:
81 (type locality—México, Morelos, 5 mi
S Alpuyeca, 3700 ft; holotype—TCWC
4921).

Sigmodon hispidus ischyrus Goodwin,
1956:8 (type locality—México, Oaxaca,
Yautepec District, Santo Domingo Chon-
tecomatlan, “El Arco” gorge of Rio
Grande, 2600 ft; holotype—AMNH
171600).—Zimmerman, 1970:446 (syn-
onymy with S. mascotensis).—Hall,
1981:740 (retention as a subspecies of S.
mascotensis).

844

Distribution.—Coastal plain, interior ba-
sins, and west- and south-facing slopes of
western México, from southern Nayarit and
southwestern Zacatecas to southern Oaxaca
and extreme southwestern Chiapas; occur-
ring at low to intermediate elevations (sea
level to 2100 m), most localites of speci-
mens examined falling between 500 and
1500 m; and absent from the highlands of
the Transverse Volcanic Range, Sierra Ma-
dre del Sur, and northern Oaxaca (Fig. 16).
In Nayarit, limited to the southeastern quad-
rant of the state, mostly in tablelands and
low mountains but also collected at low el-
evation in the valley of the Rio Grande de
Santiago (Fig. 18).

The range of S. mascotensis closely ap-
proaches that of the S. hispidus complex in
three regions where sympatry may be even-
tually recorded. Two involve relatively arid
interior valleys, the Tequisquiapan Valley,
in southern Queretaro and western Hidalgo,
and the Tehuacan Valley, in western Puebla.
The third includes the warm lowlands of the
Isthmus of Tehuantepec in western Chiapas
and southeastern Oaxaca, where, as percep-
tively observed by Goodwin (1969:199),
“Sigmodon h. mascotensis does not inter-
grade with the smaller short-tailed cotton
rats of the hispidus group [i.e., toltecus and
saturatus| that are prevalent on the Gulf
drainage.” The basis of his conviction can
be readily grasped when perusing the
AMNH series from the Tehuantepec dis-
trict.

Remarks.—In hindsight, the specific af-
finity of forms now gathered under synon-
ymy of S. mascotensis was predictable from
the early descriptions themselves. Thus,
Bailey (1902:109) characterized tonalensis,
his new subspecies of S. hispidus, as *‘Sim-
ilar to mascotensis but larger and slightly
browner,”’ and further allowed that “‘This is
merely a large form of mascotensis, from
which there is no sharp geographic sepa-
ration.”’ In similar manner, the recurring fo-
cus of contrast in the differential diagnosis
of the other synonyms has been Allen’s
(1897b) S. mascotensis (Elliot 1903, Hall

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1949, Russell 1952, Goodwin 1956), in-
cluding his own S. colimae, which he de-
scribed in the same publication. With regard
to the separation of the latter from S. mas-
cotensis, Allen (1897b:55) acknowledged
that “‘I am unable to recognize any distinc-
tive cranial features.’’ Specific recognition,
as for many other forms described in the
late 1800s, rested upon subtle differences in
pelage color or size and those posited from
actually few specimens (Allen had available
four individuals each of his new species
mascotensis and colimae).

Our examinations of types, referred spec-
imens of original type series, and other mu-
seum holdings confirm the morphological
homogeneity and probable genetic close-
ness among those populations now ar-
ranged under S. mascotensis. Allen’s
(1897b) type of S. mascotensis (AMNH
8321/6655) was consistently associated
phenetically with our Nayarit samples
(Figs. 2 and 17, Table 6), as were the four
other holotypes seen and measured (coli-
mae, tonalensis, inexoratus, and ischyrus—
Table 6). Pelage color and texture of the
latter four, as well as their cranial features,
match the general color pattern and mor-
phology of S. mascotensis. We did not ex-
amine the type of Hall’s (1949) atratus
(MVZ 100628), though Russell’s (1952)
merger of the Michoacan taxon into full
synonymy with S. hispidus inexoratus, ac-
cepted by Hall & Kelson (1959) and main-
tained by Hall (1981) as S. mascotensis
inexoratus, conveys much about the speci-
men’s inseparability from S. mascotensis.
Hall’s own illustration (1949:150, plate [X)
of the type specimen strongly suggests an
example of S. mascotensis, including pos-
session of the oblong paraptertygoid fenes-
trae; his single referred specimen, from Za-
mora (USNM 120268), is undoubtedly an
example of the species. Nor did we person-
ally study the holotype of Russell’s (1952)
obvelatus (TCWC 4921); however, all other
specimens from his original series (TC WC
4920, 4922, 4923) exhibit the characteristic

VOLUME 112, NUMBER 4

845

alleni
arizonae
mascotensis

Fig. 18. Distribution of three species of Sigmodon in Nayarit, México, based on collections reported herein
(dashed-dotted line = 100 m contour; light stipple = elevations between 1000 and 2000 m; dense stipple =

elevations above 2000 m).

morphology of S. mascotensis, as do others
in the USNM collections from Morelos.

