Catalog of the Type Specimensf Seastars (Echinodermata: Asteroidea

in the National Museum of NaturalHistory, Smithsonian Institution

CYNTHIA GUST AHEARN

I

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 572

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S M I T H S O N I A N C O N T R I B U T I O N S T O Z O O L O G Y • N U M B E R 5 7 2

Catalog of the Type Specimensof Seastars (Echinodermata: Asteroidea)

in the National Museum of NaturalHistory, Smithsonian Institution

Cynthia Gust Ahearn

SMITHSONIAN INSTITUTION PRESS

Washington, D.C.

1995

A B S T R A C T

Ahearn, Cynthia Gust. Catalog of the Type Specimens of Seastars (Echinodermata:Asteroidea) in the National Museum of Natural History, Smithsonian Institution. SmithsonianContributions to Zoology, number 572,59 pages, 1995.—The asteroid type specimens depositedin the National Museum of Natural History comprise 1183 lots representing 552 nominalspecies, subspecies, and varieties in 179 nominal genera. A taxonomic list of types reflects theircurrent taxonomic status; an annotated alphabetic list is also provided. The format includes theoriginal bibliographic citation for each species, catalog number, number of specimens,preservation, locality and collector data, synonymies, and remarks concerning status. Anappendix of locality information is presented alphabetically by research vessel. A bibliographyof relevant publications is provided. Problems concerning the status of W.K. Fisher's typespecimens are summarized in the introduction and specifically resolved in the text.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and isrecorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coralMontastrea cavernosa (Linnaeus).

Library of Congress Cataloging-in-Publication DataNational Museum of Natural History (U.S.)Catalog of the type specimens of seastars (Echinodermata: Asteroidae) in the National Museum of Natural History,Smithsonian Institution / Cynthia Ahearn.p. cm.—(Smithsonian contributions to zoology ; no. 572)Includes bibliographical references.1. Starfishes—Type specimens—Catalogs and collections—Washington, (D.C.) 2. National Museum of Natural

History (U.S.)—Catalogs. I. Ahearn, Cynthia. II. Title. III. Series.QL1.S54 no. 572 [QL384.A8] 592 s-dc20[593.9T074753] 95-3644

® The paper used in this publication meets the minimum requirements of the AmericanNational Standard for Permanence of Paper for Printed Library Materials Z39.48—1984.

Contents

page

Introduction 1Format 1Special Problems 1Acknowledgments 2

type Specimens Labeled by W.K. Fisher but not Referred to by USNM CatalogNumber in Original Publication 2

Taxonomic List of Asteroid Types Deposited in the National Museum of NaturalHistory 3

Annotated Alphabetic List of Asteroid Types Deposited in the National Museum ofNatural History 8

Appendix: List of Stations 42Literature Cited 55

in

Catalog of the Type Specimensof Seastars (Echinodermata: Asteroidea)

in the National Museum of NaturalHistory, Smithsonian Institution

Cynthia Gust Ahearn

Introduction

The asteroid type specimens in the United States NationalMuseum (USNM), now the National Museum of NaturalHistory (NMNH), comprise 1183 type lots representing 552nominal species, subspecies, and varieties in 179 nominalgenera. The author has examined and verified every typespecimen listed. This is the first comprehensive publishedreport of asteroid types in the NMNH, although speciesdescribed by W.K. Fisher and deposited in both the CaliforniaAcademy of Sciences (CAS) and the USNM have beenreported (Chaffee and Weitbrecht, 1984).

FORMAT.—The catalog is arranged as two lists, an annotatedlist and a taxonomic list. The taxonomic list is arranged byfamily in the classification outline proposed by Clark andDowney (1992:1-2); within the families, taxa are arrangedalphabetically by genus, and then by species. Currentlyaccepted generic combinations are used, and in some instances,it was necessary to create unpublished (junior synonyms)combinations in order to include the original species name inthis listing.

In the annotated list of type specimens, the arrangement isalphabetical according to the specific names used in theoriginal descriptions, and the information provided has beenstructured for consistency as follows: Species, or subspecies,Genus, (Subgenus), Author, year.page(s), plate(s), figure(s).

Type status, USNM catalog number, number of specimen(s),preservation, collecting vessel, station number, modifyinglocality, exact locality, depth, collectors), expedition, datecollected, condition of specimen(s) (where applicable), originaldepository and catalog number.