In accepting the name changes proposed
by Zimmerman (1970), Hall (1981) re-

tained most junior epithets as subspecies of
S. mascotensis, a nomenclatural formality
unsubstantiated by fresh analyses of intra-
specific variation. Our qualitative appraisal

846

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 8.—Measurements (mm) of type specimens of taxa considered as synonyms of Sigmodon mascotensis.

mascotensis colimae

tonalensis

inexoratus ischyrus

Sex i M
Age A A
TOTL

ET

LHF

ONL 34.7 36.0
ZB 19.5 19.9
IOB Ww S21
BBC 13.5 13.9
BOC 7.4 8.2
DBC a PRS
DTR 4.2 Sey
LR be. t 12.0
BR 6.4 6.8
PPE 12.4 17
LBP 6.1 6.6
BBP tS 8.0
LIF 8.7 7.8
LD 10.0 9.6
BZP 4.0 4.2
ZS 4.7 = |
LAB 5.4 6.0
LM1-3 6.12 6.61
WM1 2.18 220
DI 1.97 212
DM 6.2 6.3

of some 500+ museum specimens, collect-
ed across the range of S. mascotensis as
now defined, suggests that such future anal-
yses should devote attention to populations
from eastern Oaxaca and contiguous Chia-
pas. Even allowing for the advanced age of
the type specimen of Bailey’s (1902) tona-
lensis (Table 8), the populations it repre-
sents—for example, those from Tehuante-
pec and Guiengola, Oaxaca—are impres-
sive for their size, which approximates the
dimensions attained by adult S. arizonae. In
addition, these large Oaxacan animals typ-
ically possess a more corrugated palate and
pronounced mesopterygoid spine, again
more like examples of S. arizonae and less
like most S. mascotensis. Still, in inflation
of the bullae, size of the foramen ovale, fre-
quency of parapterygoid fenestrae, and pel-
age features, these series fit best with S.
mascotensis.

Apart from the enigmatic combination of

M M M
O A A
350.0 310
166.0 146 145.5
41.0 36 34.5
38.9 3741 37.4
22.0 20.5 215
5.8 5.0 3.2
14.9 14.3 14.3
8.1 Tey TT
12.5 10.9 R17
4.4 3:9 4.4
L397 122 13.4
7.8 7.4 i
14.0 13.9 14.0
6.3 7.4 6.5
8.3 1 8.2
S. ae 8.4
11.4 10.7 10.8
4.8 4.4 4.0
a2 5.8 a4
39 5.6 Bo
6.23 6.32 6.13
2.09 2.24 2.10
PAA | 2.14 2AM
Pe. 6.7 6.7

traits found in these southern populations,
samples of S. mascotensis from the northern
and middle parts of its range appear uni-
form and display the kinds of minor inter-
locality variation attributable to differences
in age composition, molting stage, and col-
lecting season. Further, no chromosomal
variation is thus far known for populations
distributed over this region, from Nayarit to
western Oaxaca (Zimmerman 1970, this re-
port). To our knowledge, chromosomes of
named forms from eastern Oaxaca (ischy-
rus) or western Chiapas (tonalensis) have
yet to be examined.

Ecological notes.—Sigmodon mascoten-
sis Was common in the southern part of
Nayarit, where most collecting sites repre-
sent the Arid Lower Tropical Subzone
(Goldman 1951). The region is topograph-
ically diverse with low mountains and ta-
blelands dissected by river valleys and
steeply eroded arroyos. Ridges are covered

VOLUME 112, NUMBER 4

with tropical deciduous forest that grades
into oak and pine-oak woodland associa-
tions at the upper elevations; meandering
streams and valley bottoms, rocky outcrops
and old lava flows afford ample terrain for
open grass and brush stands and mixed sa-
vanna-scrublands. Where verifiable, speci-
mens of S. mascotensis originated from the
latter habitats, as well as from around ag-
ricultural fields bordered with rank brush.
At Estanzuela, traplines that produced S.
mascotensis ran through grassland and bro-
ken pine-oak woods and along a stream-
bank covered with ferns. South of Ahuaca-
tlan, examples of S. mascotensis were cap-
tured in tall grasses interspersed among
rocks, bare soil, and scattered oaks and cac-
tus.

In addition to Sigmodon alleni and S. ar-
izonae, S. mascotensis was documented
sympatrically with Sciurus aureogaster, S.
colliaei, Spermophilus variegatus, Pappo-
geomys bulleri, Liomys irroratus, L. pictus,
Baiomys taylori, Neotoma mexicana, Ory-
zomys melanotis, O. palustris, Peromyscus
maniculatus, P. melanophrys, P. pectoralis,
P. spicilegus, and Reithrodontomys fulves-
cens. The locality of sympatry with S. al-
leni, Arroyo de Jiguite near its confluence
with the Rio Grande de Santiago (100 m),
is situated in a lowland region where a
number of intermediate- to high-elevation
Peromyscus species were unexpectedly
found to occur (Carleton et al. 1982).

Specimens examined.—Chiapas: Cuinta-
lapa, Cinco Cerros (AMNH 148825-
148828); Cintalapa, 555 m (UMMZ 92476,
94033-94035, 96306); Tonalaé (AMNH
167430, 167431); mountains near Tonala
(USNM 75144, 76089). Colima: Armeria,
sea level—200 ft (USNM 33323/45351-
33332/45360, 34185); Colima, 1700 ft
(USNM 33333/45361—33341/45369); plains
of Colima (AMNH 2646/2319, 2647/2129,
2648/2318, 2649); 3 mi SE Colima
(UMMZ 100693, 100694). Guerrero: 0.5
mi W Acahuizotla, 3000 ft (UMMZ
109205); Acapulco, near sea level (USNM
70764); Agua del Obispo, 2900 ft (UMMZ

847

108558); Chilpancingo, 4800 ft (FMNH
47550, 47552, 47554-47557, 48466,
51947-51949; UMMZ 81461-81470,
84015, 84016, 89262, 104977-104991;
USNM 70300, 70760—70763); Cuapongo
(FMNH 47553); El Limon (126530,
126531); Iguala, 750 m (UMMZ 92597,
93455-93462); near Ometepec, 200 ft
(USNM 71575, 71576); Tlalixtaquilla,
4200 ft (USNM 70234). Jalisco: Ameca,
4000 ft (USNM 82247); 6 mi W Ameca,
4300 ft (UMMZ 95495-95499); 7 mi W
Ameca, 4000 ft (UMMZ 95500); Aten-
quique (AMNH 26322-26324); Atemajac,
4500 ft (USNM 34146/46223); 4 mi NE
Autlan, 3000 ft (UMMZ 95501-95507,
96268); Estancia (AMNH 25289, 25346,
25350, 25357, 25358, 25364-25366); Hue-
juquilla, 5400 ft (USNM 90810); Lagos,
6150 ft (USNM 78969, 80348); Las Canoas
(AMNH 26317-26321, 26551-26557);
Mascota, Mineral San Sebastian, 3300 ft
(AMNH 8321/6655, 8322/6656); 0.5 mi W
Mazamitla (UMMZ 100695, 100696,
100698); Ocotlan, 5000 ft (FMNH 8651-—
8653, 8656-8658, 8660—8662, 8664-8666;
USNM_) 120153-—120159); Ojo de Agua,
near Amatl4an (AMNH 25360); Rancho
Palo Amarillo, near Amatl4an (AMNH
25353-25356); Arroyo de _ Plantanar
(AMNH 25348, 25349, 25361); 2 mi N Re-
solana, 1500 ft (UMMZ 95510-95512);
Rio Santa Maria (AMNH 25347, 25359);
Tuxpan (AMNH 26293, 26294); Zacoalco,
3000 ft (USNM 34143/46220—-34145/
46222); Zapotlan, 4000 ft (USNM 3359/
45624, 33590/45625). Michoacan: Hacien-
da El Molino (USNM 20440/35670); La
Huacana (USNM 126522—126529); La Sa-
lada (USNM 126520, 126521); 9.6 mi S
Lombardia, 1400 ft (UMMZ 110562,
110563); Los Reyes (USNM 125933-
125935); Queréndaro, 5800 ft (USNM
50201); 12 mi S Tzitzio, road to Huetamo,
1050 m (UMMZ 92214); Zamora (USNM
120268). Morelos: 5 mi S Alpuyeca, 3500
ft (TCWC 4921, 4922); Cuernavaca, 5000
ft (USNM 51164, 51165); 2 km S Jonaca-
tepec, 4500 ft (TCWC 4920); Puente de

848

Ixtla (USNM_ 126938—126940); Yautepec,
4100 ft (USNM 51166-51169); 6 mi W
Yautepec, 6000 ft (TCWC 4923). Nayarit:
8 mi S (by rd) Ahuacatlan (USNM 523952—
523955); Amatlan (AMNH 25345, 25351,
25352); Arroyo de Jiguite, Rio Santiago,
100 m (USNM 523956—-523977, 523979-
523981, 523983523986, 524386—524392);
1.8 mi NW (by rd) Coapan, 4650 ft (USNM
511701); Estanzuela (USNM 510051-—
510058, 510793-—510799); Rancho Sapoti-
to, 3500 ft (USNM 511702—-511706,
511756); 1 mi SW San José del Conde,
3000 ft (UMMZ 95492-95494); 2 mi E San
Pedro Lagunillas, east side of lake (USNM
509982509998, 510000—5 10002, 510004—
510011, 510014—510016, 510020—510027,
510034, 510038, 510041, 510046, 510048—
510050, 510766, 510767, 510770, 510772—
510774, 510776, 510778, 510779, 510784,
510786, 510787, 510789, 510790, 510792);
2 mi N Santa Isabel, 3800 ft (UMMZ
95490, 95491). Oaxaca: Agua Blanca, 4000
ft (AMNH 145823); Buena Vista (AMNH
145252); Cerro Calderona, 3000 ft (AMNH
147696); Cuicatl4an, 640 m (UMMZ
96801); Escuranos (AMNH_ 143950);
Guiengola, 3600 ft (AMNH 142522,
142523, 142530—142533, 143432, 143951,
147697-147712); Huajuapam, 5500 ft
(USNM 70233, 70235, 70236); Huilotepec
(AMNH 143430); Jalapa (AMNH 171618,
171619, 175188—175191); Juchitan (AMNH
175193—-175197); La Reforma (AMNH
175192; FMNH 14100, 14102); 5 km N
Las Cuevas (AMNH 142524—-142527); Li-
mén (AMNH 186920); Miahuatlan (UMMZ
94032); Mixtequilla (AMNH 175186);
Morro Mazatlan (AMNH 167432, 171607,
171654); Nizanda (AMNH _ 175253); Oa-
xaca, 5000 ft (USNM 68230, 68652-—
68654); Cerro Palma de Oro (AMNH
147717); Cerro Pozo Rio (AMNH 142529,
148816, 148817); Puerto Angel, 100—300 ft
(USNM 71577-71581); Reforma, 50 m and
100 m (UMMZ 96805, 96806); Rinc6én
Bamba (AMNH 148819—-148823); Sacate-
pec (AMNH_ 189288); San Antonio
(AMNH_ 143425-143427, 143477); San