Cynthia Gust Ahearn, Department of Invertebrate Zoology, NationalMuseum of Natural History, Smithsonian Institution, Wahington D.C.20560.

Remarks include any necessary verification and discussionof type status and correction of erroneous published informa-tion concerning the type(s) or type locality.

Synonymy, where applicable, includes first reference to thetaxonomic change.

In 1920, A.H. Clark, curator of Echinoderms at the USNM,began the system of assigning catalog numbers prefixed withthe letter E. Prior to this date, echinoderms shared a singleseries of catalog numbers with the coelenterate collections andno letter prefix was assigned. Both numbering systems arereflected in the asteroid type collection. Occasionally, a typespecimen was incorrectly cataloged twice, and I used which-ever catalog number was published—even if that was the morerecent number, rather than the original catalog number.

An appendix of locality descriptions is provided for typespecimens collected by a research vessel and assigned a stationnumber. The appendix is arranged alphabetically by vessel.Collection information for specimens with descriptive localityor specimens from other sources, such as individual collectors,is provided in the text.

The bibliographic information includes both publications inwhich at least one species was described and the type(s)deposited in the USNM and publications cited in the text forspecies in which some taxonomic change has occurred, ingeneric placement, in documentation of synonyms, or indesignations of lectotypes and neotypes.

SPECIAL PROBLEMS.—Nearly half of the asteroid types in the

USNM were described and deposited by W.K. Fisher between1905 and 1940. Fisher described many starfish species inpreliminary reports and then expanded his descriptions withmore specimens in his larger monographs (1911a, 1919a,1928a, 1930). Due to this practice, the USNM asteroid typecollection held hundreds of specimens that were examined byFisher for inclusion in his monographs but can not be

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

considered type material. Unfortunately, the California Acad-emy of Sciences published an asteroid type catalog (Chaffeeand Weitbrecht, 1984) based on incorrect information con-tained within the USNM collections. The original descriptionof a species, no matter how brief it may be in Fisher'spreliminary reports, must be the original citation. Only thespecimens properly documented in the original citation canhave type status, and the specimens examined by Fisher insubsequent monographs are simply voucher specimens orhypotypes and are not now included in the asteroid typecollection.

In several of his preliminary reports, Fisher designated thetype by a USNM catalog number, and in these instances, thestatus of the type material is unequivocal. However, he did notalways clearly designate types, nor did he always publishcatalog numbers in his reports. A list of type specimens labeledby Fisher but not referred to by USNM catalog number in theoriginal publication precedes the taxonornic and annotatedalphabetic listing of types herein.

Usually, Fisher described a single specimen and providedprecise measurements for that specimen; sometimes hementioned an Albatross station number or an Albatross localitythat can be matched to a station number. In these cases,although the USNM catalog number is not published in theoriginal description, the USNM holds the specimen with W.K.Fisher's handwritten type label. According to Article 73(a)(ii)of the International Code of Zoological Nomenclature (ICZN,1985), this is evidence of Fisher's intent to name the holotype,and the specimen is the holotype by monotypy.

In many of his preliminary reports, W.K. Fisher recordedAlbatross locality information without stating the associatedAlbatross station number. The annotated type list records justthe station number. Some Albatross station localities wererecorded without providing latitudes and longitudes and thesewill have the descriptive localities provided in the text ratherthan the appendix.

Due to the large number of Fisher's preliminary reports withbrief descriptions, complete references to his monographicreports are provided following the original citation for eachnew species in the annotated list.

In Parts 2 and 3 of his Asteroidea of the North Pacific andAdjacent Waters (1928a; 1930), W.K. Fisher described bothnew subspecies and new forms of various taxa, which indicatesthat he clearly intended to use both ranks for his new taxa.According to Article 45(g)l of the ICZN, the rule ofinterpretation can be applied to Fisher's intent to maintain hisinfrasubspecific ranks and therefore these are excluded fromthe asteroid type collection.

ACKNOWLEDGMENTS.—I wish to thank Frederick M. Bayer,Stephen D. Cairns, and David L. Pawson, all of the Departmentof Invertebrate Zoology, Smithsonian Institution, for theirassistance in interpretation of the International Code ofZoological Nomenclature. I would also like to acknowledge thehelp of Curtis W. Sabrosky, former Chairman of the Editorial

Committee of the 1985 edition of the ICZN, who confirmed ourinterpretations of the Code. I wish also to thank Daniel B. Blake(University of Illinois, Urbana, Illinois) and Philip Lambert(Royal British Columbia Museum, Victoria, British Columbia)who kindly reviewed the manuscript and offered usefulsuggestions for improvement.