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Dionisio, Buena Vista (AMNH 145992-—
145997); San Pedro Jilotepec (AMNH
145249, 149922, 149923); Santa Lucia
(AMNH 144572); Santiago Lachiguiri
(AMNH 145832-145835); Santo Domingo,
900 ft (AMNH 3096/2415; USNM 73316—
73318); Cerro Sombrerito (AMNH
145831); Tamazulapam, 7000 ft (USNM
70231, 70232); Tapanatepec (AMNH
177093-177100, 177980); Tehuantepec,
150 ft (AMNH 142535, 147713, 147714,
148824, 171608—-171617, 171620, 171621,
171655—171658, 175184, 175185, 175187;
USNM 9672/9508, 9673/9509, 9671/9510,
11907, 13071, 13072, 73315); 50 mi N Te-
huantepec (AMNH 174811); Teotitlan, 950
m (UMMZ 89263, 89264, 91602); Tequis-
istl4n (AMNH 143420—143424); Cerro de
Tigre (AMNH 142534, 147716); Zanatepec
(FMNH 14101). Puebla: Tehuacan (UMMZ
89250-89260, 89323; USNM 9385/8667);
2 mi W Tehuacan, 5200 ft (UMMZ
103188); Tepanco (UMMZ 89261). Quer-
etaro: Cadereyta, 2100 m (UMMZ 93737);
Tequisquiapam (USNM 78427). Zacatecas:
6 km S Apozol, 1170 m (MSU 27240); 16
km SSE Monte Escobedo, 2010 m (MSU
27241-27247); 5 km NE San Juan Capis-
trano, near Las Tablas, 1330 m (MSU
23645); 18 km N San Juan Capistrano, near
Las Tablas, 1100 m (MSU 27234-27239);
2 mi N Santa Rosa, 3850 ft (MSU 11668,
12475); Valparaiso, 6200 ft and 6500 ft
(USNM 92008-92010).

Sigmodon alleni Bailey

Sigmodon alleni Bailey, 1902:112 (type lo-
cality—Meéxico, Jalisco, San Sebastian,
Mascota; holotype—USNM 88227).

Sigmodon vulcani Allen, 1906:247 (type lo-
cality—México, Jalisco, Volcan de Fue-
go, 3050 m; holotype—AMNH 26310).

Sigmodon guerrerensis Nelson & Goldman,
1933:196 (type locality—Meéxico, Gue-
rrero, Omilteme, 8000 ft; holotype—
USNM 126936).

Sigmodon planifrons Nelson & Goldman,
1933:197 (type locality—-México, Oaxa-

VOLUME 112, NUMBER 4

ca, Juquila, 5000 ft; holotype—USNM
71918).

Sigmodon planifrons minor Goodwin,
1955:1 (type locality—Meéxico, Oaxaca,
District of Tehuantepec, 12 km NE Ten-
ango, Santa Lucia, 4000 ft; holotype—
AMNH 143429).

Sigmodon macdougalli Goodwin, 1955:3
(type locality—México, Oaxaca, 12 km
S Yautepec, Santo Tomas Teipan, 7000 ft;
holotype—AMNH 149122).

Sigmodon macrodon Goodwin, 1955:4
(type locality—México, Oaxaca, District
of Tehuantepec, 20 km W Mixtequilla,
Cerro San Pedro; 3600 ft; holotype
AMNH 142536).

Sigmodon planifrons setzeri Goodwin,
1959:447 (replacement name for S. plan-
ifrons minor Goodwin, 1955, preoccu-
pied by S. minor Gidley, 1922).

Distribution.—Pacific coastal lowlands
and contiguous slopes of western cordille-
ras, from southern Sinaloa to central Oa-
xaca (see Baker 1969, Shump & Baker
1978). In Nayarit, at low elevations (sea
level to 760 m) in the southwestern quad-
rant of the state, the deepest interior records
found along the lower valleys of the Rio
San Pedro and the Rio Grande de Santiago
(Fig. 18).

The geographic range of S. alleni, as
presently understood, is partly congruent
with the revised distribution of S$. masco-
tensis. Individuals of the former, however,
apparently prefer more mesic habitats,
which may explain the species’ occurrence
farther north into Sinaloa and at higher el-
evations, up to 3050 m and into pine-oak
vegetation (Baker 1969), as well as its ab-
sence from the upper drainage of the Rio
Balsas and Lago de Chapala, dryer interior
basins where S. mascotensis is found. Nev-
ertheless, the geographic range of S. alleni
deserves further documentation, as we have
encountered many instances of specimens
mistakenly intermixed with S. mascotensis
in museum collections.