Type Specimens Labeled by W.K. Fisherbut not Referred to by USNM Catalog Number

in Original Publication

acanthonotus, Benthopecten Fisher, 1905aciculosus, Acantharchaster Fisher, 1910cactinocles, Zoroaster Fisher, 1919bactinodetus, Astrocles Fisher, 1917dalascensis, Pseudarchaster Fisher, 1905annectens, subsp., Henricia leviuscula Fisher, 1910danoplus, Dipsacaster Fisher, 1910casthenactis, Henricia Fisher, 1910dboreal is, Dipsacaster Fisher, 1910ccalifornica, Hippasteria Fisher, 1905clarki, Ceramaster Fisher, 1910cclarki, Cladaster Fisher, 1910dclaviger, Benthopecten Fisher, 1910ccoscinactis, Anteliaster Fisher, 1923acoscinopeplus, Pteraster Fisher, 1910acrassus, Odontaster Fisher, 1905dissonus, Pseudarchaster Fisher, 1910cectenes, subsp., Myxoderma sacculatum Fisher, 1919bevermanni, Zoroaster (Myxoderma) Fisher, 1905evoplus, subsp., Cheiraster agassizi Fisher, 1910cexiguus, Solaster Fisher, 1910dexilis, Brisinga Fisher, 1905eximius, Dipsacaster Fisher, 1905fecunda, Freyella Fisher, 1905flexilis, subsp., Poraniopsis inflata Fisher, 1910dfurcilliger, Lophaster Fisher, 1905gilberti, Dytaster Fisher, 1905heathi, Hippasteria Fisher, 1905intermedius, Acantharchaster Fisher, 1910cjordani, Pter aster Fisher, 1905koehleri, Hymenaster Fisher, 1910alaetmophilus, Dipsacaster Fisher, 1910cleiopelta, Hippasteria Fisher, 1910clepidonotus, Cryptopeltaster Fisher, 1905leptocerama, Tosia Fisher, 1905longispina, Henricia Fisher, 1910dmagister, Pedicellaster Fisher, 1923amarsippus, Pter aster Fisher, 1910amicroplax, Freyellidea Fisher, 1917dmordax, subsp., Zoroaster evermanni Fisher, 1919bmultispina, subsp., Henricia leviuscula Fisher, 1910dmutabilis, Benthopecten Fisher, 1910cophiurus, Zoroaster Fisher, 1905

NUMBER 572

pectinatus, Bathybiaster Fisher, 1905pedicellaris, subsp., Acantharchaster variabilis Fisher, 1910cpenichra, Lithosoma Fisher, 1917bpenicillatus, Persephonaster Fisher, 1905perissonotus, Hymenaster Fisher, 1910apropinquus, Leptychaster Fisher, 1910cpusilla, Brisingella Fisher, 1917dpus ill us, Pseudarchaster Fisher, 1905pycnopodia, Aphanasterias Fisher, 1923bquadrispinosus, Hymenaster Fisher, 1905rhomaleum, subsp., Myxoderma platyacanthum Fisher, 1919bsacculatus, Zoroaster (Myxoderma) Fisher, 1905stereosomus, Echinaster Fisher, 1913bswifti, Mimaster Fisher, 1905synaptoma, Craterobrisinga Fisher, 1917dtemnochiton, Pteraster Fisher, 1910atenebrarius, Porcellanaster (Eremicaster) Fisher, 1905tenellus, Mediaster Fisher, 1905trigonodon, Pteraster Fisher, 1910avalidus, Cladaster Fisher, 1910cvariabilis, Acanthar chaster Fisher, 1910cvexator, subsp., Lophaster furcilliger Fisher, 1910d

Taxonomic List of Asteroid Types Depositedin the National Museum of Natural History

Class ASTEROIDEAOrder PAXILLOSIDA

Family LUIDIIDAELuidia armata Ludwig, 1905Luidia asthenosoma Fisher, 1906aLuidia avicularia Fisher, 1913cLuidia bernasconiae A.H. Clark, 1945Luidia ferruginea Ludwig, 1905Luidia gymnochora Fisher, 1913cLuidia heterozona Fisher, 1940Luidia hystrix Fisher, 1906bLuidia ludwigi Fisher, 1906aLuidia magnifica Fisher, 1906bLuidia orientalis Fisher, 1913cLuidia ported A.H. Clark, 1917cLuidia prionota Fisher, 1913cLuidia sarsi africana Sladen, 1889Luidia superba A.H. Clark, 1917d

Family ASTROPECTINIDAEAstromesites compactus Fisher, 1913aAstropecten acanthifer phragmorus Fisher, 1913aAstropecten americanus (Verrill, 1880b)Astropecten americanus var. subgracilis Verrill, 1915Astropecten articulatus (Say, 1825)Astropecten benthophilus Ludwig, 1905Astropecten brasiliensis Miiller and Troschel, 1842Astropecten callistus Fisher, 1906bAstropecten comptus Verrill, 1915

Astropecten ctenophorus Fisher, 1906bAstropecten eremicus Fisher, 1913aAstropecten eucnemis Fisher, 1919aAstropecten exiguus Ludwig, 1905Astropecten hawaiiensis Doderlein, 1917Astropecten irregularis pontoporeus (Sladen, 1883)Astropecten luzonicus Fisher, 1913aAstropecten nitidus Verrill, 1915Astropecten nitidus var. forcipatus Verrill, 1915Astropecten nuttingi Verrill, 1915Astropecten ornatissimus Fisher, 1906aAstropecten pectinatus Sladen, 1883Astropecten pedicellaris Fisher, 1913aAstropecten productus Fisher, 1906bAstropecten pusillulus Fisher, 1906bAstropecten sulcatus Ludwig, 1905Astropecten tenellus Fisher, 1913aAstropecten vappa inaequalis Fisher, 1919aBathybiaster robustus (Verrill, 1884)Ctenophoraster diploctenius Fisher, 1913aCtenophoraster hawaiiensis Fisher, 1906bCtenophoraster marquesensis Marsh, 1974Ctenopleura astropectinides Fisher, 1913aDipsacaster anoplus Fisher, 1910cDipsacaster antillensis Halpem, 1968Dipsacaster borealis Fisher, 1910cDipsacaster diaphorus Fisher, 1913aDipsacaster eximius Fisher, 1905Dipsacaster imperialis Fisher, 1917bDipsacaster laetmophilus Fisher, 1910cDipsacaster nesiotes Fisher, 1906bDytaster demonstrans Ludwig, 1905Dytaster gilberti Fisher, 1905Dytaster grandis grandis (Verrill, 1884)Koremaster evaulus (Fisher, 1913a)Koremaster evaulus spiculatus Fisher, 1919aLeptychaster anomalus Fisher, 1906aLeptychaster inermis (Ludwig, 1905)Leptychaster pacificus Fisher, 1906aLeptychaster propinquus Fisher, 1910cPatagiaster nuttingi Fisher, 1906bPatagiaster sphaerioplax Fisher, 1913aPersephonaster anchistus Fisher, 1913aPersephonaster cingulatus (Fisher, 1906b)Persephonaster euryactis Fisher, 1913aPersephonaster euryactis brevispinus Fisher, 1919aPersephonaster habrogenys Fisher, 1913aPersephonaster luzonicus Fisher, 1913aPersephonaster monostoechus Fisher, 1913aPersephonaster multicinctus Fisher, 1913aPersephonaster oediplax Fisher, 1913aPersephonaster suluensis Fisher, 1913aPersephonaster tenuis Fisher, 1913aPlutonaster agassizi (Verrill, 1880b)

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Plutonaster sirius A.H. Clark, 1917cPsilaster abyssicola (Ludwig, 1905)Psilaster andromeda florae (Verrill, 1878)Psilaster armatus Ludwig, 1905Psilaster attenuatus Fisher, 1906bPsilaster gotoi Fisher, 1913aPsilaster pectinatus (Fisher, 1905)Psilaster robustus Fisher, 1913aPsilaster sladeni Ludwig, 1905Tethyaster aulophora (Fisher, 191 Id)Tethyaster canaliculatus (A.H. Clark, 1916c)Tethyaster grandis (Verrill, 1899)Thrissacanthias penicillatus (Fisher, 1905)Tritonaster craspedotus Fisher, 1906bTritonaster evorus Fisher, 1913aTrophodiscus almus Fisher, 1917c

Family RADIASTERIDAEGephyreaster swifti (Fisher, 1905)Radiaster notabilis (Fisher, 1913a)