Remarks.—In his description of S. alleni,

849

Bailey (1902) recorded the co-occurrence
of his new species with mascotensis at San
Sebastian, Jalisco, and with major (now =
S. arizonae) at San Blas, Nayarit. Based on
these records of overlap, Bailey diagnosed
alleni as a species and relegated all other
members of his hispidus group to subspe-
cies of the latter. While explaining his
choice of patronym, he continued (1902:
113), “It [S. alleni] is the species Dr. Allen
had in mind in writing his description of
mascotensis from which he intended to sep-
arate the Colima form, but as the two spec-
imens of the species here named alleni were
not fully adult he unfortunately selected as
his type another specimen that proves to be
the same as the form to which on the fol-
lowing page he gave the name colimae.”’
Allen (1906) continued to recognize coli-
mae as distinct from mascotensis, but we
concur with Bailey’s taxonomic judgement
regarding both the synonymy of colimae
under mascotensis and their specific dis-
tinction from S. alleni. The possibility of
confusion is understandable, as we noted
above under the morphometric results, par-
ticularly when separating isolated skulls of
S. alleni from those of S. mascotensis.

The junior synonyms that have accrued
to Bailey’s S. alleni, following Baker
(1969) and Shump & Baker (1978), require
substantiation. To us, their arrangement as
a single species seems improbable. Al-
though they exhibit the reddish brown dor-
sum and softer fur typical of alleni sensu
lato, the types of planifrons and planifrons
setzeri are strikingly diminutive in all di-
mensions of the skin and skull—particular-
ly as reflected in the narrow incisors, deli-
cate molar rows, constricted interorbit, tiny
otic capsules, and shorter tail—compared
with alleni-like taxa also described from
Oaxaca (macdougalli and macrodon) or
with typical alleni from farther north (in-
cluding vulcani and guerrerensis). The di-
vergent treatment of planifrons by Good-
win (1969), who retained it as a species,
and Baker (1969), who relegated it to a sub-
species of S. alleni (along with macdougalli

850

and macrodon in full synonymy) warrants
reconciliation. It is noteworthy that Good-
win (1969) had identified both planifrons
and macdougalli from Santa Lucia, Oaxaca,
the type locality of his (1955, 1959) plan-
ifrons setzeri. Renewed study of the Oaxa-
can forms will prove critical to much-need-
ed revision of the S. alleni complex.

Ecological notes.—Other rodents cap-
tured at localities with Sigmodon alleni in-
clude Sciurus colliaei, Spermophilus annu-
latus, Thomomys umbrinus, Liomys pictus,
Baiomys taylori, Hodomys alleni, Neotoma
mexicana, Oryzomys melanotis, O. palus-
tris, Osgoodomys banderanus, Peromyscus
maniculatus, P. melanophrys, P. pectoralis,
P. simulus, and P. spicilegus. Records of
sympatry with other Sigmodon appear to
fall where coastal plain meets southern
foothills, with S. arizonae at San Blas and
with S$. mascotensis at Arroyo de Jiguite.

Specimens examined.—Nayarit: Arroyo
de Jiguite, Rio Santiago, 100 m (USNM
5235939; S25978, 9239827 S235 9GT. 525988);
4 km S Aticama (USNM 524479, 524480);
Chacala (USNM 523934); 1.2 mi S (by rd)
El Casco, Rio Chilte, 480 ft (USNM
511697-511699); 3.5 mi E (by rd) El Ve-
nado (USNM 509981); 1 km S La Villita
(524478); 1 mi S Lo de Marcos, sea level
(USNM 523936—-523943); San Blas
(USNM 89212, 89213); 3.5 mi E San Blas,
100 ft (UMMZ 100680, 100681).

Sigmodon hispidus

Sigmodon berlandieri Baird, 1855:333
(type locality—México, Coahuila, Rio
Nasas; holotype—USNM 566/1687).

Sigmodon borucae Allen, 1897a:40 (type
locality—-Costa Rica, Puntarenas, Boru-
ca; holotype—AMNH 11761/10045).

Sigmodon hispidus eremicus Mearns, 1897:
4 (type locality—México, Sonora, Cie-
nega Well, 30 mi S monument no. 204,
Mexican boundary line; holotype
USNM 60319).

Sigmodon hispidus saturatus Bailey, 1902:

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

111 (type locality—México, Tabasco,
Teapa; holotype—USNM 99998).

Sigmodon hispidus microdon Bailey, 1902:
111 (type locality—MéExico, Yucatan,
Puerto Morelos; holotype—USNM
108467).

Sigmodon borucae chiriquensis Allen,
1904:68 (type locality—Panama, Chiri-
qui, Boqueron; holotype—AMNH
18789).

Sigmodon hispidus griseus Allen, 1908:657
(type locality—Nicaragua, Chontales,
coastal lowlands; holotype—AMNH
28497).

Sigmodon hispidus confinis Goldman,
1918:21 (type locality—United States,
Arizona, Graham County, Safford, 2900
ft; holotype—USNM 204241).

Sigmodon zanjonensis Goodwin, 1932:1
(type locality—Guatemala, Zanjon, 9000
ft; holotype—AMNH 69277).

Sigmodon hispidus villae Goodwin, 1958:2
(type locality—México, Chiapas, 5 mi E
Teopisca, 6000 ft; holotype—AMNH
174799).

Acknowledgments

We thank the many curators and collec-
tions managers for their cooperation and
promptness in making available specimens
for our examination, through loans or mu-
seum visits: Guy G. Musser and Eric M.
Brothers (AMNH); Bruce D. Patterson and
Bill Stanley (FMNH); Robert M. Timm and
Thor Holmes (KU); Barbara Lundrigan and
Laura Abraczinskas (MSU); George Baum-
gardner and Rodney Honeycutt (TCWC);
and Philip Myers (UMMZ). We especially
appreciate the sleuthing efforts of Bob
Timm, who doggedly tracked down certain
obscurely reported specimens from Oaxaca.
David E Schmidt undertook the cranial
photography, line drawings, and distribu-
tion maps; his conscientious efforts and at-
tention to detail, as always, measurably en-
hance our report. The Direccion General de
la Fauna Silvestre, México, kindly provided
collecting permits. This study would not

VOLUME 112, NUMBER 4

have been possible without the coordination
and oversight provided by Don E. Wilson,
leader of the Nayarit Project. During vari-
ous periods of the Nayarit survey, Barbara
A. Bacon, Catherine A. Blount, Michael A.
Bogan, Clyde Jones, Patricia Mehlhop,
Robert Reynolds, C. Brian Robbins, Don E.
Wilson, and Bruce Woodward participated
as field workers who procured the speci-
mens reported here. Finally, we thank Joa-
guin Arroyo-Cabrales, Robert D. Bradley,
and Robert S. Voss, who improved the final
version of our study with their helpful re-
views.

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854 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Appendix.—Descriptive statistics for selected cranial and external variables and OTUs of Sigmodon.

Species OTU n Mean Range SD

Total length

S. alleni OTU 1 14 245.8 221.0—278.0 16.3

S. arizonae OTs 15 305.5 247.0—363.0 33.6

S. mascotensis OPUES 20 278.1 242.0—313.0 20.1

OTU 6 32 257.0 220.0—305.0 21.4

S. hispidus OTU 9 44 250.8 220.0—282.0 18.4

Tail length

S. alleni ORC | 14 110.2 91.0—-130.0 LO.F

S. arizonae OTru3 15 124.4 101.0—-145.0 13.6

S. mascotensis OFfriis 20 132.3 110.0-156.0 12.3

OTU 6 5p i23 96.0—-141.0 10.5

S. hispidus OTU 9 44 99.1 81.0-115.0 8.6

Hindfoot length

S. alleni OTe! Py 30:2 27.0—32.0 1.4

S. arizonae Of. 19 38.5 34.0-43.0 2:3

S. mascotensis OU a 34.1 31.0—38.0 1.9

OTU 6 33 35.0 31.0-38.0 et

S. hispidus OTU 9 | d383 31.0—-36.0 3

Mass (Weight)