Family PORCELLANASTERIDAEBenthogenia cribellosa Fisher, 191 IdEremicaster pacificus (Ludwig, 1905)Eremicaster tenebrarius (Fisher, 1905)Eremicaster vicinus (Ludwig, 1907)Eremicaster vicinus var. inermis (Ludwig, 1907)Eremicaster waltharii (Ludwig, 1905)Hyphalaster moseri Ludwig, 1905Porcellanaster nudus (Ludwig, 1907)Porcellanaster semimarginalis (Ludwig, 1905)Sidonaster psilonotus Fisher, 1913aStyracaster monacanthus Ludwig, 1907Styracaster paucispinus Ludwig, 1907Thoracaster magnus Ludwig, 1907

Family GONIOPECTINIDAE

Ctenodiscus orientalis Fisher, 1913aGoniopecten asiaticus Fisher, 1913aPrionaster analogus Fisher, 1913aPrionaster elegans Verrill, 1899Prionaster gracilis Fisher, 1913aPrionaster megaloplax Fisher, 1913a

Order NOTOMYOTIDAFamily BENTHOPECTINIDAE

Benthopecten acanthonotus Fisher, 1905Benthopecten claviger Fisher, 1910cBenthopecten cognatus (Ludwig, 1905)Benthopecten moluccanus Fisher, 1913cBenthopecten mutabilis Fisher, 1910cBenthopecten pectinifer (Ludwig, 1905)Benthopecten polyctenius Fisher, 1913cBenthopecten spinosus Verrill, 1884Benthopecten spinuliger (Ludwig, 1905)Benthopecten styracius Fisher, 1913cCheiraster (Christopheraster) blakei A.M. Clark, 1981Cheiraster (Cheiraster) diomedeae Fisher, 1917b

Cheiraster dispar (Verrill, 1915)Cheiraster enoplus Verrill, 1915Cheiraster gracilis (Verrill, 1915)Cheiraster (Christopheraster) horridus Fisher, 1906bCheiraster (Cheiraster) inops Fisher, 1906bCheiraster (Cheiraster) ludwigi Fisher, 1913cCheiraster mixtus (Verrill, 1915)Cheiraster (Cheiraster) planus Verrill, 1915Cheiraster (Luidiaster) robustus (A.H. Clark, 1917c)Cheiraster (Cheiraster) sepitus (Verrill, 1885)Cheiraster (Cheiraster) snyderi Fisher, 1906bCheiraster (Cheiraster) triplacanthus Fisher, 1913cMyonotus intermedius (Fisher, 1910c)Nearchaster aciculosus (Fisher, 1910c)Nearchaster insignis (A.H. Clark, 1916c)Nearchaster pedicellaris (Fisher, 1910c)Nearchaster variabilis (Fisher, 1910c)Pectinaster agassizi (Ludwig, 1905)Pectinaster agassizi evoplus (Fisher, 1910c)Pectinaster mimicus hylacanthus (Fisher, 1913c)Pectinaster mimicus palawanensis Fisher, 1919a

Order VALVATIDA

Family ODONTASTERIDAEAcodontaster propinquus (A.H. Clark, 1917a)Odontaster crassus Fisher, 1905Odontaster hispidus Verrill, 1880bOdontaster robustus Verrill, 1899Odontaster setosus Verrill, 1899

Family GANERIIDAEAleutiaster schefferi A.H. Clark, 1939bHyalinothhx millespina Fisher, 191 lbPerknaster sladeni georgianus Fisher, 1940Tarachaster tenuis Fisher, 1913c

Family ASTERINIDAEAnseropoda antarctica Fisher, 1940Anseropoda insignis Fisher, 1906bAnseropoda macropora Fisher, 1913cAsterina corallicola Marsh, 1977Asterina coronata euerces (Fisher, 1917b)Asterina orthodon Fisher, 1922bAsterinopsis pedicellaris (Fisher, 1913c)Mirastrella biradialis Fisher, 1940Nepanthia variabilis H.L. Clark, 1938Paranepanthia platydisca (Fisher, 1913c)Paxillasterina pompom A.M. Clark, 1983Pseudonepanthia gotoi A.H. Clark, 1916aTremaster mirabilis Verrill, 1879a

Family PORANIIDAEChondraster grandis (Verrill, 1878)Porania pulvillus insignis Verrill, 1895Poraniomorpha abyssicola (Verrill, 1895)Poraniomorpha spinulosa (Verrill, 1879a)Poraniopsis inflata flexilis Fisher, 1910dPoraniopsis japonica Fisher, 1939