S. alleni OTe t 17 JOS 49.0—106.0 15.9

S. arizonae OTUs 19 150.3 83.0—300.0 54.8

S. mascotensis OFUS Tih 99.6 63.0—-148.0 22.0

OTU 6 33 TEs 51.0—-122.0 17.8

S. hispidus OTU 9 15 89.8 68.0-144.0 18.3
Occipitonasal length

S. alleni OTU I ied! Soe 27.9-36.0 Pippi

S. arizonae OTU 3 A 32 27.440.4 ee)

S. mascotensis OTS 27 32.8 29.8—-37.0 1.9

OTU 6 40 31.6 27.5—35.3 ny

S. hispidus OTU 9 50 34.6 30.2—37.7 1.8
Zygomatic breadth

S. alleni OTe *1 20 17.8 15.8-19.8 1.0

S. arizonae OFU-3 25 20.9 17.4-23.8 1.6

S. mascotensis GEES 2H 18.7 17.0—20.7 1.0

OTU 6 45 18.3 15.6—20.7 1.0

S. hispidus OTU 9 5S [oe 16.3-21.1 1.0
Interorbital breadth

S. alleni OTU 1 19 5.0 4.6—5.6 0.1

S. arizonae OFU 3 DS Ye | 4.5—5.6 0.3

S. mascotensis OTU 5 Dey 4.9 4.5—5.5 0.3

OTU 6 A5 5.0 4.6—-5.4 0.2

S. hispidus OTU 9 59 5:1 4.45.7 ok
Breadth of braincase

S. alleni OTU 1 19 13.7 12.9-14.3 0.4

S. arizonae OEW 3 7a, | 14.9 13.9-16.3 0.6

S. mascotensis OEU'S 27 14.0 13.1—-14.7 0.4

Oru 6 A3 13.9 12.6—-14.6 0.4

S. hispidus OFU 9 33 14.4 13.3—15.4 0.5

VOLUME 112, NUMBER 4 855

Appendix.—Continued.
Species OTU n Mean Range SD
Breadth of occipital condyles
S. alleni ORI 20 3 6.7-7.8 0.3
S. arizonae OTUs 20 8.1 6.9-8.8 0.5
S. mascotensis OFS 21 7.4 6.9-8.0 0.3
OTL 6 41 yea 6.9-8.2 0.3
S. hispidus OTU 9 52 Mi, 6.1—8.0 0.3
Depth of braincase
S. alleni OTU 1 19 10.2 9.3-11.3 0.5
S. arizonae OTU.3 19 Lis7 9.9-12.9 0.9
S. mascotensis OTU 5 27 ti3 10.8—12.2 0.3
OTU 6 41 10.8 8.9-11.7 0.5
S. hispidus OTU 9 BPs ID. 9.8-11.6 0.4
Distance between temporal ridges
S. alleni OTUs 20 33 2.7—3.9 0.3
S. arizonae OFUS 24 4.4 3.3-5.0 0.4
S. mascotensis OTU 5 27 4.0 3.34.8 0.4
OTU 6 45 ed 2.54.4 0.4
S. hispidus OTU 9 53 BEV 2.5—4.1 0.3
Length of rostrum
S. alleni OTU 1 iar POST 9.1-12.5 1.0
S. arizonae OTu:3 aS TRG 8.3-13.7 KS
S. mascotensis OTs 27 10.6 9.0-12.7 0.9
OTU 6 43 10.3 8.1-11.7 0.7
S. hispidus OTU 9 50 11.6 9.8-13.0 0.8
Breadth of rostrum
S. alleni OTU 1 20 65 5.5-7.5 0.5
S. arizonae OTU 3 24 7.0 5.4-8.9 0.8
S. mascotensis Ore 5 As | 6.4 5.5-7.4 0:5
OTU 6 45 6.3 5.3-7.2 0.4
S. hispidus OTU 9 53 6.7 5.7-7.8 0.5
Postpalatal length
S. alleni OTe :-4 19 11.2 9.3-12.8 0.9
S. arizonae OTe: 3 20 12.8 9.6—-15.5 1.6
S. mascotensis OFU5S 27 bAS 10.0—13.2 0.8
OTU 6 4] 10.9 9.2-12.7 0.7
S. hispidus OTU 9 ae i a | 5.6—7.3 0.4
Length of bony palate
S. alleni OFWU 1 20 6.1 5.4-7.1 0.5
S. arizonae OTU.3 25 6.6 5.3-8.0 0.5
S. mascotensis OTW.5 pay | 6.1 5.2-6.9 0.4
OTU 6 45 6.1 5.1-6.8 0.4
S. hispidus OTU 9 53 6.6 5.6-7.3 0.4
Breadth of bony palate
S. alleni OTU 1 20 1-2, 6.7-8.1 0.4
S. arizonae OTU 3 25 8.1 6.7—9.2 0.5
S. mascotensis OTU.S 27 te 6.7—8.0 0.4
OTU 6 45 7.4 6.8-8.0 0.3

S. hispidus OTU 9 33 rea 6.7—8.5 0.3

856 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Appendix.—Continued.

Species OTU n Mean Range SD

Length of incisive foramen

S. alleni ORE 1 20 6.7 5.8-7.6 0.5
S. arizonae OTU 3 25 8.0 6.2-9.7 0.9
S. mascotensis OTU 5 27 6.9 5.9-8.2 0.6
OTU 6 45 6.7 5.5-7.8 0.5
S. hispidus OTU 9 33 7.8 6.2—9.0 0.6
Length of diastema
S. alleni OU 20 8.6 7.2-10.2 0.8
S. arizonae OTU 3 25 9.8 7.3-12.1 a3
S. mascotensis OTU 5 27 8.7 7.6—10.3 0.7
OTU 6 45 8.3 6.3—9.7 0.7
S. hispidus OFRUS 53 2.5 7.7-11.2 0.8
Breadth of zygomatic plate
S. alleni OF? 20 56 3.04.1 0.3
S. arizonae OU 3 25 4.1 3.1-5.1 0.5
S. mascotensis OLS psy 3.6 2.84.5 0.4
OTU 6 45 7 3.04.6 0.3
S. hispidus OTU 9 53 a9 2.64.8 0.4
Length of zygomatic spine
S. alleni OTW | 20 3.8 3.14.4 0.4
S. arizonae OTU 3 25 5.0 3.6—-5.9 0.7
S. mascotensis OF 5 Zh 4.3 3.34.9 0.4
OTU 6 46 4.4 3225555 0.5
S. hispidus OTU 9 33 4.5 3.2—5.5 0.5
Length of auditory bulla
S. alleni OTU 1 20 =a | 4.6-5.6 0.3
S. arizonae ORE) 3 24 6.1 5.3-6.8 0.3
S. mascotensis OTU 5 ZT SES 4.9-5.8 0.2
OTU 6 43 5.3 4.7-5.8 0.2
S. hispidus OU 9 33 6.3 5.7—7.0 0.3
Length of maxillary toothrow
S. alleni OFW 1 20 5.70 5.36—-6.13 0.19
S. arizonae OFU 3 25 6.82 6.30—7.39 0.25
S. mascotensis OFWS 27 5.95 5.59-6.78 0.26
OTU 6 46 6.23 5.80—6.63 0.23
S. hispidus OTU 9 53 6.39 6.00-6.81 0.18
Width of upper first molar
S. alleni OTU 1 20 1.90 1.76—2.03 0.08
S. arizonae OPUS 25 2.29 2.12—2.47 0.09
S. mascotensis OFU'S ZT 2.01 1.84—2.26 0.09
OTU 6 46 2.09 1.94-2.34 0.08
S. hispidus OTU 9 53 2. t3 1.91—2.32 0.08
Depth of upper incisor
S. alleni OTU 1 20 1.81 1.47-2.12 0.09
S. arizonae OFU 3 25 2.01 1.53—2.37 0.22
S. mascotensis OTUs Dy 1.81 1.52—2.07 0.08
OTU 6 46 1.85 1.47-2.17 0.15
S. hispidus OTU 9 33 1.99 1.52—2.22 0.16
Depth of mandible
S. alleni OfUa 20 5.8 5.1-6.6 0.4
S. arizonae OFS 25 6.6 5.2-7.6 0.6
S. mascotensis OTrus ps5 3:9 5.3-6.7 0.4
OTLT.G 46 5.8 4.7-6.7 0.4

S. hispidus OTU 9 53 6.4 5.47.2 0.4

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CONTENTS

Abyssal sea anemones (Cnidaria: Actiniaria) of the northeast Pacific symbiotic with molluscs:
Anthosactis nomados, a new species, and Monactis vestita (Gravier, 1918)
Tracy R. White, April K. Wakefield Pagels, and Daphne G. Fautin
Urticina mcpeaki, a new species of sea anemone (Anthozoa: Actiniaria: Actiniidae) from the
North American Pacific coast J. Susanne Hauswaldt and Katherine E. Pearson
Morphological observations on a hatchling and a paralarva of the vampire squid, Vampyroteu-
this infernalis Chun (Mollusca: Cephalopoda) Richard E. Young and Michael Vecchione
Nereidids (Polychaeta) from the Caribbean Sea and adjacent Coral Islands of the southern Gulf
of Mexico
Jesus Angel de Leoén-Gonzdlez, Vivianne Solis-Weiss, and Victor Ochoa Rivera
A new stygobitic Calanoida (Crustacea: Copepoda) of the genus Stygodiaptomus Petkovski,
1981 from the Balkan Peninsula Tomislav Karanovic
The freshwater centropagid Osphranticum labronectum Forbes, 1882 (Crustacea: Copepoda:
Calanoida) in Mexico with description of a new subspecies
M. A. Gutiérrez-Aguirre and E. Suarez-Morales
New records of isopods from the Indian River Lagoon, Florida (Crustacea: Peracarida)
Brian Kensley and Marilyn Schotte
A new species of Pseudostegias Shiino, 1933 (Crustacea: Isopoda: Bopyridae: Athelginae)
parasitic on hermit crabs from Bali Jason D. Williams and Christopher B. Boyko
Caprella kuroshio, a new species (Crustacea: Amphipoda: Caprellidae), with a redescription
of Caprella cicur Mayer, 1903, and an evaluation of the genus Metacaprella Mayer, 1903
Atsushi Mori
Palaemonella hachijo, a new species of shrimp (Crustacea: Decapoda: Palaemonidae) from a
submarine cave in southern Japan Junji Okuno
A new species of Macrobrachium (Crustacea: Decapoda: Palaemonidae), with abbreviated
development, from Veracruz, Mexico José Luis Villalobos and Fernando Alvarez
Description of a new spider crab, Maja gracilipes, from the South China Sea, with notes on
the taxonomic validity of M. brevispinosis Dai, 1981 (Crustacea: Decapoda: Brachyura:
Majidae) H.-L. Chen and Peter K. L. Ng
On the identities of two Pacific species of deep-water porter crabs, Hypsophrys longirostris
Chen, 1986, and Homologenus donghaiensis Chen, 1986 (Crustacea: Decapoda: Brachyura:
Homolidae) Peter K. L. Ng and H.-L. Chen
Neocallichirus raymanningi, a new species of ghost shrimp from the northeastern coast of
Venezuela (Crustacea: Decapoda: Callianassidae)
Juan Pablo Blanco Rambla and Rafael Lemaitre
Cucumaria flamma, a new species of sea cucumber from the central eastern Pacific (Echino-
dermata: Holothuroidea) Francisco A. Solis-Marin and Alfredo Laguarda-Figueras
A new species of rainfrog of the Eleutherodactylus cruentus group from eastern Honduras
(Amphibia: Anura: Leptodactylidae)
Jay M. Savage, James R. McCranie, and Larry David Wilson
A new species of arboreal viper (Serpentes: Viperidae: Atheris) from Cameroon, Africa
Dwight P. Lawson
Taxonomic notes on hummingbirds (Aves: Trochilidae) 2. Popelairia letitiae (Bourcier &
Mulsant, 1852) is a valid species Gary R. Graves
Identification and distribution of cotton rats, genus Sigmodon (Muridae: Sigmodontinae), of
Nayarit, México Michael D. Carleton, Robert D. Fisher, and Alfred L. Gardner
Contents, Volume 112
Index to New Taxa, Volume 112

682

687

695

714

hee.

739

746

754

759

768

778

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